University of Miami

Scholarly Repository
Studies in Tropical Oceanography University of Miami Press

1963

Systematics and Life History of the Great

Barracuda, Sphyraena barracuda (Walbaum)
Donald P. De Sylva

Follow this and additional works at: http://scholarlyrepository.miami.edu/trop_ocean

Part of the EnvironmentalSciencesCommons, MarineBiologyCommons, and the
OceanographyandAtmosphericSciencesandMeteorologyCommons

Recommended Citation
De Sylva, Donald P., "Systematics and Life History of the Great Barracuda, Sphyraena barracuda (Walbaum)" (1963). Studies in
Tropical Oceanography. Book 1.
http://scholarlyrepository.miami.edu/trop_ocean/1

This Book is brought to you for free and open access by the University of Miami Press at Scholarly Repository. It has been accepted for inclusion in
Studies in Tropical Oceanography by an authorized administrator of Scholarly Repository. For more information, please contact
repository.library@miami.edu.
 •

Frontiispiece. Sphyraena barracuda, U .S.N.M . No. 11449978, from Florida. From

George B. Goode, "“T Thhe fiisshheerriies and fiishhery industrriies of thhe United Staatteess,'”' 1188884,
Section 11,, Plate 11778. Drawing by H. L. Todd. Cover drraaw wiing by Cr aig Phillips.
 S tudie s in Tropical Oceanography N o . 1
Institute o f Marine and Atmo spheric Science s
University o f Miami

Sys tema tics and L ife H is tory of the

Great Barracuda
Sphyraena barracuda (Walbaum)

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

Un ivers ity of Miami Press

Cora l Gab les, Flor ida
 This paper may be referred to as:
Stud. trop. Oceanogr. Miami, 1: viii + 1 7 9 pp.,
32 Tables, 36 Figure s, October , 1963

Second prin ting , 1970

Prin ted in the Un ited States of America

 C o nte nts

A b stract ................................ . •........................................................................................ 1

I n t r o d u c t i o n ........................................................... . ...................................................... 2
A c k no w le dg m e nt s ..................................................................................... ,............ 3
Sy s t e m a t i c s ...................................................................................................................... 5
P h y lo g en e tic R e l a tio n sh ip s of the F am ily S p h y r a e n i d a e ................. 5
F oss il D is tr ib u tio n of the F am ily S p h y r a e n i d a e ..................................... 5
R e c e n t D i s t r i b u t i o n ........................................................................................ 6
D e lin ea tion of the G en u s S p h y r a e n a ............................................................... 7
Sy n o n y m y of Sphyraena b a r r a c u d a ............................................................. 7
M e t h o d s o f M easuri ng and C o u n t i n g ........................................................ 22
C o m p a r i so n s .................................................................................................... 24
Y o u ng S tages of the F a m i l y S p h y r a e n i d a e . ........................................... 29
K e y t o S p e c i e s o f S p h y r a e n i d a e f o r s p e c i m e n s o f .5 5 t o 1. 1 9
MM FROM THE WESTERN NORTH A T L A N T IC ............................................... 31
K e y t o S p e c i e s o f S p h y r a e n i d a e f o r s p e c i m e n s o f 1 2. 3 t o 1 3 0
MM FROM THE WESTERN NORTH A T L A N T I C ................................. .. 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 im 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
Po stlarval D e v e l o p m e nt of Sphyraena b a r r a c u d a ........................... 35
D i s c u s s i o n .................................. ...................................................................... 42
D escr ip tion and S i ze of the A d u l t s .......................................................... 42
R e p r o d u c t i o n ................................................................................................................ 43
Sexua l d im orph ism ........................................................................................ 45
C r it er ia for S tages o f Sexua l D e v e l o p m e n t ......................................... 47
A g e a n d S i ze a t M a t u r i t y ............................................................................. . 47
Sex R a t i o .......................................................................................................... . 49
S paw n i ng P e r i o d ............................................................................................ 49
Frequency an d E x te n t o f S p a w n in g ....................................................... . 51
F e c u n d i t y ............................................ ............................................................... 53
E g g s ..................................................................................................................... ' 53
D i s t r ibu ti on of the P os tl arvae in R e la ti on to H ydrograp h i c and
M e teoro log ica l C o n d i t i o 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 nd Gro w t h ........................................................................................................ 59
Cause of Annulus F o r m a ti o n ............................................ .. ................................ 59
M e t hods of Sca le Co llection and A n a l y s i s ............. . .. ................................... 59
Va lid ity of the Annu lus as a Y e a r - M a r k ..................... . • • • • ................... 61
Criteria for the In terpre ta tion of Annua l M a r k s ....................................... 61
False A n n u l i ................................................................................................................ 64
“Sk i pped” A n n u l i ..................................................................................................... 64
Grow t h of t he Y o u n g ................................................................................................ 65
Empir i ca l Grow th R a te in A d u l t s .................................................................... . 66
Ca l cula ted Grow th R a t e s ..................................... . . . . ......................................... 69
Body-sca le Rela tionsh ip ........................................................................................ 69
C a l c u l a ti o n s ................................................................................................. . * . . . . . 72
Cal cul at ed Growt h I n c r e m e n t s .......................................................................... 74
Leng t h-frequency Di st ri b uti o n s .......................................................................... 74
M o r t a l it y a nd S ur v i v a l ........................................................................................ 80
L a n d in g s of 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
Po ll ution ............................................ . ......................................................................... 92
Salinity ......................... .. ............................................................................................... 92
Dep t h .............................................................................................................................. 93
Tempera t ure ................................................................................................... .. 93
Eco l ogy of Young B a r r a c u d a ............................................................................. 94
Eco l ogy of A d u lt B a r r a c u d a .................. . .. .............................................. .. 95
Assoc ia tes and P r e d a to r s ................................. ...................................................... 96
B e h a v io r ................... ....................................................................................................... 99
Aggrega ting B e h a v i o r ............................................................................................. 99
Peck O r d e r ................................................. .. ............................................................. 1 01
F e e d i n g H a b i t s .............................................................................................................. 102
F ood H a b it s ................................................................................................................... 103

M e thods ......................... .. ........................................... ................................................. 103

Num ber of It em s of Food per S t o m a c h ...................................... ... ................ 104

vi

«
 F o o d o f Y ou n g from F l o r i d a .................................................................... 104
F o o d o f Y ou n g from B im in i, B a h a m a s ................................................... 106
F o o d of Y ou n g from A d d iti o n a l A r e a s ...................... ............ .. .............. 109
F o o d of A d u lt s from F l o r i d a .................................................................... 110
F o o d o f A d u lt s from B im in i, B a h a m a s ................................................... 112
C om par i son s a n d D iscuss ion .................................................................... 113
A ttacks by B arracuda on Hum a ns ............................................................... 121
L itera tu re R e v ie w of A t t a c k s ....................................................................... 122
P oss ib le C auses of A t t a c k s ....................................................................... 126
P o iso n in g of H um a n s by B a r r a c u d a ............................................................ 128
G enera l R e v ie w o f P o i son ou s F i s h e s ....................................................... . 128
C harac t er is ti cs o f P o i sonous B a r r a c u d a ................................................... 128
Tox i n from Barracuda F le s h ....................................................................... 133
E ti o lo g y of F ish P o is o n in g ......................................................................... 133
R el ati on t o S p a w ni n g ................................................................................... 134
R e la tio n to D i e t ........................................................................................... . 135
T ox i s O rgan ism s in the F o o d W eb of the B a r r a c u d a ........................... 135
F ood H a b it s and E c o l o g ic a l R e l a t i o n s of O r g a n is m s in

the F ood W eb of the B a r r a c u d a .............................................................. . 138

G eograph i ca l D i s tr ibu tion of T ox ic B a r r a c u d a .................................... . 139
R e la tio n of T ox i c O rgan ism s to H ydrograph i c and M e teoro lo g ica l
C o n d i t i o n s ................................................................................................... . 140
C o n c l u sio n s ..................... . .............................................................. ........................ .. .. 152
L it e r a t u r e C i t e d ........................................................................................................ 153
 SYSTEMATICS AND L IF E H ISTO R Y OF
T H E G R E A T BARRACUDA,
S P H Y R A E N A B A R R A C U D A (W A L B A U M ) 1
D O N A LD P. d e SYLVA
Ins titu te of Mar ine Science
Uni versit y of Mi ami

A b st r ac t

The great barracuda, Sphyraena barracuda (Wa lbaum ), family Sphy-

raenidae, is found in all tropical seas, with the excep tion o f the eastern
Pacific. The following nomina l specie s are considered as synonym s o f
barracuda Walbaum, 1792: p icuda Bloch and Schneider , 1801 ; becuna
Lacepede , 1803; barracuda Shaw, 1804; barracuda Cuvier, 1829; com-
merson i Cuvier, 1829; dussumieri Cuvier, 1829; agam Riippell, 1835;
affinis Riippell, 1835 ; nuageuse Lienard , 1843; p icuda Poey , 1866; snod-
grassi Jenkins, 1901; and akers trom i Whitley, 1947. S. kadenar Montrou-
zier, 1857, and m icrops Marshall, 1953, are probably synonyms.
Barracuda were collec ted off Miami, Florida , from Bimini, Bahamas,
and specimen s from the Danish Oceanographical Expeditions in the west -
t t . -
ern A lans ic were used Ontogenetic variation f in proportional
t measure
ments wa determined
s for 171 specimens rom 5.5 o 1111 , mm standard ,
length. Key , for , identification are . presented for barracuda guachancho
picudilla borealis and sphyraena t ll
Mosts males mature t a two years andll a mature t by three years. Some
female ismature a three years, and a mature a four. The spawning
season
t from tApril until October off southern Florida. . Spawning was
no
i observed bu probably occurs in the open ocean The eggs were not
dentified from plankton tows. Postlarvae f t were taken. far from shore and
near the surface, over depths o 200 o 5200 meters f
Age analyses were based upon the scale readings o 555 barracuda, s
back-calculated
t -f lengths from scale f measurements,s and upon mode in the
leng
It h requency
s distributions
s t o 2707 f specimen
fi st from southern Florida.s
t attain 10 ftoits12s inche a the end o the rf winter, t and 18 to 22 inche
a the end o fecond.
t It reaches
i t anf age o a least 14 years. Maximum
length is about 6 ee and a we gh o more s than , 100 pounds.its fi st i
The young barracuda s, drifts inshore s in pring spends t r summer n i
shallow nursery areas and move offshore it sto somewha deeper water t n
late fall. During theI econd i summer enter s the mangrove habita or the
deeper weed beds. sn ts third year it enter the coral-reef habitat. The
barracuda migrates easonally t along
. both coasts o f Florida. Some indi-
viduals may drift far s ou to i seas t
Young sometime occur n loo e aggregations, bu the adults are solitary. s
Peck order was observed among . individuals from 12 to 16 inche long
which were confined in a tank
l t t . f , f ,
Con
f ribu ion N o 220 rom The Marine Laboratory Institute ll t os Marine . , Science University
o Miami, Papers from the D A N A , Oceanographical Co ec ion f Nt o 47
l and
t . Contribution
li ti
from
f the
s Lerner Marine Laboratory The Americanti l Museum
i o Na urati , His ory Pub , ca on
of llthi paper through . as grant from the Na onai Sc ence Founda on tG-24119 is grate- f
u y acknowledged l f Thi paper represents
t s a d ssertation
f submitted to he Faculty f ot thef
Graduate
i s Schoo
. o Cornell Universi y a partial ulfillment for the degree o D oc or o
Ph lo ophy
 Young f eed 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 exist ence
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 experience s with barracuda are
discussed in relation to the factor s which are favorable to an attack. t
Poisoning in humans t who have eaten l fresh barracuda
. I is due to at oxin
t ,
in isthe flesh
s and no from bacteria poisoning n the western s A lan ic
po oning have occurred throughout t f the year. There seem to be no
relation between
s the attainmen
f o maturity
s . or the spawning
t f cycle and
the
f poisonou nature o barracuda fle h iSome 58 per cen to the, stomachs f
o adult barracuda tcontained t fishes wh ch i are reputedly oxic o which-
20 per cent were f te raodon
t , iform fishes. Ev sdence is presented for a food
chain originf o the oxin and l mechanism are discussed for the trans-
missionf o the toxin from p anktonic and benthic algae to barracuda by-
way i o l intermediate
s organisms. Variations in hydrographic and meteor s l
olog ca feature t areit believed
f to result
s in distinct
s temporal
. and patia
variation in the oxic y o barracuda a well a other fishes
I n t r o d u c t io n
s, , -
t i The barracuda t familyll Sphyraenidae
f form t a well-defineds group con
a ning abou 20 species, i t a oi which l t belong
s f t to he genu . Sphyraena. They i
are wide-ranging
s n he trop ca wai er ot , he world Some
s arel found, n-
school and seldom s i exceed 2 feet n leng h while other , :are so). itary no
madic sspecie wh ch may l reach 10 feet (Biittikofer s 1890
s 451
fis s. Most are
used a food, ands the arger tspecies are al o valued f a game
f t he s.
Very little t ha beenj writ en on thes biology o any ( o )he barracuda
Except
( ) for he two ma f t or contribution
t by Gudger
t t 19
is 1 8 and Walford
19 3 2 f the content o he ra her voluminous i sli era ure limited s, essentially
to briet and l often
t . inadequate species tlist ng and
t description and snotes
on na ura ( his ),ory Notwithstanding
it ( ), he recen
s ( taxonomic
), t revision
t f by-
Schultz 1 9 5s3 sSm h 1t9 5 6 and William 1 9 5 9 he dear h o muse st
umf t specimen
is . ha preven ed t a comprehensive,l world-wide si f taxonomic udy s
o h group Because of he comparatively arge ze o some specimen
t s f , s
and he ensuing problem o preservation f manys type specimen are no
longer
iff in
t existence. A number
st s f o nominall specie evidently sare based s upon-
d t erenf ontogenetic st ages o a sing e species. Adequate , erie represent
ta ive o ontogenetic
s f f age
t are seldom available and . consequently direc
comparison o dif t s eren species
s can t tseldom be made
s s
Mostt charac er used tto l epara e hes specie of barracuda are based ts
upon he number itof , la era ).-line scale or proportional s measuremen
l
(Schultz, t 1953; Sm h 1956iffi Since lt the tscale i . are t marked tt i yt sdeciduous,
accuras e counts t are, s oftens d cu to ob, a n O hert ,a r bu e used for
specie fi separa t ion uch a eye diameter head leng i h .maxillary lengths,
and n leng hs, show t fconsiderable
t t ontogenetic
st i f var ation s They sometimes s t
vary
iff so t much tha dif eren it is grow
l th t ad s a o a speciesl eem to repre s enst
d eren species.it Thus, i c ear ha t i uch proportiona
f comparison mu
be used w h discretion n the iden if cation o specimens. Color or color
2
patterns are seldom distinctive and the numbers of fin rays or spines are
of little taxonomic value, although Williams (1 959 ) has used color pat-
tern s w ith success for Ind ian Ocean specimens. Although a world-wide
revi sion, whi le is so urgentl y needed i n t his group, must await t he accumula-
tion of addit ona i l specimens, some conclusion s may be drawn concern ng i
the presen t taxonomic sta tu s of some of the species to elucidate a few of
their biological problems.
Of all the sphyraenid s, perhap s the mo st ha s been written about the
grea t barracuda , Sphyraena barracuda (Walbaum ), herea fter re ferred to a s
t barracuda . Found hroughou
he t t most o f the rop t cai l and ub s rop
t ca
i l
wa ter s of the world , it is an impor tan t 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
fle sh o f th is and other sphyraenid s is known to possess poisonous qua litie s
occasionally.
The following sections represent an att empt t o collate t he widely scat-
tered accounts o f the barracuda supp lemented with ori ginal dat a, and to
deduc t there from a synthe sis of the biology of th is intriguing anima l.

A c k no w le dg m e nt s

The pre sen t study is an ou tgrow th of fie ld work star ted a t Pier 5 ,
M iami, Flor ida . This famou s fish ing dock is the cen ter for char terboa t tr ip s
from M iam i to the nearby ree fs and to the Gulf S tream . It o ffer s an exce l-
lent opportunity t o make observat ions on large, pelagic fis hes whi ch are
usually difficult to obtain. During 1952 and 1953, at the suggestion of
Prof. Lui s Rene Riva s of the Departmen t of Zoology , University of M iami,
I made frequen t visit s to Pier 5 to examine ca tches of scombroid fishe s.
As a result of several outbreaks of food poisoning in man due to the
ingestion of fresh barracuda caugh t in Flor ida waters in 1954 and 1955,
t au thor became n
he i tere sted n
i study ng
i the li fe his tory o f the barracuda ,
and in 1955 and 1956 t he field work at Pi er 5 was res umed. I wish t o
acknowledge the complete cooperation of Mr. Dan Daniels, former ly dock-
master of Pier 5, Miami, and the numerous charterboat captains, all of
whom willing ly and patiently supported the fie ld observa tions. To Dr .
Edward C. Raney I am indebted for his guidance throughout the course of
t study . Ialso wish ot hank
he t Dr s. John P . Barlow , Howard E . Evans ,
and Gustav A. Swanson or many help fu l suggestion s and cr iticism s, a s
f
wel l a s the sta ff of the Depertmen t of Con servation , Cornell Un iversity ,
f mak ng
or i numerou s facil tie i s available .
I wish to thank Dr . Charles M. Breder, Jr., Dr. and Mrs. Loui s A.
Krumholz , pre sen tly of the University of Louisville , M iss Francesca R .
LaMonte, and Dr. Albert E . Parr for mak ing available the facil itie s of
t American Mu eum
he s t lHistory and he
of Na ura t Lerner Mar ne i Labora -
t a t Bim ni,
ory ‘
i Bahama s. Dr s. Krumholz and Le ster R . Aronson a ssisted
3
w it h s ei n i ng operati ons. The assist ance and knowl edge of coll ecti ng t ech-
n ic s o f the na tive staff at B i m in i greatly exped it ed the work .
Thank s are due Dr. E . Ber te lsen and the la te Dr. A Vede l Tan ing
t Car lsberg Founda tion , Copenhagen , Denm ark , or
he f mak ng
i ava ilab el an
ex t ensi ve coll ecti on o f po stlarva l sphyraen id s co llec ted by the D A N A e x -
pediti on s under the leader sh ip o f the la te Pro f. Johanne s Schm id t. Dr .
W o lfgang K lau sew itz o f the Senckenberg M u seum a t Frank fur t-am -M a in ,
G ermany , k ind ly re -exam ined severa l o f R iippe ll’s type s and repor ted on
t
hem in de at il.
I w ish to thank Dr . F . G . W a lton Smith and the staff o f the I nstit ute
o f M ar ine Sc ience o f the Un iver sity o f M iam i for prov id ing much o f the
equ ipm en t us ed in t his st udy, and f or off eri ng many us ef ul s uggesti ons t
s t
oward i he m provem t fen t o he smanu i t. cr p. Dr John . E Rll,anda i U-nsitver f
t y oi Puer , s o liR co l uppl ed i f va uab ti e n orm s a onl on evera barracuda
a ttack s and ca se s o f po ison ing .
In add ition to the per son s o f the above -m en tioned i n stitu tion s, the
of llow ng i nd i vi dua
i ls oaned
l s m
pec i en s or
f his
t study :Dr .Jame sE .Boh ke l ,
A cadem y o f Na tura l Sc ience s o f Ph ilade lph ia ; Dr . John C. Br igg s, U n iver -
sity o f Texa s; Dr . D av id K . Ca ldwe ll, L o s A nge le s C oun ty M u seum ; Dr .
Bruce B . C o lle tt e , U . S. Fis h and W il dlif e Servi ce; Mr. D ona l d S. Erdman,
D epar t m en t of Agri cult ure and C om m erce, San Juan, Puert o Ri co; Dr .t
Rober i W. t Harr
, ngl oni Jr., t Ft or da S a fe Board lt ; o H. ea h Mr. Frank J
Ma ther , III, W ood s H o l e Oceanographi c I nstit uti on; Dr. Ernest A. Lach-
ner , U . S. N a ti ona l M us eum; Dr. Gil es W. M ead, Harvard Uni versit y;
Pro f. Lu is R ene R iva s, Un iver sity o f M i ami; Dr. Leonard P. Schult z, U. S.
N a tiona l M u seum ; Dr. C. L avett Smit h, Am eri can M us eum of Nat ural
H istory ; and Dr . R oya l D . Su ttku s, Tu lane Un iver sity .
F or va luab le in form a tion on the inc idence and tox ico logy o f barracuda
po ison ing , thank s are ex tended to Dr . Bruce W . H a lstead , Wor ld L ife
Re search In stitu te , C o lton , Ca liforn ia ; Dr . Y o sh iro H a sh im o to , F isher ie s
Depart m ent, U ni versit y of T okyo; Dr. M as ao Mi git a, Tokai Fis heri es i
Exper m ent St ation, Tokyo ; and Mr. Si dney Paet ro, Broward Count y
F( ol r di a ) H ea lt h Depar m t en t. I w is h ot hankt t numerou s v ci ti
he m so f
barracuda po ison ing who d isc lo sed ex ten sive in forma tion on the na ture o f
t po ison ng
he i and he t l
oca litie s where he t fish were caugh t. Thank s are
a lso due to Dr . L ou is F . W e stbrook and h is sta ff o f the D ade Coun ty
(F lo r id a ) H ea lth Depart m ent f or bact eri ol ogi cal anal ys es of s ampl es of
barracuda fle sh .
Spec ia l apprec ia tion is expre ss ed t o M ess rs. Jos eph T. R ees e of Fort
Lauderda le and the la te A1 Pfl ueger of Mi ami f or s uppl yi ng s cal e s ampl es
from argel barracuda and or f he t u se o f a arge l number o f mea suremen ts
o f barracuda, res pecti vel y. I wis h t o express my grat itude t o Capt ain
E dd i e M oore and his crew of t he M / V P A N D A f or t heir ent husi ast i c
s
uppor t ni co llec ting adu lt pec s m i en s o f barracuda in he t Bahama s.
The fo llow ing pers on s have a ided ma ter ia lly in co llec ting or a ssisting
4
the writer in collecting specimens: Dr. and Mrs. Ilias Konstantinu; Mr. and
Mr s. William H . Peterson , Jr .; Mr . and Mr s. Neil Roseman; M iss Ka th -
leen Parker ; Dr . Frederick A. Kalber; Dr . Allan B . Lewis ; Messrs. Selwyn
J. Bein, Robert F. Burrows, Edward F. Klima, Reynolds A. Moody, Alan
Mo ffe tt, Cra ig Phil lips, Warren F . Rathjen , Jame s D. Regan, Durb in C .
Tabb, and Alfred V. Volpe. I wish to t hank Dr. Frederick A. Kal ber f or
preparing t he 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
t illu stra tion s. Dr s. Gera dl P . Cooper , University o f Michigan , Louis A.
he
Krumholz , Erne st A. Lachner , C. Richard Robins, Gilbert L. Vo ss, and
F. G. Walton Sm ith o ffered valuable criticisms of the manu scr ip t.
Finally and especially , I wish to thank my wife , Doris, for her patient
and valuable assistance dur ing the many hour s of fie ld and labora tory work .
Sy st em a t ic s
P h y lo g en e tic R e la tio n sh ip s of the F am ily S p h yraen idae .— Specie s o f the
family Sphyraenidae are charac ter ized by the ir long , pointed heads^, the ir
pike-like appearance , and the ir two dor sa l fin s. The jaw s contain well -
developed shear ing teeth , and in Atlantic specie s there are no g ill raker s.
The phylogenetic re lation sh ip s of the family have been discu ssed in de ta il
by Br idge (1 8 9 6 ), S tark s (1 8 9 9 ), Dollo (1 9 0 9 ), Goodrich (1 9 0 9 ),
Regan (1 9 1 2 ), Ribeiro (1 9 1 5 ), Jordan and Hubb s (1 9 1 9 ), Frost (1 9 2 9 ),
Gregory (1 9 3 3 ), Hollister (1 9 3 7 ), and Gosline (1 9 6 2 ). 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 w ith the c leithra 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 repre sen ts a compac t family having anato -
i l st t s t t iff tf its st l s.
m ca ruc ure no grea ly d eren rom neare re ative
il i t i ti t il i . s s
F oss D s r b u o n of he F am y S ph yraen dae — Fossil phyraenid are
s t l t
widespread ince he Lower Eocene and inc ude he genera S ph yraen odu s
( i t ), t ,
Agassiz = D c y o d u s Owen P ro o sp h yra en a Leidv, P rosph yraen a Pauca
l i . s s f t l tt s i l
and Sphyraena K e n Numerou fossil specie o he a er genu nc ude
t . l i , , f t f ; .
he following: S ugard White 1926 rom he Eocene o Nigeria 5
i , t ti f t t
fayu m en s s Dames, 1883 from he Ter ary o Egyp and he Eocene of
It ), l i s , , f
aly (Bassani, 1899 S. m a em been s s Dartevelle and Ca ier 1943 rom
t f t (s il
he Lower Miocene o he Belgian Congo ee also Henriques da S va
[ ] f l t f f i i
1956 or a closely re a ed orm rom Portugal); S. v anna Dartevelle
s , , f t ; i , ,
and Ca ier 1949 rom he Miocene of Celebes S. w eb er Casier 1954
5
from the Miocene of Java; S. (Otolithus) handsfuchsi Schuber t, 1906 , from
the Miocene of Voslau, Hungary; S. pannonica Wei ler, 1938, f rom t he
Middl e Oligocene of Hungary; S. striata Cas ier, 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 t he London Clay; S. tyrolensis Zitt el, 1890, from Aus tri a; 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 i n 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 Eng land ( 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 .
Pic tet (1 853 ) inc luded 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 na tura lly difficult to evaluate in the light of such fragmentary
knowledge. However, the distribution of fossil genera and specie s indicates
t t he
ha t amily
f Sphyraenidae wa swide spread throughou t he t warm , shallow
wa er of he Ter ary Tethy Sea (Ekman, 1953: 63 67 Agassiz ’s (1843 )
t s t ti s - ).
recognition o f Sphyraena barracuda from Cretaceou s beds would suggest
t th
ha t is specie s is a re atively
l stab el one morphologically .

Recent Distribution.— Sphyraena barracuda is presently distributed in all

trop ical, Recen t sea s with the exception of the ea stern Pacific Ocean
(Fig. 1 ), the grea t expanse o f wa ter between the Hawaiian Island s and the
western coasts o f the Americas acting as an effective barrier to disper sa l
(Ekman, 1953 :72 -77 ) for many trop ica l fishe s. Although the Equa toria l
Counter Current could conceivably supply a stock of the pelagic young,
evidently th is expanse plu s the encroachmen t of polar curren ts toward
t equator iseffective ni preven ing
he t any extensive colonization by we stern
Pacific species. The data plotted in Figure 1 include barracuda and all t he
nominal species which I consider to be synonyms of barracuda.
The barracuda may occur many miles from shoal water, particularly
abou t the oceanic island s o f the Pacific . Murphy (1914 ) repor ted th is
species about 900 miles east of Bermuda in August. Subsequently, Om-
manney ( 1 9 5 3 ), Murphy and Shomura (1953a , 1953b ), Mather and Day
(1 9 5 4 ), Shomura and Murphy (1 9 5 5 ), Spr inger and Bulli s (1 9 5 6 ), 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 dis tribution o f the barra-
cuda appears spotty due to lack o f collecting in many areas, i.e., the
South Atlantic, Indian Ocean , and the northern Au stralian coast. Its known
distribution is within the range given by Ekman (1 9 5 3 :3 ) for the mean
location of the 20 °C isotherm, wh ich limits the poleward distribution o f
tropica l species. However, some barracuda evidently to lera te somewhat
lower tempera ture s, since Cape Cod , Massachusetts , from which speci -
men s have been taken in August, ha s a mean tempera ture of about 18 .3 °C
for th is mon th (Fuglister, 1947 ). A lso , Durban , Sou th Africa, and Kyushu ,
Japan , evidently represen t the max imum poleward d ispersal o f barracuda
and have sligh tly colder tempera ture s than genera lly pre ferred .
D e lin ea tion o f t he G en u s Sphyraena.— The living members o f the family
Sphyraenidae are represented by a single genus, Sphyraena Klein, 1778.
Fowler ( 1903 ) proposed the subgenus A g r i o p o sp h yra en a for the large-
s
caled form , with 75 ot 87 scales , having a fla ttened or sligh tly concave
head, with barracuda Walbaum, 1792, as the type, as opposed to the sub-
genus S phyraena with more than 100 scales and a slightly convex head.
Smith ( 1 956 ) 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 generi c dist inctions are, however, based upon what would
appear at best to be s peci fic charact ers, s uch as t he number of gill rakers
( “one or two,” as compared wit h “one” ), length and placement of fi ns,
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 dist inctions. All these charact ers overl ap when
considered on a world-wide basis , although they suffice for western Indian
Ocean sphyraenid s. Whitley ’s (1 947 ) genu s A u s tra lu zza , although pro -
posed for a different species ( Sphyraena n ovaeho lland iae Gunther), also
seem s to be only a case o f spec ific d ifferen tia tion ; it is based upon the
position o f the dorsal and pelvic fins, and upon the number o f lateral-line
s
cales. Until a complete revision of he t family isundertaken , Ipre er f to
re ain Sphyraena a he sole gener c repre enta ive o he living specie s
t s t i s t f t
of the family Sphyraenidae.

S y n o n y m y o f Sphyraena barracuda
Sphyraena barracuda was originally described as E so x barracuda in
1792 by Walbaum from Catesby’s ( 1 743 ) pre-Linnaean “Umbla minor
marina maxillis longioribus” from the West Indies. Riippell ( 1835 ) showed
that Forskal ( 1775 ) had mistaken S. agam Riippell for E so x sphyraena
Linnaeus. Walbaum’s description was apparently overlooked for a number
7
 of years, and the We st Indian form was again described, this time by Bloch
and Schneider in 1801 a s Sphyraena sphyraena var . picuda, based upon
Parra ’s (1787 ) non-binomia l “Picuda” from Havana. Lacepede (1803 )
described S . becuna from a draw ing by Plum ier a t Mar tinique , wh ich Cu -
vier (in Cuv ier and Valenciennes, 1829 ) subsequently believed to be
identica l w ith Parra ’s “Picuda,” bu t he had evidently overlooked both
Walbaum ’s and Bloch and Schneider ’s description s. Shaw (1804 ) des -
cr ibed Esox barracuda a fter Catesby’s (1 743 ) specimen , and Cuv ier (in
Cuv ier and Valenciennes, 1829 ) later described S. barracuda based on
Shaw ’s Esox barracuda, bu t o ffered no comment; since Cuvier’s type
locality wa s Brazil he may have believed tha t because Shaw ’s specimen
wa s no t from Brazil it wa s necessarily a d ifferen t species. Swa in (1882 )
iden tif ied Shaw ’s barracuda w ith Bloch and Schneider ’s picuda. Walbaum ’s
or igina l description wen t unnoticed un til Jordan and Ru tter (1897 ) point-
ed out that barracuda Walbaum had pr iority over picuda Bloch and
Schneider , 1801. It had been recognized by mo st authors that becuna
Lacepede, 1803 and barracuda Cuvier, 1829 , were synonyms of picuda
Bloch and Schneider, 1801 , although this was doubted by Ppey ( 1 8 5 8 ),
and thu s picuda ha s been in wide use since, notwith standing Jordan and
Rutter’s ( 1897 ) statement.
Cuvier’s (in Cuvier and Valenciennes, 1829 ) description of S. commer-
soni was based upon Lacepede’s ( 1803) plat e of t he “vari et e de l a Sphy-
rene Chinoise,” and has since been loosely applied to all large, I ndo-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 (1 8 7 5 -1878 ). In the original description, Cuvier stated
that Dussumier himself believed that the species named after him was
iden tica l with a common specie s in the Red Sea , wh ich is evidently S. bar-
racuda . Bleeker (1 865 ) iden tif ied a s S. commerson i a specimen from the
Antilles which was labeled “S. p icuda” in the Leiden Museum. He com-
pared it critically with specimens from the Netherlands Indies but could
not find “ the sligh te st difference,” and, as Weber and de Beaufort (1921 )
pointed out, it is surpr ising that Bleeker did no t place commerson i in the
synonymy o f h is “S. p icuda .” Since Bleeker (1854b ) had previously sta ted
t t commerson i and dussumieri m gh
ha i t be iden ical,
t and since dussumieri
was later shown to be the same as picuda by Weber and de Beaufort
( 1 9 2 2 ), it seems strange that this point has been overlooked.
The identity of commerson i with barracuda had been proposed by
Fowler ( 1935b ) and Herre (1931 , et seq .) . Fowler had earlier ( 1928 )
s
uspected t t S . commerson i, Kner (1 8 7 9 ), a s wel l a s S . commersoni,
ha
Jenk n i s ( 1901 ) were actually barracuda Walbaum. Bleeker ( 1854b )
fur ther compared commerson i w ith barracuda; the la tter is suppo sedly
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 t he height of t he body 7 or more times i n t he
length,” all of which mi ght be att ri but ed to ontogenetic changes. All
s
pecimen s o f barracuda I have examined from the tropi cal west ern Atlantic
have fine teeth on the vomer.
Schultz ( 1953 ) tentatively placed co m m erso n i in synonymy with S.
ch inens is Lacepede, 1803 , but co m m erso n i , as understood from Cuvier’s
( 1829 ) description o f 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 com m erson i to be a probable
synonym of f iav icauda Rtippell, 1835 , yet in Smith’s description of
f iav icau da 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
t is . i f it
bu rounded Hence, co m m erso n cannot be a synonym o e her
i i i i . ( )
ch nens s or f av cauda Possibly Weber and de Beaufort 1922 had
t is i i t
recognized h since they included co m m erso n n he synonymy of
. i . .
“S p c u d a ” { —S b a rra c u d a )
Cuv er { n Cuv er and Valenciennes, 1829) also stated that in co m m er-
i i i
i t s i s , s i it ends
son he opercle end n two spinou points wherea n barracuda
i one. Ihave found th is charac ter to be var ab
n i el n
i S. barracuda from the
t ica l we stern Atlantic, being sub jec t to ontogenetic a s wel l a s indiv idua l
rop
variation, and care must be exercised not to overlook the lower point
i fl s fl .
wh ch may be concealed by a e hy ap
it ( ) t t t i ti s t fi s t isti
Sm h 1956 observed ha he wh te p of he n are charac er c
. is ( ), i i, t
of barracuda Th led Bleeker 1 854b n speaking of com m erson o
st t [t s t ]: its l, s l fi it i
a e ran la ion “In violet caudal, ana and dor a n w h wh te
ti s it s t i f i t l is
p make me h nk o Sphyraena agam Riippell, and n ac ua ity a
l t i , is its t i t
re a edl fispecies,
, i wh ch
s however, easily recognized by hr
t is sce cut-ou it
cauda n wh ch ha a quadrilobed appearance.” Compare h al o w h
s ( ) t l fi f i s
Riippell’
s 1835 description i of he cauda n o i agamit wh ch . . .ha “an
dieitemtiRande
s bildem dre halbrunde Ausschnitte t isti v er Sp zen f .” The t
wh ste I p mentioned
s, by Bleeker
, are charac er c of
), t barracuda rom he
(
We ndie ), Africa
t (Smith 1956; Isl Williams,
s 1956 ),he ltRed Seat isUSNM-
147455
t is and
j t he Hawaiian
i t and (Jenkins, t 1901
si f ta hough
s i h . char
ac er sub ec t il isto var a ion depending
s i l upon he ze o t is pect men
he t The
quadrilobed( a , only ,obviou . n arge
, I. I; specimens, bu dis
I. II, ive of
inc
barracuda , s Smith l s j 1956 ll . Fig 4b P Williams, 1956, P lower
figure) a wel ’s ( a e o
) i t i s , f t ,
Klunzinger
st t 1884iff specimen
tf den if ed al agam rom( he Red ). Sea
wa hough to be ( d )eren rom t tagam Riippel t i by Schultz i 1 9 5 3 ’s Jordan
( )
sand Evermann s 1905 doub l f ed he iden if cation of
t i Ste ndachner
t i t 1901 s s
fpecimen
t i a agam. Riippel (rom Honolulu, ) t bu fa led f o g ve he rea on
or he r decision Schultz 1953 separa ed agam rom barracuda essen-
9
tially on the lateral-line pore count, barracuda having 76 to 85, while
agam has “more than 100” pores. Smith ( 1 9 5 6 ), apparently on Klun-
zinger’s ( 1870 ) statement that agam has 85 scales, as well as Weber and
de Beaufort’s ( 1 922 ) inclusion of agam in the synonymy of “S. p i cu d a ”
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
s ing
ca l is conventiona l and doe s no t repre en
s t het rue
t sca el count. Dr .
Wolfgang Klausewitz has kindly examined t he t ype of Ruppell’s agam
in the Senckenberg Museum and in form s me tha t it ha s 81 la tera l-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 aff in is 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 ( 1 891 ) 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 t o as cert ai n if t he enti re
description is cited. Lienard’s description o f the size o f nuageuse, to 5
feet, and the coloration, wit h blacki sh spots about t he l at eral line, t he
dusky vertical fi ns, t he black opercul ar membrane and base of t he
pectoral fin all are charct eristi c of barracuda, and, i n combination wit h
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 vari et y of Esox sphyraena Linnaeus, and t hat it was not
permissible to give proper names to varieties. Poey felt that his previous
recognition o f the species ( 1858 ) and comparison of it with S. becuna
Lacepede entitled him to describe t he species as new. He als o mentioned
t t he wa s thu s ju stified since Bloqh and Schneider added nothing to
ha
Parra ’s (1 787 ) figure. Whether or not Poey is correct i n maki ng thes e
assump tion s is irrelevant, since picuda Bloch and Schneider is ant edat ed
by Walbaum’s (1792 ) description.
Jenkins’s (1901 ) original description of S. snodgrassi noted that “the
tips of the second dor sa l and cauda l fin s [are ] white” and tha t th is specie s
is “ . . . most closely rel at ed t o Sphyraena commersonii . . . ” but diff ers i
.n. being
. sl
“ , i ender n shaving at lomewha tonger
less l fi pec
, ora n and
i having the black blotches of the soft dorsal and anal fins.” The black
n
blotches, however, are typical of “commersoni” ( = 5 . barracuda ) as
10
120 60
150 E ISO o 30

.\

/ /
F1iag uu trtEe
F 1l.. Distrribbuuttionn of thhe barrrraaccuuda, S phhyraena barracuuda (Walbauunn)).. Eaach dot rrepprreeseents aa puubbllisshedd rreecorrdd of tthis sppeecies oraa nam
me considderredd aas a synonym
m.
figured in the excellent photograph by Williams ( 1 9 5 6 ). The pectoral fin
l th is an allome tr ci charac ter , and slenderne ss essentially depend s
eng
upon the physiological condition of the animal and thus cannot always
be considered diagnos tic for taxonomic purposes. Subsequen tly , Jenk in s
(1904 ) believed h is type specimen to be iden tica l with com m erson i,
although Jordan and Snyder (1907 ) stated that S. snodgrass i was “prob-
ably not the same as the East Indian S phyraena co m m erso n i ” Jordan
and Richardson (1908 ) described a 6-inch specimen o f co m m erso n i from
Aparri in the Philippines as “agreeing with S. snodgrass i 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.
com m erson i, a s 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 (1 943 ) gave an adequate description o f “sn odgrass i”
from Samoa, which he believed to be possibly identical with barracuda.
Subsequent ly, Schultz ( 1953) tent atively considered t he 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. a k ers t ro m i from a specimen 1 4 4 5 mm in
ot ta l leng ht rom
f Lowendal Island , We stern Au stra ia l . He sta ted tha t h is
s
pecimen , when keyed out, “comes down to he t We st Indian Barracuda
(S . p ic u d a ) in Weber and de Beaufort [1 9 2 2 ] . . . bu t d iffer s very much
ni the size of the eye and its ra tio s to o ther par ts o f the head , the
ventrals are much shorter than the pectorals, and the outline of the
cauda l 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
t eye size or
he f “S . p ci u d a ” (= S . barracu da) a s 6 0. ot 6 4. ni head ; my
own measurements on barracuda from the tropical western Atlantic show
a range o f from 6 .0 to 9 .2 , indicating tha t for these specimen s 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
(
1922
).
The eye of Whitley’s ( 1 9 4 7 ) type of ak ers t ro m i ( 1 2 1 0 mm) was
contained 7 . 8 times in the head ( 4 . 5 per cent o f the standard length), and
does not differ markedly from that found for barracuda of comparable
size (Fig. 2 1 ) . The remaining charact ers on whi ch a k erst ro mi is based
appear to vary wit h t he si ze of t he fis h. Specimens collected i n nearby
localities on subsequent dates and assigned to a k ers t ro m i by Whitley may
represent different species, possibly je llo , according to his description
of two o f these. He stated that a k ers tro m 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 o f 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 o f 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 leng th . Subsequen t calculation of the percen tage o f 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 para type wh ch i
was apparently a species differen t from the type . Th is seemingly cau sed
Smit h ( 1956) t o place microps i n t he synonymy of barracuda, s ince t he
scale count given by Marsha ll (1953 ) is for the second fish only.
However, there is no reason to believe that the scale coun t for this second
specimen is the same as for the type. Marsha ll’s latera l-line sca le count
of 116 scale s on the para type is far too high for barracuda or its previously
discu ssed synonyms. He concluded that “in the first specimen there was
probably some mutilation for in the second fish the caudal had the middle
rays somewha t produced as in S. je llo ” In such a large specimen it would
be expected tha t the se midd le 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 pre sen ted here for Sphyraena barracuda inc lude s 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.
s l i
: Genu Sphyraena K e n i s).
Sphyraena-Klein,, i1778 464i (ex Artedi,
, based: on Esox sphyraena L nnaeu
i Sphyraena-Rose
s). n Arted Piscium 1793 112 (based on Esox sphyraena
L nnaeu : ( s
i Sphyraena-Bloch
s). and Schneider, 1801 109 b a e d on Esox sphyraena
L nnaeu : ).
A cws-Lacepede, 1803 327 :(ex ,Plumier, M S
Sphyroena-Dumeril,
: - 1806
). 148 342 (different spelling based on Bloch and
Schneider, 1801 109 110 , : l
Sphyrena-Rafinesque
, ). 1810 34 (different spelling based on B o c h and
Schneider 1801
14
 Agrioposphyraena-Fowlev, 1903: 749 ( new subgenus, based on Esox barra-
cuda Walbaum, 1792). Smith, 1956: 41 ( elevat ed to generic status, after
Fowler , 1903 ).
Austra luzza-Whitley, 147 :131 (Orthotype for Sphyraena novaehollandiae
G un ther ).
Sphyraenella-Smith, 1956 :38 (new genus for Sphyraena flav icauda Riippell,
1835 ).
Indosphyraena-Smith, 1956: 39 (new genus for Sphyraena africana Gilchrist
and Thompson, 1909 ).
Ca llosphyraena-Smith, 1956 :42 (new genus for Sphyraena toxeuma Fowler ,
1903 ).
Pre -Linnaean Name s
Allualu or brochet-Vlaming , 1715 ( 6 ) (Indian Sea s). Valentijn, 1724 (70 )
(ba sed on V lam ing ). Renard, 1754 , Table 40, Fig. 202 (cop i ed from V lam ing ).
Umb la minor, etc. (the barracuda)-Catesby, 1743: 1, PI. 1 (Bahamas; size,
description, voracity, poisonou s qua litie s).

Post-Linnaean
: Names t t ).
Esox sphyraena-Forskkl, : 1775 16 (Zanzibar; . ( misiden ; fifica ion s
liti s).
Picuda-Parra, 1787 90; Table 35, Fig 2 Havana ood value, poisonou
qua e : - l
t Esox barracuda-Walbaum,
: t I i s). 1792 93 94 (origina description after Catesby;
ype locality Wes nd e . - , .
; Sphyraena sphyraena l var picuda-Bloch and Schneider, i 1801: 109 110 Fig
1 Table , 29). (origina description; type locality: “ n America australi,” after
Parra 1787 , I. , . l
Variete de la lSphyrene : Chinoise-Lacepede ). 1803:
, P: 8 Fig ( 3 (origina
description;s type ocality t iIndian s). Ocean Cloquet 1827 233 considered the
same a Sphyraena spe L nnaeu , , I. .
Sphyraena becuna-Lacepede t 1803:
t 325
). P 9, , Fig : 3 (original , I. description
,
after
( drawing by; Plumier a Mar inique Shaw 1804 112 P ).109 bottom i
“Becuna pike” description i from Lacepede - ( t after s, Plumier, f M S Cuvier n
Cuvier and it tValenc ennes, 1829: s 340 341 f An ille [ coast ]; o Brazil;s description;
s -
tibelieved
s). o i be the same a the Picuda o Parra 1787 ( size,, poi f ionou ). quali
e , Cuvier (n Cuvier and Valenciennes, 1831: 507 Gorea A r ca Castel- ,l
nau 1855: 6 ti “Sphyraena ). , becuna Schneider” (s snon. Lacepede;
i . Rio, de Janeiro :
Brazil;
(s co ora. on Poey i1858: ). 398 ame a S p cuda) Poey 1868 275
imilar to S vulgaris Cuv , er: - (
[ Esox]; barracuda-Shaw 1804 105 106 original description based, on Catesby
1743 type locality: Carolinas; s tivoracity,
s). alleged danger to( bathers , distribution-
in West Indies, poisonou; quali s te ).Castelnau, , 1855: : 6 Bahia ( Brazil;
, com).
moni name: “Solteira”
ls , :de crip ( ion Poey, 1868 277 Havana, Cuba
N chosi and ). Breder 1927 72 Long Island New York; description distribu-
tion, ze i l -
( Sphyraena barracuda Cuvier n Cuvier and Va enciennes, [ t];
1829: 342l 349l-
:
original description based on Esox barracuda , i Shaw ; no commen type oca
ity Brazil;f common names: f “becune ” “p cuda” also recorded from , Puertos
Rico; briesi description
). , o :internal organs . ands; skeleton, , t , poisonou
voracity
qualities, ’sze [ Cope ] 1871 472 (St f Martin’ counts colora ion comparison ’s [ ]
with s iPoey ti ). 1858 description o picuda; disagrees with Gunther 1860
de cr p on ii i : -
Sphyraenal commerson Cuvier n Cuvier and Valenciennes, 1829 352 353
(origina description , based on I. a, drawing . , by Lacepede, t the “Variete). de la
Sphyrene Chinoise ” 1803: P 8 Fig 3 from he Indian Ocean Bleeker,
15
1849a: 66 (Makassar, Ce lebe s). Bleeker, 1849b : 55 -56 (Batavia , Samarang,
Surabaya , Java , Kammel, Madura Islands , Ternate, Sindangole , Halmaheira
Island s). 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 . com m er-
son i and S . dussumier i may be iden tica l). Gunther , 1860: 338 ( “Javanese Sea;
Mo lucca Sea ; Hindo stan” ; description , synonymy ). Castelnau, 1861: 7 (Sou th
A f rica ). Bleeker, 1865: 265 -266 (identification o f a specimen labelled “S.
p icuda” from the An tille s as Sphyraena com m erson ii) . Castelnau , 1873 : 102
(Knob Island, Nor th Au stra lia ). MacLeay , 1881 : 33 (c ita tion o f Castelnau ’s
[1873 ] re ference ). MacLeay , 1882 : 33 -34 (Knob Island, North Australia,
after Castelnau [1873 ]; description, counts, co lora tion ). Kner, 1879: 44 (V iti
Islands, Ocean ia ). Day , 1875 -1878 : 342 -343 ( “Seas o f India to the Malay
Archipelago and beyond” ; counts; identity o f Bleeker ’s specimen s o f S . com -
merson i with S . dussumier i in the British Museum ; synonymy , description,
coloration, food va lue ). Day , 1889: 335- 337 (Indian Seas; description, counts,
coloration, hab ita t). Sauvage, 1891 :522 (Madagascar; common nam e ). Saville-
Kent, 1893: 293 (Thursday Island, Queensland, Australia; range, size ).
Fowler , 1900: 501 , 520 (“Sandwich Islands” ; Tah iti). Snyder, 1904 : 523
(Hawaiian Islands). Jordan and Richardson, 1908: 245 (a 6-inch specimen “from
Aparr i [Philippine s] agreeing with S. snodgrassi, from Honolulu , Hawa ii, in
every aspec t except tha t there is a slight tentacle a t the tip o f the ch in” ; counts,
measurements, ambiguity o f Cuvier ’s original de scr ip tion ). Gilchrist and
Thompson, 1909 : 255 -256 (Durban , South A f r ica ). Snyder, 1912: 495
(Ok inawa ). Weber, 1913: 150 -152 (Makassar fish market and Kangeang
Island, Du tch East I nd ie s). Maxwell, 1921: 203 ( M al aya). Fowler, 1922: 82
( H aw a ii). Barnard, 1925: 312- 316 (South Africa; description, synonymy,
distribution , common names; S. snodgrass i a synonym ). Whitley , 1927 : 4 (Fiji).
Shmidt, 1930 : 2 (R iukyu Island s). Pinter, 1930: 500 (Cey lon ; cestode
para site ). Fowler , 1931 : 325 (S . commerson i, Schmeltz [ = Kner ], 1879 , no t
o f Cuv ier ). Ommanney , 1953: 66-67 (Amirante Bank, Seychelles; breeding
season, food , common nam e ). Williams, 1956: 16-18 , PI. II, bottom (East
Africa ; synonymy, common names, distribution in East A frica , description,
color, size, seasona l occurrence, ecology , spawning, food , potentia l fishery,
migration s).
Sphyraena Dussum ieri-Quvier in Cuvier and Valenc iennes, 1831 : 508 -510 ,
Fig. 67 (original description; type locality: Maldive Islands, Indian Ocean, and
eastern coast o f Africa; food value; Dussumier believed it to be identical with
a common species o f the Red Sea ). Gunther , 1860: 339 (lie de France , Indian
Ocean; de scrip tion ). Sauvage, 1891 : 522 (Madagascar , common nam e ).
Sphyraena agam-Riippell, 1835: 99-100, PI. 25, Fig. 2 ( original description;
type locality : Red Sea; f ood value, size, Arabic name: “agam”; S. sphyraena,
Forska l 1775 , a synonym ). Gunther, 1860: 341 (Red Sea ; synonymy). Playfair ,
1866 : 78 (Zanzibar and Red Sea, based on Esox sphyraena , Forskal, 1775, no t
o f L innaeu s). Klunzinger, 1870: 822 (Red Sea ). Klunzinger, 1884: 128-129
(Red Sea; dangerous to m an ). Schultz, 1953: 281 (S. agam Klunzinger, 1884 ,
not o f Riippell, 1835 ). Sauvage, 1891: 410 -414 (Madagascar; S. nuageuse
Lienard, 1843, a synonym; common name ). Steindachner, 1901: 500 ( H ono -
lu lu ). Jordan and Evermann, 1905: 143 (S. agam , Steindachner, 1901, no t o f
Riippell, 1835 ).
Sphyraena affin is-Riippell, 1835: 98-99 (original description; type locality:
Red Sea ). Klunzinger, 1870: 822 (S. affin is Riippell a synonym o f S. agam
Riippell, 1835 ).
Sphyraena pi cuda-R ichardson , 1836: 5, 32, 34 ( b a s e d on B l o c h and
Schneider, 1801; “Gulf o f Mexico, sea o f Brazil, and at Gorea, on the coast
16
o f A fr ica” ). Poey , 1858: 164-166; 255, 398 (Havana, Cuba; description,
synonymy; differs from S. becuna Lacepede; Echeneis sphyraenarum attached
to gills o f barracuda ). Gunther , 1860 : 164 (H avana ). Poey , 1864a : 179
(his torica l accoun t o f nomenclature o f S . p icuda) . Goode , 1876 : 62 (Bermuda ;
breeding plentifully; believed DeKay’s borealis to be “closely allied to, if not
identical with, Sphyraena pi cuda”; food value, poisonous qualiti es). Goode,
1877: 292 (Bermuda; common name: “sennet” ). Goode and Bean, 1879a:
116, 146, 342 ( “South Florida,” “West Florida,” Cuba , Bermuda s). Goode
and Bean, 1879b: 146-147 ( “Wood’s H o le ,” Pensacola, and Cuba; value o f
lateral-line scales in identification; scale counts , range ). Goode and Bean,
1882 : 239 ( G u lf o f M exi co). Jordan and Gilbert, 1882: 412 ( “West Indies,
north to Florida” ; description, si ze). Swain, 1882: 307 (ident ification of Esox
barracuda Shaw, 1804, with Sphyraena p icuda Bloch and Schneider, 1801 ).
Jordan and Gilbert, 1883: 589 ( “rare at Charleston Harbor, South Carolina” ;
scale count; 18-inch specimen from 10 fa thom s). Meek and Newland , 1883:
67 -70 (Havana , Key West, “Wood ’s Ho le , Massachusetts” ; key to specie s o f
western A tlantic Sphyraenidae, synonymy , range; description o f S . p icuda
based on 40 specimens; size attained, proportiona l mea suremen ts). Goode ,
1884: 488 PI. 178 (f ood value, poisonous qualities, range, common name s).
Jordan, 1884 : 77 -79 (Flor ida Keys; method o f catching , associated spec ie s).
Jordan, 1885: 117 (Key West, Florida; food va lue ). Bean and Dresel, 1885:
167 (Jamaica; common names, coun ts). Jordan, 1886: 572 ( “West Indie s” ).
Bean, 1889: 132 , 145, PI. 2, Fig . 14 (Bee sley ’s Point , New Jersey ; occurrence
o f young , coun ts). Kendall, 1889: 308 (Garden Key, Tortugas , F lor ida ).
Jordan and Bollman , 1889 : 550 (Green Turtle Cay, Bahama s). Bean, 1890:
194, 205 (Cozumel, Yucatan; food value, ecolog ica l associates , food o f bar-
racuda ). Henshall, 1895: 213 (food fish at Key West; found in shallower
water than S . guaguanche [ s ic ]). Jordan and Evermann, 1896a: 823 (descrip -
ion,
t synonymy, distribution in A lantic
t , ood
f value, danger to bathers , common
name s). Murphy, 1914 : 2 (o ff Bermuda , latitude 31 ° 50 ' N , longitude 50 °
49 ' W; para site s). Metzelaar, 1919 : 41 (Cura sao ; common name: “snoek” ;
synonymy, range, seasonal occurrence, commercia l importance ). Weber and
de Beaufort, 1921 : 70-71 (Indo-Pacific ; believed that S. commerson i Cuvier
and S. agam Riippell are synonymou s with S. picuda B loch ). Weber and
de Beaufort , 1922 : 224 -225 (Indo-Pacific ; synonymy , description, common
names, size, coloration, range in Atlantic and Pac ific ). Fowler , 1925: 7 (Guam;
description, counts, co lora tion ). Abe, 1939: 533 (Pa lao Islands; size, distribu-
tion ). Blanc and Fourmanoir, 1953: 546 -550 (Oceania: Tahiti, Hikueru,
Apataki, Niau , Takurea, Kaukura ). Mendis, 1954: 31, 102, 103 (C ey lon ).
Morrow , 1954 : 806 (Ea st A frica , between Kenya and Tanganyika ; “several,
up to four fee t” ). Munro, 1955: 90 -91 , PI. 16 , Fig . 246 (Ceylon ; New Guinea;
description, color; range: “coa stal” ; S. commerson i considered a synonym ).
Sphyraena nuageuse-Lizn&vdi, 1843 : 64 (origina l description; type locality:
li e de France ). Sauvage, 1891: 410- 414 (Madagascar, based on Lienard;
habits; probably the same as S. agam R iippe ll).
Lucioperca kadenar-Montrouzier [M S ], 1856 (original de scrip tion ).
Sphyroena kadenar-Thiolliere, 1856: 149 -150 (based on MS o f Montrouzier,
1856; Moiou [ M e l ane sia ]).
Sphyraena kadenar-Montrouzier, 1857: 119 (description; type locality:
Kanala, New Ca ledon ia ). Jouan, 1861: 302 -303 (de scription based on Mon t-
rouzier [1856 ]: poisonou s qua lities).
Sphyraena pi cuda-Poey, 1864a: 179 (synonymy ; identification and renaming
o f Bloch and Schneider ’s p icuda , which they considered to be a variety o f
Esox sphyraena L innaeu s). Poey, 1864b : 187 -188 (synonymy , reclassification,
nomenclatoria l notes ; comparison with S. becuna Lacepede , S . barracuda
17
Cuvier, S. guaguanche [sic ] Cuvier, and S. p icudilla P oey ). Poey , 1868: 275 ,
277, 359 (Havana ; description, synonymy, poisonou s qualities; differs from
becuna Laeepede). Poey, 1875: 95 ( H avana). Poey, 1881: 334 (Puert o 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; s ynonymy). Jordan and Rut ter, 1897: 98 ( barracuda
Walbaum has priority over p icuda Bloch and Schne ider ). Smith, 1898 : 94
(Wood s Ho le, “a rare straggler” ; young from Quisset Harbor ). Jordan and
Evermann, 1900: PI. 128, Fig. 3 4 9 ). Gilbert, 1900: 164 (Maceio, Brazil).
Evermann and Marsh, 1902: 115- 116 ( Puert o Rico: San Juan Market, San
An tonio Bridge, Mayagiiez, Ensenada del Boqueron, Hucares, Fajardo; f ood
va l ue). Fowler, 1903: 749 (San Domingo, Puerto Rico; coloration; proposal
o f subgenus Agri oposphyraena f or this species on basis of the large scales and
the shape o f 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 Thomp son, 1905:
236 (Bush Key, Tortugas, Florida; size ). Bean, 1906: 41 (F latt’s and Cable
Town, Bermuda ). Fowler, 1906: 92 (young from Marquesas and Bdca Chica
Key, Florida; adults from Bahia Honda Key, F lorida ). Kendall, 1908: 68
(Wood s Ho le, Massachu se tts). Vincen t, 1910: 46 (Trinidad; taking trolling,
with seine, and with harpoon). Rosen, 1911: 52 ( Nass au, Bahamas; common
in Bahamas; young from tidepoo ls). Starks, 1913: 41 (Na ta l, Brazil). Sum-
ner, Osborn, and Cole, 1913: 447 (Wood s Ho le , Massachu se tts). Fowler,
1915a: 50 (Santo Dom ingo , West Ind ie s). Fowler, 1915b: 532 (possibly seen
at St. Kitts, West I nd ie s). Ribeiro, 1915: 2 (Braz il). Fowler, 1916: 401 (Toro
Point, Colon, Canal Zone ). Hildebrand, 1916 : 306 (first record for Beaufort,
Nor th Carolina, in “the past two years ”). Ribeiro, 1918: 45 Brazil; s ynonymy,
de scrip tion ). Fowler, 1920: 155 (identifie s young recorded from Bee sley ’s
Point, New Jersey as Sphyraena p icuda Bloch and Schneider by Bean [1889 ]
as S. barracuda) . N icho ls, 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; de scrip tion ). Breder, 1927: 8,
13, 24 (Cuba; f ood includes worm eels, other barracuda). Fowler, 1927: 264
(Philippines: Vigan; Iloko s Sur Province ; Luzon ). Fowler and Bean, 1927:
5 ( Benkoelen Beach, Sumat ra). Ni chols 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: Hono lulu , Hilo, and Waialua; synonymy, dist ribut ion).
Fowler , 1929: 603 ( brief description of head from market in Hong Kong,
Ch ina ). Truitt, Bean, and Fowler, 1929: 62-63 (Worcester County, Maryland;
range, size, f ood value, danger t o bat hers). Ni chols, 1929: 224 (San Juan,
Puerto Rico). Jordan , Evermann, and Clark, 1930: 255 ( range, common names,
s
ynonymy ). Parr , 1930 : 45 Rum
( Cay and Crooked Island , Bahama s). Fowler ,
1931: 325 (Honol ul u, Hawaii; same as S. commersonii, Kner, 1879). Herre,
1931: 5 (Tenibuli , Solomon Island s). ?MacCoy, 1931: 17 (Massachusetts; “a
nine-inch little barracuda Sphyraena barracuda DeKay , from Menemsha Pond ,
November, 1930” ). Fowler, 1932a: 5 (Suva, Fiji; count s, co lora tion ). Fowler,
1932b: 397 (Gasparee Island, Trinidad; co lora tion ). Beebe and Tee-Van ,
1933 : 89 (Bermuda; diagnosis, field ). characters,
, size,: coloration,
- distribution ;,
small individuals “nots uncommon”
f Gregory 1933 261 268 (relationships
description and s ku ll). Fowler, 1934 : 399 (re ference to previous
[ ] figure o the . iji). ,
work 1932a reporting S barracuda from F Beebe and Hollister 1935:
18
213 (Un ion Island, Grenadines , British West Ind ie s). Fowler , 1935a: 276 -277
(Hong Kong, China; size, counts; range: “Red Sea , Madagascar, Malaya, Eas t
Indies, Philippines, Riu Kiu, Polynesia, Hawaii, also tropical America” ).
Fowler, 1935b: 440 (doubts if S. barracuda really occurs i n South Af rica,
since Castelnau’s [1861 ] description could have equally applied to S. langsar
Bleeker, 1854a ). Fowler , 1936 : 578 -579 (N iger River, Africa ; based on speci-
men described by Gunther [1860 : 336 ]; description, coloration, synonymy;
range: “tropical parts o f the A tlan tic” ). Herre, 1936 : 100 (Fiji, Solomon
Islands, Mindanao , Tinian; description, counts , color, habit s; size: “a t least
1800 mm” ; record o f a 106-pound barracuda from M indanao ). 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 , co lora tion ). Delsman ,
1941: 74 (Cay Sal, Bahama s). Fowler, 1941: 225 (Pivers River, Beaufort,
and Cape Lookou t, Nor th Carolina; “common” off Sou thpor t). Longley and
Hildebrand, 1941: 69 -70 , PI. 2, Fig . 2 (Tortugas, Florida; size, food , ecology ,
feeding habits , color, counts, mea suremen ts). Rohl, 1942 : 392 (coa st o f Vene -
zue la ). Fowler , 1944 : 443 , 462 (Panama; Jama ica ). Fowler , 1945 : 128 -129 ;
185, 290, Fig. 75 (Marquesas Keys, Lake Worth, Key West, Card Soilnd,
Boca Raton, Florida ; Magnolia Beach, South Carolina; distribution, re ference s).
Baughman, 1947 : 280 (Po in t Isabel, Texa s). Breder , 1948b : 114 -115 ( “West
Indies straggling to Massachusetts” ; danger to man ; size, line drawing ). Schultz,
1949: 109 -110 (Pun ta Gorda, Venezuela ; size ). Cadenat, 1950: 149 (Senegal
River, A f r ica ). Fowler, 1952: 90 (Port-au-Prince, H a iti). Mather, 1952: 120,
125-128 (Trujillo Bay, Guaimoreto Lagoon, Puerto Castillo, Honduras; Puerto
Barrios, Guatemala; barrier ree f off Cay Bokel and at G lover Reef , Swan
Islands; Carmen, Mexico; sport fishing technics; “one o f the mos t common
reef fish in the northwestern Caribbean” ). Fowler, 1953: 63 (Barranquilla,
Colombia; co lora tion ). Herre, 1953: 235 -236 (Indo-Pacific localities; common
names, synonymy, range, litera ture ). Murphy and Shomura, 1953a: 3, 44
(Kingman Ree f, Palmyra Island , latitude 6 ° 13' N , longitude 163 ° 05 ' W, at
15-40 fa thom s). Murphy and Shomura, 1953b: 29 (near Phoenix Islands , lati-
tude 0 1 ° 18' N , longitude 179 ° 55 ' E ). Schultz, 1953: 279 -280 , 293 , PL 23a
(Bikini, Eniwetok, Guam , Rota ; keys, synonymy , description , coloration o f
living
f and preserved
t I i s). specimens, eco logy ; , should: be compared - ( s with specimensf
rom the Wes nd e , Mather t and Day 1954 f 178 188 off ). hore records o
barracuda from Brazil the ; An illes, and l south o Cape s C odi Reid, 1954:t 31- s).
32 (Cedar Key, : Florida description;; arge specimen rare ; n shallow swa li er
it ).
Kilby, 1955 212 (Bayport, Florida - 48-mm
( specimen temperature, a n y f
Shomura and Isl Murphy,
). 1955: 39: 49 longline
( t catches from the vicinity o f
Christmas). and Erdman, ,1956 : 321, Puer , o Rico; spawning, s occurrence o
young Iversen and Isl Yoshida ). 1956 13 19 26 (longline : catche
, near ,Christmas
Island
( lf and
f Palmyra , and
st ti Springer
s). ,and Bullis,
: 1956
- 69 109, 113 122, 124
G u; o Mexico , s 10 a ts). on Inger 1957 374 375 , (Sandakan,
: North
, Bor-
neo
( counts mea uremen Iversen and Yoshida 1957 21, 22, 24 31, 32,
longline and troll ;catches from Christmas, s). Washington, and : PalmyraI. I islands
and Kingman s Reef two offshore record Williams, , 1959 107,, P D (Easts
Africa; ynonymy, [ ]).description, distribution, ecology taxonomy comparison
with Smith 1 9 5 6 i : - (
Sphyraena snodgrass ; -Jenkins, 1901 387 388 ; original ldescription; s type
locality:
. Honoluluii . type specimen: USNM 49693 “most c o ely related tos
S. commersoni, ”) Fowler, 1903: 749 (Hawaiian ; Islands,
s .Tahiti; same , a
S commerson Fowler i 1900; description, ).color “clo , e to S barracuda ( ap-t
parently
. differing
i n the shorter
, is maxillary” s Jenkins
. 1904:i. 438 believed tha
S snodgrass Jenkins, 1901 the same a S commerson Jordan and Ever-
19
 TABLE 1
M e a sur e m e nt s e x p r e sse d as r a t io s of b o dy par ts a nd m e r is t ic da ta of Sphyraena barracuda a nd sy no ny m s
ASSIGNED TO IT.
barracuda barracuda barracuda barracuda akerstrom i
Western Atlan tic Marshall Islands Indian Ocean Red Sea Western Australia
Schultz (1953 ) Smith ( 1956 ) USNM 147455 Whitley ( 1947 )
Number o f 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
Pelv ic fin length 9. 8-13.2 — 9. 5- 13.2 5.1 13.0-13.1
In head length:
Length o f snout 2 1 2.2 2.2 2 .2-2.1 2. 2 2.0-2.1
. - 2.2
1.7 1.8-2.2 — 2.3 1.8-2.0
Length o f maxillaries -
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
Dep th cauda l 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 pectora l base
o pelv c base 40 45 43 44 43 75 50 —
t i . - . . - . . - . . 3 . 2 - 3 .4
Eye in postorbital 2 .2-3.8 2 3 -3 .1 2.7 -6 .1 # 2.1
Dorsa l elements V +U8 V+U8 V + I ,9 4 V + I ,i, 9 V + i, 9
Anal element I,i,7 -8 U7 1,8 Li,8 i,8
s
Pectoral elements ii,11-12 ii,11-12 i ,l 1-ii, 10 ii-12 i , 13-14
Transverse scales 1 0 /1 / 9 to 13 /1 /14 — 10 /1 /9 — 9 / 1 /1 5 to 12 /1 /14
Lateral-line scales 75-87 76 -85 75 -90 84 76 -83
 ?

TABLE 1— (C O N T IN U E D )
m icrops picuda snodgrass i agam
Queensland “Indo- Pacific,” Hawaii, Red Sea,
Marshall ( 1953 ) Weber and de Beaufor t (1922 ) USNM 46963 Riippell (1835 )3
Number o f specimen s 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
Pecto ja l fin length 11.6 — 8.4 __
Pelvic fin length 13.1 — 8.9 __
In head length:
Length o f snout 2.1 2.2
Length o f maxillaries 2.2 2. 2- 2. 4 2.3 __
nterorbital
I 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 element s ii,8 ii,7 I,i,8 ii,7
Pectoral elements — 13 ii-11 i-13
Transverse scales — 13 /1 /14 1 0 /1 /1 0 11 /1 /13
Lateral-line scale s — “± 8 0 ” 79 81
8 Personal communica tion from Dr. Wol fgang Klausewitz.
bConverted from total length.
c Specimen gutted.
mann, 1905: 141-143 (Hawaiian Island s). Jordan and Seale, 1906: 218
(H aw a ii). Jordan and Snyder, 1907 : 208 ( “probably no t the same as the East
Indian S. commerson ii” \ counts ; young near shore and in mulle t pond s).
Iordan and Jordan, 1922: 29 -30 (common in Hawaii ; common name: “kaku” ;
size; should be compared with other large specimen s o f the South Sea s).
Barnard, 1925: 312 (synonym o f S. com m erson i ) . Jordan and Evermann,
1926: 3, 8 (Hawaii ; common name, habitat, range into fresh wa ter ). Schultz,
1943: 82-83; 256 -257 (Can ton Island; comparison needed with S. barracuda;
Phtheirich thys mu ltirad ia tus Schultz from S. snodgrass i) .
Sphyraena akerstrom i-Whitley , 1947: 131 -136, PL 11, Fig. 1 (original de-
scription; type locality: Lowenda l Island, Western Australia; extensive counts
and measurement s on four specimens; relationships, food , gonads; clo se “to
Weber and de Beau for t’s Sphyraena p icuda”) . Schultz, 1953 : 283 (S. akers trom i
Whitley the same a s S . barracuda Wa lbaum ). Smith, 1956 : 41 (same a s
barracuda ) .
Sphyraena ra/crops-Marshall, 1953: 54 -55 , Fig . 2, PI. 2 (identification as
barracuda questionable; origina l description; type locality: Moreton Bay, South
Queensland , Au stra lia ). Smith, 1956 : 41 (S. m icrops Marshall is the same a s
Agr ioposphyraena barracuda [W a lbaum ]).
Agrioposphyraena barracuda (Walbaum )-Smith , 1956: 40 -41 , PL I, Figs.
4a -4c (new combination; Fow ler ’s [1903 ] subgenus raised to generic level,
based on Esox barracuda Walbaum, 1792 ; synonymy , description, keys, mea s-
urements, range in Indo-Pacific, coloration, danger to man , food value ,
on togene tic change s in proportiona l mea suremen ts).

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 ( 1 9 2 8 ), Schultz ( 1 9 5 3 ), and Smith ( 1 9 5 6 ), accounts for
some o f the problems in comparing individua ls 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., akerstrom i Whitley; microps Marshall; snodgrassi
Jenkins) for which data are available and which are evidently synonyms of
barracuda. Data in the literature are often presen ted 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 apprec iated unless measuremen ts are presen ted for fish o f
comparable sizes, or unless the mea surements 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 (1 9 5 6 ).
s s
Methods of Measuring and Counting.— The following method were u ed
l s ts:
in making proportiona mea uremen
( ) l t : f s t f
1 Standard eng h tip o snout with mouth clo ed to midpoin o hypural
22
 TABLE 2
M e a sur e m e nt s of Sphyraena barracuda e x p r e sse d as p e r c e nt a g e s o f t h e sta nda r d l e ng t h f r o m the w e ste r n
A t l a n t ic O c e an ( in p a r e nth e se s) a nd the w e st e r n I n d i a n O c e a n ( S m ,i t h )
1956

( ) ( ) ( . ) ( ) ( ) ( ) ( ) ( )
Standard Length mm 60 6 1 110 111 5 150 149 180 182 370 369 900 8 6 1 1345 1111
t ( . ) ( . ) ( . ) ( . ) ( . ) ( . ) ( . )
Dep h 14 14 4 15 13 0 15 15 8 15 14 0 15 14 9 14 13 7 14 14 9
( . ) ( . ) ( . ) ( . ) ( . ) ( . ) ( . )
34 36 2 35 36 0 33 28 7 30 29
( .0 )
Head 38 40 0
( . )
37 37 0
( . ) ( . ) ( . ) ( 3 . 3 ) 30 ( . )
3.6 33
Eye 6.5 (6 .7 ) 5.8 58
(6 .0 ) 5.3 ( . )
5.2 ( 5 .7 ) 5 (4 .6 ) (3 .9 ) 2.7 ( . )
7 6 8 ( .7 )
6
Interorbital
t width 4.5 ( 4 .4 ) 5 ( 4 .6 ) 5 ( 6 .7 ) 6.5 ( 5 .3 ) 6.8 ( 5 . 4 ) 7.5 ( . )
Snou length 17 ( 17. 5) 16.5 ( 16. 7) 16 (16 .1 ) 15 (15 .3 ) 15 (14 .9 ) 11.8 (1 4 . 3 ) 11 (14 .6 )
12 14 4
Postorbital
l length 15 15 2
( . )
14 (13 .9 ) 12.3 ( 1 4 . 3) 13 (15 . 1 ) 13 (13 .0 ) 11.5 (10 .4 ) ( . )
Pectora
l f fin
l length 10 11( .0 ) 10 (10 .1 ) 11 (10 .3 ) 11 (11 .0 ) 12 11 ( .9 ) 11 1(1 .5 ) 12 11
( .1 )
Pe vic
t in ength 10 ( 9 .7 ) 10 10
( .8 ) 11 ( . 3 ) 10.5 1(0 .2 ) 10
11 ( 8 . 8 ) 7.5 ( 8. 6) 7.5 (7 .6 )
Dep h 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 ( 4 2 .1 )
Second predorsal
l length 75 ( 73 .0 ) 75 (71 .8 ) 72 ( 71 . 1 ) 72 (72 .5 ) 73 ( 70 .2 ) 67 ( 72 . 1 ) 70 ( 7 2 .5 )
Prepectora length 37 (37 .2 ) 37 (35 .5 ) 35 (35 .1 ) 35 (33 .8 ) 35 (31 .2 ) 30 (30 .9 ) 29 (3 1 .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 o f erect first dorsal fin.
(3 ) Second predorsal length : tip o f snout to origin o f second dorsal fin.
(4 ) Prepectoral length: tip o f snout to origin o f erect pectoral fin.
(5 ) Prepelvic length: tip o f snout to origin o f erect pe lvic fin.
(6 ) Preanal length: tip o f snout to origin o f anal fin.
(7 ) Head length: tip o f snout to posteriormost part o f fleshy tip o f opercle.
(8 ) Maxillaries: tip o f snou t to posterior border o f maxillary including the
premaxillary and maxillary bones.
(9 ) Snout length: tip o f snout to anterior rim o f fleshy orbit.
(1 0 ) Eye diameter: horizontal distance between rims o f fleshy orbit.
(11) Interorbital width: least width o f bony portion o f skull measured be-
tween orbits.
(1 2 ) Greatest depth: greatest depth measured at origin o f spinous dorsal fin
(fish with stomachs obviou sly distended with food are excluded ).
(1 3 ) Caudal peduncle : least depth.
( 1 4) Pectoral fi n length: length from upper base of erect pectoral fi n to ti p
o f longest ray.
(1 5 ) Pelv ic fin length: length from origin o f erect pelv ic fin to tip o f long -
est ray.
(1 6 ) Length o f spinous dorsal fin: length from origin o f first dorsal fin to
tip o f longe st depressed spine.

Scale counts were taken according to the methods o f 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 ( 1 956 ) data of barracuda from the
western Indian Ocean (Table 2) . It is seen t hat i n specimens of compar-
able size most measurements agree well for the two regions. Possible
sexual dimorph ism , if any , ha s no t been resolved bu t would pre sumab ly
lead to a mis int erpret ati on of t he dat a. My measurements of large specimens
in which the sex is identified are few, but at the extreme ranges those for
which I have dat a are females. Whitley’s akerstromi, which is shown to
be a s ynonym of barracuda, is repres ent ed i n Figures 2 and 3 and,
f
ortunate yl , h is two specimen s repre sen t both sexes. A comparison of thes e
i t t t t ts f t s
lim ted da a reveals ha he body par o he female eem to be consis-
t sli tl l i l ti t st t
ten ly though only gh y arger n re a on o andard leng h and head
l t .
eng h
s i l ts s t
Difference n proportiona measuremen between specimen from he
I s , ) t f t
Atlantic and ndian ocean (Smith 1956 are mos noticeable or he
i ls ); t is is t i
larger indiv dua (Table 2 h evidently due to he discrepancy n
i t st t s f t (
size n he large of he specimen compared rom he Atlantic 1111
t I i s ( ). I s i ls i
mm) and he nd an ocean 1345 mm n the e two indiv dua m nor
s i t t , s t t , i l t .
difference occur n he eye diame er nou leng h and postorb ta leng h
, t t it l t s s i t
However he in erorb a wid h seem consistently maller n he Atlantic
t t is fl ts iff i f s ,
specimens, bu possibly h re ec a d erence n method o mea uring
24
��������.....--,
50
45

40
45
40 .. '· ..--
�-�. "iti.
• ("A'.':i... B
35 ·"

30 . - � ·.
, -'t'J· •

751 ,," .;>,;:..,·,...... e ,, ··...:·: ..

• ••. : ••••• :,.�.• ,. �..., .¡'
· '\,:"'·.t'.·:. •
j 45
• ·.1.t;·•.;,· � •
. "-�
'·�· Á<!!
K
,:,,.,._·. a-'�·

l
10 ·.: ... 40
• . ,"J

•
��"
5 ·,•.·.1.o��·•.•..•...•.•.......;:�,l.",,
.�
.
O

•
'ºf '\ 35 •

10 ; •• ·• .

: ,." .
.
..
:�.
l
f .......,::�, ··-,:�..: ,.911..-. ,. .¡,.f,<o

.... . . .
5t . ' ··:-- ... . E

'� . . .

4C

35
 30 .,
25._......��--'����.. _ ,
1100 1
1000
0 10v0v0
rv OI 1100 110000 11000000

S TANDDAARRD
D LENG
GTH
H, M
MM
M. SSTAN
NDDAARRD LEN
NGGTH,M
MMM..

FIiGg Uu Rr E
F e 2. Ch anges in prropo rtion a l measuremcents w ith grrowt h of Sphyraena
barracud a (includ
om g synonyms akerstromi, micro ps, and snnooddggrraassssii)).. M easure-
ing
ments arre expressed as perccentage o f standard lleenngth. Dots arre western

Atla ntic spcecimens of barra cu da ; cirrcles aarree Innddiian Ocea n specimen s off barra-
cuda frrom Smith (19 95566));; solid ttriiaanngles arre ak erstrom i Whitley , 11994477, from
Western Australia ; solid sqquare iis microps Marshall, 11995533,, from Quueensland,
Australia; cross is barracuda,, U SN M 11447744555, from Djid da , Red Sea; solid
dia mond is snodg ra ssi JJenkin s, 1199001, USN M 46963, frrom Hon olulu , Hawaii.
A. Firrst prredorrsa l length. B. Prrepectoral llenngth. C. Secon d prredorrsal llength.
D. Interorbital width. E. Least! dep th ca ud al pceduncle. F F .. Len
ngth of ddorsal

spines. G. Head lleen

ngth. H. Snout length. I. Eye diameter. JJ. Prreeaanal length.
K. Preppeellvic length. L. Pect oral fiin length. M. Length of max illaries. N. Greeat-
est bod y depth. O. Pelvic f in length.
25
 since Smith ( 1956 ) did not state his method. Also noticeable is the marked
discrepancy in snout length between the two largest specimens. Perhaps
t is indicate s a rea l dif ference between he
h t indiv dua i ls from the two
regions, since, as suggested by the plots of ak ers t ro m i in Figures 2H and
3D, the difference does not seem to be sexual for this character.
Ratios, measuremen ts, 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 o f 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
t da at o f Schultz (1 953 ) since he presen ed
he t ra io t s bu t did not g ve i
original measurements.
Differences between measurements which cannot be attributed to size
d erence s and in which there is no overlap in ranges occur in ak ers tro m i
iff
and the western Atlantic specimens of barracuda in interorbital width
and in distance between dorsal fin origins. Marshall’s ( 1 953) type of
m ic ro p s evidently ha s a longer pec toral fin than does Smith ’s (1956 )
barracuda o f comparab le size from the Indian Ocean. However, the
proportions presented here for m i cro p s 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 merist ic counts f rom vari ous regi ons, includi ng pres umed
synonyms of barracu da, are comparabl e. The fi n- ray counts are pres ent ed
as taken from the lit erat ure. The pelvic and spinous dors al fi n- ray counts
showed no vari ation, t he f ormer being consistently 1, 5. Combining tot al
dorsal fin elements for ease of comparison ( cf. de Sylva, 1955: 30- 31), t he
count obtained is 10 or 11, with t he USNM specimen from t he Red Sea
t t f ll s is l
having he highes count. The range or a specimen 9 or 10 ana
s t t t t ts i s .
element and ha for he pec oral elemen var e from 12 to 15 Whitley
( ) s t fi - t i s
1947
f t
gavei. aI omewhat s higher pec oral sn ray f coun n, this det cription l
os akerss rom n we tern t Atlantic
, specimen it o( barracu ) da he la era i -line
cale t vary f from
f s 75 o s87 although t I Sm h 1956 . listed a maxstmum t
coun
t t o 90 s or pecimen from
t he ndian
l s Ocean
t il st ti since
However, f he a ed s(
ha he) ha counted f ,
abou 90 s sca e l on he t is ilu ra on o Riippell’, I
1 835 type s o agam t perhap t t t he inc uded h s n his count. However t s
previou s t ly pointed ou s hat he type itof agam ha only 81 scales, and. hu
perhap s f he t ranges s pre en l s ed by Sm t h should be somewhat t lower The
range
iff o
t ran s. ver e sca e around he body appear o be comparable for
d eren i i areal ts i s i s
Or g na measuremen for barracuda from var ou reg on have been
t i s . s ts f
plo iteds nt Figure t t2 and s 3 When points repre t enting
i measuremen
fi s, or,
reg on o her han he we tern Atlantic are plo ted n these gure rough
26
 55 •
� ·�.. 5

-•·-.
25
A
45
20 '

4
15
35 ' "
IO 2S'f,
•
5 25

• 20
•
25 • •
20
o
.
. '•
',
15
•
15 •. 1 ••
. , ·: ''::.:::;. ;�
�,·1� ·
8

•
55
' .
:..·_::.<�·.
.• . • ...."'.,;·!;. ,-,,.�.
10 ., .
·.¡, .
so
4 • .o·
.. • ·•
'/:":o' ' ,
.
",...
45
4
..
30
25 ' . ·"',·¡'·º .
. • $0, ••
e ••• G
. • ·º r -,
35 ',, '" '('
.' :,''.,-"
s-
. .'
55
25
H
45 20
.... •.·
40 o • 15

�
'-���IO,-��IOO,. ,_�...i
.
HHEEAD
O LEN
NGGTTHH,,M
MMM. HEEADO LEN
H NGTH,MM
M.
FIiGg Uu Rr E
F e 3. Cha n ges in prrop orrtiona l meas urements w ith grrowth of Sphyraena
barracuda (including synonym s ak erstromi, m icrop s, and snood
dggrraassssii)).. Measure-
ments ar e e xpr essed as p ercenrtages of the head length. D Dots ar e w western
Atlantic specimens o f barracud da; circles are Ilnndian Ocean specimens o f b arr a-
cuda from Smith (19 95566));; solid triannggles arre akerstromi Whitley, 1199447, fírornr

Westerm
n A ustralia ; solid sqqua rre is microps Ma rrsh all, 1199553, from Queens]land,
Australia; crross is barracuuddaa, U SN M 114477445555,, from Djidda, Red Sea; solid
diiaamo nd is snodgrassíi JJeennkiinnss, 11990
011,, U SN M 469 63 , from Honolulu, Haw aii.

A . Pelvic fin length. B

8. Ilnterorbital wid th. C. Pect oral fiin length. D. Snout
lleenngth. E. Length of maxillaries. FF.. Least depth cau da l peduncle. G
O. Greatest
body dep t h. H. Eye diameter.
visual comparisons may be made. If these figures are compared with the
data pres ent ed in Table 1, a number of f eat ures do not become evi dent
unless viewed graphically. In comparison with the western Atlantic
specimens, Sm ith ’s (1956 ) two large st fish have a somewhat shor ter
second predor sa l leng th , preana l leng th, prepelvic length, and snout length.
These comparison s are magn ified when the head leng th 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 (1 956 ) specimens of barracuda from the Indian Ocean differ
racially from those of the western Atlantic on the basis of these two
charac ter s. A stati stica l comparison o f specimen s from the two reg ion s
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 ) ak ers tro m i, which is said to differ from p icu da ( =
b arracu da) in the “s ize o f the eye and its ra tio s to o ther parts of the
head,” as well as in the length of the maxillary bones and the pectoral fin,
is seen to approximate t he points for West I ndian specimens very well .
The latter specimens show s omewhat great er dept h and int erorbit al widt h,
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 o f measuring.
Although Marshall’s ( 1953 ) description of m ic ro p s is inadequate for
analysis, possibly this species does not belong in the synonymy of barra-
cuda . The spec ific name, m ic ro p s , on which Mar sha ll apparently based his
diagnostic charac ter , doe s not refer to a differen tia ting character, as the
eye diameter falls well within the scatter of points represented by
barracuda . However, apparent distinction s 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
t premaxil lary , max llary
he i , and supramax illary . In h is illu stra tion (P I. 2,
Fig 2 he showed what appears to be an elongated supramaxillary,
. )
reach ing far past the eye, but with the maxillary considerably shortened.
Since the juxtaposition o f the two bones as depic ted by Marshall does not
i s I it s s t t t
occur n any phyraenid which have seen, eem ha he type of
i t i s t t i t l
m c ro p s may be malformed and he max llary hor er han n he norma
, t s i is l ; i t is s
situation and he upramax llary norma ly developed n h ca e
s ll t t i . t
Mar ha may have measured only to he end of he max llary If he
t t ti f t s i s ( I. ,
measuremen were taken to he p o he upramax llary a shown P 2
i . ) t t l si il t t i
F ^ 2 he measuremen would be more near y m ar to ha found n
t s s t t is si .
o her phyraenid a h ze
Unless Marshall’s type has a malformed pectoral fin, it is difficult to
explain the difference i n the relative length of the fins of barracuda and
m i cro p s of comparable size, unless there is a sexual difference, or unless
28
m icro p s can be considered as actually representing a valid species on the
ba sis o f this single character. Tentatively , however, I follow Smith (1956 )
in placing m i c ro p s in the synonymy of barracuda .
Neither the Red Sea specimen nor t he t ype of sn odgrassi Jenkins f rom
Honolulu seem s to diff er i n any res pect f rom t he wes tern Atlantic speci-
mens of barracuda, although perhaps diff erences i n proportional measure-
men ts would not be disclosed at s uch a s mall si ze. Both t he pectoral and
pelvic fins do appear to be somewhat longer i n snodgrassi i n comparison
with the western Atlantic species, bu t probably not significantly so.

Y o ung S tag es of the F am il y S p h y r a e nid a e

The smallest postlarval (Hubbs, 1943) specimen of barracuda collected
by me measured 11.9 mm st andard lengt h 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 Schm id t (1929 ) and Jespersen and Taning (1 9 3 4 );
data for hydrographic stations occupied where s phyraenids were collected
are summarized i n Table 7.
A graded series o f 171 barracuda collected from southern Florida and
Bimini, Bahamas, as well as from the DANA localities encompass a
si ze range of fis h from 5.5 mm ( to base of urostyle) to 1111 mm standard
lengt h. I n meas uring specimens, an ocular micrometer was used for those
up to 28. 7 mm, dial cal ipers readi ng to t he 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 o f 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
I
slands, Jama ca i , and Bermuda ; two indiv duai ls ten atively
t t ed
iden if i by
me as sph yraen a Linnaeu s occurred in a collection from the nor thea stern
coast of South America. Additional specimens from these areas represent
either b orea li s De Kay or pi cudill a Poey. These closely rel at ed species pose
a difficult problem at this time, but since boreali s is apparentl y rest ri ct ed
to the mainland of North America I am therefore tentatively ass uming
t th
ha t is fifth specie s ni the DANA collections is p cudi illa .
Through he courtesy of Dr Ernes A. Lachner, three drawings of
t . t
young sphyraenids by Dr. Louella E. Cable have been loaned to the writer
f rom t he fil es of t he United St at es National Museum. Two of thes e
repres ent guachancho and boreali s; the t hi rd drawi ng is of a specimen
9 mm total length which is labelled (<Sph yraen a barracu da ” from Beaufort,
North Carolina. Although I have not seen the specimen, this drawing is
29
 u

}<JiO
F gUuR
rEe 4. Ph o tograph s showim
ng
g pat term
n i n th rree specries of Sphyraen a from the
western N orth Atlantic Ocea n. A . Sphyraaeena bo rea lis, 6 6 .0 mm standard
lengthh; collected by S herrma n C. Bishop at Firre Islan d Inlet, Lon g Ilsland , New
York , Aug ust 1111,, 1199338, CornelJlI U niversit y N o. 627 7 . B . Sphyra ena barracuda,
6 2 .7 m
mmm standard length; collected by Do nald P. and Dooris D O. de S ylva at

Bimini, Bahamas, JJuully 1133,, 1199556. C. Sphyra ena g u achancho , 6 3.3 mm standard
length; collected by Donald P. de S ylva an d Nardo Zaias at Bear Cu1t, Key

Bisca yne, Mia mi, F

Aloorid a, Septemb er Payne. 26 ,
1199552. Photogrraap h by Douglass M..
M
 of a young specimen of S. borealis . The fleshy tip o f the lower jaw separates
it from barracuda , and the row of large, distinct melanophores along the
middorsal ridge, and the lack o f 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 shor t, and the head lack s the pigmentation o f S . sphyraena of
comparab le size ; fur thermore , 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 o f Brazil. Slight differences in pigmentation and size o f the eye are
noted between Dr. Cable’s drawing and the DANA specimen s o f 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, allopat ri c
species, a tent ative key is present ed for t he ident ifi cation of postl arva l
sphyraenid s for spec fied
i size group s. On yl two po stlarva l specimen s of
S. sphyraena have been i dent ifi ed, both of whi ch were i n poor condition;
thus, characters based on these specimens should be used with caution.
Characters used to ident ify t he young cannot be used to s eparat e t he
adult s since they may not be present i n t he young. The key is s ubdi vided
roughly into fish from 5.5 to 11.9 mm, f rom 12.3 t o 130 mm, and fis h
greater than 130 mm. Since t here is no pronounced metamorphosis i n
h t isfamily , and because here
t isconsiderable var ai ion
t ni he
t color pa ttern
f s f
o specimen o comparable ze here si , t is ac ua ly no trenchant break
t l
between he key those specimen wh ch fall near t he ext remes of t he
t s; s i
ranges should be checked using both keys.

K ey to S p ec ies of S p h y r a e n id a e for S p e c im e ns of
I
5.5 TO 11.9 M M FROM THE W ESTERN N O R T H A TLA N T C
la . Head compara tively long , about 34 -39 per cen t o f the standard length;
body not heavily pigmented; distinct melanophore patterns on dorsal and
ventral ridge and along posterior part o f lateral line only; fleshy tip o f
lower jaw developed to a length less than half the eye d ia m e te r .............2
l b. Head comparatively short, less than 30 per cent of the standard length;
f
body heavily pigmented on posterior part f o body and s about supraoccipital f
region; fa distinct mid-lateral; row o t melanophore f j along most ot the
length
i ot the caudal peduncle fleshy ip o lower aw , nearly equal o eye
d a m e e r ....................................Sphyraena sphyraena European barracuda
. s l t t
2a Melanophore along middorsa t ridge no no iceably larger (than i s. surround-
, );
ing ones, scattered
l and no distinctly arranged ; tiseriallyf F g 5B s C
supraoccipita region t ll not noticeably pigmented p o snout with light
pigmentation, if a a .........................................................................................3
. l s -
2b Middorsa line straddledt by relatively llarge, distinct melanophore f s ft ar
ranged serially from s t poin above pectora
f s ft l origin to end o
s, if o dorsal-fin
base; melanophore( i . a); bases f o o dorsa and anal s fin present,
l small
;
and
f diffuse F g 5 A a ew large melanophore on occipita region tip
o snout generally heavily pigmented ............................................................ 4
31
 •'
� .•. • ..••••••• ··.-... :.1,".\ ••
• ·• •• ' • . • ''• • • ' .. : :f�.· :;'Se, T

•..·.·�
·. ·. ... ;,----._.
··
_;.,:
e
.

-- c;g -----
-- . ·--�-��===�.,,_-_._.._...·_
�

o -

-��
�- ·:·.. -····· .... ·-·-··". --. ....
':fA
.;;; -
4!!ib F
F1iGgUuRr ce
F 5. Draw ings shoow
wing pigment arry ddifffeerreenncces iinn tthhee postlarrvae of tthhrreee

sppeecciies of Sph yra ena from tthhee western North Atlantic. A. Sphyraena picudilla,
ddorsal aspcect. B . S phyraena barracuuda, dorssal assppect. C. Sphhyra ena guachann--
cho, dorsal asppect. D. Sphyra ena picud illa, ventral aassppect. E. Sphhyyrraena
barracuda, ventral asppeeccct. F
F .. Sphyraena gu
uaacchhaanncchho, ventraal aassppeecct. Alil
specimens aarre abboouuct 9 mm sttaanndard Jlength.
3322
3a. A row o f melanophores along midventral ridge extends posteriad only
slightly past tips o f apressed pectora l fins; tip o f lower jaw no t noticeably
conica l or projecting beyond snout; pigmentation on tip o f lower jaw and
snout sparse ; lower jaw obliquely rounded in dorsa l view; teeth nearly
vertical in lateral view; faint pigmentation on ly on either side o f dorsal
midline; melanophores at base o f dorsal and anal fins rather large, dense
F( gi . 6 A D - ); preorbita l pigmentation usually distinct, subtriangular ni
outline; no hourglass bands on posterior sec tion o f trunk; snout rounded
above , dorsal aspect o f frontal region sigmoid in lateral view , particularly
in specimens less than 7 mm long; lower posterior margin o f opercle
without spines; opercle ends in a single point; maxillary series nearly
straight when v iewed laterally . . . . Sphyraena barracuda , grea t barracuda
3b. Midventral row o f melanophore s extends posteriad from isthmus to ven t
(F ig . 5 F ); tip of lower jaw projects noticeably beyond snout ( as compared
to barracuda), ending in a bluntly conical, fleshy knob; lower jaw ti p
heavily pigmented; snout and lower jaw tip acute in dorsal view ; teeth
directed obliquely backward, rather than vertically; faint pigmentation
only on either side o f middorsal line, melanophores at bases o f soft dorsa!
and anal fins diffuse; two hourglass-shaped crossbands on sides in larger
specimens: the first begins about two-thirds o f the distance between the
pectoral fin origin and the anal fin origin, the second commences just be-
hind rear o f anal fin base; preorbital pigmentation generally no t distinctive;
dorsa l profile o f snout nearly straight i n l ateral view; lower posterior
margin o f opercle with three weak spines; opercle ending i n two points;
maxillaries rather strongly arched in lateral v i e w ..........................................
Sphyraena guachancho , guaguanche
4a. Preorbita l pigmentation extending forward to snout tip as a line of
melanophores; dorsal profile o f snout nearly straight in lateral view; caudal
base with a vertica l series o f melanophores ; eye larger in specimen s o f
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 i n specimens
o f comparable s i z e .........................Sphyraena borea lis, northern barracuda
K ey to S p ec ies of S p h y r a e nida e f or S p e c im e ns f r o m
1 2.3 to 130 m m S ta nda r d L e ng t h f r o m th e w e s t er n
N ort h A tl a nt ic
l a . Spinous dorsal fin placed well behind origin o f pectoral fin; pectoral fin
reaches origin o f pelvic f i n ................................................................................2
lb . Spinous dorsal fin placed above or slightly in fron t o f origin o f pelvic fin;
pectoral fin doe s no t reach origin o f pelvic f i n ...............................................3
2a. Last rays o f soft dorsal and anal fin longer than anterior rays; teeth
noticeably directed backwards; scales, when completely formed, 108 to
114 in lateral line; interorbita l noticeably convex in large specimens, with
raised longitudinal frontal and parieta l cre sts; 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 o f it, its scales noticeably l arger t han the s ur-
rounding scales; circumorbital and opercular region with silvery reflections
in preserved specimens; upoer surface o f tip o f lower jaw blackish ; tip s
o f middle caudal ray s usually blackish with center rays darkest, stippled;
three hourglass-shaped band s on body (F ig . 4 C ) : the first forms a dusky
saddle on back beginning at about the last spine o f the first dorsal fin
and extends posteriad to abou t one-third the interdorsa l distance ; a second,
wider saddle extends vertically to below dorsal base, become s narrow at
 the lateral-line region, and thence again flares at region o f anal base; a
third hourglass mark occurs ha lfway between the second dorsal fin and
the caudal fin ba se, similar to the first two; margins o f dorsal fins blackish
with pale bases; pectoral fin dusky, other fins pale; upper margin o f
opercle p a l e .......................................... Sphyraena guachancho, guaguanche
2b . Las t ray s o f soft dorsa l and ana l fin equal to or shorter than anterior ray s;
teeth nearly vertica l in la tera l view in most specimens; scales, when com -
pletely formed, 75 to 87 in lateral line; interorbital area flattish to concave,
without raised longitudinal ridge s; maxillaries nearly straigh t in la tera l
view; opercle ends in a single point; l ateral line whitish, not distinctly
raised or noticeable; sides o f head seldom with silvery reflections , and, if
presen t at all, only about opercular bones; t ips of middle caudal rays
diaphanous; outer caudal fin ray s dark , except for their tip s; a sub tr i-
angular, darkly pigmented region overlaps base o f middle caudal ray s
and caudal base; side s with seven to 12 variable markings which form a
longitudina l band in fis h less than 30 mm; these form contiguous H-shap-
ed blotches in fish to about 60 mm (F ig . 4B ); in larger individuals some
o f the blotches may extend to and coales ce with the dorsal pigmentation;
soft dorsal fi n with white tip in large specimens of t his size range; anal
fin base pale; base o f pelvic fin s pale, re st o f fin dusky; pectoral fin pale
except at its base; upper margin o f opercle d u s k y ..........................................
Sphyraena barracuda , great barracuda
3a. Scales very s mall, about 150 to 160 i n l ateral line; fleshy tip of lower jaw
well developed; pigmentation on back and sides obscure, not forming a
distinctive p a tt e r n .................... Sphyraena sphyraena , European barracuda
3b. Scale s larger, from 110 to 135 in the lateral line; p igmentation on back
and sides well developed, forming a series of quadrate saddles on back,
alternating with subquadrate rectangle s along sides (F ig . 4 A ) .................. 4
4a. Eye large in specimens o f comparable size, averaging 8.0 per cen t o f
tandard
s length ni pecimen s s o f about 50 mm , and 7 0. per cen t in speci-
men s o f abou t 100 mm; interorbital space convex; median ridge o f frontal
groove no t d e v el o p e d .......................................... Sphyraena picudill a, sennet
4b . Eye relatively smaller in specimen s o f comparable size, averaging 7 .0 per
cen t in specimen s o f 50 mm standard length, and 6 .0 per cent in specimen s
o f 100 mm ; interorbita l spacefl attish ; median ridge fo fronta l i groove
p r o m n e nt ...................................... Sphyraena boreali s, northern barracuda
K ey t o t h e S p ec ies o f S p h y r a e nida e fo r S p e c im e ns
LONGER THAN 130 M M FROM THE W ESTERN N O RTH A T L A N T IC
la . Origin o f spinous dorsa l fin definitely inserted behind origin o f pelvic fin;
tip o f apressed pectoral fin reache s origin o f pelvic fin; tip o f lower jaw
without a distinct, fleshy t i p ........................ .....................................................2
lb . Origin o f spinous dorsal fin above or slightly in fron t o f origin o f pelvic
fin; tip o f apressed pectora l fin falls considerably short o f origin o f pelvic
fin; tip o f 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 cle ft to the hypural fold; when fi ns are depressed, the last
rays o f the so ft 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 s ection o f the trunk in s pecimens 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
o f pelvic and anal fin black; tips of middle caudal rays black; size to
about 24 inches; Wood s Hole , Massachusetts, to Brazil and throughout the
34
 Gu lf o f Mex ico and Caribbean Sea, drifting to the eastern A tlan tic ...........
Sphyraena guachancho , guaguanche
2b. Scales large, 75 to 87 in the lateral line; when fins are depressed, the
anterior rays o f the soft dorsal and anal fi n 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 o f
trunk ; adults deep green to steel gray on back, sometime s reflecting
purplish hues; mostly silvery on sides, abruptly becom ing white along
ventral surface; upper sides with 18 to 22 oblique, backward-directed
dark bars in life; lower sides with several to many inky blotche s which
are variable in size and posi tion (F ig . 7 D ); so ft dorsal, anal , and
caudal fins black to viole t, with whitish tips; pectoral fin pale except for
its upper margin, which is black; upper border o f opercle dusky to black ;
length to about 6 feet; Woods Ho le , Massachusetts , south to Rio de
Janeiro, Brazil, and throughout the Gu lf o f Mex ico and Caribbean Sea ;
also occurring in a ll other tropica l waters o f the world excep t the eastern
Pacific ............................................. . Sphyraena barracuda , grea t barracuda
3a. Scales comparatively large, 110 to 135 in the lateral line; depth o f body
12.0-14 .3 per cent o f the standard l e n g t h ..................................................... 4
3b. Scale s very small, from 150 to 160 in the lateral line; depth o f body 10.0-
11.1 per cent o f the standard length; length to about 24 inches; eastern
Atlantic from the Bay o f Gascogne to Mossamedes, Angola , and the
Mediterranean and Black seas; also recorded from Bermuda and Rio de
Janeiro , B r a z i l........................ Sphyraena sphyraena , European barracuda
4a. Eye large in specimens o f comparable size, averaging 6.3 per cen t in
standard length in specimen s 150 to 200 mm, 6 .0 per cent in specimen s
250 mm , and 5.5 per cen t in specimens 300 mm ; body depth abou t 13.9 -
14.3 per cent o f standard length; scales in lateral line 107 to 116; inter-
orbital area convex; median ridge o f fronta l groove little developed ; length
to about 18 inches; Tortugas, Florida, Bermuda, and throughout the
An til le s, south to Rio de la Plata, Uruguay . . .Sphyraena picudilla , sennet
4b. Eye smaller in specimen s o f comparable size , averaging 5.3 per cen t in
standard length in s pecimens 150 mm s tandard length, 4. 9 per cent i n
specimens 200 t o 250 mm, and 4. 7 per cent in specimens 300 mm; body
depth about 12 .0 -13.0 per cent o f standard length; scales in lateral line
from 118 to 135; interorbital area fl at tis h; 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 Domi ngo, and Rio de
Janeiro .......................................... Sphyraena borea lis, 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 ustration showing the postlarval developmen t of S . sphyraena
ill
from the Med iterranean by Lo Bianco (1 9 3 7 ), the illu stra tion s of S .
pinguis and an uniden tified spec ies by Uchida , e t al. (1958 : 46 , 47 , P I. 44,
Figs. 2-4 , P L 47, Figs. 1 - 8 ), and Orton’s ( 1955 ) work on the develop-
men t of argen tea apparen tly constitute the reference s on this subject. Illus-
t tion s o f young spec men
ra i s of barracuda are ound f ni Fow erl (1 9 4 5 ),
Gordon ( 1 9 4 6 ), and Breder ( 1 9 4 9).
Essentia l to a study o f the systema tics of Sphyraena barracuda is a
knowledge of ontogenetic change n s propor i ona ti meal uremens For
purpo ts.
e s of s uillrast onti graph , showing
s change n s ire a lonti to het head
length (Fig. 3 ), as well as in relation to standard length (Fig . 2 ) , are
35
presented. When head lengt h is used as a base, it tends to magni fy
variation, and thus it appears to be a bett er indicator of i nt ers pecifi c
differences, since, depending upon t he species, diff erent 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 developmen t in Sphyraena barracuda from 5.5 mm to 11.9 mm
standard length. A . 5.5 mm standard length ; co llec ted by R /S DAN A ,
Station 1293-V , 17 °43 'N , 64 °56 'W , April 17, 1922 . B. 6 .6 mm standard
length; co llec ted by R /S D A N A , Station 952 , 17 °55 'N , 64 °48 'W , May 12 ,
1921. C. 8 .6 mm standard length; collected by R /S D A N A , Station 1352-V ,
35 ° 42 ' N , 7 3 ° 4 3 /W, May 21, 1922. D . 11.9 mm standard length; collected by
Donald P. de Sylva, 1 mile southwest o f the harbor entrance o f North Bimini,
Bahamas, June 6, 1956.
36
 seldom be disclosed by an examination of t he regressi on on st andard l engt h.
Figure 2 indicates that of the proportional measurements taken, only
preanal length and pectoral fin length grow isometrically with an i ncreas e

F i g u r e 7. Drawings showing changes in pigmentation and body form with

postlarval developmen t in Sphyraena barracuda from 17.2 mm to 790 mm
standard length. A . 17.2 mm standard length; co llec ted by Donald P. de Sylva,
1 mile southwest o f the harbor entrance o f Nor th Bimini, Bahamas, May 12,
1956. B. 23.7 mm standard length; collec ted by Donald P. and Doris D. de
Sylva, at beach east o f 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 o f Florida
No . 7072 . D. 790 mm standard length; collec ted by Doris D. de Sylva, north
o f North Bimini, Bahamas, 25 ° 48 'N , 79 °17 'W , July 18, 1956.
37
in the lengt h of t he fis h (Martin, 1949). Most pronounced allomet ry is
seen in changes in the first predorsal, prepectoral, and prepelvic distances,
and in the head and pelvic fin leng ths. A s might be expected, inflec tions
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 o f
t
he size range of he
t barracuda a fter it ha s reached 20 mm .
In the following discussion of the development of barracuda, reference
is made to Figures 2 to 7; body lengt h ref ers to st andard lengt h unless
otherwise stated. Original measurements are presented by de Sylva ( 1958 :
332 - 345 ).
In the s mallest specimen examined, 5.5 mm (measured to t he base of
the uro style in th is specimen), the yolk has evi dent ly been abs orbed f or
some time, and t he t eet h are comparatively well developed. The st omach
o f this specimen contained fish fragments , indicating tha t at this stage it is
a lready preda tory . Orton (1955 ) noted tha t 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 o f barra-
cuda depict ed i n Figure 6A mi ght perhaps be about six or seven days old.
The snou t and max illarie s are ex tremely shor t, with the la tter repre sen t-
ing 12.2 per cent of the leng th. The snou t profile is sigmoid in outline and
the snout tip is rounded. The large eye represents 24 per cent of the head
leng th , wh ich in turn is 36.6 per cent of the body length. The body is
re la tively deep at this stage, and repre sen ts 42 per cent of the head leng th
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 dor sa l sur face and a s a ser ie so
f
concen t melanophores along e her
rated it desi of het ana ln fibase (Fig.
5 E ). 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 , w ith no traces of suppor ting ray s. Myomere s 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 adu lt (Fig . 7 D ), and , n
i fac t, is somewha t heterocercal, with the lower
lobe the longer.
A t 6.6 mm (Fig. 6B) t he eye has noticeably decreased i n si ze, and
repre sents 19.3 per cen t of the head leng th . Relative depth has increa sed
slightly, as has the pectoral fin length, the base of which is still evi dent
38
as a fl es hy pedicel. The snout has lengthened noticeably, from 12.2 t o
18.1 per cent of the body length. The maxillaries are well developed
and are expanded at t he posterior margi n. Preorbit al pigmentation has
become still more distinct, although the supraoccipital melanophores, at
leas t in th is specimen , are fewer than in the earlier stage . The sub - and
interopercular bones are beginning to increas e in si ze and extend f urt her
ventrad. Body pigmentation has increas ed considerably, but is still concen-
trated along the middorsal and midventral lines, and a band of melano-
phores occurs as a broken line along t he pos terior part of t he t runk
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 fi ns are enclosed i n a fl es hy s heat h whi ch ext ends abou t
one-third of the height of the rays. The urostyle has now become nearly
ver tica l and the hypura l plate is fu lly formed , although a trace o f the
upturned urostyle remains. Onl y a sli ght i ndentat ion is seen in t he caudal
margin.
Little change occurs between 6.6 mm and 8 .6 mm (Fig. 6C) except
t t he
ha t shape of he t snou t is near yl stra gh
i t ni la era
t l view . The re al tive
decrease in depth is not as great as the figure indicates, but there is a
decrease in the size of the eye and a sligh t increa se in the leng th of the
head, snout, and maxillaries. At this stage the maxillaries reach their
maximum mean length rel ative to t he body l engt h. The fis h closely
resembles the adult in general body outline, notwithstanding the absence
o f the pelvic and spinous dorsal fins, and the teeth are longer and more
robu st. The ana l and sof t dor sa l fin s are re latively longer , and the fle shy
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-
ta tion is concen trated posteriorly along the la tera l line, and a t the
i
m ddor sa l and m dven
i tra l sections; the la tter pigmentation ha s increa sed
and ha pread an er ad . Branching and the formation of str iae in the ou ter
s s t i
ray s o f the caudal, so ft dor sa l, and ana l fin have increa sed considerably ,
and striae in the pectoral fin rays are first eviden t at this stage. The
urostyle is still visible through the flesh, but the increased thickness o f
t fle sh obscure s he
he t myomere coun t ni he t an er t or
i runk
t section .
A 11.9 mm a noticeable change ha occurred and he fish has
t s t
assumed distinctive juvenile characters (Fig. 6 D ). The pelvic fins are
noticeable and represent 2.3 per cent of the body length; they appear
fi
r sta t8.8 to 10.1 mm .
The spinous dorsal fin first appears at between 10.1 and 11.9 mm; its
depressed s pines repres ent 1.9 per cent of t he body lengt h when 11.9 mm.
The head is rel atively longer t han i n previous stages, and t he eye corres-
pondingly smaller. Cephalic pigmentation is marked about t he s nout,
i l i , it
supraocc pita reg on and along the pre - and postocular reg ions, w h
39
 sugges tion s o f an incipient band along the head from t he snout to t he
preopercle. The s ub- and int eropercle have completely covered t he
branchiostegal tips, but t he l att er bones are still vis ible t hrough t he
opercular bones. Although mi ddors al pigmentation is more extensive t han
at the previous stages, the conspicuous changes are seen i n t he ant eri ad
encroachment of t he melanophores along t he lower si des of t he body whi ch
appear as s mall, indistinct blotches, rat her t han as general ized, stellat e
melanophores . Fur ther cro ss striations occur in the ti ps of t he fi n, and
fi
r st sign of branching ni these ray s is seen ; also , there is a defi nit e
indication of a bifurcation of the caudal fin.
A t 17.2 mm (Fig . 7 A ), 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 dors al and pelvic fi ns, and t he
rays of the latter have already begun to show striae at their tips. The
melanophores are now more diffuse along t he cephalic rerion. These
coalesce along the snout to become confluent with the lateral pigmentation
of the trunk, s uggest ing a diff us e, l ongi tudinal band; t he cont rast between
th is dark band and the wh iti sh ven tra l area is striking. The cauda l fin
has as sumed a defi nit e bifurcat e appearance wit h acut e lobes appearing as
a transitional stage between those illustrated in Figures 6D and 7B.
At 23.7 mm (Fig. 7 B ), the lateral band of melanophores is beginning
to break up on the interradial membrane. The soft dorsal and anal fin
a ssume a juvenile shape and are well formed . Relative maxima are seen
ni het leng th s o f the prepelvic , head , and snou t. The ra te o f increase n i
t leng th of the depressed dor sa l spine s ha s dropped o ff ra ther sharp ly .
he
Deep forking of t he caudal fi n is noticeable, and it has revert ed to t he
rounded-lobe situation . The tee th are longer and sturd ier , and there is
evidence of specialization o f the anteriormo st teeth into longer canine s.
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
la tera l-line section o f the cauda l peduncle, bu t are less noticeable than in
borealis (see Nichol s and Breder , 1927 :73 ).
Noticeable changes other than in proportional measurements and in the
exten t o f scalation and color pa ttern occur between the stage il lu stra ted
ni Figure 7B and tha t o f the 213-mm specimen o f Figure 1C. A tran sition
occurs in the pa ttern from a partially broken longitudinal band through a
s s i tt t lt tt .
serie of H-shaped blotche pr or to a aining he adu pa ern The
l t
horizonta connections between these are even ually lost and form oblique,
s t s . ft s l, l,
wavy band along he upper side and back The so dor a ana and
l i ti s, t t fi is s .
cauda lobes may show wh te p and he pec oral n pale to du ky
fi s its t l s i i s.
The pelvic n ha cen ra ray blackish with wh te marg n
t st s i s
Between he two successive age depicted n Figure 1C and 7D
s t l t
numerou inky blotches appear on he cauda peduncle and along he
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
i the grea te st dep th , while the ra te of change in the pec tora l leng th and
n
t in erorb
he t ita l rema ni s unchanged between Figure s 7C and 7D. For
excellent photographs o f living specimens of barracuda abou t 1 to 2 inches
long, see Gordon ( 1946 ) and Breder ( 1 9 4 9 ).
The adult is represented by an illustration of a specimen 790 mm
standard length (Fig. 7 D ). 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 t he back to a s eri es of 18 to 22 cross bars
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 ti p 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 ( 1 952 ) show the variation in the number and position of these
blotches.
Significant proportional changes which occur between the juvenile and
he
t adu lt (Figs. 2 and 3 ) are seen ni he t decreased re ative
l leng h
t s of he
t
prepelvic distance, head, prepectoral, and diameter o f the eye. A somewhat
esser
l ra te o f change with increased leneth is observed n i the leneth o f the
nout,
s leng ht of spinou sdor as l fin , and pelvic fin . Nearly isometric grow ht
between the two stages is observed for the length of the preanal, the depth
o f 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 t he change i n the t ail from a f orked
ot a scalloped marg in . Three more or less semicircul ar indentat ions are
observed which give the margin a quadrilobed appearance; this characteris-
41
tic is even more noticeable i n l arger specimens. This was noted by
Riippell ( 1 8 3 5 ) for agam and by Bleeker ( 1854b) f or com m ersoni, bot h
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 t he ant eri or border of t he eye i n t he j uvenile to a
line passing through the posterior margin of the eye in a very large adult.
In the early st ages t he maxi llaries fall f ar s hort of the ant eri or border of
the eye.
D iscu ss io n .— Allometric growth, or heterauxesis (Martin, 1949 ), is notice-
able in barracu da , 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 allome try is seen, after the 20-mm stage is reached ,
in the rel ati ve lengt h of t he head, maxillaries, s nout, prepect oral, fi st r pre-
dorsal, and prepelvic, and in eye diameter; positive allometry occurs in
the body depth, interorbital width, and pectoral fi n l engt h. Isometric
growth occurs only in preanal length. By using head length as a base
length (Fig. 3) it is possible to fit to these dat a an ass ort ment of parabolic ,
hyperbolic, and thi rd- degree polynomial curves, but s ince head l engt h
shows negative allometry itself, s uch curves are diffi cult t o visualize i n
term s o f body grow th .
The inflections noted in the growth data may reflect changes in the life
his tory of the barracuda . Although ecological aspects are treated under a
subsequent heading , it seems appropriate to interject here some comments
on the dis tribu tion o f the young barracuda . It appear s tha t most spawn ing
occurs at a considerable dis tance from land , probably over deep wa ter
between the deeper reefs and the edge o f the continental slope, and that
t t s t j ts t
he young are subsequently ran por ed both by ma or curren to o her
s ti l t
region and inshore by da movement. Postlarvae up to abou 12 mm
f t s t i t
occur ar ou to ea bu only barracuda 15 mm long and above ev den ly
i litt l t s. It is s t t t t is
occur n ora wa er ugges ed ha during h shoreward move-
t t s s t
men they undergo a moderate ran formation which include a ra her
i t t f , t f t ,
sudden increase n he exten o pigmentation he lengthening o the eeth
t t s s s l fi s, s s i
he appearance of he pinou dor a and pelvic n a well a a rap d
i st t f t t l fi s, t
development n function and ruc ure o he ver ica n he pelvic, and
l fi s. t it s tt t f , t
cauda n By s he time l ha a ainedtta leng l h; oit is18 ttot 22
is tmms ihe
barracuda
i s ha genera ly adapted t to sa li ora lifef t . a h l rant ititon,
n habit and ecology t when he
t graph
f show an in lec , ions Sexua ma ur y
which
f t occurs
i a a fork s leng h. o 460 to 660
). mm doe not appear to be
re lec ed n the graph (Figs 2 and 3
i ti i t lt . ( )
D escr p o n a n d S ze o f he A d u s — Jordan and Evermann 1 896
t f t lt . ,
presen
s l ed an extensive description s f ot he adu t barracuda Adequate
upp ementary description o a quali ative na ure are available for both
42
 the young and adu lt fish in Breder (1948a , 1949 ), Bleeker (1854b ),
Day (1875 -1878 , 1889 ), Fowler (1 9 2 5 ), G ilchris t and Thompson (1 9 0 9 ),
Meek and Newland (1 8 8 4 ), Meek and Hildebrand (1 9 2 3 ), Poey (1 8 5 8 ),
Schultz (1943 , 1953 ), Sm ith (1 9 5 6 ), Weber and de Beaufort (1 9 2 2 ),
and Whitley (1 9 4 7 ). Quantita tive charac ter s are listed in Table s 1 and 2 .
The max imum size a ttained by the adu lt barracuda ha s been one o f
speculation. Most estimates of the great size attributed to this species
can be traced to the writi ngs of earl y nat uralists. Labat ( 1724) report ed
barracuda mea suring 18 to 20 feet long from the Antilles, and Fermin
(1 769 ) recorded specimens 15 feet long f rom t he coast of Suri nam.
Bti ttiko fer ( 1890 ) recorded a 10-foot specimen of S. jello from Liberi a,
and Vincent ( 1910 ) measured a specimen of barracuda from Tri ni dad
which was 8 V2 feet long. Popular stories often relate the sighting of 7- or
8-foo t barracuda in the Florida Keys , bu t such fish must be uncommon ,
for no fish longer than 63 inche s to tal leng th occurred in the record s o f
Mr. A1 Pflueger from 1953 through 1957.
The largest fis h f or whi ch I have meas urements was a female caught
off Miami Beach on March 18, 1956, whi ch weighed 39 pounds and 9
ounce s and wa s 55.5 inches total length. Gudger’s ( 1 918) 38-pound
barracuda was 55 inches long, and Herre ( 1936) report ed a s pecimen
o f barracuda from the Philippines wh ich was 62 inches long and weighed
25 kilograms (55 pound s). Fish weighing upwards of 50 pounds are
evidently no t rare in the Florida Keys, although I have seen none under -
water , e ither in the Florida Keys or in the Bahamas , wh ich would exceed
40 or 50 pounds. These gi ants are not oft en caught by angl ers bu t
are occasionally taken by s kill ed s pearfis hermen. Fish weighing 45- 50
pounds seem to represent t he maxi mum weight ent ered i n local fis hi ng
tournaments ni M am i i dur ngi recen t year s.
The world-record barracuda caught on hook and line which is recog-
nized by the In terna tiona l Game Fish Association (LaMonte, 1952 )
weighed 103Va pounds and was 66 inches in total length, wit h a girth of
3 1 ^ inches. Although it was only 4 inches longer t han t he speci men
reported by Herre ( 1 9 3 6 ), it weighed nearly twice as much. This record
fis h was taken at West End, Bahamas, i n 1932 by Mr. C. E. Benet.
However, Herre ( 1 936) report ed a 106-pound specimen of barracuda
caugh t w ith rod and ree l by Genera l 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 o f the young by Breder (1949 ) and Erdman
( 1 9 5 6 ). Malpas ( 1926 ) and Williams ( 1 956 ) examined gonad samples
from S. commerson i ( = S . barracuda) from the Indian Ocean. Goode
( 1876 ) mentioned that S. picuda ( = 5 . barracuda) commonly spawned
in Bermuda , and Holder (1924 ) sta ted tha t barracuda in the Florida Key s
43
 1.i.---SV
�--AT
V
.---BA
L

'-----GB
;J----SP
;¡ ST
1,1---SB
,I----FB

F1i0gUu R
F rE
8. IInnterna!l or gans in situ of a riippe male ssppecimen of Sphyraena barra-
e
cuda, 8 27 mm f ork length, collected from B Bimini, Bahamas, on JJuully 7 , 1199556
6..
Scale two-fifths naturral size. SV V== sinuus venos us; AT T== attrriiuum; Y
V== ventr icle;

BA
A== bulbus arrtteeriioosus; L-
= liver; PC
C== pyloric caecae; GBB=
= galll bllaadder;
SPP=
= spleen; ST
T== stomach ; SBB=
= swim bbllaadder; FB
B== fat boddies; DT
T== diigges--
tive tract; T=
= ttestis; AO
O== aorta.
4444
 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 na ds o f Sphyraena barracuda
Female s
Stage 1 “Resting” ; ovary full o f immature eggs only , no t showing up as distinct
granules; ovary pink ( young individual s) to reddish (older indivi-
dua ls), gelatinous and translucent; blood vessels generally no t visible;
lumen easily visible.
Stage 2 Ovary ripening, full o f maturing egg s; pinkness fading to a yellowish
color; ovary opaque; blood vessels beginning t o appear as faint, reddish,
transverse stripes; transverse fold s beginning to appear.
Stage 3 Ovary nearly ripe; small eggs are visible whi ch 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 fo ld s 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 consi stency ; vein s st ill purplish bu t
less turgi d; transverse f olds obscure; ovary has lost its whitish color
and becomes redder as it approaches s tage 1; oft en difficult to distin-
guish between late stages o f 5 and early stages o f 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 no t flow out when testes
are pressed.
Stage 3 Testes thicker than stage 2; b lood vessels further developed; milt flows
out readily when ducts are pressed.
Stage 4 Spent; testes slender and flaccid;blood vessels reduced ;milt no t present.

spawned in t he “earl y spri ng. ” Fowler ( 1905) collected young barracuda

from New Jersey, and ( 1945 ) recorded speci mens to 2 inches i n l engt h
from Key We st, Florida , in December. LaMonte (1 951 ) reported that
“t he United St at es Fish and Wildlife Servi ce at Beaufort [ Nort h Carolina]
reports many barracuda from six to eight inches long, at whi ch si ze they
are ma ture o ff the Beaufort-Morehead City section . . . .” However, it
seems that the species referred to here is confused with another sphyraenid,
since no ma ture indiv idua l o f S. barracuda less than 20 inches long ha s
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. j ello and S. obtusata f rom t he
Philippines.
Sexual D imorphism .— There is no sexual dimorphism in general appear-
ance and no appreciable difference in the sexes in the length-weight
45
1-JiGgUu Rr E
F e 9. Hristologitcal sections o f the gon a d s otf S �phyraen a barracud d a. A. S ec-
tion of immature ovarry (x x3399)),, 544 mm , collect ed JJuuly 1188,, 11995566. B . Secticon of
nearly rriippee (stage 3)) ovary (x x55..55)),, 1122008
8 mm, collectecd July 8, 11995555.. C. Section
of riippee ovary ((xx55..33)),, 1111449
9 mm, collecte d JJu ullyy 9, 1199555. D. Section of ripe

tesntiss (xx3355)),, 971 mm, collected JJune 8, 1199556. Mea su rrem e nrts aare fork l[engths.
S pecimens rreprresen lted in A, B
8, and C w ere collected alt Bimini, Bahamas, and
specimen rreprresented iin D O was collected at Ca t Cay, Bahhaamas. Photographs
by Do uglass 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 tran sver se ly and placed in wa ter , the edges tend to cur l. The te stis
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 Developmen t.— The criteria given in Table
3, modified somewhat from Walford’s ( 19 3 2) st udy of S. argentea, have
been found t o be s at isf actory f or ident ifying t he vari ous st ages of gonadal
development in S. barracuda.
Gonads were preserved i n Bouin’s fl ui d and subsequently st ained wit h
toluidine blue and picric acid. Photographs of histological sections of a ri pe
male (stage 3 ) and stages 1 , 3 , and 4 of a female are presented as Figure 9.
Age and Size at Mat urit y.— 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 o f two years. Some female s first ma ture at three years at a length
o f about 580 mm; most do not mature until an age o f four years at a
length of about 660 mm. Mal pas ( 1 926) found ri pe specimens of S.
commerson i ( = barracuda) from Ceylon which measured from 350 to

TABLE 4
Sex ratio o f 463 s p e c im e n s ofSphyraena 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.
Month Number Number of Number o f Per cen t Per cent Ratio,
examined female s male s female s 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_AUG.
-i=P-

-R = r i SEP.

--- 1 □ arfwlrj-T- OCT.

: nov.

0300 1100
FORK LENGTH , MM
F i g u r e 10. Relation between size and maturity o f Sphyraena barracuda . Speci-
mens were co llec ted from Pier 5, Miami, Florida , from August , 1955 , through
Augu st, 1956 . Block s above horizonta l axe s are male fish ; tho se below axe s are
f
ema el fi h
s . Shaded blocks represent fish ni spawning condition; open block s
represent non-spawning fi sh .
48
410 mm. If these specimens were indeed barracuda, they matured much
earlier than this species does in the western North Atlantic.
Sex Ra tio .— The sport fishery at Pier 5, Miami, from August, 1955,
through August , 1956 , yielded 287 female s and 176 males, a ra tio of
8 to 5 (Table 4) . Monthly fl uctuations i n t he sex rati o 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
t males, so ha
he t tthey are no t available to he t ang er
l . During het mon h t s
when the males outnumbered the females the ratio was lower than
when the reverse was true.
%
Spawning Per iod .— The percentage of individua ls 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 i d 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 of 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)
— —
— 66.7 — (1)
Mar. 33.3 (1) (2) (3)
May — — 100.0
(9)
—
— — 100.0 — (9)
June (4) (4)
July — — 89.5
(17 ) 10.5 (2) (19)
Aug. — — 89.8 (44) 10.2
(5) (49)
Total — 56 — — 102 — (7) (167
( ) (3) ( ) )
Female S age 1 2 3 4 R pe 5 Spen To a
s t ( i ) ( t) t l
Aug. 25.0 — 75.0 — —
(1) 15.4 19.2 (3) 23.1 — (4)
Sep. 42.3 11 (4) (5) (6) (26)
Oct. 75.0 ( (9)) --- — 8.3 16.7
(2) (12)
Nov. 95.2 (20) 4.8 — — (1) —
— (1) — — (21
17)
Dec 100 0 17 (18)
Jan.. 100..0 (18) — — — —
( )
Feb. 84.6 ((11)) 15.4 (2)
— — — (13)
Mar 60.7 (17) 28.6 (8) 10 7 3 — — (28
Apr . 64 3 9 71 21.5 (3) 7.1 1 — 14)
May. 7.1 ( )
1 21.5 (13) .
42 8 (6) 21 5 ( 3) 71 1 (14)
June 6.5 (2) 32.3 ( )
10 .
22.6 (7) 19.3 (6) 19.3 (6) (31)
Ju y .
25.0 (4) 6.3 ( 1) 12 5 (2) 31.2 (5) .
25.0 (4) (16)
l
Aug 28.0 21( ) 6.7 (5) 16.0 12
( ) .
24.0 ( )
18 ( )
25 3 19 (75)
To a. ( )
— 141 .
— ( )
35 . (41)
— — (40) . (32)
— ( )
289
t l ( ) ( ) ( ) ( ) ( ) ( )
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-
t reache s 73 -74 °F (Bumpus, 1957 ) for the Miam i reg on
ure i (Fig . 11 ).
The first ri pe mal e barracuda, meas uring 1035 mm fork lengt h , wa s
observed on March 31, 1956. By May, 11 per cent of the males sampled
were ripe. In July and August a few mal e barracuda were taken whi ch
had recently ceased spawning (stage 5 ). By October, 64 .7 per cent
o f the males were still ri pe; by November, all mal es had completed t hei r
spawning activities. There appeared to be no peak time of spawning in
the males, as milt could be expres sed from t he testes of mat ure mal es
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 fi rst ri pe female was
taken on April 18, 1956, meas uring 1002 mm f ork lengt h, and was t he only
ripe female observed i n t he April sampl es. The l arger females appeared
to ripen earli er, as indicated in the data for May (Fig. 10). In May,
21.5 per cent of t he females were i n a spawni ng condition. By m i -d
October, 91. 7 per cent of t he females had completed t hei r s pawni ng.
Reproductive activities s topped toward t he end of October. Spawning ceased
when the wat er t emperat ure had dropped f rom a mean maxi mum s ummer
temperature of 83. 8° to 77 °F.

UJ
cr
3

cSr
UJ
o.
2
UJ
I—
cr
UJ
i

UJ

F i g u r e 11. Relat ion between spawning season of Sphyraena barracuda and

mean water temperatures adjacent to Miami, Florida . Solid line represents per-
centage o f males in spawning condition ( N = 2 0 3) ; broken line represents per-
centage o f f emales in spawning condition ( N = 2 8 4) . Dott ed line is monthly
mean water temperature at Fowey Rocks Lighthouse, southeast of Miami,
Florida . Temperature data from Bumpus (1 9 5 7 ).
50
 The peak o f 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 o f greater spawning
ac tivitie s are eviden tly re flec ted in the occurrence of peak s in the grow th
o f the young-of-the-year.
Postlarvae were taken from December through June in the Virgin
Island s area (Table 6 ) . Fowler (1 945 ) recorded specimen s 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 f ormer
da te is near ly a month earlier than the date when fis h of a si mil ar si ze
were collected from Florida waters.
Williams ( 1 956 ) found ripe individuals of S. commersoni ( ^barracuda )
in East African waters from September to May. Ommanney ( 1953 ) took
specimens of S. commerson i from the Seychelles Islands with nearly ripe
eggs in May, and ripe females occurred in November; ripe males were
taken in December.
Frequency and Ex ten t of Spawning.— Kisselevich ( 1 9 2 3 ) used the fre-
quency distribution o f ova diameters to ascertain the frequency and extent
o f spawning in clupeid fishes, and Walford ( 1 932 ) successfully utilized
this method for Sphyraena argentea. Hickling and Rutenberg ( 1936 )
postulated that the breeding cycle o f a fish could be predicted by an
analysis of the ova diameters, and that species having a sharply d em 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
o f a fish having an extended spawning period are characterized by a series
o f 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 D A N A EXPEDI TIONS FROM JU N E , 1920 , THROUGH M A Y ,
1922, i n t h e V i r g i n 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 ort h C a r o l in a a nd A ndr o s I sl a n d B aha m as
(
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 —
1111 aD
12 l 1 _ 3 1 1 __
Tota 6 11 28 20. 7 1
Mean length 8.4 9.1 8.8 75 8.3 7.0
51
 In order to determine first if the eggs in one part of the barracuda’s
ripe ovary were larger t han in anot her part, 559 ova di amet ers were
measured from the ovary of a ri pe female whi ch meas ured 1077 mm f ork
length (Fig. 12 A - D ). Ova showing varyi ng st ages of mat urit y appear to
be similar in size , regard le ss o f where they were taken , since the peak s
o f the polygons correspond in each succeeding section . Carbine (1 944 )
found tha t the middle par t of the ovary of the nor thern pike, Esox luc ius ,

OVA D IAMETERS, MM
F ig u r e 12. Frequency distribution o f 559 ova diameters from different sec tion s
o f a ripe ovary (stage 4 ) o f a 1077-mm specimen o f 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 o f frequencie s o f ova diameters from section s 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 l argert han about 0.5 mm i n diameter form a s eri es of
low peaks which are nots harpl y marked. This suggests an almost continu-
ous suppl y of gradually mat uring eggs which contribute to t he general
stock of eggs throughout t he spawni ng season. Walford ( 1 932) found
evidence that a single female of S. argentea spawned several times per
season. The extended spawni ng season postulated here for S. barracuda
is corroborated by an examination o f the gonads throughout the year
(Table 5 ) , and by the abundance o f barracuda less than 1 inch long
from April to November from southern Florida waters (Fig. 18 ). Erdman
( 1 956 ) noted the presence of young barracuda over the same period
in Puerto Rican waters.
Fecund ity .— Walford (1932 ) calculated the number o f eggs produced
by a single female of S. argentea . He was able to separate mature and
imma ture egg s in the ovary of S. argentea, but this distinction generally
is no t marked in barracuda (Fig. 12 ). However , in some large indiv idua ls
of barracuda, the differences among t he ova diameters were s uffi ci ent t o
permit a rough calculation of fecundity. The number of eggs i n a s ampl e
was counted, weighed, and t he number of eggs per gram calculated. A
barracuda 895 mm fork lengt h was estimated to contain about 560, 000
mature eggs, and one meas uring 1011 mm fork lengt h held about 670, 000
mature eggs. However, the assessment of fecundity in a species whi ch
s
pawn s severa l time s over an extended period seems to have litt le
biological value unless it could be determi ned 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 expres sed
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 i n diameter (Walford, 1932; Orton, 1955).
The modes o f the peaks shown in Figure 12 occur at approximately
0 .14 mm, 0. 30 mm, 0. 48 mm, and 0. 70 mm. Evidently, t he ovary oft his
individual either was not dead ripe, or some eggs had already been spawn-
ed, as most eggs depicted (Fig. 12) are s maller t han t he trans lucent ones.
Barnhart ( 1 9 2 7 ) and Ort on ( 1955) stat ed t hat t he egg of S. argentea
is . is s i l’s ii
pelagic Th wa also noted for Sphyraena (sphyraena?) by N ko k
( : ), t ( ) ( ) s
1954 4 0 1 bu Delsman 19 3 0 believed Sphyraena species? ha
. . s s i i t t t t t
uch big egg n the r pe ovary ha one may conclude a once ha
t s t
he egg are nott pelagic.”ts t s tifi ll
Numerous
t a emp
s f wereis made
( )o fertilize and hatch( eee ),ar tca lly
using he method o Morr 1 954 and Oppenheimer 1 9 5 5 bu a
s i .
experiment fa led
53
 Distribution of the Postlarvae i n Rel ation to Hydrographic and Meteoro-
logical Conditions.— Hydrographi c 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 y d r o g r a p h i c methods and collecting gear are

TABLE 7
H y dr o g r a p h ic a nd m e t e o r o l o g ic a l o b se r v a t io ns at s t a t io n s o c c u p ie d
wher e p o stl a r v al sp ec im e ns o f Sphyraena barracuda wer e c o ll ec t e d b y
DANA e x p e dit io ns to t he t r o p ic a l w e ster n A t l a n t ic O c ean from
1920 to 1922 . D ata ar e su m m a r iz e d f r o m S c h m id t (1 9 ).
29
SampleWater Wind
Station t depth, depth, Wea- fand
number Da e Latitude Longitude Hour meters meters ther orce
/ / ° ' ( )
848 6 / 6/ 20 18 °00 'N 6 4 0 4 1 ^ 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 6 4 0 4 8 ^ 2300 200 (4800 ) squally ESE 4-5 -
952 5 /13 21 17 ° 55 ' N 6 4 0 4 8 ^ 0200 400 4800 squally ESE 4 5
1186X I 12 / 1/21
/ 17 ° 59 'N 6 4 0 4 1 ^ 0045 50 3160 cloudy N E 3
1190VII
I 12 /13 /21 17 °59 N 6 4 0 4 5 ^ 0400 100 3160 cloudy calm
1190IIIX 12 /13 /21 17 °59™ ' 6 4 0 4 5 ^ 0400 50 ( 3160 ) cloudy calm
1191 I 12/ 14/ 21 17° 49' N 6 4 0 5 4 ^ 1850 100 ( 1000) cloudy ENE 3
1191VIII 12/ 15/ 21 17 49 N 6 4 0 5 4 ^ 0400 100 1000 cloudy EN E 3
1192 V 12 15 /21 17043 ° ™ ' 6 4 0 5 3 ^ 1935 100 410 squally SE
1195VII 1// 3 /22 19 °01 'N 6 5 0 2 3 ^ 0350 50 ( 5250 ) cloudy N N E 4
1216V 1/ 29 /22 18 22 N 7 8 0 3 8 ^ 0100 0 (2000 ) cloudy N E 3
1256V 3 / 3 /22 17043 ° '™ 6 4 0 5 6 ^ 1920 100 (1000 ) cloudy ESE 6
1257V I 3 / 6 / 22 17° 43 'N 6 4 0 5 6 ^ 0010 100 1000 cloudy E 4
1266bI 3 / 1 3 / 2 2 17 ° 45 ' N 6 4 0 5 5 ^ 1900 800 950 cloudy N E 4
1266bV II 3 / 13 / 22 17 °45 ' N 6 4 0 5 6 ^ 1900 100 950 cloudy N E 4
1266VIII 3 / 13 / 22 17 ° 45 'N 6 4 0 5 6 ^ 2215 50 950 cloudy N E 4
1266VI 3 /13 /22 17 45 'N 6 4 0 5 5 ^ 2215 25 ( 950 ) cloudy N E 4
1270 V 3 16 22 17043 N 6 4 0 5 6 ^ 1800 600 1000 squally SE 3
1238 IV 2 /1 1 /2 2 26 1 3 'N 7 8 0 4 8 ^ 0400 0 ( 620 ) cloudy S 2
1273 V 3 / 24 /22 17043 'N 6 4 0 5 6 ^ 1915 300 ( 1000 ) cloudy ENE 4
1273V 3 / 2 4 / 2 2 17043 'N 6 4 0 5 6 ^ 1915 100 ( 1000 ) cloudy ENE 4
1278V 3 /29 /22 17043 'N 6 4 0 5 6 ^ 1900 100 (1000 ) cloudy ENE 5
1280V 3/ 31 /22 17043 ° 'N 6 4°0 5 6' ^ 2015 100 (1000 ) cloudy ENE 4
1281XI 4 / 1/ 22 17° 43' N 64I 56 I W 2330 50 ( 1000 ) clear E 1
1284V 4 / 6/ 22 14 ° 38 ' N 6 0 6W 0300 0 ( 800 ) cloudy ENE 2
1286V 4 / 7/ 22 15 17 'N 6 1 0 2 9 ^ 2100 50 ( 200) squally ENE 3
1289X1I 4 /1 5 /2 2 17043 'N 6 4 0 5 6 ^ 0400 50 (1000 ) squally E 3
1292V 4 1 6 2 2 17043 N 6 4 0 5 6 ^ 1930 50 1000 cloudy ESE 3
1293VI 4 /1 7 / 2 2 17043 'N 6 4 0 5 6 ^ 1940 100 ( 1000 ) cloudy E 3
/ / ' ( )
1293V 4 /1 7 /2 2 17o43 N 6 4 0 5 6 ^ 1940 50 (1000 ) cloudy E 3
13521 5 2 1 2 2 35042™ 7 3 0 4 3 ^ 2130 300 5000 cloudy SW 4
1352IV 5 /2 1 /2 2 35 ° 42 'N 7 5 0 4 3 ^ 2130 100 ( 5000 ) cloudy SW 4
1352V 5 /21 /22 35042™ 7 5 0 4 3 ^ 2130 50 ( 50 0 0 ) cloudy SW 4
a Dep h given in parentheses were estimated from U . S. Hydrographic Office
t s
Chart 1290 and Na tiona l Geographic Society “Map o f the West Indies,” 1954.
54
 TABLE 7— (C O N T IN U E D )

Sea Surface Number Other

Station and Temp., salin- Densi- 0 2 of specie s o f
number force °C . ity, %0 ty, o-t m l/ L specimen s 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 —
1190VIII E 3 — — — — 1 —
1190IX E 3 — — — — 2 —
1191III EN E 3 — — — — 1 —
1191VTI EN E 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 pi cud illa ( 1 )
1257V E 3 — — — — 1 pi cudilla ( 6 )
1266bII SE swell 25 .14 35 .84 23 .95 5 .24 2 —
1266bV SE 25 14 35 84 23 95 5 24 2 p cudilla 2
1266VII SE 25 .14 35 .84 23 .95 5 .24 2 i — ( )
1266V SE 25 .14 35. 84 23 .95 5 .24 2 —
1270 VIII SE 3 —. —. —. —
. 2 p cudilla 2
1238 IV SE 2 23 60 — — — 1 i — ( )
1273 IV ENE 3 — . — — — 1 —
1273VI EN E 3 — — — — 2 p cud a 1
1278V ENE 4 24 70 — — — 4 i
picud illa (3 )
. guachancho
ill (1 )
1280V ENE 3 24 80 — — — 6 guachancho (1 )
1281X E 1 24 .90 — — — 11 picud a (2 )
. ill
guachancho (1 )
1284V NE swell 26 00 — — — 2 guachancho (1 )
1286VI E swell 26 .10 — — — 1 p cud a (1 )
1289X1 E 3 25 .30 — — — 1 i —
ill ( )
1292V S swell 25 .40 — — — 2 —
1293VI S swel 25 .40 — — — 1 —
1293V S swell 25 .40 — — — 5 —
13521 I SE swelll 25 .60 36 08 23 99 — 3 —
1352 V SE swell 25. 60 36.08. 23. 99 — 1 —
I
1352V SE swell 25.60 36.08 23 .99 — 1 —
. .

described by Schm d 1929 and Jesper en and Taning 1 9 3 4 All

tow were made a ia tvessel ( )
speed o 2 knots s and each la ted( two ). hour
withs he exceptiont o a 30-minutef hau a , S a ion 1271s a 90-minute s,
hau at Sta ion 1191 and f a 180-minute hau l at Sta t t ion 1186 , A pecimen
werel takent w h a 200 , cm conical ram nl net t except
t or .those
ll s collecteds
a Stat ion 1284it V 1292 - V and , st1293i V a, which f150-cm nets were
t
opera ted sand a -StaI, ion 1270-1V
- I, a wh ch - aI, 150-cm
t k net wa opera ed
t , mum dep
The max t ht a which he, ne t i hed given in sil Table 7,s bu since
t
i t t t t fis is
he nets were no o he closing type he ac ua depth o cap ure o he t
t t f t , t t l f t f t
55
postlarvae was not ascertai ned. The DANA vessels operat ed in t he
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
Isl ands, but a few were collectedf romt he wat ers off Jamai ca, Mart inique,
Dominica, Andros Isl and, Bahamas, and Cape Hat teras, North Carolina.
When the frequency o f postlarval specimens o f barracuda taken by the
nets was plotted, two distinct peaks of abundance occur (Fig. 13): one
at 4 AM, t he second at 7 PM. This would at fi rst suggest a di urnal
mi grati on. However, when the tot al number of st ations occupied is
compared with the distribution o f collections in which the barracuda occur-
red, it is apparent that t he great est number of barracuda was collected at
the hours when most collecting was undertaken. Even so, the almost
complete absence of barracuda in collections made duri ng daylight hours
does not preclude the occurrence of a limited vertical migration. But barra-
cuda of all sizes are agile and speedy s wimmers, and perhaps they are
able to dodge the approaching net in the daylight hours.
A s might be expected in the open-ocean environment, only slight
dif ference s in sa linity and tempera ture were noted at the DANA stations
(Table 7 ); the surface salinity values were between 34 .98 and 36.08%c.

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

h i
K z
s 8

HOURS
F ig u r e 13. Relat ion between hours at which postlarval specimens of Sphyraena
barracuda were collec ted by plankton net s (dashed li ne) and hours at which
plankton nets were operated (so lid line ) by D A N A vessels, 1920- 1922, in the
tropical western A tlantic Ocean. Da ta are from Schmidt (1 9 2 9 ).
56
Surface tempera tures ranged from 23 .60 to 26 .80°C , and were typica l
of the trop ical wes tern A tlantic. Dissolved oxygen values in surface
wa ters from the localities a t which barracuda were collected, although
limited in the number of observa tions , were al l high.
Informa tion on the depths of wa ter a t the DANA sta tions is supple-
men ted by U.S . Hydrograph ic Char t 1290 , and the Na tiona l Geograph ic
Society’s Map of the Wes t Indies . All pos tlarva l barracuda were taken over
depths greater t han 200 meters, and the max imum dep th was 5250 me ters .
I collected postl arva l barracuda measuring 11.9 to 17.9 mm near
Bimini , Bahamas , re latively far from shore . These were taken a t the
surface by d ipnet and p lank ton tows over depths of nearly 200 me ters , a t
a local ity where a current ri p exists between t he litt oral wat ers and the
F lori da Current. Postl arval barracuda have also been t aken by me from
shallow-wa ter hab ita ts . I t is uncer ta in if these were spawned near the
location of their cap ture or had been transported there by curren ts and
t
ides. Weber (1913 : 150-152) collected a 13-mm arva l of S phyraena
(spec ies?) and four specimens of S. je llo which measured 47 to 55 mm in
l
ength , far offshore in the Ha mahe
l i Sea, Du ch
ra t Eas t Indies . Beebe and
Tee-Van (1 928) found specimens of barracuda of 20-26 mm in length ,
from March to May , which were collected severa l miles from Ha iti. On
these bases , barracuda probab ly spawn over deep wa ter near the junc ture
between the ocean ic and coas ta l c ircu la tion , so tha t the eggs and larvae
may be subsequen tly en tra ined in bo th coas ta l and offshore wa ters .
In two of the DANA Coll ect ion hauls, t hree sphyraeni ds from t he
wes tern Nor th A tlan tic occurred toge ther ; in these , p icu d illa and guachan -
cho were taken with barracu da , not wit hst and i ng t ha t t he adults of these
three species have considerabl y di fferent habit at s and verti cal id ts ribu t ions
.
t tl
Possibly he pos arvae of each species occurred in a d fferen s ra um i t t t
or density l ayer in t he wat er column and all were brought t oget her as
the ne t was re trieved .
L e ng t h - w e ig h t R e l a t io n

Specimens from bot h Fl ori da and Bimini t aken over a 17-mont h peri od
were utilized in t he calcul ati on of t he l engt h-weight rel ati on. Possible
rac ial and seasonal differences between t he barracuda of t hese areas might
be i ncluded in t he calculat ions. Usi ng t he lengths and weights of 59
females , 37 males, and 54 i ndivi dual s of undet ermi ned sex, t he empi rical
re la tion for t he t wo measurement s are present ed in Fi gure 14. From these
da ta , the lengt h-weight rel ati on for t he male i s W = 1.56 X 10~6 L 2-84589,
and for t he female i s W = 2. 67 X 10~6 L 2-92134 . The relatively small
t i ti
exponents observed here are charac er s c of a slender species such as the
barracuda , which con trast with t hat of a chunkv, heavy-bodi ed fi sh such
as the little tuna , E u t h yn n u s all ett erat u s, in which the exponent i s 3. 16
(de Sylva and Ra th jen , 1961) .
The calcula ted regress ion lines are approx ima te ly equal, excep t a t abou t
57
 TOTAL LENGTH, INCHES

FORK L E N G T H , MM

F i g u r e 14. Length-weigh t relation o f 150 specimens o f Sphyraena barracuda

from Florida and Bimini, Bahamas, collec ted from April, 1955, through August,
1956 . Open circles and solid line represent male fi sh (N = 3 7 ) ; solid dots and
broken line represent female fish ( N = 5 9 ) ; triangles represent 54 fish o f un -
determined sex, and for which the equation has no t been calculated nor the
slope fitted to the points.
58
 900 mm when the curves intercept. Males are heavi er t han the females at
the same length. The emp irica l da ta fit closely t he cal cul at ed regressions
for smaller fish, although poi nts representi ng fi sh great er t han 600 mm are
more sca ttered abou t the slopes.
The following mean conversion fac tors have been ca lcu la ted from the
length of over 200 barracuda which measured from 17 to 1111 mm
s tandard length from the western Nor th A tlan tic:
T o O b t a in : M u lti p l y by: T o O b t a in : M u lti p l y by:
S tandard from fork 0.919 Fork from to ta l 0.901
S tandard from t ot al 0.829 Tot al from st andard 1.207
Fork from s tandard 1.089 To tal from fork 1.110

A g e a nd G r o wth
With the except ion of Wal ford’s (1 932) study on t he age of S. argent ea
pub li shed dat a are not available on age assessment of any of the species
of Sphyraen idae . Cockerell (1914) used his bri ef study and descri ption
of the scales of S phyraena p i cu d a ( = S. b a rra cu d a ) to draw conclusions
on the phy logene tic affinities of the Percesoces.
C au se o f A n n u l u s F orm a tion in B arracuda . — The lit erature on the use of
scales and ot her hard part s for age det ermi nati on of fishes i s profuse. Most
references given by Graham (1929b) and Lagler (1 956) deal with t em-
perat e forms, where t he t heory is usually held t hat slackened body growth
due t o decreasing wat er t emperat ure i s reflected in a correspondi ng de-
crease in the growt h rat e of t he scale, as reflected in the relative spacing of
the circuli . In trop ica l and sub trop ica l species, a marked slowing of growth
generally does no t occur as a resu lt of the smaller changes in wa ter tem-
perat ure. Menon (1 953) believed t hat t emperat ure change was not
necessary for annul us formati on in the scales of tropi cal fishes, bu t t ha t the
period ic rhy thm of spawning seemed to be sufficient p e r se to cause for-
mati on of spawning marks on the scale. Chevey (1 933) no ted t ha t a
tempera ture difference migh t cause an annulus in trop ica l species, although
the slowing of growth of the scale wou ld be less pronounced than in
temperat e species. He concluded t hat a t emperat ure difference of at l east
4 to 5°C seemed necessary to cause growth slowing and a resul ting annual
check in the scale growth of trop ica l fishes. The barracuda is considered
a trop ica l species (Ekman , 1953 : 4 6 -4 7 ) , and although there are seldom
sudden drops in wa ter tempera ture during w in ter mon ths in southern
F lorida , the da ta of Bumpus (1957) show a mean annua l tempera ture
change of 6.5 °C for the surface wa ters sou theas t of Miami, F lor ida . Thus ,
if the assump tions of Chevey (1 933) are applicab le to the barracuda , the
annua l tempera ture change in the wa ters of sou thern F lor ida wou ld be
adequa te to cause annulus forma tion in the scales of the barracuda .
M e t h o d s o f Sca le C o llec tion and A n a l ys i s . — Scales were collected from
barracuda caught off Miam i and brough t in by sport fishermen to P ier 5,
59
F i g u r e 15. Monthly increment from last annulus to anterolateral margin o f
scale based on measurements o f 316 specimens of Sphyraena barracuda of
age-classe s I through V I from southern Florida . Circles represent mean values
for combined age-classe s for each month. Da shed lines are mean ra te o f deposi -
tion o f scale increment. Bracket A indicates probable resorption at scale margin
prior to annulus formation ; bracket B represents probable period o f annulus
f
ormation .

Miami. Smaller fish which generally do not ent er the anglers’ catch were
collected from the wat ers south of Miami, using hook and line, seine, and
ro tenone . A large number of scales was collected and lengths recorded by
t dermis
ax i ts Joseph T. Reese of For t Lauderda le and t he l at e A1 Pflueger
of Miami, Fl ori da. A series of scales was collected from t he left side of
each barracuda , midway between the l at eral line and the base of t he first
dorsa l fin. Thi s area has an unusually high number of regenerat ed scales,
bu t, fort unat el y, enough were collected so t hat perfectly formed scales were
usually ob ta ined .
Because thickness and opacity prec lude read ing the scale itself, p las tic
impress ions were used (G a thm an and Dawson , 1948) . Bu tler and Smith
(1953) concluded tha t a lthough there may be significan t differences in
s ize between the orig ina l scale and its impress ion , errors in compu ting
back-ca lcu la ted body lengths are w ithin the range of errors ob ta ined in
measur ing the fish and its scales. Emp ir ica l ages of 555 fish were assigned
by scale read ing irrespective of the fish length , sex , or date of cap ture .
All scale read ings were made using a pro jec tion device (Phil lips and Web-
ster , 1959) . Agreemen t between first and second read ings on a sample
of 100 scales was over 95 per cent . Using this me thod , annulus forma tion
a t the pos tero la tera l marg in of the scale may be identified (Figs . 16 and
17).
The age analyses were poo led for 488 indiv idua ls collected over a
60
period of 13 months. Ages were recorded as the number of wi nt ers com-
pleted using Roman numera ls , as is customary, t o designate t he number of
annuli presen t. The las t day of w in ter was arb itrar ily regarded as March 1 .
Scales in which the annulus was omitted were consi dered as having the
annulus if ad jacen t annu li of ot her barracuda belongi ng t o t he same si ze
group had the annulus presen t. These scales were not used for growth
calcula tions .
V a li d it y of t he A n n ul u s as a Y e a r-m a rk. — Lagl er (19 5 6 ) present ed cri -
ter ia for the es tab lishmen t of a valid annul ar mark. These assumpt ions
necessitate t hat only one mark i s deposited per year and at about t he same
same time every year.
To obt ai n the approxi ma te time of annu lus forma tion, the i ncrement
from the las t annu lus to the an tero la tera l marg in of the scale was summed
for fish belonging to age groups I through VI , and the mean ob ta ined for
each mon th . The trends of the mean month ly incremen ts were fitted by
eye in F igure 15. Annu lus forma tion occurs some time between January
and March ; from December through January there appears to be a slight
resorp tion of the scale marg in .
The time of annulus forma tion based upon scale incremen t is corrobor-
a ted by an exam ina tion of the scales from 10 young barracuda collected
from November to June (Tab le 8 ) . Young specimens collected in No-
vember and early December showed no annulus , bu t a check in the
growth of the scale appeared to have occurred recently in f ish collected
on December 31 and on January 4. However , an indiv idua l collected on
January 7 showed no such mark . No young were availab le during February ,
bu t two specimens collected on March 19 had well-defined annuli , with
numerous add itiona l circul i laid down pas t annulus forma tion . Observa-
t
tions on samples of young and yearlings collected from March o Novem-
l t t l i
ber disclosed no annuli supp emen ary to hose a d down between January
.
and March
l ti i i l
The re a on between age and s ze of barracuda based upon emp rica
i i . i t t
scale analysis l s given i inl F gure , 19 F sh assigned l i o het samet age t group
in genera t have , sim lar
l engths t , although over ap s grea due i . he slow
o
growth ra e differentia grow i t h and
t the prolonged l spawning
t per
l odl t Modes
in the length-frequericy
, d i s ribu ions
t agree wel with l he ca
t cu a ed and
observed ages t t although
t t f sh less
i t han t two . years tand o der han 10 years
are
t difficul
i o de ec tin the d s ribu ions t For he youngert age groups
here s good agreemen among . calcula
l ed growths of differen aget groups
of the t,same year’s , collections
t, Ca
, culations . based iti
upon, samples aken in
August 1955 and Aueus l 1956 agree well In tadd ton young-of-the-year i i i t
tdo no showl annuli l t (Tab e 8) beforel the
. leng h of he f sh nd ca ed by
he back-ca cu a ed growth (Tab e 11)
it i t t t ti l . it i
C r erl a fo rt hei Inti erp re a o n o f A n n u a M a rk s —, The cr er ta used for ,
annu us de erm na on in fishes by Creaser (1 9 2 6 ) Van Oos en (1921
61
 ,
'

FIiG
F gUuR
rEe 1166.. IIm
mppressions of scales from specíimens of Sphyr,aena barracuda
ccollected frrom sou therm n Florid a. A . Fema le, 7 56 mm fork length, shhowing four
annuli (x x77.66));; collected April 5, 1199556. B . Male, 9 1111 mm fork length, shhoow
wing
eight annuli (x6..99));; collected Decemb err 22, 1199555.
62
 l-liGgUuRr Ee 1177.. Ilmpression s of scales from specimens of S phyra ena barracu da col·-
F
lleeccted from southern Florida. A. S ex undetermiin need, 1122445 mm fork length,
s�howing n
rninnee aannnnuulli (x
x66.2));; collected March 1l., 1199556. B . S ex und etermined,
1122559
9 mm fork length, shho win g 1144 annnnu
uli (x
x77.00)); collected August 1122,, 1199556.
63
1929) , Beckman (1 9 4 3 ) , Kest even (1 9 5 0 ), and Lagl er (1956) have
been used here.
False A nnuli.— Sprugel (1954) summari zed the possible causes of forma-
tion of false annuli. These were comparati vel y common in barracuda, and
of the 515 scales aged, 25.8 per cent had from one t o four false annuli, and
nearly t hree-fourt hs had one false annul us (Tabl e 9 ). Fal se annuli seemed
to be most preval ent i n t he younger fish. Tabl e 9 shows t hat 46. 7 per
cen t of the false annuli occurred in fi sh under two years, 75. 7 per cent of
the fish were younger t han four years, and no fish showed false annuli aft er
he
t eighth year of life . Because it was considered tha t the di st ri buti on of
he
t false annuli migh t merely reflect the abundance of the respective age
groups, t he frequency of t he observed ages in rel ati on t o the frequency of
false annu li was compared. There i s a di sti nct preponderance of young
fish wit h false annuli, whereas t he age-frequency di st ri buti on i s composed
essentially of fish o lder t han two years of age.
“S k i p p ed ” A nnuli.— Sigler (1 949) not ed t hat some white bass, Roccus
chrysops, fail ed t o form an annulus at the expect ed time. Thi s was found
in barracuda also (Tabl e 9 ). Slowed growt h due t o a l ack of food does
no t explain this phenomenon in t he barracuda, which has an abundance of
food availab le at all times, and food i nt ake t hroughout t he year shows
l
ittle var ai tion (F gi . 3 3 ) .

TABLE 8
R e sul ts of t he e x a m in a t io n of sc ales of 10 y o u n g - o f - t h e -y e a
r sp ec im e ns o f Sphyraena barracuda F l o r id a
from so u t h e r n to
ASCERTAIN PROBABLE PERIOD OF ANNU LU S FORMATI ON.

Collection date Fork length (mm) Description o f scale

,
November 15 1955 161 N o annulus formed
December 2, 1955 163 N o annulus formed
December 15, 1954 153 N o annulus formed
December 31, 1949 175 Annulus just being formed; crossing-
over along posterolateral margin o f
scale
December 31, 1949 172 Annul us 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 N o annulus formed
March 19, 1952 164 Annulus formed, with f our circuli l aid
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 g e d is t r ib u t io n in r e l a t io n to the num b er of f alse a nnuli a nd “ sk ip p e d”
a n n u l i o b se r v e d in the sc a les o f 519 s p e c im e ns Sphyraena barracuda
of
c ollec ted f r o m so ut h e r n F l o r id a from A ug ust , 1955, t h r o u g h A u g u s t , 1956 .
No. Per cent
Age Number o f Per cent of Number o f Per cent o f fish fish
class false annuli false annuli “skipped” annuli “skipped” annuli examined examined
O 22 13.0 16 34 .0 8 1.5
57 33 7 10 21 3 19 3.7
I 27 16.0
. 9 19.2
. 38 7.3
II 22 13.0 7 14.9 53 10.2
IV
III 16 9.5 2 4.2 78 15.0
V 14 8.3 — — 67 12.9
VI 8 47 2 4.2 67 12.9
VI 3 1. 8 1 22 66 12.7
VIII — —. — —. 47 9.1
IX I — — — — 31 6.0
X — — — — 18 3.5
X — — — 13 2.5
XII
I — — — — 8 1.5
XII — — — — 4 0.8
X VI — — — — 2 04
Tota
I 169 100.0 47 100.0 519 100.0
.
l

Grow h of the Young — Young barracuda were commonly observed in he

shallow t wa ers of he .F or da Keys from May un November af er which t
ime hey move t nto
t somewha
l i deeper wa er where til hey become , t difficult
to
t collect.
t Seine icollections were t made ast often as possib t e each mon h
bu collecting dur ng he w n er months yielded few smal l barracuda t ,
Despt te he r observed i tabundance
i t barracuda are difficul o l collect with.
seinesi because
t i of he r wariness , t t
From field and tmuseum i collections,
. 244 young barracuda were measur-
ed; these represen collections made from 1945 hrough 1956 (F g 18)
Var a ons in growt h from year to year may be obscured t by he groupings
i . .
of ali tida a under at single mon h’s growth t
l tApr o Oc ober young
From t barracuda . grow o an average eng h of
abou 4 inches il tin s andard
t , length with a range of abou t IV2 o 11 l inches
t
The tvar a on in he t observed grow , h of he young appears t t o be due.
essentiallyi tito prolongedt spawning hroughou t t he season F ort example,
he mean grow h ra e (Fig 18) ncreases t t t
steadily from . Apr hrough
tJuly hen dropst o a tvalue only . i
slightly h gher han he mean value il t during
May, t Th s appears t due pr mar y o hei occurrence t t of relatively large
.
numbers iof very small, recently i ilha tched t fish
The var a ons in length among t he young. in a given mon h are con-
siderable iand ti means are smoo hedt by a runn ng average of t hrees pre-
sented by, a dashed line The relatively t i
abrup slackening in the tgrowth, ra e
. t t
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 o f 244 young-of-the -year
F ig u r e
specimen s o f Sphyraena barracuda co llec ted from southern Florida from 1945
through 1956 . Da shed line represents mean growth rate obtained by moving
average o f threes. Number in parentheses represents number in each mon th ’s
samples. Horizon tal bar represents mean; vertical line represents range o f varia-
tion; longer rectangle represents one standard deviation on either side o f the
mean; shorter rectangle represent s two standard error s on either side o f the
mean.

of the young fish no ted by Walford (1932 : 28-30) in Sphyraena argen tea
was no t evidenced in S. barracuda .
Those specimens of barracuda hat ched l at er in the season grew at a
somewha t slower rat e. Presumably, this reflects a decrease in abundance
of arva and pos arva fishes, on which young specimens of barracuda
lfeed as he season progresses Bu generally here s ittle change in the
seasona l grow h ratle of lyoung specimens of barracuda on he basis of da a
, edt in F gure 18
presen . t, , t i l
l t t t t
Emp r tca Grow i h R a e. n A d u s — Adu s as used in the arb rary sense
here represen barracuda more han 12 months old. Emp r ca da a are
basedi iupon lt .
l scalet read tngsi of barracuda lt
collected a P er 5 Miami, it F or da
t t
from Augus 1955 hrough Augus 1956 (Fig 19) Of 488 specimens i i l t
151 were males and 222 were females; the sex of he rema n ng 115 iwas,
i t i , l
no determ ned t, , t t, . . ,
The emp rica growth ra e of males, females, andt specimens i i whose sex
wast no de ierm . ned are shown in F gure 19 using he me hod of Hubbs and
i l t
66 t t i i t t
Hubbs (1 9 5 3 ). Dat a for fi sh ol der t han nine years are few, but in those
less t han nine years t here appears t o be no significant di fference i n the
growth rate between the sexes. The only depart ure in t he growt h rat e
between the sexes is for cl ass-III fish; however, a t -t est i ndicates t hat the
difference is no t significan t for tha t class ( t = 0 . 2 7 , P > 0 .5 0 ) .
The lengths of male and female barracuda assigned to age-class I are
no t represent ati ve of t he popul ati on. These fish are i mmat ure, and only
the l arger indivi duals have developed sexually t o a stage whereby t he sex
can be identified by fi eld met hods. Thus, males (410 mm) and females
(400 mm) of age-class I appear to have a great er mean l engt h t han the
specimens whose sex is unknown (364 m m ). When all specimens belong-
ing to age-class I are grouped, the mean fork l ength i s 378 mm, which i s
more represen ta tive of the popu la tion .
The barracuda is a relatively long-lived species; two individuals weigh-

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

1400.

1
1000-

8 00-

600.
I
f
400 .
M ALE S F E MA L ES COMBINED

1
200 -
0* T T~T I T
VI V II V III IX XI XII
AG E CLASS

F i g u r e 19. Empirical growth rate of Sphyraena barracuda from southern Flor -

ida , based on scale analysis of 488 specimens collect ed from August, 1955 ,
through August, 1956. Horizontal bar represents mean; vertical line represents
range o f variation; longer rectangle represents one s tandard deviation on either
side o f the mean; shorter rectangle represents two standard errors on either
side o f the mean. In each age-class, stippled rectangles are males, obliquely
ruled rectangles are females, and solid rectangles comb ine males, female s, plus
fish o f undetermined sex.
67
 14

·/
1200 .. ;··
. .r:
. .,:}. ,:;. · .
-, "t,t� •
."..'l: .
·.::;f .. ·
··�·;·,/
... :.¿, .
", ;�,·
1
80 ·.-
¡<",'.
• ::¡,'.('.'.
! '• ..
,:r-!' ..�:-sv , .
_""','
z

"'
o:
60 ·t' ¡..'·' .
¡ .
�
Í .'
200
I
.' ',.
''
'' '
'
o.¡.,.,�������������
o 100 200

ANTEROLATERAL SCALE RAOIUS. MM (,26.7)

FIiGgUuRr E
F e 20. R
Relation between anntterriorr scale rraaddiiuus iinn m
mm (x
x 266..77)) aannd fork

length of 29 5 sppeeccimens of S phyra ena barracuud a collected from southern Flor--

ida frrom April, 1199555, tthrough August, 1199556. Relation iiss exprressed bbyy tthe eqquuaa-
tion, y = 30.4 + + 4 . 02 x .

6688
 TABLE 10
L e ng t h o f sp e c im e ns of Sphyraena barracuda c ollec t ed f r o m so ut h e r n
F l o r id a at the t im e of sc a l e f o r m a t io n in ar ea b et w e e n b a se of
SP INOUS DORSAL F I N AND LATERAL L I NE .

Fork length ( mm ) Descrip tion o f area

28. 0 N o scales visible
30.0 N o scales visible
30.1 N o scales visible
30.2 Scale pocket s present but no scales visible
30. 4 N o scales visible
31. 0 Scale pockets present but no scales visible
31.8 Scales developed
32.9 Scales deve loped
35. 0 Scales well deve loped
38. 0 Scale s well developed

ing 26 and 39 pounds had 14 annuli. Since t he barracuda i s known t o reach

a weight of 106 pounds (Herre, 1 9 3 6: 1 0 0 ), it i s probabl e t hat it lives
for much longer than 14 years .
Ca l cu lat ed Grow t h R a t es.— Calcu lated growth rates more accurate ly depict
the leng th of the fish a t each year than do emp rica i l grow ht ra es
t because
of incremen t in grow t h of the scale beyondt hel as t annulus . Me t l hods of
back-ca cul ati ng growt h from scale analysis have been studi ed by Graham
(1 9 2 9 a) , Van Oos ten (1929 , 1953) , H ile (1936 , 1941) , Cohen (1 9 5 4 ) ,
Lag ler (1 9 5 6 ) , and o thers .
Body-sca l e R e la tionsh ip .— In calcula ting growth rates, it is assumed tha t
t
hroughou t life he
t scale grows a t he t same ra et as het fish . Van Oos en t
(1 929) made use of scale magnifications w hou rever ng o he ac ua l
it t ti t t t
scale measuremen ts in the back-ca lcu la tion of body grow th , and this
me thod has been followed in the presen t study.
I t appears t hat t he vari ance of the i ndependent vari abl e (body l engt h)
is corre la ted with t he dependent vari able (scal e l engt h). Alt hough this
var iance could be reduced by t he use of l ogari thmic t ransformati ons, Whit -
ney and Car lander (1 956) consi dered it sufficiently accurat e t o di sregard
the assump tion tha t the variances of the body lengths are the same for
each scale length.
The body-sca le re lati on for 295 barracuda coll ect ed i n 1955 and 1956
from the wa ters ad jacen t t o Miami, Fl orida, i s shown in Fi gure 20. It was
assumed here tha t the re la tion is cons tan t among the year classes w ithin
t he sample, and t hat t here was no difference between t he sexes. The
equa ti on y = 30. 4 -f 4. 02 x was obt ai ned for t he general model
y = a -f bx, with a correl ati on coefficient ( r) equal to + 0. 9 7; an r 2 value
of 0 . 94 suggests t hat l ineari ty account s for 94 per cent of t he vari ance i n
the fork engl ht (Y ) due ot scale eng l ht ( X ) . There si probab ly very
69
 TABLE 11
Su m m ar y of m ean c a l c ul a t e d f o r k l e ng t h s b a se d o n sc ale m e a sur e m e nt s of Sphyraena barracuda c ollec te d
from so u t h e r n F l o r id a from A ug ust, 1955, t h r o ug h A ug ust , 1956.
Observed MALES
No. mean Calculated fork lengths (mm) at end o f year o f 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
15 530 286 452
II 13 656 265 457 595
III
V 28 715 275 448 578 675
I
V 17 786 260 421 552 655 734
V 25 853 263 434 561 657 745 818
VI 18 905 267 444 569 674 758 831 892
VII 9 940 242 419 534 631 734 811 869 918
III
X 4 1040 261 438 567 706 811 891 942 1019 1050 — — — —
I
X 786 832 891 953 — — —
2 1060 271 394 494 608 665 729
X 3 1106 284 442 564 662 748 834 896 956 1008 1053 1089 — —
I
X 2 1260 336 476 572 676 783 867 948 1017 1017 1113 1060 1197 1131
III
To a 140
t l:
Ca cu a ed mean ork eng h 269 440 566 663 747 825 890 945 1013 1042 1077 1197 +b
l l t f l t : 78 65 55 68 29 35
Grow h ncremen 269 171 126 97 84 + +
t i t:
Ca cu a ed mean eng h Augu 404 525 629 712 789 864 913 973 1047 1057 1095 — +
l l t
Number o h l t , st: 140 136 121 108 80 63 38 20 11 7 5 2 3
f fis :
aOne ma e o age c a XIV wa no nc uded becau e ca e wa no aken rom he ame oca ion on he body a o her
bData n lu cf en - l ss s t i l s s l s tt f t s l t t s t s.
i s ffi i t.
 TABLE 11— (C O N T IN U E D )
Observed FEMALES
No. mean Calculated fork lengths (mm) at end o f year o f 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
13 510 260 418
II
23 607 263 427 561
III
V 27 728 261 431 562 677
I
V 25 785 269 436 556 659 738
V 27 848 277 432 556 659 749 824
I
V 25 941 278 428 569 687 777 855 913
II
V 22 970 282 440 556 663 755 833 900 947
III
X 15 1014 282 425 551 671 752 829 899 944 989 — — — —
I 637 741 821 884 951 1000 1038 — — —
X 9 1040 277 405 551
X 9 1094 259 429 559 670 760 838 907 963 1017 1067 1096 — —
I
XII 4 1140 244 409 545 654 739 814 873 922 982 1034 1027 1128 —
X 1 1160 262 447 555 620 686 755 821 882 948 1002 1037 1098 1133
III
To a 205
Catcul: a ed mean ork eng h 271 429 558 667 753 833 900 947 996 1047 1072 1122 +b
l l t f l t : 50
Grow h ncremen 271 158 129 109 86 80 67 47 49 51 25 +
Ca cuta ed i mean t:eng h Augu 409 508 623 722 796 873 934 971 1020 1073 1085 1147 +
l l
Number o t h l t , st: 205 199 186 163 136 111 84 60 38 23 14 5 1
f fis :
bDa a n u c en
t i s ffi i t
little, if any, curvil ineari ty (Fig. 2 0 ), and because of the rat her high
corre lati on coefficient and the relatively small vari ance t here seemed t o be
no need to test for linearity of regression .
The i ntercep t value of 30 .4 mm represen ts the fork leng th of the fish
when the scales were first deposited in the area of scale sampling. This
calcula ted observati on is in good agreement with empi rical observations
of young f ish . Scales are first deposited on the cauda l peduncle in young
barracuda a t abou t 25 mm fork length , and are la id down progressively
an terad along the la tera l- line region. Exam ina tion of 10 young barracuda
showed tha t scales were presen t between the first dorsa l fin and the la tera l
line, the loca tion a t which scales were taken from the adu lt fish , at
between 31 .0 and 31.8 mm fork length (Tab le 10) . A lthough no b arra-
cuda less than 200 mm fork length was used in compu ting the body-sca le
re la tion , emp irica l evidence on the approx ima te time of scale forma tion
would suggest t hat the rel ati on remains approxi mat el y li near even at t he
smalles t stages.
Ca l cul ati ons. — Fork lengths at t he end of each year t o annulus deposition
(approx ima te ly February) were back-cal cul at ed using Fraser’s (1920)
equati on:
ln = (L — C .F .) sn + C .F .,
S
where L = fork length of fish a t cap ture ,
C .F . = correc tion fac tor correspond ing to fork leng th a t which scales
were deposited in the area of samp ling ( = 30 .4 m m ) ,
S = length of scale at cap ture (x 26.7 in this s tudy) ,
sn = d is tance to any annulus from focus (x 2 6 .7 ) ,
ln = fork leng th of fish when annulus was deposited .
The number of annuli on each scale was recorded, and measurements
were made from the focus t o t he ant erol at eral margi n of t he scale in mm
( x 2 6 .7 ) to ob ta in scale scale l ength (S ), and from t he focus t o each
annulus (sn) along this same line. Since scales from t he same area of t he
barracuda show a cert ai n amount of variability in si ze and shape, from
t
hree to six scale impress ions were measured on each slide and t hei r
measurements averaged to reduce the vari ance about t he regressi on line
of the body-sca le re la tion (Fig . 2 0 ) .
The same scales were used for back-ca lcu la ted grow th compu ta tions
as were used for emp irica l observa tions .
Ca culations based on 140 males and 205 females from Miami, F or da
l
are summar zed in Tab e 11 Modera e y good agreemen among calcula l i ed,
lengths a each annulus suggests again ha these marks are yearly t in
i l . t l t
na ure D tfferences in ca cu a ed lengths a teach t age are remarkab y similar
for he sexes and are al within the expec ed range of error The lclass-VII
t . i l l t t
t
females (N =, 39) are calcula
l ed o have ta length 10 mm grea
. er han ha
t t t t t t
of the males (N = 38) and the class-IX males (N = 11) are 17 mm
72
 FO RK L E N G T H , MM
600 800 1000 1200 1400
' ------ 1 . . . ...... . ...........................I .,...
I
SER
JM L 1

OCT
n W ln J ln ,
.
l 1 f fl , NOV
—T —f^l—T ——1— — n nn■ 1
ii i
r y - Irn - n p r n n n n_____BE£_
J
AM
.
-O__ CLn n
f
a
.. FEB

nnnn
.r r- -, .
n— H nn n APR 1
MAY 1
JUNE
j Eh_
fm J
ULY
I -----
FLOR DA
....
BAHAMAS

i' i
n b M vHr

' ........ i | i | jll l| i

I(
""T""! " 11 11111rj i■ ■pi •■ ■• • 11•■■11■111
,
TTTII » »

0 200 400 600 800 1000 12200 1400 1600

. T O T A L L E Ns G fT H MM s f
. i t s l-
F i g u r e 21 Length-frequency measurement o 1091 specimen o Sphyraena
i ( ) ( ss l )
barracuda Histogramst enclosed , n solid
. lines represen
s monthly sample coi
lected n southern Florida
t N = 8 6 0 by seining le than 300 mm ong , i and,
from , angler catches
, a Pier 5 Miami Histogram , enclosed in dashed lines n
July sample represen 231 i barracuda collected from Bimini, Bahamas n July
l1955 and July 1956. Area sbetween broken t vertical lines represents
s range
generally not sampled n Florida due to gear selectivity. Back-calculated mean
engths based on scale analysi for Augus are indicated by arrow on length-
frequency distribution for August. 73
larger than the females (N = 2 9 ). The difference between t he males and
females of age-class X II i s considerabl y great er t han in any ot her age
classes, bu t a di screpancy in t he calculat ed growth rat e of t he males for
t class (N = 2) may be due to the small sample involved. The differ-
his
ences between the mean growth rat e of the sexes of the cl ass-VII group
was found to be non-significant (t = 0. 04, P >. 5 0 ); similarly, sexual
difference for class IX was no t significant ( t = 0 .49 , P > .5 0 ) .
Barracuda do not exhibit cont rasti ng regions of rapi d and slow growth
from summer to win ter, even t hough t here i s some evidence for this in the
length frequencies for O-class fish (see above under Grow th of the Y ou ng) .
Where data were adequa te , slight evidence for Lee’s phenomenon , or the
apparen t change calculat ed from different age groups (Van Oosten, 1929),
was evidenced in the back-cal cul at ed growths for the first fi ve year classes
of male barracuda bu t this was no t observed for the females.
Ca lcu lat ed Growt h Increment s. — The calculat ed growths of barracuda
from F lor ida are summarized in Tabl e 11. The i ncrement s between suc-
cessive years are approxi mat ely the same for the sexes. The great est i n-
cremen t i s during t he first year, when the calculat ed growth for males and
females is 269 mm and 271 mm , respectively .
Leng th- jrequency D is tr ibu tions. — The observed length frequencies of
young and adult barracuda collected from sout hern Fl ori da are shown i n
F igure 21. Young barracuda (i .e., t hose less t han 12 months ol d) were
mostly collected by seine from 1945 t hrough 1956; the adults were largely
from P ier-5 anglers’ catches.
In May, young are observed in t he 80-mm class i nterval. By t he follow-
, t t
ing March the young have become oo large o be collected by seining, and
t i , i i , i t
he r absence ndicated by vertical, broken lines in F gure 21 s cons rued
l
as being due to gear selectivity. C ass-I fish are observed even in the
i t i ti , t t t
small frequency d s r bu on for May where hey appear a he 2 80-mm
i l; i t t t i .
nterva age-class II s represen ed a he 460-mm class nterval
t t, , i t i t t
The tisample aken in Augus 1956 s the. bes est ma e of he relative t
propor on tof each age group in the sample A scale of t values represen -
ing calculat, ed i mean i fork lengths for each age, t correc ed ito give lengths
for Augus
. s super
t mposed above the leng
t l h-frequency
t d agram i t for t this
month Agreemen t between the peaks inl he eng h-frequency d s ribu ion
and the calculat ed lengths aret .remarkab y similar (age-classes IV through
IX ) wheret da a are adequa e , ,
Leng
i h frequencies
, , of 231
, barracuda
, collected in Bimini iBahamas t t
durt ng July
l i 1955 and July 1956 are superimposedi on the .d s ribu ion l
of he lF or da ispecimens for July as i dashed lines in F gure 21 it The re a-
tively
i arger s ze t of the liO-class f sh . from Bimini suggests e her more
rap d t growth i t i eti or ear er spawning
ra i . , i t
In the d s r bu on for Julyl (Fi g 2 1 ) a comoar t t son between t he barra-
cuda aken in Bimini and F or da indicates ha most of he individuals
74
 1953
VA-J---------- 1---------- 1---------- L ______ 1_______ 1___ _

JAN . L
J ______ n r. L

I n n n i— i ^ |

n i---
» P --- 1 r
H n h v j ' j " V . r t

SI
F
“

F
. j l F ' U „ " 1
i

O „
i
. n

R
E ^i —i i n rH rn - *'— P1 ----1 n
n J U N
n
E
I
’
B
M „
U i
, J V i , . - r l

^nnPs r
if -
^

A ______ OH__ CLD_r 1 1 n r rn n

- -. ....
« P

_a______ ___ o o _r n ___ a___ lzj __ □__

0 - i
1 n n n — H n — n n n NOV. F
0 fll i i
D E C .1
m— n M U n n n n
i ' i
L
r 1 1 1 1 1
i
0 10 20 30 40 50 60
T O T A L LENGTH , INCHES
F i g u r e 22. Length-frequency measurements o 501 specimens o Sphyraena
f
barracuda from taxidermist records from southern Florida taken f n 1953
’s i .
75
 1954
M ^ m i u f c U A i J U t 11111111111 h i

r—i fl n

JL J k a ^ r
-" t
x 0-
— 5 3 " " t
JD__£L
" 4
l t/
£ o
t
M __CL r-T T T K
r 3. rfil
uj 0 1 n n d nP
2CD 5
l
3 J

AUG.
JL . / ■ V
n n fl ^ l - T J U i n , ^ ! SEP~ |
0
J ------------ cm---- n ib ----- 11 r—'In-n - h
0
r A rJ l n P *l nn N0V' E
0

0 i/n n n HI r M “ S’ I
0 10 20 30 40 50 60
T O T A L LE N G TH , IN C H E S
23 Length-frequency measurements o 682 pecimen o Sphyraena
F ig u r e
barracuda .from axidermist records from southern
f s
Florida s f n 1954.
taken
t ’s i
76
 1955

JA N .

0
__n nJVT-A-jinfi
5 FE B .
i
J ---------mD—
J LLI n r> H
n l l nn l n» r "v u " ^^ 1 ^ 1 a K

0
j __ - - . 0 „n n lJ Rn
, MAR^
1<> 5M
77/ 0-1i j=D aJ L
°0 l CL -0 — n,. ^ J W M U n n n " ^
ll_ - l
o 1 p A p i»j x ^ — r
.
DC
ULY
m— a M
LlI
t. J
m n n k
r-i i I i
AUG
.
nrHnnn
. SE P
_______ rJ ■ nn ^V Un .
,] I - '-' l OCT
fin.
n n nn a— an.
1 .
NOV
0 \ nn n n n n n j i Ol

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

t ..

M iy n n mi mn ii m n
fi'i'i i r i i | 11 i |i i 8 i|ii ii i i i [ i i i i i i|i i
0 10 20 30 40 50 60
T O T A L L E N G T H , INCHES
24 Length-frequency measurements of 672 specimens of Sphyraena
F ig u r e
barracuda .from taxidermist’s records from southern Florida taken n 1955
exc us ve of Pier-5 samp es i ,
l i l .
77
 1956

ii
f i i J i 11111 >>11>>>a **ii Ia h iJLi j l ii In i,u u u l i i ii ii 111Ii i
5
0
jn-rx.
5
0
n_ ■n
5 p/UI MAR
0 .n...nn j=£L d±£2 £ a
APR .
5
n n n o nj y J W
i
xco t5 M AY
**■ 0 JQ____ O__ D J 1 Q I i
m =L

u.
0 o

JULY
c

m
5
0 £L nn J V N S n
5
AUG .
1 10
5
0 ax
SEP
5 .

0 J
D C H n
3-
-EL cQfl oN o
,
0 nnn n n 0C T
-

0 - EL
5 DEC
.
0 ^TT r 111111r p n n 111 J1 ......
i | ti ..
0 10 20 30 40 50 60
T O TA L L E N G T H NCHES
F ig u r e 25
, I
Length-frequency measurements of 852 spec mens of Sphyraena
barracuda .from taxidermist’s records from southern Florida
i taken n 1956
exc us ve of Pier-5 samp es i ,
l i l .
78
 TO T A L 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 ,
1 _ rtn ~ r' u ’a _ L
0
X
CO
1 MAY .
0
Li. 1 JUNE
U_ 0
1 JULY
UJ
1 —On

1
CD

-
u

12 H
I SEP
.
! !
t EESnZTCEEXX
II
II tlt s
I

0
1 CT F
0NOV„„
0 1 .
12 DEC
i ■■ .. r - 9
4//0 200r ... 400 600 800 1000 1200 1400
FORK L E N G T H MM
,
26 Length-frequency measurements of 2707 spec mens of Sphyraena
F ig u r e
barracuda .from taxidermist’s records from southern Florida
i taken n 1953-
1956 exc us ve of Pier-5 samp es Back-calculated mean lengths based ion sca e
, for
ana ysis l iAugust are indicated
l . by arrows l
l .
79
 of age-class I and many of those from age-class II are absen t from the
F l ori da length frequencies. What may be a dom inan t year class can be
followed in the d is tr ibu tion throughou t the year by re trac ing the moda l
growth from age-classes IX (1 947) and X II (1 9 4 4 ) . Age-class V also
appears to be par ticu lar ly abundan t in the Augus t samples, bu t it is no t
possible t o retrace its growth .
Tot al lengths, measured t o the nearest inch, of 2707 barracuda caught
between Pal m Beach and t he Fl ori da Keys (not including thosel anded ta
P ier 5) were made available by t he l at e Mr. A1 Pflueger. The dat a were
recorded by mon ths for 1953 through 1956 , and are shown in F igures
22 to 25 . The compos ite da ta for these years , added by months , are given
in F igure 26 .
A compar ison was made between calculated lengths and length-fre-
quency modes for the mon th of Augus t, based upon P ier-5 samples.
No ca lcul at ed lengths or modal values are available for O-class fish, and
ages o lder t han X I were not obt ai ned by back-calcul at ions. In most cases
he t moda l grow ht ra et si slightly h gher i t
han e ither he t observed or
ca cu a ed grow h ra es and he calcula ed grow h ra e is higher than
l l t t t , t t t t
tha t of he t observed . These d screpanci i may be due in par t ot he
es t pres-
ence of weak year classes in the d is tribu tions . Nevertheless , it appears tha t
the grow ht ra es t using the two different me hods t are in relatively good
agreemen t.
As a supp lemen tary me thod for tes ting the validity of check marks as
annua l marks , H ile (1941) stressed the impor tance of trac ing year classes
t
hrough severa l years on he t assump ion
t t t weak or strong year classes
ha
shou ld be apparen t over this per iod . In order to exam ine this hypo thes is ,
t month yl da at have been summed ot ob at ni he
he t annua l ot at l number
landed (F gi . 2 7 ) . Emp ri ca i l lengths a t each age are fitted to the graph
and each age class is traced back through succeeding years . If it can be
assumed tha t the mean ra te of growth was approx ima te ly equa l for tax i-
derm is t’s samples from 1953 through 1956 , and tha t this ra te is the same
t i t
in he P er-5 samples, he modes of each succeeding year class can be
t i t t
raced over a four-year per od when mean calcula ed leng hs are assigned
t t t t , , ,
o each year class. I would appear ha in 1944 1946 1947 and 1948
ti l ,
compara ve y strong year classes were produced whereas in 1945 and
l t tl . i
1949 poor y represen ed year classes apparen y occurred This s in
t t t i i . .
modera e agreemen with he P er-5 samples (F g 2 1 )

M or t a l it y a n d S u r v iv a l
i t i t l t t
R cker (1 948) reviewed he h s orica aspec of the developmen and
i t t ti t
n erpre a on of “ca ch curves” derived from a knowledge of the age
i t i ti it l ti . ll t
d s r bu on w hin a popu a on sample Rounsefe and Everhar (1953)
t t t i t t
presen ed assump ions which mus be satisfied pr or o the estima ion of
t t l t lit t . lt l t t
o a mor a y ra es A hough al of the assump ions may no be fulfilled,
80
 TOTAL LEN G TH , INCHES

C A L C U L A T E D AGE
F igu r e27. Succession of year-classes in length-frequency measurements of
2707 specimens of Sphyraena barracuda from taxidermist’s records from
southern Florida taken in 1953-1956, exclusive of Pier-5 samples. Oblique
lines connect success ive year c lasses in which each age c lass was spawned .
Back-calculated mean lengths based on scale analysis are indicated by arrows.
81
it seems feasible to a ttemp t to construct a rough ca tch curve for barracuda .
The catch curve of the P ier-5 barracuda is presen ted in terms of the
to tal age classes and the logarithmic transforma tion of the ir frequency in
the sport catch (Fig. 2 8 ). The ages are based here upon empi rical age
analysis, rat her t han on back-cal cul at ed rat es. As Ri cker (1948) poi nt ed
out, the ascending ( lef t) leg of the curve and dome (cen tra l por tion)
represen t age classes which are taken in less frequen t propor tions than are
older fish, whereas the descending (r igh t) leg of the curve represen ts
fish which are vul nerable to t he fishery. If t he descendi ng legi s a straight
line, a un iform ra te of surviva l is assumed . A concave descending leg
suggests a decrease in nat ural mort ality with age, as well as a popul ati on
which may be overfished ; a convex descending leg seems to indicate an

>-

AG E C L A S S
F ig u r e 28 . “Catch curve” based on age composi tion o f 488 specimen s o f
Sphyraena barracuda caught on hook and line from southern Florida from
August, 1955, through August, 1956.

increase in nat ural mort ality with age, and, conversely, an underexpl oit ed
popul ati on. Keepi ng in mind t hat all of the assumpt ions for mort alit y esti-
ma tes canno t be satisfied, the catch curve (Fig. 28) suggests t hat, since
the descending leg is convex, t he barracuda popul ati on suffers l ittle fishing
mor ta lit y; rat her, it would appear t hat the fishery i s actually being under-
explo ited. However, Ri cker (1948) not ed t hat it i s difficult t o distinguish
between a decrease in fishing pressure upon t he popul ati on, and eit her an
i
ncreas i mor tality with age, in which the more rapi dl y growing fi sh die
ng
off sooner, or incomplet e recruit ment appeari ng in the ol der age groups.
Such a decrease in fishing pressure may be reflected in t he age composi tion
of the barracuda due to t he general inabi lit y of t he angler t o cat ch these
large fish.
Survival estima tes were ob tained using the me thod ou tlined by Jackson
(1 9 3 9 ) . Consi dering fi sh of only age-class IV and ol der because these
fish are theoreticall y i ncompletely vul nerable, the t ot al survival of fi sh
between age-classes IV and X III is 81 per cent, or a mort alit y of 19 per
82
 TABLE 12
C a l c u l a t io n of m o r t a l it y a n d s u r v iv 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
of sc al es 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 i n e , f r o m P i e r 5, M i a m i , F l o r i d 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. 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 ( 1 9 ).
39
l
Annua mortality Per
s cent Per cent
between ages Ratio urvival mortality
I /
M 4 6 1 /4 7 8 96 4
II-III 4 2 7 /4 5 9 93 7
IIII-IV 3 7 6 /4 2 5 88 12
V-V
- I 3 0 3/ 3 7 4 81 19
V VII 238 /3 0 1 79 21
VI-V
II- I 1 7 0/ 236 72 28
VIII-IVII 1 12/ 168 67 33
VI - X 7 3/ 1 1 0 66 34
X XI 4 3 /7 1 61 39
XI-X II 2 6 /4 1 63 37
XII X III 1 3/ 2 4 54 46
XIII--X I 6 /1 1 55 45
X X V 2 4 50 50

t t t l t lit l .
cent due to bo h na ura and fishing mor at yt (Tab e 12) i Such a survival
ra e seems
i . exceedingly
lt ihigh, and implies t t that lthe fishery l s highly under-
explo ted A hough it , s tconceivable ti t ha o a l surviva for all age classes
approaches this value t he es tma ed surviva . between age-classes X II
and X III of 55 per cenl does no . tseem plausible t t The samples t i for
t younger
t t
fish arei corresponding y higher t I appears l i ha the mos t mpor
t an fact or
respons
t t bleit for this
t i apparen t.high survivai sti the neglec t of he assump i ion
ha
t recru men s i .constan An exam
t t na on of
i he descend ti ng leg of
he catch curve (F tg i 28) lindicates ; ha the po nts
l i represen t ng each
t i age
class are t somewha l , rregu ar this suggests a rep en shmen of cer a n age
groups a interva s which tends tol give thet appearance i of, at high survival. it t
If the stock annually was overexp l oited in he Miam areal he . recru men
from unfishedt areas wou d supply t t a fresh t stock iof alt ages i
t The ca ch curve , indicates t t ha fish i aret not explo tedl ito tiany , degree pr t orl
o age-class
t , IV and i ha the l ht ghes ra e of exp . o a on tor na ura
mort ality t occurs n fishes , o der han age-class
t tVI Since mos males are,
ma
t ure a ll age-class i II and females ma ure
t it t a age-classes t l III t and IV i
heoretica
t t i yt these i f sh thave hadt an oppor un yi to ispawn ai eas once i pr or ,
oit he r en rance nto the tspor t fishery.
i To ma nt a n a bas c spawn t ng stock
would thusi appear . ha here s no need for he managemen of barracuda
in the Miam area
L a nd ing s of B a r r a c uda
t i ll li t l
A few barracuda are it;aken commerc a yt by hand
l nes andt by ro
l ling,
with live or artificial ba some are caugh in hau seines, ramme nets
83
and runaround gill nets. The present -day commerci al cat ch i s t aken
essentially by anglers who sell t hei r fi sh t o dealers. The report s do not
include the many barracuda t aken for persona l use.
The U.S. Fish and Wildlife Service (1952-1957) repor ted the annua l
commerc ial ca tch of barracuda , all of which is taken from F lor ida wa ters .
Some are taken by angling in the Caro linas and along the nor thern Gu lf
of Mexico bu t are not given in the statis tical records. The commerc ial
catch for selected years from 1918 through 1956 i s given in Tabl e 13.
The barracuda is an impor tan t recrea tiona l species. A long the eastern
coast of F lor ida , Ellis (1 957) found tha t in the number of char terboa t
trips when fish were caught, barracuda were exceeded only by do lphin
( Coryphaena hi ppurus ) , l ittle t una ( Eut hynnus all ett erat us ), and king
mackere l ( Scom berom orus cavall a). Barracuda ranked seventh in numbers
of sport fish caught.
Since 1950, t he report ed commerc ial landings of barracuda have been

TABLE 13
Com m e r c ia l l a n d ing s o f Sphyraena barracuda f r o m F l o r i d a f o r s e l e c t e d
YEARS, 1918 -1956 , AND TOTAL FOOD F ISH , 1950 -1956 . D ATA FROM
U .S . D e p ar t m e nt of C o m m er c e (1 9 2- 0 1 9),
38 A n d e r s o n a
( - ), .n d.
P o wer 1 9 4 0 1 9 5 4 (a n d U
- S F).i s h a nd W il d l if e S er vi
ce
1952 1 9 5 7
f t l
, Average f f , , Average
Landings o price per To a s s
price per
barracuda, Value pound, ood ish Value pound,
Year pounds, dollars .
cents pounds3 dollar cent
.
1918 7775 329 4 .4 b — __
1923 ,
1600c 64 40 — — —
1927 ,
1000c 20 20 — — —
1929 16 000 560 3.5 — — —
1932 18 720d 749 4-0 — — —
1933 4245 f 180 4.2 — — —
1934 4245ef 180 4.2 — — —
1939 , 500 5 1.0 — — —
1940 , 200 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 50 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 70 64, 580, 484 8, 154, 828 .
12.6
1955 20 305 1015 51 66 635 464 7 979 452 12.0
1956 s 14,463 s, 723 , 5.0 l 66 780 403 7 356 316 11 1
i f . t
st l . t .
aExclude alewives,
t . scad menhaden sharks, adyfish -and freshwater catfish.
bNot ncluded or these years. eMonroe Coun y only
cWest coa on y fPalm Beach Coun y only
dDade Coun y only gExcluding fresh and saltwater catfish.
84
decreasing (Tab le 13; Fig. 2 9 ) , although landings of food fish is increased .
The relative ca tch of barracuda taken commerc ia lly by counties (Tab le
14) remained roughly the same until the summer of 1954, when t he catch
repor ted i n Broward Count y sharpl y decreased. The l andi ngs report ed in
Palm Beach were cur ta iled la te tha t year , and in Apr il, 1955, the landings
repor ted for Dade Count y abruptl y ceased. These decreases in report ed
landings can be traced t o a series of ou tbreaks of ich thyosarcotox ism
caused by barracuda which occurred along the sou theas tern coas t of
F lori da from 1954 to 1956 (Pae tro , 1956 ; de Sylva, 1956) . Because a
Dade Coun ty law which forbids the sale of barracuda has been rigidly
enforced since this period, dealers have report ed few or no l andings. No
such decrease was observed in the catch report ed from P inellas Coun ty on
t wes ern
he t coas t of F or
l da
i , where ch i hyosarcotox
t i eviden ly
sm t does no t
occur , and where the sale of barracuda is no t proh ib ited by law.
During semi-weekly visitst o iP er ,5 Miam i (Tab l e 1 5,) t he grea
t es
tt
number was aken in August (16. 7 individuals per vi sit ), and the fewest
during February (3 .4 per v is it) .

29. Commercia l monthly landings o f Sphyraena barracuda from Flor -

F ig u r e
ida waters, 1950 to 1957 . Da ta are based on U .S . Fish and W ild life Service
“Florida Landings.”

A compar ison of the commerc ia l and spor t land ings of barracuda is

made (Figs . 30-31) . The large number landed in the commerc ia l catch
dur ing December , January , and February con tras ts with the relatively few
caugh t for troph ies for the same per iod . A lthough some barracuda are
repor ted in commerc ia l catches in Augus t, the ir numbers are propor tiona lly
l
ower than in the P ier-5 da ta for this mon th . Th is may reflec t the d ispersa l
of barracuda from the F lor ida Keys to the Gu lf Stream , as well as a
slackening of fishing pressure in the Keys due to an abundance of
mosqu itos . From Sep tember through November a decrease in commerc ia l
land ngs
i si observed , followed by an increase in he t ca ch
t with he t re urn
t
t t t
of he angler o he Keys
85
 96

48
1953
o
96

:X: 48 1954
(/)
IL
IL
o
o 96 1955
a::
w 48
a:,
:::¡;;
::::,
z o
144
1956
96

48

JAN. MAR. MAY JULY SEP. NOV.

FIiGgUu Rr E
F e 30. Relative monthly catch of Sphyraena barracuda by anglers in

southern Florida in 119953-119956, as reeñfleected in n umber o f f ish from taxidermist'’s

records.
MIiO
M gRrA
aTt IiO
oNnS
s

Reéccoords from commercial catches suggest tha t barr acu da migra te

northward in the sprin g and southward in the fall (T

T abble 1144)).. Other

factors which m a y en ter here include: ( 11 ) an onshore movement in t he

8866
spring, with barracuda in the sout hernmost regions moving in earliest;
(2 ) a coastwise change in fishing pressure as t he weat her 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 weat her
seems unlikely , since in the sout hern part of Fl ori da t he reverse has been
observed . The remain ing possibilities would depend upon t he intensity of
and type of fishing carri ed out in the nort hern coast al areas. Angling i s
w idespread in F lorida throughou t the year , and it would be expected tha t
at least a few barracuda would be reported from the nor thern coastline
during the winter mon ths if they were available .
Migrations of sphyraenids have been repor ted by o ther au thors . Williams
(1 956) believed t hat along t he East Afri can coast S. commersoni ( = 5 .
barracuda) made local migrations correspondi ng t o t he spawning peri od
during the monsoon season. Meek (1916) sta ted tha t barracuda (spec ies?)
migrate periodically to t he coast (localit y?) in the summer and rema in
there un til the end of the year. Migrations were no ted by Villadolid (1940)
for S. p icuda ( = 5 . barracuda) in the Ph ilippines , and by N iko l’skii (1954 :
401) for Sphyraena (sphyraena?). Walford (19 3 3 ) s ta ted tha t S. argentea
makes a spring m igration , evidently for spawning , as far nor th as Ven tura
Count y, Cal ifornia. On t he basis of tagging studies, Springer and McErl ean
(1 961) repor ted tha t, generally , only large specimens of t agged S. barra-
cuda moved any grea t d istance in F lor ida waters .

fei
si
<?
ss f e 40
So

F i g u r e 31. Relative abundance o f angler-caught specimens o f Sphyraena bar-

racuda from southern Florida , taken in 1953 -1956 (broken lin e ), commercia l
landings o f S. barracuda from Florida (so lid lin e ), and commercia l landings
o f total food fish from Florida (da shed lin e ), taken in 1950 -1956 . Angler -
caught specimen s o f 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
Com m e r c ia l l a n d ing s o f b a r r a c u da , 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 i s h a nd W il d l if e S e r v i c e “ F l o r i d a L a n d i n g s/ ' 1952- 1956.

West coast coun tie s East coast coun tie s
(nor th ) (sou th ) (nor th )
e ie
ard s
e e ch Lu
nr roward rtin John
as r o c ia
on
alt ell lli o d lu
ee
. W Pin L C M D PalmB Ma S V
o a ea . rev o .
. t t
B B S
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 — — —

. 1400 140
June. — — — — — 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 ( 170ti 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 — — — — — —

con nued
88
 TABLE 14 (con tinued )

West coa st coun tie s East coast counties

(nor th ) (sou th ) (nor th )
s e iec
n er ro ard in ard ias
lla ch John
lto e lli on d ow Lu lu
ne art
e
. Wa Pi Le Co M D Br PalmB M S B V S
a ea t. rev o t.
.
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 —
i t186i t l t t i ti
i t i .
t tit l t t t t
Insl gh nto, t t e causes
he possib t of he nor t hward m gral on of the
barracuda t may
. be gained by referr tng o F gure 32 i Since
l tt barracuda t are
t t l t
caugh in quan y from Pa m Beach sou hward hroughou he year
. i l i i .
(Tab e 14) counties o he nor h which repor only seasona catches are
it il t i i t t t t t
represen ed The catch from 1952 hrough 1956 s p o ed by mon h for
t , t t l t i l
each coun y nor h of Pa m Beach in which barracuda were repor ed by the
i t t t t .
U.S F sh and Wildlife Service in “F or da Land ngs” (1952-1957)
t t l
Arb rar y using he m dpo n of each coun y on he eas coas where
t t .
barracuda were repor ed I have represen ed on he ef vert ca axis its
t l i t t l t
d s ance from a sou hern baseline, he Monroe Coun y line The area
included between the dashed lines represen s regions nor h of Pa m Beach
which repor ed barracuda during the mon h indicated Superimposed on
the graph are he mean month y so herms for the coas a wa ers along the
89
eastern coast, the values for which were taken from Fug lis ter (1 9 4 7 ) .
Barracuda are t aken in Martin Coun ty in February , and in Apr il and May
they move northward and appear in St. John Coun ty , a d istance of abou t
330 miles nort h of Monroe Count y. Shortly t hereaft ert hey begin a gradual
dr ift sout hward, and by Sept ember they have moved as far south as t.S
Luc ie Coun t .y On the sou t hward migrationt hey are found in Mar ti t n
Coun y until December, with some remaini ng near the sout hern boundary
of this county unt il January. Barracuda move nort hward (or offshore?)
ou t of the Pal m Beach regi on in June and ret urn i n Sept ember. These
migrations probab ly include only a limited segmen t of the popu la tion .
This suggests t ha t barracuda prefer tempera tures h igher than 74°F .,
as no f ish were repor ted in the commerc ia l ca tch nor th of this iso therm .
In May, the m igra tion seems to follow the 7 7 9 iso therm to S t. John
Coun ty where the barracuda appear in June . Before the tempera ture has
reached 80° in July , some of the barracuda are a lready moving southward .
By Sep tember they have moved as far south as St. Luc ie Coun ty , where
t mean so
he i herm
t for this mon ht si 80° . Since here
t si a movemen t of

TABLE 15
N um b er of sp e c im e ns ofSphyraena 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 , from A ug ust, 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
no t m a de f r o m A p r i l 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. Dat e No . Dat e .

No t e
Da .
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 Tota l 649
25 10 22 3 22 4 No . tr ip s 83
26 1 25 4 25 3 N o . per
28 11 27 5 29 8 tr ip 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
 M EA N I S OT H E R M S (°F)
58° 66° 74° 80° 80° 74° 58°

F ig u r e32. Nort h- sout h drift of Sphyraena barracuda in relation t o annual

temperature changes in Florida east-coast waters, inferred from northernmost
locality in which specimen s o f S. barracuda were reported in commercia l
landings by U.S . Fish and Wildlife Service “Florida Landings.” Mean monthly
isotherms taken from Fuglister (1 9 4 7 ). Vertica l axis at right shows principa l
countie s north o f Palm Beach where S. barracuda is landed.

some of t he fish out of the Pal m Beach area in June t o Marti n Count y
when the t emperat ure i s above 82°, adult barracuda may avoid t empera-
tures much h gher
i than his
t figure.
A similar bu less well-marked m igra tion is no ted for barracuda along
t
the west ern coast of Fl ori da (Tabl e 14). Duri ng most of t he year, they
are l anded i n Pinellas Count y (St. Pet ersburg a re a ), but t hese landings
are generally no t extensive pri or t o March or Apri l, even t hough a number
of individuals seem t o be resi dent t here t hroughout t he year. Duri ng May,
August, and Sept ember, barracuda have been report ed as far nort h as
Walt on Count y at the nort hern Fl ori da “panhandl e, ” approxi mat el y 600
miles nort h of Monroe Count y. Large barracuda migrate i nt o t he nort hern
Gu lf of Mexico duri ng t he summer mont hs (Fig. 1 ), and are t aken by
spearfishermen along the Mississippi coas t; these fish eviden tly d isappear
91
from the coas ta l wa ters with the adven t of cold wea ther.
A coastwise rat e of migration of about 2 miles per day was calculat ed
for barracuda from the da ta in Tabl e 14, for fish on bot h t he east ern and
western coasts of F lorida.
E colo g y
Barracuda are abundant about the reefs, shallow flats, and mangrove
hab ita t of F lori da. They are common along t he sout heast ern, wi ndward
side of the F l ori da Keys, but occur i nfrequent ly in Fl ori da Bay and among
the Ten Thousand Islands and nort hward. The barracuda occurs bot h
inshore and offshore, although north of Miami it i s more often found
offshore, presumab ly because it associates with t he warm Gul f St ream which
subsequen tly swings away from coas ta l waters . In litt oral wat ers, the
barracuda shows definite preferences at vari ous ages and si zes in its
choice of hab itat.
Po llu tion .— Alt hough pol lution of the coastal waters adjacen t to Miami
is no t yet extensive, sewage disposal, dredging operati ons, siltation of
reefs, and forma tion of spoil banks have changed some of the environ-
ment, parti cul arl y in Biscayne Bay, which was formerl y more favorable
t o barracuda and its food. Similar operati ons are altering the environmen t
in the same way in the F lori da Keys. Such alterations or destruction of
the hab ita t are of grea t impor tance in limiting the range and abundance of
a species, probab ly far more than is the inefficient harves ting by man .
Living cora l reefs, the associates of which adu lt barracuda so greatly de-
pend upon for food , are par ticu lar ly suscep tib le to m inor physica l changes
(Kuenen , 1950 : 4 14-419) , and some reefs in the M iam i-For t Lauderda le
area which were formerly flourishing have been since affected adversely
by sil tation and probab ly by po llu tion .
Sa li n it y.— Adu lt barracuda t end t o avoid wat ers of low salinity. The
salinity of the coast al wat ers adj acent t o Miami, where adult barracuda
are still somewhat common, i s typically oceanic. The mean salinity of these
wa ters is abou t 35% o , with a range of abou t 24 to 36.6%c (Do le, 1914 ;
Dole and Chambers , 1918) . Fresh wa ters from the Miami R iver and the
Coral Gables Cana l extend into lower Biscayne Bay, Miami (Smith , e t a l.y
1950) . Juvenile barracuda freely en ter the lower reaches of these wa ters
in which the salinity is compara tive ly low , bu t I have never heard of
adults being taken from them 1. Kilby (1955) collec ted a 48-mm specimen
of S. barracuda from Baypor t, F lorida , in wa ter having a salinity of \6 .1%0.
Ne it her Regan (1 9 0 8 ) nor Gunt er (1942) report ed barracuda from the
fresh waters of Cent ral or Nort h Ameri ca, but Thompson [1944?] report ed
t t the barracuda (species?)
ha . . ascends up into qui te bracki sh
t
wa er . . .” in Br sh Honduras Bryan (1915 : 349ff) and H ia tt (1947)
iti .
s a ed ha barracuda is a serious preda tor in the mullet ponds of Hawa ii,
t t t t
l , - t t sit f
On July
i 29 1963, a 25 pound sbarracuda was caugh
s t in ,the tStuden Lake, Univer y o
Miam by Mr. Ray ll Taylor. N o alinity
f value wa ob ained bu these waters are reportedly
fresh and are we above the area o salt intrusion.
92
 which are presumabl y bracki sh. Jordan and Evermann (1 926) report ed
t t in Hawa ii barracuda actually en ter “ . . . mout hs of streams, some
ha
of t hem extendi ng t hei r range i nto fresh wat er. ” Boul enger (1901: 363)
recorded S. guachancho from the fresh wat ers of t he Congo Ri ver, and
Irv ine (1947 : 195-197) also listed this species from t he fresh wat ers of
Africa . Reeve (1912 : 273) stat ed t hat barracuda (speci es?) penet rat e
the upper wa ters of the Gamb ia to a d istance of 160 miles from the sea .
D e p th .— Young barracuda are found usually in less t han 10 feet of wat er,
bu t adults may swim in somewha t deeper st rat a. A school of sphyraeni ds
was observed by Quilici (1954 : 67) at a dept h of 120 feet. The species
canno t be identified with cert ai nt y even from his excellent phot ograph
bu t it appears to be S. obt usat a. Mr. A1 Pflueger i nformed me t hat during
the Ocean Reef F ishing Con tes t off Key Largo, Fl ori da, in August, 1955,
severa l barracuda of abou t 15 pounds were t aken on live bait at a depth
of 240 feet. I have seen a barracuda near the bott om at a dept h of 160
fee t off West End, Bahamas, which weighed perhaps 25 pounds. Large
indiv iduals are t aken by chart erboat men by still-fishing with live baits in
t deep wa ters off Miami. Bu t mos t barracuda are t aken at or near the
he
surface , perhaps because most angling for oceani c species i s by surface
trolling . Dur ing longlining exp lora tions , U.S. Fi sh and Wildlife Service
personne l have t aken barracuda at dept hs of over 200 feet in the
mid-Pacific.
Tem pera t ure .— The barracuda seems to be sensitive t o sudden t emperat ure
changes . Gal loway (1 941) observed dead barracuda t o be “pl enti ful ” at
Key West, F l ori da, duri ng t he cold wi nt er from 1939 t o 1940, when the
wa ter dropped to a low of 43°F in January. Bumpus (1957) gave 71. 6°F
as the mean January wa ter tempera ture for Key West.
The apparent avers ion of barracuda to wa ter colder than 74°F in
F ori da wat ers has been previ ously suggested as a possible fact or control ling
l
the ir migrations. Dur ing per iods of cold wea ther in F lor ida , Mr. R ichard
Meyer of Miami informed me tha t large barracuda are found in dense
aggregations in the warm-wa ter outflow of a power p lan t on Biscayne
Bay, Miami.
Bumpus (1957) gave mean maximum wa ter tempera tures (Augus t)
for Fowey Rocks L ighthouse , sou theas t of Miami, as 83 .8°F , and for
Dry Tort ugas, Fl ori da, as 85.1 °F. Alt hough the barracuda in southern
Fl ori da wat ers seem to t ol erat e t hese high wat er t emperat ures, t hose nort h
of Miami woul d appear to avoid t hem (Fig. 3 2 ). In Bimini harbor, small
barracuda were observed to become sluggish when t he water t emperat ure
exceeded 86°F. The shallow bank wa ters east of Bimini are subject to
wide vari ations in t emperat ure (Newell and Imbrie, 1955), sometimes
droppi ng to t he middle forties ( ° F ) . Dr. Louis A. Krumho lz informed me
t hat in January, 1956, a low of 57°F was recorded, and in July, 1956, t he
t t t t lt
empera ure rose o 9 8 °F in these lagoon wa ers of Bimini. A hough
93
barracuda are sometimes common in such areas , they perhaps avoid
extremes of tempera ture by moving in and ou t with the tida l waters .
In July 1956, Dr. Krumholz and I observed barracuda about 12 inches
long swimming near the Lerner Mari ne Laborat ory dock at Bimini in
wat er about 10 to 15 inches deep, t he t emperat ure of which was 88. 7°F
at t he surface. The barracuda were in the company of needlefish ( S trongy -
l ura ), moj arra ( G erres ), and mullet ( M ugil ) but did not seem t o be
feeding upon these fishes. These associates were seen to swim i nto wat ers
as shallow as 3 or 4 inches deep in which the surface t emperat ure was
93 . 2°F , bu t the barracuda did no t follow them into these warm wa ters ;
they had prev iously been observed to do so when the wa ter was 80.6 to
82 .4°F . On August 29, 1955, I collected a number of 2- to 4-inch
barracuda from wa ter 2 to 3 feet deep at the Matheson Hammock wading
beach, Biscayne Bay, Fl ori da, where t he t emperat ure was 94-95°F. On
o t her occasi ons when small barracuda were collected at the same pl ace ,
wa ter t emperat ures commonly exceeded 88°F. Alt hough the young would
thus appear to to lera te h igher tempera tures t han t he adults, adult barracuda
seldom en ter the very shallow wa ters regardless of wa ter tempera ture .
E c o l o g y o f Y o u n g B arracuda . — Tempera ture evidently plays a ma jor role
in determ in ing the seasona l movements of bo th the young and adu lt
barracuda . As ide from the migra tion pa ttern which was discussed above,
and which apparen tly is par tia lly a func tion of tempera ture , seasona l
changes in wa ter tempera ture seem to be respons ible for movements
between shallow and deep wa ter. These movements are corre la ted with
grow th and are reflected in the appearance of cer ta in items in the food
cha in (Tab les 16 , 3 0 ) . The young occur in abundance along the shallow
beaches as la te as mid-September . The ir d isappearance from the shallows
i
ndicates tha t young f sh
i overwin er t in deeper wa er t where here t si a
l t t
sma ler empera ure range and changes occur less rap d y i l .
Duri ng t he warmer months, young barracuda were t aken most often
jus t below the surf, and over sandy or weedy bo tt om. Here t he det rit us
of mangrove seeds and l eaves offers prot ecti on, as well as an excellent
i t t .
vantage po n from which o forage Young barracuda find food and shelter
, . , t
in floating Sargassum spp. (Breder 1949) Occasionally I have aken
i tit
young barracuda assoc ated with such a small quan y of Sargassum or
t t t l t.
C y m o d o c e a ha it would seem o afford no concea men
t ti t l t tl
The young often find pro ec on among he b ades of ur e grass
l i i i , l ,
(T ha ass a) and its assoc ated algal flora such as L aurenc a C au erpa and
l i i . : t l tt
P o ys ph on a Waite (1923 109) no ed a remarkab e similarity in pa ern
. t t t li t l
in young specimens of S ob u sa a from Aus ra a and he b ades of
t t t t t
dugong (C y m o d o c e a ? ) grass in which he young live, and s a ed ha
. t t , t i l t , t i
when
i danger hrea ens , t ihey immed ate y upend i hemselves
t l he r
snouts d trected downwards he tr tails i moving in un son l with he l ades
b .
of grass o which the stripes on he f sh bear a remarkab e resemb ance ”
94
 Rosen (1 911) observed young barracuda in tidepools in the Bahamas .
Such niches are impor tan t for the young bo th as a source of shelter and
for the rich gobiid fauna which serves as food for young barracuda .
Loose aggregations of young barracuda , especially dur ing per iods of
feeding, have been observed by me. Such groups usually compr ise from
10 to 30 individuals , bu t occasiona lly large numbers of individuals have
been report ed. Longley and Hil debrand ( 1 9 4 1: 6 9- 7 1) not ed t hat on
one occas ion in the Tor tugas . . abou t a thousand young up to 150 mm
in length were observed in one school.” Gary Benne tt, of Cocoa , F lor ida ,
reported in his “Trou t Cap ital Fishing News ,” da ted Augus t 6 , 1 9 5 6 , tha t
at the west end of Canaveral H arbor and along the jetties there were
“ thousands of very small baby barracudas . . .” less than 1 Vi inches long .
Large numbers of young are no t common on the wes tern coas t of
Flori da. Rei d (1 954) and Kilby (1955) repor ted only a few young
barracuda from the Cedar Keys-Baypor t area as a resu lt of severa l years’
intensive collecting.
Eco l ogy of A d ult Barracuda. — Large adults are comparati vel y rare along
the eas tern coas t of F lor ida nor th of Cape Canaveral. They seldom venture
into shallow, sandy localities found here where envi ronment al changes are
often severe. Barracuda more t han 3 feet long are “l one wolves” of the
sea, are usually solitary bo t h i n reef and offshore areas, and constant ly
appear to be searching for food (Sauvage, 1891; Gudger, 1918). In
con trast t o younger stages, adults usually do not aggregate; on occasion ,
groups of two or t hree large i ndividuals have been observed t o cruise slowly
abou t. During the wi nt er, these l arger barracuda are more prone t o
aggregate.
In Bimini , Bahamas , barracuda to 4 or 5 fee t long have been observed
abou t the oolitic islands, along the deep channe ls , and , in the summer, at
the junc tion of the Gu lf Stream and the coas ta l wa ters or in the Gu lf
Stream . Dur ing July , 1955 . and July , 1956 , large barracuda were observed
and taken on hook and line southwest of Bimini , where they occur over
extensive turt le-grass communit ies , usually a t high tide . With low tide ,
t
hey were seldom caugh t in these shallow regions, bu t could be aken t on
the leeward side of the islands, which is steep and cora lliferous , and where
t
hey were seldom akent on high tide . In the F or
l da
i Keys, such a preference
t t t i l ,
seems o reflec he r avoidance of cora stacks and barracuda were seldom
observed swimming abou t the jagged branches of e lkhorn cora l (A cropora
pa lma ta) and s taghorn cora l (A . cerv icorn is) , par ticu lar ly when heavy
swells might dash such long-bod ied species as barracuda about. Thus , the
smaller reef fishes may find ample pro tec tion among the branches of these
i
mposing cora l barr ers
i (see Longley and H ildebrand , 1941 , PI . 2, Fig . 1) .
Large barracuda were often observed hovering nearly motionless jus t
off the bo ttom , or near the surface (Longley and H ildebrand , 1941 , PI. 2,
Fig. 2 ) . They also occur commonly a t the junc ture between the turt le-grass
95
beds and t he extensi ve sand pat ches fringing t he reefs. Ot hers may occupy
shallow t renches a short di st ance from t he reefs. These poi nt s are excellent
places from which to forage for reef-dwelling prey.
It seems possible to offer some hypo theses on the seasona l movements
of barracuda in F lor ida wa ters using severa l independen t cr iteria . Under-
wa ter observat ions on the di st ri buti on of various age classes in different
hab itats have been corre la ted with (1 ) catches in seine hauls , (2 )
observa tions made while angling for barracuda , and (3 ) informa tion
from charterboat men and ot her anglers who are famili ar with t he seasonal
occurrence of saltwa ter fishes in southern Florida.
Postl arval barracuda i nitially occur in the shallow grass flats from late
spring to early fall. Dur ing thei r first wi nt er, when t hey are about 6 t o 8
inches long, barracuda appear to move offshore i nto grass flats which are
approx ima te ly 1 t o 2 fat homs deep; at this si ze t hey are seldom if ever
found in reef areas . A t the end of t hei r first wi nt er, at a length of 8 t o 10
i
nches, they rema in in the shallows. Some en ter mangrove habit at s or the
deep channe ls between the keys. By t he beginni ng of t hei r second wi nt er
a t an average l engt h of 20 i nches, t hey move offshore t o t he coral reef, in
depths of from 3 t o 10 fat homs. By t he beginni ng of t hei r t hi rd year of
life, the maj orit y are mat ure, and alt hough most remai n on the reefs, some
en ter the shallow mangrove habit at by l at e spring. They overwint er on
the reefs, and in spring many migrate nort hward or are found in the
Gu lf S tream. The adults remai n scatt ered until Sept ember when they
gradua lly re turn to the reefs. These fall concent rati ons on t he reefs are
coincident with t he sout hward migrati on along t he Fl ori da coast. Duri ng
the winter, in the deep channel s of the Fl ori da Keys, chart erboat men have
observed large aggregations of barracuda which t hey claimed would
seldom take a bait .

A sso c i a t e s an d P red at o rs. — The adults of the four common sphyraeni ds in

the wes tern Nort h Atl anti c do not occur in the same habit at. Adult speci-
mens of guachancho are often t aken in 12 t o 20 fat homs of wat er by
shr imp trawls in the Gu lf of Mexico, according to H ildebrand (1954 ,
1955). This species is common enough at Veracruz, Mexico, t o ent er into
the commerci al fishery (Dr. Henry H. Hil debrand, personal communica-
ti on). Bot h b oreali s and pi cu dill a occur in deeper st rat a in t he west ern
A tl anti c t han does barracuda. Where barracuda i s not common, the ot her
l i t
three species rep ace it in the food chain n shallow wa er.
The young of barracuda and boreali s were t aken t oget her by me, and
on ot her occasi ons specimens of barracuda and guachancho occurred i n
the same seine haul. Small specimens of guachancho are not often report ed
and may be overl ooked, although t hei r patt ern i s sufficiently di sti nct to
i i t i t . .
nsure den if ca ion (Fig 4 )
i t t i t t
The mpor an groups of fishes and nver ebra es 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, IM PORTANT ORGANISMS ASSOCIATED WI TH 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 da a t t h is s iz e a r e a b s e n t o r
RARE IN TH IS N IC H E . O R G A N IS M S P R E C E D E D BY ( 1 ) ARE PREDATO RS W H IC H
C O M P E T E W I T H BARRACUDA FOR FOOD. O R G A N I S M S P R E C E D E D BY ( 2 ) ARE
E A TE N BY BARRACUDA . O R G A N IS M S PR E C E D E D BY ( 3 ) C O M P E T E W IT H A N D
ARE E A T E N BY BARRACUDA O . RGAN SM I S P R E C E D E D BY 4 ( )N E T H
I ER COM
P E T E W T IH NOR ARE E A T E N 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 ti forme s
Engraulidae 2 Clupei -
Belonidae ( yg .) formes + +
Syngnathiformes 2 Gerridae
Tet raodon ti- Scaridae
forme s (yg .) (yg)
4 Mug ilidae
t l i s
13-18 Gerridae Belonidae - 2 Atherinidae
A herinidae .) Lutjanidae
t s 2 C upe forme
t
Engraulidae Cyprinodon
- 2 Cephalopoda +
Scaridae i(juv fi'forme
s 3 Synodon idae
Synodontidae Clupei 3 Belonidae
Pomadasy dae orme
Gerridae
Mugilidae
Carangidae
19-30 2 Mug lidae 1 Ser o a spp.
2 Lutjanidae
i 2 Pomadasy
i l dae
2 Clupeidae 2 Te raodoni orme
3 Synodontidae 2 t
Lutianidae tif s
+ 3 Belonidae 2 Hemiramphidae
2 Cephalopoda
3 Synodon idae
3 Belonidaet
3 Serranidae
3 Carangidae
3 Scombridae
4 Elasmobranch
ii
>30 Scom berom orus 1 Scom beromorus
spp spp
Ser o .a spp. 1 Coryphaenidae
.
Lutjanidae
i l 1 Is iophoridae
large indiv duals 2 Cephalopoda
t
Serranidae
( i ) 3 Eu hynnus
(large indiv duals 3 S ronpy
t ura
+ + Tetraodon orme i ) t raph ldoma
Hemiramphidae
tif s 4 Acan hocyb
i um
Cephalopoda 4 Thunnust spp
i
Eu hynnus 4 K a suwonus .
t
Caranx spp. t
4 Exocoe dae
Belonidae ti
4 Elasmobranchii
Elasmobranchii

97
various hab it ats occupied by barracuda of differen t sizes in sou thern
F lorida, and the relationsh ip between the barracuda and these organisms ,
is summarized in Tab le 16. Large barracuda inhab iting reefs occupy the
same preda tory posit ion as the amber jack (Ser io la dumerili), large jacks
( Caranx spp .) , and various species of mackere ls ( Scomberomorus spp .) .
Needlefish ( S trongylura spp .) also compete for food , bu t to a lesser exten t
than o ther 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 ma jor preda tor w ith the amber jack . O ther carn ivores
which occur in this hab it at are groupers (Ep inephelus, M ycteroperca ,
Cepha lopho lis), large snappers (Lu tjan idae) , little tuna (Eu thynnus) , and
sharks ; sharks apparen tly do no t compe te w ith barracuda for the same
t
ype of food . The barracuda si capab el of ak t ng
i and killing large prey
with its immense gape and powerfu shear ng jaws can ingest much larger
l, i
fish than can snappers , jacks , scombrids , and small groupers .
In the offshore, pelagic habit at, common predat ory species i nclude
sa lfish (Isti ophorus), dol phins (Coryphaena sp p. ), and t una-li ke fishes
i
(Thunnus spp., Kat suwonus). These offshore predat ors compet e only t o a
limited extent with the barracuda (unpubli shed d at a ). Within t he sandy-
shore hab it at, young barracuda have t he l east compet ition, for t he small
needlefish are the only common preda tors .
Gudger (1 918) discussed the presence of a remora (Echenei s naucrates)
on barracuda from Fl ori da wat ers, and I have seen echeneids on barracuda
underwa ter bu t the species could not be identified. Poey (1865) descri bed
Echenei s sphyraenarum from t he gill s of barracuda from Cuba, and
Schultz (1943) descri bed a specimen of S. snodgrassi ( = 5. barracuda)
from Can ton Island in the Phoen ix Islands.
Many popul ar tales rel ate the animosity which supposedl y exists between
barracuda and sharks. This i s suggested by Wri ght ’s (1948) observat ions
with a group of Bri tish frogmen in t he Bahamas. He report ed t hat a
barracuda att acked a nurse shark (Gi ngl ym ost oma ci rrat um) which had
been speared through a gill slit and was b leed ing profusely . Ogilby (1915 :
8, 47) no ted t ha t in Aus tra lia, S. ob t usa ta has been known to drive off
large sharks. Dur ing the course of abou t 40 underwa ter tr ips to the F lorida
Keys and the wes tern Bahamas , I never saw sharks and barracuda toge ther ,
although they are bo th common in these waters .
Evi dently, few predat ors are sufficiently l arge and swift t o capt ure
large barracuda , a lthough I was t old by a spearfisherman t hat he once
found a small barracuda in t he stomach of t he spotted jewfish (Epi ne-
phelus it ajara), report edl y a rat her sluggish species. Kl unzinger (1884)
report ed t hat t unas feed upon sphyraeni ds, and ot her aut hors have not ed
t
races of small specimens of Sphyraena in the d iet of several species of
Pacific tunas . I have t aken two barracuda of 20-30 mm from t he stomach
of do lphin (Coryphaena h ippurus) from P ier 5, Miami.
98
 B e h a v io r
Typical coloration and positions when res ting are described for young
barracuda by Breder (1948a , 1949) , and by Long ley and H ildebrand
(1941 : 6 9 -71) . Schlesinger (1911) briefly discussed the fins and genera l
swimming behav ior of S. sphyraena . The young of barracuda closely re-
semble twigs, and on severa l occasions have been near ly overlooked as
t seine hauls were being exam ned
he i . Young and uveniles
j l
b end well with
the ri backgrounds in na ure t , and can change he t ri pa ttern accordingly.
Over sand bo ttom they are ghostly pale , bu t over sand mixed with tur tle
grass they have irregu lar , greenish bands ; the trans ition may be no ted as
a barracuda swims from one type of hab ita t to ano ther . The b lack
marg ins of the cauda l fin and opercu lar flap are conspicuous during bo th
of these phases .
B arracuda are swift swimmers. Gray (1957) repor ted t ha t the speed
of S. barracuda was clocked in a labora tory tank a t 27 miles per hour .
A slow drift is characteristic of indiv iduals when no t actively feeding.
Breder (1926) described the ir locomotion as “pike-like movemen ts” with
great “flexibility in con tinuous swimming.”
Large barracuda were often observed to lurk just benea th the surface
in the wa ters of Bimini and the Florida Keys. H ia tt (1 947) stated t hat
“barracuda habitually appear ha lf asleep and float in one positi on close
ot the surface, often mak ng i slow movements from side ot side. When
t
hey are d si urbed
t t
hey move with extreme rap di ity ”.
On several occasions I have not ed large individuals swimming erraticall y,
with a list or with a distinct, convex flexure in the back . Since these fish
appeared only momentarily , it was no t known if it represen ted a temporary
behav ior pa tt ern or was a deformity . Wrigh t (1948) no ted such a hump-
backed indiv idual in the Bahamas , and spearfishermen have observed them
ni he
t F lor ida Keys.

Aggregati ng B ehavi or.— In barracuda from 10 to 30 inches long, two

types of aggregating behav ior were observed : feeding and defensive.
For examp le, a typica l haun t of barracuda from 10 to 15 inches long is
a qu ie t, shallow area , abou t 2 to 6 feet deep , with sand bo ttom 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 terr itory . Occasionally one wou ld dar t
ou t to ensnare a passing fish bu t re turned to its terr itory . Aggregating
behav ior to feed begins aft er the onset of flood tide, coinci dent with the
shoreward movemen t of schools of small atherini ds, clupeoi ds, and gerrids.
These schools of forage fishes rema in compac t and seldom sca tter when a
single barracuda rushes a t them . Gradua lly the barracuda leave the ir
respective territories, congregate , and harass the schools of small fishes,
i i i i ,
p ck ng off stragglers. Such behav or has been descr bed by Gudger (1 9 1 8 )
, .
Mowbray (1 9 2 2 ) and Gordon (1 9 4 6 )
99
 When a school of prey i s swimming over sand bott om, seven or eight
barracuda have been observed t o li e paral lel to one anot her over a weed
bed, facing the prey. This apparent abandonment of t errit ori alit y was also
no ted on numerous occasi ons when a lure was cast into a weed bed con-
ta ining barracuda . If the lure was ret rieved rapi dl y, often one barracuda
would dart out t o t ake the lure. However, if the l ure was ret rieved slowly
a number of barracuda would converge from the weed bed t oward the
lure bu t made no a ttemp t to strike a t it. When the l ure was slowed or
stopped the barracuda would slow or stop; in the l att er i nstance they usu-
ally re turned to the weed beds.
The second type, designa ted as defensive behav ior, was observed in
barracuda from 18 to 30 inches long. This reac tion seems to be due to the
presence of an enemy. A t Ca t Cay , Bahamas , on two occasions barracuda
formed a semicircle abou t me as I en tered the shallow wa ter , bu t remained
abou t 10 to 15 feet di st ant. As I approached , they gradually retreated ,
and the reverse was true. On each occasion, when I attemp ted to spear
one it would dar t away with grea t speed bu t immed iate ly re turned to its
approx ima te place in the formation . Several barracuda swam a t the
unre tr ieved spear; they merely bumped it with the snout, bu t made no
a ttemp t to bite or slash.
Ward (1907 : 313) report ed t hat schools of sphyraeni ds (speci es?) off
Aus tra lia . . rival t hose of herri ng and mackerel for numbers and
denseness.” Walford (1932 , 1933) and Ni kol ’skii (1954: 401) briefly
men tioned schooling behavior in Sphyraena. Schooling or aggregating
behav ior seems t o be confined to the small species of Sphyraena, or to
the juvenile forms of the large species .
Barracuda less than 2 fee t long are sometimes found in we ll-marked
i ,
s ze aggregations even within the same age class. Spawning barracuda
, t t t
may congregate int schoolst of i the same sex based on my observa .ions ha
individuals caugh in a res r cted locality are often of the same sex
t t
On two occasionst t a Bimini hree large individuals were observed to
cruise
i t slowly , at he surface, drifting offshore with the tide. In the second .
ns
l ance
t the hree were swimming within inches of one another The ,
arges barracuda
t was swimming slightly ahead of and between the others
and .was aken ton hook and line; it was a 26-pound . female with nearly i ripe
eggs On the o her occasion i tnone was caught Gudger (1918) t descr, bedl ithis,
accompanying
t t t behav
t or in hree barracuda tobserved a iTortugas . Ft or tda t
excep ha the hree individuals were abou ...t i the same s
i ze He no ed ha
the two
t individuals on
t l i either side
. . had “ he r heads n the region
. of the
righ andt left pec ora i f ns .” of the middle barracuda This may
represen pre-spawn ng activity, since the barracuda from Bimini had
nearly ripe ovaries.
t i i i
A ype ; t of accompanying behav or tt st observed when a barracuda s
hooked o her individuals may be a rac ed and appear frenzied as they
100
swim beside the tethered compan ion , and occas ional ly will slice off the
tail of the hooked fish.
Peck O rder.— Barracuda evidently are no t very compatib le, and it would
seem tha t where aggregations of indiv iduals occur , some type of social
stab ilization wou ld develop. Ev idently, a h ierarchy exists in the juveniles ,
as suggested in the following observa tions made at Bimini at t he Lem er
Marine Labora tory .
On July 11, 1956 , three barracuda which had been caugh t on hook and
line were placed in a concre te poo l, measuring approx ima te ly 4 fee t wide
by 2 fee t deep by 6 fee t long. The fish inc luded a 450-mm ma le , a 398-mm
female , a 397-mm male , and a 306-mm indiv idua l (fork leng ths) whose
sex could not be determ ined subsequen tly by dissection. The first t hree
barracuda were p laced in the tank w ithin a few minu tes of one another.
Each wen t to and rema ined in a separa te com er of the tank , brea thed
rap id ly , and a ttemp ted to ma in ta in ba lance . Each lay near the bo ttom
and close to the side of the t ank. Within a few seconds after t he t hi rd
barracuda was p laced in t he t ank all had recovered . The three individuals
raced at one ano her bu t no, con
t ac was t tmade and ,af er a few seconds
again re t ti
red ot separa et corners of the ankt .The two f sh
i which measured
397 mm and 398 mm lashed a t and sideswiped one ano ther ; the ir snouts
glanced off a t an angle jus t below the base of the first dorsa l fin. This
“jousting” was repeated four times , and then one struck a deep , glancing
blow which opened a gash about 1Vi inches l ong and about V4 i nch deep
on the side of t he ot her fish. Agai n they hit each ot her with a sideswipe,
a lthough this time t here was no gash inflicted, and they t urned once more
and a ttacked . They slowed momentarily, cl amped t he tips of t hei r jaws
t
oge t , rocked the ir heads sideways for severa l seconds, and slowl y
her
released t hei r hol ds. Thi s was repeat ed once more, and t hen t hey ret urned
ot he
t ri respective comers .
A t this time the larges t fish lef t its corner and faced the adversary
which had inflicted the gash. Bo th “ jous ting” and jaw-c lamp ing were re-
peated. The larger fish wen t to the corner prev iously occupied by the
sma ller barracuda . The entire episode occurred in approx ima te ly two
minu tes . A four th barracuda (306 mm) was p laced in the tank with the
o thers , and it en tered the unoccup ied corner . The three orig ina l indiv idua ls
congregated in one corner while the smalles t barracuda occupied the d i-
agona l corner . No a ttemp t was made by any to fight with this newcomer ,
nor was any fighting observed up to the time when the observa tions ceased
abou t 15 minu tes la ter.
Duri ng the present i nvestigations, several barracuda were caught which
bore scars below the spinous dorsal base which might have been obt ai ned
in such a battle. This may also account in part for t he high percentage of
regenera ted scales which I found in this reg ion of the body .
Mowbray (1 922) wrot e t hat in the F lori da Keys, two adult male barra-
101
cuda which had been pl aced in t he live well of a boat inflicted severe
gashes on one anot her soon aft er t hey were confined. Mr. Craig Phillips,
formerl y of t he Miami Seaquari um, i nformed me t hat shortl y aft er i nt ro-
duc ti on i nto a t ank 80 feet in di ameter, t wo l arge barracuda fought, split
each o t her’s head open, and bot h died.
It is not known at what size this fighting commences. Barracuda less
than 1 inch long will devour or a ttemp t to ea t one ano ther when p laced in
a 40-gallon aquarium tank . Cann ibalism was no ted by De Kay (18 4 2 : 3 9 -
40) in young specimens of S . borea lis .

F e e d in g H a b it s

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

more read ily when in the company of congeners than when only one
specimen is presen t. Bo th young and adu lt barracuda have have been
observed to swallow prey e ither head or tai l first, or to grasp the prey a t
the midsection , fold it, and swallow it . On o ther occas ions , the prey may be
seized at the midsection and sliced in two or more pieces , and the ta il
section swallowed; this is more common in larger individuals . The litera-
ture on the func tiona l morpho ogy
l of feeding in this family si limited to a
reference by Yasuda (1960) on S. p ingu is .
Barracuda from abou t 100 to 300 mm were fed por tions of frozen
shr imp ( Penaeus ) . Usua lly they allowed the shr imp to fall to the bo ttom ,
where it was scru tin ized for a few seconds before being held loosely in the
jaws momen tar ily pr ior to swallowing. Those from which food was w ith-
held were quite ravenous and took the shrimp as soon as it touched the
surface of the wa ter.
Barracuda of all si zes feed upon moving organisms, and t he stomach
content s suggest t hat t hey depend pri maril y upon sight t o obt ai n t hei r
food . Thei r predil ecti on i s for fi sh which flash or show some type of
abnorma l, erra tic movements .
Barracuda are generally daytime feeders, and probabl y do not move
about much at night. At Bimini, small cypri nodont s and gerrids i nt roduced
a t ni ght into a t ank contai ning several barracuda were still t here several
hours l at er, whereas under daytime conditions t hey were readi ly consumed.
Barracuda caught earl y in t he morni ng usually had empt y 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 in teres t in lures or fishes used as ba it.
Duri ng 1961 and 1962, several peri ods were spent i n the west ern
Bahamas and the Fl ori da Keys using wat erproof ni ght lights over reefs
and in deeper wat er. On one occasion, a barracuda of abou t 3 feet long
approached t o wit hin 10 yards of t he l ight (150 watt s) but was apparentl y
dazed by the light and slowly dri ft ed away. On numerous night-divi ng t ri ps
to the F lori da Keys, Walter A. Starck, II, and William P. Dav is, of the

102
Institute of Marine Science, Univers ity of Miami (persona l commun ica-
ti o n ) , observed a barracuda on only one occas ion ; this too appeared dazed
by the ir lights as they swam underwater.
Relatively little noc turna l tro lling or drift fishing is carr ied on in sou thern
F lorida , and thus deductions based upon t he l ack of specimens cannot be
considered conclusive. Bu t Springer and Bullis (1956) recorded barracuda
t aken on longline gear in the nor thern Gu lf of Mexico between the hours
of 10 PM and 1 AM. Williams (19 5 9 ) report ed t hat all of t hei r t roll
catches of barracuda off Eas t Africa were made during dayligh t hours ,
although o t her sphyraen ids were predom inan tly n ight-time feeders. La-
Monte (1952) stat ed t hat members of t he family Sphyraeni dae often feed
a t night , and Doyle (1 953) referred t o barracuda (speci es?) caught by
n igh t angling off East Africa. Pal ombi and Sant arelli (1 9 5 3: 2 4 ) st at ed
tha t in the Med iterranean Sea, the commotion made by small fishes which
have been att ract ed t o the surface lights used by fishermen att ract s the
a tt enti on of Sphyraena spet ( —S. sphyraena). Ommanney (1953) and
Williams (1956) not ed t hat large numbers of S. j ell o were t aken at night,
primarily dur ing periods of full moon, off Ami rant e Bank, Seychelles, by
hand line , bu t specimens were never taken by trolling .

F ood H a b it s

Excep t for t he studi es of Hi att (1 947) and Williams (1 9 5 6 ), little has

been found in the lit erat ure regardi ng the food of t he barracuda. Barracuda
stomachs were collected mos t of the year in sout hern Fl ori da from t he
spor t fishery. A dependence on such a fishery leaves much t o be desired,
since if the fish does stri ke a l ure or bait it implies t hat t he if sh must i have
been hungry or ncited. Thus exami nati on of t he stomach may cause an
underes tima tion of the quan tity of food taken under norma l circums tances .
M e t h ods. — Stomachs of 901 barracuda were exami ned from vari ous l o-
calities in the t ropi cal west ern Atl anti c Ocean. Nearl y all adult fi sh exam-
ined (l onger t han 300 mm) were caught on hook and line with live or
artificial ba it, and a few juveniles were taken in seines .
Al l young fish cap tured by me were taken e ither with a 10-foo t “Com-
mon Sense” V^-inch mesh cott on minnow seine, or with a 30-foot nylon
seine of V^-inch mesh.
A t Bimini , most stomachs of adults were exami ned in the field wi thin
a few minutes of cap ture. A few adults which were caugh t on char ter
boats at Bimini and all of the adults examined from P ier 5, Miami, had
been caugh t from one-hal f t o eight hours pri or t o the time of dissection.
There was no evidence from presen t observat ions t hat barracuda re-
gurgitate food in t hei r stomachs when t hey have been caught on hook and
line, such as occurs in cer ta in scombrid and o ther fishes.
Food is here considered as any type of solid ma tter inc luding fish bones ,
flesh, scales, squid beaks and “pens ,” algae , and vascu lar p lan ts . 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 A TL A N T IC . Y O U N G WERE COLLECTED FROM
1 9 2 0 t h r o ug h 1 9 5 6 ; a d u l t s w e r e c o l l ec t ed f r o m A p r il , 1 9 5 5 ,
th r o ug h A ug ust , 19 5 6 .

Adults
Young mm
( 3 0 0 -1 2 1 0
( 5 .5 - 2 9 9 mm fork length) fork length) Tota l
No. items
per
stomach Frequency Per cen t Frequency Per cen t Frequency Per cen t
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

recogn izab le ma ter ia l was recorded as “unident ifi able matt er. ” If only
chyme was present, the stomach was considered empty. Food i s report ed
as the number of stomachs conta ining a parti cul ar item. Vol umet ri c dat a
were no t taken .
Num ber of It em s of F ood per St omach. — On t he average, each stomach
con ta ined only one item (Tabl e 17), and this usually filled the stomach
area . The tendency for a stomach t o have more t han one item was only
somewha t more pronounced in t he young barracuda; 33 per cent contai ned
more t han one item, and in the adults 25.3 per cent did so. One 850-mm
barracuda , caugh t near Bimini, held chopped remnant s of worm eel s,
Ah lia egmon tis, i ncl uding 11 heads. A barracuda 56 mm long, from Bis-
cayne Bay, Miami, con ta ined 16 postlarval a therin ids which ranged from
11 to 20 mm in t ot al length. In most i nstances when young barracuda
had ingested more than one item of food, t he organisms were the post -
larvae of schooling fishes such as at heri ni ds, engraul ids, or clupeids. Bean
(1 8 9 0 ) , Gudger (1 9 1 8 ) , Longley and Hil debrand (1941 : 69-71), Gor-
don (1 9 4 6 ) , and Legand (1952) not ed t hat the barracuda feeds upon
schools of small fishes, but did not gi ve the si ze of t he barracuda nor the
prey.
The percentage of stomachs conta ining food in young barracuda is given
in Tabl e 18.
Food of Young from F lor ida .— The organisms found in the stomachs of
154 young barracuda from sou thern F lor ida are given in Tab les 19 and
20. Fishes occurred in 95.5 per cen t of the stomachs which con ta ined food.
104
 TABLE 18
P e r c e ntage 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 i d a , 1945 t h r o u g h
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 i n p a r e n t h e s e s .

No. %
Fork exam- with
eng
l ht (mm) Jan . Mar . Apr . May June Ju yl Aug . Sep t. Oc t. Nov . Dec . ned
i food

0 -2 4 100( 1) — 100 (1) 0 (2) 100 (1) 100 ( 3) 59.1(2 2 ) — — 100 (1) — 31 6 4 .6

2 5 -4 9 — — 8 3 .3 (6 ) 1 0 0 (3 ) 8 3 . 3 (6 ) 8 9 . 9 (9 ) 60 (3 1 ) — 8 5 .8 ( 7 ) 1 0 0 (3 ) — 68 7 9 .5

5 0 -7 4 — — 100 (1) 100 (2 ) 1 0 0 (1 0 ) 50 (2 ) 7 1 .4 (7 ) 1 0 0 (3 ) 6 6 .7 ( 3 ) 75 (3) 100 ( 1) 32 8 7.5

75 -9 9 — — — — 100 (1) 100 (8) 100 (2) 1 0 0 (1 ) 1 0 0 ( 2) 1 0 0 (3) 0 ( 1) 18 9 4. 5

1 0 0 -1 2 4 5 0 (2 ) — — —■ 50 (2 ) 100 (3 ) 1 0 0 (3 ) — — 50 (2 ) — 12 8 1.8

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

1 5 0-17 4 — 10 0(1) 100 (1) — — 100 (2) —■ — — 100 (3 ) 50 (2) 9 8 8. 9

1 75 -1 9 9 100(1) — 100 ( 2) — — — 100 (1) — — 100 (3) 50 (2) 9 8 8.9

2 00-224 — 1 0 0 (1 ) 10 0 (1) — 3 3 . 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 7 5 ( 4 ) 1 00 (2 ) 9 1 .7 (1 2 ) 7 1 .4 (7 ) 8 2.6 (23 ) 9 3.1(2 9) 6 9.6 (6 9 ) 100(4) 8 3 .3 ( 1 2 ) 8 1.5(26 ) 4 2 . 9 ( 7 ) 1 95 7 9 .0

No. items 5 3 11 6 19 27 54 4 10 22 4 1 65 —
No. stomachs __
with food 3 2 11 5 19 27 48 4 10 22 3 154
 Food preferences are no t marked , and the young barracuda ea t
wha tever is available in the appropr ia te s ize class. However , cer ta in
groups of fishes are taken frequently (Tab le 2 0 ) . Gob io ids , clupeoids , and
atherin ids were by far the mos t impor tan t items of food, and occurred
in 69.1 per cent of the stomachs. Gob io id fishes excep ted , the striking
fea ture in the d ie t of the young barracuda is the predom inance of small ,
schooling fishes of the littora l regions. Gob io id fishes, however, occur in
t
idepools and on sand flats in grea tes t abundance , where they find shelter
in bivalve shells and echino id tests.
Inver tebrates as a group were a m inor food item of the young, and
occurred in only 3.9 per cen t of the stomachs.
F ood of Young from Bi mi ni, Bahamas. — Coll ect ions of young barracuda
were made in July, 1955, and July, 1956, mostly along the east ern shore
of North Bimini , in front of the Lerner Mari ne Laborat ory. The difference
between the two years in relative percentage contai ning food i s not
significant (ch i-square = 2 .64) at the 5-per cen t level (Tab le 2 1 ) .
Fishes were found in 97 per cen t of the stomachs of the young which
con ta ined identifiable food ; of these , 30 ( 4 6 % ) were identifiable.
The kinds of food taken by young barracuda from Bimini in 1955 and
1956 are shown in Tab les 22 and 23 . Gob iids, a therin ids , gerrids, cyprino-
dontids, and clupeoids were found in 26 (8 6. 7 % ) of t he 30 stomachs
which conta ined identifiable fishes, with a therin ids the most common .

TABLE 19
Su m m ar y of fo o d f o u n d in the s t 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 i d a ,
1945 th r o ug h 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 . Tota l
Gobio ide i 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
Cyprinodontiforme s 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
Tota l 12 11 6 19 27 55 4 10 22 166

106
 TABLE 20
F ood it e m s f o und in the st om a c hs of 154 y o ung s p e c im e ns ( l e ss than 300 m m ) of Sphyraena barracuda
c oll ec te d in so uth e r n F l o r id a , 1945 t h r o ug h 1956. N um bers ar e st om ac hs in w h ic h it e m s wer e fo und .

Pisces:
Clupeidae — — — — — 6 — — — 6
Redear sardine, Harengula humeralis — 1 — — — — — — — 1
Harengula sp. 1 — 1 — — — — —1 3
Dwarf herring, Jenkinsia lampro taen ia — — — 1 — 2 — — — 3
Engraulidae — — — 1 — 1 — — — 2
Striped anchovy , Anchoa hepsetus — — — — — — —1 3 4
Cyprinodontidae
Rainwater killfish, Lucania parva 1 — — — — — — — — 1
Poeciliidae
Mosquitofish , Gambus ia affinis 1 — — — — — — — — 1
Sailfin molly , Molli enesi a latipinna — 1 — — — — — — — 1
Hemiramphidae — — — — — 3 — — — 3
Gerridae 1 — — 1 — 1 — — — 3
Silver jenny, Eucinost omus gula 1 — — — — 1 — — — 2
Coryphaenidae — — — — — — 1 — — 1
Sparidae
Pinfish, Lagodon rhombo ides — — — 1 — — — — — 1
Scaridae 1 — — — — — — — — 1
Callionymidae
Lancer dragonet, Calli onymus bairdi — — — — — 1 — — — 1
Blenniidae — — — — — 1 — — — 1
Gobiidae 1 — —1 5 3 — — — 10
o
00

( I )
TABLE 20— C O N T N U E D
. . . .
Item Dec.-Mar Apr. May June July Aug Sept. Oct N ov Total
, i i
Clown goby M , crogob
i us gulosusi
ll — — — — 3 — — — — 3
Highfin
t goby
i s Gob
. one us ocean cus — — — — 1 — — — — 1
Ba hygob
s us p — — — — 1 — — — — 1
Gobie ocidae — — — 1 — — — — — 1
Atherinidae 1 1 1 2 1 5 — — — 11
Hardhead silverside, A therinomorus stipes — — — 2 — 1 — — 2 5
Unidentifiable fishes 4 6 3 8 1 0 28 2 9 1 1 81
Arthropoda-Crustacea
Grass shrimp, Pa laemonetes 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
 A t Bimini, the food of the smallest barracuda consis ted mainly of
gobiids, whereas those over 3 inches long fed pr imaril y upon schooling
a therin ids and clupeoids . Perhaps this indicates a trans itiona l stage between
niches a n d / o r types of food available . Where surf is absen t, the qu ie t,
shallow-wa ter zone inhab ited by very small barracuda varies from abou t
1 to 12 inches deep, and is often very ri ch i n det rit us which offers
excellen t camouflage for young barracuda. Thi s niche is often ri ch in
gobiids, cypr inodont ids, and small gerrids, but t he abundance of schooling
fishes is no t real ized until somewhat deeper wat er i s encount ered, where
barracuda of slightly l arger si ze are abundant, but are less gregarious t han
in shallower waters .
A partl y digested barracuda which measured abou t 90 mm had been
swallowed tail first by a 291-mm barracuda ; this was the only instance of
cannibalism in the exam ina tion of 901 stomachs .
F ood of Young from A dditi on al A rea s. — The stomach content s of 39
barracuda from museum collections were exami ned by me from various
localities in t he t ropi cal west ern Atl anti c (Tabl e 2 4 ). The food of si x
postl arval barracuda rangi ng from 5.5 t o 12. 1 mm in st andard length,
collected by t he DANA vessels from t he Vi rgin Isl ands, consisted of bits
of fish.

TABLE 21
t f st s t i i f i s i s (l sst
P ercen age o om ach con a n ng ood n 89 y o u n g pec m en e
) f
h a n 300 m m
ll t
o Sphyraena barracuda c o
f i i i,
ec ed rom B m n
s, i l ,
B aham a
,
l ,
n Ju y
. 1955 a n d Jfu iy i1956
i ls
N um ber o
s t i
nd v dua repre en ed n
.
p a r e nt h e se s
t l l l
Fork
l To
. a Tota. Tota t
ength, , nof with no per cen
f
mm July 1955 July, 1956 ood examined with ood
- ( )
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 4 2 .8 ( 1 4 ) 6 7 .5 (19 ) 19 33 57 .6
100-124
- 8 4 7 1( 3 ) 8 4 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 299l —. ( ) —. ( ) — — —.
Tota
. 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 o f stomach s o f 4 4 6 s pecimen s o f Sphyraena barracuda
from Pier 5 , Miami, Florida, which con tained food , taken from August , 1 9 5 5 ,
through Augu st, 1 9 5 6 .

Food of A d u lt s from F lori da .— From August, 1955, t hrough August, 1956,

I examined 446 stomachs of adu lt barracuda . Of these fish, 185 (4 1 .6 % )
con ta ined food (Tab le 2 5 ) .
Mos t fishes taken are assoc ia ted w ith the coral-reef hab ita t (Tab les
26 and 2 7 ) . Fishes accoun ted for 93 .3 per cen t of identifiable food . Of
these 71.1 per cen t were identif iab le a t least to genus.
Where samples were adequa te for compar ison , stomachs taken from
Oc tober through March con ta ined only slightly less food than those taken
in the remain ing mont hs (Fi g. 3 3 ). The smallest young and the l argest
adults more often bad food in thei r stomachs (Fig. 3 4 ) .
F our groups compr ised near ly 64 per cen t of the identifiable fishes
taken by adults . The Be loniformes , mainly hem iramphids , were the mos t

TABLE 22
Su m m a r y o f fo o d it em s f o u n d in s t o m a c hs o f 70 y o ung sp ec im e ns ( l e ss
than 300 m m ) of Sphyraena barracuda c ollec ted f r o m B im in i , B a h a m a s i,
n July , 1955 , a nd J u l y, 1 9 5 6. N u m b e r s a r e st o m a c h s in w h ic h it em s
WERE FOUND .

Group 1955 1956 Total

Gob io ide i 2 6 8
Atherinidae 2 4 6
Gerridae 4 1 5
Cyprinodont iformes — 4 4
Clupeiforme s 2 1 3
Scaridae 1 1 2
Sphyraenidae — 1 1
Opisthognathidae 1 — 1
Uniden tified fishes 8 26 34
Invertebrates ____ 1 1
Plant material 1 — 1
Uniden tified matter 8 4 12
To ta l 29 49 78

110
impor tan t single group, and account ed for 24 ( 2 2 % ) of the i dentifiable
fishes (Tabl e 2 6 ). Fi shes of the order Tet raodonti formes (Pl ect ognat hi )
ranked second i n i mport ance as a group, and occurred in 23 stomachs .
These were followed by carango id and mugilid fishes.
I t was not always possible to distinguish between fish t aken as bait s
and those t aken nat urall y, alt hough in t he former inst ance hook marks
were usually diagnostic. Species of the families Hemi ramphi dae and
Mugil idae are popul ar baits (for sailfish), and those of t he families Carangi -
dae and Scombridae are usually still-fished or drifted live over the reefs.
Of the reef-dwelling te traodon tiform fishes, members of the family
Balistidae were mos t important, and occurred in 13 stomachs ; d iodontids

TABLE 23
It em s o f f o o d f o u n d in th e st om a c hs o f 70 y o ung sp ec im e ns ( l e ss t h a n
300 m m ) of Sphyraena barracuda c ollec ted from B im in i, B a ha m a s , in
J u l y, 1955, a n d J u l y, 1956. N um b er s ar e st om ac hs in w h ic
h i t e m s w e r e f o u n d.

Item 1955 1956 Total

Pisces:
Elopidae
Ladyfish, E lops saurus — 1 1
Clupeidae
Harengu la sp. 1 — 1
Dwar f herring , Jenkinsia lamprotaen ia 1 — 1
Cyprinodontidae
Cyprinodon baconi — 2 2
Poeciliidae
Gambusia (manni?) — 2 2
Gerridae 1 1 2
Silver jenny , Euc inostomus gula 1 — 1
Cuban mojarra, Gerres havana 2 — 2
Scaridae — 1 1
Striped parrotfish, Scarus croicensis 1 — 1
Gobiidae 1 2 3
Ba thygob ius spp. — 3 3
Clown goby, M icrogob ius gulosus 1 — 1
Darter goby, Gob ione llus bo leosoma — 1 1
Opisthognathidae
Mottled jawfish, Opis thogna thus 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 mos t common . To my knowledge , the te traodon tiform
fishes are never used as bait .
The size of t he food ingested oft en bore no rel ati on t o t he si ze of the
barracuda . A barracuda which measured 657 mm had eat en a lizardfish
which was 37 mm long. Remoras of less t han 4 i nches were t aken from
barracuda which were over 3 feet long. Needlefish which were al most
as long as t he barracuda were usually found folded over several t imes
in the stomach of the preda tor .
F ood of A du lt s from B im in i, Bahamas .— Of the stomachs of 132 adults
wh ich were exam ined from Bimini , 78 (5 9 .1 % ) con ta ined food (Tab le
2 8 ) . Barracuda in 1955 con ta ined food more often (7 5 .8 % ) than d id
t hose capt ured in 1956 (5 3 . 5 % ) . Barracuda t aken in 1955 were generally
smal ler and were t aken in shallow-wat er mangrove habit at s; t hose t aken
in 1956 were l arger and were caught in somewhat deeper wa ter and
about oolite reefs.
Fishes accoun ted for 84.2 per cent of the identifiable items found in
64 stomachs of adu lt barracuda taken from Bimini in 1955 and 1956
(Tab le 2 9 ) . The tetraodon tiform fishes were the most impor tan t group,

TABLE 24
F ood f o u n d in st o m a c hs o f 29 y o ung 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 ollec ted f r o m 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
I s l a n d s, a nd B im in i a nd 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.

N o . o f stomachs
with item
Clupeidae
Harengula sp. 1
Belonidae
S trongylura sp. 1
Cyprinodontidae
Striped killifish, Fundulus majalis 1
Poeciliidae
Gambusia sp.
Gerridae 1
Sparidae 1
Gobiidae 2
Blenniidae 1
Atherinidae
f ll tt
Ree silverside, A ane a harringtonensis
i ti 1
Hardhead silverside, Ather nomorus s pes 1
Mugilidae t s 1
Uniden ifiable fi hes 14
Arthropoda
Insecta, Blattidae 1
Plant material
i
Ang osperm flower 1
l 29
Tota
112
 1 — '— I— '— I— •— I— '— I— I—
0- 200- 400- 600- 800" 1000- 1200
99 299 499 699 8 99 1099 1299
FO R K L E N G T H , M M .
F i g u r e 34. Percentage o f stomachs of 641 specimens of Sphyraena barracuda
from southern Florida, arranged by size group, which contained f ood. Fish
less than 300 mm long were collect ed from 1945 through 1956; fis h longer
than 300 mm were taken from August , 1955 , through August , 1956 .

with the puffer occurri ng most frequently. Carangids, princi pal ly the bar
jack and bigeye scad , ranked second ; parro tfishes were third .
Compari sons and Discussion. — The lit erature on the food habits of
sphyraen ids conta ins mostly random observa tions , so tha t it is difficult
to draw conclusi ons about the relative i mport ance of cert ai n items as food.
Bean (1890) no ted t ha t barracuda from Yuca tan fed upon schools of
Harengula and Stolephorus. Legand (1952) observed tha t in New Ca ledo-
nia, S. p icuda ( — S. barracuda) and S. je llo fed upon schools of A therina
forskali. He added t hat bot h species fed upon mullet ( Mugil sp. ) which
frequent ed the mangrove habit at. Mowbray (1 922) not ed t hat barracuda
fed upon schools of mullet, when “barracuda are seen in the great est
numbers .” Breder (1927) found a 10-mm barracuda in the stomach of
a l arger barracuda, and t hree in the stomach of ano ther i ndiv idual taken
from Cuba. He also recorded Synodus and a myrid eel. Beebe (1928)
observed a barracuda feeding upon a fairy bass le t ( Gramma sp .) in
Bermuda waters , bu t such instances of a barracuda feeding upon corallico-
lous fishes are rare . Ommanney (1953) no ted tha t five of 12 stomachs of
S. commerson i ( = S . barracuda ) from the Seychelles Islands conta ined
specimens of Le thr inus ramak , and ano ther con ta ined an eel. Williams
(1956) work ing in the same region and upon the same species , collected
mostly fishes, i ncluding a beloni d, a l utjani d, a scombrid, and a clupeid;
stoma topods ( Squilla sp .) and squid were also found.
Longley and Hil debrand (1941: 69-71) observed t hat on occasion
squid were eaten by barracuda i n t he Tortugas, Fl ori da, region. Malpas
(1926) found small fishes, as well as a cephal opod (Sepi a), in the stomach
of Sphyraena acuti pinnis, and two specimens of Bali st es miti s in specimens
of S. ob tusa ta from Ceylon. Chacko (1949) also found Sep ia in the
113
 TABLE 25
P e r c e ntage of st o m a c h s c o n t a in ing fo o d in 446 adult sp ec im e ns o f Sphyraena
barracuda c oll ec ted f r o m P ie r 5, M ia m i, F l o r id a , from A ug ust, 1955, t h r o u g h
A u g u s t, 1956. N um b er of in d i v 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.
l t ,
Fork . t. t. . . . . .
eng h
mm Aug Sep Oc 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 50 4 66.7 (3) 42.9 (7)
1100-1199 — 100 2 — 0 ( (1) — .
1 100 1 0 2 33 3 3
1200 1299 100 — — — — — — —
t l ( ) ( ) ( ) ( ) ( ) . ( ) ( ) ( )
To a 71.4 14 42.4 33 34.5 29 50 36 32.5 26 46 7 30 42.9 14 44 25
f it s
No.. ost em s 10 15 10 23 14 14 6 13
No it omach
f
w h ood 10 14 10 18 8 14 6 11

Fork
l t , . it
eng h . l . Noi %w h
mm - Apr May June Ju y Aug exam ned food
( )
300 399 — — ( ) — 100 ( )
(1) —. ( ) 1 100.
400-499
- — 50 2 —. ( ) 0 (1) 12 5 (16) 29 31.1
500-599 — — () 33 3 (3) 0 (3) 30. (10) 41 29.3
600 699 — 50 (2) 0 (1) 25 (8) 31 3 16 64 39 1
700-799
- — ( ) 75 (4) 42.8 (7) 25 (8 ) 41.2 ((34 )) 94 40.4
800-899 50 4 75 4 50 8) 22.2 9 45 20 89 43.8
900-999 100. ((3)) 33.3 ((3)) 72.8(11 0 (1) 40 (25) 63 49.2
1000-1099 66 7 (3) 25 (4) 50 (4))
( 50 (4) 54 5 11 )
. ( 47 53.2
1100-1199 0 3 0 2 50 2 — 0 (1 ) 17 29.4
(100 )
1200 1299 100 (1) — — — — 1
Total 57.2(14) 47.6(21) 52.7(36) 38.9(35) 36.9(133) 446 41.6
No. o f items 12 10 25 12 57 221 —
No. stomachs
with food 8 10 19 9 49 186 —

stomachs of 51 specimens of S. ob tusa ta , which ranged in length from

17 to 45 mm, from Ind ian waters ; clupeids, atherin ids, mugilids, and
lep tocephalus larvae were also eaten . Inger (1955) collected a gobiid
and a percoid (A m bassis) from a 400-mm specimen of S. je llo from
Borneo. Ah Kow (1950) found anchovies ( St ol ephorus sp. ) t o be a mai n
food item in the di et of Sphyraena j ell o from the Si ngapore Straits during
January , Oc tober, and December; o ther food taken included squids and the
larvae of decapod crus taceans . Rough ley (1916) sta ted tha t in Aus tra lia
S. novaeholl andi ae . . plays havoc with . . . garfish [Hemi ramphus],
yellowtail [Trachurus], and mulle t [M u g ill ” Gudger (1918) found 5
114
 TABLE 26
Su m m ar y of f o od it em s in st om a c hs of 186 a dult s p e c im e ns of Sphyraena barracuda c ollec ted from P ie r 5,
M ia m i, F l o r id a , from A ug ust , 1955, th r o ug h A ug ust , 1956 . N u m b e r s ar e st o m a c h s in w h ic h it em 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

Clupeiforme s 1 2 4 — 1 8
Scombridae 2 2 — 3 7
Pomadasyidae 1 1 — — — — — 2 — — 1 — — 5
Anguilliforme s — — 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 c ollec te d at P ie r 5, M ia m i ,
F l o r i d 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 em s wer e f o 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
Atlan tic round herring,
E trumeus sadina — — — — — — — — —- 2 — — — 2
Harengula sp. — — — — — — — — — — 3 — — 3
Synodontidae
Inshore lizardfish,
Synodus foet ens — — — — — — — — — — — — 1 1
Muraenidae
Gymno thorax sp. — — — — 1 — — — — — — — — 1
Ophichthyidae — — 1 — — — — — — — — — — 1
Sharptail eel,
M yrich thys acum inatus — — 1 — — — — — — — — — 1 2
Belonidae
Houndfish ,
S trongylura raph idoma — — — — — — — 1 1 — — — — 2
S trongylura sp. — — — — — — — — 1 — — — 2 3
Fla t needlefish, Ab lennes hians — — — 1 — — — — — — — — — 1
Hemiramphidae — — — — 1 2 — 2 — 1 — — 1 7
Ballyhoo ,
Hem iramphus brasiliensis — — 2 3 — — 1 1 1 — 1 1 — 10
Exocoetidae — — — — — — — — — — — 1 — 1
Serranidae — — — — — — — — — — 1 — — 1
Ep inephelus sp. — — — — — — — — 1 — — — — 1
Sand perch,
D ip lec trum formosum — — — — — — — — — — 1 — — 1
 TABLE 27— (C O N T IN U E D )

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

Lutjanidae — — — — — — — — — — — — 1 1
Carangidae — — 1 — — — — 1 1 — — — 4 7
Rough scad, Trachurus latham i — 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, Aux i s thazard — — — — — — — — — — — — 1 1
Ophidiidae
Short-bearded cusk eel,
Lepoph id ium brev ibarbe — — — — — — 1 — — — — — — 1
Nome idae
Man-of-war fish,
N om eus gronowi — — — — — — 1 — — — — — — 1
Mugilidae
117Mullet, Mug il spp. — 1 1 1 3 — — 1 — — 1 4 — 12
Bothidae — — — 1 — — — — — — — — —
1
 TABLE 27— ( C O N T IN U E D )
00
Item Aug. Sept. Oct. Nov . Dec . Jan. Feb. Mar. Apr. May June July Aug. Total

Echeneidae
Marlinsucker,
Remora osteoch ir
Balistidae
Orange filefish,
A lu tera schoepfi
Gray triggerfish,
Batistes capriscus
Ocean triggerfish,
Can th iderm is sufflamen
Tetraodontidae
Diodontidae
Striped burrfish,
Ch ilomyc terus schoepfi
Porcupinefish, D iodon hystrix
Unidentifiable fishes
Reptilia-Chelonia
Mollusca-Cephalopoda
Echinodermata-Ophiuroidea
Basket star,
A s trophy ton muricatum 1 1
Plant material
Turtle gra 1 1
ss, estudinum
Thalassia t
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 fragmen ts in the stomach of a 55-inch
barracuda from Tor tugas , inc luding ha lf of a margate (Haemu lon a lbum) .
Beebe and Tee-Van (1 928) no ted a 55-mm parro tf ish (Scarus cro icens is)
in the stomach of a 200-mm barracuda from Ha iti, an 80-mm specimen
of Chl oroscombrus chrysurus in a 400-mm barracuda , and a 70-mm
specimen of Ocyurus chrysurus in a 697-mm barracuda .
In the western Nor th A tlan tic, Sphyraena boreali s, a small species, has
been found to con ta in young fish and young gas tropods (Summer , e t al.,
1913), and shrimp (L in ton , 1905) . A ther in ids and mysids are the
princi pal food of juveniles of borea lis t aken from the Del aware R iver
estuary and adj acent areas (de Sylva, et al ., 1962).
Of the 901 barracuda of all sizes exam ined by me during the presen t
inves tiga tion from various l ocalities in the t ropi cal west ern Atl anti c, 529
(5 8 .7 % ) con ta ined food . F ishes were found i n 82. 2 per cent of these
stomachs , p lan t ma ter ial in 2.8 per cent, i nvert ebrat es in 2. 6 per cent,
and un iden tif iab le ma tter in 12.2 per cent. Gobii d and at heri ni d fishes
ranked first and second as food of t he young in bot h t he Bimini and t he
F lori da region. Gerri ds ranked t hi rd i n Bimini, whereas t hey were sixth in
i
mpor t
ance in F lor ida . Cypr inodon ts .were fourt h in Bimini, but seventh
in F lori da waters. Invert ebrat es, which ranked fift h in Fl ori da, were t ent h
in i mport ance i n Bimini. Pl ant mat eri al occurred fewer t imes in Fl ori da
t
han in Bimini.
Bel oniformes were more i mportan t in the diet of adult barracuda from
F lor ida than any o ther ma jor group , bu t only ranked four th in Bimini .
The Te traodonti formes ranked first in Bimini , and second in F lor ida as
food. Carang ids ranked second and th ird from Bimin i and F lor ida ,
respectively , bu t scar ids were of considerab ly grea ter impor tance in
Bimini than in F lorida. Mugilids, which were so commonly found in
F lor ida barracuda , occurred only once in 38 stomachs a t Bimini , and
a lthough mulle t are compara tive ly scarce in Bimini, the ir abundance in
barracuda from F lor ida seems to reflect the ir wide use as ba it. This
seems to be true also of the Be loniformes and the Carang idae , which
were so commonly found in the F lor ida barracuda. Small scombrids and
pomadasy ids , which figured prom inen tly in the d ie t of the F lor ida fish , did
no t occur in those from Bimini .
The differences in the type of food taken by adults in F lorida and
Bimini would seem to reflect a difference in hab it at s, rat her t han food
preferences of the barracuda . The wa ters of sou theas tern F lor ida suppor t
luxur ani t growths of massive and branch ng i cora sl whereas Bimin i si
surrounded by extensive sandy stretches which are less conducive as
subs tra te to the grow th of cora l reefs (Parr , 1933 ; Kuenen , 1950 ; Newell
and Imbr ie , 1955) .
A definite change occurs in the type of food ea ten with a change in
size (Tab le 3 0 ) . A ther in ids , clupeo ids, gerrids, and cyprinodon tids occurred
in 76 .2 per cen t of the stomachs of the young. These same groups accoun ted
119
 TABLE 28
P e r c e ntag e of st o m a c h s c o n t a in in g fo o d in 132 a dul t sp e c im e ns of
Sphyraena barracuda c ollec ted f r o m B im in i , B aham as , in J u l y, 1955, and
Ju 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 i n p a r e n t h e s e s.

Fork
length,
mm 1955 1956 Total
300-399 7 1 .5 (1 4 ) 53 ( 1 7 ) 61.3 (31 )
400-499 50 (4 ) 50 (6 ) 50 (1 0 )
500-599 100 (4 ) 75 (4) 87.5 (8)
600-699 100 (3 ) 100 (4 ) 100 (7 )
700-799 100 (3 ) 30 ( 2 0 ) 43 .5 (23 )
800-899 33.3 ( 3 ) 6 4.7 (1 7 ) 60 (2 0 )
900-999 100 (1 ) 60 . 9 ( 2 3 ) 62 .5 (2 4 )
1000- 1099 — 50 ( 6 ) 50 ( 6)
1100- 1199 — 0 ( 2) 0 ( 2)
1200-1299 100 (1 ) 100 (1 )
Total 7 5 .8 (3 3 ) 5 3.5 (9 9 ) 59 .1 ( 1 3 2 )

TABLE 29
F oo d it e m s f o u n d in the 78
st om a c hs o f a du l t sp e c im e ns o f Sphyraena
barracuda c ollec ted f r o m B i m i, n i B a h a m, a s in J u, l y 1955
, a nd J u, l y .
1956 .
N um bers ar e st om a c hs in w h ic h it e m s wer e fo und
I ( ) ( ) t l
tem 1955 N = 2 5 1956 N = 5 3 To a
:
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
Holocen tr idae
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 gra, Cymodocea manatorum 1 — 1
Unidentifiable ss,
matter 2 7 9
Total 29 56 85

for only 5.3 per cen of he food of he adu barracuda In adults he

t t t lt . , t
Beloniformes , Te tradon tiformes , Carang idae , Mugil idae , and miscellaneous
Perc iformes are more impor tan t and accoun ted for 66 .7 per cen t of the
food . However , these groups are presen t in only 6.5 per cen t of the
stomachs of the young. These changes in d ie t reflec t in par t a change in
hab ita t with grow th of the barracuda . The adu lt barracuda shows definite
preferences for two ma in categories of fishes: compara tive ly fast-swimming
fishes of the surface or mid-dep ths , such as hem iramph ids , carangids , and
scombrids ; and the sluggish, reef-dwelling te traodon tiform fishes . The
scarcity or absence of such active, reef-dwelling forms as lutjan ids ,
serran ids , labr ids , and pomacen tr ids in the d ie t seems no tewor thy .
From a survey of the litera ture , and based on persona l observa tions ,
it appears that, in general, sphyraeni ds which do not att ai n a l arge si ze
feed mai nly on small fishes which school, and upon cephal opods, whereas
he
t large species of Sphyraen idae which tend t o be solit ary feed upon
arger
l fishes which are essentially non-schoo ing l in na ure
t .
A ttac ks by B a r r a c uda on H u m a ns
Perhaps t he mos t colorfu l accoun ts of barracuda are those which involve
he
t ir alleged vorac ity . Of the early repor ts by explorers and na urat lis st , de
Rochefor t (16 6 7 ) appears t o be one of t he first t o record t he barracuda’s
ferocity , and he believed tha t in add ition to its desire for human f lesh ,
its tee th con ta ined po ison . Gudger (1 918) gave an excellent and amusing
summary of the popu lar beliefs , citing the works of de Rochefor t (1 6 6 7 ) ,
121
 TABLE 30
Su m m ar y of fo o d it em s f o u n d in s t om a c hs o f 529 sp ec im e ns of Sphyraena bar-
arcuda from 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 1956 ,
th r o ug h
SHOWING CHANGES IN FOOD H A B I T S W ITH GROWTH. N U M B E R S ARE STOMACHS I N
WHICH ITEMS WERE FOUND.

Fork length (mm)

0 - 100- 200- 300- 400- 500- 600- 700- 800- 900- 1000- 11GO- 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
A her n dae 17 11 — 2 30
Gerr
t dae i i 3 5 3 1 12
C upei orme 14 5 — 1 2 — 1 1 — 3 — — — 27
Scar
l dae if s — 2 1 2 1 2 — 1 1 1 1 — — 12
Bo hi dae — — — — — — 2 1 — — — — — 3
Betoniforme
i 4 — — — 3 3 9 4 3 4 2 1 — 33
Telraodon s
t orme - 3 4 5 5 9 X 27
Angutif li orme s 2 3 2 —i — — 7
Carango il f de s 1 — — — — 1 2 4 4 6 4 — 2 24
Scombr dae i i 3 2 2 1 — — 8
Mug dae i 1 2 3 3 1 2 — — 12
O herili he 5 3 — — 1 — 2 4 7 2 3 1 1 29
Unt den fis s ab e
he tifi l
ihropoda 106 26 3 7 7 17 9 14 10 6 205
Arfis s
Cru acea 3 1 2 6
Mot u stca _ _
Cepha opoda 2 _ _ _
1 2 1 1 7
P anll sma er a 3 — — 3 1 1 1 5 2 — — — — 16
Un den ab e l
l mat er t i l 7 11 1 5 4 4 4 6 11 8 6 2 69
To ia tttifi l 184 82 8 21 12 21 44 49 57 51 26 5 3 563
t l
Du Ter re (1 6 6 7 ) Sloane (1 7 0 7 ) Damp er (1 7 2 9 ) L aba (1 7 4 2 )
Ca esby t (1 7 5 4 ) and , Ferm n (1 7 6 9, ) i , t ,
A
t acks have ,been au heni ca ed by. the unm s akab e oo h marks of
a barracuda
tt As Gudger t andti Bredert (1928) po in ted out, l t its t cleaver-like
bite can be .dist nguished from ha of a shark in i t ha it consists of two
near y para e irows of oo h marks t t whereas hat of t a shark leaves a,
parabo
l c woundll l of jaggedt t oo h marks , The shark
t t also ends o clamp
its preyli and ear off flesh with t ta sawing . motion Pho ographs t of t a barra-
cuda’s b te are t given in Gudger and Breder .(1 928) t and Tanous and
ano (1 i9 6 0 ) and ha of a shark n Bolin (1 9 5 4 )
ll My research , andt observa
t ions ndicate
i ha he . barracuda depends
almos entire y upon sight and t usually
i makest at single
t a ack wh ch may
be severet bu l s seldom fa al. In general, a shark s fur her
tt s mulatedi by
the presence t of i b ood drawn t so ha repea ed a acks i occur
t tiwhich often
resu in fatalitiesl t t t tt
L eralt ure R ev ew . of A a c k s — The first au hen ca ed repor by a barra-
cuda
it upont a human
i appears
tt .to be ha of P t keti(1873 t 286t 2 9 4 ) who
122 t t i : , ,
rela ted persona l experiences on Mauriti us; he was att acked by a large
barracuda which he was able t o harpoon aft er a second attack. Gunt her
(1 8 7 7 ) , on the au thor ity of Andrew Garrett, wrote t hat barracuda were
dangerous to bathers in the Indo-Pacific. Klunzinger (1884) reported tha t
a native diver in the Red Sea died of wounds aft er having been att acked
by a specimen of Sphyraena agam ( —S. barracuda ) . A native of Cocos-
Keeling At oll whose calf muscles had been t orn away by a barracuda
was observed by Wood-Jones (1902: 322ff). Hol der (1908, 1924) stat ed
tha t in the F lor ida Keys a man was a ttacked repeat edl y by several barra-
cuda . C iting ano t her i nst ance, he not ed t hat a barracuda, which had been
p laced in the well of a sloop and forgot ten, att acked and mai med a man
who en tered it later. Ho lder also men tioned an episode a t Sea Horse Key,
Bahamas , where a man suffered a severe bi te on t he arm while swimming.
Gudger (1918) report ed the case of a man having been bitt en on the
foot by a barracuda as he was walking on a shallow reef beside deep
wa ter. Mowbray (1922) s ta ted tha t a young woman while approach ing a
buoy when swimming in Fl ori da wat ers was att acked by a barracuda and
b led to death. Waite (1923) remarked t hat in Sout h Ameri ca a champi on
lady swimmer lost her leg and d ied as the resu lt of an att ack by a large
barracuda . Two young girls were att acked by barracuda while swimming
off Miami Beach, Fl ori da, according to Forbi n (1 9 2 4 ). l
A valuab e det ail ed and i llust rat ed report was made by Gudger and
Breder (1 928) of an a ttack by a barracuda upon a man swimming in
shallow wa ter on the A tlan tic side of Panama . They also referred to a
young woman who was fa tally a ttacked by a barracuda off St. Petersburg ,
F lorida, some years pr ior to 1928. If the locality is correc t, this repor t
differs from that report ed by Waite (1923) or Forbi n (1 9 2 4 ), although
it may be identical with tha t report ed by Mowbray (1 9 2 2 ) . Irvine (1947)
gave an accoun t of a man who lost bo th hands af ter he was b itten by a
barracuda off Ghana . In popu lar accoun t, Schultz and S tern (1948) no ted
t t “barracuda have r pped
ha i off the calves of swimmers [and] have
ampu ta ted legs and arms with a single b ite .” S torr (1957) repor ted tha t
he was a ttacked by a barracuda which had apparen tly become irr ita ted by
t white shoes of ano her
he t diver. Accord ng i ot S orr t (persona l commun -i
ti , t tt tt
ca on) he a ack pa ern was no comp eted t l .
Coppl eson (1 9 5 9 ), in his study of shark att acks in Fl ori da wat ers, listed
t tt i t t ,
two fa alities a r bu ed o barracuda bites one in 1947 upon a swimmer
t t. t , t l t
a S Augus ine and a second a Key West in 1952. Copp eson repor ed
tt , t ,
an a ack on a swimmer off Durban Sou h Africa which was believed to
. t l tt t t
be by a barracuda A fa a a ack on a ba her in 1958 a Key West was
t tt . t
repor ed by Hu on (1 959) as possibly due to a barracuda A four h
tt , t l, l tt .
a ack non-fa a was isted by Hu on as occurring in 1953 off Juno Beach
ll t i
Tanous
t and anol (1 i 960) . presen
t i ed case i h storiest ofi four attacks
, by
preda
t t ory fishes in F or dat From he r descr, ptions of hei njuries I believe
ha these were all due o barracuda bites I have descr bed two of these
123
 in some detail in the following section. Of t he remai nder, one att ack
occurred in t he vicinity of Miami Beach i n Sept ember, 1949, and was
inflicted upon the leg of a 37-year-o ld woman while she was swimming in
t af ternoon . The o ther case involved a 49-year-o l d man who was bi tten
he
on the foot at approxi mat el y 6: 45 PM in August, 1955, in t he vicinity
of Miami Beach.
On May 7, 1953, t he Miami (Fl ori da) Heral d report ed t hat a boy was
b itten on the foo t while swimming close to a school of mulle t abou t 50 to
75 feet off Hau lover Beach , Nor th Miami. Severa l stitches were requ ired
to close five or six deep cuts. It was report ed t hat t he att acker was a
barracuda (Tanous and llano , 1960) .
The Miami Heral d report ed on May 8, 1955, t hat a 15-year-ol d boy
was att acked underwat er by a 4-foot barracuda which he had speared
off Crandon Park , Miami. Af ter dragging him a few fee t through the
water, the barracuda broke loose from the spear and att acked his right
side, leaving four large too th marks which requ ired 23 stitches .
On September 18, 1955 , according to Mr. Tom Morgan (persona l
commun ica tion) of the Co llier Coun ty News, Nap les, F lori da, a young
girl was severely b itten abou t the arm and fingers in Ba teman Cove on
t south side of Marco Pass by a f sh
he i which may have been a barracuda .
The Miami Heral d carri ed an arti cl e on July 16, 1956, about a
19-year-o ld girl who was a ttacked while floating in an inner tube abou t
15 miles nort h of Myrtl e Beach, Sout h Carolina. The physici an who
trea ted her repu ted ly descr ibed the in jury as a “multiple , tear ing wound
of the lower lef t leg, ankle and foot, severe and extens ive,” which in his
op inion was due to a barracuda ra ther than a shark .
The Miami Heral d for Jul y 19, 1956, pri nt ed a similar story in which
a 38-year-o ld woman was “ . . . slashed repea ted ly and savagely by a
barracuda in the shallow surf off . . . Miami Beach .” She was b itten six
t imes on her left leg, officials said, and once slightly on the calf of her
r ieh t leg. The victim suffered from shock and loss of bl ood, and was
subsequen tly hospitalized . The att endi ng physici an believed t hat t he bite
was t ha t of a barracuda and not of a shark. There were conflicting report s
on the number of a ttackers (Tanous and llano , 1960) .
The New York Heral d Tri bune of July 17, 1957, report ed t hat “a shark
or barracuda att acked and killed a swimmer . . . ” off Atl anti c Beach,
Nort h Caro li na, on July 16, 1957. The victim, who was an except ional ly
good swimmer, shouted t o a li feguard t hat a shark had bitt en him.
Subsequen t exam inati on of the wounds by i cht hyologists and physici ans did
no t resolve the iden tity of the assa ilan t.
Dr. John E. Randall, present ly of t he Uni versi ty of Puert o Ri co,
informed me tha t on March 5, 1957 , a man speared a 5-foot barracuda
south of Key Bi scayne, Fl ori da. The fish broke t he line and t urned to
a ttack , at which poi nt the man rai sed his ri ght arm t o prot ect himself,
with the resu lt tha t he was slashed deeply on the forearm , requ ir ing abou t
124
 70 stitches. Ano t her att ack was report ed by Dr. Randall as follows: in
Augus t, 1926, a man was swimming about 50 yards off Miami Beach, in
murky wa ter which was over his head. The man sustai ned t hree bites on
his left arm, which was subsequent ly amput at ed. The surgeon removed
barracuda tee th from the wounds .
The Tampa Tribune carr ied an art ic le in May 3, 1959 , repor ting an
instance off Miam i where a barracuda leaped from the wa ter and b it a
3-inch gash in the neck of a fisherman who was sitting in a small boa t,
with the result t hat six stitches were requi red for t he wound. No additional
details were repor ted .
On Jul y 15, 1960, t he Miami Heral d report ed t hat a 34-year-o ld skin
di ver was att acked twice by a barracuda about Va -mile off Pompano
Beach , F lori da, in about 20 feet of wat er. The 15-pound barracuda made
four rushes a t the diver , two of which ended in attacks, resul ting in severe
lacera tions of the knee and wri st which requi red 31 stitches. The diver
wore no bright ly colored apparel or jewelry, nor had he aggravated the
barracuda in any way. Ne it her t he barracuda nor ot her fish had been
speared . Visibility was descr ibed as “fa ir .”
From t he lit erat ure and newspaper accounts I have listed 29 separate
a acks att ri but ed t o barracuda upon humans ; 19 are documented, two
tt
are probabl e, and t he remaining eight are possibly due to barracuda. The
report s of Pi ke (1 8 7 3 ), Kl unzinger (1 8 8 4 ), Wood-Jones (1 9 0 2 ) , Ho lder
(1908, 1924), Gudger (1 9 1 8 ), Gudger and Breder (1 9 2 8 ), Irvi ne
(1 9 4 7 ), Coppl eson (1 9 5 9 ) , H utt on (1 9 5 9 ) , and Tanous and ll ano
(1960) cont ai n evidence as t o t he i dent ity of t he att acker. Personal report s
from Randall, and newspaper account s of May 7, 1953, off Haul over
Beach, Miami, t hat of May 8, 1955, of the assault off Miami, t hat of July
19, 1956, on t he att ack off Miami Beach, and t hat of July 15, 1960, off
Pompano Beach are due to barracuda. The report on the att ack of July
16 , 1956 , off Sou th Caro lina , and tha t off Durban , Sou th Africa , in
1946 report ed by Copp leson (1 959) were probab ly made by barracuda .
One of t he att acks (1 952) report ed by Coppl eson and one (1958) by
Hu tton were probab ly made by barracuda. The i dent ity of t he species off
Nap les , F lor ida , on Sept ember 18, 1955, i s questionable; t he nat ure of
the wound and the single att ack suggest t hat it was a barracuda. Lit erat ure
repor ts of Mowbray (1 9 2 2 ), Waite (1 9 2 3 ), Forbi n (1 9 2 4 ), and one of
the cases in Gudger and Breder (1 928) are lacki ng in detai ls and locality
of t he attack. Accordi ng t o dat a present ed by Coppl eson (1 9 5 9 ), possibly
these may refer to the same att ack; it i s debat abl e t hat they were due t o
barracuda . The identity of the att acker in t he fat ality off Nort h Caroli na
on July 16, 1957, is questionable.
In t he t ropi cal west ern Atl anti c, barracuda i s the only sphvraeni d which
a ains sufficient si ze t o inflict a wound and it probabl y made most of t he
tt
a ttacks listed above. In ot her areas, it i s possibl e t o i dentify t he att acker
i i , . , . j ll i . j t i
on an a pr or basis since only S barracuda S e o Cuv er, S ors er
125
Valenciennes, and S. obt usat a Cuvi er att ai n appreci abl e si ze. In general,
the d is tr ibu tions of these large species do no t overlap .
Poss ib le Causes of A tta c k s .— An interes ting accoun t of a group of abou t
40 British frogmen and t hei r experiences with sharks and barracuda i n
the Bahamas is given by Wright (1 9 4 8 ), who stat ed t hat the barracuda i s
feared by the natives because it can come into much shallower wat er t han
can a large shark . He concluded t hat under cert ai n conditions barracuda
and sharks should be feared, but t hat normal ly they were harmless. His
inferences were as follows:
“ (1 ) A man in the wa ter, alone or in a group , acting quietly and
smoothly , did no t release the a ttack pa ttern of e ither shark or barracuda .
(2 ) B lood w ithou t movemen t did no t release a feed ing pa ttern in shark
or barracuda in the Bahamas . (3 ) F ish b lood and jerky , rap id movements
re leased the a ttack pa ttern in bo th shark and barracuda . (4 ) Jerky , rap id
movements alone re leased the a ttack pa ttern in bo th shark and barracuda .”
The frenzied reacti on of a barracuda t o a vi brati ng or bl eedi ng fi sh on
the end of a spear i s a rat her well-known but unnervi ng experience t o
spearfishermen . Wright (1948) report ed a case where a barracuda rushed
in and seized a small, bl eedi ng sting ray and j erked the ray and spear
ou t of the m an’s hand and made off with bo th . Off Ca t Cay, Bahamas ,
I speared a Nassau grouper (Ep inephe lus s tria tus) which subsequen tly
wrenched the spear from my hand and became wedged under a rock
ledge. During my efforts to dislodge it, the grouper struggled violently
and em itted croak ing sounds , and short ly four barracuda from 3 to 4
feet long appeared in the vicinity. When dislodged the grouper was
bl eeding profusely and swimming erraticall y from the weight of the spear,
which was s til l 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 F lor ida Keys, while collecting specimens I have no ted
tha t parro tf ish , groupers , and grunts , in par ticu lar , all flap rap id ly and
b leed profusely when speared , and groupers and grunts also emit croaki ng
sounds , and often barracuda are att ract ed; however, only in the instance
descr ibed above was the specimen carri ed off. An i nj ured fish or live
grun t with severa l d iagona l cuts is a well-known ba it for large barracuda
in the F lor ida Keys.
A flashing obj ect i s quickly di scerned by a barracuda; hence, in wat ers
i it i l t
nhab ed by barracuda it s advisab e o remove shiny objects such as a
, l t, i t . i , t t i t
ring brace e or wr stwa ch On one occas on in ra her urb d wa er off
, l i , t t t
Key Largo F or da a barracuda abou 2 fee long charged a and hit
t
my face mask which may have reflected the sun’s rays. In ano her
t t i l ti i t
ins ance I was pho ograph ng some cora forma ons n abou 10 feet of
t l t
wa er using a camera enclosed in a ucite case when a barracuda abou
t . t t t
3 V2 fee long appeared I approached suddenly to within abou 4 fee of
t t i t
the camera buti soon ll backed
t off o a d stance of abou 15 feet tt and . began
to swim erra ca y a first and eventually in a figure-8 pa ern As the
126
camera was poi nt ed downward, the barracuda showed no excitement,
but each time the camera case was di rected t oward the fish it showed
interes t. On severa l occasions it approached the camera rapi dl y but t urned
sharp ly away and swam in a wide circle. Its behavi or was simi lar t o
tha t of a barracuda confron ted with a speared f sh i .
On about 40 occasions when swimming abou t Bimin i and Ca t Cay,
Bahamas, and in the Fl ori da Keys, I encount ered barracuda on nearly
every trip. They ranged in si ze to perhaps 5 feet, but on no occasi on was
I ever attacked . However, on any occasi on it was possible t o elicit an att ack
pa tt ern by spear ing a small grun t or parro tfish when barracuda were close
by.
If present in the area, barracuda congregated immed iate ly upon one’s
ent rance into the wat er, parti cul arl y if it was accompani ed by splashing ,
t l t t i .
bu short y hereaf er they usually d sappeared Occasionally a stray would
be encountered and would follow a t a d istance of perhaps only 10 or
15 feet as I moved from one reef to the next .
Based on lit erature and newspaper repor ts , as well as persona l experi-
ences , there is only one instance known in which an unprovoked , in the
str ictest sense, barracuda has a ttacked a swimmer who was completely
submerged (Ju ly 14 , 1960) . In the May 8 , 1955 repor t of an a ttack on
a 15-year-old boy off Crandon Park , Miami, and the sim ilar a ttack on
the man off Key B iscayne in 1957 the a ttacks were made by an in jured
barracuda. Numerous barracuda are speared in the F lor ida Keys annually ,
bu t few, i f any, att empt to att ack t he diver. Miss Kat hl een Parker
(persona l commun ica tion) informed me tha t a barracuda slid down a
spear and made a slash in a boy’s leg which requ ired 12 stitches. A lthough
I have never speared a barracuda weighing over 8 pounds , divers who
have taken t hem up to 50 pounds i nformed me t hat they were never
menaced or a ttacked af ter the fish was speared .
There are 15 separat e instances where barracuda were definitely or
probabl y i nvolved in att acks where the swimmer was splashing at or near
the surface , swimming in turb id wa ters , swimming t oward dusk when
visibility is reduced, or a combi nati on of these fact ors. If t he “possibl e”
a tt acks are added, t hen anot her seven att acks can be considered as resul ting
from splashing or swimming. Such attacks, although they might be
considered ones of “mi st aken i dentit y,” probabl y should be considered as
being provoked in the sense t hat t he victim was splashi ng or flashing
suddenly under conditions which elicited an att ack patt ern. Probabl y
barracuda are less able to discern t hei r prey in t urbi d wat ers, and in such
situations the risk of a ttack is h igher than in c lear waters .
In summary, conditions which appear favorabl e t o an att ack by an
unprovoked barracuda are: (1 ) i f the swimmer were splashing at the
surface of the wat er; (2 ) i f the wat er were t urbi d; or (3 ) a combi nation
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 of H um a ns by B arracuda
F ish and Cobb (1954) wrot e t hat t here i s probabl y more controversy
over the toxic properti es of t he family Sphyraeni dae t han in any ot her
group of poi sonous fishes. Some specimens of barracuda are poi sonous,
while o thers are safe t o eat, accordi ng to where and when t hey are caught,
and t he size of the fish. Since poi soni ngs can occur from eating a freshly
caugh t barracuda , the toxin is evidently not rel at ed t o bact eri al decompo-
sition , and is probabl y associ ated with cert ai n fact ors in its life hi st ory
and ecology. Thus it seems apropos t o investigate some of these fact ors
which may be respons ible for its occasionally po isonous na ture .
References to poisonings by the barracuda which date back several
hundred years are found in papers by Gudger (1918 , 1930) . The subjec t
of poisonous aquati c organisms has been reviewed in some detail by the
wr iter in his doct oral di ssert ati on (de Sylva, 1958) on fil e in the Cornell
Un ivers ity library , and the discussion here in will be limited essentially to
the po isonous proper ties of barracuda and its food .
G en era l R e v i e w o f P o i son ou s F i sh es.— Resumes on the historical back-
ground of po isonous fishes are found in the works of Moreau de Jonnes
(1 8 2 1 ) , Dumer il (1 8 6 6 ) , Poey (1 8 6 6 ) , Pe llegrin (1 8 9 9 ) , Byam and
Arch ibald (1 9 2 1 ) , Phisalix-P icot (1 9 2 2 ) , Hoffmann (1927 , e t s e q . ) ,
Pavlovski i (1 9 2 7 ) , Ma tsuo (1 9 3 4 ) , Maass (1 9 3 7 ) , H iyama (1 9 4 3 ) ,
Ha ls tead (1 9 5 3 ) , Phillips and Brady (1 9 5 3 ) , Ha ls tead and Lively
(1 9 5 4 ) , Whee ler (1 9 5 4 ) , F ish and Cobb (1 9 5 4 ) , Lo ison (1 9 5 5 ) , Jones
(1 9 5 6 ) , Mills (1 9 5 6 ) , Ebro in (1 9 5 6 ) , Randa ll (1 9 5 8 ) , and Bartsch , e t
. t i
al. (1 9 5 9 ) Cer a n groups of fishes are more often repor ed o be po -
t t i
t . l i
sonous when ea en These include arge indiv duals of he phv o^ene ically t l t
l t i , i , i , tj i ,
unre a ed families Sphyraen dae Carang dae Serran dae Lu an dae Mu-
i , t t t ti .
raen dae and mos families of he order Te raodon formes
i i i t t t t
F sh po son ng in the sense used here refers o ha in humans due o
i , ; l ,
the ingestion of flesh which s perfectly fresh (Fereuson 1823 Dun op
1917; Cl ark, 1920; Yasukawa , 1934; Jones, 1937 ; Gilman, 1942; Hi yama,
1943; Goe and Hal st ead, 1955), and in which l eucomai ne-li ke symptoms
are produced . Th is type is generally known as i cht hyosarcotoxi sm (Hal -
, i t t t i l t tl ti
stead and Lively, 1954) and as c gua era in he rop ca wes ern A an c
ll, .
(Randa 1958)
t i ti i . t l
C h arac er s cs o f P o so n o u s B arracuda — The mos commonly imp icated
i i . . t
member of the family Sphyraen dae s S barracuda O her species which
i j ll i
have been cited as being po sonous are S. e o Cuv er (Fonssa<?rives and
i t, , . t i i i , ; , ,
de Mer cour 1861) S fo rs er Cuv er (H yama 1943 Smith 1947) and
t t t i l t i i ,
S. sph yraen a L. when caugh in he rop ca wa ers of West Afr ca (Irv ne
. i t t
1947) Sphyraen ds which are considered safe to ea generally do no a
tt i l i . t
a n a arge i , s ze Barracuda
i t weighing
t l less han 3 pounds are considered
non-tox c andt it s known i ha arger specimens
i of a given i species t, are;
more likely o be po sonous (Fonssagr ves and de Mer cour 1861
128
Jouan , 1863b ; Poey , 1866 ; Dumer il, 1866 ; Pellegrin , 1899 ; Rogers , 1899 ;
Nichols, 1912; Cl arke, 1920; Byam and Archi bal d, 1921; Walker, 1922; l
B , oedorn; and Hakansson, ;1926 Hoffmann ; 1927 , O ’Neill, 1938 Gilman
1942; Strong, 1942; Hi yama, 1943; Paet ro, 1956; de Sylva, 1956; Cal vo
and A lvarez , 1957). It also appears t hat t he l arger t he barracuda, the
more violen t will be the action of the toxin on humans (Yasukawa , 1934 ;
H iyama , 1943 ; Smith , 1947) .
In the barracuda as well as in species of some ot her families, t he visceral
organs and b lood are though t to be mos t po isonous . Gun ther (1 880) and
Mills (1956) believed tha t po isonous fishes are often edible if the head
and intestines are removed immed iate ly af ter cap ture . Yasukawa (1934)
observed tha t the loca tion of the po ison var ied with each species. The
v iscera, par ticu lar ly the gonads , and the skin are known to be mos t toxic
in tetraodon tiform fishes (Remy , 1883 ; H iyama , 1943 ; Tani, 1945) , bu t
the flesh i s often toxic, depending upon the stage of the reproducti ve cycle.
Fortuna te ly , barracuda po ison ing is infrequen t, and deaths are rare .
However, mor talities resu lting from the ingestion of fresh barracuda have
been repor ted by Gu icheno t (1843 : 164) , Mor ton (1 8 6 8 ) , Dun lop
(1 9 1 7 ) , and Ca lvo and Alvarez (1 9 5 7 ) . Poey (1 866) and C larke (1920)
each repor ted two fat alities, Morice (1 958) report ed t hree, and Norman
and Fraser (1938 : 172-173) no ted tha t an en tire family per ished after
having eaten barracuda . Von Fraenke l and Kr ick (1 9 4 5 ) , citing a native
san itary officer in the Mar ianas Islands , no ted tha t during the German
occupa tion there was an 8 to 10 per cen t mor ta lity ra te among the natives
who had been poisoned from eating barracuda . In February , 1961 , a
43-year-o ld native of Bimini, Bahamas , died following a mea l of fresh
barracuda , and there is a repor t of an Amer ican sailor who died of ana l
rup ture following a severe case of po ison ing from a barracuda caugh t in
Cuba .
Non-fa ta l cases of po ison ing resu lting from eatinq fresh barracuda are
repor ted by Moreau de Jonnes (1 8 2 1 ) , Sauvage (1 8 6 5 ) , Dumer il (1 8 6 6 ) ,
Poey (1 8 6 6 ) , Ga tewood (1 9 0 9 ) , Mowbray (1 9 1 6 ) , Gudeer (1 9 1 8 ) ,
Cas te llan i and Cha lmers (1 9 1 9 ) , C larke (1 9 2 0 ) , Byam and Arch iba ld
(1 9 2 1 ) , Hoffmann (1927 , 1929) , Hoffmann and Embil (1 9 2 8 ) , O ’Neill
(1 9 3 8 ) , G ilman (1 9 4 2 ) , H iyama (1 9 4 3 ) , Von Fraenke l and Kr ick
, , i t , t , i
(1 9 4 5 ) Arcisz (1 9 5 0 ) Hash mo o (1 9 5 6 ) Pae ro (1 9 5 6 ) Ebro n
, , l l , ,
(1 9 5 6 ) de Sylva (1 9 5 6 ) Ca vo and A varez (1 9 5 7 ) Morice (1 9 5 8 )
l .
and E lington (1 9 5 9 )
t i i t i l
Corrobora ed po son ng records by fresh barracuda in the rop ca
t tl ti t lit t ,
wes ern A an c which have appeared in he era ure supplemented by
i ti t i
nforma on supplied me by victims of 13 separa e inc dents of barracuda
i i , t l i , t l t
po son ng mostly from sou hern F or da which involved a eas 89 per-
l l t i , t
sons, appear t in Tab e 31. In alt lpar ts of the Car :bbean t here twere 32
cases
t repor l ed which involved a eas t 249 persons
t t hese do no include
hose by E lington (1959) who repor ed a leas 50 cases of barracuda
129
- TABLE 31
o A u t h e nt ic a t e d r e c o r ds of p o is o n in g in h u m a ns from e a t in g b a r r a c uda in the t r o p ic al w e ster n A t l a n t ic ,
COLLATED FROM THE LITERATURE AND S U PP LE M EN T ED BY PERSONAL INTERV IEWS .

Date o f Dat e o f Locality o f No . o f persons Size o f

poison ing 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. ( mal e) b
“Middle o f 1926” “Previous San An ton io , Cuba 23 ? c
afternoon”
Oct., 1927 “Previous day” Rio Almendares, Cuba 8 ? c
April, 1931 ? Camajuam, Cuba 36 ? c
f f ,
“Middle o 1931” “A few hours Havana Cuba 1 ? c
, be ore”
May., 1944 ? Virgin Islands 2 ? d
Aug. 1945
, ?. , f t.
Virgin Islands , 3 !/ ? d
Aug 8 1945 Aug 6 1945 Harbor o S Thomas 3 4 2 ft. d
Virgin Islands, near sub-
, marine base
July 11 1951 ? li s
Terre-de-Bas, 15 ? e
eff de Saintes, Guadeloupe
,
April, 1954 Same day , O Miami, Florida 5 “Large”
/ s. f
h
May 12 , 1954 May 11 1954 Ft. Lauderdale, Florida 5 61 2 lb f
June 27 1954 ? Hillsborough Inlet, Pompano 7 ?
, , .
Beach, Florida s. f
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 lb
Boca Raton,l Florida
,
Aug., 1954 ? Rancho , Ve oz 21 ? g
., Havana Cuba s.
Nov. 1954
, Oct.,. 1954
, Off Miami, Florida 4 10 lb 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 o f Date o f Locality o f No . o f persons Size o f

po isoning 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 mile s 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 o ? ?
f
Cocoa Beach, Florida i
Feb 21 1959 Feb. 20 1959 5 mile north o 5 10 lb
. , , Foweys Rocks, f s. i
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 Governmen 12 3 specimens h
. , . , t
Cut, Miami, Florida 4 20 lbs. ,
Oct 31 1962 Same day 1 3 mile off 9 -
10-12 lbs. h
. , -
Palm s
Beach Inlet
aWalker, 1922
bGilman 1942
cHoffmann
, and Embil, 1931
dArcisz 1950
eEbroin,, 1956; Morice, 1958
Paetro, 1956
fgCa vo and Alvarez, 1957
hPersonal
l interview or newspaper accoun
'Randall personal communica ion t
, t
poisoni ng in t he Cayman Isl ands over 18 mont hs, but no furt her
da ta are available on dates . All symptoms were d iagnos tic of ich thyosarco-
toxism ra ther than bac ter ia l po isoning . Undoub ted ly there are many
unrepor ted cases, since barracuda is usually sold under o ther names , and
because mild cases are often unrepor ted or are b lamed upon bacteria l
poisonings.
A samp le of flesh was obt ai ned from a barracuda about 3 feet long
which had po isoned a family of four. It was caught on July 8, 1955, in
the vicinity of Fowey Rocks, sout h of Miami, and was immedi atel y pl aced
on ice bu t was not cleaned until the family reached home. Shortly aft er
eati ng the fi sh t he members of t he family experi enced typical symptoms
of fish po ison ing ; the effects were fel t for abou t four months .
Dr. Lou is F . Wes tbrook of the Dade Coun ty Depar tmen t of Hea lth
made a bacterio logica l analysis of a sample of the frozen f lesh and found
t t an a pha-
ha l t
ype t ococcus
S rep t , which could no t have caused he t severe
symptoms repor ted , was the only pa thogen ic m icroorganism isolated .
Pae tro (1956) repor ted tha t a sample of f lesh from a barracuda which had
po isoned five persons was found to be negative for organisms commonly
assoc iated with bac ter ia l food po ison ing when analyzed by the F lor ida State
Board of Hea lth . O ther workers such as Yasukawa (1934) and Arcisz
(1950) who have a ttemp ted to iso late pathogenic bac teria from the fresh
flesh of toxic fishes have me t with negative results.
Identifica tion of po isonous barracuda has no t been accomplished (de
Sylva, 1956; Cal vo and Alvarez, 1957), ot her than by feeding the flesh
to experiment al animals. Popu lar beliefs by natives to distinguish toxic
barracuda include black ish or purplish color at the base of the tee th ,
blackening of silver coins or spoons when cooked with the flesh, the
bitt erness of the liver, the presence of a milky substance under the scales ,
and loose or deciduous scales (P lee in Cuvier and Valenciennes , 1829;
Dumeri l, 1866 ; Gudger , 1918 ; Byam and Arch iba ld , 1921 ; Ha ls tead and
Lively, 1954) . Mowbray (1916) and Gudger (1918) report ed t hat
barracuda having b luish backs ( “bl ue-back” ) are considered poi sonous.
The t erm “bl ue-bl ue” i s applied at Bimini, Bahamas, t o barracudat hough t
i , l t t t l
to be po sonous and E lington (1959) repor s ha the f esh of poisonous
barracuda in the Cayman Isl ands has a bluish or bl ue-bl ack appearance in
contrast to the norma l light grey.
A widespread belief among t he natives from vari ous part s of t he worl d
is tha t a lean fish , inc luding barracuda , par ti cul arl y one i n which the liver
tas tes b itter and peppery , is po isonous (Laba t, 1724 ; Damp ier , 1729 ;
Ferm in , 1769 ; Poey, 1866; H iroa, 1938) , bu t tha t fat fish are safe to
ea t (Mills , 1956) . The f lesh of barracuda is of excellen t flavor and is
preferred to o ther species by natives of Cayman Isl ands, according to
Thompson [1945?!, even though barracuda are often poi sonous t here
(E lling ton , 1959). Victims of barracuda poi soni ng have often commented
upon the unusually fine flavor (O ’Neill, 1938) , or tha t it was the bes t
132
 fish ever t ast ed (Paet ro, 1956). The only ot her comment on t he uni queness
of the fish made by any of the victims seems to be t hat the fish was
unusually difficul t to scale or to skin. Mills (1 956) wro te tha t po isonous
indiv iduals . . are dry and hard on cooking and the skin creases like
rubber . If the flesh of a f ish is fa tty it is always safe .” In Cuba , it is
believed tha t fresh barracuda are po isonous which have scales which s tand
out or are deciduous , and in which the mea t is soft. However , none of the
above is infallible , and Von Fraenke l and Kr ick (1 945) s ta ted tha t even
in the Mari anas Isl ands, where t he incidence of fish poi soni ng is high,
the natives are unab le to ident ify a fish which is poisonous.
Tox in from Barracuda Fl esh. — In some of t he t et raodonti form fishes, t he
chem ica l na ture of t he poi son has been studi ed by Takahashi and Inoko
(1 8 8 9 ) , Tahara (1 9 1 0 ) , and Ishi hara (1 9 1 8 ). In ot her groups the
chemica l na ture of t he t oxin i s unknown but i s t hought t o be of an
alka loid type which produces effects similar to tha t of curare (Strong ,
1942; Simmons, e t al., 1944) or aconitine (Ross , 1947) .
A virulent poi son was isolated by Hash i mo to (1956) from a speci-
men of S. pi cuda ( = S. barracuda) about 1 met er long, presumably
from the t ropi cal west ern Pacific. The barracuda, which was preserved in
good cond ition after it was caught, had po isoned abou t 30 persons of five
families . The symptoms in those afflicted were typically those of ich thyo-
sarco toxism . Hash i mo to found t hat t he skin of this i ndivi dual produced
toxic symptoms when fed to animals , bu t he refra ined from deriving
conclusions abou t t he toxicity of t he skin because it was t aken from the
boiled fillet. Hashi mo t o concluded t hat “ . . . t he poi son i s of qui te a new
t
ype, which has never been found in aqua ti c ani mal s,” and differed
from the para ly tic po ison found in the family Te traodon tidae (Tahara ,
1910; Hash imo to and Migita, 1951) , shellfish (G ibbard and Naubert,
1948; Hash imo to , e t a l., 1950 ; Hash imo to and Migita, 1950 ; M cFarren
and Bart sch, 1960), and t he hi st ami ne-l ike substances i sol ated from
scombroids (Ha l st ead and Lively, 1954). Hashi mot o (1956) found t hat
t
he toxin was no t harmfu l when adm in is tered to mice, but had severe
effects upon cats. Poi sonous barracuda may have a more marked effect
upon cats than upon humans , and i ngest ion of even small amount s of flesh
invar iab ly ends with the dea th of the ca t (Locke , 1675 ; Hoffmann , 1929 ;
Hoffmann and Embil , 1931 ; O ’Neill, 1938 ; G ilman , 1942 ; H iyama , 1943 ;
Pae tro , 1956 ; Larson and R ivas , 1962) . O ’Neill (1 938) report ed tha t in
the V rgin
i l
Is ands a fisherman’s ca t which had ingested some of he t b ood
l
of a barracuda was para lyzed in the legs the nex t day and eventually died.
H iyama (1 9 4 3 ) no ted tha t a ca t which was fed cooked flesh from the
reg ion of the pelvic f in of a 91-cm barracuda died abou t 14 hours la ter ;
however, ano ther ca t which was fed raw flesh from the ta il of the same
fish d id no t die.
E tio logy of Fish Po ison ing .— The theor ies which have proposed to accoun t
for the occasional toxicity of barracuda , as well as o ther species, are
133
summar ized by G il man (1 9 4 2 ), Hal st ead (1 9 5 3 ), Paet ro (1 9 5 6 ), de Sylva
(1956 , 1958) , and Randa ll (1 9 5 8 ). Knowledge of the etiological fact ors
responsible have progressed little beyond M art yr’s conclusi on in 1530 (i n
Gudger , 1930) t hat barracuda become poi sonous because t hey feed on
the berr ies of the manch inee l p lan t, H ippomane manc inella L . (Gudger ,
1930; Calvo and A lvarez, 1957).
Of the various hypotheses which have been proposed t o account for
the origin of the t ox i n(s) in barracuda, only two seem pl ausi ble: (1 ) the
po ison occurs as a resu lt of spawn ing (Cos ta Mandry , 1933 ; Strong , 1942 ;
Brown , [1945?]) , or (2 ) the tox in is acqu ired through the food
(M oreau de Jonnes , 1821 ; Dumeri l, 1866 ; Gun ther , 1880 ; Rogers, 1899;
Pellegrin, 1899 ; Yasukawa , 1934 ; Hi yama , 1943 ; Ha lstead and Lively,
1954).
Rel a tion to Spawni ng .— It is known t ha t the gonads of te traodon tid fishes
become poisonous duri ng t he spawni ng season, aft er which some of the
toxin may diffuse to o ther organs , as well as the flesh and the skin (Remy ,
1883; Yasukawa , 1934 ; H iyama , 1943; Tan i, 1945). Ye t t here is little
evidence a t hand t o disclose why such predat ory species as cert ai n scom-
brids, serranids , carangids , and lutjan ids are more ap t to be poisonous
when large. The widespread belief tha t barracuda weighing less than 3
pounds are non-poi sonous (Poey, 1866; Hoffmann, 1927; Hi yama, 1943,
and o thers) seems generally to be t rue. Thi s arbit rary figure of 3 pounds
migh t correspond to t he onset of mat urit y, since females mat ure at a
weigh t of from 3.1 t o 3.9 pounds (580 t o 660 mm fork l engt h); however,
males ma ture earlier, at a weight of from 1. 3 to 1.9 pounds (460 to 500
m m ) . Thus while it i s possible t hat a toxic substance develops in the
female a t the onset of mat urit y, t he males would also be expect ed to be-
come toxic a t mat urit y. If this were t rue, however, cert ai nl y a few poi son-
i
ngs by sexually ma ture males weighing less t han 2 pounds would have
occurred .
Watanabe (1946) has suggested tha t when fishes caugh t only in a cer-
ta in locality are po isonous these represen t ma ture fish which have schooled
for spawning. This exp lana tion would no t, however , expla in why only one
of two fish of iden tica l size and taken from the same place can cause
poisoning. H iyama (1943) found no differences in toxicity between the
sexes in a large number of po isonous indiv idua ls from widely differen t
families. Some freshwa ter fishes, no tab ly Lep isos teus and some European
cyprinids, are among those which have po isonous ovaries (Phisa lix-P ico t,
1922; Pavlovskii , 1927) . A lthough Brown [1945?] no ted tha t in the Lee-
ward Islands t he ovaries of cert ai n fishes, i ncl uding t he barracuda, were
sometimes regarded as dangerous t o eat, Ackerman (1 9 5 2 ) wrot e t hat in
F lori da, ripe ovaries of barracuda are considered a delicacy and are eaten
w it hout harm . But t he spawni ng habit s of most t ropi cal mari ne
, t i l t ti
species are unknown and here s little evidence re ating o the produc on
t i t
of ox n o spawning.
134
 The spawning season of barracuda i n sout hern Fl ori da i s late March to
mid-Oc tober . The di st ri buti on of poi soni ng by freshly eat en barracuda
which caused 30 separate instances of poi soni ng t aken from t he dat a i n
Tab le 31 is as follows:
MONTH JA N . FEB . MAR . APR . MAY JU N E JU L Y
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
no t include the spawning season, and hence it seems unlikely t hat spawning
is respons ible for toxin produc ti on in barracuda. However, work by Takat a
(quo ted in Randa ll, 1958) i ndicating t hat t oxin could be mai nt ai ned in
living fish flesh for at l east 14 months may conceivably i nval idat e the
above analysis.
Rel a ti on t o Di et. — That barracuda acquire a t oxin from eating poi sonous
food is support ed by a large body of ci rcumst anti al evidence. Cert ai n
trop ica l mar ine animals appear to be po isonous nearly all of the t ime i n
some localities . Yet t here seems t o be no evol uti onary patt ern followed
w ithin a group i n the development of a t oxin, with t he excepti on of t et ra-
odon tiform and anguilliform fishes. The former group incl udes t he families
D iodon ti dae, Tet raodonti dae (sensu l at o), Bal ist idae, Monacant hi dae, and
Os trac ii dae, as well as the primitive t et raodonti form families Acant huri dae
and Chae todon ti dae. However, families such as t he Cl upeidae, Sphyraeni -
dae, Carangidae , Scombridae, Serrani dae, Lutj ani dae, Muraeni dae, and
Scaridae are no t closely rel ated; t hese groups and t he t et raodonti form fishes
accoun t for mos t of the po isonings .
We must consider the po isonous princ iple of the organ isms in re lation
to t hei r habit s a n d / o r positi on i n the food web. Those who have att empt ed
to explain fish poisoning in light of the l att er have often oversimplified the
food web in the t ropi cal reef environment. I have att empt ed to present
the generalized i nt errel ati onshi p among organi sms found on a typical coral
reef in southern F l ori da (Fig. 3 5 ). The complexity of t his rel ationship
i
ndicates tha t severa l pa ths may be taken in the t ransmi ssion of a t oxin
from its source to the u ltima te consumer .
Tox ic Organ isms in the F ood Web of the Barracuda .— The following dis-
cussion pert ai ns only to species found in shallow, reef areas where the
incidence of toxic organ isms is high. The families Sphyraen idae , Carang i-
dae, Scombridae, Muraen idae, and large individuals of the families Ser-
rani dae and Lu tjan idae include the most i mportan t fish-eating teleosts, and
are also the most frequently i ncri minated in outbreaks of fish poisoning,
save for the tet raodon tiform fishes. A few of the many references on the
po isonous na ture of preda tory fishes are those of Dumer il (1 8 6 6 ) , Mow-
bray (1 9 1 6 ), Dunlop (1 9 1 7 ), Cl arke (1 9 2 0 ) , Byam and Arch iba ld
(1 9 2 1 ) , Gregory (1 9 2 5 ) , Hoffmann (1 9 2 7 ) , Pavlovski i (1 9 2 7 ) , Cos ta
135
o F
figure
s FISH-EATING TELEOSTS
o
u
th
a rg e nd
(l i vi dua
i sl o fthe genera S p h y ra e n a S
, e r oi al ,C a ra n x ,Scom beromorus ,Ep nephe
i us
l , rca ,Lutjanus)
e r3
n5.
FG
lore
idaen
a DETRITUS-FEEDERS A N ,D BROWSERS . ( INVERTEBRATE-FEEDERS
r ( i . , i ,
l
. iz
ed S p h a e ro d e s .Balistes M onacanthus
, l ) i
Haemulon small serran ds
)

re Acanthurus C h aeto d o n A u tera ca ra n g d s and lutjanids

la
to
i
n
s
am
o
n
g
o
g ar
n
smi
s
o
fe t
h
co Z O O PLA N K TO N
r \ copepods zo ea
al ( , )
-r
ee
f
habitat DETRITUS ( l BENTHIC, A LG AE ,
C h o roph yceae M yxoph yceae PHYTOPLANKTON
lli ) ( , i fl ll )
cora ne Rhodophyceae M yxoph yceae
D no a ge a ta
Mandry (1 9 3 3 ) , Yasukawa (1 9 3 4 ) , Mann (1 9 3 8 ) , O ’Neill (1 9 3 8 ) ,
G ilman (1 9 4 2 ) , H iyama (1 9 4 3 ) , Brown [1945?] , and Ha ls tead and
Lively (1 9 5 4 ) .
Among the fishes which are essentially detr itus-feeders , the te traodon ti-
form fishes occupy a wide variety of hab ita ts bu t are all closely assoc iated
with the cora l-reef hab ita t. The highly po isonous na ture of this group is
t ed
rea t specifically by Anderson (1 7 7 6 ) , Pappe (1 8 5 3 ) , R chardson i
(1 8 6 1 ) Jouan (1 8 6 7 ) Remy (1 8 8 3 ) Takahash and Inoko (1 8 8 9 ) ,
, , , i
Seale (1 9 1 2 ) , Kerr (1 9 1 2 ) , Bryan (1 9 1 5 ) , Ish ihara (1 9 1 8 ) , Phisalix-
P icot (1 9 2 2 ), Nagai and Ito (1 9 3 9 ) , Larsen (1 9 3 9 ) , H iyama (1 9 4 3 ) ,
Tan i (1 9 4 5 ), Yudki n (1944 ; 1945), Hash imo to (1 9 5 0 ) , C lark and Gohar
(1 9 5 3 ) , Jones (1 9 5 6 ) , and Benson (1 9 5 6 ) .
Those feeding on i nvert ebrat es such as pomadasyi ds, l abri ds, and small
individuals of t he families Serrani dae, Lutj ani dae, and Carangi dae have
no t been often i mpl icated in cases of poi soni ng, although when small some
pomadasy ids and lutjan ids are said to be po isonous in the Virgin Is lands
(Wa lker , 1922) . When large serranids , lutjan ids , and carangids are pr in-
cipally fish-eaters.
Coral-feeders are represen ted by numerous members of the family
Scaridae, and cert ai n balistids and t et raodonti ds. Cases of poi soni ng by
scarids and balistids have been recorded by Sonnerat (1 7 7 4 ), Cl oquet
(1 8 2 1 ) , Moreau de Jonnes (1 8 2 1 ) , Va lenc iennes in Cuv ier and Va len-
ciennes (1 8 3 9 ), Poey (1 8 6 6 ), Dumeril (1 8 6 6 ), Hill (1 8 6 8 ), Gunt her
(1 8 8 0 ) , Tybr ing (1 8 8 7 ) , Dunl op (1 9 1 7 ), Cl arke (1 9 2 0 ), Byam and
Arch iba ld (1 9 2 1 ) , Walker (1 9 2 2 ), Pavlovskii (1 9 2 7 ), Simmons, et al.
(1 9 4 4 ) , Ha ls tead and Bunker (1 9 5 4 a), and Mills (1 9 5 6 ). The scarid
genera Scarus and Pseudoscarus, as well as several rel at ed genera, have
often been imp licated in cases of death, but the geographi c vari ati on in
toxicity is marked , perhaps more so than in any o ther group .
The nex t groups in the food chain depic ted in F igure 35 are ben th ic
invert ebrat es and the forage fishes. The following i nvert ebrat es reput edl y
have caused po isoni ng in humans:
Coelen tera tes (Ca lme tte, 1908)
Po lychae tes (Burrows , 1945)
Prawns (MacGowan , 1887)
Crabs (D u Ter tre , 1667 ; Sloane, 1725 ; Damo ier , 1729 ; Poey, 1866 ;
Jouan , 1867 ; Roger, 1895; Banner , 1952)
Spiny lobsters (Dun lop , 1917; C larke, 1920)
Gas tropods (Poey , 1866; Banner , in Banner and Randa ll, 1952)
Pelecypods (G ibbard and Nauber t, 1948 ; Riegel, e t a l , 1949 ; Hash i-
moto , e t a l , 1950 ; Edwards , 1956)
Ho o huri ans and echinoids (Frey , 1951; Nigrelli and Zahl, 1952;
l t
Cata la , 1957)
The above groups are essentially browsers and filter-feeders , and serve
as food for a number of fishes. Hash imo to , e t al. (1950) have iso lated a
137
 toxin, venerupi n, from Japanese mussels, and toxins have been ext ract ed
from Atl anti c mussels (Gi bbard and Naubert, 1948) and Pacific mussels
(R iege l, e t al., 1949) .
Food habits of pl ankti vorous filter-feeding fishes, such as the clupeids ,
in tropi cal wat ers are poorly underst ood. Since they strain large quantities
of zoo- and phy top l ankt on from the waters, they probabl y ingest a number
of toxic pl ankt on organisms, parti cul arl y in t he t ropi cs where bl ue-green
algae and dinoflagellates are the domi nant phyt opl ankt ers. In New Cal e-
don ia and in cer tai n part s of the t ropi cal west ern Atl ant ic, M el ett a ( —Cl u~
panodon) venenosa and M el ett a thrissa ( —Opi st honema ogli num) are well
known for the ir poisonous na ture (Locke , 1675 ; Moreau de Jonnes , 1821 ;
C loque t, 1821 ; Ferguson , 1823 ; S tephenson , 1838 ; Mon trouz ier , 1857 ;
Fonssagrives and de Mer icour t, 1861 ; Hill, 1868 ; Gun ther , 1880 ; Remy ,
1883; Rogers , 1899 ; Dun lop , 1917 ; C larke , 1920 ; Simmons, e t al., 1944 ;
Mills, 1956). Ingest ion of these species often rapi dl y causes fatalities.
Certai n species of Harengula, not abl y macrophthalma and humeralis, also
acquire toxic properties in some areas (Duhamel du Monceau, 1782;
Hill, 1868; Remy, 1883).
The toxicity of phytop lank ton has been reviewed by Mason and Whee ler
(1 9 4 2 ) , Steyn (1 9 4 5 ) , Presco tt (1 9 4 8 ) , Louw (1 9 5 0 ) , V inberg (1 9 5 4 ) ,
Ingram and Presco tt (1 9 5 4 ) , Dawson , e t al. (1 9 5 5 ) , and Habekos t, e t al.
(1 9 5 5 ) .
Dawson, et al. (1955) and Habekost, et a l (1955) have shown tha t
cert ai n marine green and blue-green algae, princ ipa lly Cau lerpa and
Lyngbya , as well as some coralline red algae which are toxic t o l aborat ory
mice, are import ant in the diet of some toxic marine fishes (e.g., Acant hu -
rus, Chae todon , Abudefduf , A ro th ro n ) .
From the above review, the presence of toxins in various mar ine organ-
isms is wel l estab lished , and the hypo thes is tha t a basic toxin is transm itted
t
hrough severa l pa hs
t of the food web of mar ne i organisms should be n- i
vestigated pharmaco og ca ly Studies by Takahash and Inoko (1 8 8 9 ) ,
l i l . i
Ish ihara (1 9 1 8 ) , G ibbard and Nauber t (1 9 4 8 ) , Hash imo to and Migita
(1 9 5 1 ) , V inberg (1 9 5 4 ) , and Banner , e t al. (1960) indicate tha t the
pharmaco log ica l proper ties of the toxins in various troph ic levels may
differ , although the generic toxins w ith in a troph ic level are generally
similar. Much work remains to prove tha t these toxins do no t undergo
modification during transference between successive troph ic levels , in spite
of the apparen t pharmaco log ica l differences among these toxins.
Alt hough the symptoms of these toxins are not ident ical, with the pos-
sible except ion of shellfish and puffer poi soning, the symptoms fall within
the cigua tera-l ke i syndrome .

Food H ab it s and Ecol ogi cal Rel ati ons of Organi sms i n the Food Web of
the Barracuda .— A number of groups which i nclude poi sonous species are
eaten by the barracuda (Tab le 3 0 ) . These include members of the Te tra-
138
odontiformes, Gobiiformes (Pavlovskii, 1927), C lupeiformes, Carang idae ,
Scaridae, Be loniformes (Moreau de Jonnes , 1821 ; C loquet, 1821) , Scom-
bridae , and Anguilliformes . Barracuda over 3 pounds generally cause
poisoning , and the impor tan t d ie tary items of the adult are the Te traodon-
tiformes , Beloniformes , Carang idae , and Scaridae, although there is doub t
as to the import ance of the species of Beloniformes as na tura l food.
Te traodon tiform fishes occurred mos t often in the d ie t of the adult barra-
cuda in sou thern F lor ida and Bimini , being found in 20 per cen t of the
stomachs which conta ined identifiable f ishes . Species which have repu tedly
caused po ison ing in humans were found in 58.1 per cen t of the stomachs
of the adults which con ta ined identifiable food (Tab les 26-30) .
Geographica l D istribu tion of Tox ic Barracuda .— One of the most perp lex-
ing problems in the study of poisonous barracuda is t hei r bi zarre geo-
graphi cal di st ri buti on wi thin a relatively small area. Nat ive fishermen from
such widely scattered po ints as New Ca ledonia , the Mar ianas Islands , and
the Bahamas are report ed to insis t tha t barracuda , as well as many o ther
species, caugh t on one side of a reef or island are safe to eat, while they
are poisonous on the o ther side of the reef only a shor t d istance away
(Jouan, 1861, 1863b; Rogers, 1899; Pellegrin, 1899; Mowbray, 1916;
Walker , 1922 ; Yasukawa , 1934 ; H iroa, 1938 ; Mann , 1938 ; Thompson ,
[1945?]; Brown [1945?]; Mills, 1956) . Yasukawa (1934) observed tha t
poisonous fishes may or may no t be toxic at di fferent Pacific islands or even
at different places on the same reef, and suggested t hat perhaps a parti cul ar
type of food was found only in cer ta in places. Hi yama (1 943) was evi-
dently the first to advance the t heory t hat t here was a rel ati on between the
d istribu ti on of poi sonous fishes and t hat of coral reefs, more specifically the
nema tocys ts of the coral polyps. Daget (1954) not ed t hat a puffer, Tet rao-
don fahaka, which is po isonous elsewhere in its range, is eaten without ill
effects in the upper Niger R iver, where no cora ls exist.
Ecological re lationships in the food web of poisonous fishes have been
briefly d iscussed by Yasukawa (1 9 3 4 ) , Mann (1 9 3 8 ) , Brown [1945?],
Ha ls tead and Lively (1 9 5 4 ) , Mills (1 9 5 6 ) , and Randa ll (1 9 5 8 ) . These
workers have a ttemp ted to pursue a single tra in of events which would
explain the occurrence of toxins in al l poisonous fishes. Randa ll (1958)
presen ted an intriguing ecological hypo thes is on the origin of toxins in
the trop ical mar ine env ironmen t which par tia lly accoun ts for the geograph-
ical paradox involved. He considered tha t toxic b lue-green algae are merely
the firs t organisms growing in norma l ecological succession, and tha t po i-
sonous fishes are found where new subs tra te is being colonized by such
.
aleal growth This hypothesis nicely explains the sometimes sudden appear-
i i .
ance of po sonous fishes following anchor ng of ships or reef destruction
, i t t i l
More recentlv Dawson (1959) also arr ved a the hypo hes s of ecologica
t i it i i , i t
succession. While this heory s mer or ous in most nstances it s no
t l t i ,
adequa e to explain the etiology of an area which harbors toxic p ank on c
139
pelagic, as well as benthi c, organisms, and hydrographi c, chemical, and
me teoro logica l fac tors mus t also be considered .
I t i s not suggested t hat toxins may be t ransmitt ed from algae in only
one way, even though the symptoms produced might be similar. Thus,
po isoning by barracuda might occur from its having eat en poisonous fi shes
which had eat en poi sonous i nvert ebrat es; these could have become poi son-
ous from filtering out large quantities of phyt opl ankt on or from browsing
on toxic, benthi c algae. Barracuda could also become toxic from ingesting
te traodon tids which had become po isonous as a result of toxins produced
in the ir gonads dur ing the reproduc tive cyc le .
Brown [1945?] reported upon the d istribu tion of po isonous and non-
poisonous areas in the Windward and Leeward islands of the Car ibbean .
He concluded tha t in most cases the w indward or eas tern steep , exposed
areas have a repu ta tion for po isonous species and the leeward or shallow-
bank areas are regarded as safe for the same species of fish a t the same
time. Wa tanabe (1946) re la ted tha t native Ok inawans believe tha t red
snapper , Lu tjanus va ig iens is , inhab iting shallow places where the curren ts
are weak are non-poisonous , while those living in deep localities within
swift curren ts are po isonous . G ibbard and Nauber t (1948) observed tha t
mussels which had become po isonous from ingesting toxic , p lank ton ic
dinofiagellates in offshore areas were consistently free from po ison inshore .
The re la tion between toxic mussels and hydrography has been repor ted on
by Wald ichuk (1958) and Prakash and Medcof (1 9 6 2 ) .
Al l po ints in the trop ical western A tlan tic for which informa tion (Tab le
32) is available designating so-called poi sonous and non-poisonous areas
are p lotted in Fig. 36. Most refer to the edibility of the barracuda, but each
po in t may also pert ai n to ot her species such as carangids, l utj ani ds, or
serran ids ; t et raodonti form fishes are not i ncluded. The prevai ling ocean
curren ts , based upon Whiteleather and Brown (1 9 4 5 ) , and the 100- and
1000-fa thom curves are also shown. With few except ions, localities where
fishes are said to be consistently poisonous are charact eri zed by a steep
grad ient along the coastal shelf, and strong, onshore currents. Conversely,
localities where fishes are said never to be poi sonous are generally charac-
ter ized by a relatively gentle-sloping coas ta l shelf assoc iated with weak
currents . Where fishes are occasionally poi sonous, the areas have a steep
coasta l shelf grad ien t bu t lack the strong , onshore curren ts , or the converse .
In the Pacific, Randa ll (1958) observed t hat poi sonous fishes were often
found in areas of seasona l estuarine runoff, which he ascri bed to t he suc-
cession of euryhal ine bl ue-green algae following the destruction of steno-
, ,t i i
haline algae from runoff. In the West Indies however here s no ndication
t t i ,
ha poisonous fishes are ever assoc ated with estuaries and it appears
t t .
ha here areas having estuarine runoff never harbor toxic organisms
l ti i i t i t l i l i
Re
ti a . on of Tox ,c Organ sms o Hydrographt c and
ti M e eoro og ca Cond -
ons — Basically the problem of oxin produc on seems to be one of
140
 TABLE 32
L o c a l it ie s wher e f ish e s ar e r e p u te dly p o is o n o u s or n o n - p o is o no us 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 Mex ico communication
December
(sometimes)
l i ,
Barracuda A ways Havana, Cuba Ro g 1957
ti i i i,
Barracuda, Some mes B m n Bahamas Personal
grouper communication
i , , ,
•‘All spec es but Hormigasl Banksj Bahamas
, Banks Poey 1866
groupers” Islandes as Mu eres
Sanda de Campeche
i , ,
“All f shes” “Always”? Formigas Anegada — Rogers 1899
, ,
Barracuda, cero ? Bahamas
: (general);
i Grand Turk Bermuda;
; Grandi Turk
l Mowbray 1916
carangids Island north s de more often Island south s de ess
often
t t. ,
Barracuda. , East and sou ;h of S Thomas
; — O’Neill, 1938
Caranx latus Virgin
t Islands
i Culebrita
; north
t
other caran- coas of V eques Island eas of
gids; large king Culebra Island
mackerel
Barracuda “Always” Northern coas of Cuba South coas of Cuba Hoffmann 1931
t t ,
Barracuda Never Is e of Pines Cuba Personal
l , communication
 TABLE 32— (CONT INUED )

Species When poisonous Poisonous locality Non-poisonous locality Authority

A ll 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 o f Sail Rock
Lachnolaimus
Barracuda — Yuca an Bean, 1890
t
Red snapper and Windward ground and bank around Re Grenada; S Lucia; Brown 1945?
“other fishes” s
donda; Nevis Statia Saba between St.- Dom nica “practically
t. [ ]
;
Kitts and Statia, ;
and o ;the east and i
unknown , excep for
t
southeast o S Bar windward o Mont- barracuda whicht has a
erra eastf and
t. southeast
ts; o the f Saba doubtful reputation”
s
Bank t; Anguilla seaward o Dog f Island water c o e to ;
Flat ;Cay and ;“outside waterf Ragged; Anguils al s
Island, Bahamas ’’; l
Barracuda ? t.
Montserrat; S Kitt s — Brown [1945? ]
A ll fishes Always Islands and cay s along Norman Island to — Clarke, 1920
Virgin Gorda, British Virgin Islands; to
the north and northeast o f Necker Island;
waters to the southeast at Westend , Tor -
tola; Barbados; Turks Island, Virgin
Islands
Harengula St. Christopher’s ( “one end o f island” ) St. Christopher’s Clarke, 1920
humeralis ( “another end” )
Harengula Occasionally South side o f Jamaica North side o f Jamaica Clarke, 1920
humeralis
 TABLE 32— (CONT INUED )

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 o the S Martin-St Bar Carupano and Puerto Morice 1958
thelemy islands; Terre-de-Bas, e de a Cruz, Venezuela
f
Saintes, Guadeloupe t. . - ,
li s l
Barracuda April Leeward shore o Guadeloupe in chan- Bay o Basseterre Ferguson, 1823
Op sthonema nel be ween S Kit and S Eustatia (Opisthonema); Barba-
,
large carangid f ; f
dos (barracuda)
i , t t. t t.
Some carangid s Nor hea end o Barbados Ferguson, 1823
Barracuda s Smith
t Bayst and fLuongo Cay, east and Gilman, 1942
amberjack southeas o S Thomas, Virgin Islands,
,
king mackerel, respectively
mal , t f t.
“mackerel” .
s l
Barracuda Luis Pena Cay, Culebra Island Virgin Gilman 1942
Islands
, ,
Barracuda Never Bermuda Goode 1876
Barracuda ? S Chr opher and Nevis Castellani
, and
Chalmers, 1919
t. ist
 TABLE 32 (CONT INUED )

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda Occasionally Rio de Janeiro, Brazil — Fonssagrives and
ll l l s f f de Mericourt, 1861
,
s. t s t
A arge fishes A way North coast o Cuba and Jamaica; most South coast o Cuba Hoffmann, 1927 1929
over 3 lb other northern coasts in the Wes Indie and Jamaica; mos
iother southerns coasts
n West Indie
Barracuda Never ,
Cartagena; Islas del Personal i ti
Rosario Colombia commun ca on
Barracuda Occasionally? “Reefs off Key West are regarded with Jordan, 1885
su spicion.”
Barracuda, porgy, Bahamas Locke , 1675
grouper, king
mackerel,
cowfish, puffer,
Op is thonema
Barracuda Never — Key West, Florida Lonnberg, 1894 ;
Mowbray , 1916 ;
Holder , 1924 ;
Schroeder , 1924
Barracuda Never — Tortugas, Florida Gudger, 1918
Barracuda, king ? North side o f Turks Islands, Bahama s South side o f Turks Mowbray, 1916
mackerel, three Islands, Bahamas
specie s o f Car-
anxy red rock-
fish tiger
rockfish
,
Barracuda Havana and Ma anza Cuba Nichols, 1912
t s,
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-pois onous locality Authority

Barracuda May to October Santo Dom ingo — Pavlov skii, 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 ,
(s . )
Scomberomorus ? Barbados — Rogers 1899
p?
Barracuda ? — Spanish Honduras Strong, 1942
, [ ]
Barracuda Occasionally British Honduras
f — Thompson
s , [ 1944?]
Barracuda ? West end o Grand Cayman Island “Other parts” Thomps on, [ 1945?]
Lachnolaimus
, ? Turks and Caicos islands — , t. Thomp on 1945?
? s,
Barracuda small — Cha Cha Towns S Walker, 1922
carangids Thoma Virgin Is-
si lands; Hondura
s
Barracuda and Occa onally Martinique — Moreau de
other specie s , , , Jonnes, 1821
, t
Barracuda Occa ionally Fajardo and Humacao Puerto Rico Aguadilla Arecibo Personal
s t f f
Ponce1 Mayagiiez ,
communica ion
Anchovie ? s Nor h shore o Jamaica t , ll
South shore o f ,
Poey 1866
, s Jamaica t ;
145 Alway Jabor Island, Jaluit A oll Marianas; A other islands o Hiyama
Barracuda
1943
Saipan Mariana Jaluit A oll 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 A to ll, Tuamotu Archipelago Harry, 1953
some snappers,
groupers,
parrotfish
All specie s ? Leeward side o f Canton Island, Phoenix Halstead and
Islands Bunker, 1954a
“Fishe s” ? Leeward side o f Hull Island, Phoenix Halstead and
Islands; northwest, leeward side o f Sidney Bunker, 1954a
Island
“Fishes” Lagoon o f Johnson Island Outside o f lagoon Halstead and
Bunker, 1954b
Barracuda Never Hawaii Jordan and
Evermann 1905
Nearly all species Alway Hood Island near Mangareva ,
Hiroa, 1938
C upanodon ? s ,
New Caledonia Islands o the north o Jouan, 1863b
venenosa New Caledonia
l t f
Barracuda New Caledonia — Jouan, 1863b
Barracuda August to March New Hebride — Mills, 1956
snapper
, s
s
 TABLE 32— (CONTINUED )

Species When poisonous Poisonous locality Non-poisonous locality Authority

Snappers and ? N ew Caledonia; north coast o f Australia; Sikiana, Solomon Is- Mills, 1956
“other species” Fiji; Micronesia; Ellice Island; French lands; Tonga ( “no
Oceania; Philippines and Hawaii; es pec- poisonous fishes” )
ially at Lolowai, Aoba, N ew Caledonia
Mugil cephalus ? Aoba , Maewo, Espiritu Santo; Maske- “Elsewhere in the Mills, 1956
leynes; Malekula group”
“Reef cod” 9 Aoba, Maewo, N ew Caledonia Espiritu Santo, Mills, 1956
New Caledonia
“Large, elongate Always Tuamotus — Pellegrin, 1899
fishes”
“Fishe ” ? Jaluit A o Mariana Carolines: Palau, Yap Yasukawa 1934
s t ll, s Ponape Truk, Kusaie, ,
“F he ” Commonly Canala, N ew Caledonia on east coa La Foa, N ew Ca edo Loison 1955
is s ( st) nia on, west coa l - ,
“Fishe ” ? Outside the harbor a Saipan ( st)
Inside the harbor a Yasukawa, 1934
s t Saipan t
“Fishe ” Occasionally “A certain sec ion o the ree near the r Randall, 1952
s northern part t o Ono
f oa A oll f Gilbert
Islands] wa po[ fonou ”t t ,
Tetraodontids ? s ois Caroline
Outside reefs s Islands and Inside reefs o Caro- Yasukawa 1934
Tinian f line Islands and
f Tinian ,
A fishe ? — Nauru and Ocean Yasukawa, 1934
ll s islands
“Fish and “Extremely rare” — Gilber Islands Catala, 1957
shellfish” t
 ( I )
TABLE 32— CONT NUED
i
Spec es When poisonous Poisonous locality Non-poisonous locality Authority
ll it ,
Barracuda ? — , ?Seyche es Islands Sm h 1956
Barracuda ? lie de Maurice off Madagascar — Lienard, 1843
Coryphaena ? Madagascar ; Madeira Western Atlantic Poey , 1866
Ocean
Sphyraenidae ? — Japan Hiyama, 1943
Barracuda ? — ?Cey lon Ma lpas , 1926
S . sphyraena Occasiona lly Free Town , Sierra Leone Mediterranean Irvine , 1947
Barracuda ? Marshal l Islands — Sm ith , 1947
Tetraodon fahaka Never — Upper waters of Niger Daget , 1954
River
Tetraodontiform ? A1 Ghardaqa, Egypt Suez, Egyp t Clark and Bohar,
fishes 1953
1A spillage o f crude oil near Ponce in July, 1962, was f o llowed by an outbreak o f fish pois oni ng in late 1962 ( pers onal com muni cati on).
primary produc tivity or, more specifically, why cer ta in localities are more
ap t to produce toxic algae than are o thers . Since the toxic b lue-green algae
and dinoflagellates are also the dom inan t phy top lank ters in trop ica l wa ters
and are the base of the food chain , it is only necessary to determ ine the
fac tors limiting abundance of phy top lank ton and ben th ic algae in general,
i.e., light, tempera ture , nu tr ien ts , and p lank ton spores. The first two would
no t appear to be limiting fac tors in clear, warm trop ica l seas. Dissolved
nu tr ien ts , however , are relatively scarce in the eupho tic layers , since tur-
bu lence and upwelling processes are seldom opera tive in trop ica l wa ters .
In trop ica l areas where there is a dear th of e ither nu tr ien ts or phy top lank-
t , he
ers t ncidence
i of toxic organisms migh t be low , whereas po sonous i
areas migh t represen t areas of high produc tivity .
With respect t o the possible effects of current s upon an area, it has
a lready been suggested t hat the sides of reefs or i slands which are exposed
to an onshore curren t more often have typically po isonous organ isms than
do the leeward sides. Similarly, cora l reefs flourish far be tter 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
H i yama (1 9 4 3 ), t hat the di st ri buti on of poi sonous mari ne organi sms cor-
responds generally t o t he di st ri buti on of reef-bui ldi ng coral s, as well as
cora lline algae, which flourish along areas exposed t o onshore current s or
onshore componen ts of these curren ts , is re inforced .
With respec t to mechanisms which migh t influence the supply of nu-
rt ents
i , physica l fac ors
t advecting dissolved nu rt en i st ot eupho tic wa ers
t ,
such as turbul ence and upwelling, are rarel y operati ve in t ropi cal seas;
ne ither is there any indication from the preva iling curren ts in re la tion to
the d is tr ibu tion of po isonous organ smsi (Fig . 36) tha t any upwelling exists.
In fac conditions appear o be dea in mos t ins tances for sinking of wa ter
t, t i l
along the exposed faces of reefs and islands (Sverdrup , e t a l., 1946 : 501 ,
Fig. 124) . Such conditions would be idea l for the opera tion of the “p lank-
t on-trap” mechanism descr ibed by C larke [MS] . He no ted tha t many
phy topl ankt ers are able t o adjust t hei r level of flotation, and t he swim-
ming activity of the organ isms perm its them to aggregate a t var ious levels
in the wa ter column . A wa ter mass converging a t the shore line wou ld sink,
ye t positively pho to tac tic organisms such as dinoflagellates would tend to
rema in in the upper layers of the wa ter resu lting in a “p lank ton trap .” I t
would no t be necessary for the curren t to move a t righ t angles. In terac tion
with preva iling winds could supply an onshore componen t, similar to tha t
dep ic ted by Sverdrup , e t al. (1946 : 500-501) . Loca l var ia tion in the pro-
ductivity could thus be caused by slight varia tions in winds , ocean curren ts ,
or bo ttom topography , with a resulting var ia tion in the toxicity of organ-
isms from one p lace to ano ther .
In po isonous terres trial p lan ts , Muenscher (1951 : 16-17) no ted tha t
there is a similarity in toxic symptoms produced in animals as a resu lt of

149
the ir feeding on cert ai n crop pl ant s, grasses, weeds, and algae, and t hose
produced by feeding the animals pot assi um nit rat e. Pl ant s may store con-
siderable quan tities of po tass ium n itra te in ce ll sap . Muenscher sta ted tha t

3 6 . Distribution of l ocalities i n the tropical western Atlantic based

F ig u r e
on literature reports and personal communications where fi shes are reputed to
be “poisonous” or “non-poisonous.” Current patterns are based on Whiteleather
and Brown (1945).
150
 “the toxic quality of such vegetation i s not due t o the nit rat e directly bui
when the n it rat e i s ingested it i s reduced to nit rit e which i s t oxi c. ” Muen-
scher also no ted tha t “the accumulati on of pot assi um nit rat e varies greatly
in differen t kinds of p lan ts. The quantit y of pot assi um nit rat e may be af-
fec ted by the amounts of nitrogen and pot assium in the soil and the amoun t,
of ligh t coming in cont act with a growing pl ant. A fi eld may be ‘spott y, 5
some areas producing po isonous p lan ts while o thers may be harm less .”
Even though terres tr ia l and aqua tic situations may no t be d irectly compar-
able, I suggest an analogy between the spotty d is tr ibu tion of po isonous
mar ine organisms and tha t of terres tr ia l p lan ts , with a common underly ing
principle. Perhaps such poisonous aqua tic areas could be due to a d ifferen-
tial accumu la tion of nu tr ien ts , with the “toxic princ ip le” in mar ine organ-
isms merely being due initially to various levels of n itra tes . Possibly , local
variations in the current s or nut ri ent supply could favor the growth of
toxic species.
Studies by Thomson , e t al. (1957) and Hughes , e t al. (1958) have
thrown new light on t oxin producti on in bl ue-green algae, which suggest
tha t bac ter ia presen t in the po lysacchar ide shea th of t he bl ue-green algae
may be active in toxin produc tion . These and subsequen t studies by Banner ,
e t al. (1 9 6 0 ) , which showed tha t areas of toxic seaweed in Hawa ii do
no t harbor poisonous fishes, indicate tha t a species of b lue-green algae
may vary spatially and temporally . In light of Muenscher’s (1951) studies,
an analysis of the re la tionsh ip between hydrograph ic and chemica l fac tors
to the toxicity of bac ter ia assoc iated with b lue-green algae , as wel l as cora l
and o ther algae symbionts, migh t be revealing.
The significance of the steep , exposed face of a reef or island in de ter-
mining the presence of po isonous organisms is no t clear. Randa ll (1958)
suggested tha t con tinua l break ing of reef ma ter ials migh t offer a new sub-
strate . Perhaps the grea ter dep th of the euphotic zone on the exposed side
perm its a significantly grea ter produc tivity , which is fur ther enhanced by
the reduced t urbi dit y of these deeper st rat a. Anot her possibility is t hat
along steep faces where onshore curren ts converge, the resu ltan t sinking
of wa ter would be more pronounced in deep wa ter , thus perm itting the
forma tion of a more d iscrete “p lank ton trap .” Fur thermore , a small bu t
steady suoply of nu tr ien ts a n d /o r phy top lank ters would be available to
the benth ic organisms growing along the vert ica l, exposed surface of a
steep reef. The prevailing curren ts could also ac t by supply ing cer ta in
nu trien ts such as n itra tes or phospha tes which are favorable to the pro-
duction of phy top lank ton and sessile algae , or reefs with phy top lank ton .
On the windward side of islands where coral growth is generally more
l
uxur i t, a steady supply of p ank
an l on t si brough t onshore by the preva ling i
. i i t i
currents Reef-bu ld ng corals which con a n symbiotic algae some of ,
which belong to the toxic dinoflagellate group , utilize the me tabolites of
the algae which migh t be concen tra ted by the cora l polyps. These me tabo-
lites could be taken up when the cora l was eaten by fishes such as scarids
151
and t et raodon tiform fishes, which have already been shown to be po ison-
ous and are ea ten by barracuda.
Certa in localities may become toxic because t hey were favorable t o
accumula tions of wind- or current -dri ven algae or dinoflagellates (Mi gi ta
and Kanna , 1957; Waldi chuk, 1958), which are eat en by organi sms on
which barracuda feed. Freshwat er bl ue-green algae accumulat e along
w indward shores af ter a gentle bu t steady breeze (Moh ler , 1941 ; V inberg ,
1954), and the toxicity of the algae increases as the algae decomposes
(Steyn, 1945) .
C o nc l u s io ns

With t he above sequence of events in mind, it becomes clear how some

of the paradoxes in t he probl em of poisonous barracuda fit into t he ecolo-
gical scheme. I t has been already po inted ou t tha t barracuda less than
3 pounds are non-tox ic . A p laus ible exp lana tion for this cryp tic figure is
seen in an event which occurs a t abou t the time the barracuda reaches a
weight of 3 pounds. Thi s i s a movement from a shallow-wat er, grass-bed
hab it at to the deeper wat ers of the coral reefs, which i s associ ated with a
change in behavior and food habits. There i s a rat her abrupt change in t he
diet of the barracuda (Tab le 30) at this size from small fishes such as
atherin ids and gobiids to te traodon tiform , carangid , and belon iform fishes.
It is at this t ime in t hei r li fe when the barracuda themselves are first
considered poisonous.
This change in di et would explai n why only l arger barracuda are toxic.
For this same reason, it would explai n why sphyraeni ds which do not
reach a l arge size are seldom if ever toxic. Thus, S. pi cudill a and S. gua -
chancho, which seldom exceed 2 feet in length, are commonly eat en
t
hroughou t het t ca
rop i l wes ern
t A tlan tic w ithou t il l effects (Gu cheno
i t,
; , ; i , . t
1843 Poey 1858 N chols 1912) O her small species such as S apon ca, . j i
S. obtusa ta , and S. novaeholl andi ae are eat en t hroughout the t ropi cal Indo-
Pacific region, and S. sphyraena is of great importance as food in the
Med it erranean Sea where it is eat en wit hout ill effects (Pal ombi and San-
tarelli, 1953) . S. p ingu is and S. nigrip innis are prized food fish in Japan ,
where poisoning by t hem is unknown (Hi yama, 1943). None of t he above
species att ai ns a l arge si ze. If t he onset of sexual mat urit y were t he cause
of po isonings , surely some large individuals of the sma ller species would
have been incrim ina ted , bu t apparen tly this has no t occurred . Evidently ,
the lack of po ison ing by these sphyraen ids in par t reflects the ir differen t
d ietary habits, for these non-toxi c species feed essentially upon cephal opods
and small a therin ids and clupeo ids none of which is po isonous excep t for
the specia l cases of certain trop ical clupeids of the genera Ov is thonema ,
Harengula, and Cl upanodon. The i mport ant Cali forni a food fi sh Sphyraena
argentea is eaten safely, and even t housh it reaches a length of 3 V2 feet
and a weight of at l east 10 pounds, the l ack of poi soni ng by it seems t o
be due to the dearth of po isonous fishes along the Ca liforn ia coast , which
152
again would appear to be re la ted to the absence of cora l-reef fauna and
flora in the tempera te Ca liforn ia wa ters .
The po isonous na ture of only large indiv idua ls belong ing to o ther
predatory groups , such as the amber jack , king mackere l, lu tjan ids , and
serranids could be expla ined in tha t only the large indiv iduals of these
groups are capab le of swallowing large te traodon tiform fishes. Po isoning
in humans by large barracuda is often more severe than when po ison ing
resu lts from ea ting smaller barracuda (Yasukawa , 1934 ; Smith , 1947) ;
this is true also for te traodon tids (Remy , 1883 ; Tan i, 1945) . This suggests
t t the oxin
ha t si e ither cumu ative
l in the f shi or hat t arger
l t
preda ory i
f sh
are able to inges t larger , and therefore presumab ly more toxic , prey ,
whereas in prey such as the te traodon tiform fishes the amoun t of toxin
produced may be a func tion of the s ize of the reproduc tive organs .
Ano t her enigma is why barracuda are more often po isonous than are
o ther preda tors . I t is suggested tha t the te traodon tiform fishes, which are so
impor ant t in he
t barracuda’s diet, d scourage
i by het ri protective covering,
or canno t be ea ten by , mos t o ther preda tory species which lack bo th the
gape and the sharp , shear ing tee th of the barracuda . The hard integumen t
which is developed in the adults of these slow-swimming crea tures and
t ri large s ze
he i offers little opposit on i ot het barracuda .

L ITERATURE C ITED
A be, T o k ih a r u
1939. A list o f the fishes o f the Palao Islands. Stud. Palao trop. biol. Sta.,
( 4 ) : 523 -583 .
,
A B il l
c ker m a n
i ’f ’i . . .
1952. , Go n ishin n Florida Amer Pub. Co., Miami, 120 pp
A g a s s iz L o u is
s s f ss s s s.
1843. Recherche ur les poissons t l, o iles. . Pois on cycloide Imprimerie
. de Petitpierre, Neucha
f ss e 5: f 1-160 it.
1845 Report on the: o il- fishes . o the London clay. Rep. Br Ass., 14th
, meeting, York 279 310
Ah Kow . T ham f i f f its.
1950 The ood . ff.and feed ng relationships
( ): . o the fishes o Singapore Stra
Colon
, . O
. Fish.. . Publ., 1 35 pp
A n d e r s- o n A W a n d E A fP o w e r . ., . .
1940 . Fishery( statistics, , , o , the , United , States.
). Statist Dig U S Fish Wildl.
1954 Serv., , 2 18 25 27 30 34 36, 3 9
A n d e r s o n W il l ia m t f s s fis i .
1776. An accoun - . o ome poisonou h n the South Seas Phil. Trans.,
66:, 544 552
A r a m b o ur g C a m il l e f s s s - -
1952. Les isi ).
vertebres t s,ossile . des gisement de Phosphate . ,
(Maroc , Algerie:
Tun . e No e Div Mines Geol. Serv geol., Maroc 92 Pisces
, 1-275
A r c is z . W il l ia m fis . s . . .
1950 Ciguatera: ( ): tropical. h poisoning Spec. ci Rep. U S Fish Wildl.
, Serv.,
. 2 7 23 pp.
B a n n e r. A H. a n d J o h n E R a n d a l l f t ,
1952 Preliminary t llreport on marine
, ( ) : biology. study o Onotoa A oll Gilbert
Islands. A o Res. Bull. 1 3 62 pp
153
 P. J. S c h e u e r , S . S a s a k i , P. H e l f r i c h , a n d C . B . A l e n d e r
B a n n e r , A . H .,
1960. Observations on ciguatera-type toxin in fish. Ann . N . Y. Acad . Sci.,
90 ( 3 ) : 770 -787 .
B ar b o ur , T ho m a s H .
1905. No te s on Bermudian fishes. Bull. Mus. comp. Zool. Harv., 46 ( 7 ) :
109 -134 .
B a r na r d, K eppel H.
1925. A monograph o f the marine fishes o f South Africa. Ann. S. Afr. Mus.,
21 , Part 1: 1-418.
B a r nh a r t , P er c y S.
1927. Pelagic fish eggs off La Jolla, California. Bull. Scripps I ns t. Oceanogr.
tech., 1 ( 8 ) : 108 pp.
B a ss a n i, F r a nc e s c o
1899 La ittiofauna de Calcare eocen co d Gassino n Piemonte Atti
. Accad Sci. l Napo 9, Seri 2, i1 3
mat. 1-42. i .
B a u g h m a n J o h. n L fis. li, . ( ):
1947. Fishe, not. previously reported from Texas with m cellaneou notes
s species. Copeia 4 280
on other , is s
, ( ): .
B e a n T ar leto n H
,
1889. .
Report on the fishes observed
n Great Egg Harbor Bay New Jersey during
the summer o 1887 Bull U i S Fish Comm., for 1887 , 7 ,
129-152 f . . . . , :
1890. No e on . fishes co ec ed a Cozumel, Yucatan by the U S Fish
t s
Commission with lldescriptions
t t o new species., Bull U . S . Fish
Comm., or ,1888, 8: 193-206 f . . .
f .
1906 A catalogue o the fishes o Bermuda, with note on a collec ion made
. in 1905 or thef Field Museumf Publ. F e d Mus.s 1 0 8 7 2t 21 89
f . i l ( ), ( ) : - .
B e a n T ar leto n H a nd H G D r e se l
, . . .
1885 A catalogue o fishe received from the Public Museum o the Insti-
. tute o Jamaica, f s descriptions o Pr s poma approxf mans and
with
f
Tylosurus euryops two new species. Proc. f iUti S nat. Mus.,i for 1884,
7 151 170 , . .
: - .
B e c k m a n W il l ia m C
,
1943 Annulus .
formation on the scales o certain Michigan game ishes
. Pap Mich. Acad Sci., 28: 281 312 f f .
. . - .
Beeb e W il l ia m
,
1928 Beneath tropic ea a record o diving among the coral reefs o
. Haiti. G P Putnam
s s; Sons, New fYork, xiii + 234 pp f
. . ’s .
Beeb e W il l ia m a n d G l o r ia H o l l is t e r
,
1935. The fishes o Union Island, Grenadines British West Indies, with
f o a new specie o tar-gazer.
the description , Zoologica , N . Y ., 19
f s f s
6
( ): 209 - 224 .
B eeb e W il l ia m a n d Jo hn T ee- V an
,
1928. The fishes o f Port-au-Prince Bay, Haiti. Zoologica , N . Y ., 10 ( 1 ): 1-
279.
1933. Field book o f the shore fishes o f Bermuda. G . P. Putnam’s Sons, New
York, 337 pp.
B e nso n J er o m e
,
1956. Tetraodon (blow fish ) poisoning . J. forensic Sci., 1 (4 ) : 119 -126 .
B erg L ev S
,
1955. .
Sistema ryboobraznikh i ryb, nynye zhivu shchykh i iskopayemykh .
[Classification o f fishes both recent and fo ssil.] Trav. Inst. zool.
Acad. Sci. U.R .S.S ., 20: 1-286. Rev. Ed.
154
B l a nc , M a u r ic e a n d P ie r r e F o u r m a n o i r
1953. Poissons d ’ Oceanie rapportes par M. G. Ranson en 1952. Bull. Mus.
Hist, nat., Paris, Ser. 2, 25 ( 6 ) : 546 -550 .
B l e e k e r , P ie t e r
1849a. A contribution to the knowledge o f the ichthyologica l fauna o f
Celebes. J. Indian Archipelago , 3: 65-1 A.
1849b. Bijdrage tot de kennis der Percoiden van den Malajo-Molukschen
Archipel, met beschrijving von 22 nieuwe soorten. Verh. batavia.
Genoo t., 22: 1-64.
1854a. Specierum piscium Javanensium novarum vel minus cognitarum diag -
noses adumbratae. Natuurk. Tijdschr. Ned.- Ind ., 7: 415 -448 .
1854b. Nalezingen op de ich thyologie van Japan. Bijdrage tot de kennis der
Sphyraenoiden van den Indischen Archipel. Verh. batavia. Genoo t.,
26: 1-22.
1865. Description de quelques espece s de poissons du Japon du Cap de
Bonne-Esperance et de Suriname, conservees au Musee de Leide. Ned .
Tijdschr. Dierk. , 2: 250 -269 .
B l o c h , M a r c u s E. a n d J o h a n n G. S c h n e i d e r
1801. Systema ichthyologiae inconibus ex illustratum. Bibliopolio Sande-
riano Commissum, Berlin, Vol. 1, lx + 584 pp.; Vol. 2, 110 pis.
B l o e d o r n , W . A . a n d E. H a k a n s s o n
1926. Some observations regarding Culebra. Nav . med. Bull., Wash., 24:
534 -544 .
B o l in , R o l f
1954. Report on a fatal attack by a shark. Pacif . Sci., 5 ( 1 ) : 105-108.
e o r g e A.
B o ul e ng e r , G
1901. Les pois sons du bassin du Congo. Etat independant du Congo ,
Brussells, lxii 4- 532 pp.
B r e d e r , C h a r l e s M ., J r .
1926. The locomo tion o f fishes. Zoologica , N . Y., 4 ( 5 ) : 159-297 .
1927. Scientific results o f the first oceanographic expedition o f the “Paw-
nee.” Bull. Bingham oceanogr. Coll., 1 ( 1 ) : 1-90.
1948a. Observations on coloration in reference to behavior in tide-pool and
other marine shore fishes. Bull. Amer . Mus. nat. Hist., 92: 285 -311 .
1948b. Field book o f marine fishes o f the Atlantic Coast from Labrador to
Texas. G. P. Putnam ’s Sons, New York, 331 pp.
1949. On the relationship o f socia l behavior to pigmentation in tropical
shore fishes. Bull Amer. Mus. nat. Hist., 94: 85 -106 .
, .
B r id g e T h o m a s W
1896. The mesial fins o f ganoids and teleosts. J. Linn. Soc. (Zoo l.) 25:
530-599 ; 600-602.
, .
Brown H erb ert H
[1945?] The fisheries o f the Windward and Leeward Islands. Bull. Developm .
Welf. W. Ind., ( 2 0 ): 97 pp.
,
B r y a n W il l ia m A
.
1915. Natural history o f Hawaii, . . . Hawaiian Gazette Press, Hono lulu ,
596 pp.
, .
B u t t iko f e r J
1890. Reisebilder aus Liberia. Vol. 2. D ie Bewohner Liberia’s. Thierwelt.
E. J. Brill, Leiden , 510 pp.
,
Bum p u s D ean
1957. Surface water temperatures along Atlantic and Gulf coast s o f the
United States. U. S. Fish Wildl. Serv., Spec. sci. Rep. ( 2 1 4 ): 153 pp.
155
B ur r o w s, W il l ia m
1945. Periodic spawning of “pal ol o” worms in Pacific waters. Nature,
Lond., 155 ( 3 9 2 4): 47- 48.
B u tl er , R o b ert L. a nd L lo yd L. S m i t h, J r.
1953. A method for cellulo se acetate impressions o f fish scale s with a
measurement o f its reliability. Progr. Fish Cult., 15 ( 4 ) : 175-178.
B y a m , W i l l i a m a n d R . G. A r c h i b a l d , e d s .
1921. The practice o f medicine in the tropics. Oxford Univ . Press, London,
1 : 790 -798 .
C a d e n a t , J.
1950. Poiss ons de Mer du Senegal. Inst, f rang. Af r. Noi re, Initiations
Africaines, 3: 5- 345.
Calm e t t e, A lb ert
1908. Venoms, venomou s animals and antivenomous serum-therapeutics.
John Bale Sons and Daniels son , Ltd., London , 403 pp., 124 figs.
C alvo, R af ael and A lb er t o A l var ez
1957. Recordativo sobre la ciguatera. Mar y Pesca, 1 ( 6 ) : 32-36 .
C a r b i n e , W. F.
1944. Egg production of the northern pike, Esox lucius L., and the per-
centage survival o f eggs and young on the spawning grounds. Pap.
Mich. Acad . Sci., for 1943 , 29: 123 -137 .
,
C a sie r E d ga rd
. i . t .
1944 V. Les genres Tr chiurides Winkler (s str.) et Eu richiurides nov
l’ s ’
VI. Sur de sSphyraenodus de ( tEocene
t i fet i ur ).el presence
. d un sphy-
t.
raenide dan ):le Bruxellien Lu e ien n er eur Bull Mus. Hist. na
. Belg. 20 ( 11 t 1-16.
l l’ l .
1946 La faune t. ich hyo , ogique
. I, ( de): Ypresien de a Belgique. Mem Mus.
Hist. na Belg. s Ser l’ 1 0 4 1-267. s .
1950. fContributionf a etude des poissons is li fossile de la Belgique. 6 La
aune des( ormations
): . dites “pan e ennes.” Bull. Mus. Hist. nat.
Belg., 26 4 2 1-52
. s t
1954. Appendix In Leriche, M., Les faune, ich hyologiques marines. du
Neogene des Indes orientales. Schweiz palaeont. Abh., 70: 1-21
, .
C a s t e l.l a n i A ldo a n d
f A l b er t J C h a l. m e r s
1919 Manual
. , o tropical. medicine Bailliere, Tindall, and Cox, London,
Ed 3 x + 2436 pp
, .
C a s t e l.n a u F r a n c o i s L d e L a p o r t e s d.e . . l’ , . . . -
1855 Animaux . . nouveaux , ous, rare( ) , de
t Amerique . du Sud Pois
. sons P Bertrand s Pari s7 2 l’ Par f i 3, 112 pp . . t fils,
1861 Memoire , ur. les poisson de A r que australe J.-B Bailliere e
. Paris 78 pp t f t s s
1873 Contribution. to the ich hyology o. Australia. VI. , No e - on . fi hes from
,
Knob Island
. .
Proc. zool. acclim Soc. Victoria 2: 37 158
C a t a l a. R e n e L A : f .
1957 Report . on the
( ): Gilbert Islands
. some aspects o human ecology Atoll
,
Res Bull., 5 9 187 pp
C a t e s b .y M a r k l f s;
1743 The . . . natura - history. o Carolina, Florida .and the Bahama Island
1731
is. 1743 Privately published for C Marsh, London, 2 vols.,
220 p
, . I.
C h a c k o. P f s f f lf f
1949 Food and eeding . habit: o- the
. fishes o the Gu o Manaar. Proc.
Indian Acad Sci., 2 9 B 83 97
156
C h e vy, Pa ul
1933. The method o f reading scales and the fish o f the intertropical zone .
Proc. fifth Pacif. Sci. Congr., 5: 3817 -3829.
C l a r k , E u g e n i e a n d H. A . F . G o h a r
1953. The fishes of the Red Sea: order Plectognathi. Publ. Mar. biol. St a.
Ghardaqa, ( 8 ) : 80 pp.
C l a r k e , G e o r g e L.
[MS] Ecological relations possibly invol ved i n the red tide. March 21,
1950, Univ. Miami, Inst. Mar. Sci. [ library]. Typescript.
C l a r k e , T. L. E.
1920. Some observations on fish poisoning in the British Virgin Islands.
W. Ind. B u ll, 17: 56-67.
C l o que t , H ip p o l y t e
1821. Ich thyolog ie . In Dictionnaire des science s naturelles. . . . F. G.
Levrault, Paris, 22: 550 -554 .
1827. Ichthyologie . In Dictionnaire des science s naturelles. . . . F. G.
Levrault, Paris, 50: 233 .
C o c k e r e l l , T h e o d o r e D . A.
1914. Observations on fish scales. Bull. U. S. Bur. F is h , 32: 119-174.
C o h e n, D a n ie l M.
1954. Age and growth studies on two species o f whitefishes from Point
Barrow, Alaska. Stanf. ichthyol. B u ll, 4 ( 3 ) : 167-188.
C o l b y , M a l c o l m , Jr .
1943. Poisonou marine animals n the Gu o Mexico Proc. Tex. Acad
S c for 1942 s 26: 62 70 i lf f . .
C o p e E d w a i, rd D , - .
1871., .
Contribution to the ich hyology o the Lesser Antilles. Trans. Amer
phil. S o c new S er 14: t445 483 f .
1875. Synopsis ,o the vertebrata , -o the . Miocene o Cumberland County,
New Jerseyf Proc. Amer phil f S o c 14: 361 364 f
C op pleso n V M . . . , - .
1959. Shark , . attack
. Angu and Robertson, Sydney 266 pp
C ost a M a ndr y O . s , .
1933. Food , po. oning in Puerto Rico Puerto Rico J publ. H h 9 1
44 89
C r e a s e r C h a r l e sisW . . lt , ( ):
1926 The - structure
. and growth o the scales o fishes in relation to the
, interpretation . o their li e history, with specia re erence to the sun
. fi h Eupomo s gibbosus Misc. f f
Publ. Mus. Zool. Un v M ch
1 7 1-82 f f l f -
C u v i e r G eso ,r g e s a n d Atic h i l l e V a l.e n c i e n n e s i . i ,
1829 Histoire( ): naturelle . de poissons. F. G. Levrault, Paris, Vol. 3
1831 , Histoire naturelle des poissons. F. G. Levrault, Pari Vol. 7
1839. Histoire naturelle dess poissons. F. G. Levrault, Paris Vol. 14
D a g e t .J s, .
1954.. Les poisson du N ger superieur. Mem Inst fran?. ,A r. No .re 3 6
, . 1-391.
D a m e s W i l h e l m B»s i . . f i , ( ):
1883., ber eine tertiare Wirbelthierfauna von der westlichen Inseln des
t)
Birket-el-Qurun in Fayum Aegyp en S B preuss Akad W
6: 129 153 P 3 Fig 12 ( t ). . . . . iss,
D a m p i e r W i l l i-a m , I. , . .
1729 , Dampier voyages consisting o a new voyage round the world, a
supplement to the voyage around the world, Two voyage to Cam-
. peachy ’s Ed by ;John Masefield, f E P Du on N Y vii + 624 pp
s
, ... . . . tt , . , 157.
D ar tevelle, C. a nd E d g a r d C a s ie r
1943. Les poisson s fossiles du Bas-Congo et des regions voisines . Ann .
Mus. Congo beige, (Ser. 3 A ), 2 ( 1 ) : 200 pp., 16 pis.
1949. Les poissons f ossiles du Bas-Congo et des regions voisines. ( Deux
par tie). Ann. Mus. Congo beige, (Ser. 3 A), 2 ( 2) : 201- 256, Pis.
xvii-xxii .
D a w so n, E. Y al e
1959. Changes in Palmyra At oll and its vegetation through activities of
man, 1913 -1958 . Pacif. Nat., 1 ( 2 ) : 1-51.
D a w so n, E. Y ale, A. A. A leem , a nd B r uc e W . H alste a d
1955. Marine algae f rom Palmyra Island with special ref erence to the
feeding habits and tox icology o f reef fishes. Occ. Pap. Allan Hancock
Fdn., ( 1 7 ) : 39 pp.
D ay
, F r a n c is
1875- The fishes o f India, . . . Bernard Quaritch, London, 2 vols., xx + 778
1878. pp., 198 pis.
1889. Fishes, ln W. T. Blanford, ed., The fauna of British India, including
Ceylon and Burma. London , 2 vols., 341 figs.
, .
d e B e a u f o r t L F.
1926. On a collection o f marine fishes from the M iocene o f South Celebes.
Jaarb. Mijnw. Ned.-Oost -Ind., 54e annee, Verh., for 1925: 117 -148 .
, .
D e K ay J a m es E
1842. Zoology o f New York, . . . Vol. 1, Part 4. Fishes. W. A. White
and J. Visscher , Albany, 415 pp., pis.
, . .
D elsm a n H C
1930. Fish eggs and larvae from the Java Sea. 16. Amph ipr ion percula
C. V. Treubia, 12 ( 3 - 4 ): 367 -370 .
1941. Resultats scientifique s des croisieres du Nav ire -fico le Beige “Mer­
cator .” Mem. Mus. Hist. nat. Belg., Ser. 2, 3 ( 2 1 ) Part 3: 47 -82 .
,
d e S yl v a D o na l d P
.
1955. The o eology and phylogenetic relationships o f the blackfin tuna,
st
Thunnus atlanticus ( Less on ). Bull. Mar. Sci. Gu lf & Carib., 5 ( 1 ) :
1-41.
1956. Poisoning by barracuda and other fishes. Spec. Serv. Bull., Univ.
Miami Mar. Lab., ( 1 3 ): 1-9. [Mimeographed.]
1958. The lif e history and systematics of the great barracuda, Sphyraena
barracuda ( Wa lbaum ). Doc t. Diss., Cornell Univ ., 374 pp.
d e S y l v a , D o n a l d P. a n d W a r r e n F. R a t h j e n
1961. Life history notes on the little tuna, Eut hynnus alletteratus, from
the southeastern United States. Bull. Mar. Sci. Gulf & Carib., 11
( 2 ) : 161-190.
, . , . , .
d e S y l v a D o n a l d P., F r e d e r i c k A K a l b e r a n d C a r l N S h u s t e r J r
1962. Fishes and eco logical condition s in the shore zone o f the Delaware
River estuary, with notes on other species collec ted in deeper water.
Inf. Ser. Un iv . De l. Mar. Lab., Publ., ( 5 ) : 1-164.
D ole
, R ic h a r d B .
1914. Some chemical characteristics of sea-water at Tortugas and around
Biscayne Bay, Florida. Pap. Tortugas Lab., 5: 71- 78.
D ole, R ic h a r d B . a nd A lf r ed A . C ham b er s
1918. Salinity o f ocean-water at Fowey Rocks, Florida. Pap. Tortugas
Lab., 9: 299 -315 , PI. 99, Figs. 5-6.
D ollo
, L o u is
1909. Les teleos teen s a ventrales abdominales secondaires. Verh. zool. -bot.
Ges. Wien, 59: 135 -140.
158
D o yle, A d r ia n C o na n
1953. Heaven has claws. Random Hou se , New York, 245 pp.
D uha m el d u M o nc e a u, H e nr y L o u is
1782. Traite general des pesches, et histoire des poiss ons qu’elles fournis-
sent, . . . Saillant e Lyon, Paris, Part 2, Sect. 3.
D u m e r il , A ndre
1806. Zoologie analytique, ou methode naturelle de classification des ani-
maux. Allais , Paris, xxxiii + 344 pp.
D u m e r il
, A ug us t
1866. Des poissons veneneux. Ann. Soc. linn. Maine-et-Loire, 8. [Translat-
ed by W. E . O’Shaughnessy in Ann . Mag . na t. Hist., Ser . 3, 20:
153-167 , 1867 .]
, . .
D unlop W R
1917. Poisonous fishes in the West Indies. W. Ind. Bull., 2: 159 -166 .
Du T e r t r e , J e a n B. s .... s l ll , s, .
1667., Histoire generale des Antille Thoma o y Pari Vol. 2
E b r o in A ry
s s fis f .
1956. The venomou , and poisonou . h o Guadeloupe i Lai Peche Inde-
pendante,
i . July (Sept., ) : Oct., Nov . Published n English n Publ. Fish.
,Car. b. Comm., 3 1-14, 1959
E dw a r ds H I t i i f .
1956. The e iology and ep - dem . ology o paralytic shellfish poisoning. J
, Milk Tech., 19: 331 335
E k m a n. S v e n f . ,
1953 Zoogeography
. o the sea Sidgwick and Jackson, Ltd., London 417
pp, . .
E l l i n g .t o n A C s i . . . ( ): - .
1959 , Poisonou
. fishes n the Caribbean area W Ind Fish. Bull., 6 15
E l l i s R o b e r t W f fis ’s . .
1957. Catches o . h byi charter . boats( on ): Florida
, east. coast Spec. Serv
, Bull., Univ. Miam Mar Lab., 1 4 1-6 Figs. 1-3 [Mimeographed.]
E r d m a n D o n a lt dfisS s . . lf
1956. Recen ( ): h- record . from Puerto Rico. Bull Mar Sci. Gu & Carib.,
6 , 4 315 340 . .
E v e r m a n n B a rst o n Wf a n d H o m e r C M a . r s h. . , ,
1902. Thet fi: hes- o ,Puerto is. Rico. Bull U S Fish Comm., for 1900 20
Par
, 1 51 350 P i-xlix.
F e r g u s o n W il l ia m s f . ,
1823. On the poisonou - . fishes o the Carribee Islands Trans roy. Soc.
, Edinb., 9: 65 79
F e r m i n P h il ip p e l
1769. Description generale,. . historique, . . . geographique et physique , de a
, colonie. de Surinam . E von Harrevelt, Amsterdam 2 vols.
F i s h C h a r l e si J sa n d M a r y C C o b b f
1954. Nox. ou marine . . animals .o the central ( ): and . western Pacific Ocean.
Res , Rep.. U S Fish Wildl Serv., 3 6 45 pp
F o n s s a g r i v e s M a sn ds L e r o y d e M e r i c o u r t s s s
1861. Recherche . ur les poissons toxicophore t : - exotique
. de pays chauds.
, Ann Hyg. publ., Paris, 16, Par 2 326 359
F o r b i n V. l s , ( ): - .
1924. , La Sphyrene de a mer de Antilles. Nature Paris, 52 1 47 48
F o r sk a l P e h r s , ,
1775. Descriptione . . . animalium avium , amphibiorum,
, piscium insectorum, is.
vermium, Carsten Niebuhr editor Copenhagen, 164 pp., 43 p
159
F o w ler , H e n r y W.
1900. Contributions to the ich thyology o f the tropical Pacific. Proc. Acad .
nat. Sci. Philad., for 1900, 52 , Part 3: 493 -528 .
1903. New and little known Mugilidae and Sphyraenidae. Proc. Acad. nat.
Sci., Philad., for 1903, 55 , Part 3: 743 -752 .
1905. The fishes o f New Jersey. Rep. N . J. Mus., for 1905: 35 -477 .
1906. Some cold -blooded vertebrates o f the Florida Keys. Proc. Acad. nat.
Sci. Philad, for 1905 , 58 , Part 1: 79 -113 .
1915a . A list o f Santo Dom ingo fishes . Copeia , (2 4 ): 49 -50 .
1915b. Fishes o f Trinidad, Grenada and St. Lucia, British West Indies. Proc.
Acad. nat. Sci. Philad., for 1915, 67: 520 -546 .
1916. Cold-blooded vertebrates from Costa Rica and the Canal Zone. Proc.
Acad . nat . Sci . Philad., for 1916 , 68: 389 -414 .
1920. A list o f the fishes o f New Jersey. Proc. biol. Soc. Wash., 33: 139-
170.
1922. A list o f Hawaiian fishes. Copeia , (1 1 2 ): 82 -84 .
1925. Fishes lof, (Guam , Hawaii, Samoa and Tahiti. Bull. Bishop Mus.,
Honolu u 2 2 ): 38 pp.
1927. No te s on the Philippine fishes in the collec tion o f the Academy .
Proc. Acad . nat. Sci. Philad., 79: 255 -297 .
1928. The fishes o f Oceania. Mem. Bishop Mus., Honolu lu , 10: 540 pp.
1929. No te s on Japanese and Chinese fishes. Proc. Acad. nat. Sci. Philad.,
31: 589-616.
1931. The fishes of Oceania. Suppl. 1. Mem. Bishop Mus., Honolulu, 11
( 5 ) : 314-381.
1932a. Fishes obtained at Fiji in 1929. Occ. Pap. Bishop Mus., Honolulu ,
9 ( 2 0 ): 1-13.
1932b. Fishes obtained by the Barber A s phal t Company i i n Trinidad and
Venezuela n 1930. Proc. Acad. nat. Sci. Philad., for 1931, 83:
391 -410 .
15(34. The s f .
( ) : fishe- o . Oceania Suppl. 2. Mem Bishop Mus., Hono ulu 11
. l ,
. 6 385 466
s f f . t t.,
1935a A (synopsi ): -o the . fishes o China Par 5, continued. Hongkong Na
. 5 4 s 304 319 . . i t l -
1935b Fishe obtained by Mr H. . W Bell-Marley chiefly i. n Na a and Zulu - .
. land from 1929 f to 1932 Proc.. Acad.
. nat. . Sc Philad., 86: 405 514
( ):
1936 Marine. fishes o West Africa Bull Amer Mus. nat. Hist., 70 1
. 605 pp f
1938 The fishes o the . George . Vanderbilt ( ) :South Pacific Expedition, . 1937.
Monogr.f Acad nat. Sci Philad., 2 349 pp., f 12 pis . .
1941. A list o (the ) : fishes- known . from the coast o Brazil. Arch Zool. S
. Paulo, 3 f6 115 184 ( ).
1944 Resultss o the. Fifth. George . Vanderbilt( )Expedition
: - , 1941is. The
fi hes. Monogr f Acad nat
f Sci. Philad., 6 57 t 529 20 p .
1945. A study. o the . fishes o the southern ( ): Piedmon and coastal fi s. plain
Monogr Acad f nat. Sci. Philad.,
. . 7 . vi + 408 pp., 313 , g ( ): -
1952. The. fishes o Hispaniola Mem Soc cubana Hist, nat. 21 1 81
115
. f . - .
1953 The shore fishes o the Colombian Caribbean Caldasia, 6 (27): 43 73
, . .
F o w l er H e nr y W a nd B ar to n A B ea n
t s i , . .
1927. Not. e on fishes ( obtained
): . n Sumatra Java and Tahiti. Proc. U S
, .
na Mus., 71 10 1-15
F r a se r C M cL ea n f , . . ,
1920. Growth . , rate o the. Pacific salmon. Trans roy Soc. Can., Ser 3
Sect 5 13: 163-226
160
F r e y, D a v id G.
1951. The use o f sea cucumber s in poisoning fishes. Copeia, ( 2 ) : 175 -176.
F r o s t , G. A l l a n
1929. A comparative study of the otoliths of the Neopt erygian fis hes
(
con tinued ). (N o . 18 . Percomorphi. [ Concluded.] Mugili oidae.)
Ann . Mag. nat. Hist., 10th Ser., 4: 120-130.
1934. Otoliths of fishes from the Lower Tertiary Formation of Southern
England. III. Percomorphi, Scleroparei. Ann. Mag. nat. Hist., 10t h
Ser., 13 ( 7 6 ) : 426 -433 .
, .
F u g l is t e r F r e d e r ic k C
1947 . Average mon thly sea . surface temperatures
,
f
( ) : o the western North
Atlantic
, . .
Ocean. Pap phys. Oceanogr. 10 2 1-25.
G allo w ay J C
. f f - s t
1941 Lethal effect o the cold , ( )winter
: - o .1939 40 on marine fi hes a Key
West,
, Florida.
. Copeia 1 118 119
G atewo o d J a m e s D
. l . . s
1909 Nava . hygiene P Blakiston’ Sons and Co., Philadelphia, xiv +
779
, .
pp. . , .
G a t h m a n C A a nd C har les E D a w so n J r
. f s f fis
1948 The s use o the
. plastic laminating ( pres
): for the
. mounting o h
cales. Quart. J Fla. Acad. Sci., 10 4 153-154
, . .
G ib b a r d J a n d J N a u b e r t
. . . .
1948 Paralytic shellfish , ( ) : poisoning
- . on the Canadian Atlantic Amer J
publ. Hlth. 38 4 550 553
, .
G
. C h a r l e s fH
il b er t
t I . f .
1900 Results . o the . Branner-Agassiz
. - Expedition
. o Brazil. II The ishes
Proc Wash Acad Sci., 2: 161 184
, . . . .
G i l c h r i s t J F D a n d W W. T h o m p s o n
s f s f . . :
1909. Description ; -o fi. hes from the coast o Natal. Ann. S Afr Mus., 6
145-206 213 279
, .
G il m a n R o b er t L
. f fis i s :
1942 A reviewf o h poisoning n the Puerto Rico-Virgin Island . area a
report o ten cases . occurring on Culebra Island. Nav med. Bull.,
Wash., 40: 19-27
, . .
G o e D o n R a n d B r uc e W H a lste a d
f fis
1955. A case, o h poisoning s from Caranx ignobilisf Forskal from Palmyra t -
Island f with comment onf the t sensitivity
. o the, mouse-injection
( ): - . ech
nique or the screening o oxic fishes Copeia 3 238 240
, .
G . G eorge B
o o de
f t f . . . . .
1876 Catalogue
( ): o . he fishes o the Bermudas, Bull. U S nat. Mus.,
. 1 5 82 pp f , s f
1877 A preliminary catalogue o the reptiles f fishe and f leptocardians
s o
the Bermudas, with. descriptions
- o. four species o fi hes believed to
be new. Amer. J Sci.,
f 14: 287 298 . s s
1884. Natural
i history
f o useful aquatic animals
, t The
, fisherie
f and fi fhery
ndustries o the United States. Sect. 1 Par 3 The ood fishes o. the
United States. Government Printing Office, Washington, 895 pp
, . .
G o o de G
. eor ge B
f T a rlll e ttio n H f B e a n
and i lf f
1879a Catalogue t o . a .co ec , on o fishes obtained f n the Gu o Mexico.
by. Doc or J W Velie - with descriptions o seven new species. Proc
U S. nat. Mus., 2:ll333 345.f
1879b. Catalogue o a co ec ion o fishes sent from Pensacola , Florida, and
f t
vicinity , by Mr. Silas Stearns, with description o f six new species.
Proc. U. S. nat. Mus., 2: 121-156.
161
 1882. A list of the ( 2 9 6) species of fishes recorded as occurring in the
Gu lf o f Mexico. Proc. U . S. nat. Mus., 5: 234 -240 .
G o o d r ic h , E d w in S.
1909. Vertebrate Craniata ( cyclost omes and fis hes). In Lankester, E. R.,
A treatise on zoo logy. Adam and Charles Black, London, 9: xvi +
518 pp.
G o r do n, M yr o n
1946. Barracuda— tiger-of-the-sea. Nat. Hist., N . Y ., 55 ( 1 ) : 27 -32 .
G o sl in e , W il l ia m H.
1962. Systematic position and relationships o f the percesocine fishes. Pacific
Sci., 16 ( 2 ) : 207-217.
G r aham , M ic h a e l
1929a. Studies on age-determination in fish. Part 1. Fish. Invest. Minist. Agr.
Fish., Ser. 2, 11 ( 2 ) : 50 pp.
1929b. Studies on age-determination in fish. Part 2. A survey o f the litera-
ture. Fish. Invest. Minist. Agr. Fish. , Ser. 2, 11 ( 3 ) : 50 pp.
G r ay
, Jam es
1957. How fishes swim. Sci. Amer., 197 ( 2 ) : 48-54.
G r egory
, C l if f o r d R .
1925. Three cases o f food poisoning attributed to eating freshly caugh t fish
— yellow jacks. Nav. med. Bull., Wash., 23: 316 -320.
G r e g o r y , W i l l i a m K.
1933. Fish skulls: a study o f the evolu tion o f natural mechanisms. Trans.
Amer. phil. Soc., 23 ( 2 ) : 75 -481 .
G udg e r , E ug e ne W .
1918. Sphyraena barracuda; its morphology, habits and history. Pap. Tor-
tugas Lab., 12 ( 4 ) : 53- 108.
1930. Poisonous fishes and fish poisonings, with special reference to c igua -
tera in the West Indies. Amer. J. trop. Med., 10: 43-55.
G u d g e r , E u g e n e W. a n d C h a r l e s M. B r e d e r , J r.
1928. The barracuda ( Sphyraena ) dangerous to man. J. Amer. med. Ass.,
90: 1938-1943.
G u n t her , A l b er t C . L. G .
1860. Catalogue o f the Acanthopterygian fishes in the co llec tion o f the
British Museum. Taylor and Francis, London , 2: xxi + 548 pp.
1877. Andrew Garrett’s Fische der Siidsee, . . . J. Mus. Godeffroy, 4
( 2 ) : 129-260.
1880. An introduction to the study o f fishes. Adam and Charles Black,
Edinburgh, 720 pp.
G u ic h e no t , A l p ho nse
1843 Pece de Cuba. In Sagra Ramon de la Historia isica politica y
. natura s de a sla de Cuba ,A Bertrand, Paris , Vol. 4. f ,
G u n t e r G o r d oln l I . . ,
1942 , A list o the fishes o the mainland o Nor h and Middle America
. recordedf rom both ffreshwater and sea f water
t Amer Midi. Na
28 2 305 f 326 . . t.,
H a b e k o s t R (o b)e: r t C - I a.n M F r a s e r a n d B r u c e W H a l s t e a d
1955 Tox , co ogy—. ,Observations. on toxic
, marine algae.. J Wash Acad. Sci.,
. 45 i4 l 101 103 . .
H a l s t e a d B r( u )c:e W - .
1953. Some , general
. considerations o the problem o po onou fishes and
ichthyosarcotoxism Copeia, 1 f 31 33 f is s
H a lste a d B r uc e W a nd N o . r m a n C (B )u:n k e r- .
1954a. A , survey o . the poi onous. fishes o the Phoen x Islands Copeia,
1 1 11 f s f i .
( ): - .
162
 1954b. A survey of the poisonous fishes of Johnston Island. Zoologica, N. Y.,
39 ( 2 ) : 61 -77 .
H a l s t e a d , B r u c e W. a n d W. M. L i v e l y , J r.
1954. Poisonous fishes and ichthyosarcotoxism. U. S. Forces med. J., 5
( 2 ) : 157-175.
H ar r y, R ob ert R .
1953. Ichthyological field data o f Raroia Atoll, Tuamotu Archipelago.
A to ll Res. Bull., ( 1 8 ) : 190 pp.
H a shim ot o , Y o s h ir o
1950. On the toxicity o f a puffer, “na sh i-fugu.” Bull. Jap. Soc. sci. Fish.,
16 ( 2 ) : 43 -46 .
1956. A note on the poison o f a barracuda, Sphyraena picuda Bloch and
Schneider. Bull. Jap. Soc. sci. Fish., 21 ( 1 1 ) : 1153-1157.
H a s h i m o t o , Y o s h ir o , K . K a n n a , a n d A . S h io k a w a
1950. On shell-fish poisons. II. Paralytic poison (preliminary repor t). Bull.
Jap. Soc. sci. Fish., 15: 771 -776 .
H a s h i m o t o , Y o sh ir o a nd M asa o M i g it a
1950. Inadequacy o f acidulated alcohol s with hydrochloric acid as solvent.
Bull. Jap. Soc. sci. Fish., 16: 77 -85.
1951. The assay method o f puffer poison. Bull. Jap. Soc. sci. Fish., 16:
341- 346.
H e nr iq ue s da S i l v a, G.
1956. La faune M iocene de l’lle du Prince. Mem. Mus. min. geol. Univ.
Coimbra, 42: 29-53.
H e n s h a l l , J a m e s .A
1895. Not es on fishes collec ted in Florida in 1892. Bull. U . S. Fish Comm.,
for 1894, 14: 209 -221.
H e r r e A W. C. T.
, .
1931. A check list o f fishes from the Solomon Islands. J. Pan-Pacif. Res.
Instn., 6 ( 4 ) : 4-9.
1934. Not es on fishes in the Zoological Museum o f Stanford University . I.
The fishes o f the Herre 1931 Philippine Expedition with descriptions
o f 17 new species. The Newspaper Enterprise , Ltd. , Hong Kong,
106 pp.
1936. Fishes o f the Crane Pacific Expedition . Publ. Field Mus., ( 353 ) ,
zool. Ser. ( 2 1 ) : 427 pp.
1953. Check list of Philippine fishes. Res. Rep., U . S. Fish Wildl. Serv.,
( 20 ) : 977 pp.
H i a t t , R o b e r t W.
1947. Food- chains and f ood cycle i n Hawaiian fis h ponds.— Part 2. Biotic
interaction. Trans. Amer . Fish. Soc., for 1944, 74: 262 -280 .
H i c k l i n g , C. F. a n d E. R u t e n b e r g
1936 . The ovary as an indicator o f the spawning period in fi shes. J. Mar.
biol. Ass. U. K., new Ser., 21 ( 1 ) : 311 -318 .
H il d e b r a n d
, H e nr y H.
1954. A study of the fauna of the brown s hrimp ( Penaeus azt ecus Ives)
grounds in the western Gulf o f Mexico. Publ. I ns t. Mar. Sci. Univ.
Tex., 3 ( 2 ) : 233 -366 .
1955. A study o f the fauna o f the pinkf shrimp ( Penaeus. Iduorarum s .
Burkenroad)
. .,grounds
( ) : in the
- Gulf
. o Campeche. Publ n t. Mar
Sci Univ. Tex 4 1 172 232
,
H i l d e b r a n d S a m u e l F.
. . i l t , . ,
1916 The U. S Fisheries . Biolog
, ca Station a- Beaufort
. N C. during
1914 and 1915 Science new Ser., 43: 303 307
163
 1941. An annotated list o f salt and brackish water fishes, with a new name
for a menhaden, found in North Carolina since the publication o f
“The fishe s o f North Carolina” by Hugh M . Smith in 1907. Copeia,
( 4 ) : 220 -232 .
H il e , Ralph
1936 . Age and growth o f the cisco , Leuc ich thys ar ted i (LeSueur ), in the
lakes o f the northeastern highlands, Wisconsin . Bull. U. S. Bur. F is h ,
( 6 8 ): 211 -317 .
1941. Age and growth of the rock bass, Am bl oplit es rupestris ( Rafi nes que),
in Nebish Lake, Wisconsin. Trans. Wis. Acad. Sci. Arts L e tt, 33:
189-337.
H il l , R ic h a r d
1868. On poisonou s fishes. Proc. sci. Ass. Trinidad, 1 ( 5 ) : 210 -223 , 227-
228.
H ir o a , T e R a ng i (P et er H . B uc k)
1938. Ethno logy o f Mangareva. Bull. Bishop M u s, Honolulu , ( 157 ) : 301.
H iy a m a , Y o sh io
1943. Report o f an investigation o f poi sonou s fishes o f the South Seas.
Translated from the Japanese language by W . G. van Campen from
Spec. P ub l, Niss en Fish. Exp. S t a , Odawara Branch, 137 pp. Spec.
Sci. Rep. U . S. Fish. Wildl. Serv , ( 25 ) : 188 pp., published Wash-
ington, D . C , 1950 [without original pla te s].
H o f f m a n n , W. H .
1927. La Ciguatera, die Fischvergiftung von Cuba. Arbeiten i iber Tropen-
krankheiten und deren Grenzgebiete. Bernard Nocht, Hamburg.
1929. Los peces venenosos de Cuba y la Ciguatera. Rev. chil. Hist, n a t,
33: 28-30.
1931. La Ciguatera— enfermedad producida por peces venenoso s de Cuba .
Rev. M ed , 6: 212 -216.
, . .
H o f f m a n n W H a n d V i c e n t e A. E m b i l
1928. Observaciones sobre unos casos de ciguatera. Rev. Med. Cir. Habana,
33 ( 1 8 ) : 844-851.
H o l d e r , C h a r l e s F.
1908. Big game at sea. Outing C o , New York, 352 pp., 32 pis.
1924. The big game fishes o f the United States. The Macmillan C o , New
York, 435 pp.
,
H o l l is t e r G l o r ia
1937. Caudal skeleton o f Bermuda shallow water fishes. II. Order Per-
comorphi , Suborder Percesoces: Atherinidae, Mugilidae, Sphyraeni-
dae. III. Order Iniomi: Synodontidae . Zoologica , N . Y. 22 ( 3 ) :
265 -279 ; ( 4 ) : 385 -399 .
, .
H ub b sC arl L
1943. Terminology o f the early stages o f fishes. Copeia, ( 4 ) : 260 .
, .
H ubbs C arl L a n d C lark H u b b s
1953. An improved graphical analysis and comparison o f series o f samples.
Syst. Z o o l, 2 ( 2 ) :49-56; 92.
, .
H u b b s C a r l L a n d K a r l F. L a g l e r
1947 Fishes o f the Great Lakes region. Bull. Cranbrook Inst. S c i,
.
( 26 ) :xi -1- 186 pp.
, . , . . , .
H ug h e s E O P R G o r ham a nd A Z e h nde r
1958 Toxicity o a unialgal culture o f M icrocysti s aeruginosa. Can. J.
. f
M i crob io l, 4 : 225 -236 .
, .
H u t t o n R o b ert F s s . . , ( ): .
1959. The Florida hark tory. Fla. State Bd Cons., educ S er 1 3 3 7 pp
164
Ing e r , R o b er t F .
1955. Ecological no tes on the fish fauna o f a coasta l drainage o f Nor th
Borneo. Fieldiana , Zool., 37: 47 -90 .
1957. Repor t on a collec tion o f marine fishes from Nor th Borneo. Fieldiana,
36: 341-405 .
I n g r a m , W i l l i a m M. a n d G. W . P r e s c o t t
1954. Tox ic fresh-water algae. Amer . Midi. Na t., 52 ( 1 ) : 75 -87 .
I r v i n e , F r e d e r ic k R .
1947. The f ishes and fisheries of the Gol d Coast. The Crown Agents for the
Colonies, London, xv + 352 pp.
I sh ih a r a , F u sa o
1918. Ober die phys iologischen Wirkungen des Fugutoxins. Mitt. med. Fak.
Tokio , 20: 375 -426 .
I v e r s e n , E d w i n S. a n d H o w a r d O . Y o s h i d a
1956. Longline fishing for tuna in the Central Equatorial Pacific, 1954.
Spec. sci. Rep. U. S. Fish Wildl. Serv., ( 1 8 4 ) : 33 pp.
1957. Longline and troll fishing f or tuna in the Central Equatorial Pacific,
January 1955 to February 1956. Spec. sci. Rep. U. S. Fish Wildl.
Serv. , (20 3 ) : 38 pp .
J a c k s o n , C . H . N.
1939 The analysis o an animal population J Anim Ecol. 8: 238 246
J e n k i n .s O l i v e r P f . . . , - .
1901 , De crip ion . o fifteen new specie o fishes from the Hawaiian
. Islands.
s tBull.
s Uf S Bur Fish. or 1899 s f19: 387 404
1904 Report on collec . ion
. .o fishes
, f made , n the Hawaiian
- . Islands, with
. descriptions o new t sspecies.f Bull U i S Bur Fish., for 1902, 22:
417 511 f . . . .
J e s p e r s e n P - a n d. A V e d e l T a n i n g
1934 ,Introduction
. . to the reports from the Carlsberg Founda on ocean
. ographical expedition round the world 1928 30 Dana ti Rep., ’s 1 -
130 pp - . ( ):
Jo ne s J D
.
1956 , . Observations
. on h poisoning n Mauritius. Proc. Soc. Arts Sci
. Mauritius, 1: 367 fis 385 i .
Jo ne s S - .
1956 , . Some deaths due to h poisoning ich hyo arco oxism n India
. Indian J med. Res., 44fis2 353 360 ( t s t ) i .
. ( ): - .
Jo n e s S B
1937. , . Annals
. o Anguilla S Kitts, Briti h Wes Indies, 78 pp
f . t. s t .
J o r d a n D a v id S t a r r
1884., The fishe o Florida Keys Bull U S Fish Comm., for 1884 4:
77 80 s f . . . . ,
1885 L - o. fishes collec ed a Key West, Florida, with notes and descrip-
tions. Proc. U S nat. Mus., 7 103-150
1886.. Aistpreliminary
f list ot het fishes o the West Indies. Proc. U S na
Mus., 9: 554 608 . . : .
1904 No e on fishes collec f t ed n the fTortugas Arch pelago Bull. .U t.
S
Fish Comm., -22: . 539 544
J o r d a n. D a vti ds S t a r r a n d C h a r lt e s Hi B o l l m a n n i . . . .
1889 Lis o fishes collec -ed a. Green Turtle bay, n he Bahamas, by
, Charles L Edwards, with .descriptions o three new species. Proc.
. U tS nat. f Mus., or 1889 t t 549 553
11: i t
. f
. . f , - . 165
Jo r da n, D a v i d S t a r r a n d B a r t o n W. E v e r m a n n
1896a. A check-lis t o f the fishes and fish-like vertebrates o f North and
Middle America. Rep . U . S. Comm . Fish, for 1895 : 207 -590 .
1896b. The fishes o f Nor th and Middle America. Bull. U . S. nat. Mus., ( 47 )
Part 1: v-lx + 1240 pp.
1900. The fishes o f Nor th and Middle America. Bull. U . S. nat. Mus.,
Part 4: iii -ci 4- 3137 -3313 + 392 pis.
1905. The shore fishes o f the Hawaiian Islands, with a general accoun t o f
the fish fauna . Bull. U . S. Fish Comm., for 1903, 23: 1-574, 138 pis.
+ 229 figs.
1926. A check-list of the fishes of Hawaii. J. Pan-Pacif. Res. Instn., 1 ( I ) :
3-15.
J o r d a n , D a v i d S t a r r, B a r t o n W. E v e r m a n n, a n d H o w a r d W. C l a r k
1930. Check-list o f the fishes and fishlike vertebrates of North and Middle
America north o f the northern boundary o f Venezuela and Colombia.
Rep . U . S . Comm . Fish., for 1928, Par t 2: 1-670.
Jo r da n, D a v id S ta r r a nd C ha r l e s H . G il b e r t
1882. Synopsis of the fis hes of Nort h America. Bull. U. S. nat. Mus., ( 1 6) :
Ivi -f 1018 pp.
1883. Notes on a collec tion o f fishes from Charleston, South Carolina,
with descriptions o f three new species. Proc. U. S. nat. Mus., 5:
580-620.
J o r d a n , D a v i d S t a r r a n d C a r l L. H u b b s
1919 . Studies in ich thyology : a monographic review o f the family o f
Atherinidae or silversides. Leland Stanf. Univ . Publ., Univ. Ser. ,
( 4 0 ) : 1-87.
J o r d a n , D a v i d S t a r r a n d E r ic K. J o r d a n
1922. A list o f the fishes o f Hawaii, with notes and descriptions o f new
species. Mem. Carneg. Mus., 10 ( 1 ) : 1-92.
Jo r da n, D a v id S tar r a nd R o b e r t E. R i c h a r d s o n
1908. Fishes from the island s o f the Philippine Archipelago . Bull . U . S .
Bur. Fish. , for 1907, 27 : 233 -287 .
,
Jo r d a n D a v id S ta r r a nd C lo udsley R u t t er
. ll ti f i
1897 A co ec on o fishes made. by Joseph Seed : Roberts- . n Kingston,
Jamaica. Proc. Acad. nat. Sci Philad., for 1897 91 133
,
Jo r d a n D a v i d S t a r r a n d A l v in S e a l e
f . . . . . . . , - ,
1906. The fishes ofi s.Samoa, Bull U S Bur Fish. 25: 175 455
20 pis., 111 g
, .
J o r d a n D a v id S t a r r a n d J o h n O S n y d e r
. t s f f .
1907 No . e. on fishes is ., o Hawaii, , with descriptions
- . o new species. Bull
U S Bur, F h for 1906 26: 205 218
, . .
Jordan
. D a v i d fisS t a r r a n d fJ C T h o m p s o n . ll. . .
1905 The h , fauna o - the . Tortugas Archipelago Bu U. S Bur Fish.,
for 1904 24: 231 256
,
Jouan
.H e n rti s s s - i .
1861 No e ur quelques . espece de pois ons - .de la Nouvelle Caledon e
. Mem. Soc.
t s s Sci nat.
s Cherbourg, 8: 241 308l - i -
1863a No et sur quelque s animaux. observes a. a .Nouvelle t. Caledon e: pen .
den le annee 1861 et 1862 Mem. Soc Sci na l Cherbourg,- 9 115 i .
1863b. Supplement
. a .la description
t. des :poissons- de . a Nouvelle Caledon e
Mem t sSoc. Sci nas Cherbourg, 9 185 187 . . . t.
1867. No e ur quelque poissons - . nuisables du Japon Mem Soc. Sci na
Cherbourg, 13: 142 144
K e nda ll, W il l ia m C.
1891 . List o f Brachyura, M oll us ca and f ishes coll ect ed. In Adams, A. C.
and W. C. K endall, Report upon an i nvesti gati on of t he f ishing
grounds off the west coa st o f F lor ida . Bull. U . S. F is h Com m ., f or
1889 , 9: 289 - 3 1 2 .
K e nda ll, W il l ia m C.
1908 . Fauna of N ew England. 8. List of t he Pis ces. Occ. Pap. Bost on Soc.
nat. H ist., ( 7 ) : 1 -152 .
K err , W . M .
1912 . A no te on a cas e o f fish po is on i ng in G uam. N av. med. Bull., Was h.,
6 ( 3 ) : 401 - 4 0 2 .
K e s te v e n , G. L .
1950. A n exami nati on of cert ain as pects of t he m et hodol ogy and t heory
o f fis hery bi ol ogy. Coun. sci. i ndustr. Res. Aust rali a. Cronull a, 211
pp. [ M i m eographed .]
K i lb y , J o h n D .
1955. The fishes o f two G u lf coa sta l marsh areas o f Fl or ida . Tu lane Stud.
Zoo l., 2 ( 8 ) : 175 - 247 .
K i s s e l e v i c h , K . A.
1923. Mat eri als on t he bi ol ogy of t he Cas pi an herrings. 1. T he f ertilit y of
the V ol ga - Cas pi an herrings. Trav. Lab. icht h. Ast rakhan, 5 ( 1): 1- 55.
K l e in , J acob T h e o do r
1778. N eu er Schaup la tz den Na tur nach den r ich tigsten Beobach tungen . . . .
M. G . W e i dm ann s, Le ipz ig , V o l. 6.
K l u n z ing e r , C ar l B e nj a m in
1870. Synopsis der Fis che des Rot hen Meeres. I Theil. Percoii den- M ugili o-
iden. Verh. zoo l-bo t. Ges. Wi en, 20: 6 6 9 - 834 .
1884 . D i e Fis che des Rot hen Meeres. Ei ne kritis che R evisi on mit Besti m-
m ung s- Tabe llen. Stuttgart, Part 1, 133 pp.
K ne r , R udo l f
1879 . P isce s. In Schme ltz , J. D . E ., Wirbe lth iere (A n im a lia Ver tebra ta )
und N ach trage zu Ca ta log v & v i au s den iibr igen Th ierk la ssen . Mus.
Gode ffroy , Ca ta l ogue 7, L. Fr i edrichsen and Co ., Hamburg , v- viii +
98 pp.
K ue ne n, Ph. H.
1950. Marine geo l ogy . John W iley and Sons, N ew York , x + 4 68 pp.
L ab a t , J ean B a p t is t e
1724 . N o u v ea u voyage aux Isl es de l ’Amer i que . P. H u sson e t al., The
H ague, V o l. 1.
L a c e p e d e , B . G . E. d e
1803. Hist oi re nat urell e des poiss ons. Chez Pl ass an, I mprimeur- Librai rd,
Paris, V o l. 5, 802 pp.
L agler, K arl F.
1956 . Fre shwa ter fishery b io logy . Wm . C. Brown, D ubuque , 421 pp.
L a M o n t e, F r a nc e s c a R .
1951. A preli mi nary s urvey of marine angli ng in Nort h Carolina. In Tayl or,
Harden F ., Survey of marine f isheri es of N ort h Caroli na. U ni v. Nort h
Caroli na Press, Chapel Hill, x ii + 555 pp.
1952. Marine game fishes o f the world . D oub leday and Co ., I nc., N ew
York , 190 pp.
L a r s e n, N il s P.
1942. Te t rodon po is on i ng in Hawaii. Proc . sixt h Pac if. Sci. Congr ., f or
1939 , 5 : 417 - 4 2 1 .
L a r so n, E dw a r d a nd L u is R e n e R iv a s
1962. Barracuda (Sphyraena barracuda) po is oni ng . Fed . Proc ., 21 ( 2 ) : 36.
167
L egand, M.
1952 . £ t a t act uel et pers pecti ves de l ’indust rie du poiss on dans l es terri-
to ire s franga is du Pacifique sud. Cyb ium , ( 7 ) : 1 -23 .
L e id y , J o se p h
1855. I ndi ca ti ons o f t we l ve s pec ie s o f f o ssil f ishes. Proc. Acad . nat. Sci.
Phil ad., 7: 3 9 5 - 397 .
L i e n a r d, E l iz e
1843 . Tre i z iem e rapport. Soc . H ist. nat. Maurice .
L in t o n, E d w in
1905. Parasit es of f ishes of Beauf ort, N ort h Caroli na. Bull. U. S. Bur.
F is h ., f or 1904 , 24: 323 - 428 .
L o B ia n c o , S a l v a t o r e
1937. P la t e 35 . In Fauna e f lora del G o lf o di N apoli. Monogr., N apoli
Staz. zoo l. Pub l., ( 3 8 ) , p la te s.
L [ o c k e ] , J.
1675. On po is onou s f ish in the Bahamas I slands. Phi l. Trans ., 10: 312.
L o n n b e r g , A x e l J o h a n E in a r
1894 . List of f ishes obs erved and coll ect ed in Sout h Florida. Of vers.
Ve t s . . t ., - ; - .
is
L o on G
, . en kAkad Forh S ockh 51: 93 107 109 131
1955. P o(is onou ):
s f ishes of t he Sout h Pacif ic. Quart. Bull. S. Pacif. C omm.,
- .
5 4 28 31
l , illi . l . il
L ong ey W a m H and S a m ue F H d eb rand
. s st ti t l f t f s f t t s. . t s
1941 A y ema -c ca, a ogue . - .o he ishe o he Tor uga Pap Tor uga
Lab., 34: 1 331 Pis 1 34
L o u w , P. G . J.
1950. T he acti ve con stit uent o f t he pois onou s algae, Mic rocystis t oxi ca
St ephens. S. A f r . industr. Chem ., 4: 62 - 66 .
M a a s s , T. A .
1937. Tabu lae b io log icae . G ift-tiere . N . V . Van de Garde and Co ., The
Hague , V o l. 13.
M a c C o y , C l i n t o n V.
1931. Mus eum not es: f ishes. Bull. Bost on Soc. nat. Hist., ( 5 8 ) : 3- 22.
M a c G o w a n , D a n i e l J.
1887. N o t e s upon fish and t he f isheri es. Pois onou s fish in Chi na. Bull. U . S.
Bur. Fis h., f or 1886 , 6: 130 - 131.
M ac L eay, W il l ia m
1881 . D es cri pti ve cat al ogue of Aust rali an f ishes. F. W. Whit e, Sydney.
2 vo ls. in 1; Part 2, 323 pp.
1882. Des cri p ti ve cat al ogue o f t he f ishes o f Aust rali a. Proc. Linn. Soc.
N . S. W ., 6: 1- 138; 2 0 2 - 387 .
M a lp a s , A . K.
1926 . The mari ne bi ol ogi cal s urvey of t he litt oral wat ers of Ceyl on. Bull.
C eyl on Fis h., 2: 13- 165.
, .
M a n a c o p P R.
1936. The. s exua l t it
ili .ma. uri., y o
f s om e com m erci a l f ishes caught in Manila
- .
Bay Ph pp J Sc 59: 383 391
, .
M a n n W L.
. is is i l - i i Isl s . . .,
1938 F h po on ng in Cu ebra V rg n and area. N av med Bull
s ., - .
Wa h 36 : 631 634
,
M a r r J o h n C.
1955 . The us e of morphomet ri c dat a in s yst emati c, racial and relat ive
grow t h st ud ie s in f ishe s. Cope ia , ( 1 ) : 23 -31 .
s ll,
M ar ha T o m C.
1953. I ch t hyo log ica l no te s. D ep . Harb . Mar . Qd., ( 2 ) : 1 -73 .
168
M a r t in , W . R.
1949 . T he m echani cs of envi ronmen tal cont rol o f body f orm in fishes. Publ.
Ont. Fis h. Res. Lab., biol. Ser., ( 5 8 ) : 1-91.
M a so n, M . F. a nd R. E. W heeler
1942 . Ob serva tion s upon the t ox ic ity o f blue -green algae. Fed. Proc. Amer .
Soc . biol. Chem ., 1 ( 2 ) : 124.
M a t h e r , F r a n k J., I l l
1952. Sport fis hes of t he vi ci nit y of t he G ulf of Honduras, cert ain Carib-
bean Isl ands, and Carmen, M exi co. Proc. G ulf & Carib. Fis h. Inst.,
Four th Ann . Sess., f or 1951 : 118 - 128 .
M a t h e r , F r a n k J ., I l l a n d C. G o d f r e y D a y . .
1954. Ob servati ons o f pelag ic f ishes o f t he tropical A tl anti c. Copeia , ( 3 ) :
179 - 188 .
M a t suo , R ik u ic h i
1934 . Report of an i nvesti gati on on t he pois onous f ishes of Jaluit Isl ands.
Tran sla ted from the Japane se language by W. G . van Campen , from
Med . Trea tise , Sou th Sea s Co lon . Office , (2 ) : 309 -326 . U . S . F ish
Wildl. Serv. [ M i m eographed .]
M a x w e l l , C. N .
1921 . M a layan fishe s. J. Straits Br. R. A siat. Soc., 84: 179 - 280.
M c F a r r e n, E. F . and A . F . B ar tsc h
1960 . A pp lica ti on o f the para ly tic shellfis h pois on assay t o po is onou s fishes.
J. A ss. off. agric. Chem . W a sh ., 43: 548 - 554 .
M eek, A l e x a nde r
1916 . The m igra ti ons o f fish. Edward Arnold , London , 427 pp.
M e e k , S e t h E. a n d S a m u e l F . H i l d e b r a n d
1923 . The mar ine f ishes o f Panama . Publ. F ie ld Mus., 15 ( 2 1 5 ) , Part 1:
330 pp.
M e e k , S e t h E. a n d R o b e r t G . N e w l a n d
1884 . A rev iew o f the Amer ican spec ie s o f the genu s Sphyraena . Proc .
Acad . nat. Sc i. Ph ilad ., f or 1884 : 67 -75 .
M e n dis, A . S.
1954 . F is he s o f C ey lon ( a ca ta log , key and b ib liography ). F ish . Res. Sta.,
D ep . F is h ., Bu ll., ( 2 ) : 1 -222 .
M e no n , M. D e v id a s
1953 . The de term ina tion o f age and grow th o f fishe s o f tropica l and sub -
trop ica l water s. J. Bombay nat. His t. Soc ., 51 ( 3 ) : 623 -635 .
, J an
M etzel aar
1919 . Over trop isch A tl antis ch viss chen. A. H. Kruyt, Am st erdam, 314 pp.
i it , . .
M g a M and K K anna
1957. On t he s ource of s hell-f ish pois on at t he Lake Hamana. 1. Att em pts
at experi ment all y rendering s hell-fish t oxi c. Bull. Jap. Soc. sci. Fis h.,
23 ( 4 ) : 215 - 2 2 1 .
M il l s
, A . R.
1956 . P o iso n o u s fish in th e sou th P ac ific . J. trop . M ed . ( H y g .) , 59: 9 9 -103 .
M o h e r W A.
l , .
1941 . Een b lauww ieren - phenom een aan het strand van Ba likpapan . Natuurk.
Tijdschr. N ed .-I nd ., 101 ( 3 ) : 75 - 79 .
M o nt r o u z ie r , P e r e X a v ie r
[M S ] Essa i sur la faune de l’ile du W ood lark ou M o iou . 1856 .
1857 . E ssa i sur la faune de l ’ile du Wood lark ou M o i ou . Imprimerie de F.
D um ou lin , Lyon , 226 pp.
M o r eau de J o n n e j, A l e x a ndr e
1821 . R echerche s sur le s po isson s tox icophore s de s Inde s Occ iden ta le s.
Impr imer ie de M ignere t, Paris, 34 pp.

169
M o r ic e , J ean
1958 . A ni m aux marins com esti bl es des A ntill es f rangais es. ( Oursi ns,
Cru stace s, M o llu sque s, Po isson s, Tor tue s e t C e t a ce s). Rev. Trav.
Off . Peche s marit., 22 (1 ) : 85 -104 .
M o r r is , R o b ert W .
1954. Some as pects of t he probl em of reari ng marine f ishes. D oct. Diss.,
Stanf. Un iv . Pub lished in 1956 in Bull . Inst , oceanogr ., M onaco ,
( 1 0 8 2 ) : 1-61.
M or r o w, Jam es E.
1954 . F ishe s from East A f r ica , wit h new records and des cri pti ons o f t wo
new s pec ie s. A nn . Mag. nat. H ist., Ser. 12, 7: 7 9 7 - 820 .
M o r t o n, C har l es
1868 . D iarrhoea as a result o f fish po ison ing at Na ssau , Bahama s. Statist .
Rep. Hlth . N avy , Lond., f or 1865: 107- 108 .
M o se l e y, H e nr y N.
1839 . N ot es by a nat uralist on t he “C hall enger, ” bei ng an account of . . .
the voyage o f H . M . S. “C hall enger” . . . , in t he years 1872 - 1876 ,
. . . M acm illan and Co ., London , xv i + 620 pp.
M o w b r ay, L o uis L.
1916 . F ish po ison ing (ich t h y o t o x is m u s). Bull. N . Y . zoo l. Soc ., 19: 1422 -
1423 .
1922 . The barracuda. Bull. N . Y . zoo l. Soc ., 25: 143 - 144 .
M uller , J o h a nne s a nd F r a nz T r o sc h e l
1848 . F ishe s. In Schomburgk , Rober t H ., The h istory o f Barbados; . . . .
Longman , Brown , Green , and Longman , London , xx + 722 pp.
M ue nsc h e r , W al ter C.
1951 . Po is onou s plants o f t he U n it ed States . Rev . Ed., The Macm illan Co .,
N ew York , xv ii + 277 pp.
M unr o , I a n S. R.
1955 . T he marine and fresh water f ishes o f Ceyl on . D ep . ext ern. Affairs,
Canberra , xv i + 351 pp ., 56 pis .
M u r r h y , G a r t h I. a n d R i c h a r d S. S h o m u r a
1953a. Longli ne fis hing f or deep -s w i mm i ng t unas in t he cent ral Pacif ic, 1950-
51 . Spec . sci. Rep . U . S. F ish Wildl. Serv ., ( 9 8 ) : 75 pp.
1953b. Long line f ishing f or deep -sw im imng t unas in the central Pacific ,
January-June 1952 . Spec. sci. Rep. U . S. F is h Wildl. Serv., ( 1 0 8 ) :
32 pp.
M ur r h y , R ob er t C .
1914 . N o t e s on pe lag ic fishes. Cope ia , ( 6 ) 1-3.
N a g a i, J. a nd T . Ito
1939 . Chemi cal st udy of f ugu ( Sp heroides) pois on; nit rogen dist ribut ion i n
purifi ed pois on. J. Bi ochem., T ok yo, 30: 23 5- 2 3 8.
N e w e l l , N o r m a n D . a n d J o h n I m b r ie
1955 . B iogeo log ica l reconna issance in the B im in i area, Grea t Bahama Bank .
Trans. N . Y. Acad . Sci., Ser. 2, 18 ( 1 ) : 3 - 14 .
N ic h o l s , J o h n T.
1912 . N o t e s on Cuban fishes. Bull. Amer. Mus. nat. H ist., 31 ( 18): 179- 194.
1921 . A li st o f Turk Island fishes, with a descr ip ti on o f a new flatfish. Bull .
Amer . Mus. nat. H ist., 44 ( 3 ) : 21 - 24 .
1929 . The f ishes of Puert o Ri co and t he Vi rgi n Isl ands: Branchi ost omi dae
to Sci aeni dae. Sci. Surv. P. R., 10 ( 2 ) : 161- 295.
N ic h o l s, J o h n T. a nd C h a r l e s M. B r e d e r, J r.
1927. The mari ne f ishes of N ew York and s out hern N ew Engl and. Zool-
og i ca , N . Y ., 9 ( 1 ) : 1-192 .
170
N ig r e l l i, R oss F . a nd P a ul A . Zahl
1952. Som e b i o l og ica l charac teristic s o f ho lo thur in . Proc Soc . exp . B i o l,
N . Y , 81: 379 - 380 .
N i k o l ’s k i ,i G eorg V.
1954. Chast naya i khti ol ogi a. [ Speci al i cht hyol ogy.] Gos udarst v. I z d , “Sovi et-
s kaya N auk a, ” M os cow, 458 pp.
N or m a n, J o h n R.
1957 . A draft synop sis o f the orders, f am ilie s and genera o f recen t fishe s and
fish-l ike ver tebra te s. Brit. Mu s. (na t. H is t.), 649 pp . [M im eo -
graphed .]
N o r m a n, J o h n R. a nd F. C. F r a s e r
1938. G ian t f ishes, wha le s and do lph in s. W. W. N or t on and C o , N ew York ,
376 pp.
O g i l b y , J. D o u g l a s
1915. The com m erci a l f ishes and f isheries o f Queensland . Brisbane.
Om m a nne y , F. D .
1953. The pel agi c f ishes and a not e on t ow netti ngs. In Wheel er, J. F. G.
and F. D. Ommanney. Report on t he M auriti us- Seychell es fis heri es
survey , 1948 -49 . Co lon . Off . F ish . P u b l, 1 ( 3 ) : 148 pp.
O ’N e i l l , J. B.
1938. Food po is on ing in the first marine brigade, f leet mari ne f orce , Cu le -
bra, P. R. N av. med. B u ll, W a s h , 36: 629 - 631.
O f p e n h e i m e r , C a r l H.
1955 . The eff ect o f marine bact eri a on t he devel opm ent and hat chi ng of
pe lag ic fish egg s, and the con tro l o f s uch bacteria by an tib iotics.
Cope ia , ( 1 ) : 43 - 49 .
O r t o n , G ra c e
1955. Early deve lopm en ta l stages o f the Ca lif orn ia barracuda, Sphyraena
argent ea Girard . Ca lif. F is h Gam e , 41 ( 2 ) : 167 - 176 .
P a e t r o, S id n e y
1956 . F ood po ison ing cau sed by the great barracuda . Publ. Hlth. R e p ,
W a s h , 71 ( 9 ) : 933 - 937 .
P a l o m b i, A rt ur o a nd M a r io S a ntar elli
1953. Gli animali com m esti bili dei mari d’lt ali a. Ul ri co H oepli, Mil ano,
vi + 349 pp.
P a p p e K a r l W. L.
,
1853 . Synop sis o f the ed ib le fishe s o f the Cape o f G ood H ope . Van de Sand t
de V illier s and Tier , Cape T ow n , 34 pp.
P ar r , A l b ert E.
1930. T e leo stean shore and sha llow -wa ter f ishes from the Bahamas and
Turk s Is land . Bull . B ingham oceanogr . C o ll, 3 ( 4 ) : 1 -148.
1933. A geograph ic -eco log ica l ana ly sis o f the sea sona l change s in tempera -
ture cond iti on s in s hall ow wat er al ong t he A tl anti c Coast of t he
U n it ed Stat es. Bull Bi ngham oceanogr . C o l l , 4 ( 3 ) : 1-90.
1956 . On the or igi na l varia tes o f t axonom y and their regre ss ions upon size
in fishes. Bull. A m er . Mu s. nat. H is t, 11 0 ( 5 ) : 373 - 397 .
P ar r a, A n t o nio
1787 . Des cri pci on de dif erent es pi ezas de hist ori a natural, las mas del
ramo mariti mo, . . . I mpren ta Capitama Genera l, Havana , 197 pp.
P a uc a , M ir c e a
1935 . Fis che aus dem Mi ozanen D azitt uff Siebenbii rgens. Bui. Soc. roman.
G e o l , 2: 2 26 .
P a v l o v s k i i, E v g e n i i N .
1927 . G iftti ere und i hre G ifti gkeit. G ust ave Fis cher, Jena, xv i + 516 pp.
171
P e l l e g r in , J a c q u e s
1899 . Le s po isson s veneneux . The sis 510 . O llier -Henry , Paris, 121 pp ., F ig s.
1 -16 . (R ev iew ed in Rev . sc i., Paris, Ser. 4 , 14: 178 -180 , 1 9 0 0 ).
P h i l l i p s , C r a i g a n d W i n f i e l d H. B r a d y
1953. Sea pests: pois onou s or harmf ul sea lif e o f Fl or ida and the West
Indies. Un i v. M iam i Press, Coral Gab les, 78 pp.
P h i l l i p s , E l m e r S. a n d D w i g h t A . W e b s t e r
1960 . Improvemen ts in a micropro jec tor f or fish scale s. Progr. F is h Cu lt.,
22 ( 1 ) : 24 - 29 .
P h i s a l i x - P i c o t, M a r ie
1922 . A n im aux ven im eux e t ven in s. Chap . 9. Po isson s tox icophore s. M a s-
son e t C ie ., Paris, Vo l. 1, xxxv -f 656 pp.
P i c t e t , F r a n c o is -J u l e s
1854 . Traite de pal eont ol ogie ; . . . . J.-B. Bailliere, Paris, Ed. 2, Vo l. 2, 727
pp.
P ik e , N ic h o l a s
1873 . Sub-t ropical rambles in t he land o f t he Aphanapt eryx . . . . Harper
and Bros., N ew York , 5 1 0 pp.
P in t ne r , T heo do r
1930. W enigbekannte s und unbekann tes von Russ elbandwurmen . S. B.
Akad. Wiss. W ien, 139 ( 7 ) : 4 4 5 - 5 3 7 .
P l a y f a i r , R. L a m b e r t
1866 . Sphyraeni dae. ln Playf air, R. Lambert and Al bert C. L. G. Gunt her,
The fishes of Zanzibar. John Van Voorst, London, 153 pp. + 21 pis.
Plum ie r , C har les
[M S ] Zoographia americana , p isce s et vo la tilia con tinen s, auc tore R. P. C.
Plumier. [ca . 1 6 9 5 ], 3 vols.
P oey y A lo y, F e l ip e
1858. Poiss ons de Cuba, es peces nouvell es. M emori as sobre la hist oria
natural de l a Isl a de Cuba, . . . I mprent a de l a Vi uda de Barci na,
Havana , Vo l. 2: 115 -441 .
1864a . Enumera tion o f the fish de scribed and figured by Parra, sc ien tifica lly
named by Feli pe Poey . Translated and edit ed by J. C. Brevoort. Proc.
A cad . nat. Sci. Philad., f or 1863 : 174 - 180 .
1864b. D e scr i p ti on s des po iss on s nouve ll es ou peu connu s. Proc . Acad . nat.
Sci. Philad ., for 1863 : 180 -188 ; 227 .
1866 . Ci guatera; memoria sobre la enf ermedad ocasi onada por l os peces
veneno s o s. Repert. fis.-na t. Isla de Cuba, 2: 1-24; 57 - 69 .
1868 . Synopsis pis ci um cubensi um. Revista del os ti pos Cuveri anos y Val en- i
c enni anos corres pondi ent es a l os peces de la Isl a de Cuba. Repert.
fis - nat. Isla de Cuba , 2 : 2 7 9 - 468 .
1875 . Enumera tio p isc ium cuben sium . An . Soc . esp. Hist, nat., 4: 75 - 1 6 1 .
1881 . Peces. In Gund lach, Juan, A pun te s para la f auna puerto-riquena. An .
Soc . esp. H ist, nat. , 10: 317 -350 .
P r a k a sh , A . a nd J. C . M edc o f
1962 . Hydrograph ic and m e teoro log ica l fac tor s affecting shellfis h t ox ic ity
at Head Harbour, N ew Bruns wick, J. Fis h. Res. Bd. Canada , 19 ( 1 ) :
101 - 112 .
P r e s c o t t , G e r a l d W.
1948 . Obj ecti onabl e algae with ref erence t o t he killi ng of fish and ot her
animals. Hydrob io log ia , / ( 1 ) : 1 -13 .
Q u i l i c i, F o l c o
1954 . The b lue con tinen t. Rinehart and Co ., N ew York , 246 pp.
172
R af in e s q u e - S c h m ,
a l tz C o n s t a n t i n e S.
1810 . I nd ice d ’ittio log ia sic iliana . . . . Pre sso G iovann i del N obo l o ,
Messina , 70 pp.
R a n d a l l , J o h n E.
1952 . I nvestigati on o f t he i cht hyo f auna o f Onot oa, Gil bert Isl ands. In Ban-
ner, A . H . and John E. Randa ll, Prelim inary repor t on marine
b io logy study o f O no toa A to ll, G ilber t Islands. A t o ll Res. Bu ll., ( 1 3 )
Part 2: 43 - 62 .
1955 . F is he s o f the G ilber t Islands. A to ll. Re s. Bu ll., ( 4 7 ) : x ii + 243 pp.
1958 . A rev iew o f cigua tera , trop ica l fish po ison ing , with a ten ta tive ex -
p l ana ti on o f its cau se . Bull. Mar. Sci. G u lf & Carib ., 8 ( 3 ) : 2 3 6 - 267 .
R e e v e , H e n r y F.
1912 . The Gamb ia ; it s h istory , . . and a de scr ip tion o f the birds, beasts
and fishes f ound therein. Smith and El der, London , 287 pp.
R e g a n, C . T a t e
1908. Bi ol ogi a Cent rali- Ameri cana. Pis ces. Tayl or and Francis, London,
xxx ii + 293 pp., 26 pis.
1912 . N o te s on the c la ssifica tion o f the te leo stean fishes . Proc . int. zoo l.
Congr ., f or 1907 , 7 : 838 - 853 .
R e id , G eorge K ., Jr .
1954. A n eco l og i ca l st udy o f t he G u lf o f M ex ico fi shes in the v ic in ity o f
Cedar Key, Fl ori da. Bull. Mar. Sci. G u lf & Carib ., 4 ( 1 ) : 1 -94 .
Rem y, C.
1883 . Sur le s po isson s tox ique s du Japon . C. R. Soc . B io l., Paris , Mem .,
Ser. 7, 35: 1- 28.
R e n a r d, L o u is
1754 . Hist oi re nat urell e des pl us rares curi osit ez de la Mer des I ndes.
Po isson s, ecrev isse s e t crabe s, . . de s Isle s M o luque s, e t sur le s co te s
des terres Au stra le s . . . . Rein ier and Josue O ttens, Am sterdam , 7 ,
2 vols. in 1, xv iii- b 100 col. pis.
R ib e ir o , A l ip io d e M ir a n d a
1915 . Fauna bra silien se— Pe ixe s. V o l. 5 (E leu therobranch io s a s p irophoro s)
— Phy s oc listi. Pogonocoe li. Arch . Mus. nac. R io de J., 77 : 1-4.
1918 . Fauna bra silien se— Pe ixe s. Vo l. 5 , Part 3. B ib liographia e ind ice .
A rch. Mus. nac. R i o de J., 21: 37 - 227 .
R ic h a r d s o n, J o h n
1836 . The fish. In Fauna Borea li-Amer icana ; or the zoo logy o f t he northern
parts o f British A m eri ca , . . . Richard Bentl ey, London, Part 3,
326 pp.
1861. On the po is onou s e ffect o f a small porti on o f t he li ver o f a D i od on
inhab iting t he seas o f sou thern A f r i ca . J. Linn. Soc . ( Z o o l.), 5 :
2 1 3 - 216 .
R i c k e r , W i l l i a m E.
1948. M et hod s o f esti m ati ng vital st atisti cs o f fish popul ati ons. Ind. U ni v.
Publ., ( 1 5 ) : 101 pp.
R ie g e l , B y r o n D ., W a r r e n S t a ng e r , D o na ld M. W ik h o l m , Ja m es D. M old,
a nd H er m a n n S o m m er
1949 . Para ly tic s hellfi sh po ison . V . Primary s ource o f po ison , mar ine
p l ankt on organis m , Gonyaulax cat enell a. J. bi ol. Chem., 177: 7- 11 .
R o c he f o r t , C h a r l e s C e sa r de
1667 . H isto ire na ture lle et mora le des lie s An tille s de l’Amer ique . Chris tofle
Fourmy, Lyon , V o l. 1, xviii + 583 pp.
173
R o hl. E dua r do
1942. Fauna de scrip tiva de Venezue l a . Peces. Caracas, 4 3 2 pp.
R o se , A nto n F e r d in a n d
1793. Pet ri Art edi Angerm anni a— s ueci s ynonymi a nomi num pis ci um, . . .
G re ifswa ld , Ed . 2.
R oger, G eor ges E ug e ne H e nr i
1895. Les i nt oxi cati ons. In Bouchard, Charl es, ed., Trait e de pat hol ogi e
generate. Ma sson et C ie ., Paris, V o l. 1.
R o g e r s , J a m e s M.
1899 . Wes t Indian po is onou s fishes. Pop. Sc i. M on ., 55: 680 - 685 .
R o ig , M a r io
1957 . La “p icua” o p icuda . Mar y Pe sca , 1 ( 3 ) : 51 -52 .
R o se n, N il s
1911 . Cont ri buti on t o t he f auna of t he Bahamas. A ct a U ni v. lund., Af d. 2,
7 ( 5 ) : 1-72.
R o ss, S. G.
1947. Pre lim inary repor t on fish po is on ing at Fann ing Is land (Cen tra l Pa -
c ifi c ). Med . J. Au st., 34: 6 1 7 - 621 .
R o u g h l e y , T h e o d o r e C.
1916 . F ishe s o f Austra lia and the ir t echno logy . W. A . G u llick , Sydney ;
Technol. Mus., t ech. Educ . Ser., ( 2 1 ) : x v i + 2 9 6 pp.
1951. F is h and fis heries o f Au stra lia . A ngu s and Rober tson , Sydney , 343 pp.
R o u n s e f e l l , G e o r g e A . a n d W. H a r r y E v e r h a r t
1953. Fis hery s ci ence: its m e t hods and app lica tions. John W iley and Son s,
N ew York , xii + 4 4 4 pp.
R u p p e l l , W i l h e l m P.
1835. N eu e W irbe lt hiere zu der Fauna von A byssi ni en gehor ig , . . . F is che .
S. Schmerber , Frankf urt-am -M a in , 2 vols., 95 pis.
Sa uv a g e , H e nr i E.
1865 . Les po iss on s et le s cru stace s. In Brehm , A . E ., Merve ille s de la
nature . Par is, 6: 91 -618 .
1891. H istoire na turelle des poiss on s. In Grandi di er, A lf red , H ist oire
phy sique , nat urelle et po litique de Madaga scar . LTmpr imerie N a -
tiona l e, Paris, 16: 543 pp.
S a v i l l e - K e n t, W il l ia m
1893 . The great barrier reef o f Austra lia ; its produc ts and po ten tia litie s.
W . H. A llen , London , xv ii -f 387 pp.
SCHLESINGER , G U N T H E R
1911 . Schw i mmen und Schw imm typen flinker Meere sfische . K o s m o s, S tu tt-
gart, 8: 2 8 4 - 287 .
Sc hm id t , J o h a nne s
1929 . In troduc tion to the oceanograph ica l repor ts inc lud ing li st o f the sta -
tion s and hydrograph ica l ob serva ti ons. The D an is h “D ana”— exped i-
tion s 1920 - 1922 in t he N or t h Atl anti c and t he G u lf o f Panama.
Oceanogr . Rep. “D ana” Exped ., ( 1 ) : 1 -87 , Pis. 1-6.
SCHROEDER, W I LL I A M C .
1924. Fis heri es o f K ey West and t he clam indu stry o f sou thern Fl ori da.
Rep. U . S. Comm . F is h ., f or 1923 , App . 12 : 74 pp.
S c h u b e r t , R. J.
1906 . D ie F ischo to lithen de s o sterre ich -ungar ischen Ter tiar s. III. Be schre ib -
ung der ubri gen Ot olit hen des ost erreich- ungarischen Tert iars. Jb.
geol. Reichs anst., ( B u nd e s an st.), Wi en, 56: 643- 706 .
174
S c h ul t z , L e o na r d P .
1943. F is he s o f t he Phoen i x and Samoan island s co llec t ed in 1939 during
t he exped ition o f the U . S. S. Bu s hne ll. Bull. U . S. nat. M us., ( 1 8 0 ) :
316 pp.
1949 . A f ur ther con tr ibu tion to the i ch t hyo l ogy o f Venezue la . Proc . U . S.
nat. M us., 99: 1 -211 .
1953 . Sphyraen idae . In Schu ltz , Leonard P ., Ear l S. H era ld , Erne st A .
Lachner, Ar thur D . We lander , and Loren P. W ood s, F ishe s o f t he
Mars ha ll and Mar iana s island s. Bull . U . S. nat. Mu s., ( 2 0 2 ) :
xxx ii + 685 pp.
S c h ul tz , L e o na r d P. a nd E d it h M . S t e r n
1948 . The way s o f f ishes. D . van No strand Co ., N ew York , 2 6 4 pp.
S e ale, A l v in
1912. Ed it orial: Som e po is onou s Ph ilipp ine fishes. Ph ilipp . J. Sci., 7D:
2 89-29 3.
S h a t t u c k , G e o r g e B ., e d .
1905. The Bahama Islands. The Macm ill an Co ., N ew York, xxx ii + 630 pp.
Shaw , G eorge
1804 . Genera l zoo l ogy or sy stema tic natural h istory , . . . P isce s. Thom a s
D av is on , London , V o l. 5, Part 1, x + 2 5 0 pp.
Sh m id t , P y o t r I u.
1930 . A check -list o f the f ishe s o f the R iu - K i u Islands. J. Pan - Pac if. Res.
In stn ., 5 ( 4 ) : 2 -6 .
S h o m u r a , R i c h a r d S . a n d G a r t h I. M u r p h y
1955 . Long line fish ing f or deep -sw i m m ing t una s in t he central Pacific, 1953 .
Spec. sci. Rep. U . S. F is h W ild l. Serv., ( 1 5 7 ) : 70 pp.
S i g l e r , W i l l i a m F.
1949 . L if e h istory o f the w h it e bass, Le pi be m a chrysops ( R a fi n e s q u e ), o f
Spirit Lake, Iowa . I owa St. Co ll. A gr ic ., Ent. econ . Zoo l. Sect., Res.
Bull., ( 3 6 6 ) : 2 0 1 - 2 4 4 .
S i m m o n s , J a m e s S ., T o m F. W h a y n e , G a y l o r d W . A n d e r s o n , H a r o l d M.
H o r a ch , and c o lla b o r a t o r s
1944 . G l oba l ep idem io logy . J. B. L ipp inco tt, Ph ilade lph ia , 504 pp.
S lo a ne , H a ns
1725 . Na t ura l h istory o f Jama ica . London , V o l. 2.
S m ith , F. G . W a l t o n , R o b e r t H . W i l l i a m s , a n d C h a r l e s C . D a v is
1950. A n eco l og ical s urvey o f t he s ubt ropi cal i ns hore waters adjacent to
M iam i. Eco logy , 31 ( 1 ) : 119 -146 .
Sm i t h, H ug h M.
1898. The f ishes f ound in t he vi ci nit y o f W oods H o l e . Bull. U . S. Fis h
C om m ., f or 1897 , 17: 85 - 111 . *
S m i t h , J. L. B.
1956 . The f ishes o f the f am il y Sphyraenidae in the W e stern I nd ian Ocean .
D ep . Ich th ., Rhode s U n i v ., ich th . Bu ll., ( 2 ) : 37 - 46 , P is. 1-2.
S m i t h , R. O.
1947 . Survey o f the fisheri es o f the f orm er Japane se manda ted Island s. F ish .
Lea fl., W as h., ( 2 7 3 ) : 105 pp.
S n y d e r , J o h n O.
1904 . A ca ta l ogue o f t he s hore ns nes coll ec t ed by t he st eamer “Al bat ross ”
abou t the H awa iian Island s in 1902 . Bull. U . S. Fis h. C om m ., 22:
5 1 3 - 538 .
1912 . The fishes o f Ok i nawa , one o f t he R i u -K i u Island s. Proc . U . S. nat.
Mu s., 42: 4 8 7 - 5 1 9 .
175
S o n n e r a t , P ie r r e
1774. Sur quel ques poiss ons de l ’ll e de France qui em pois onnent ceux qui
le s mangen t dan s un cer tain temp s de l’annee . J. Phy s. Ch im . H ist,
nat., 3: 227 - 2 2 9 .
S p r in g e r , S t e w a r t
1943. Sharks and t heir behavi or, wit h parti cul ar ref erence t o ei ght genera
i p lica ted in repor ts o f a ttack s on man . Coord . Re s. D eve op
m l .,U .S N.
a v y Emerg .R e cues Equ pm i Sec
. t.,W a hs ng
i on
t ,D .C., 31 pp.
S p r ing e r , S t e w a r t a nd H a r v e y R . B u l l i s , J r.
1956 . Co llec tion s by the O R E G O N in the G u lf o f M ex ico . Spec . sci. Rep .
U . S. F is h W il d l. Serv., ( 1 9 6 ) : 134 pp.
S p r ing e r , V ic t o r G. a nd A nd r e w J. M c E r l e a n
1961 . Taggi ng of great barracuda, Sphyraena barracuda ( W al bau m).
Trans. Amer . Fish. Soc ., 90 ( 4 ) : 4 9 7 - 5 0 0 .
S p r ug e l, G eor ge
1954 . Grow t h o f bl uegills in a l ake wit h parti cul ar ref erence t o f als e an-
nuli. Trans. A m er. Fis h. Soc., f or 1953, 83: 58- 75.
S t a r k s , E d w in C .
1899 .The o st eo log ica l charac ters o f t he fishes o f t he s ub-order Perce s oces.
Proc. U . S. nat. Mus., f or 1899 , 22: 1- 10.
1913. The f ishes o f t he St anf ord expediti on t o Brazi l. St anf. U ni v. Publ.,
( 1 2) : 77 pp. + 15 pis.
S t e in d a c h n e r , F r a n z
1901. F ische aus dem S tillen Ozean . Ergebn isse e iner Re ise nach dem
Pacific ( Schau i nsland 1 8 9 6 - 9 7 ). Denk schr . Akad . Wis s. W ien , for
1900 , 70: 316 - 318 ; 4 8 3 - 5 2 1 .
St ep h e ns o n , J ohn
1838 . M edi ca l zool ogy and mi neral ogy, . . . John Churchill, London,
v i + 345 pp.
S t e y n , D ouw G .
1945 . Po is on ing o f an ima ls and human be ing s by algae . S. A fr . J. Sc i., 41:
24 3 - 2 4 4 .
S t o r r , J o h n F.
1957 . Progress of recovery of t he com m erci al s ponge beds of Fl ori da. Proc.
G u lf & Carib. Fish. I nst., nin th Ann . Sess., f or 1956 : 120 - 127 ; 133 -
135.
S t r o n g , R ic h a r d P .
1942 . Stitt’s di agnosis, preventi on and t reat ment of t ropi cal dis eas es. The
B l ak iston Co ., Ph ilade lph ia , Vo l. 2, v ii + 87 2 - 1 7 4 7 .
S u m n e r , F r a n c i s B., R. C . O s b o r n , a n d L. J. C o l e
1913 . A b i o log ica l survey o f the waters o f W ood s H o le and v ic in ity . Secti on
3. A cat a l ogue o f t he marine f auna. Bull . U . S. Bur. F is h ., 31,
Part 2: 547 - 7 9 4 .
S v e r d r u p H a r a l d U ., M a r t i n W. J o h n s o n , a n d R i c h a r d H . F l e m i n g
1946 . The oceans: t heir physi cs, chemist ry and general bi ol ogy. Prenti ce
H all, N ew York , x + 1087 pp.
S w a in , J o se p h
1882 . A n i dentifi cati on of t he s peci es of f ishes des cri bed in Shaw’s “G en-
eral Z o o logy .” Proc . Acad . nat. Sci. Ph ilad ., f or 1882 : 303 - 309 .
S w a in s o n , W il l ia m
1839 . The nat ural hist ory of f ishes, amphi bi ans and rept iles, or m ono-
cardian ani mals. In Lardner, Di onys us, Cabi net Cycl opedi a, Long-
man , Orme , Brown , Green , and Longm an , London . 2 : 1 -452 .

176
T a ha r a , Y o s h is u m i
1910. Uber das Te t rodongilt. Bi ochem . Z , 30: 255 - 275 ; 506.
T a k a h a sh i, J u n t a r o a nd Y o s h it o I no ko
1889 . Exper i m ente lle Un ter suchungen iiber das Fugug ift. E i n Beitrag zur
Kenn tn is der F ischg ifte . Arch. exp . Path. P harm ak , 26: 401 - 418 ; 453 .
T a n i, I w a o
1945. T ox i co l og i ca l st udie s on Japanese ball oonfis h. Imp. Chem . Corp.,
T okyo , 2 ( 3 ) : 1 -103 .
T a n o u s , J o h n H. a nd Pa ul I la no
1960 . Preda tory fish bites in Sou th Florida . Amer . S u rg , 26 ( 7 ) : 443 -4 4 5 .
T h i o l l ie r e , V ic t o r J.
1856. Partie ich thyo log ique de la faune de File Wood lark ou M o iou
( M e la n e si e ). Revue et m is e en ordre. I mpr imer ie de Barret, Lyon,
89 pp. [Ba s ed on MS o f Mont rouzier, 1856.]
T ho m pso n, E. F.
[ 1 944? ] The f isheri es o f British Honduras. Bull. D eve l opm . W elf. W. I n d ,
( 2 1) : 32 pp.
[ 1945? ] The f isheri es o f Cayman Islands. Bull. D eve l opm . W e lf. W. I n d ,
( 2 2) : 33 pp.
T h o m s o n , W. K , A . C. L a i n g , a n d G . A . G r a n t
1957. T oxi c algae. I V. Is ol ati on of t oxi c bact erial cont ami nants. Det. Res.
K ing st on Lab . ( C a n a d a ), R e p , ( 5 1 ) : 1-7.
T r u i t t , R e g i n a ld V , B a r t o n A . B e a n , a n d H e n r y W . F o w l e r
1929 . The fi shes o f Maryland . Maryl and Cons erv. B u ll, ( 3 ) : 120 pp.
T y b r ing , O sc a r
1887. Pois onous fish. Bull. U. S. Fish C o m m , f or 1886, 6: 148- 151.
[Transla ted f rom N or s k Fis ker iti dende , Bergen, Oct ober, 1885.]
U c h i d a , K e i t a r o , S a d a h i k o I m a i, S a t o s h i M i t o, S h i r o F u j i t, a M a s a a k i
U e n o , Y o i c h i S h o j i m a, T e t s u s h i S e n t a, M a s a h a r a T a h u k u, a n d Y o s i e D o t u
1958. Studie s on the egg s, larvae and j uven ile o f Japanese fishes. Ser. 1.
Second Lab. Fish. B i o l, Fish . D e p , Fac . A g r i c , Kyu shu Un iv .,
89 pp., 86 pis.
U. S. D e p a r t m e n t o f C o m m e r c e
1920 - F is hery indu strie s o f the Un ited States. F isher ies Docum en ts.
1938 .
U . S . F is h a nd W il d l if e S e r v ic e
1952 - F lor ida landing s. Dep t. Int., Branch comm . F ish ., and Fla . St. Bd.
1957 . Con serv .
V a l e n t i j n , F r a n c o is
1724. Bes chryvi ng van oud- en ni euw O ost-l ndi en, vervatt ende een naauw-
keurige en u itvoer ige verhande ling van Neder land s m ogen theyd in
d ie gewe sten , . . , Dordrech t and Am sterdam , 5 v o ls , 1050 pis.
V a n O o ste n, J o hn
1921. The whit efis hes ( Coregonus clup eaform is) . A st udy of t he s cal es of
whitefis hes o f known ages. Zoo log ica , N . Y , 2 ( 1 7 ) : 380 - 412 .
1929. L if e history o f t he lake herring ( Leucichthys arted i LeSueur ) o f
Lake Huron as reveal ed by its s ca les, w ith a criti que o f the sca le
m e t hod . Bull . U . S. Bur. F is h ., 44: 265 - 428 .
1953. A modifica ti on in the t echni que o f com pu ti ng average leng t hs f rom
the s cal es of fishes. Progr. Fish Cult., 15 ( 2) : 85- 86.
177
V a ug h a n, T ho m a s W .
1918. The temperat ure o f t he Fl ori da coral reef tract. Pap. Tort ugas Lab.,
9: 319 - 3 3 9 , Fi gs. 7 -9.
V l L L A D O L I D , D EOGR AC I A S V .
1940. Philippi ne f isheries and probl em s o f t hei r cons ervati on. Proc. sixth
Pacif. Sci. Congr ., 3: 369 - 389 .
V i n b e r g , G . G.
1954 . Tok siche sk ii fitop lank ton . A dv . mod . B i ol., M os cow , 38, Part 2 ( 5 ) :
216 - 226 . [ Transla t ed f rom t he Russi an by G. Bel ov, Nat. Res. Counc.
Canada, tech. Trans l. TT - 549 : 25 pp.]
V i n c e n t , H.
1910 . The sea fish o f Trinidad . J. J. L ittle , Inc ., N ew York , 97 pp.
V l a m in g , C o r ne il l e de
1715. Zee - Tooneel, ( 6 ) .
V o n F r a e n k e l , P. V . a n d E. S. K r i c k
1945. F ish po is on i ng by barracuda in the Marianas. N av . med. Bull., W a sh.,
44 ( 2 ) : 42 7 - 4 3 1 .
W a i t e , E d g a r R.
1923. The f ishes o f Sou th Australia . R. E. E. Roger s, A de la i de , 2 40 pp.
W alb a um , Jo h a n n J u l iu s
1792. Petri Art edi Sueci Genera pis ci um in quibus s yst ema . . . . I cht hy-
o log iae , Part 3. G re ifswa ld , 723 pp.
W a l d ic h u k , M ic h a e l
1958. Shellf ish t ox ic ity and the weat her in t he Strait o f Georgi a during
1957. Fish. Res. Bd. Canada , Progr. Rep., Pac if. Coa st Sta., ( 1 1 2 ) :
10 -14 .
W a l f o r d , L i o n e l A.
1932 . The C a liforn ia barracuda ( Sphyraena argentea). I. L ife his tory o f
the Ca liforn ia barracuda . II. A b ib liography o f barracuda s (Sphy -
raeni dae ). Fis h Bull., Sacremen to , ( 3 7 ) : 6 - 132 .
1933. The lif e hist ory o f the Calif orn ia barracuda and its beari ng on the
con serva tion o f the fisher ie s. Ca lif. F ish Gam e , 19 ( 1 ) : 37 - 40 .
W a l k e r , F. D .
1922 . F ish po ison ing in the V irg in Island s. N av . med . Bull ., Wa sh., 1 7 :
193 - 202 .
W a r d, T h o m a s
1907. Rambl es of an Aust rali an naturalist. [ Writt en by Paul Fount ai n
f rom not es s uppli ed by Thom as Ward.] J. Murray, London, 343 pp.
W a t a na b e, M a sa o
1946. Cas es of pois oni ng by t he reef fish, Lutjanus vai gi ensi s. Trans lated
from het Japane se anguage l by W. G . van Campen , romf Shor tRep N.
o .6 Re
, s.Inst .nat. Re ource
s s,Tokyo .Spec .sci. Rep .F ish .,U .S F.
h isW d il Serv
l. .,2 5( )2:0 9 2 1-7 pub
, lis W a h ng
hed s i ont D, C.,
. 1950 .
W eb er , M ax
1923. Di e Fis che der Si boga- Expediti on. In Uit komst en op zool ogis ch,
bo tan isch , oceanograph isch en geo log isch geb ied ver sameld in Neder -
lands ch Oost-I ndi e 1899- 1900 aan boord H. M. Si boga . . . . E. J.
Brill, Le iden , V o l. 57 , 7 10 pp., 12 pis., 123 figs.
W eb er , M ax a nd L. F. de B e a uf ort
1921. Con tri bu ti ons to t he know ledge o f Indo- Austra lian fishes. Zoo l.
M eded., 6 , Part 1: 70 - 71 .
1922 . The fishes o f the Indo -Au stra lian Arch ipe lago . A . J. Brill , Le iden ,
4 : 4 1 0 pp.
178
W e il e r , W.
1938 . N eu e U n t er s uchungen an M itt el oli gozanen Fis chen Ungaras. G eol.
hung . Ser. pa leon t., 15: 1-31.
W h e e l e r , J. F. G .
1954. The probl em o f po is onou s f ishes. Rev. Agri c. Mauri ce, 33: 132 - 139 .
W h i t e , E r r o l I.
1926 . Eocene f ishes f rom Ni geri a. Wit h an appendi x on t he ot olit hs by G.
A ll en Frost. Bull. geol. Surv. Ni geri a, 10: 7- 89.
W h i t e l e a t h e r , R ic h a r d T . and H er b er t H . Brown
1945. A n experi m ent al f ishery s urvey in Tri ni dad, T obago and British
G u i ana, wit h recom m ended i m provem ents in m et hods and gear.
A ng l o - A m eri can Carib. Com m ., W as hi ngt on, iv + 130 pp.
W h itle y , G ilb e r t P.
1927 . A check -list o f fi shes recorded from F ijian waters. J. Pan - Pac if. Re s.
In stn., 2 ( 1 ) : 3-8.
1947 . N ew sharks and f ishes f rom Western Austra lia . Part 3. Aust. Zoo l.,
11: 129 - 150 .
W h i t n e y , R ic h a r d R . a n d K e n n e t h D. C a r l a n d e r
1956. I nterpretati on o f body -s ca le regressi on f or com pu ti ng body lengt h
o f fish. J. W il d lif e M gm t., 20 ( 1 ) : 21 - 27 .
, F r a n c is
W il l ia m s
1956. Pre lim inary survey o f the pe lag ic fishe s o f Ea st A fr i ca . Co l on . Off.
F is h . Pub l., ( 8 ) : 68 pp.
1959. The barracudas ( genu s Sphyraena) in British East A f r i can waters.
Ann . Mag. nat. H ist., 13, Ser. 2, ( 1 4 ) : 9 2 - 128 .
W o o d -Jo n e s
, F.
1902. Corals and a t olls. Low e ll Reeve and C o ., London , xx iii + 329 pp.
W o o d w a r d , A r t h u r S.
1901. Ca ta l ogue o f the f ossil f ishes in t he British M us eum . London . Long-
mans and Co ., London , V o l. 3.
W r ig h t , B r u c e W .
1948 . Relea ser s o f attack behavior patt ern in shark and barracuda. J. W il d-
lif e M gm t., 12 ( 2 ) : 117 - 123 .
Y a suda , F .
1960. The f eed i ng mechan is m in s om e carni vorou s fishes. Rec . oceanogr .
Wks. Japan, 5 ( 2 ) : 153 - 160 .
Y a s u k a w a , T a ka shi
1934 . Report of an i nvesti gati on of pois onous fis hes wit hi n t he j uris di cti on
f t i f t t- l. sl t f
o he Sa pan branch o he G overnm en G enera Tran a ed rom
t s l . . , f t i s
he Japane e anguage by W G van Campen rom Con ag ou
is s s s. I st., . I l., i . i ., .
D ea e Re n D ep m m uno T okyo mp U n v Spec sci.
., . is ., ( ) : - , lis s i t ,
Rep U S. F h Wildl. Serv 25 189 197 pub hed W a h ng on
. ., .
D C 1950
, .
Y . W atr r e n H
u d k in
is i . . i . ll., ( ) : - .
1944 Te raodon po on ng Bull B ngham oceanogr C o 9 1 1 16
. f i -i i it t i s f t
1945 The occurrence l o a. card
. ll.o nh b . or in he
si l., ovar
( )e: o - he . puffer,
Spheroides macu atus J ce comp Phy o 25 2 85 95
, .
Z it t el K arl A vo n
. ii l t l i . t il t t . is .
1890 Grundz
i ge der
. Pal aon
ss o. og e i II. Ab e ung
l Ver
, ebra .,a Rev ed by F
Broil and M Sch o er M un ch and Ber in 971 pp 2048 figs.
179