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NOAA Technical Report NMFS Circular 432

Synopsis of Biological Data

on Bonitos of the Genus Sarda

May 1980

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FAO Fisheries
Synopsis No. 1 18

NMFS/S 118

SAST - Sarda
1.75(01) 001

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

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NOAA Technical Report NMFS Circular 432

Synopsis of Biological Data

on Bonitos of the Genus Sards

Howard O. Yoshida

May 1980

FAO Fisheries Synopsis No. 118

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O

U.S. DEPARTMENT OF COMMERCE

Philip M. Klutznik, Secretary

National Oceanic and Atmospheric Administration

Richard A. Frank, Administrator

National Marine Fisheries Service

Terry L. Leitzell, Assistant Administrator for Fisheries

The National Marine Fisheries Service (NMFS) does not approve, rec-
ommend or endorse any proprietary product or proprietary material
mentioned in this publication. No reference shall be made to NMFS, or
to this publication furnished by NMFS, in any advertising or sales pro-
motion which would indicate or imply that NMFS approves, recommends
or endorses any proprietary product or proprietary material mentioned
herein, or which has as its purpose an intent to cause directly or indirectly
the advertised product to be used or purchased because of this NMFS
publication.

CONTENTS

Introduction 1

1 Identity 1

1.1 Nomenclature 1

1.11 Valid name 1

1.12 Synonymy 2

1.2 Taxonomy 3

1.21 Affinities 3

1 .22 Taxonomic status 4

1.23 Subspecies 4

1.24 Standard common names, vernacular names 5

1.3 Morphology 5

1.31 External and internal morphology 5

*1.32 Cytomorphology

1.33 Protein specificity 7

2 Distribution 7

2.1 Total area 7

2.2 Differential distribution 10

2.21 Spawn, larvae, and juveniles 10

2.22 Adults 16

2.3 Determinants of distribution changes 16

*2.4 Hybridization

3 Bionomics and life history 17

3.1 Reproduction 17

3.11 Sexuality 17

3.12 Maturity 17

3.13 Mating 18

3.14 Fertilization 18

3.15 Gonads 18

3.16 Spawning 18

3.17 Spawn 20

3.2 Preadult phase 21

3.21 Embryonic phase 21

3.22 Larvae and adolescent phase 21

3.3 Adult phase 24

3.31 Longevity 24

3.32 Hardiness 24

3.33 Competitors 24

*3.34 Predators

3.35 Parasites, diseases, injuries, and abnormalities 24

3.4 Nutrition and growth 27

3.41 Feeding 27

3.42 Food 27

3.43 Growth rate 29

*3.44 Metabolism

3.5 Behavior 32

3.51 Migrations and local movements 32

3.52 Schooling 32

3.53 Responses to stimuli 33

4 Population 33

4.1 Structure 33

4.11 Sex ratio 33

4.12 Age composition 33

4.13 Size composition 33

4.14 Subpopulations 37

4.2 Abundance and density 38

*4.3 Natality and recruitment

*4.31 Reproduction rates

iii

*4.32 Factors affecting reproduction
*4.33 Recruitment
4.2 Mortality and morbidity

4.5 Dynamics of population 41

4.6 The population in the community and the ecosystem 41

5 Exploitation 42

5.1 Fishing equipment 42

5.2 Fishing areas 42

5.3 Fishing seasons 43

5.4 Fishing operations and results 44

6 Protection and management 47

6.1 Regulatory measures 47

Literature cited 47

*No information available.

Synopsis of Biological Data on Bonitos of

the Genus Sarda

HOWARD 0. YOSHIDA1
ABSTRACT

Published and some unpublished information on the biology and resources of the three species of
Sarda, S. australis, S. chiliensis, and S. sarda, are compiled, reviewed, and analyzed in the FAO
species synopsis style.

INTRODUCTION

In response to a growing demand for tuna, increased
effort has been expended over the years to harvest the
commercially important large tunas throughout the
world's oceans. The effort has increased to such an extent
that some of the tunas are in danger of being overex-
ploited. Examples of these are the yellowfin tuna, Thun-
nus albacares, in the eastern tropical Pacific, which has
been under management now for a number of years, and
the albacore, T. alalunga, of the South Pacific Ocean
(Skillman 1975). To meet the demand for tuna and at the
same time avoid the overexploitation of important
resources, attention has been focused on tuna resources
that are relatively underutilized, such as the skipjack
tuna, Katsuwonus pelamis. In addition, some of the
smaller tunas or tunalike species such as the bonitos,
Sarda spp., which up to now have had mixed acceptance
in the marketplace, are gaining more attention.

The bonitos are not an entirely unutilized resource.
The total world catch of bonitos has ranged from about
92,200 to 140,500 t (metric tons) in the 10-yr period from
1964 to 1973 (Food and Agriculture Organization of the
United Nations 1970, 1974). As indicated above,
however, bonitos have varying acceptance throughout
the world. In the Mediterranean and Black Sea
countries, Sarda sarda is of great importance and has
been the object of a fishery for many years (Demir 1963).
In Chile, in contrast to earlier years when S. chiliensis
contributed insignificant amounts to the total fish
processed, more and more bonitos have been processed
into canned products in recent years (Barrett 1971). In
California waters, S. c. lineolata has been exploited since
the beginning of the century and there are indications
that the California Indians utilized this species before
the arrival of Europeans (Klawe2). It does not have much
commercial value, however, and the California tuna in-

Southwest Fisheries Center, National Marine Fisheries Service,
NOAA, Honolulu, HI 96812.

2W. L. Klawe, Inter-American Tropical Tuna Commission, La Jolla,
CA 92037, pers. commun. February 1978.

dustry has accepted bonito more from necessity than by
choice (Frey 1971). The flesh of S. orientalis is considered
rather soft and inferior in quality, and except in Kyushu,
where it is caught as an adjunct to the mackerel and
other pelagic species, bonitos are not especially sought
after in Japan (Kikawa and Staff of the Nankai Fish-
eries Research Laboratory, Kochi, Japan 1963). And in
Australia, the edible qualities of S. australis are not
highly regarded (Grant 1972).

The purpose of this report is to review and analyze all
the information available on the biology and resources of
the bonitos throughout the world, following the FAO
(Food and Agriculture Organization of the United
Nations) species synopsis style (Rosa 1965). This report
is based on published papers, in general, and the FAO
species synopses on the bonitos (Demir 1963; Kikawa
and Staff of the Nankai Fisheries Research Laboratory,
Kochi, Japan 1963; Idyll and de Sylva 1963; Silas 1963)
were heavily relied on. Ancieta (1964) also prepared a
species synopsis on S. chiliensis in Peruvian waters in the
FAO species synopsis style.

1 IDENTITY
1.1 Nomenclature
1.11 Valid name

The confusion on the number of valid species in the
genus Sarda has been cleared and, basically, four allo-
patric species are now recognized (Collette and Chao
1975). These are the southeast Australian Sarda australis
(Macleay 1880), the eastern temperate Pacific Sarda
chiliensis (Cuvier 1831), the tropical Indo-Pacific Sarda
orientalis (Temminck and Schlegel 1844), and the Atlan-
tic Sarda sarda (Bloch 1793) (Fig. 1).

The Australian bonito was originally described as
Pelamys australis Macleay 1880. Type-locality: Port
Jackson, Sydney, Australia. Holotype: Macleay
Museum, University of Sydney F-333, now at Australian
Museum, Sydney.

The eastern temperate Pacific Sarda chiliensis was

(a)

Figure 1.— The four species of Sarda: (a) Sarda
australis (from Serventy 1941a, plate 4); (b)
Sarda ehiliensis (from Frey 1971); (c) Sarda
orientalis (from Kikawa and Staff of the Nan-
kai Regional Fisheries Research Laboratory
1963, fig. 1); (d) Sarda sarda (from Demir 1963,
fig. la).

first described as Pelamys ehiliensis Cuvier 1831. Type-
locality: Valparaiso, Chile. Holotype: Museum National
d'Histoire Naturelle, Paris A.5608.

The tropical Indo-Pacific bonito was first described as
Pelamys orientalis Temminck and Schlegel 1844. Type-
locality: Japan. Three syntypes are in the Rijksmuseum
van Najuurlijke Historie, Leiden, of which Boeseman
(1947, 1964) selected the largest as lectotype, RMNH
2286.

The Atlantic bonito Sardo sarda was first described as
Scomber sarda Bloch 1793. No types known to be extant.

1.12 Synonymy

The synonymies given below are modified versions of
those given by Collette and Chao (1975) and do not dis-
tinguish objective and subjective synonyms.

Sarda australis (Macleay)

Pelamys australis Macleay 1880 (Port Jackson, Sydney,

Australia)
Pelamys schlegeli McCoy 1888 (Prince Phillip Bay, Vic-
toria, Australia)
Pelamys chilensis (not of Cuvier 1831) Ogilby 1893 (New

South Wales, Australia)
Sarda chilensis (not of Cuvier 1831). Waite 1904 (New

South Wales, Australia)
Sarda ehiliensis (not of Cuvier 1831). McCulloch 1922

(New South Wales, Australia)
Sarda orientalis (not of Temminck and Schlegel 1844).

Lord 1927 (Tasmania, Australia)
Sarda australis. Walford 1936
Sarda ehiliensis australis. Roughley 1951 (Queensland,

New South Wales, and Victoria, Australia)

Sarda chilensis australis. Silas 1964 (eastern Austra-
lia)

Sarda chiliensis (Cuvier)

Following Collette and Chao (1975) the synonymy for
S. chiliensis is split for the southeast Pacific S. chiliensis
chiliensis and the northeast Pacific S. chiliensis lineolata
(see section 1.23 Subspecies).

Sarda chiliensis chiliensis (Cuvier)

Pelamys chiliensis Cuvier in Cuvier and Valenciennes

1831 (Valparaiso, Chile)
Pelamys chilensis. Giinther 1860
Sarda chilensis. Starks 1906 (Callao, Peru)
Sarda chiliensis. Walford 1936
Sarda sarda chiliensis. Buen 1958 (Chile)
Sarda chilensis chilensis. Vildoso 1963 (Peru)
Sarda sarda chilensis. Sanchez and Lam 1970 (Peru)
Sarda chiliensis chiliensis. Kuo 1970 (Peru)

Sarda chiliensis lineolata (Gerard)

Pelamys lineolata Girard 1859 (San Diego, Calif.)

Pelamys chilensis. Giinther 1860

Sarda chilensis. Jordan and Gilbert 1882

Sarda lineolata. Walford 1936

Sarda stockii David 1943 (Santa Monica Mountains,

Calif.)
Sarda chiliensis. Chabanaud 1944
Sarda chiliensis lineolata. Kuo 1970 (Calif.)

Sarda orientalis (Temminck and Schlegel)

Indo-West Pacific

Pelamys orientalis Temminck and Schlegel 1844 (Ja-
pan)

Pelamys chilensis (not of Cuvier 1831), Day 1878

Sarda chilensis var. orientalis. Steindachner and Doder-
lein 1884 (Japan)

Sarda orientalis. Jordan and Snyder 1900 (Tokyo)

Sarda chilensis (not of Cuvier 1831), Jordan and Snyder
1904 (Honolulu)

Sarda chiliensis (not of Cuvier 1831), Fowler 1938
(Honolulu) •

Sarda orientalis serventyi Whitley 1945 (Western
Australia)

Eastern Pacific

Sarda chilensis (not of Cuvier 1831), Gilbert and Starks

1904 (Panama City)
Sarda chiliensis (not of Cuvier 1831), Herre 1936

(Galapagos)
Sarda velox Meek and Hildebrand 1923 (Panama

City)
Sarda orientalis. Fraser-Brunner 1950

Sarda sarda (Bloch)

Scomber sarda Bloch 1793 (Europe)

Scomber mediterraneus Bloch and Schneider 1801

Scomber palamitus Rafinesque 1810 (Palermo, Sicily)

Scomber ponticus Pallas 1811 (Crimea)

Thynnus pelamis. Risso 1826 (Nice)

Thynnus sardus. Risso 1826 (Nice)

Thynnus brachypterus Cuvier 1829

Sarda sarda. Cuvier 1829

Pelamys sarda. Cuvier in Cuvier and Valenciennes 1831

(Cape Verde Islands and Brazil)
Palamita sarda. Bonaparte 1831
Pelamis sarda. Valenciennes 1844 (Canary Islands)
Sarda pelamys. Gill 1862
Sarda mediterranea. Jordan and Gilbert 1882 (both sides

of Atlantic)
Sarda pelamis. Smitt 1892 (Scandinavia)

1.2 Taxonomy

1.21 Affinities

Suprageneric

Phylum Chordata
Subphylum Vertebrata
Superclass Gnathostomata
Class Osteichthyes
Subclass Actinopterygii
Order Perciformes
Suborder Scombroidei
Family Scombridae
Subfamily Scombrinae
Tribe Sardini

Generic

Genus Sarda Cuvier 1829.

The generic concept of Collette and Chao (1975) is

followed.
Sarda Cuvier 1829:199 (type-species Scomber sarda

Bloch 1793 by monotypy).
Pelamys Cuvier in Cuvier and Valenciennes 1831:149

(type-species Scomber sarda Bloch 1793 by original

designation).
Palamita Bonaparte 1831:173 (substitute name for

Pelamys Cuvier 1831 preoccupied by Pelamys Oken

1816 in Reptilia, Hydrophiidae; therefore, takes the

same type-species Scomber sarda Bloch 1793).
Creotroctes Gistel 1848, p. X (type-species Scomber

sarda Bloch 1793; substitute name for Sarda Cuvier

1829).

The species of Sarda all have several stripes dorsally,
ranging from horizontal to oblique in orientation. The in-
testine runs straight from the stomach to the anus and
there are two intermuscular bones on each side of the
back of the skull. The bony caudal peduncle keels are
well developed as in higher tunas, but are divided into

anterior and posterior sections on each vertebra; the
spleen is large and prominent in ventral view; the right
and left lobes of the liver are both much longer than the
middle lobe. The body is fusiform instead of being more
laterally compressed. Collette and Chao (1975) com-
pared the similarities and differences of the genus Sarda
with the other genera of bonitos within the tribe Sar-
dini.

Collette and Chao (1975) recognized four allopatric
species of Sarda: S. australis, S. chiliensis, S. orientalis,
and S. sarda.

Specific

four

A summary of characters distinguishing the
species of Sarda is given in Table 1.

A key to the species of Sarda as adapted from that
given for the Sardini by Collette and Chao (1975) is pre-
sented below.

2a Total gill rakers on first arch 8-13; supramaxilla

narrow . . S. orientalis (Temminck and Schlegel)

2b Total gill rakers on first arch 19-27;

supramaxilla wider 3

3a Total gill rakers on first arch 19-21; pectoral
rays 25-27, modally 26; teeth sometimes pres-
ent on vomer; length of first dorsal base 315-
343 thousandths of fork length; maxilla 503-

539 thousandths of head length

S. australis (Macleay)

3b Total gill rakers on first arch 23-27; pectoral
rays 22-26, modally 24 or 25; teeth never
present on vomer; length of first dorsal base
267-314 thousandths of fork length; maxilla

460-503 thousandths of head length

S. chiliensis (Cuvier)

1.22 Taxonomic status

Key to the species of Sarda

la Spines in first dorsal fin 20-23; total vertebrae

50-55 S. sarda (Bloch)

lb Spines in first dorsal fin 17-19; total vertebrae

43-46 2

As already noted four allopatric species of Sarda are
recognized (Collette and Chao 1975).

1.23 Subspecies

Two subspecies of the eastern temperate Pacific Sarda

Table 1. — Summary of characters showing differences and similarities among the four species of Sarda . (From Collette

and Chao 1975, table 17.)

Character (reference)

S. sarda

S. australis

S. chiliensis

S. orientalis

Lamellae in nasal rosettes

22-33 (x 26.5)

34-39 (i 37.2)

21-30 (i 25.4)

25-36 (i 31.9)

Vomerine teeth present

sometimes

never

Upper jaw teeth

16-26

18-30 (x 23.5)

12-20 (i 15.5)

Lower jaw teeth

12-24 (x 16.0)

11-20 (£ 14.5)

4-25 (x 19.2)

10-17 (x 13.0)

Palatine teeth

8-21 (i 12.3)

7-14 (i 10.7)

9-22 (i 15.2)

8-19 (i 11.9)

Supramaxilla width

intermediate

wide

narrow

Ectopterygoid-dorsal portion

pointed

slightly expanded

Hyomandibular spine-condyle

projects beyond
condyle

short

projects beyond
condyle

Angle of hyomandibular spine

about 90°

greater than 90°

less than 90°

Elliptical ceratohyal window

present

only slight
depression

Ventral surface of glossohyal

depression present

no depression

Gill rakers

16-23

19-21

23-27

8-13

Vertebrae

50-55

43-46

Pleural ribs

19-23

Intermuscular bones

31-45

32-36

Keels on vertebrae number

5-10

5-8

First closed haemal arch

13th-15th vertebra

12th-14th vertebra

Length of haemal prezygapophyses

postzygapophyses

prezygapophyses

and postzygapophyses at precau-

longer than

dal -caudal junction

prezygapophyses

postzygapophyses

Dorsal spines

20-23

17-19

Dorsal finlets

modally 8

modally 7

modally 8

Anal rays

14-17 (modally 15)

12-15 (modally 14)

14-16 (modally 15)

Anal finlets

modally 7

modally 6

modally 7

modally 6

Total anal elements

19-23

18-22

20-22

(modally 21-22)

(modally 20)

(modally 21)

Supracleithral notch

wide angle

almost 90°

wide angle

Pectoral rays

23-26

25-27

22-26

(modally 26)

(modally 24-25)

Vertical wing of pelvic girdle

shorter and

narrower and

wider

longer

chiliensis are recognized: Sarda chiliensis chiliensis
(Cuvier) for the southeastern Pacific population and
Sarda chiliensis lineolata (Girard) for the northeastern
Pacific population (Collette and Chao 1975).

1.24 Standard common names, vernacular names

The common and vernacular names of the species of
Sarda are given in Table 2.

1.3 Morphology

Collette and Chao (1975) made a detailed study of the
morphology of the species of Sarda including color
pattern, scales, morphometry, meristics, soft anatomy,
and osteology.

1.31 External and internal morphology

Sarda australis

Probably in part because of its restricted geographic
range there have been no studies on the use of morpho-
logical characters to differentiate subpopulations or on
the geographic variation in the morphology of S.
australis. Collette and Chao (1975) presented data on
morphometric characters of S. australis (Table 3) which
give some indication of individual variation in the char-
acters examined.

Sarda chiliensis

Although the temperate northeast (S. c. lineolata) and
southeast (S. c. chiliensis) Pacific populations of Sarda
chiliensis are completely separated geographically, there
are few differences between the two populations and
anatomically they are virtually identical (Collette and
Chao 1975).

Godsil (1954) presented meristic and morphometric
data from the northeast Pacific population of S.
chiliensis. In a later study, Godsil (1955) also presented
similar data for bonitos from the southeast Pacific popu-
lation and made a comparison of the specimens from the
two populations. His conclusion was that such differ-
ences as existed between specimens of the northeastern
and southeastern populations should be considered as
varietal or population differences and that both varieties
should be assigned to the same species, S. chiliensis.

Collette and Chao (1975) discussed the results of
earlier investigators (Walford 1936; Hildebrand 1946;
Kuo 1970) who tried to distinguish the two populations
by the use of various morphometric characters. They
noted that among the meristic characters, the total
number of vertebrae was the best character to distin-
guish the southeast and northeast populations of S.
chiliensis (Table 4). The mean vertebral count of the
northeast population was slightly higher than that of the
southeast population. After considering all the published
data and their own data, Collette and Chao concluded
that "The available data does not convince us that the

northeast and southeast Pacific populations are sub-
species. However, as the populations are genetically iso-
lated from each other and there are some significant dif-
ferences, there is practical value in using the available
subspecific names, and there is ample historical prece-
dent for the name lineolata for the northwest popu-
lation."

Sarda orientalis

The distribution of Sarda orientalis is widespread in
the Indo-Pacific and it is likely that independent popu-
lations occur in the many different areas where the
species is found. Therefore, as Collette and Chao (1975)
indicated, subspecific or populational differences are
possible.

Silas (1964) gave meristic and morphometric data
from S. orientalis collected at Vizhingam on the Kerala
coast of India. He summarized his data by 5 cm size
groups but did not make any conclusions as to any dif-
ferences in body proportions or meristics in the different
size groups. Silas also compared published meristic data
on S. orientalis from various areas in the Indo-Pacific but
refrained from drawing any conclusions on subspecific or
populational differences. Based on the scattered ma-
terial available, Collette and Chao (1975) concluded that
there appear to be no significant anatomical or meristic
differences between any populations of the species. Mor-
phometric data on S. orientalis from Japan and the east-
ern tropical Pacific are given in Table 5. Godsil (1955)
also made detailed morphological studies on S. orientalis
from the eastern tropical Pacific.

Sarda sarda

A detailed analysis of the geographic variation in S.
sarda was done by Collette and Chao (1975). They com-
pared S. sarda occurring in five geographic areas: North
and South America, northeast Atlantic (Scandinavia,
Atlantic Europe, and the Azores), Mediterranean Sea
(including the Black and Adriatic Seas), and the Gulf of
Guinea (extending south to South Africa). They found
varying meristic and morphometric differences between
and among the populations in the five areas. For exam-
ple, the meristic characters were similar for the two west-
ern Atlantic populations as they were for the Mediterra-
nean and Gulf of Guinea populations. In summary, they
concluded that there was at least as much justification
for recognition of subspecies in S. sarda as in S.
chiliensis, if names had been available for the popula-
tions. They stated that more study is needed on this
problem. Morphometric data on S. sarda from North
America, the Mediterranean, and the Gulf of Guinea are
given in Table 6.

Demir (1964) presented meristic data on S. sarda from
Turkish waters. He found that fin ray counts in the
ventral and caudal fins did not vary. However, there were
variations in the number of rays in the other fins. He also
found variation in the number of vertebrae, gill rakers,
and teeth. He computed the mean, standard deviation,

Table 2. — Common and vernacular names of Sarda spp.

Standard

Species

Country

common names

Vernacular names

Sarda australis

Australia

Australian bonito

Horse mackerel, little bonito

Sarda chiliensis

United States

Pacific bonito

Chile

Bonito

Peru

Bonito

Sarda orientalis

Australia (Western)

Oriental bonito

India

Oriental bonito

Vari choora (Malayalam)

Mauritius-Seychelles

Brasse-a-dents (Creole name)

Somali

Sinufa

South Africa,

Bonito

Republic of

Japan

Hagatsuo

Kitsunegatsuo, Hohzan,
Sujigatsuo, Sabagatsuo,
Shimagatsuo, Tozan

United States

Bonito

Sri Lanka (Ceylon)

Thora-baleya

Sarda sarda

Albania

Palamiti

Algeria

Bonite

Bonito, Bonite a dos raye
Palamita, Rsela

Bulgaria

Palamud, Turuk

Lakerda

Canary Islands

Bonito

Bonite

Denmark

Rygstribet Pelamide

France

Bonite a dos raye

Pelamide commun, Pelamide,

Greece
Germany

Israel
Italy

Palamida, Toriki
Pelamide

Sarda
Palamita

Libya

Balamit

Madeira Islands

Cerda

Malta

Palamit

Monaco

Palamida

Morocco

Cerda, Bonito

Portugal

Bonito, Serra

Rumania

Pelamida

Spain

Bonito

South Africa

Bonito

Sweden

Pelamide

Syria

Palamet

Tunisia

Palamid

Turkey

Palamut, Torik

United Kingdom

Pelamid

U.S.S.R.

Pelamida

United States

Common Bonito

Yugoslavia

Polanda

Conite, Pelamido, Pelamida,
Palamida, Boniton, Bonicou,
Bonnicou, Boussicou, Boussicon

Pelamyda, Doriki, Touliki,
Ternata, Koini

Bonite, Unechter bonito,
Mittellandischen bonite,
Rygstribed pelamite

Palamita sarda, Pelamida,
Palamide, Palamida, Palamita,
Palamito, Palamitu maiaticus,
Palametto, Pilamitu, Palamia,
Paamie, Pirantuni, Pisantuni,
Tombarello, Parantuni, Strombo.
Strumbo, Scurma, Sangulu,
Sgamiru, Sgonfietto, Cuvarita,
Cavaritu imperiali, Bonnicou,
Tunnachiu

Blamto

Plamtu, Palamita, Plamitu,

Palamia
Piramida, Paramida

Lacherda

Bonitol, Bonitu, Cerda
Katankel, Sarrajoa
Rygstrimmig pelamid

Pelamid, Balamit, Toumbrel,
Rsela

Belted bonito, Stripe-backed

pelamis
Lacherda
Atlantic bonito, Bonito,

Boston mackerel. Bone jack.

Bloater, Skipjack
Polandra, Palovnic, Pastrica,

Sarica, Sargasto, Tombarel,

Trup lacherda

Table 3.— Morphometric characters of Sarda australis. Upper set of
numbers are measurements expressed in thousandths of fork length,
lower set as thousandths of head length. (From Collette and Chao
1975, table 18.)

Character

Range

Fork length (mm)

Snouth — A
Snout — 2D
Snout — ID
Snout — P ,
Snout — P*

P,-P:
Head length
Max. body depth
Max. body width
F; length
P, length
P, insertion - vent
P_. tip - vent
Base ID
Height 2D
Base 2D
Height anal
Base anal
Caudal spread
Snout (fleshy)
Snout (bony)
Maxilla length
Post orbital
Orbit (fleshy)
Orbit (bony)
Interorbital width

Snout (fleshy)
Snout (bony)
Maxilla length
Post orbital
Orbit (fleshy)
Orbit (bony)
Interorbital width

195-526

Fork length

662-698
581-605
251-276
281-312
258-281
104-125
259-279
221-240
127-169
112-135

76-118
340-393
260-311
315-343

77-95

88-118

72-92

48-88
238-277

88-103

76-88
131-150
120-136

32-41

56-84

58-72

Head length

342-381
291-324
503-539
478-508
134-148
231-287
226-266

349

674

586

263

296

267

116

267

231

141

121

374

290

326

103

259

139

L30

361
305
518
492
137
246
249

20
20
20
20
20
19
20
16
17
19
20
18
18
20
19
20
20
20
11
20
20
20
20
19
20
19

20
20
20
L9
20
19

and standard error of the mean for each meristic charac-
ter with varying counts.

1.33 Protein specificity

See also section 4.14.

Cushing (1964) reported on the use of lectin in the
investigation of antigens of tunas and other species. As
Cushing pointed out, lectins are not antibodies per se but
proteins possessing antibodylike properties. The most
well known source of lectin is the seeds of plants of a
variety of species. Cushing cited unpublished results of
experiments which showed that lectins can be useful re-
agents in detecting antigens in fishes. The lectin ob-
tained from seeds of a plant, Dolichos bifloris, that is
routinely used to distinguish the human A! subtype was
used to show that a significant difference exists between
the frequencies of positive fish occurring in samples of S.
chiliensis of larger and smaller sizes. The cause of this
difference has not been determined.

Barrett and Williams (1965) determined the blood
hemoglobin level of nine specimens of S. chiliensis. The
hemoglobin level ranged from 11.2 to 15.3 g/100 ml and
averaged 12.9 g/100 ml.

2 DISTRIBUTION

2.1 Total area

The distribution of the four species of Sarda as given
below and in Figure 2 was adapted from Collette and
Chao (1975). The geographic classification and codes
given by Rosa (1965) were followed.

Sarda australis

ISEW (Indo-Pacific, central)

610 Australia. 614 Victoria; 615 New South Wales; 616
Queensland; 617 Tasmania; 618 Norfolk Island.

Sarda australis is known only from the east coast of
Australia and Norfolk Island. Off the east Australia
coast, it is common from about the Capricorns (Queens-
land) to Sydney or even Gabo Island (Whitley 1964). The
westernmost record for the species is from Port Fairy,
Victoria (Serventy 1941b).

Sarda chiliensis

INE (Pacific, NE)
ISE (Pacific, SE)

200 North America. 212 British Columbia; 220 Alaska;
231 Washington; 232 California.
300 Latin America. 311 Mexico; 342 Peru; 343 Chile.

This species is found only in the eastern Pacific, where
it is separated geographically into north and south tem-
perate populations. The usual range of the northeast
population is from about Point Conception, Calif., to
Magdalena Bay, Baja California. There is a record of this
species from Socorro Island in the Revilla Gigedos. It is
uncommon north of Point Conception but it has been
recorded from off the Farallon Islands and Eureka,
Calif.; off Puget Sound, Wash.; off the east coast of Van-
couver Island, B.C., Canada; and in coastal Alaska in
Clarence Strait northwest of Ketchikan and off the Cop-
per River.

The range of the southeastern population is along the
coast of South America from Mancora, Peru, in the north
to Valdivia, Chile, in the south.

Sarda orientalis

ISE (Pacific, SE)

ISEW (Indo-Pacific, central)

ISW (Indian Ocean)

100 Africa. 154 South Africa; 156 Malagasy Republic.

Table 4. — Comparison of morphometric characters in two populations of S a rda chilien-
sis, northeast Pacific (S. c. lineolata) and southeast Pacific (S. c. chiliensis). Upper set
of numbers are measurements expressed as thousandths of fork length, lower set as
thousandths of head length. (From Collette and Chao 1975, table 19.)

Northeast Pacific

Southeast Pacific

Character

Range

Fork length (mm)

207-587

375
Fork lengtr

24
i

325-672

498

Snout — A

642-674

656

631-672

654

Snout — 2D

553-595

573

551-585

569

Snout — ID

243-281

269

268-290

279

Snout — P2

275-318

295

280-323

303

Snout — P,

247-279

266

260-300

275

Pi-Pi

106-121

112

105-131

118

Head length

248-275

263

259-292

272

Max. body depth

179-232

210

177-230

210

Max. body width

98-167

131

116-154

134

P, length

99-132

116

125-152

138

P2 length

65- 87

65- 91

P2 insertion • vent

329-367

353

327-368

346

P2 tip - vent

248-331

276

237-352

269

Base ID

278-314

297

267-303

286

Height 2D

64- 98

82-116

Base 2D

71-115

80-109

Height anal

58- 89

77-107

Base anal

61- 84

61- 88

Caudal spread

196-300

234

228-289

258

Snout (fleshy)

87-119

86-102

Snout (bony)

71- 83

75- 89

Maxilla length

115-136

126

121-143

130

Post orbital

125-144

139

131-150

142

Orbit (fleshy)

25- 44

27- 36

Orbit (bony)

47- 63

Interorbital width

57- 69

62- 82

Head length

Snout (fleshy)

339-368

353

331-363

348

Snout (bony)

283-308

297

279-329

299

Maxilla length

460-503

481

463-489

477

Post orbital

499-548

526

504-544

523

Orbit (fleshy)

102-168

119

102-141

115

Orbit (bony)

190-238

218

174-229

205

Interorbital width

213-278

239

233-302

257

300 Latin America. 311 Mexico; 314 Costa Rica; 315
Panama; 341 Ecuador (Galapagos Islands); 342 Peru.

400 Asia. 423 India; 424 Sri Lanka; 437 Philippines;
438 Khmer Republic (Cambodia); 451 Japan.

600 Oceania. 612 Western Australia; 660 U.S.A.
(Hawaii).

sian Gulf, in the Seychelles Islands and Aldabra Island,
and along the coast of Natal, South Africa, south to
Durban. It has also been recorded from Eilat at the
northern end of the Gulf of Aqaba in the Red Sea.

Sarda sarda

This bonito occurs in widely scattered locations in the
Indo-Pacific and Pacific. There is a population of S.
orientalis in the tropical eastern Pacific that is confined
to the coastal areas between the tip of Baja California,
Mexico, and Ecuador and around the Galapagos Islands.
It occurs in Hawaii but is not common there. In Japan,
the species occurs along both coasts of Honshu and is
most abundant along the coasts of Kyushu. The species
has been recorded from the coast of China and also from
the Philippines.

In the Indian Ocean S. orientalis has been recorded
from southwest Australia, along both coasts of India and
from Sri Lanka, from Muscat at the entrance to the Per-

ANW (Atlantic, NW)

ASW (Atlantic, SW)

ANE (Atlantic, NE)

ASE (Atlantic, SE including Bay of Biscay,

Mediterranean Sea, Sea of Marmara, Black Sea)

100 Africa. 110 northwestern area; 111 Tunisia; 112
Algeria; 113 Morocco; 114 Canary Islands; 115 former
Spanish Sahara; 140 western central area; 141 Islamic
Republic of Mauritania; Republic of Senegal, Republic
of Guinea; 142 Cape Verde Islands; 144 Ghana; 151
Angola; 152 South-West Africa; 154 Republic of South
Africa.

Table 5. — Comparison of morphometric characters in populations of Sarda orientalis
from Japan and the eastern tropical Pacific. Upper set of numbers are measurements
expressed as thousandths of fork length, lower set as thousandths of head length.
(From Collette and Chao 1975, table 20.)

Japan

East tropical Pacif
Range x

Character

Range

Fork length (mm)

342-560

432
Fork length

354-613

472

Snout — A

674-703

694

662-703

678

Snout — 2D

596-614

606

569-596

582

Snout — ID

273-308

286

274-311

288

Snout — P2

293-316

303

299-321

310

Snout — P,

272-292

281

277-299

290

Pi-P2

109-118

113

105-118

114

Head length

268-286

278

266-294

284

Max. body depth

221-244

234

193-236

213

Max. body width

143-151

146

127-153

144

P, length

104-125

115

119-134

127

P2 length

70- 78

81- 91

P2 insertion - vent

374-419

392

353-384

367

P2 tip - vent

305-322

311

265-302

280

Base ID

285-327

306

282-302

292

Height 2D

75- 82

89-101

Base 2D

85-111

88-107

Height anal

61- 85

84- 97

Base anal

66- 78

73- 83

Caudal spread

168-234

214

192-259

236

Snout (fleshy)

86-103

98-105

101

Snout (bony)

80- 97

83- 91

Maxilla length

141-149

145

146-156

150

Post orbital

128-147

139

141-151

146

Orbit (fleshy)

32- 60

34- 40

Orbit (bony)

29- 65

60- 68

Interorbital width

67- 73

71
Head length

65- 79

Snout (fleshy)

306-368

344

348-369

357

Snout (bony)

288-344

308

291-317

303

Maxilla length

510-529

522

512-557

528

Post orbital

476-525

503

494-553

512

Orbit (fleshy)

102-152

136

120-147

130

Orbit (bony)

210-234

220

210-238

226

Interorbital width

251-263

256

230-283

251

200 North America. 217 Nova Scotia; 235 Southern
States; 238 Southern Atlantic States.

300 Latin America. 311 Mexico; 321 Trinidad; 331
Colombia; 332 Venezuela; 351 Brazil; 353 Argentina.

400 Asia. 411 Lebanon; 413 Israel.

500 Europe. 510 Scandinavia; 511 Denmark; 514
Norway; 521 Netherlands; 524 France; 531 Ireland; 534
Scotland; 541 Azores, Madeira; 542 Spain, Balearic
Islands; 543 Italy; 553 Greece; 555 Socialist Republic of
Romania; 556 Turkey.

700 Union of Soviet Socialist Republic. 710 Crimea.

This bonito is found on both sides of the tropical and
temperate Atlantic Ocean, in the Gulf of Mexico, and in
the Mediterranean and Black Seas. In the western Atlan-
tic off the east coast of the United States its usual north-
ern limit is Cape Ann, Mass. However, S. sarda has been
recorded from Casco Bay, Maine, and from several
localities along the outer coast of Nova Scotia. It occurs

off Florida but is uncommon off Miami and the Florida
Keys.

Off the Atlantic coast of South America the species is
recorded from Colombia and Venezuela and from about
Rio de Janeiro, Brazil, to Buenos Aires, Argentina.

Biogeographical and natural characteristics of areas

No detailed description of the habitat of the four
species of Sarda will be presented here. From all indica-
tions the species of Sarda are inhabitants of the coastal
or the pelagic neritic province. Sverdrup et al. (1942) set
the vertical border separating the neritic from the
oceanic province at the edge of the continental shelf.
Thus the neritic zone would include all waters of depths
<200 m and accordingly may extend far seaward where
the continental shelf is wide or may extend only a short
distance where the shelf is narrow. "The chemical con-
stituents of the sea water in the neritic province are more

Table 6. — Comparison of morphometric characters in populations of Sarda sarda from
North America, the Mediterranean Sea, and the Gulf of Guinea. Upper set of numbers are
measurements expressed as thousandths of fork length, lower set as thousandths of head
length. (From Collette and Chao 1975, table 21.)

North America

Mediterranean
Range x N

GulfofGuin
Range i

Character

Range

Fork length (mm)

228-500

362

260-504

376

305-443

379

Fork length

Snout — A

646-690

668

641-685

660

648-666

657

Snout — 2D

570-594

582

563-593

579

570-585

578

Snout — ID

262-275

270

257-284

266

262-306

298

Snout — P,

286-304

296

280-302

288

267-284

275

Snout — P,

259-284

269

255-274

263

253-273

267

P1-P2

105-118

111

94-114

104

107-118

111

Head length

256-272

264

251-268

259

253-278

271

Max. body depth

200-224

214

197-216

205

195-228

217

Max. body width

96-171

135

115-148

131

P, length

96-138

115

105-127

116

120-136

130

P2 length

73- 84

76- 86

77- 87

P2 insertion - vent

348-403

366

356-379

370

341-388

357

P2 tip - vent

269-302

282

269-297

285

252-293

266

Base ID

291-330

311

301-323

311

298-323

311

Height 2D

68- 90

85-117

81- 99

Base 2D

85-113

93-112

104

92-112

107

Height anal

68- 98

68- 85

77- 91

Base anal

63- 86

69- 90

66- 89

Caudal spread

204-235

222

226-270

247

223-289

253

Snout (fleshy)

76-100

88- 98

88-101

Snout (bony)

78- 93

74- 83

75- 85

Maxilla length

131-141

136

127-136

131

125-145

138

Post orbital

130-142

136

126-136

132

133-144

138

Orbit (fleshy)

27- 40

27- 34

31- 36

Orbit (bony)

35- 64

53- 62

53- 64

Interorbital width

59- 73

59- 64

58- 65

Head length

Snout (fleshy)

354-374

360

346-367

358

346-366

353

Snout (bony)

289-342

309

289-312

303

293-308

301

Maxilla

503-529

514

497-511

505

494-523

509

Post orbital

494-541

516

503-521

511

489-519

506

Orbit (fleshy)

100-149

121

104-133

118

116-132

122

Orbit (bony)

203-248

222

210-232

220

204-228

220

Interorbital width

216-275

242

231-245

238

212-245

231

variable than in the oceanic. Salinities are usually lower,
sometimes markedly, and undergo seasonal or sporadic
fluctuations such that many of the inhabitants are more
or less euryhaline in nature — that is, able to endure wide
ranges of salinity." (Sverdrup et al. 1942.)

Laevastu and Rosa (1963) determined the distribu-
tion of various species of tunas and the temperature
range in which the species were found. For the species of
bonitos they gave a general temperature range of 12°-
25° C and the temperature range of the fisheries for
bonitos as 15°-22°C.

2.2 Differential distribution

2.21 Spawn, larvae, and juveniles

Sarda australis

There is no information on the distribution of fertil-

ized eggs of S. australis. The capture of the larvae and ju-
veniles also has yet to be recorded in the literature.

Sarda chiliensis

The distribution of the fertilized eggs of S. chiliensis in
the eastern Pacific is not well defined. Barnhart (1927)
collected bonito eggs in a plankton net off La Jolla, Calif.
Orton (1953b) described the early embryonic stage of Pa-
cific bonito but gave no capture locality for her samples.
Sokolovskii (1971) made a more detailed study of the
distribution of fertilized eggs of the Pacific bonito off
Baja California (Fig. 3). As noted by Sokolovskii, S.
chiliensis lineolata in the spring of 1966 spawned mostly
in the shallow waters of the coastal zone. He found fewer
fertilized eggs outside the 200 m isobath. He also de-
termined that the eggs are found only at the surface, for
no eggs were collected in vertical plankton hauls.

Records of capture of juvenile S. chiliensis (Table 7)

•

*®o

u->

t/1

\s>

(1)

<yi

-C

i/>

CO c/) CO (/)

Figure 3. — Distribution of the fertilized eggs of Sarda chiliensis
lineolata off Baja California in the spring of 1966, based on tows with
the ichthyoplankton net. 1 — from 1 to 50; 2 — from 51 to 500; 3 — more
than 500 in one tow of the egg net; 4 — ichthyoplanktonic stations;
5 — isotherms. (From Sokolovskii 1971.)

distribution of the fertilized eggs of S. orientalis. Mito
(1961) gave generalized descriptions of fertilized eggs of
the species in what he termed the Scombrina, including
S. orientalis. However, Mito gave no capture locality for
his samples. Except for this fragmentary bit of informa-
tion on the planktonic fertilized eggs of S. orientalis off
Japan, there is nothing on the distribution of their eggs
in the other areas of the Pacific and Indo-Pacific where
this species occurs.

Records of larval S. orientalis are few. Gorbunova
(1963) recorded the capture of a larva 11.56 mm long
from off the northwest coast of Australia in the Indian
Ocean. Elsewhere in the Indian Ocean, Jones (1960) re-
ported specimens 80, 89, 174, and 262 mm in total length
from Vizhingam on the west coast of India. In the Paci-
fic Ocean, juveniles ranging in length between 75 and 100
mm are reportedly caught in waters adjacent to south-
ern Kyushu (Yabe et al. 1953), and Kishinouye (1923) re-
ported a specimen 170 mm taken off Wakayama Pre-
fecture, Japan. Juveniles from the stomachs of predators
have been recorded from Hawaii (Honolulu Laboratory,
Southwest Fisheries Center, unpubl. data.)

Yabe et al. (1953) reported that 75-100 mm S. orien-
talis are caught in waters around southern Kyushu. They
also reported on S. orientalis landed at the port of
Aburatsu, Japan, ranging in size from 161 to 348 mm.
Preadult S. orientalis occasionally appear in large
numbers in the Indian Ocean. Jones (1960) reported
large catches offish ranging in length from 160 to 200 mm
off the west coast of India between Trivandrum and Cape
Comorin. Gnanamuttu (1966) recorded the occurrence of
S. orientalis ranging from 240 to 260 mm in total length
off the northeast coast of India. In the waters around Sri
Lanka the commercial fishery lands fish ranging up-
wards from 20 cm (Sivasubramaniam 1969).

include two larvae 2.9 and 3.5 mm long; Klawe (1961b)
could not determine whether they were S. chiliensis or S.
orientalis. Other than the prelarval stages which Barn-
hart (1927) and Orton (1953a, 1953b) reported, and the
two larvae recorded by Klawe, the only other larvae of
Sarda reported from the eastern Pacific are those by
Sokolovskii (1971). Sokolovskii reported the capture of
six larvae ranging from 7 to 13.8 mm at an ichthyo-
plankton station in the waters off Baja California. It can
be seen from Table 7 that juvenile S. chiliensis in the
Northern Hemisphere have been found off Baja Cali-
fornia between lat. 22° and 26° N. In the Southern Hem-
isphere they have been found along the coast of Chile and
Peru between lat. 13° and 25°S. Vildoso (1966) sampled
preadult S. c. chiliensis from the Peruvian fishery rang-
ing upward from 20 cm, and Barrett (1971) reported
bonito of about 15 cm taken with anchovy catches at
Arica, Chile. In southern California, preadult S. c. lineo-
lata 15.2-25.4 cm (6-10 in) are first seen by the bait
fishermen in the early summer (Frey 1971).

Sarda orientalis

There is hardly any information in the literature on the

Sarda sarda

The distribution of fertilized eggs of S. sarda has been
studied in greater detail in the Black Sea and the Sea of
Marmara. Mayorova and Tkacheva (1959) reported on
the distribution of S. sarda eggs in the Black Sea in 1956
and 1957 (Fig. 4). They noted that in 1956 bonito eggs
were found in great numbers in the surface layer, the
maximum catch amounting to 12,000 eggs per "10-min
catch," presumably made using a plankton net. At a
depth of 5 m the number of eggs taken did not exceed
5,000 eggs/10-min tow. In 1957 the overall abundance of
fertilized eggs was less than in 1956, and the vertical
distribution was also different in that the 5 m depth pro-
duced more eggs than the surface layer. Demir (1963)
noted that available data indicate that S. sarda eggs
have been found everywhere in the Black Sea, the loca-
tions varying from year to year.

The fertilized eggs of S. sarda have also been found in
the Sea of Marmara (Fig. 5) (Demir 1963). It can be seen
from Figure 5 that in June and July of 1959 most of the
fertilized eggs were found in the eastern portion of the
Sea of Marmara.

Table 7. — Records of capture of juvenile Sarda chiliensis from the eastern Pacific.

1961b, table 2, 1962.)

(From Pinkas 1961, table 2; Klawe

Date

General locality

Latitude Longitude

Method of
capture

Size range

Fork length

(mm)

No. of
speci-

Northern Hemisphere

17-18 May 1947

Off La Jolla, Calif., United

Night light,

—

States

dip net

5 Aug.

1951

100 miles NW of Cape San Lazaro
Baja Calif., Mexico

25°35'N

113°56'W

Night light,
dip net

5 Aug.

1951

Off Baja Calif., Mexico

25°35'N

113°56'W

Night light,
dip net

12 Aug.

1951

Off Ballenas Bay, Baja Calif.,
Mexico

26°29.5'N

133°29.2'W

Plankton net

2.9,3.5

15 July

1953

South of Cape San Lazaro, Baja
Calif., Mexico

23°47'N

112°25'W

Night light,
dip net

25-41

18 July

1953

Southwest of Cape San Lazaro,
Baja Calif., Mexico

23°16'N

112°45'W

Night light,
dip net

19-41

11 Apr.

1955

South of Cape San Luca9, Baja
Calif., Mexico

22°52.8'N

109°53.7'W

Night light,
dip net

16.7

11 July

1956

Southwest of Cape San Lucas,
Baja Calif., Mexico

23°35'N

112°11'W

Night light,
dip net

33.0-33.5

12 July

1956

Southwest of Cape San Lucas,
Baja Calif., Mexico

22°20'N

112°27'W

Night light,
dip net

24-48

17 July

1956

Southwest of Cape San Lucas,
Baja Calif., Mexico

22°47'N

112°14'W

Night light,
dip net

54.5

5 Apr.

1960

Off Baja Calif., Mexico

23°19'N

110°22'W

High speed
net

Southern Hemisphere

20 Feb.

1951

OffPt. Lobos, Peru

125-130

3 Jan.

1956

OffPt. Negra, Peru

Bait net

143-164

10 Dec.

1957

Independencia Bay, Peru

14°14'S

76°12'W

Bait net

128

13 Dec.

1957

Off Ilo, Peru

17°38'S

71°23'W

Bait net

121

15 Dec.

1957

Off Ilo, Peru

17°38'S

71°18'W

Bait net

200.9

18 Dec.

1957

Off Ilo, Peru

Bait net

70-135

21 Dec.

1957

South of Santa, Peru

09°31'S

78°26'W

Bait net

38.0-51.0

31 Dec.

1957

OffPt. Pichalo, Chile

19°35'S

70°16'W

Night light,
dip net

1 Jan.

1958

Off Ilo, Peru

17°47'S

71°30'W

Bait net

86.8

3 Jan.

1958

OffPt. Dos Reyes, Chile

24°30'S

70°49'W

Night light,
dip net

3 Jan.

1958

Off Pt. Dos Reyes, Chile

24°36'S

71o01'W

Night light,
dip net

36-44

14 Jan.

1958

Southwest of Fraile Pt., Peru

13°14.8'S

77°55.5'W

Night light,
dip net

Feb.

1958

Off Chimbote, Peru

Bait net

89-129

6 May

1958

Almejas Bay, Baja Calif., Mexico

Bait net

103

1 Feb.

1959

Sama Cove, Peru

Bait net

111

15 Mar.

1959

Off Ilo, Peru

135-160

16 Mar.

1959

Off Barranca, Peru

173, 199

Identity uncertain.

It is of interest that very little is known of the distri-
bution of fertilized S. sarda eggs in the Mediterranean
Sea. Demir and Demir (1961) noted that planktonic fer-
tilized eggs of S. sarda were unknown from the Medi-
terranean Sea. They pointed out that the eggs reported
on by Sanzo (1932) were eggs that were extracted from a
female gonad and artificially fertilized and reared. More
recently, however, Duclerc et al. (1973) reported on the
collection of fertilized S. sarda eggs near the Balearic
Islands in the Mediterranean Sea. Thus, as far as can be
determined from the literature, the distribution of ferti-

lized eggs of bonito, is not very well defined in the Medi-
terranean Sea.

Except for the report of the collection of the fertilized
eggs of S. sarda off the coast of Massachusetts by Sette
(1943), nothing is known of their distribution in the
Atlantic Ocean.

Mayorova and Tkacheva (1959) stated that S. sarda
prelarvae and larvae occur in the same regions as the
fertilized eggs in the Black Sea. Figure 4 shows that fer-
tilized eggs of S. sarda are widely distributed in the
Black Sea, thus indicating that larval bonito are also

Egg-net
catches (number
per 10 minutes)

EH) ot to AO 100
KM— 200
2(H— 500
5W— 1000

m— zoon
O over 2000
CD not found

- II- 50

- 51-100

- iui-200

- c'ul-JOO

- }OI-5oO

- 5uI-lOuO

Figure 4.— Distribution of the fertilized eggs of Sarda sarda in the Black Sea, in June 1956 (a) and June 1957 (b). (From Mayor-
ova and Tkaeheva 1959, figs. 1, 2.)

widely distributed. The larvae were more abundant at a
depth of 5 m than at the surface. Mayorova and
Tkaeheva also noted that juvenile S. sarda 10-95 mm
long were distributed over a vast area of the Black Sea in
waters from 80 to 100 miles from the coast. In the Sea of
Marmara, however, plankton tows did not reveal any S.

sarda larvae although fertilized eggs in all stages of de-
velopment were plentiful (Demir 1963).

Elsewhere, larvae of 5. sarda have been reported from
the Mediterranean Sea northwest of Oran by Ehren-
baum (1924) and in the Alboran and Mediterranean Seas
by Buen (1930, 1932). More recently, freshly hatched

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prelarvae were collected near the Balearic Islands by
Duclerc et al. (1973). In the Atlantic Ocean, larvae have
been found off Cuba (Gorbunova and Salabarria 1967),
and a juvenile measuring 34 mm has been recorded off
the South Carolina coast (Klawe 1961a). Two juvenile S.
sarda 64 and 67 mm in total length were also captured in
the Gulf of Mexico (Klawe and Shimada 1959).

Sarda sarda from 6 to 18 cm long are captured in vary-
ing numbers in the Dardanelles, the Bosporus, the Sea of
Marmara, and the Black Sea nearly every year in July-
August (Demir 1963). The abundance of preadults in this
size range apparently is a good indicator of the future
abundance of fish 25-38 cm long which are the basis of an
important Turkish fishery that takes place starting in
September in the same areas. Preadults 8-12 cm long
have been captured in large numbers in the Aegean Sea
(Serbetis 1955), and Belloc (1954) discussed the occur-
rence of S. sarda 7.5-20 cm long caught in the Gulf of
Naples and the Gulf of Catania and 15-25 cm long fish in
the Bay of Biscay.

The capture of preadults 20-30 cm long has been re-
ported from the Atlantic coast of France (De la Tourrasse
1957), and 25-35 cm fish have been taken commercially
together with mackerel in the Miramichi estuary in the
Gulf of St. Lawrence (McKenzie 1959). Klawe and
Shimada (1959) reported the capture of two specimens 64
and 67 mm long in the Gulf of Mexico. Young S. sarda
12.7-15.2 cm (5-6 in) long have been taken off Orient,
N.Y. (Nichols and Breder 1927).

2.22 Adults

Sarda australis

Information on the seasonal and annual variations in
the occurrence of S. australis on the east coast of
Australia is fragmentary. Serventy (1941b) stated that
this species apparently is only a summer visitor to Vic-
toria. Munro (1958) noted that S. australis was more
plentiful north of Sydney in the winter; and according to
Grant (1972), it commonly occurs in schools in inshore
waters of Queensland, especially in winter. However,
Marshall (1964) stated that S. australis is very common
on the coasts of Queensland and New South Wales and is
found in great schools throughout the year.

Sarda chiliensis

For the northeast population of S. chiliensis MacCall
et al. (1976) suggested an uneven geographical distribu-
tion of various age groups. They noted that, roughly,
older fish were more available offshore and around Mex-
ico although large fish were also taken in the Santa Bar-
bara, Calif., area in the fall.

As for the southeast population of 5. chiliensis, An-
cieta (1963) found that off the coast of Peru 82.2% of the
total landings during 1951-60 were made in the central
zone, from Chimbote to Pisco; 11.8% were made in the
southern zone, from San Juan to Ilo; and 6% were made
in the northern zone, from Mancora to Salaverry. How-

ever, Mejia et al. (1971)3 noted that beginning in 1962
there has been a gradual overall decline in the annual
Peruvian landings of 5. c. chiliensis, most notably at the
central zone port of Callao; in contrast the landings at
the northern zone port of Paita increased somewhat.

Off the Chilean coast S. c. chiliensis are taken from the
northern boundary of Chile to as far south as Talca-
huano, but most are taken off the northern ports of Arica,
Iquique, and Antofagasta, especially between Iquique
and Mejillones (Barrett 1971). It was also seen that
almost two-thirds of the annual Chilean landings of S. c.
chiliensis are made in the last third of the year.

Sarda orientalis

Along the southwest coast of India, fully mature adult
S. orientalis are found from May to September followed
by the juveniles from October to November. Slight an-
nual fluctuations in seasonal occurrence are also seen
(Silas 1964).

Off the south coast of Sri Lanka (Ceylon), S. ori-
entalis appears in the commercial catches throughout
the year. Mature fish are taken off the south and south-
west coasts primarily between September and February,
and the juveniles appear off the west coast more often
from June to August (Sivasubramaniam 1969). Gen-
erally, S. orientalis appears in the commercial catches
most frequently off the south coast, to a lesser extent off
the west and northwest coasts, and least frequently off
the southwest and northeast coasts.

Sarda sarda

The seasonal occurrence of S. sarda in various locali-
ties is summarized in Tables 8 and 9.

2.3 Determinants of distribution changes

Along the California coast there appears to be a re-
lationship between the latitudinal distribution of S.
chiliensis and sea temperature (Radovich 1963). An up-
surge in the average daily S. chiliensis catch of party-
boat anglers took place with the onset of a warm water pe-
riod off California in 1957. Radovich suggested that S.
chiliensis moved northward into California waters during
warm years. He also noted that during warm years,
bonitos spawned successfully in California waters, thus
further increasing the local population.

From all indications it appears that all the species of
Sarda are closely associated with the coast, where the
ocean environment may be subject to rapid changes.
Demir (1963) stated, however, that S. sarda cannot toler-
ate sudden changes in environment, but they can adapt
to gradual changes in temperature ranging from 12° to

'Mejia, J., M. Samame, and A. de Vildoso. 1971. Revision de la
pesqueria del bonito en quas Peruanas. Fourth Session of FAO Panel of
Experts for the Facilitation of Tuna Research, La Jolla, Calif., 8-13
November 1971. (Mimeogr.)

Table 8. — Fishing seasons for Sarda sarda.

Area

Fishing season

Peak months
of fishing

Source

Black Sea

Bosporus, Sea of Marmara,

Dardanelles
Aegean Sea (around Greece)
Eastern Mediterranean (off Spain)
Mediterranean (off Tunisia)
Eastern tropical Atlantic
Eastern Atlantic (off Morocco)
Eastern Atlantic (Bay of Biscay)
Western Atlantic (Gulf of Maine)

May-October
Throughout the year

Throughout the year
May-June

Throughout the year
October-May
Throughout the year
Mid-April to mid-May
June-October

April-May,
September-December

May-July

Demir(1963)
Demir (1963)

Serbetis (1955)

Gruvel (1931)

Postel (1955b)

Furnestin et al. (1958)

delaTourrasse(1957)

Bigelow and Schroeder (1953)

Table 9. — Fishing seasons in the Spanish fishery for Sarda sarda.
(From Rodriguez-Roda 1966, table 2.)

Area

Fishing season

Peak months of
fishing

Bay of Biscay

(along Spanish coast)
Eastern Atlantic

(off northwest coast

of Spain)
Eastern Atlantic

(southwest coast of

Spain)
Western Mediterranean

(southeast coast of

Spain)
Western Mediterranean

(east coast of Spain )
Western Mediterranean

(northeast coast of

Spain)
Western Mediterranean

(Balearic Islands)
Eastern Atlantic

(Canary Islands)

April-November
April-December

January -December

January-December

January-December
January-December

July-October
July, August

September, October

August-October

May, September
June-November

May-September
August, September

to identify the sex of another bonito except by beha-
vioral characteristics. The bonito is normally hetero-
sexual; however, Vildoso (1960) found cases of hermaph-
roditism in S. c. chiliensis in Peruvian waters. It is of
interest that Magnuson and Prescott observed that one
fish exhibited both a "wobbling" and "following" beha-
vior. They attributed the anomalous behavior of this
particular fish to a possible misidentification and did not
speculate on the possibility of hermaphroditism or other
causes for the aberrant behavior.

As is the case with the other species of bonitos, the
sexes cannot be distinguished in S. sarda. Apparently,
however, some Turkish fishermen can distinguish the
sexes by rubbing the skin anterior of the anus with a
fingertip. The skin is said to be smooth in the females
and rough like emery paper in the males (Demir 1963).

3.12 Maturity

Sarda australis

27 °C and salinities between 14 and 39%o. Sarda sarda
have been taken in the Miramichi estuary, Gulf of St.
Lawrence, where the condition must surely be less than
oceanic. Sarda orientalis have been found in waters rang-
ing from 13.5° to 23°C (Kishinouye 1923).

3 BIONOMICS AND LIFE HISTORY

3.1 Reproduction

3.11 Sexuality

All four species of Sarda are usually heterosexual, and
there are no apparent external anatomical differences be-
tween the males and females. In Sarda c. lineolata, how-
ever, Magnuson and Prescott (1966) noted a sexual di-
morphism in the behavior of this species in captivity.
Magnuson and Prescott noted that some S. c. lineolata in
their observation tank were "wobblers" and that others
were "followers." They postulated and subsequently de-
termined that the "wobblers" were females and the "fol-
lowers" were males. They also determined by their
observations that a male bonito was apparently unable

The literature contains very little information on
Sarda australis. Serventy (1941b) reported on three fish
caught in January and February 1939 off Wilson's
Promontory and Port Albert. The fish weighed between
1.8 and 2.3 kg (4 and 5 lb); and two, which were females,
had "relatively large roes." Whitley (1964) noted that S.
australis sometimes venture into Victoria in the summer
(January to April) and that some of these fish had large
roe. He also found that fish ready to spawn were found in
February and March off New South Wales.

Sarda chiliensis

Barrett (1971) determined the gonad index (GI) (GI =

t^tX 102, where w

weight of both ovaries in grams and

W = body weight in grams) of bonitos and found that
female S. c. chiliensis off the coast of Chile initially
reached sexual maturity at a length of 51 cm. For the
bonitos off the coast of Peru, Vildoso (1966) determined
that the size at first spawning ranged from 47 to 53 cm.
Kuo (1970) stated that in the Northern Hemisphere
population of S. chiliensis, the females attained sexual
maturity at 51 cm and that the fish was 5 yr old at that
length.

Sarda orientalis

Although the size of S. orientalis at first spawning has
not been clearly determined, a 38.6 cm specimen posses-
sed residual eggs which were presumably remnants from
an earlier spawning (Silas 1964). However, Silas indi-
cated that specimens in "ripe running condition or had
already spawned, some showing signs of recovery"
measured between 480 and 605 mm from samples col-
lected in 1960 and 1961 at Vizhingam, India. The age of
these specimens was not determined. Rao (1964) found
four specimens ranging in size from 48 to 55 cm with
running-ripe ovaries in June, August, and September
1959, also from Vizhingam.

Sarda sarda

The bonito in the Sea of Marmara and the Bosporus
usually attain sexual maturity at the end of the second
year, although in some years they may be sexually ma-
ture in a year. Two-year-old fish range from 52 to 57 cm
and 1-yr-old fish from 42 to 48 cm.

In the Black Sea, part of the stock of S. sarda attains
sexual maturity in the second year of life. These 2-yr-old
fish are said to vary from 33 to 50 cm. Off the coast of
Dakar in the eastern Atlantic, the size at first maturity is
392 mm for the males and 370 mm for the females. These
fish are less than a year in age.

3.13 Mating

As Magnuson and Prescott (1966) pointed out, pelagic
schooling fish were assumed to have no discrete court-
ship behavior and no pairing and were believed to shed
eggs and sperm promiscuously while gathered in large ac-
tive schools. To the contrary, Magnuson and Prescott
found that S. c. lineolata do exhibit courting and pairing
behavior, if only temporarily, and observed pairs of fishes
in a sequence of behavior leading to a simultaneous and
adjacent release of eggs and milt. The pair of bonitos re-
leased the gametes during a circle swimming behavior in
which the male swam in tandem with the female in a
circular path.

Although some behavioral patterns have been
observed only in 5. c. lineolata, it would not be unreason-
able to expect that the other species of Sarda also be-
have in a similar manner.

3.14 Fertilization

External in all four species of Sarda.

3.15 Gonads

Sarda australis
No information.

Table 10. — Fecundity of Sarda sarda.

Size of

Fecundity

Area

fish

(No. of eggs)

Reference

Black Sea

40-50 cm

700,000-1,000,000

Zusser (1954)

60-70 cm

1,500,000-2,000,000

70 cm

6,000,000

Black Sea

—

450,000-1,000,000

Slastenenko (1956)

Black Sea

—

700,000-1,000,000

Krotov (1957)

Black Sea

56-65 cm

732,160-3,233,580

Mayorova and Tkacheva

(2-3.8 kg)

(1959)

Eastern

60 cm

900,000

Postel (1955a)

Atlantic

Sarda chiliensis

Vildoso (1963a) estimated the number of ova spawned
by a 600 mm (3 kg) bonito at half a million per spawning
season. According to Vildoso, spawning is fractionary.
Kuo (1970) estimated that the fecundity of the northern
S. chiliensis ranged from 104,900 to 894,200 eggs for fish
from 47.6 to 63.7 cm in fork length. He indicated that the
fecundity increased exponentially with size of fish.

Sarda orientalis

Silas (1964) made fecundity estimates for five mature
specimens of S. orientalis from the Indian Ocean. He
estimated that the females produce 0.08-0.15 million
eggs/spawning and 0.24-0.64 million eggs/spawning
season. Rao (1964) estimated that the bonito spawns
0.21-0.28 million eggs/spawning and 0.91-1.15 million
eggs/spawning season in the Indian Ocean.

Sarda sarda

Demir (1963) summarized the fecundity determina-
tions for S. sarda made by various investigators in the
eastern Atlantic (Table 10). The fecundity estimates
range from 450,000 to over 3,000,000 eggs.

3.16 Spawning

Sarda australis

Whitley (1964) indicated the occurrence of fish that
were ready to spawn in February.

Sarda chiliensis

The spawning season for S. c. chiliensis in Chilean
waters begins in September, is at a maximum in October
and November, and is well over before April (Barrett
1971). The spawning season off the coast of Peru is very
similar to that off Chile; peak spawning of bonito extends
from October to February (Vildoso 1966). According to
both authors, larger, older bonito mature earlier in the
spawning season than do the younger fish (Fig. 6).

For the Northern Hemisphere population of S.
chiliensis, Kuo (1970) investigated several different ap-
proaches to determine the spawning season, including

NOV. a

APR.o o o

SEPT. x'
JULY o--'

46 48 50 52 54 56 58 60 62 64 66 68
LENGTH (cm)

Figure (i. — Mean monthly gonad indices, GI =

10 , by length

class, of female Sarda chiliensis chiliensis sampled from the com-
mercial landings at Iquique, September 1968 to July 1%9. (From
Barrett 1971, fig. 5.)

the annual cycle in the development of intraovarian
oocytes, the annual cycle of frequency distribution of
ovum diameters, and the annual change in gonad index.
All of the different methods indicated that the spawning
season for the Northern Hemisphere S. chiliensis in
southern California is from May through July. Kuo
speculated that off southern Baja California spawning
may start as early as April and may continue into
August. He also indicated that the larger fish tend to
spawn earlier in the season and longer than the smaller
fish. Frey (1971), however, attributed an early spawning
season to S. c. lineolata between southern California and
northern Baja California waters. He stated that the
bonitos spawned between January and May in these
waters.

The results of an ichthyoplankton survey off Baja Cali-
fornia seem to corroborate the conclusions of Kuo (1970).
On the basis of the distribution of fertilized eggs, Soko-
lovskii (1971) showed that S. c. lineolata spawned in a
relatively large area near the coast (Fig. 3). In early
March the most intensive spawning occurred in the
southern area and in early April it occurred in the north-
ern area. The depth of the ocean in the spawning area
ranged from 40 to 150 m, and practically no bonito eggs
were found outside of the 200 m isobath. Larval bonito
were captured at only one station during this survey.

Most of these observations tend to support Klawe
(1961b) who, on the basis of the capture of larval and ju-
venile S. chiliensis, had stated that the spawning of this
species takes place in the warmer season off California,
Baja California, Peru, and northern Chile.

Evidence has also been gathered which suggests that 1-
and 2-yr-old S. c. lineolata spawn in areas influenced by
warmwater discharges during spawning periods when the

water is cold. Tag returns have shown that small
numbers of young fish stay in these areas for as long as 2
yr and provide a small amount of recruitment even in
cold years (Collins and MacCall 1977).

Sarda orientalis

The literature contains very little information on 5.
orientalis spawning in the eastern Pacific and around
Hawaii. Klawe (1961b) reported on two larval forms of
Sarda from off Baja California that could not be identi-
fied to species. The distribution of the northeast popula-
tion of S. chiliensis overlaps or nearly overlaps that of the
population of S. orientalis at the location of capture of
the larvae; therefore, the larvae could be either of the two
species.

Nothing is known about the spawning of S. orientalis
around Hawaii except that a few juveniles have been
found in the stomachs of predators (Honolulu Labora-
tory, Southwest Fisheries Center, unpubl. data), which
provides evidence of reproduction.

No formal studies on the spawning of S. orientalis in
Japanese waters have been made. Kikawa and Staff of
the Nankai Regional Fisheries Research Laboratory
(1963) presumed that spawning occurs in the coastal
waters of the tropical zone in the Lndo-Pacific. There ap-
parently is some local unrecorded knowledge of bonito
spawning in Japan because Harada et al. (1974) re-
ported that the spawning season is May-June. They
caught mature S. orientalis on 15 May and 10 June in
traps set near the coast of Oshima Island and used these
fish successfully in an artificial fertilization experiment.
Mito (1961) gave descriptions of the fertilized eggs of S.
orientalis found in plankton collections but gave no col-
lection details. Kishinouye (1923) and Yabe et al. (1953)
reported on the capture of juvenile bonito in southern
Honshu and Kyushu waters.

Off the coast of Vizhingam in the Indian Ocean, S. ori-
entalis spawn from April to September and possibly in
other months of the year. Ova diameter frequency distri-
butions indicate the possibility that individual females
spawn several batches of ova (Fig. 7) during the spawn-
ing season (Silas 1964).

Gorbunova (1963) reported on the larva of S. ori-
entalis from the Indian Ocean off the northwest coast of
Australia and generalized that spawning is limited to the
autumn-winter period. However, this conclusion is
based, apparently, on the capture of one larva of S. ori-
entalis in December. Gorbunova probably has the
seasons confused, for December off the northwest coast of
Australia would be closer to summer in the Southern
Hemisphere.

Sarda sarda

Demir (1963) stated that S. sarda spawning begins in
mid-May, reaches a peak in June, and lasts at least to
the end of July. He assumed that the spawning season is
the same in the Black, Marmara, and Aegean Seas.

Indications are that the spawning season in the Medi-

Table 12.— Relation between the stage of sexual maturity and the
gonadosomatic index (GI =~tf * 100, where w is gonad weight and
W is fish weight) for Sarda sarda landed at Barbate, Spain in 1964.
(From Rodriguez-Roda 1966, table 26.)

10 20 30 40 50 60

OVA DIAMETER ( IN MICROMETER DIVISIONS )

Figure 7. — Ova diameter-frequency polygons of ripe ovaries of the
oriental bonito, Sarda orientalis, from the southwest coast of India.
The ripe ova were sampled from two of four ripe fish examined from
June to September 1959. (From Rao 1964, fig. 3.)

terranean is also from May to July. However, Demir
(1963) noted that Dieuzeide et al. (1955) found that the
spawning season off the coast of Algeria is from March to
May.

Rodriguez-Roda (1966) made observations on the sex-
ual development of S. sarda landed in May, June, and
July 1963 and 1964 in the Spanish fishery based at the
Mediterranean ports of Barbate and Tarifa. By gross
examination of the gonads, he classified the males and
females according to six stages of sexual maturity: I, Im-
mature; II, Early maturing; III, Mature; IV, Pre-
spawning; V, Spawning; and, VI, Spent. Data on the
monthly percentage distribution of fish in the various
stages of sexual maturity were given by Rodriguez-Roda
(Table 11). He also computed the GI of the fish for his

Males

Females

Stage of sexual

No. of

maturity

Mean

Range

fish

Mean

Range

fish

I (immature)

0.56

0.42-0.69

II (early maturing)

0.94

—

0.85

—

III (mature)

4.15

0.96-6.90

4.44

1.77-12.15

IV (prespawning)

—

5.39

3.56-9.40

V (spawning)

—

VI (spent)

—

2.15

—

w
1964 samples using the equation GI = xy • 100, where w is

the weight of the gonads and W is the weight of the fish,
and determined the relation between the stage of sexual
maturity and the GI (Table 12). Rodriguez-Roda noted
that the smallest fish with prespawning and spawning
gonads were 39.5 cm for the males and 40.5 cm for fe-
males.

In the eastern Atlantic near the coast of Dakar the
spawning season extends from December to June, in-
cluding peaks in January and April, and June to July in
Moroccan waters. Less is known about S. sarda spawning
in the western Atlantic. Bigelow and Schroeder (1953)
stated that the bonito spawns south of the Gulf of Maine
in June. However, Sette (1943) reported the collection of
bonito eggs near Martha's Vineyard, Mass., in July.
Further south off the South Carolina coast Klawe
(1961a), based on the capture of a singe 34 mm bonito,
stated that this species spawns in the winter. The cap-
ture of a larval bonito (Gorbunova and Salabarna 1967)
off the coast of Cuba and a juvenile specimen in the
central Gulf of Mexico (Klawe and Shimada 1959) indi-
cates some spawning activity in these areas.

3.17 Spawn

The fertilized eggs of the bonitos have been described
for all the species in the genus except Sarda australis.
Except for differences in size, the fertilized eggs at the
various developmental stages appear to be quite similar,
and it is doubtful if the species could be separated. As ex-
amples of differences in size of the fertilized eggs among
the species of Sarda, Demir (1963) found that for S. sarda
in the Sea of Marmara, the eggs, which were measured
after preservation in 4% Formalin, varied between 1.18
and 1.55 mm in diameter. The fertilized eggs of S. c.

Table 11. — Monthly percentage distribution of Sarda sarda in the various stages of sexual development,
Barbate and Tarifa, Spain, in 1963 and 1964. (I, immature; II, early maturing; III, mature; IV, prespawning;
V, spawning; VI, spent.) (From Rodriguez-Roda 1966, table 24.)

Males

Females

No. of

III IV

fish

III IV

fish

May

7.02

1.75

87.72 3.51

30.00

15.00

55.00 —

June

—

73.81 23.81

2.38

—

2.38

—

64.29 30.95

—

2.38

July

—

100.00

—

54.17 41.67

—

4.17

Uneolata from off the coast of Baja California measured
from 1.4 to 1.8 mm in diameter (Sokolovskii 1971). How-
ever, no mention is made of whether the eggs were
measured before or after preservation. Harada et al.
(1974) reported that the fertilized eggs of S. orientalis
were from 1.32 to 1.45 mm in diameter. Here again it is
not clear whether the eggs were measured fresh or after
preservation. These fertilized eggs were the result of a
successful experiment in artificial fertilization, the first
record of success in artificial fertilization for this species.

Because the fertilized eggs of the species of Sarda are
similar in appearance, as a typical example Demir's
(1963) description of the eggs (Fig. 8) of S. sarda is given
below.

The fertilized planktonic eggs of S. sarda are spheri-
cal, transparent, and, except for the oil globule, color-
less. The yolk sac is homogeneous and finely granulated,
and varying numbers of oil globules of different sizes are
present on its surface.

Although yellow and black pigmentation are present
on the developing embryo, only the black pigments (me-

lanophores) remain after preservation. The melano-
phores on the yolk are found on the surface of the sac. On
the embryo they initially appear on the dorsal part of the
body and later on the head and tail.

3.2 Preadult phase

3.21 Embryonic phase

See section 3.17.

3.22 Larvae and adolescent phase

The size of a newly hatched larva of S. chiliensis is
given by Barnhart (1927) as 3.75 mm. Newly hatched lar-
vae of S. orientalis measured 4.1-4.3 mm (Harada et al.
1974). As a typical example of the development of the
larvae of bonitos, a description of the larval and post-
larval development of S. sarda is given below. The
description is taken from a translation of Padoa (1956).

Embryonic development is rapid and the larva hatches
only a day after the closure of the blastopore.

Figure 8. — The fertilized eggs of Sarda sarda
at various developmental stages. (Note: The
developmental stages were not defined by the
authors.) (From Demir and Demir 1961.)

The mouth of the larva is not yet open, and the eye
without pigment at hatching (Fig. 9a). The yolk sac and
fin folds are large, and several oil droplets are present in
the posterior part of the yolk sac. The anal opening is lo-
cated a little less than midway between the head and
tail. The pectorals are barely outlined. There is a total of
about 50 body segments, including 15 preanal and 38
postanal segments.

The yolk is reduced to less than half in a 4.6 mm speci-
men 2 days after hatching (Fig. 9b). The mouth is open,
the anus has moved forward, and the number of preanal
segments is decreased from 15 to 11 and the postanal
increased to 39. The pectorals are well developed and
membranous; the hypurals are absent. Melanophores on
the trunk are limited to the caudal ventral margin; other
melanophores are present in the curve of the peritoneal
cavity and a few are found on the profile of the snout.

The yolk is nearly used up in a 4.68 mm specimen 4
days after hatching (not illustrated). The snout is con-
siderably longer and there are well-defined teeth. There
is a posteriorly directed triangular spine at the pre-
opercular margin. The anus is displaced even farther
forward to about the 9th segment. The pectorals are un-
changed and the hypurals are still absent. The melano-
phores persist on the caudal ventral margin; the number
and size of melanophores are increased on the dorsal
peritoneal walls and the head.

The yolk is exhausted in a 4.20 mm specimen 6 days
after hatching (Fig. 9c). The snout is a little more dis-
tinctly pointed and is changing to the subconical form.
The preopercular spine is pointed and evident; the pre-
anal distance is about 34% of the total length. The pec-
torals are membranous and the rounded margin reaches
about the level of the anus; the hypurals are still absent.
There are a few irregularly spaced melanophores on the
ventral margin of the trunk. The melanophores persist on
the peritoneal curve, along the thoracic girdle, and on the
upper profile of the head; they are also present at the
apex of the snout and in the middle of the mandi-
ble.

Melanophores are present on the apex of the snout, on
the profile of the head, on the peritoneal curve, and along
the thoracic girdle in a 7.2 mm specimen (Fig. 10c).
Black pigments are almost completely lacking on the
caudal trunk and limited to a few dispersed elements
along the ventral margin. Two series of preopercular
spines (an anterior of three and a posterior of six), of
which the two central ones are very long, are now de-
veloped; two other spines are present on the posterior
margin of the otocysts. About 10 pointed teeth are on
each side of the maxillary and the mandible. Outlines of
about 13 rays, the future second dorsal and anal fins, can
be counted. The preanal distance now represents 42% of
the total length and the distance from the snout to the

Figure 9. — Prelarvae of Sarda sarda: (a)
4.32-mm prelarva (from Padoa 1956, fig.
309); (b) 4.6-mm prelarva (from Padoa
1956, fig. 310); (c) 4.20-mm prelarva (from
Padoa 1956, fig. 311).

2 n

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3 E

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£ W

to *C

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H 3

fa W

insertion of the anal fin, 65Tr; there are about 20 seg-
ments in the postanal region.

In a 26.5 mm specimen, the preopercular spines are
well-evident but small (Fig. lOf). The caudal fin is now
forked; the pectoral and ventral fins are relatively small.
The membrane that is continuous with the second dorsal
and anal fins still unites the finlets. The first dorsal is
covered with black pigments except for a clear space at
the base of the 10th to the 13th rays; the other fins are
transparent. Fine black pigments are on the snout, the
nape, and the dorsal region of the trunk where six verti-
cal bands, which extend from the dorsal edge to about
the middle of the flanks, begin to appear. A dark hori-
zontal line separates the pigmented dorsal portion from
the ventral, unpigmented portion. In a 32 mm specimen
the preopercular spines are smaller but still evident (Fig.
lOg). The membrane still joins the finlets and the fin
coloration is unchanged. The pigmentation on the head
is increased and the six vertical bands are quite pro-
nounced.

Illustrations of postlarval S. sarda given by Vodianit-
skii and Kazanova (1954) are also shown in Figure
10.

Pinkas (1961) and Klawe (1961b) gave descriptions of
postlarval S. chiliensis, and Jones (1960), Gorbunova
(1963), and Harada et al. (1974) described larval and
postlarval S. orientalis (Figs. 11, 12).

Except for S. chiliensis and S. orientalis in the eastern
Pacific, where there may be some overlap in the distri-
bution of these two species, the problem of identifying
and separating the larval and juvenile species within the
genus does not occur because the species are allopatric.
As for the larval forms of S. chiliensis and S. orientalis in
the eastern Pacific, Klawe (1961b) stated that it would
be impossible to separate the two species without a com-
plete developmental series of both species. Klawe fur-
ther noted that it should be possible to distinguish ju-
venile S. chiliensis and S. orientalis from each other
based on gill raker counts even at a relatively small size.
Pinkas (1961) pointed out other characters (number of
teeth on lower jaw and presence or absence of posterior
gill teeth) that could be used to separate S. chiliensis
>24 mm from similar sized S. velox ( = S. orientalis).

Once the juveniles acquire the full adult complement
of certain characters, which seems to be at a relatively
small size (Pinkas 1961; Klawe 1961b), the summary of
distinguishing characters (Table 1) for the four species of
Sarda compiled by Collette and Chao (1975) indicates
that the juveniles should be separable without difficulty.
Sarda australis, S. chiliensis, and S, orientalis can be
separated from each other on the basis of gill raker
counts while S. sarda can be distinguished from all the
others by the number of vertebrae.

3.3 Adult phase

3.31 Longevity
See section 4.13.

3.32 Hardiness
See section 3.52.

3.33 Competitors

Sarda chiliensis

The relationships of S. c. lineolata with the other
species occupying the same areas off southern California
are not clear. However, recent changes in the species
composition of pelagic fish stocks off California shores
may involve the bonito as well as Pacific sardine, Sar-
dinops sagax, and northern anchovies, Engraulis mordax
(Frey 1971). Presumably these changes would involve the
ability of the various species to compete with one another
for food and space. Behavior studies on S. c. lineolata
indicate that they can compete very well with other
species in obtaining food. In a fish tank containing Pa-
cific barracuda, Sphyraena argentea; yellowtail, Seriola
dorsalis; and tarpon, Megalops atlantica, S. c. lineolata
was the first species to reach and ingest food tossed onto
the surface (Magnuson and Prescott 1966). Magnuson
and Prescott noted that the high speed of S. c. lineolata
gives it an advantage over the other species in a
"scramble" type of competition.

Sarda orientalis

In the Indian Ocean adult S. orientalis are often
caught together with Euthynnus affinis, Auxis spp.,
Thunnus tonggol, and Scomberomorus spp. and pre-
sumably compete with these species for food (Silas 1963).
Demir (1963) stated that all predatory fishes and dol-
phins are potential competitors of S. sarda, including
Delphinus delphis ponticus, Tursiops truncatus,
Scomber scombrus, Trachurus mediterraneus, Poma-
tomus saltatrix in the Black Sea, and additionally,
Scomber japonicus, Auxis rochei, and Euthynnus
alletteratus in the Sea of Marmara.

3.35 Parasites, diseases, injuries, and ab-
normalities

Records of parasitism on bonitos include a trematode,
Didymozoon pelamyzis, from the gill lamella of S. sarda
from the Black Sea, Sea of Marmara, and the Medi-
terranean (Dawes 1946; Demir 1963). Larvae of the
cestode Callitetrarhynchus gracilis have been found in
the body cavity and the isopod Livoneca sp. from the gill
lamellae of S. sarda (Postel 1954). Parasitic copepods
Caligus sarda (Pearse 1952) and Ceratocolax euthynni
(Vervoort 1971) have been found on S. sarda. Silas (1967)
and Silas and Ummerkutty (1967) produced a detailed
listing of the parasites of scombroid fishes, including
Sarda (Table 13).

Two stripeless S. c. lineolata have been recorded from
waters off La Jolla, Calif. (Matsumoto et al. 1969). These
specimens possessed all the meristic characters typical of
S. c. lineolata except that they lacked stripes.

Figure 11.— Postlarval Sarda chiliensis: (a) 16.7 mm fork length (from Pinkas 1961, fig. 4); (b) 33.0 mm fork length
(from Pinkas 1961, fig. 5); (c) 42 mm (from Klawe 1961b, fig. 3); (d) 160 mm (from Klawe 1961b, fig. 4).

E ~

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_- E

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Table 13.— Parasites of Sarda. (Modified from Silas 1967; Silas and Ummerkutty 1%7.)

Monogenetic

Digenetic

Copepods

trematodes

Cestodes

Sarda chiliensis

Caligus bonito

Caligus mutabilis

Pseudocycnus

appendiculatus

Sarda orientalis

Caligus bonito

Capsala ca

ballerio

Bucephalopsis cybii

Caligus coryphaenae

Caligus productus

Sarda sarda

Caligus bonito

Capsala pel

imydis

Aponurus tschugunowi

Grillotia erinaceus

Caligus mutabilis

Hexostoma

pricei

Atalostrophion sardae

Lacistorhynchus

Caligus pelamydis

Hexostoma

thynni

Bucephalopsis arcuata

tenus

Caligus productus

Dinurus barbatus
Hirudinella clavata
Lecithochirum

caudiporum
Lecithochirum texanum
Nematobothrium

pelamydis
Opecoelides vitellosus
Rhipidocotyle

angusticollis
Tormopsolus orientalis?
Unitubulotestes sardae

Scolex pleuronectis
Tentacularia bicolor
Tentacularia coryphaena
Tetrarhynchus

megabothrium
Tetrarhynchus

scomber-pelamys
Tetrarhynchus sp. ''

Species incertae sedis.
"Indicates doubtful record.
Forms unidentifiable.

3.4 Nutrition and growth

3.41 Feeding

Magnuson and Heitz (1971) examined the relation be-
tween the gill raker morphology and the food habits of
scombrid fishes, including S. c. lineolata. In the past, the
high diversity of food organisms in the stomachs of scom-
brids gave rise to the opinion that scombrids are non-
selective feeders. However, Magnuson and Heitz pointed
out that selectivity does exist in terms of food size in
scombrid food habits in that the stomach contents of
small and large fish of the same species were dissimilar.
They suggest that larger predators have a reduced ability
to catch small prey (crustaceans) because of a relatively
large gap between the gill rakers. Among scombrids of
the same size S. c. lineolata and S. orientalis had the
largest gill gaps (1.8-3.3 mm). The relative unimpor-
tance of crustaceans in the food habits of S. c. lineolata
noted by Pinkas et al. (1971) could be related in part to
the large gill gaps in this species. The data presented by
Magnuson and Heitz (1971) also indicate a relationship
between the number of gill rakers and the gill raker gap,
i.e., the greater the number of gill rakers, the smaller the
gill raker gap. Sarda orientalis have fewer gill rakers than
S. chiliensis (Collette and Chao 1975), and as Magnuson
and Heitz's data show, the gill raker gap is greater in S.
orientalis. Therefore, it would be expected that the diet

of S. c. lineolata would contain a larger proportion of
smaller organisms than that of S. orientalis.

3.42 Food

Sarda australis

Only a few miscellaneous observations are available on
the feeding habits of the Australian bonito. Particularly
during the winter months, S. australis occurs commonly
in schools in the inshore coastal waters of Queensland
and feeds on hardyheads, Pranesus ogilbyi; pilchards,
Sardinops neopilchardus; and anchovies, Engraulis
australis (Grant 1972). Munro (1958) indicated that they
also feed on mackerel scad, Trachurus mccullochi.

Sarda chiliensis

A detailed study of the food habits of S. c. lineolata
was conducted by Pinkas et al. (1971) based on a total of
1,498 stomachs collected in 1968 and 1969 from fish cap-
tured in nearshore waters of southern California and
Baja California. Their study clearly showed that the
northern anchovy, Engraulis mordax, was the major food
item in the diet of S. c. lineolata. The common squid,
Loligo opalescens, ranked next in importance, and mis-
cellaneous fishes and a few crustaceans made up the
small remainder of the diet of S. c. lineolata (Table 14).

Table 14.— Food of Sarda chiliensis lineolata, 1968 and 1969. (From Pinkas et al. 1971.)

Frequency

Percent

Volume

Percent

frequency of

Food items

Number

number

(ml)

volume

occurrence

Fishes

Engraulidae

Engraulis mordax

4,159

75.5

11,356.4

75.9

462

56.3

Scomberesocidae

Cololabis saira

0.5

190.8

1.3

1.9

Gadidae

Merluccius productus

0.4

6.8

<0.1

1.8

Carangidae

Trachurus symmetricus

0.3

185.6

1.2

1.6

Sciaenidae

Genyonemus lineatus

<0.1

38.0

0.3

0.1 +

Seriphus politus

+0.1

1.3

<0.1

0.2 +

Embiotocidae

Brachyistius frenatus

<0.1

10.0

<0.1

0.1 +

Cymatogaster aggregata

<0.1

14.0

0.1

0.1 +

Zalembius rosaceus

<0.1

9.0

<0.1

0.1 +

Scorpaenidae

Sebastodes spp.

0.2

5.8

<0.1

0.6

Stromateidae

Peprilus simillimus

0.1 +

195.0

1.3

0.7

Bothidae

Citharichthys sordidus

<0.1

0.6

<0.1

0.1 +

Unidentified fishes

765

13.9

223.8

1.5

185

22.5

Subtotal, fishes other than

Engraulidae

859

15.6

880.7

5.9

247

30.1

Subtotal, all fishes

5,018

91.1

12,237.1

81.8

709

86.4

Cephalopods

Loligo opalescens

448

8.1

2,690.7

18.0

207

25.1

Onychoteuthis boreali-japonicus

<0.1

3.7 s
9.9

0.1 +

Unidentified cephalopods

0.3 +

2.0

Crustaceans

. 0-2

Pleuroncodes planipes

<0.1

5.6

0.5

Crab megalops larvae

<0.1

0.1

0.1 +

Unidentified animals

0.3 +

4.7 I

2.0

Subtotal, all invertebrates

(except Loligo opalescens)

and unidentified animals

0.8

24.0

0.2

4.8

Totals

5,510

100.0

14,951.8

100.0

—

There was a seasonal variation in the importance of the
common squid in the diet of S. c. lineolata. Squid was
most important in the first and second quarters of the
year but was less important in the third quarter of the
year. This seasonal pattern of the importance of squid in
the bonito's diet was related to the seasonal con-
centrating behavior of squid during its reproductive cy-
cle.

Sarda orientalis

Information on the food and feeding habits of S. ori-
entalis in Japanese waters is fragmentary. Yabe et al.
(1953) examined 18 stomachs from juvenile S. orientalis
in southern Kyushu waters. Fifteen of the 18 stomachs
were empty, 2 of the stomachs contained Engraulis
japonicus remains, and 1 stomach contained unidentifi-
able fish remains. Yokota et al. (1961) examined
stomachs of 24 S. orientalis caught in about the same
area and found a total of five saury, one Sphyraena sp.,
one squid, and one carangid in the stomachs.

Kumaran (1964) investigated the food habits of S. ori-
entalis in the Indian Ocean based on 43 specimens rang-
ing in length from 85 to 305 mm. The most important
single food item was Anchouiella commersonii. Fishes of
lesser importance in the diet were Leiognathus insidi-
ator, Decapterus russelli, and Sardinella sp. Kumaran
noted that the variety of food organisms was smaller in
the stomachs of S. orientalis as compared with Euthyn-
nus af finis and Auxis. He gave as the reason for this the
fact that all the S. orientalis specimens were collected
from a single locality. Sivasubramaniam (1969) looked at
the food-bearing stomachs of 11 S. orientalis caught in
the waters around Sri Lanka. Nearly 60% by volume of
the food consisted of a clupeoid fish, 18% cephalopod
mollusk (squid and octopus), 15% decapod Crustacea,
and 1% miscellaneous items including unidentified
fishes.

Sarda sarda

More information is available on the food and feeding
habits of S. sarda, including information on larval and

juvenile stages. Larval S. sarda about 5 mm long start
active feeding even before the yolk sac is completely ab-
sorbed. Larvae >6-7 mm feed on relatively large organ-
isms, including fish larvae, but prefer copepods. Ju-
veniles >18-20 mm feed on the larvae of anchovy, horse
mackerel, and S. sarda (Mayorova and Tkacheva 1959).
Zusser (1954) reportea a Pseudocalanus in the stomach
of a 7.2 cm juvenile and a 4.5 cm long anchovy in a 8.7 cm
juvenile. Demir (1963) found a single 3.5 cm horse
mackerel from the stomachs of 44 juvenile S. sarda rang-
ing in length from 6.5 to 16 cm taken in the Black Sea,
the Sea of Marmara, and the Bosporus from 1957 to 1959.
Because of the preponderance of empty stomachs, Demir
believes that the juveniles, like the adults, regurgitate
their food at the moment of capture.

Demir (1963) stated that adult S. sarda is an insati-
able predator that feeds diurnally. Feeding in Turkish
waters is much more vigorous in the early morning and
towards evening. Demir noted that the feeding season is
usually from the second half of April to the end of Octo-
ber in Turkish waters. Sarda sarda adults primarily feed
on smaller schooling fishes, the species depending on the
locality (Table 15).

In the western Atlantic in the Gulf of Maine, S. sarda
prey on mackerel, alewives, menhaden, other small
fishes such as launce and silverside, and squid (Bigelow
and Schroeder 1953). Boschung (1966) examined the
stomachs of 52 S. sarda taken during a fishing tourna-
ment in the Gulf of Mexico and noted that the fish had
fed on a variety of fishes and invertebrates. They in-

Table 15. — Food of Sarda sarda in the Black Sea, Sea of Marmara,
and the eastern Atlantic. (From Berg et al. 1949; Postel 1954;
Zusser 1954; Slastenenko 1956; Ionescu et al. 1958; Demir 1963.).

Black Sea

Engraulis encrasicholus
Spratella sprattus phalerica
Ammodytes cicerellus
Scorn ber scorn brus
Trachurus mediterraneus
Mullus barbatus

Pomatomus saltatrix (young)

Sarda sarda (young)

Mugilspp.

Atherina spp.

Gobiidae

Sea of Marmara

Important food items
Engraulis encrasicholus
Spratella sprattus phalerica
Scorn ber scombrus
Scomber colias

Less important food items
Smarts alceda.
Trachurus trachurus

Clupea pilchardus
Ammodytes cicerellus
Trachurus mediterraneus

Mullus barbatus
Atherina spp.

Boops boops

Atlantic Ocean near Dakar, Senegal

Sardinella sp.
Engraulis sp.
Scomber colias
Ammodytes sp.
Planktonic crustaceans

Caprella

Euphausiidspp.

eluded: a clupeid (probably Harengula pensacolae)
harvestfish, Peprilus paru; spot, Leiostomus xanthurus
anchovies, Anchoa sp.; mackerel, Scomberomorus sp.
sea robin, Prionotus sp.; squid, Loligo sp.; shrimp,
Penaeus sp.; and unidentified fishes.

Sarda sarda can swallow relatively large prey and the
adults and juveniles are both cannibalistic. There is a
record of a 38 cm S. sarda from the stomach of a 63 cm
fish (Zusser 1954).

3.43 Growth rate

Sarda australis

There is almost nothing in the literature on the age and
growth of S. australis except that they grow to 91.4 cm
(36 in) (Ogilby 1954).

Sarda chiliensis

In the waters between southern California and north-
ern Baja California S. c. lineolata spawns between Janu-
ary and May and the pelagic fertilized eggs take about 3
days to hatch. Young S. c. lineolata are first observed in
the early summer when they are 15.7-25.4 cm (6-10 in)
and by the following spring they are about 38.1 cm (15 in)
long and weigh from 0.7 to 0.9 kg (1.5 to 2 lb). In the fall
these fish may weigh 1.4 kg (3 lb) or more and by the fol-
lowing year in May, they may weigh 2.7-3.2 kg (6-7 lb).
When they are 4 yr old, they are 71.1 cm (28 in) and
weigh about 5.4 kg (12 lb) (Frey 1971). These data are
probably the result of random observations and not a
systematic study.

Kuo (1970) made a detailed study of the growth of S.
chiliensis from southern California waters and from Peru
based on otoliths (Fig. 13). Kuo fitted the von Bertalanf-
fy growth function to his data and found that the length-
at-age of bonito from the two areas did not differ for age
1, 2, and 6 fish but that there was a significant difference
in size offish 3-5 yr old (Table 16). The growth equations
obtained for the northern and southern populations, re-
spectively, were:

Lt = 2,661 [1 - e

-0.038lt+ 06011

and

Lt = 1,014 [1 -e-0154'^

where L, is fork length in millimeters and t is actual age
in years.

Campbell and Collins (1975) also determined the age
and growth of the northern population of S. chiliensis
based on a sample of 3,139 fish ranging in length from 23
to 79 cm landed at the canneries in San Pedro, Calif.
These fish were caught in an area between central Baja
California and Point Conception, Calif., from 1968 to
1974. Like Kuo (1970), Campbell and Collins used
otoliths to age the fish and used the von Bertalanffy
growth function to describe the growth as shown below:

x 50

I-
©

CAMPBELL 8 COLLINS (1975)

/, X Sarda chiliensis lineolato .
/Y \e m ur k, i hKUO 1970)

.' . Sarda chiliensis chiliensis

5
AGE

Figure 13. — Growth of Sarda chiliensis from southern California and
Peru. (Note: The abscissa of the Campbell and Collins curve is age
class.) (From Kuo 1970; Campbell and Collins 1975, fig. 4.)

Table

16. — Age-length data for the subspecies

of Sarda chili

msi's.

(From Kuo 1970, table 14.)

Mean

Confidence

length

Standard

interval

Age

Species

(mm)

error

(mm)

lineolata

152

0.95

150-154

222

chiliensis

151

1.99

147-155

141

lineolata

252

1.28

249-254

219

chiliensis

256

2.76

250-261

137

lineolata

347

1.70

344-351

212

chiliensis

364

6.02

352-376

lineolata

433

2.26

429-438

182

chiliensis

480

7.01

466-494

lineolata

509

3.19

503-515

133

chiliensis

550

7.27

535-565

lineolata

585

4.79

576-595

chiliensis

604

9.18

585-623

lineolata

652

13.00

625-678

chiliensis

—

lineolata

756

12.10

728-783

chiliensis

—

'lineolata = Sarda chiliensis lineolata. [Northern Hemisphere]
chiliensis = Sarda chiliensis chiliensis. [Southern Hemisphere]

L, = 76.87 [1 - e

•0.6215it-H0.410>'

where Lt is fork length in centimeters and t is age at cap-
ture, which is about 0.375 yr (September) older than the
given age.

The length-age relationship predicted by the curve for
fish taken in the fishery is shown below:

Fork length (cm)

44.9

59.7

67.6
71.9

74.2
75.4

The growth curve is also shown in Figure 13.

The maximum size of S. chiliensis in the Northern
Hemisphere is about 101.6 cm (40 in) and about 11.3 kg
(25 lb). There is an unverified record of a 16.8 kg (37 lb)
specimen, but bonito over 11.3 kg are rare (Frey 1971). In
light of these observations, the Lr of 2,661 mm obtained
by Kuo (1970) for the Northern Hemisphere bonito seems
unreasonably large. On the other hand, as noted by
Campbell and Collins (1975), the L„ predicted by the von
Bertalanffy growth function as fitted to their data was
less than what they had observed in the California fish-
ery and much less than the maximum length of S.
chiliensis (101.6 cm) in the literature. They noted that
the discrepancy could have been caused by the lack of
fish larger than 79 cm in their sample.

It is of interest also that the growth curves obtained by
Kuo (1970) and Campbell and Collins (1975) for the
northern population of S. chiliensis are quite different.
The results obtained by Campbell and Collins indicate a
rapid growth rate during the first 3 yr and a tapering off
to a relatively low rate thereafter. Kuo's growth curve on
the other hand shows only a slight curvature and sug-
gests almost linear growth for the northern bonito
population (Fig. 13). The age-length data given by
Campbell and Collins (1975) and Kuo (1970) are also
quite different. The lengths at the different ages as given
by Campbell and Collins are larger for all the ages than
those given by Kuo.

The maximum size of 5. chiliensis in the Southern
Hemisphere is not well defined. Barrett (1971) sampled
the fish landed in the Chilean fishery and noted that the
maximum size of fish landed was 74 cm. In an earlier
study Buen (1958) found fish as large as 79 cm. Mann
(1954) stated that Chilean bonito grows larger than 80
cm.

Sarda orientalis

Very little information is available on the age and
growth of S. orientalis. Yabe et al. (1953) presented data
on the mean lengths of six samples of S. orientalis rang-
ing in length from 161 to 348 mm that were sampled at
Aburatsu, Japan, over an irregular period from 26 August
to 17 October 1950 (Table 17). The mean lengths offish
from the six samples were plotted against sampling date,
and a straight line was drawn by eye through the data
points (Fig. 14). If the line is a reasonable representation
of the growth of S. orientalis between 161 and 348 mm, it
indicates a mean growth of about 2.6 mm/day. Harada et
al. (1974) made observations on the growth of larval and
juvenile S. orientalis in artificial fertilization experi-
ments. They determined that the fertilized eggs hatched
in about 50 h in water temperature ranging from 20° to
24°C and that the newly hatched larvae were 4.1-4.3 mm
in total length. The larvae grew to 14 mm in total length

Table 17. — Lengths of Sarda orientalis landed at Aburatsu, Japan
in 1950. (From Yabe et al. 1953, table 11.)

Body length (mm)

Table 18.— Monthly length data for Sarda sarda from the Black Sea.
(Data from Demir 1963.) (Note: The author did not define the length
dimension.)

Date

1NO. 01

fish

Range

Mean

Author

July

Aug.

Sept.

Oct.

205-221

214

h(cm) - -

Aug. 26

Sept. 1

241-249

245

Zusser(1954)

—

19-27

27-29.5

25-36

Sept. 2

233-243

239

Tkacheva (1958)

—

21-33

27-36

36-40

Sept. 8

161-270

239

Mayorova and Tkacheva (1959)

—

21-33

27-37

36-41

Sept. 9

254-277

263

Demir (1963)

6 L6

12-32

25-38

33-42

Oct. 17

333-348

339

AUG.

Figure 14.-

SEPT. OCT.

-Growth of juvenile Sarda orientalis in Japanese
waters. (Data from Yabe et al. 1953.)

in 10 days, to 74 mm in 20 days, to 106 mm in 30 days, to
219 mm in 42 days, and to 290 mm in 99 days after
hatching. This growth rate represented the fastest
growth under the conditions provided in the experi-
ment. The growth rate of larval and juvenile S. ori-
entalis indicated by these experiments was about 2.9
mm/day during the 99-day period and compares favor-
ably with that suggested by the data presented by Yabe
et al. (1953). Harada et al. (1973) presented some data on
the growth of S. orientalis kept in artificial enclosures.
Thirty-three S. orientalis 40 cm long and 675 g in weight
grew to 50 cm and 1,500 g in about 4 mo.

Regarding the maximum size of S. orientalis in the In-
dian Ocean, the large specimens off the southwest coast
of India are generally less than 70 mm long (Silas 1964).
Smith (1949) stated, however, that S. ehihensis (= S.
orientalis) grows to at least 101.6 cm (40 in). In Japanese
waters S. orientalis grows to about 80 cm and 1.5-3.0 kg
(Kishinouye 1923).

Sarda sarda

Various investigators have presented monthly length
data for young S. sarda from the Black Sea (Table 18).

The data were given as the ranges of lengths sampled
during monthly periods, and the midpoints of the length
ranges were plotted to represent the growth of 3-42 cm S.
sarda in the Black Sea (Fig. 15). As would be expected,
there is some variation in the data presented by the
various investigators, but the variation was relatively
small. In gross terms these data suggest that S. sarda
grew about 252 mm in 90 days (7.5-11 cm in July to 30.5-
38.5 cm in October) or at a growth rate of 2.8 cm/day.

Demir (1963) summarized the age-length relation of S.
sarda determined by various authors (Table 19). Subse-
quent to the publication of Demir's paper, Kutaygil
(1967) published some data on age-length determina-
tions on S. sarda (Table 20). Kutaygil used otoliths to
age the fish and also back-calculated the lengths of the
fish at various ages. He noted that bonito from different
year classes had different growth rates.

Zusser (1954), in contrast to the other investigators,
gave age-length relations for fish up to 9 yr of age. Demir
(1963), however, remarked that the age-length relations
given by Zusser were probably erroneous. Although sev-
eral age-length relations for S. sarda are available, as far
as it is known, no one has tried to fit a growth curve to
these data.

J 20

x x ZUSSER0954)

0 Q TKACHEVA (1958)

v 7 MAYOROVA a TKACHEVA0959)

o o DEMIR (1963)

a A MEYER (1956)

JULY AUG SEPT OCT.

Figure 15. — Growth of juvenile Sarda sarda in the Black Sea.

:si

Table 19.— Age-length relation in Sarda sarda. (From Demir 1963, table 4.) (Note:
The length dimension was not defined by the author.)

Age

Author

- Length (cm) -

Zusser (1954)

25-37

33-50

42-54

50-62

56-67

60-70

63-76

70-78

74-85

31.5

41.5

48.8

56.2

61.2

66.5

69.5

73.5

76.2

Numann (1955)

38-41

53-57

60-64

—

Nikolskii (1957)

35.3

55.1

64.2

72.5

—

Tiirgan (1958)

30-40

50-55

55-60

60-65

—

Nikolov (1960)

38.85

52.6

(74-75)

—

Table 20. — Calculated lengths for each age of Sarda sarda. (From
Kutaygil 1967, table 26.) (Note: The author did not define the length
dimension.)

No. of
fish

Age

Sample

1957

(1955)

44.0

59.1

—

(1954)

45.4

57.9

64.8

—

(1953)

47.6

59.2

64.5

68.7

Granc

mean =

45.1

58.3

64.9

N =

Jan. 1958

(1954)

42.9

53.3

60.6

64.2

Feb. 1958

(1954)

41.1

52.9

59.2

63.8

The largest S. sarda is about 85 cm in the Black Sea.
Of S. sarda in the western Atlantic, Bigelow and
Schroeder (1953) stated that they grow to about 91.4 cm
(36 in) and 4.5-5.4 kg (10-12 lb). Hammond and Cupka
(1975) noted that S. sarda seldom exceeds 6.8 kg (15 lb).

3.5 Behavior

3.51 Migrations and local movements

See also section 5.3.

The seasonal migrations of S. sarda from the Aegean
Sea through the Sea of Marmara to the Black Sea have
been well documented. The migration routes were pre-
sumably determined by the development of the fisheries
in these areas and were verified by tagging experiments
(Demir 1963). The migration from the Aegean Sea to the
Black Sea starts toward the end of April and lasts to the
beginning of June or later. The return migration from the
Black Sea to the Aegean Sea usually starts in Septem-
ber and lasts to the end of November. Some of the bonito
schools returning from the Black Sea may stay in the Sea
of Marmara but others continue on to the Aegean Sea.

The S. sarda in the Adriatic Sea apparently make a
north-south migration within that sea down to around
the Greek islands (Belloc 1954). The bonitos leave the
upper Adriatic in November-December and travel along
both shores of the Adriatic, arriving in Greek waters in
February-March. The return migration starts around
August. In addition, there apparently are schools of
bonito that remain the whole year round in Greek waters
and in waters along the east coast of Sicily.

The relation of the bonitos in the Aegean Sea-Sea of

Marmara-Black Sea complex with the Adriatic Sea
bonito and with the bonitos in the other parts of the
Mediterranean is not clear. Turkish investigators imply
that the stock in the Aegean Sea-Sea of Marmara-Black
Sea complex originates in the Aegean Sea and the Sea of
Marmara (Demir 1957) and remain within this complex.
In the eastern Pacific, over 11,200 S. c. lineolata have
been tagged and released since 1968 along the coast from
Monterey Bay, Calif., to Cape San Lazaro, Baja Cali-
fornia (Collins and MacCall 1977). Recoveries of tagged
fish have indicated that they move randomly in local
California waters, although there was a definite move-
ment down the southern California coast in the winter
and a northward movement in the late summer and fall,
apparently in response to changes in sea temperature.
Their tagging study also indicated that heated water dis-
charges from coastal electric generating stations strongly
influence the migration of young S. c. lineolata. Fish tag-
ged in heated water plumes either remained in the area
of the discharge or tended to migrate to another heated
discharge area.

3.52 Schooling

All species of Sarda generally appear to be schooling
fish. Sarda australis commonly occur in schools in in-
shore coastal waters of Queensland, Australia (Grant
1972). Frey (1971) stated that the S. chiliensis of the
Northern Hemisphere is a pelagic schooling fish.
Tominaga (1943), however, observed that S. orientalis in
Japanese waters do not aggregate densely, rarely come to
the surface of the sea, and always swim around reefs or
near a cape where the current is strong, and that they
never go out to the high seas. Conflicting observations
are available on schooling of S. orientalis in the Indian
Ocean. Silas (1963) stated that schools of adults and
young appear along the southwest coast of India. Si-
vasubramaniam (1969), however, noted that S. ori-
entalis are very seldom seen in surface schools of mixed
tunas and when they are caught from mixed schools, the
catch never exceeds six fish. All of the observations on S.
sarda indicate it is a schooling fish. Demir (1963) stated
that S. sarda gathers in dense schools of many thousands
of fish of about the same size. Sarda sarda in the western
Atlantic is a schooling fish, traveling in large aggrega-
tions. It is usually found at the surface, although oc-
casionally it is caught near the bottom (Idyll and de

Sylva 1963) . Of 5. sarda that occurs in the Gulf of Maine,
Bigelow and Schroeder (1953) wrote, "The bonito is a
strong, swift, predaceous inhabitant of the open sea and
like its tribe travels in schools."

3.53 Responses to stimuli

A detailed, systematic study on the behavior of captive
S. c. lineolata in a large fish tank at Marineland of the
Pacific, Palos Verdes, Calif., was made by Magnuson
and Prescott (1966). The courtship and spawning be-
havior of S. c. lineolata, as reported by Magnuson and
Prescott, was discussed earlier in sections 3.11 and 3.13.
Further results of their observations are detailed below.

About 25 S. c. lineolata out of 60 that were caught near
Marineland of the Pacific became established in the tank
and were used in the observational program. Of the 25
survivors, 10 fish lived for 38 mo in captivity. The tank
used was part of a public display and contained more
than 800 other fishes of over 40 species. Part of the daily
routine included five 15-min shows in which a diver
entered the tank to feed the fish. Magnuson and Pres-
cott noted that the S. c. lineolata "appeared habituated
to show announcements and background music as well as
to sounds made by tapping on the tank windows and
sides." These observations indicate that S. c. lineolata
are able to adapt to various environmental conditions.

In the Marineland of the Pacific tank, S. c. lineolata
swam continuously against the current averaging 88.2
cm/s at a tail-beat frequency of 1.42 beats/s when not
feeding or courting. Sarda c. lineolata apparently are less
powerful swimmers than skipjack and yellowfin tunas. In
fish about the same length (bonito, 57 cm; skipjack tuna,
57 cm; yellowfin tuna, 52 cm) Magnuson and Prescott
noted that at four tail beats/s S. c. lineolata traveled only
170 cm/s whereas Katsuwonus pelamis averaged 230
cm/s and T. albacares averaged 240 cm/s.

Magnuson and Prescott observed nine miscellaneous
behavior patterns in S. c. lineolata: mouth closure (long),
mouth closure (short), snap, yawn, quick swim, lean,
bend, jerk, and defecation. They discussed in detail the
possible functions of all these miscellaneous behavior
patterns. They hypothesized that mouth closure (long)
movements could have been associated with olfaction or
gill ventilation; snaps following a yawn or a quick swim
with a drinking movement; leans may have a function as
a social releaser in schooling; and bends and jerks may be
associated with food passing through the alimentary
canal.

No such detailed behavioral observations have been

Table 21.— Sex ratio of Sarda sarda landed in Spain. (From
Rodriguez-Roda 1966.)

Ratio

Year

Month

Port

Males

Females

(Male:Female)

1963

June

Tarifa

1:1.06

1964

May

Barbate

1:0.35

1964

June

Barbate

1:0.96

1964

July

Barbate

1:1.33

made on other species of Sarda. Inoue et al. (1967) were
able to maintain S. orientalis up to 438 h in a pool. Inoue
et al. (1970) determined that S. orientalis was negatively
phototactic to both sunlight and artificial light. The
swimming speeds of the fish (size not given) ranged from
0.3 to 0.56 m/s.

4 POPULATION

4.1 Structure

4.11 Sex ratio

Sarda australis

No information.

Sarda chiliensis

Kuo (1970) obtained sex ratio data on S. c. lineolata
that were caught from San Diego waters during a 1-yr pe-
riod between 1964 and 1968. The percentage of females in
the monthly samples varied from 37.5 to 70.3% and aver-
aged 49.9% for the year. He found that the monthly sex
ratios did not differ significantly from 1:1 except for the
sample from the month of May.

The percentage of female S. c. chiliensis from eight
samples of 100 bonito each obtained from the commer-
cial catch landed at Iquique, Chile, varied from 47 to 65%
between September 1968 and October 1969 (Barrett
1971). Barrett noted that more females were present in
the catch during September-October spawning season,
but stated that more data were needed to verify this
observation.

Sarda orientalis

No information.

Sarda sarda

Rodriguez-Roda (1966) determined the sex ratio of four
samples of S. sarda landed at the ports of Barbate and
Tarifa, Spain (Table 21), in 1963 and 1964. The females
in the samples ranged from 26 to 57.1%. Postel (1955b)
presented data on the monthly sex ratio of S. sarda from
the eastern tropical Atlantic (Table 22). In the Gulf of
Mexico a sample of 52 S. sarda was composed of 31 males
and 21 females (Boschung 1966).

4.12 Age composition
See section 4.13.

4.13 Size composition

Sarda australis

There is no information on the size or age of the fish ex-
cept for generalized observations. The species grows to

Table 22. — Sex ratio of Sarda sarda from the eastern tropical
Atlantic. (From Postel 1955b.)

Ratio

Month

Males

Females

(Male:Female)

January

1:0.42

February

1:0.79

March

1:1.08

April

142

115

1:0.81

Mas-

148

172

1:1.16

June

1:1

>
o

July

—

August

—

September

—

o
111

October

1:2.67

November

1:0.71

December

1:1.20

111
o
K
Ul
Q.

Total

422

419

1:0.99

about 90 cm (3 ft) but the average size of fish caught is
about 40 or 45 cm (16 or 18 in) (Marshall 1964). Grant
(1972) stated that the usual size taken is 1.8-2.3 kg (4-5
lb).

Sarda chiliensis

The length-frequency distribution of S. c. lineolata
caught by sports fishermen from southern California
waters shown in Figure 16 was prepared from data pro-
vided by C. M. Kuo4 and is based on part of the S. c.
lineolata sample he used in his study (Kuo 1970). Kuo
measured 929 S. c. lineolata from southern California
waters ranging in length from 331 to 750 mm. Both the
male and female length-frequency distributions were
similar: a single prominent mode was present between
510 and 540 mm.

MacCall et al. (1976) presented S. c. lineolata length-
frequency data for the California commercial and party-
boat fisheries in 1973 (Fig. 17). They stated that various
segments of the fishery exploit different parts of the
bonito population, which indicates an uneven geo-
graphical distribution of various age groups. It was indi-
cated that, generally, the older fish were more available
near Mexico and in offshore waters, although large fish
were also taken in the fall off Santa Barbara. The party-
boat fishery caught bonito <60 cm but the long-range
party boats fishing off Mexico took older fish. The com-
mercial fishery took larger fish. It is of interest that the
modal size taken by the party boats during the period
1964-68 was larger than that taken in 1973. Recruitment
strength is highly variable in southern California waters,
and because the party-boat fishery takes only young fish,
the catch per effort and the size composition of the catch
reflect this variability and may account for the differ-
ence in modal sizes (MacCall3).

Barrett (1971) presented bonito length-frequency data
from Chilean landings and noted that the larger bonito,

Presently with Oceanic Institute, Makapuu Point, Waimanalo
96795.

MacCall, A. D., California Department of Fish and Game, c/o Na
tional Marine Fisheries Service, La Jolla, CA 92038, pers. commun
February 1978.

FEMALES
(N = 2I2)

^tH

j=tl

10-

MALES

(N = 202)

r-TTh

rrhJrfr-

□d

35 40 45 50 55 60 65

LENGTH (cm)

Figure lf>. — Length-frequency distribution of Sarda chiliensis
lineolata captured in the sport fishery in southern California.

U.S. COMMERCIAL

CALIFORNIA A
N= 1,502 '
W= 6 45 lbs i

1973

MEXICO
N = 365
W = 7.70 lbs

vIEAN LENGTHS OF AGE GROUPS I

m nz zvrzn

U.S. PARTYBOATS

LOCAL
l\ N = 1,235
I W= 2.87 lbs

y '

LONG RANGE OFF MEXICO
N = II6
W= 8.79 lbs

30 40 50 60 70 80

LENGTH (cm)

Figure 17. — Length-frequency distribution of Sarda chiliensis
lineolata taken by the California commercial party-boat fisheries.
(From MacCall et al. 1976, fig. 9.)

in about the 62 cm modal group, made up most of the
catch in September-October 1968 (Fig. 18). Most of the
bonito in the April, August, September, and October
1969 landings, however, were smaller, 48-52 cm, and the
larger fish were present only in July and August. Barrett

56 60 64
LENGTH (cm)

Figure 18. — Length-frequency distribution of Sarda chilienss chili-
ensis sampled from the commercial landings at Iquique, Chile,
September 1%H to October 1%!). Numerals in upper left corner of
each panel indicate the month and year, number offish, and number
of samples. Unshaded distribution is from Buen (19S8). (From Bar-
rett 1971, fig. 4.)

further noted that the preponderant 72-74 cm modal
group in 1953 reported by Buen (1958) was almost absent
in the 1968 and 1969 samples, which indicated that the
older fish were no longer present in the fishery in 1968
and 1969.

Vildoso (1962) and Barrett (1971) determined length-
weight relationships for the southern S. chiliensis

population and Campbell and Collins (1975) for the
northern population (Table 23). In computing length-
weight relationships, many investigators use a logarith-
mic transformation to linearize the data. Beauchamp
and Olson (1973) pointed out that a bias is inherent in
this procedure because the largest values are compressed
on the logarithmic scale and provided a procedure to cor-
rect for this bias. As they pointed out, corrections for this
bias have been outlined in the past but seldom used in
practice and it is most likely that the length-weight re-
lationships in Table 23 have not been corrected for bias.

Sarda orientalis

Sivasubramaniam (1969) presented length data for S.
orientalis from the Sri Lanka fishery (Fig. 19). The aver-
age size of adult S. orientalis taken off southwest India
was about 45 cm (Silas 1963).

Data on the size composition of the stock of S. ori-
entalis in Japanese waters are almost nonexistent. Yabe
-et al. (1953) presented measurement data on a small
sample of S. orientalis landed at Aburatsu, Japan (Table
17).

Sivasubramaniam (1966) determined the length-
weight relationship for S. orientalis from the waters
around Sri Lanka in the Indian Ocean (Table 23). The
length-weight relationship is based on a sample of 25
fish, and although Sivasubramaniam did not give the
size range of the fish, his figure 14 indicates that the fish
ranged from about 24 to 51 cm.

Sarda sarda

Black Sea landings of S. sarda in the spring of 1955
were dominated by a group of fish centered at a length of
45 cm (Fig. 20). In the spring of the subsequent 2 yr, the
same group of fish (1956, 55 cm; 1957, 60 cm; 1958, 65
cm) dominated the catch. This group of fish was the
result of a strong year-class that originated in 1954 (May-
orova and Tkacheva 1959). Artuz (1959) also noted the
same phenomenon. He sampled the landings at the
Istanbul fish market and found that 3-yr-old fish (1954

Table 2.1. — Predictive length-weight relationships, W = aib, of Sarda. (Note: Various length and weight units
were used in the original regressions, shown in the table. For ease in comparison, the constants were converted
where needed so that the regressions are in terms of weight in grams and length in centimeters.)

Species

Sex

Size range of fish

No. of

Weight

Length

fish

(g)

(cm)

Source

595

40-73

0.0118

3.02

Barrett (1971)

—

0.0152

2.958

Sivasubramaniam (1966)

165

—

40-55.5

0.0149

2.971925

Rodnguez-Roda (1966)

0.009088

3.09749

Campbell and Collins
(1975)

2,824

200-7,675

29-77

0.009611

3.08338

Campbell and Collins

(1975)

0.009376

3.08962

Campbell and Collins
(1975)

565

—

0.006491

3.19

Vildoso (1962)

513

—

0.006311

3.19

Vildoso (1962)

S. c. chiliensis
S. orientalis
S. sarda
S. c. lineolata

S. c. lineolata

S. c. chiliensis
S. c. chiliensis

Sexes com-
bined
M
F

WEST COAST

N = 20

SOUTH COAST

N = 27

18-22 22-26 26-30 30-34 34-38 38-42 42-46 46-50 50-54
FORK LENGTH (cm)

Figure 19. — Length-frequency distribution of Sarda orientalis
around Sri Lanka (Ceylon) (1967-68). (From Sivasubramaniam
1969, fig. I.)

O 40

I 1 1 I 1 1

1955
N = 1,286 A

II
h . 1

1956
N = 1,149

1957

N = 10,881

! i i i r

1958

N = 5,167

a :

1 I : _

\ ■

30 35 40 45 50 55 60 65 70 75
LENGTH (cm)

Figure 20. — Length-frequency distribution of Sarda sarda in the
Black Sea, spring 1955-58. (Data from Mayorova and Tkacheva
1959, table 1.)

year class) dominated the landings in May 1957. His
sample showed that the May 1957 landings were com-
posed of 10.5% age I fish (1956 year class), 28.7% age II
fish (1955 year class), 53.2% age III fish (1954 year class),
and 7.6% age IV fish (1953 year class). Mayorova and
Tkacheva (1959) noted that a rich year class in 1938 also
dominated the fishery in the Black Sea from 1938 to
1945.

The 1963 length-frequency distribution of S. sarda
landed at Barbate and Tarifa, Spain, showed a strong
mode centered at around 41 cm and lesser modes be-
tween 49 and 59 cm and 59 and 69 cm (Fig. 21). The 1964
sample had only two modes: one at about 43 cm and the
other at about 51 cm. Rodriguez-Roda (1966) also pre-
sented a weight-frequency distribution of S, sarda (Fig.
22), which showed only two modes for the 1963 sample
and only one well-defined mode for the 1964 sample. The
1964 data were also summarized by month (Fig. 23). The
length-frequency distribution showed two well-defined
modes in May, June, and July whereas the weight-fre-
quency distribution showed only a single mode during
those months. It is of interest that the relative propor-

iii
1963

— i
\
\

i \

I \ . 1964
\ \ / \

1 ^

/ ;
/ /
/ i
/ /

/ /
/ /
J i
/ /

\ V
\ \
\ V
\ \
\ \
\ \

\ \ ^

V v /"' — \

^■— T ,~^r^ "~

35 40 45 50 55 60 65 70

LENGTH (cm)

Figure 21. — Sarda sarda length-frequencv distribution. Barbate

/,, + 2/„+/,1Tl

and Tarifa, Spain. Data smoothed by formula /„ = .

(From Rodriguez-Roda 1966, fig. 11.) 4

tion of fish in the larger mode in the length-frequency
distribution appeared to decrease from May to July. The

O 20-

K A

1 1 1 1

/\ / \

/ \ \

A \

/ \ '

/ /
/ /
/ (
/ /
-/ '
/ /

/ \

; \

/ \

1963 \

\
\

\ .-1964

/ /
/ /
7 /

/
\i
i

■

\ V
\ \
\ \
\ \
\ \

0 12 3 4 5

WEIGHT (kg)

Figure 22. — Sarda sarda weight-frequency distribution, Barbate

/n-,+2/„+/„+.

and Tarifa, Spain. Data smoothed by formula fn =

(From Rodriguez-Roda 1966, fig. 14.) 4

O 20-

40 50

LENGTH (cm)

I 2 3

WEIGHT (kg)

Figure 23. — Sarda sarda length (Barbate and Tarifa) and weight
(Barbate) frequency distribution 1964. (From Rodriguez-Roda
1966, figs. 12, 15.)

length-frequency distribution of male and female S.
sarda were similar except that the modes were displaced
(Fig. 24).

45 50

LENGTH (cm)

Figure 24. — Sarda sarda length-frequency distribution, Barbate
and Tarifa, Spain, arranged by sex. (From Rodriguez-Roda 1966,
fig. 13.)

Postel (1955a) presented data on the maximum size of
S. sarda sampled in monthly periods in the eastern
tropical Atlantic. The maximum lengths ranged from 450
to 690 mm for the males and 443 to 714 mm for the
females.

Rodriguez-Roda (1966) computed the length-weight
relation of S. sarda landed at the southern Spanish port
of Barbate (Table 23).

4.14 Subpopulations

It appears that the S. c. lineolata found off northern
Mexico and southern California comprise a single homo-
geneous stock. Tagging experiments conducted by the
California Department of Fish and Game indicate that
the bonito does not make long migrations. Although
some tagged fish have traveled as far as 600 miles, most
of the tagged fish have been recaptured in the vicinity of
release; fish that traveled long distances moved from
Mexico to southern California waters in the summer and
back to Mexican waters again in the winter (Frey 1971;
Collins and MacCall 1977).

From all indications it appears that the Northern and
Southern Hemisphere populations of S. chiliensis are
completely separate with little or no interchange. They
are geographically separated from each other and there
are certain meristic and morphological differences be-
tween the two populations. For example, the Northern
Hemisphere population of bonito averages more verte-
brae than the Southern Hemisphere population (Vildoso
1963b; Kuo 1970; Collette and Chao 1975).

Some preliminary work has been done on S. chiliensis
to determine if protein differences attributable to genetic
variation could be useful in identifying population
units. Barrett and Williams (1967) experimented with

gel-electrophoresis of the soluble eye lens proteins of
bonito in an attempt to find such genetically controlled
differences. They did find polymorphisms of the soluble
lens proteins for the bonito. They also calculated the
gene frequencies together with their expected distribu-
tions and found conformity to the Hardy-Weinberg prin-
ciples. However, Barrett and Williams also found that
the distributions of the apparent phenotypes were re-
lated to the lengths of the bonito. They concluded, there-
fore, that ontogenetic factors caused the observed poly-
morphism and that gel-electrophoresis of the soluble eye
lens proteins, under the conditions used in their experi-
ment, was not a useful technique in differentiating popu-
lation units of bonito.

Smith (1971) examined the electrophoretic patterns of
nuclear lens proteins from S. c. lineolata and also found
polymorphism in the protein patterns. He suggested that
there could be another explanation, as stated by Eckroat
and Wright (1969), for the observed distribution of
phenotypes than that suggested by Barrett and Williams
(1967). Eckroat and Wright (1969) suggested that two
separate fractions inherited as present/absence differ-
ences could account for the distribution of the three
phenotypes observed by Barrett and Williams (1967). If
this situation obtained, then the distribution of the
observed phenotypes would indicate that there were two
different breeding populations represented in the catches
sampled.

It is not clear whether the stocks of S. c. chiliensis off
Peru and Chile are independent of each other or whether
they form a single homogeneous population. The center
of bonito abundance off Peru is from Chimbote to Pisco
(Ancieta 1963), which is centrally located between the
north and south borders of Peru. The center of abun-
dance off Chile is closer to the northern boundary of
Chile near Arica, Iquique, and Antofagasta (Barrett
1971). In other words, the stocks of S. c. chiliensis off the

20
18

m 12

X
Q

8 8

6
<\
2

— 1 — 1 1 1
X

/ \

/ ANNUAL \

^^ \ Y

^^"^ a. \ Y

3o-~^ ,o-,:nov

y°

y ■•■•. y ^O^ \.

2°- ■■-,>. ^-^--r~

S*' In

"**■ ^ ^*v>-^-J^-»

^C" .-°

*•.. O"..^ ^o-- "y

'"***"-o

"■f

1965

1966

1967

1968

1969

Figure 25. — Apparent abundance of Sarda chiliensis chiliensis,
for individual quarters and years, by year, from data for the
bonito vessels. (From Barrett 1971, fig. X.)

coasts of Chile and Peru are at least contiguous with each
other. In his attempts at determining a stock production
model for the Chilean bonito, however, Barrett (1971)
apparently considered only the Chilean stock. He listed
as one of his recommendations, however, that future
studies should include a "determination of relation, if
any, with the Peruvian fishery for bonito."

No information is available on subpopulations of S.
orientalis and S. sarda. The coastal distribution of all the
species of Sarda, the migration patterns of S. c. lineo-
lata, and the disjunct distribution of S. orientalis and S.
sarda suggest the existence of discrete subpopulations.
Based on scattered anatomical and meristic data, there
appeared to be no difference in the populations of S. ori-
entalis, although some morphometric differences were
found in samples from Japan and the eastern Pacific.
Similarly, a geographical comparison of S. sarda samples
showed no conclusive differences among the various
populations (Collette and Chao 1975).

4.2 Abundance and density

Sarda australis

Marshall (1964) said that S. australis, along the coasts
of Queensland and New South Wales, are found in "great
schools throughout the year." Grant (1972) stated that
this species occurs in schools in Queensland waters espe-
cially during the winter months.

Sarda chiliensis

In the Chilean bonito fishery Barrett (1971) examined
the relations among fishing effort, yield, and apparent
abundance for the period from January 1965 to Decem-
ber 1969. Barrett recognized three different types of ves-
sels, according to species objective, that fish bonito in the
Chilean fishery: the anchovy, bonito, and tuna vessels.
As the name implies, the anchovy and tuna vessels fished
for bonito when anchovy and tunas, respectively, be-
came less available and bonito more available. The
bonito vessels fished primarily for bonito, although they
did also capture other species. Barrett used the data from
the bonito vessels for his analysis. The data for the entire
monitored fleet, however, showed the same trends as for
the bonito fleet after they were standardized to that of
the bonito vessels. He noted a steep downward trend in
total catch, an upward trend in relative fishing intensity,
and a corresponding decline in apparent abundance in
the fishery (Figs. 25, 26, 27). The trend appeared to level
off in 1969. Barrett stated that the decline probably re-
sulted from the effects of the fishery and not from fish-
ery-independent factors.

MacCall et al. (1976) related the indices of abundance
of the Northern Hemisphere S. chiliensis obtained by
Squire (1972) on aerial surveys with catch per unit of ef-
fort (CPUE), in terms of catch in numbers of fish per
angler, for the California party-boat fishery. MacCall et
al. assumed that the aerial survey concentrated on the
commercial fishing grounds because the surveys were

b(J
50

1 1 1 4 1 1 1 1 1 ! 1 I

/\ t l\ TOTAL CATCH

/ \ \ l\

\ \ 1 \

j \a / 1 / 1

o
_l
*:

^J 1 M /

W \y

i — i — i — i — i — r

RELATIVE FISHING INTENSITY

1 i 1 I 1 1 1 1 1 ! 1 1 1 1 1 1 1 1 1
A CATCH PER UNIT OF EFFORT

t! i i b — <r

Figure 26. — Sarda chiliensis chiliensis total catch, and relative
fishing intensity and apparent abundance (from data for the bonito
vessels), quarterly from 1965 to 1%9. Relative fishing intensity
for 1969 is not shown because data for total catch in 1969 were not
available. (From Barrett 1971, fig. 7.)

designed to aid the commercial fishery, and thus re-
flected changes in the stock occurring prior to exploita-
tion by the commercial fishery. Also, since the length-
frequency data showed that the party boats exploited
younger bonito than the commercial fishery, they further
assumed that the party-boat CPUE provided an index of
prerecruit abundance of fish before they are exploited by
the commercial fishery approximately 3 yr later. They
computed a "combined" party-boat CPUE index which
took into consideration mortality and recruitment and
related it to the aerial survey day index for the period
1963-72 (Fig. 28). They concluded that the party-boat
CPUE appears to be a valid indicator of recruitment to
the commercially exploitable segment of the bonito
stock, at least during this period. The party-boat CPUE
from 1936 to 1973 indicated recruitment was very low be-
fore 1957, after which it increased sharply (Fig. 29) (Mac-
Call et al. 1976). MacCall et al. further stated that S. c.
lineolata, for unknown reasons unrelated to fishing, be-
came scarce during the early 1940's and that sub-
sequently and until 1956, the party boats were de-
pendent on migratory fish. They noted that after 1956
"... large quantities of bonito moved into California

1966

1 1 1 1

^^v^teai

BONITO VESSELS

N%s

\ vo>

S.J965

\ \. 1967
\ \

\ N
\ N.

V \ 1968

• o

1 1

l ; i i

18 19 20 21 22 23 24 25 26 27 28

RELATIVE FISHING INTENSITY ( DAYS X I02)

Figure 27. — Apparent abundance of Sarda chiliensis chiliensis in
relation to relative fishing intensity from data for the bonito vessels
only (dashed line) and for the fleet monitored by IFOP (Instituto de
Fomento Pesquero) (solid line), 1965-68. (From Barrett 1971, fig. 9.)

2 3 4

1965

1966

1967

1968

1969

LU
Q

ol972

0 0.5 1.0 1.5 2.0

AERIAL SURVEY DAY INDEX

Figure 28. — Regression of aerial survey day index against CPUE
adjusted for mortality and recruitment for Sarda chiliensis lineo-
lata. (Note: Although the authors did not state how the line was fit,
presumably, the least squares method was used.) (From MacCall et
al. 1976, fig. 12.)

waters and became re-established as a locally spawning
population." During the 1960's young S. c. lineolata were
abundant inshore and sports fishermen landed record
numbers; however, since 1969 the sport catch has fallen
drastically (Collins and MacCall 1977).

Sarda orientalis

The mean monthly landings and catch rates for S. ori-
entalis in waters off southern Japan (Fig. 30) showed that

g 1.5

CPUE INDEX
(WEIGHTED BY AREAS)

IC/If

1935 1940 1945 1950 1955 I960 1965 1970 1975
YEAR

Figure 29. — Party-boat Sarda chiliensis lineolata CPUE as an index
of annual recruitment. (From MacCall et al. 1976, fig. 13.)

JAN FEB • MAR APR MAY JUNE JULY AUG SEPT. OCT NOV. DEC.

Figure 30. — Seasonal landings and catch rates of Sarda orientalis
in southern Japan. A — Mean monthly catch per trip by trolling
in the Aburatsu, southern Kyushu, Japan, fishery for Sarda ori-
entalis, 1970-73. B — Mean monthly landings of Sarda orientalis
at Tosashimizu and Muroto, Kochi, Japan, by various fishing
methods 1967-73. (Data provided by S. Kikawa, Far Seas Fish-
eries research laboratory, and T. Koto, Nansei Regional Fisher-
ies Research Laboratory, Japan.)

ability or abundance of S. orientalis off the coast of
southern Japan.

Sarda sarda

Wide fluctuations in the landings of S. sarda in the
Black Sea-Mediterranean Sea area are a characteristic of
this fishery (Fig. 31), and investigators in this area have
been trying to find the causes of the fluctuations. Artuz
(1959) noted that the landings at the Istanbul fish
market showed fluctuations following a 9-yr cycle during
the period from 1936 to 1958 (Fig. 32). As stated earlier,
part of the fluctuations were explained by the entry into
the fishery of rich year classes. However, Mayorova and
Tkacheva (1959) pointed out that although relatively
good year classes develop in certain years, these fish fail

YEARS

Figure 31. — The catches of Sarda sarda recorded at the Istanbul
fish market, Turkey, 1909-60 (1 pair = two fish). (From Demir 1963,
fig. 13.)

> HYPOTHETICAL FLUCTUATION CURVE

> SMOOTHED VALUES ACCORDING TO THE FORMULA

3
2-

i i : ' ' ' ' ;

9""9 —VERY GOOD YEARS-
l i
/ \
1 \
'R '

-f

1
1
1

1
I
1

1 1 1

l\ 1
/ \ 1

/ V

/ \ i
/ \ 1 '

i \\

f \ o— GOOD YEARS— f

\ \ '

1
1
1

\
\

S
\
i

1
1

\ i 1
\ t '

V i i
Vi A '
\\ /\ '

I
\
\

//
//

\
\

L
\

V / \ ' 1

_MEDIUM V^>^/ \L-J

YEARS \ 1
\ 1
1 /
. /
| /

v \
\ \

\ °

\ I

\ 1

//
//

//
il
if

\ 1
\ 1
\ 1
\ 1

—BAD YEARS — 'v J

I 1 1 I 1 I I 1 I I

r ^ti

i i

! 1

1 1

both the landings and catch per trip of S. orientalis were
relatively higher during the fall-winter months, indi-
cating that there are seasonal differences in the avail-

YEARS

Figure 32. — Fluctuation pattern of Sarda sarda landed at the Istan-
bul fish market 1936-58. (From Artuz 1959, fig. 1.)

to return to the Black Sea. They concluded that ". . . the
rich yield of young fish in the Black Sea is not always fol-
lowed by an increase in the abundance of large pelamid."
Artuz (1959) also indicated a possibility of an inverse
relation between the relative abundance of S. sarda and
the mackerel, Scomber scombrus, in Turkish waters
from 1940 to 1956 (Fig. 33).

1 1 1 1 1 1 1

1 1 1 1

Sarda sarda

I06 PAIRS

0 o ° K •

\
\
\
\

\
s

\ f

\ /
\ /
\ /

\ /

\ /
\ /
\ /
y

' \
/

/ i

;
/
/ Uskumru

1 1 1 1

v°-

**o—

-o io5 kg

[III

1955

YEARS

Figure 33. — Alternative occurrence of Sarda sarda (pairs) and
uskumru, Scomber scombrus, (kg) landed from Turkish waters
during the period 1940-56. (From Artuz 1959, fig. 5.)

4.5 Dynamics of population

Barrett (1971) was unsuccessful in developing a stock
production model, following Schaefer (1954), for the
Chilean bonito fishery. He cited as the cause of this the
fact that the necessary assumptions for this type of
model were unsatisfied. One of the requisite as-
sumptions of the model is a stable age and size distribu-
tion in the population. The rapidly occurring changes in
the fishery since 1965 have likely made this assumption
untenable, according to Barrett. Another factor that af-
fected the development of the model was the apparent
discontinuous seasonal availability of some size classes of
bonito in the fishery. Barrett concluded that he could not
determine whether the fishery was stabilized or in a state
of overfishing, and therefore recommended an acceler-
ated research program on the Chilean bonico.

MacCall et al. (1976) developed a surplus production
model for the Northern Hemisphere bonito in California
waters (Fig. 34). They used total catch data and a com-
bination of CPUE for the party-boat fishery and aerial
survey index for the period 1963 through 1972 to develop
the model. The model indicated that the Northern Hem-
isphere bonito is being harvested at or above the maxi-
mum sustainable yield. MacCall et al. cautioned,
however, that the assessment was confounded by the
possibility of a density independent decline in recruit-

ment. They noted that the California catch of S. c. lineo-
lata in 1973 appeared to be greatly in excess of the
equilibrium yield. They stated, however, that con-
clusions drawn from the production model must be
viewed with caution because bonito fishing in California
waters is influenced by ocean temperatures.

Because of the lack of necessary data, MacCall et al.
did not attempt a yield per recruit analysis.

Collins and MacCall (1977) determined a spawner-
recruit relationship for S. c. lineolata during the period
from 1964 to 1974 and found no simple functional curve
to describe the relationship. They found that in some
years a large spawning biomass was related to larger re-
cruitment than did a small spawning biomass; however,
in some years poor recruitment was also related to a high
spawning biomass as well as to a low spawning bio-
mass.

3 0

15 20 25 30 35
EFFORT INDEX (XIO7)

40 45

Figure 34. — Sarda chiliensis lineolata equilibrium yield and abun-
dance in California waters. (From MacCall et al. 1976, fig. 14.)

4.6 The population in the community and the
ecosystem

MacCall et al. (1976) analyzed trophic interaction be-
tween forage (northern anchovies, Engraulis mordax,
and Pacific sardines, Sardinops sagax) and game fish
(yellowtail, Seriola dorsalis; bonito, Sarda c. lineolata;
albacore, Thunnus alalunga; barracuda, Sphyraena
argentea; and white seabass, Cynoscion nobilis) in Cali-
fornia waters. It appeared that there was little evidence
that either forage or game fish biomass showed strong de-
pendence on the abundance of the other. It appeared
more likely that the abundance of both was determined
largely by external conditions.

5 EXPLOITATION
5.1 Fishing equipment

Sarda australis

Sarda australis are generally taken incidentally by
trollers who use the fillets as snapper bait or by sports
fishers who use it as bait for billfishes and sharks (Grant

1972).

Sarda chiliensis

The California fishery uses various kinds of gear to
catch S. c. lineolata including trolling gear and purse
seine. The largest landings are made by the local purse
seiners and the trolling fleet lands a lesser amount. The
high-sea purse seiners may occasionally catch bonito to
fill out the loads of other tunas (Frey 1971).

In a study of the California sport fishery it was found
that S. c. lineolata made up the biggest part of the fish
catch by a single species (Pinkas et al. 1968). Bonito are
taken in party and private boats, from piers and jetties,
and from the shoreline. They will strike at most bait and
lures, and fishing techniques for bonito vary from still
fishing to trolling.

The Chilean bonito fishery prior to 1964 was conducted
almost entirely on an "artisanal, semi-industrial day-
fishery basis" primarily from the ports of Iquique and
Antofagasta (Barrett 1971). The vessels used in this fish-
ery are known as "faluchos" and are about 10 m long.
They fish principally with floating gill nets or with small
purse seines called "boliches," which are hauled manual-
ly. During the period from 1964 to 1966, two modern 36 m
tuna/bonito seiners and eight other bonito/tuna seiners
entered the fishery. All the new purse seiners in the
bonito fishery have power launches and power blocks and
use nylon nets ca. 549-732 m (300-400 fathoms) long and
ca. 73-110 m (40-60 fathoms) deep.

The fishing gear and vessels of the Peruvian fishery
have been described by Ancieta (1963).

Sarda orientalis

In Japan this species is taken by various gear including
trolling gear, pole and line, purse seines, and set nets.
There is no exclusive fishery for S. orientalis in Japan; it
is taken together with species that inhabit or enter the
coastal waters of Japan in multispecies fisheries employ-
ing various gear. In the Philippines S. orientalis are
taken mostly in traps (Warfel 1950). In the Indian Ocean
fishery S. orientalis are taken primarily by drift nets (gill
nets) in India and Sri Lanka.

Sarda sarda

Sarda sarda are taken by trap net, ring net, gill net,
trammel net, purse seine, beach seine, and hook and line
(Demir 1963).

Figure 35. — Fishing ports of the Peruvian and Chilean fish-
eries for Sarda chiliensis chiliensis. (Adapted from Barrett
1971, fig. 1.)

5.2 Fishing areas

Sarda australis

There apparently is no well-developed fishery for S.
australis in Australia. It occurs on the eastern coast of
Australia along Queensland, New South Wales, and as
far south as Port Fairy in Victoria. Serventy (1941a)
stated that ". . . the bonito appears to be numerous
enough to be regarded as a commercially important tuna,
and a fishery could be maintained throughout the year
on the eastern coast." Marshall (1964) noted that S.
australis is destined to be of economic importance in
Australia some day. There seems to be a difference of
opinion on the food quality of 5. australis. Grant (1972)
stated that the flesh of the Australian bonito is dark red
and its edible qualities are not highly regarded, whereas
Marshall (1964) claimed that the flesh is light-colored, of
delicate flavor, and of good canning quality.

Sarda chiliensis

The Northern Hemisphere S. chiliensis is most abun-
dant in the area from Point Conception, Calif., to
Magdalena Bay, Baja California (Frey 1971), and pre-
sumably this is the area in which the California-Mexico
fishery for this species is conducted. Although the
market for this species is limited, this bonito has been
commercially fished in California waters since around
1900. However, the percentage of the California catch

made north of the California-Mexico border has been de-
clining. From 1965 to 1968, over 90% of the California
catch was made north of the border; during 1969-74, it
was reduced to 70%. There are two semi-independent S.
c. lineolata fisheries in the northeastern Pacific: a south-
ern California fishery conducted by small purse seiners
and a fishery conducted by the high-seas tuna fleet in
southern Baja California (MacCall see footnote 5).

The Southern Hemisphere population of S. chiliensis
is exploited by the fisheries of Peru and Chile. The Peru-
vian fishery extends along the coastline from the port of
Mancora (ca. lat. 04°S, long. 81 °W) to the port of Ilo (ca.
lat. 17°30'S, long. 71°15'W) (Ancieta 1963), and the
Chilean fishery extends from Arica (ca. lat. 20°15'S,
long. 70°15'W) to Talcahuano (ca. lat. 37°45'S, long.
73°W) (Barrett 1971) (Fig. 35).

most abundant in the coastal waters of Kyushu (Kikawa
and Staff of the Nankai Fisheries Research Laboratory,
Kochi, Japan 1963).

In the Indian Ocean minor fisheries for S. orientalis
have been reported off the southwest tip of India (Silas
1964), in the Gulf of Aden along the coast of Somalia
(Laevastu and Rosa 1963), and around Sri Lanka (Si-
vasubramaniam 1969).

5.3 Fishing seasons

Sarda australis
No information.

Sarda chiliensis

Sarda orientalis

Fisheries for this species are not very well developed
throughout its distributional range. The FAO Yearbook
of Fishery Statistics (e.g., FAO 1974) does not show any
S. orientalis landings. The S. orientalis found in the
tropical eastern Pacific between central Baja California
and Peru is of little commercial importance and when
caught may enter the catch statistics grouped with S.
chiliensis (Pinkas 1961). This species is uncommon
around Hawaii, and when landed the catches are not
identified in the statistics published by the State.

In Japan S. orientalis occurs in waters south of central
Honshu along both the Pacific and Japan Sea coasts. It is

i — |

i — .

f— 1 | — 1

i — |

Jl _

JAN. FEB MAR. APR MAY JUNE JULY AUG SEPT. OCT NOV. DEC.

Figure 36.— Mean monthly landings and shipments into California
of Sarda chiliensis lineolata, 1968-72. (Data from Heimann and
Carlisle 1970; Pinkas 1970, 1974; Bell 1971; Oliphant and Marine
Fisheries Statistics Staff. Marine Resources Region 1973.)

The mean monthly landings of S. c. lineolata in Cali-
fornia for the 5-yr period from 1968 to 1972 indicate that
the fishery is seasonal (Fig. 36). Although bonito was
landed throughout the year in this 5-yr period, peak
landings occurred in August-October.

Bonito is landed throughout the year in Chile and
Peru, but both fisheries have seasonal peaks (Fig. 37).
The seasonal peak is in October-November in Chile,
whereas it occurs in January in the Peruvian fishery.

PERI

- -

i— i

- -

—

. _

CHILE

1 1

□ 1951,1958,19
O 1965-1968

59 8 1963

R i

'ft;

(-1

r-fl

771

: .

1! :,

ft]

JAN. FEB. MAR APR MAY JUNE JULY AUG SEPT OCT, NOV. OEC.

Figure 37. — Mean monthly landings of Sarda chiliensis chiliensis
in Peru, 1951-59 (data from Ancieta 1963, table 1) and Chile (from
Barrett 1971, fig. 2.)

Sarda orientalis

In the Indian fishery stray specimens may be taken in
April. May, and June; however, the main fishing season
appears to be from about July to September (Silas 1963).
In nearby Sri Lanka, 5. orientalis appears in the catches
throughout the year off the south coast. Mature fish are
taken along the northeast coast from June-August and
off the south and southwest coasts between September
and February. The juveniles appear off the west coast
usually during the southwest monsoon in June-August
(Sivasubramaniam 1969).

Sarda sarda

The fishing seasons for S. sarda in various areas are
summarized in Table 8. Rodriguez-Roda (1966) sum-
marized the fishing seasons in the Spanish fishery by
various geographical areas in the eastern Atlantic and
the western Mediterranean (Table 9). Apparently the
fishing seasons in the Black Sea- Aegean Sea areas are re-
lated to the migrations of S. sarda. There is disagree-
ment on the exact routes of the migration in certain
areas, and the routes are unclear in other areas, but it is
certain that S. sarda does migrate back and forth from
the Aegean Sea, Sea of Marmara, and the Black Sea. Ac-
cording to Demir (1957), schools of S. sarda spend the
summer in the Black Sea and the winter in the Sea of
Marmara and the Aegean Sea. Thus, the fishing season
of May-October coincides with the time the migrating S.
sarda are in the Black Sea in the summer. The fishing
peaks of April-May and September-December in the Sea
of Marmara coincide with the spring migration of the
bonitos from the Aegean Sea through the Sea of Mar-
mara to the Black Sea and the fall return migration from
the Black Sea to the Aegean Sea.

No information is available on the migration of S.
sarda in the other areas of the Mediterranean and in the
Atlantic Ocean. The seasonal development of the
Spanish fishery in various areas of the western Medi-
terranean, however, suggests that S. sarda may be mi-
grating to and from the western Mediterranean (see
Table 9).

5.4 Fishing operations and results

The four species of Sarda are the bases of many fisher-
ies throughout the world. Detailed information on the
fisheries, however, is available only from a relatively few
fisheries. Sarda australis, being found only along the
coast of eastern Australia and around Norfolk Island east
of Australia, is the object of a small incidental fishery in
Australia. According to statistics published by FAO in
the Yearbook of Fishery Statistics, S. c. lineolata is the
basis of fisheries by Mexico and the United States. Sarda
c. chiliensis is exploited by Chile and Peru. Landings of
S. orientalis do not appear in the FAO Yearbook of
Fishery Statistics (see FAO 1974). Relatively small
amounts of the species are taken throughout its distri-
butional range and it is not the basis of any well-de-

Sarda sarda

1965 1966 1967 1968 1969 1970 1971 1972 1973

Figure 38. — Annual world landings of Sarda chiliensis and Sarda
sarda, 1965-73. (Data from FAO 1974.)

veloped fishery. Sarda sarda, on the other hand, appears
to be exploited throughout its distributional range in
fisheries of various sizes. In the Atlantic Ocean, the FAO
Yearbook lists 18 countries reporting landings of S.
sarda, mostly from the eastern Atlantic. Thirteen
countries report landings of S. sarda from the Medi-
terranean and Black Seas. The total landings of S. chili-
ensis ranged from 53,300 to 94,100 t during the period
1965-73 and were greater than the total landings of S.
sarda in each of 9 yr. The landings of S. sarda ranged
from 25,000 to 65,700 t during the same period (Fig. 38)
(FAO 1974).

Sarda australis

As can be expected from an incidental type fishery, the
annual landings of S. australis were erratic and in some
years no fish were landed at all during 1955-73 (Fig. 39).
Information is meager on the utilization of the S.
australis that are landed. They apparently are not un-
common in the wholesale fish market of Sydney,
Australia (Collette6).

Sarda chiliensis

The landing figures of S. c. lineolata in California and
Mexico show that Mexican landings make up only a
small proportion of the total in the Northern Hemis-
phere (Fig. 40). From 1965 to 1973, the California
landings constituted from about 77 to 100c"c of the total
landings of bonito north of the equator in the eastern Pa-
cific. The annual landings have been erratic, and to some

''Collette, B. B., Systematics Laboratory, National Marine Fisheries
Service, NOAA, National Museum of Natural History, Washington, D.C.
20560, pers. commun. February 1978.

2 -

— 1 . i i i i i r 1

ZJ "

NO CATCH
NO CATCH
NO CATCH

[-]

Zl
ZZ\

NO CATCH
NO CATCH

NO CATCH

III . .

ID 113 IP (C tC

m m m m in to

en 0) <x> en en <D

^fjcDcocDcur^r-r-

oicDoioicjioicfiai

Figure 39.— Annual landings of Sarda australis in Australia. (Data
from Australia. Fisheries Branch, Department of Primary Indus-
try 1957-74.)

<2 800

600

400-

?00

1955

1970

Figure 41.— Landings of Sarda chiliensis lineolata by the Cali-
fornia party-boat fleet. (Data from Frey 1971; Oliphant and Marine
Fisheries Statistics Staff. Marine Resources Region 1973.)

TOTAL

t CALIFORNIA

MEXICO

/ \

1920

1930

1940

1950

I960

1970

Figure 40. — California and Mexico landings of Sarda chiliensis
lineolata. (Data from Bell 1971; Frey 1971; Oliphant and Marine
Fisheries Statistics Staff. Marine Resources Division 1973; FAO
1974; Pinkas 1974.)

extent reflect the relative availability of the species in
southern California and Mexican waters. It was believed
that up until the early 1970's the landings did not reveal
the condition of the stock because the bonito is caught
incidentally or in lieu of more desirable species in the
California fishery (Frey 1971). However, it is now be-
lieved that the recent small landings in California strong-
ly indicate a low level of bonito abundance (MacCall see
footnote 5).

The California party-boat landings of S. c. lineolata
(Fig. 41) probably also better reflected the availability of

bonito rather than the condition of the stock up until the
early 1970's. The party-boat fishery is a multispecies
fishery and many other species of fish in addition to the
bonito are taken. Furthermore, the bonito is under a
form of management in that sports fishermen may not
possess more than 10 bonito/day. There is some indica-
tion that water temperature may affect the relative
availability of bonito to the party-boat fishery in Cali-
fornia waters. Party-boat landings of bonito were lower in
the 6-yr period 1948-53, which were cold-water years in
California (Young 1969). More recent data indicate that
party-boat landings reflect the recruitment strength to
the northern segment of the stock and therefore indicate
the condition of the stock itself a couple of years later;
also, catch per angler is a better measure of young fish
abundance than is total landings (MacCall see footnote
5).

The landings of S. c. chiliensis in the fisheries of Peru
and Chile (Fig. 42) indicate that the bulk of the landings
is made in the Peruvian fishery. Chile's share of the
Southern Hemisphere landings of bonito varied from
about 2 to 15% of the total annual landings from 1965 to
1973. The annual Peruvian landings except for minor
fluctuations increased steadily from 1941 and reached a
peak of 104,000 t in 1961. The landings, with minor
fluctuations, have been declining since then. Except for
1955, when the landings of bonito amounted to 7,500 t,
the annual landings in Chile were <5,000 1 during the pe-
riod from 1940 to 1963. The landings increased sharply in
1965 and reached a peak in 1966 as the result of the entry
of several new, modern purse seiners into the fishery in
1964-66 (Barrett 1971). They have, however, declined in
subsequent years.

970

Figure 42. — Landings of Sarda chiliensis chiliensis in Peru and
Chile. (Data from Ancieta 1963; FAO 1965, 1974; Barrett 1971.)

Sarda orientalis

The landings of this species are not identified in the
statistics published by the Japanese Government. Ap-
parently, landing records are maintained by prefectural
"fishery guidance stations," but these statistics are pre-
sumably not published or not readily available.

Table 24 gives the annual landings made at four ports
in southern Japan, Tosashimizu, Muroto, Murotomi-
saki, and Aburatsu, from 1967 to 1974.

There apparently is a minor fishery for S. orientalis in
the Philippines. Warfel (1950) made a survey of the out-
look for the development of a tuna industry in the Philip-
pines. He presented some statistics showing the landings
(number of fish) of S. orientalis at three Philippine
markets: 369 at Batangas, 2,308 at Iloilo, and 6,762 at
Zamboanga.

Sivasubramaniam (1969) concluded that the apparent
abundance of S. orientalis around Sri Lanka was so low
that it was not worth an attempt to develop a fishery foi
this species. He noted that not more than a few hundred

Table 24. — Annual landings (in kilograms) of Sarda orientalis in foui
southern Japan ports. (Data provided by S. Kikawa, Far Seas Fish-
eries Research Laboratory, and T. Kato, Nansei Regional Fisheries
Research Laboratory, Japan.) (ND = No data.)

Landings

in kilograms

Year

Tosashimizu

Muroto

Murotomisaki

Aburatsu

Total

1967

37,462

1968

22,171

1969

30,024

1970

30,718

25,461

56,179

1971

256,939

42,595

31,732

331,266

1972

229,722

41,220

9,994

280,936

1973

299,621

13,152

11,294

8,426

332,493

1974

11,710

pounds were taken in 1964 and estimated that the annual
catch was about a ton in 1969.

Sarda sarda

There are commercial fisheries of varying sizes almost
throughout the entire distributional range of S. sarda in
the eastern and western Atlantic and in the Mediterra-
nean and Black Seas (Table 25).

Table 25.-

-Countries with fisheries for Sarda sarda.
from FAO 1974.)

(Compiled

Countries exploiting Sarda sarda in the eastern Atlantic:

Angola

Morocco

Bulgaria

Portugal

Equatorial Guinea

Romania

German Democratic Republic

Spain

Greece

U.S.S.R.

Countries exploiting Sarda sarda in the western Atlantic:

Argentina Mexico

Brazil United States

Grenada Venezuela
Martinique

Countries exploiting Sarda sarda in the Mediterranean and Black Seas:

Algeria Romania

Bulgaria Spain

Cyprus Tunisia

Greece Turkey

Italy U.S.S.R.

Malta Yugoslavia
Morocco

111!'

/ \ MEDITERRANEAN S

/ 1 BLACK SEA

1 / WESTERN
EASTERN / ATLANTIC
ATLANTIC 1/

L /

^Xr*'" ^-° — °^ /

1 1 1 1 1 1 1 1 1

1965 1966 1967 1968 1969 1970 1971 1972 1973

Figure 43. — Annual landings of Sarda sarda in the Atlantic and
Mediterranean-Black Seas. (Data from FAO 1974.)

Table 26. — Annual landings of Sarda sarda (thousands of metric tons). (Data from

FAO 1974.)

1965

1966

1967

1968

1969

1970

1971

1972

1973

Eastern Atlantic:

North

2.7

1.6

0.8

2.1

1.9

0.8

6.1

1.5

Central

1.9

3.0

1.3

1.2

1.9

0.9

0.5

9.0

South

1.3

1.6

1.7

0.4

0.9

1.2

0.7

0.6

0.5

Total

5.9

7.3

6.3

2.5

4.2

5.0

2.4

7.2

11.0

Western Atlantic:

North

0.1

0.0

().()

0.1

0.0

Central

0.3

0.4

0.5

0.8

0.7

1.0

1.2

1.8

South

].;>

2.0

4.6

3.8

5.4

7.6

6.1

5.7

4.0

Total

1.9

2.4

5.1

4.6

6.3

8.4

7.1

6.9

5.8

Mediterranean and Black Seas:

26.9

22.1

41.1

25.3

55.2

10.6

22.9

21.9

22.0

The annual landings of S. sarda in the eastern and
western Atlantic and in the Mediterranean -Black Seas
during the period from 1965 to 1973 show that except in
1973 when the eastern Atlantic landings exceeded 10,000
t, the landings in the eastern and western Atlantic never
rose above this figure (Fig. 43, Table 26). The fisheries of
Spain and Portugal make up the landings in the eastern
Atlantic. In the western Atlantic the bulk of the landings
is made by the fisheries of Argentina and Brazil. The
landings in the Mediterranean and Black Seas fluctu-
ated from 10,600 to 55,200 t during the period from 1965
to 1973. Turkey contributed 71.7 to 91.3% of the annual
landings during this period.

6 PROTECTION AND MANAGEMENT

6.1 Regulatory measures

There are no measures regulating the commercial fish-
eries for any of the species of Sarda. The State of Cali-
fornia imposes a bag limit of 10 S. c. lineolata per day for
recreational fishermen (Frey 1971), and is presently seek-
ing to impose regulations on the California commercial
fishery (MacCall see footnote 5).

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1963. Effects of water temperature on the distribution of some
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RAO, K. V. N.

1964. An account of the ripe ovaries of some Indian tunas. Proc.
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RODRIGUEZ-RODA, J.

1966. Estudio de la bacoreta, Euthynnus alleteratus (Raf.) bonito,
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ROSA, H., JR.

1965. Preparation of synopses on the biology of species of living
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SANZO, L.

1932. Uova e primi stadi larvali di Pelamys sarda C. V. R. Com.
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1954. Some aspects of the dynamics of populations important to
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SERBETIS, C.

1955. Migrations des pelamides et des maquereaux de la mer
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SERVENTY. D. L.

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1943. Biology of the Atlantic mackerel (Scomber scombrus) of
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1963. Synopsis of biological data on oriental bonito Sarda ori-
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[In Russ.

FISHERIES SYNOPSES

; series of documents, issued by FAO, CSIRO, INP, and NMFS, contains comprehensive reviews of present knowledge on
species and stocks of aquatic organisms of present or potential economic interest. The Fishery Resources and Environment
Division of FAO is responsible for the overall coordination of the series. The primary purpose of this series is to make existing
information readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention to gaps
in knowledge. It is hoped that synopses in this series will be useful to other scientists initiating investigations of the species con-
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Synopses in this series are compiled according to a standard outline described in Flb/S1 Rev. 1 (1965). FAO, CSIRO,
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already issued will also be most welcome. Comments on individual synopses and requests for information should be ad-
dressed to the coordinators and editors of the issuing organizations, and suggestions regarding the expansion or modification
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Consolidated lists of species or groups covered by synopses issued to date or in preparation will be issued from time to time.
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chased from National Technical Information Service, U.S. Department of Commerce, 5285 Port Royal Road, Springfield, VA
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The following synopses in this series have been issued since January 1977:

FIR/S 1 1 3 Synopsis of biological data on the perch, Perca fluviatilis and flavescens

NMFS/S 116 Synopsis of biological data on the red porgy, Pagrus pagrus (Linnaeus)
NMFS/S 117 Synopsis of biological data for the winter flounder, Pseudopleuronectes

americanus (Walbaum)
NMFS/S 123 Synopsis of biological data on the rock crab, Cancer irroratus Say
NMFS/S 122 Synopsis of biological data on tunas of the genus Euthynnus

December 1977
May 1978

November 1978

May 1979

October 1979

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