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:3 Hole, MA C2543 NOAA Technical Report NMFS Circular 436

Synopsis of

Biological Data on

Frigate Tuna, Auxis thazard,

and Bullet Tuna, A. rochei

n

January 1981

FAO Fisheries Synopsis No. 124 NMFS/S 124 SASJ -Auxis rochei: 1,75(01)023,03 Auxis thazard: 1,75(01)023.01

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415. A basis for classifying western Atlantic Sciaenidae (Teleostei: Perciformes). By Labbish Ning Chao. September 1978, v -i- 64 p., 41

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416. Ocean variability: Effects on U.S. marine fishery resources

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417. Guide to the identification of genera of the fish Order

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tables.
NOAA Technical Report NMFS Circular 436

Synopsis of

Biological Data on

Frigate Tuna, Auxis thazard,

and Bullet Tuna, A. rochei

Richard N. Uchlda

January 1981

Marine Biological Laboratory/ Woods Hole Oceanographic Institution

MAY 6 ms

Woods Hole, MA 02543

FAO Fisheries Synopsis No. 124

U.S. DEPARTMENT OF COMMERCE

Philip M. Klutznick, 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 .

Nomenclature 1 •1.11 Valid

1.12 Synonymy 1

1 .2 Taxonomy 1 1.21 Affinities 1

1 .22 Taxonomic status 4

1.23 Subspecies 4

1 .24 Standard common names, vernacular names 4

1.;? Morphology 5

1.31 External morphology 5

*1.32 Cytomorphology

1 .33 Protein specificity 7

2 Distribution 12

2.1 Total area 12

2.2 Differential distribution 13

2.21 Spawn, larvae, and juveniles 13

2.22 Adults 14

2.3 Determinants of distribution changes 14

2.4 Hybridization 15

2.41 Hybrids; frequency of hybridization, species with which hybridization occurs; methods of hybridization 15

*2.42 Influence of natural hybridization in ecology and morphology

3 Bionomics and life history 15

3.1 Reproduction 15

3.11 Sexuality 15

3.12 Maturity 15

3.13 Mating 16

3.14 Fertilization 16

3.15 Gonads 16

3.16 Spawning 16

3.17 Spawn 22

3.2 Preadult phase 26

3.21 Embryonic phase 26

3.22 Larvae phase 26

3.23 Adolescent phase 27

3.3 Adult phase 31

*3.31 Longevity

3.32 Hardiness 31 3.33 Competitors 31

3.34 Predators 32

3.35 Parasites, diseases, injuries, and abnormalities 33

3.4 Nutrition and growth 34

3.41 Feeding 34

3.42 Food 34

3.43 Growth rate 36

3.44 Metabolism 37

3.5 Behavior 38

3.51 Migrations and local movements 38

3..52 Schooling 38

3.53 Responses to stimuli 39

4 Population 41

4.1 Structure 41

4.11 Sex ratio 41

*4.12 Age composition

4.13 Size com|X)sition 43
4.2 Abundance and density 45

4.21 Average abundance 45

*4.22 Changes in abundance

*4.23 Average density

*4.24 Changes in density

4.3 Natality and recruitment 48

4.31 Reproduction rates 48

*4.32 Factors affecting reproduction

4.33 Recruitment 48

4.4 Mortality and morbidity 48

*4.41 Mortality rates 4.42 Factors causing or affecting mortality 48

*4.43 Factors affecting morbidity

*4.44 Relation of morbidity to mortality rates

*4.5 Dynamics of population

*4.6 The population in the community and the ecosystem 5 Exploitation 48

5.1 Fishing equipment 48

5.11 Gears 48

5.12 Boats 50

5.2 Fishing areas 51

5.21 General geographic distribution 51

5.22 Geographic ranges 51

5.23 Depth ranges 51

*5.24 Conditions of the grounds

5.3 Fishing seasons 52

5.31 General pattern of seasons 52

5.32 Dates of beginning, peak, and end of season 52

5.33 Variation in date or duration of season 52

5.4 Fishing operation and results 53

5.41 Effort and intensity 53

5.42 Selectivity 53

5.43 Catches 54

Literature cited • • • 57

•No inlormation available.
Synopsis of Biological Data On

Frigate Tuna, Auxis thazard, and Bullet Tuna, A. rochei

Richard N. Uchida

ABSTRACT

This s>'nopsis of biological and technical data on frigate tuna. Auxis (hazard, and bullet tuna, A.

rochei, includes information on identity, distribution, bionomics, life history, population, and exploitation. Over 200 published and unpublished reports, up to and including those published in 1978. are covered. INTRODUCTION

Unlike a number of tuna species in the world's oceans which are heavily exploited and possibly are being

harvested near the upper limit of rational utilization, frigate mackerels (or frigate and bullet tunas; see Klawe

1977), the most primitive genus of the higher tunas

(tribe Thunnini), give every indication of being an

underutilized fishery resource. In planning for the ra- tional utilization of this resource, it is imperative to have on hand information concerning the biology of the

species and estimates of present and potential catches. The purposes of this paper, therefore, are to update our

current knowledge of the species by bringing together and abstracting all the available information on the biology and fisheries for frigate and bullet tunas and to review and evaluate in depth the results of past

research, indicating in particular where conflicting evidence exists.

Scomber thazard Lacepede 1802 (original description;

off coast of New Guinea) Scomber bisus Rafinesque 1810 (original description; Palermo) Thynnus rocheanus Risso 1826 (original description; Nice) Auxis taso Cuvier and Valenciennes 1831 (original description; New Guinea) Auxis vulgaris Cuvier and Valenciennes 1831 (original description; Mediterranean) Auxis tapeinosoma Bleeker 1854 (original description; Japan) Auxis thynnoides Bleeker 1855 (original description; Ternate) Auxis rochei. Giinther 1860 Auxis thazard. Jordan and Evermann 1896 Auxis hira Kishinouye 1915 (original description; Japan)

Auxis nuiru Kishinouye 1915 (original description; Japan)

Au.vis bisus. Cadenat 1950

1 IDENTITY

1.1 Nomenclature

1.2 Taxonomy

1.21 Affinities

Auxis thazard (Lacepede) 1802 Auxis rochei (Risso) 1810

1.12 Synonymy

It it not possible at this time to assign all the various names that have appeared in the literature to either thazard or rochei. The following names are from Rosa

(1950), de Beaufort and Chapman (1951), Collingnon

(I960),' Williams (1963),' Idyll and de Sylva (1963), and

Jones (1963) and are listed in chronological order.

'Southwest Fisheries Center. National Marine Fisheries Service. NOAA. P.O. Box 38.30. Honolulu, HI 96812.

"Collingnon. .J 1960. Report on /4uiis t/iazard in the eastern Atlantic. llnFr.l CCTA Symposium on Thunnidae. Dakar. 12-27 December

1960. a p. IMimeogr.) (Engl, transl. available Southwest Fish. Cent.. Natl. Mar. Fish. Serv.. NOAA. Honolulu, HI 96812.) 'Williams. F. 1960. A symposium of existing Itnowledge on the

fishes on the genus Auxis Cuvier, 1829 in the Indian Ocean. CCTA

Kingdom Animalia Phylum Chordata Subphylum Vertebrata

Superclass Gnathostomata

Class Osteichthyes Subclass Actinopterygii

Division Teleostei Cohort Acanthopterygii Order Perciformes Suborder Scombroidei Family Scombridae

Subfamily Scombrinae

Tribe Tunnini

{Thunnus, Katsuwonus,

Euthynnus, Auxis)

Symposium on Thunnidae. Dakar, 12-27 December 1960, 13 (Mimeogr.l
Genus Auxis Cuvier 1829 (type-species: Scomber

rochei Risso) by subsequent selection by Gill (1862). The description of the genus Auxis under "Les Scombres" first appeared in Cuvier's Regne Animal in 1829. From the time of Cuvier's work to the present, however, the genus Auxis has had several reclassifications. For example, in classifying the genus Auxis, Kishi- nouye (1915) first included it in the family Thunnidae.

However, in a later work (1917), he placed Thunnidae

and Katsuwonidae in a new order called Plecostei and

placed the families Scombridae and Cybiidae in the

order Teleostei. The genus Thunnus fell under Thunnidae and the genera Katsuwonus. Euthynnus. and

Auxis came under Katsuwonidae (Kishinouye 1923). The primary characteristic subcutaneous blood vessels; a secondary characteristic of Plecostei was the presence of well-developed subcutaneous blood vessels: a secondary

characteristic was the development of dark red lateral tissues in relation to the subcutaneous blood vessels. Many scientists disagreed with Kishinouye's new

order and its subdivision. Takahashi (1924) argued that

Plecostei was established only on partial differences in the highly variable vascular system and cannot exist on

an equal status with the other four orders of Teleostomi. Jordan (1923) and Herre (1953) placed the scombroid fishes in two families—Scombridae and

Thunnidae. Fraser-Brunner (19.50), on the other hand,

rejected any division of the family Scombridae arguing

that attempts to subdivide this family "have resulted in arrangements which are artificial and have left the clas- sification in an uneasy, shifting state.'" de Sylva

(1955), Collette and Gibbs (1963b), and most other re- cent workers recognize a single family Scombridae with

various subdivisions. Collette and Chao (1975) placed Auxis in the tribe Thunnini of the subfamily Scombrinae.The following description of the genus Auxis is from Jordan and Evermann (1905): "Body oblong, plump,

most naked posteriorly, anteriorly covered with small

scales, those of the pectoral region enlarged, forming a

corselet; snout very short, conical, scarcely compressed; mouth rather small, the jaws equal; teeth very small, mostly in a single series, on the jaws only; tail very

slender, depressed, with a rather large keel on each side;

first dorsal short, separated from the second by a consi- derable interspace; second dorsal and anal small, each

with 7 or 8 finlets; pectorals and ventrals small; no air- bladder; branchiostegals 7; pyloric coeca dentritical; gill- rakers very long and slender, numerous; vertebrae 39 in number, peculiarly modified . . . ." Auxis is the most primitive genus among the higher tunas that have developed a prootic pit and a partial subcutaneous circulatory system (Collette and Gibbs

1963b). All the Thunnini, except Auxis. have a common

cutaneous artery that divides into dorsal and ventral branches lateral to the aorta; in Auxis, however, the dorsal and ventral branches originate separately with the

latter being ver>' poorly developed (Collette 1978). The

haemal spines of the thoracic vertebrae do not form a haemal arch and the first vertebra is not sutured to the

cranium as in the higher members of Thunnini. Also, compared with Euthynnus. Katsuwonus, and Thunnus,

Auxis lacks the frontoparietal fenestra, which is an additional pair of openings present in the cranium and has

a lateral countercurrent system for heat exchange that is not as well developed phylogenetically (Collette 1978).

All members of the tribe Thunnini have a swim bladder

as juveniles; however, the bladder degenerate with growth in Auxis, Euthynnus, and Katsuwonus.

Several other characters distinguish members of the genus Auxis. the smallest of the higher tunas, from other scombrids (Collette and Gibbs 1963b; Fitch and Roedel

1963; Williams 1963). In Auxis, there is a single interpelvic process which is between and about as long as the pel- vic fins. In other tunas, but not in other scombrids, the

interpelvic process is bifurcate and much less than half the length of the pelvic fins (it is single but small in (Irammatorcynus and Gymnosarda). The number of dorsal and anal finlets (seven or eight) distinguishes Auxis from .Scomber, which has only five of each. Auxis can

also be distinguished from Rastrelliger by the lack of a

corselet and a body entirely covered with moderate-sized

scales in the latter.

Species — Auxis thazard (Lacepede) 1802

The following description of A. thazard (Fig. 1, uppe^r photo) is from Williams (1963). Major morphological

features described under the genus are omitted from the account of the species.

"Depth 3.9 to 4.5, head 3.2 to 3.8 in standard length. Eye 5.0 to 5.85 in head, 1.25 to 1.66 in snout and 1.25 to

1.7 in the flatly rounded interorbital space. Snout 3.62 to 4 in head. Maxilla reaches to a point under the anterior

half of the eye and is 3 in head. Single row of small

pointed teeth in each jaw, none on palate. Jaws almost

equal. First and second dorsal spines subequal, equal to snout and eye; following spines rapidly decreasing in

size, eighth usually shorter than the pupil. Second dorsal

fin very low, about three times its base distant from first dorsal; first ray of second dorsal about 5 in head. Anal

similar to second dorsal, first ray about 5.2 in head. Pectorals short, roughly triangular, about 2 in head and

shorter than postorbital; origin of pectoral before that of

first dorsal. Pelvics thoracic, about 2.5 in head, origin somewhat behind that of pectorals. Caudal lunate, upper

lobe about 1.8 in head. Body naked except for the corselet of scales anteriorly. Rear margin of the corselet runs from base of second dorsal to above end of pectoral; thence there is a posterior prolongation of the corselet along the lateral line; below the pectoral tips the corselet margin curves to above the pelvic base from where it turns posteriorly and finishes well behind the tips of the

pelvics. The prolongation of the corselet along the lateral

line tapers abruptly between first and second dorsal fins, and under the origin of the second dorsal is not more than

4 irregular scale rows wide. Scales large and imbricated above pectoral base. Gill rakers about 1.75 in length of

gill filaments."
Figure 1.—Auxia t/uuard (upper photo) from the eastern Pacific, collected at Morgan Bank, Baja California, and A. rochei (lower

photo) from near Santa Catalina Island. Calif. (Fitch and Roedel 196.'!)

Species — Auxis rochei (Risso) 1810 The description of A. rochei (Fig. 1, lower photo) is from Jones (1958), who originally referred to the specimen as A. tapeinosoma.

"Body robust, rounded, almost circular in cross- section. Dorsal outline moderately tuid evenly curved. Ventral outline evenly curved when fresh but slightly flattened abdominally after preservation in formalin. "Height 5.38 in sttmdard and 5.55 in furcal [= fork] length. Head 3.77 in standard and 3.88 in furcal length. Snout, pointed 3.9 in the head, longer than eye diameter. Eye 4.91 in the head, 1.27 in the snout, 1.27 in the almost flattened interorbital space. Mouth moderate,

oblique, end of maxillary reaching vertical from anterior margin of eye. Jaws nearly equal, the lower jaw pro- jecting almost imperceptibly beyond the upper. Teeth

small, pointed in a single row on both jaws, none on

palate. Branchiostegals 7. Gill rakers long and slender, 45 in first gill arch. "Two dorsal fins separated by interspace slightly shorter than head length. First dorsal roughly triangular with 10 spines, anterior spine longest. Second dorsal small with 13 rays. Dorsal finlets 8. Anal fin small, with

2 spines, 11 rays. Anal finlets 7. Pectorals roughly tri- angular, reaching vertical from the base of first ray of

first dorsal. Ventral thoracic, axillairy scales equal in length to ventrals. "Body naked except for the corselet of scales which

taper gradually to 9-10 irregular scale rows at vertical through second dorsal and end as a narrow line at vertical below second dorsal finlet. Scales large and imbri- cated above pectoral base. Caudal peduncle slender with

feebly developed lateral keels. Lateral line somewhat undulating and without a distinct arch."
1.22 Taxonomic status A specimen of frigate tuna, collected by Commerson

off New Guinea in 1768, was first described by Lacepede

in 1802 as Scomber thazard. In 1810, Risso and Rafinesque, working independently, named the Mediterranean form Scomber rochei and Scomber bisus, respectively.

Gill (1862) subsequently designated rochei as the typespecies ioiAuxis Cuvier. Systematists generally followed Giinther (1860) and Jordan and Gilbert (1882) in using

Risso's rochei instead of Rafinesque's bisus, but no one

has indicated which name was described first. Further- more, the International Commission of Zoological Nomenclature has never ruled as to which name, rochei

or bisus, is valid (Fitch and Roedel 1963). Over the years, other names appeared in the literature. These include Thynnus rocheanus Risso, 1826 (Mediterranean), A. vulgaris Cuvier and Valenciennes, 1831 (Mediterranean), A. taso Cuvier and Valenciennes, 1831 (New Guinea), A. tapeinosoma Bleeker, 1854 (Japan), and A. thynnoides Bleeker, 1855 (Temate). In 1915, Kishinouye described two additional names. The species he named hira had a short corselet which ended slightly posterior to the pectoral fin. The other he named maru

had a long corselet which extended to the anal fin. Kishi- nouye stated that the maru is probably the same species

as thazard, but could not say whether thazard corre- sponded to hira or maru. Furthermore, he believed that

Bleeker's (1854) tapeinosoma could be maru, but the

figure and description of this species were unclear and he was unable to make a positive identification. Therefore, Kishinouye described them as new species, hira and

maru.

Fraser-Brunner (1950) disagreed with Kishinouye and

recognized only a single, worldwide species, thazard, but

others, such as Wade (1949), Cadenat (1950), and Jones

(1958), recognized two species of Auxis. Wade (1949) used Bleeker's name tapeinosoma for a long-corseletted

pacific form. Matsumoto (1959, 1960a), on the other hand, finding the nomenclature of the long-corseletted form confused, resurrected the name thynnoides. He

argued that tapeinosoma of Wade (1949) and Herre and

Herald (1951) appeared to be a misnomer because it is actually a short-corseletted form. Furthermore, Matsu- moto regarded hira and maru of Kishinouye and tape- inosoma of Bleeker as synonyms of the short-corseletted, worldwide thazard.

1.23 Subspecies

None.

1.24 The standard common names, vernacular names

The standard common names and vernacular names of A. thazard were abstracted from the lists published in Fiedler (1945), Rosa (1950), Collingnon (1960—see foot- note 2), FAO (1960, 1976), Idyll and de Sylva (1963), Uchida (1963), Williams (1963), Instituto del mar del

Peru (1971^), Miyake and Hayasi (1972), Klawe (1977), and Roberts et al. (1977).

Country

Algeria

Common and vernacular names

Auxide, Scunno, Bisu, Melva, Mel-
Morocco New Zealand Norway

Papua New

Guinea

Peru

Phillipine Is- lands

Portugal

Sarawak

Seychelles Somalia (Mijurtein coast) South Africa Spain

Spanish Morocco

Sri Lanka

(Ceylon) Sweden

Taiwan

Thailand Turkey

Union of South

Africa Union of Soviet

Socialist Republic United Kingdom

Unites States

Venezuela

Yugoslavia

International organization

Food and Agri- Frigate tuna, Auxides, Melvas

culture Organization of the United Nations (FAO)

International Frigate tuna, Auxide, Melva

Commission

for the Conservation of Atlantic Tunas (ICCAT)

Jones (1963), Uchida (1963), Instituto del Mar del Peru (see footnote 4), FAO (1976), and Klawe (1977) pro- vided the following common and vernacular names of A.

rochei = A. tapeinosoma = A. thynnoides = A. maru.

Country Common and vernacular names

Australia Long corseletted frigate mackerel, Maru frigate mackerel

Melva

Frigate tuna

Auxis Deho

Melva, Macarela

Frigate mackerel, Tunungan, Manko, Mangki) (Marinao, Samal, Vis- aya. and Tao Sug) Gayado, Judea, Serra, Cachorra, Benito Tongkol

Bonite folle Tubani (Somali)

Frigate mackerel

Melva, Visol, Melvara, Bis, Benito

del Norte, Macaela Melva

Frigate mackerel, Rogodwa

(Sinhalese), Alagoduwa

Auxide

Chien yu PlaO

(jobene

Frigate mackerel. Bee hoo

Auksida, Makrelevyi tunets

Plain bonito

Frigate mackerel. Bullet mackerel. Boo hoo, Frigate tuna Cabana negra Trupac, Tunjcic, Rumbac

Ecuador

India: Malayalam

(North)

Malayalam

(South)

Tamil

Japan

Peru Union of Soviet

Socialist Republic United States

International organization FAO

ICCAT

Betellita

Kuttichoera (means small tuna and

generally applied to A. thazard and young of £. affinis) Urulan-choora (means rounded tuna and applied to A. thazard also from which this species is not generally distinguished)

Eli-choorai (means ratlike tuna), Kutteli-choorai (means small ratlike tuna) Marusoda, Marusodakatsuo. Marugatsuwo, Marumejika, Magatsuwo,

Manba, Mandara, Chiboh, Dainanpo, Nodoguro, Rohsoku, Subota, Uzawa. Maiika, Soku, Soda, Subo. (Marumedika, Marugatsuo, Magatsuo, and Dainanbo are variations

in spelling of some of the above

names.) Other names mentioned by Rosa (1950) are Kogatsuo, Kukarai, Kobukura

Melva, Fragata Skumbrievvi tunets, Auksida

Bullet mackerel. Bullet tuna

Bullet tuna

Bullet tuna To avoid confusion throughout the remainder of this paper, A. rochei will be used instead of A. tapeinosoma,

A. thynnoides, or A. maru to describe the long-corseletted form. For the short-corseletted form, A. thazard

will be used, but there is considerable confusion in the

literature because several authors used thazard in the

belief that there was only a single worldwide sijecies of Auxis (Fraser-Brunner 1950; Rivas 1951). This usage must be translated into Auxis spp. where reference is made to the Pacific forms and into A. rochei in the Atlantic based on Fitch and Roedel's (1963) interpretation or Auxis spp. based on Richards and Randall's (1967) docu- mentation of A. thazard in the Atlantic.

1.3 Morphology

1.31 External morphology The fin ray counts, together with the gill raker and

vertebral counts, are given for A. thazard and A. rochei

in Tables 1 and 2, respectively. It should be noted that some investigators did not separate the two species of Auxis at the time the meristic counts were made and,

therefore, their counts have not been included. One of the sources of confusion in identifying Auxis to
Table 1.—Meristic characters of the short-corseletted frigate tuna, AuxU thazard, by various investigators.
<^

k

(Q)

Figure 2.—Outlines of corselets drawn from actual specimens

(Jones 1963). (a) Auxia rochei; (b) A. thnzard - typical short- corseletted condition; (c) A. thazard - intermediate condition.

Among internal characters that have been a source of confusion is the gill raker count. Wade (1949) and Herre and Herald (1951) pointed out that A. thazard in the western Pacific normally have fewer gill rakers than A.

rochei. And A. thazard taken in the eastern Pacific and

the Indian Ocean also have counts that are definitely lower (Mead 1951; Jones 1963). Godsil (1954), on the

other hand, reported that all the A. thazard taken off Baja California and the Galapagos Islands have high gill raker counts similar to those reported for A. rochei by

Wade (1949) and Herre and Herald (1951). Matsumoto

(1959), however, believed that the number of gill rakers, by itself, is not a reliable character in identifying the two

species of Auxis. But Jones and Silas (1964) suggested

that the gill raker counts could be useful; in case of doubtful identification from external characters, a combination of gill raker counts and corselet width should

facilitate specific identity. Jones and Silas (1964) have used body cross section as an aid to identification but CoUette and Gibbs (1963a) have warned that this character is difficult to use (Fig.

3). Also suggested by Jones and Silas was the position of the visceral organs (Fig. 4). Godsil (1954) and Yoshida and Nakamura (1965) noted that other prominant dif- ferences between the two species occur in the skeletal structure. In A. thazard, the temporal crests diverge anteriorly so that they are not parallel to one another (Fig.

5) and the width of the skull is wider in relation to body

length (Fig. 6). Yoshida and Nakamura also noted that the length of the anterior branch of the haemal processes was longer and touching on the preceding arch of the 24th to the 28th vertebrae in A. thazard (Fig. 7).

1.33 Protein specificity

Taniguchi and Nakamura (1970) examined muscle

protein of A. thazard and A. rochei by the cellulose acetate electrophoretic method to determine whether

specific divergence occurred between species. They

found five components in the electropherograms of both

species, but some components were not common to both. The genus Auxis, they concluded, contains two distinct species based on external and internal morphological

characters although they are closely related. To analyze muscle protein polymorphism in Auxis col- lected from the coastal region of Kochi Prefecture, Japan, Taniguchi and Konishi (1971) used starch-gel electrophoresis and detected differences in protein

specificity between A. thazard and A. rochei. They con- cluded that whereas no individual variation could be rec- ognized in electropherograms of 11 specimens of A.

Table 4. —Comparison of gill raker counts for 84 Auxis thazard from three geographical

localities (Fitch and Roedel 1963).
Table 5. —Selected measurements o{ AuxU rochei from five geographic localities (Fitch and Roedel 1963).

u ^

q; PI E EE

"

.S c

3 _

o

sc

10
Figure 5.—JJoraal siev of §kull ofAuxis rochei (ca. 2" cm SL) on the left and A. thazard (ca. .')0 cm SI.) on the right. .Vote that the temporal crests of the A. rochei

skull are parallel with each other and with the supraoccipital crest, whereas in A. thazard the temporal crests diverge anteriorly so that the three crests are not

distinctly parallel (Yoshida and Nakamura 1965).

-
2 DISTRIBUTION

2.1 Total area

The genus Auxis is distributed worldwide in tropical and subtropical waters.. The confusion surrounding the

identification of the two species of Auxis is reflected in their reported distribution in the world's oceans. Fitch and Roedel (1963) concluded from their study that A.

rochei was cosmopolitan in distribution whereas A. thazard was restricted to the Pacific. Collette and Gibbs

(1963a) expressed uncertainty about the presence of both species of Auxis in the Atlantic but stated that both probably do occur there. Richards and Randall

(1967) independently confirmed that adult A. thazard do occur in the Atlantic. The distribution of A. thazard cannot be separated from that oi A. rochei at the present time because of dif- ficulties in the past in distinguishing one species from

the other (Yabe et al. 1963). Therefore, the reported dis- tribution in the literature for either A. thazard or A. rochei is questionable and needs to be critically reex- amined. The following discussion takes into account the

distribution of both species. Figure 8 shows the distri- bution of Auxis adults in relation to water tmd land

areas. In the Pacific Ocean, Auxis occur off the coast of the United States between Santa Catalina Islcmd and San

Clemente Island off California southward into the east- em tropical Pacific extending as far south as lat. 18° S

(Radovich 1961; A. Ch. de Vildoso^). They have also been reported from the Hawaiian (Matsumoto 1960a) and Marquesas Islands (Nakamura and Matsumoto

1967). In the western Pacific, they occur off the coast of Japan as far north as Hokkaido, off Korea, off the coast

^A. Ch. de Vildoso. Institute del Mar. Calla December 1975.

Peru. pers. coramun.

of China mainland near southern Manchuria and Ningpo, around Formosa, and in waters surrounding the Ryukyu and Bonin Islands (Rosa 1950). Southward,

Auxis have been reported from Samoa Islands and Papua New Guinea (Jordan and Scale 1906), the Philip- pine Islands (Herre 1953), along the eastern and southern coasts of Australia (Scott 1962; Laevastu tuid Rosa

1963; Whitley 1964), Tasmania (Lord 1927), and New

Zealand (Roberts et al. 1977). The usual latitudinal range reported for Auxis in the

tropical and subtropical waters of the Atlantic Ocean is from lat. 45°N to 35°S (CoUignon see footnote 2; Miyake

and Hayasi 1972). In the eastern Atlantic, they occur

infrequently as far north as Bergen, Norway, and the canal of Oslo Harbor (Rosa 1950) and in waters around

the British Isles (Went 1955, 1956, 1958, 1967; Rae 1963; Went and Kennedy 1969; Wheeler and Blacker 1969). Auxis also occur in the Mediterranean and Black Seas. Southward, they are found in waters off the Republic of South Africa and offshore around Ascension and St. Helena Islands. In the western Atlantic, the northern- most occurrence o{ Auxis is off Barnstable, Mass., in the Gulf of Maine (Mather and Gibbs 1957). Southward in the western Atlantic, Auxis have been reported from

the Gulf of Mexico, the Caribbean Sea, and the Atlan

tic as far south as Mar del Plata, Argentina (Lopez

1961).

In the eastern Indian Oceam, Auxis have been

reported from the east coast of India and southward

along the Indonesian Archipelago to Cape Leeuwin near

the southern tip of Western Australia (Rosa 1950; Jones and Silas 1964; Nair et al. 1970). The western Indian Ocean distribution of Auxis extends from the west coast

of India offshore to the Maldive and Laccadive Islands, the coast of Iraq in the Persian Gulf (Mahdi 1971), the Red Sea (Ben-Tuvia 1968), and from the Gulf of Aden

southward to the coast of Natal in the Republic of South

Africa (Williams 1963; Nair et al. 1970).

Figure 8.—The distribution of Auxis spp. adults in relation to water and land areas in the world's ocean.

12
2.2 Differential distribution

2.21 Spawn, larvae, and juveniles The larvae ofAuxis, like the eggs which precede them,

are displaced from the area of spawning due to drift of the ocean currents (Matsumoto 1958, 1959). Except in a few areas where currents are swift, the actual displace- ment of eggs and larvae appears to be relatively insig- nificant. Whereas the distribution of adult Auxis is usually as- sociated with land masses, that of the larvae has been

described as not only coastal but also oceanic. Figure 9 shows the localities of capture of Auxis larvae in the

world's oceans. The actual differences in larval and

adult distributions, however, may not be real. The

adults are usually reported to occur in coastal waters because most of the fishing is done there. But plankton

hauls conducted in waters far from land masses have shown that larval Auxis occur in oceanic as well as

coastal waters. Matsumoto (1958, 1959) suggested that

the localities where larvae of about 3 mm occur prob- ably represent actual spawning sites; therefore, it can be

expected that adult Auxis also occur in the oceanic

regions of the world's oceans. Watanabe (1964), on the

other hand, studying tuna and billfish stomach contents, concluded that Auxis are coastal dwellers. The distribution of juvenile Auxis, 10-20 cm SL (stan- dard length), agrees well with that of the larvae, es- pecially near land masses, but in the oceanic regions the presence of juveniles have not been well documented

(Fig. 10). Yabe et al. (1963) pointed out that the usual method of using a midwater trawl to collect juveniles has not been successful quantitatively; rather, more in- formation can be obtained through examination of stomach contents of large tunas and billfishes. That juvenile Auxis occur mostly in waters close to land masses is brought out in Table 7 which shows the number of juvenile Auxis taken by midwater trawl in Hawaiian waters. Higgins (1970) observed that in July- September 1967, the catch rate of juvenile Auxis

reached 3.5 individuals/tow, the highest among all the

20* <0* 60* 90* 100* 120*

20* 40* eo* 120* MO- teo* lao* i6o* iw 120*

Figure 9.—Localities of capture of larval Auxia (Yabe et al. 1963).

Figure 10.—Localities of capture of juvenile (10-20 cm SL) Auxis (Yabe et al. 1963).

13
Table 7. —Total number of juvenile Auxis and (in parentheses) the average number per tow collected by a 12 m wide by 8 m high mid- water trawl during the day (1200-1800), night (2000-0200). and morning (0400-1000), RV Townsend Cromwell cruise 32, 12 July-25 September 1967. The trawl, fished at a depth of 100 m during deep

hauls and 20 m during shallow hauls, was towed at a speed of 1.5 m/s (Higgins 1970). Area and
general. Salinity, likewise, showed no relationship with

catches of larval Auxis, but Klawe et al. (1970) reasoned

that this lack of relationship may be due to the narrow

range of salinity encountered during the period that the study was made. Salinity, therefore, cannot be dis- counted as a factor affecting distribution. In fact, Richards and Simmons (1971) determined that larval Auxi's were found in a relatively narrow salinity range from 33.27.. to 35.47~ compared to that of other tunas, with the exception of little tunny, Euthynnus alletteratus. Concerning the effects of light intensity, Higgins

(1970) observed that juvenile Auxis were more

vulnerable to the midwater trawl at night than during

daylight hours. Furthermore, they were more readily caught in shallower than in deeper tows, although the data suggested that they may occur as deep as 100 m.

Strasburg (1960) reported that larval Auxis occur at the

surface very infrequently during daylight hours but are often found there in large numbers at night. Noting that

this pattern of an increase in catch at night could result from either net avoidance or vertical migration, Stras- burg reasoned that if net avoidance only were involved, one would expect the catch to be essentially constant at

night. The results of his study proved otherwise; therefore, he concluded that vertical migration appears to be

the major factor causing the increase in surface catch at night. Contrarily, in a later study by Klawe et al. (1970),

it was observed that larval Auxis show no significant

diel movement in the water column. From a statistical te^t they conducted for time of day as well as for inter- action between time of day and type of tow, Klawe et al. (1970) found no interaction between type of tow and

time of day; therefore, they concluded that no vertical movement had occurred. They reasoned that any ability larval Auxis may have had to avoid the sampling gear was independent of time of day.

2.4 Hybridization

2.41 Hybrids; frequency of hybridication; species with which hybridization occurs; methods of hybridization

There has been considerable progress in studies on egg and larval development of tuna and tunalike fishes in recent years. Most impressive are the experiments by

Japanese scientists on artificial fertilization of eggs and

the subsequent rearing of the larvae. Among the tuna

species that have been used in these experiments were

yellowfin, skipjack, frigate, and bullet. In 1970, the Japan Fisheries Agency coordinated a 3-vr cooperative program to artificially fertilize and rear tunas (Ueyanagi

et al. 1973). The experiments were conducted through

the cooperative efforts of Tokai University, Kinki

University, Shizuoka Prefectural Fisheries Experimental Station, Mie Prefectural Owase Fisheries Experi- mental Station, Nagasaki Prefectural Fisheries Experi- mental Station, and the Far Seas Fisheries Research

Laboratory of the Japan Fisheries Agency. Experiments conducted at Kinki University Fisheries Experimental Station (1974) involved crossfertilization of A. thazard and A. rochei. The ripe eggs taken from A. thazard caught by set net at Kashino, Oshima, on 17 July 1973 at 0530 were fertilized and

transported to the experimental station at Shirahama. At 1350 on the same day, there were about 35,120 viable eggs which averaged 0.86 mm in diameter and about

7,(XX) dead eggs. Twenty-five thousand viable eggs were

placed in a 3-ton (3,000 liter) tank and 10,000 in a 1-ton

(1,(XX) liter) tank. Water temperature in the tanks was

about 26°C. Larvae started to emerge the next day, approximately 30 h after fertilization and all hatching was completed during the day. The larvae were kept in seawater with Chlorella which had a density of

400,000/cc. Aeration was also provided. Rotifers were added to the tank 2 days after hatching and marine

plankton was added the next day. All the larvae in the 3- ton tank died after 6 days. In the 1-ton tank, although

mortality was heavy, there were some survivors. At 14 days, the water was circulated, and at 18 days after hatching, artemia and eggs of frozen yellowtail, Seriola quinqueradiata, were used to supplement the marine

plankton. Mortality continued and the last survivor, which attained a length of 4.8 cm and a weight of 0.907 g died after 31 days.

3 BIONOMICS AND LIFE HISTORY

3.1 Reproduction

3.11 Sexuality

Like all other scombrids, Auxis are heterosexual. There are no externally visible characters that aid in distinguishing males from females. Internally, the

paired and elongated gonads are nearly symmetrical and are suspended by mesenteries extending almost the

entire length along the roof of the abdominal cavity. At

the posterior end of the abdominal cavity, the gonads

extend along both sides of the anal fin. This posterior extension, according to Kishinouye (1923), is due to the narrowness of the abdominal cavity.

3.12 Maturity

Studies of gonads of troll-caught fish from waters of Kaneohe Bay, Oahu, in the Hawaiian Islands indicate that frigate tuna are nearly mature at a size of about 35 cm (Tester and Nakamura 1957), but in Japanese

waters, they have been reported to reach maturity at about 29 cm. Yasui (1975) investigated the use of liver weight as a possible index of sexual maturity. He plotted

liver weight against body length of frigate tuna caught

off the Izu Islands and off Mera, Shizuoka Prefecture, and observed a point of discontinuity at a body length of 29 cm. His data also showed that 97% of the fish <29 cm

were caught after September whereas 95% of those

larger than that were caught before August. The repro- ductive index, according to Yasui, was highest in July.

15
Observing that almost all fish >29 cm had high repro- ductive indices, Yasui hypothesized that liver weight is related to oogenesis and that it can be used as an indi- cator of sexual maturity.

In the Spanish trap net fishery located in and around

the Strait of Gibraltar, the size of A. rochei at first spawning is 35 cm for the females and 36.5 cm for the males (Rodriguez-Roda 1966). Rodriguez-Roda classi- fied the testes and ovaries according to the develop- mental stages as follows:

I Immature n Maturing

in Ripening

rV Prespawning V Spawning

VI Spent

Rodriguez-Roda noted that in May, a large proportion

of the males and females were in stage III. In June- August some showed development in stage FV, but by

September about one-third of the males and one-fourth

of the females sampled had gonads that apf)eared to be

spent (Table 8). Rodriguez-Roda noted that gonad clas- sification based on gross characteristics such as he used

could lead to misleading conclusions; however, he indi- cated that the relative gonad weight (gonad weight in relation to body weight) tended to confirm his conclusions.

125 g, respectively. According to Rao, the smaller of the two fish may "have been captured while in the act of spawning since most of the ripe ova were already lost by

the time it was examined."

3.16 Spawning

Earlier, it was pointed out that although the planktonic eggs and larvae of Auxis become displaced from

the area of spawning due to ocean currents, the dis- placement is insignificant. Therefore, localities where

larvae of 3 mm or less occur probably represent actual spawning sites. In the eastern tropical Pacific, Auxis larvae are the most abundant among all the scombrid larvae collected. Ahlstrom (1971) identified 1,563 out of 1,919 scombrid

larvae in the EASTROPAC collection as Auxia spp. Based on this information and the distribution of the

catches of larval Auxis, Klawe (1963) observed that off Baja California, spawning occurred in coastal waters whereas to the south it appeared to be in more oceanic

waters away from continents or islands. The most northerly area of spawning appeared to be near Cedros Island and at the head of the Gulf of California. To the south, Klawe found the limit of spawning to be off Point Santa

Elena in Ecuador. He delimited the general area of

Table 8. —Percentages of male and female Auxis rochei from the Spanish trap net fishery classified as I

- immature, II - maturing, HI - ripening, IV' - prespawning, V - spawning, and VI - spent in May-September

in 1958, 1961, 1963, and 1964 combined (Rodriguez-Roda 1966).
Figure 12.— Relative abundance (number of larvae per 1,000 m^ of water strained at the surface) of Auxis in May-October, based on plankton collections made in the eastern tropical Pacific. Dashed lines enclose area under investigation by Inter-American

Tropical Tuna Commission in 1952-59; black dots indicate positions of plankton hauls vrith zero catches. (Klawe 1963.)

Table 9. —Average number of Auxu larvae caught per hour during

different months off Cape Blanco, Costa Kica, with a 1 m (silk or nylon) net towed at 1.0 m/x (Klawe 196.3). Month
Figure 13.— Relative abundance (number of larvae per 1,000 m' of water strained at the surface) of i4iu;i» in November-April, baaed on plankton collections made in the eastern tropical Pacific. Dashed lines enclose area under investigation by Inter-American

Tropical Tuna Commission in 1952-59; black dots indicate positions of plankton hauls with zero catches. (Klawe 1963.)

OCT. NOV. DEC. JAN. FEB. APR AUG.

Figure 14. —Average monthly catches, log,, (number caught =1), of larva Auxia during cruises in October- February, April, and

August to the entrance of the Gulf of California (Klawe et al. 1970).

Aaxis have also been reported to spawn in waters around the Marquesas Islands (Nakamura and Matsu- moto 1967). From samples of tuna larvae collected at the

diel variability station (station occupied for 24-h

periods), Nakamura and Matsumoto found that skip- jack tuna, Katsuwonus pelamis, larvae were most numerous followed by Auxis larvae. Because currents around the Marquesas Islands are suspected to be weak, Auxis larvae as well as those of other tunas could not have drifted very f£u- from the spawning site; therefore,

18
Table 10. —Number and size ranKe of larval Auxis thiizaril (:) mm or less) collected in 1-h horizontal hauls (three 1 m nets towed simultaneously at different depths) and

3U-min oblique hauls (surface to 200 m) by the KV Hugh M. Smith in Hawaiian and

equatorial waters (Matsumoto 1958).

Cruise and
20
spent gonads were taken in June-July and July-August,

respectively (Suzuki and Morio 1957). This was con- firmed by Hamada, Morita, Ishida, and Takezawa

(1973), who examined the gonad condition factors of A. rochei caught in waters off Kochi Prefecture and sug- gested that spawning took place in late June. They also examined the gonad condition of A. rochei taken off Taiwan in June-July and found that the gonad index was very high indicating perhaps that spawning was imminent. Concerning the larvae, Yabe and Ueyanagi

(1962) collected those of A. rochei from May through

July near Japan whereas Yokota et al. (1961) obtained

larval A. rochei, measuring 3-5 mm TL (total length), from south of Shikoku and Kyushu in June-August but most were taken in July. Farther south, however, in the Celebes and South China Seas, larval A. rochei were col- lected predominantly during January and February (Yabe and Ueyanagi 1962).

It has also been established that Auxis spawn in the Indian Ocean (Bogorov and Rass 1961) and that spawning extends for at least 8 months from August to April; however, there is no information on their spawning activity for the rest of the year. Among the tuna larvae col- lected by the Dana Expedition from south of the equator between lat. 3° and 24°S at long. 50°E, Jones and Kumaran (1963) found 131 larval A. thazard, indicat- ing that this species spawned in this area in DecemberJanuary. North of the equator in the Laccadive Sea, they also found larval A. thazard in the samples indi- cating a spawning period in January-April, slightly later

in the year than in the south. Rao (1964) added that among samples of A. thazard he collected from commercial landings, the percentage of maturing and mature

ovaries was highest in March. In August and

September, the majority of the fishes examined were ac- tually in spawning condition and some individuals completely spent. Samples obtained subsequently in November contained only spent fish. Although evidence indicates that A. thazard spawn in the Indian Ocean, present data concerning spawning of A. rochei are still too meager to draw definite conclusions. Jones and Kumaran (1%3) found 20 larval A.

rochei in the Dana collection from the Indian Ocean. All were collected in December-January from three loca- tions between Madagascar and the African coast. Matsumoto (1959), who examined the collection made by

the Dana from the eastern Indian Ocean southwest of the Sunda Archipelago in August-November, recog- nized the presence of two types of Auxis larvae. Gor- bunova (1963) confirmed Matsumoto's observations, reporting that two types of Auxis larvae were distin- guished in the Soviet collection made in the Indian Ocean. From the information available, Jones and Kumaran (1963) hypothesized that low latitudinal areas

of the eastern Indian Ocean southwest of the Sunda

Archipelago and the western Indian Ocean between Madagascar and the coast of Africa are possible spawning grounds of A. rochei. Gorbunova (1965a, 1965c) col- lected A. thazard larvae of 5-11 mm in February and March in the Bay of Bengal.

In the Atlantic Ocean, tuna larvae have been col- lected off the west coast of Africa, mainly in the region of Dakar at the western tip of Africa, and off Takoradi in the Gulf of Guinea (Kazanova 1962; Richards 1969). Among the various species of larval tuna and tunalike

fishes collected, those identified as Auxis were most

numerous. Kazanova had at his disposal a series of Auxis larvae some as small as 3 mm. Furthermore, in addition to larvae identified as ^4. thazard, there were

others which were similar to A. thazard but differing in some characteristics. Kazanova referred to them as the second of the two groups (Type II of Matsumoto 1959) of Auxis larvae. Therefore, it can be inferred that these

regions of the Atlantic are also spawning grounds of the two species of Auxis.

Certain areas in the Mediterranean Sea have also been suggested as possible spawning sites of Auxis sp. Ehrenbaum (1924) reported Auxis spawning from July

to September in the Mediterranean. Off Greece, spawning of Auxis was also reported to occur from June to September and in the Gulf of Catania (Sicily) in June and July (Belloc 1954). Declerc et al. (1973) reported capturing recently hatched larvae of Auxis in waters surrounding the Balearic Islands, located off the Mediterranean Spanish coast. In August-September

1971, 140 larval Auxis were captured by the RV Ichthys and in June-July 1972, an additional 52 Auxis larvae were taken. According to the authors, larval Auxis was

the most abundant among the tuna larvae captured which included Thunnus thynnus, T. alalunga, and

Sarda sp. The presence of the planktonic stages of all these tuna species in the plankton-net catches provided

evidence that these species spawned in waters near the

Balearic Islands. The waters around Tunisia have yielded Auxis in a mature state, and this bit of evidence has led Postel

(1964) to believe that spawning probably takes place in Tunisian waters. Postel examined the gonads of Auxis

caught in madragues (traps) off Tunisia, found them

mature, and concluded that spawning occurs in JuneAugust. For Auxis that were taken off Algeria, Postel

also determined that they spawn there probably in August. Off the northwestern coast of Africa, Cape

Verde Islands and Dakar have been mentioned as spawning sites of Auxis (Frade and Postel 1955; Kazanova 1962). Gorbunova (1965c) reported capturing about 6(X) Auxis larvae at one station in the Gulf of Aden.

In the western Atlantic, the Gulf of Mexico appears to be a large spawning ground for Auxis. Hayasi (1972) reported collecting tuna larvae identified as yellowfin, bigeye, skipjack, and Auxis in gulf waters (Fig. 17). Klawe and Shimada (1959), who collected juvenile Auxis in March-April and in June-August from a large number of stations spread widely over the Gulf of Mexico, stated that the north central gulf region is definitely

a spawning locality for this species. The area southeast

of the Mississippi River Delta is another possible spawning site because most of the larvae were captured there, but Klawe and Shimada pointed out that the collection 21
Figure 17. —Plankton stationB occupied during cruises of the RV

Shoyo Maru in 1969-70. Solid triangles represent stations at which AuxU larvae were collected. (Hayasi 1972.)

may reflect sampling intensity rather than actual dis- tribution of young Auxis. In waters adjacent to the gulf extending from Cape Hatteras to Cuba and particultirly

in the Strait of Florida, larval Auxis have shown up in collections made in February-March (Tibbo and

Beckett 1972) and in May-June and in August (Klawe

1961) indicating the possibility that spawning occurred during these months in this region.

3.17 Spawn

The eggs of Auxis are pelagic. Rao (1964), who ex- amined ovaries of A. t hazard from the Indian Ocean,

observed that ripe eggs flowed freely out of the ovaries when slight pressure was applied on the abdomen. Fresh

ripe eggs were perfectly spherical, had a colorless homogeneous yolk mass, and had an average diameter of

0.97 mm (range of 0.88-1.09 mm). Preserved eggs were somewhat translucent and had an average diameter of 0.86 mm (range 0.78-0.98 mm). Each ripe egg contained

a fairly large, spherical, single oil globule which

averaged 0.22 mm (range 0.21-0.22 mm) in diameter. Like other scombrids, female Auxis do not extrude all their ripe eggs during spawning. Microscopic examination by Yoshida and Nakamura (1965) of ovaries from

eight A. thazard and five A. rochei caught in Hawaiian

waters revealed that both species had eggs in various

stages of resorption. One specimen of A. thazard still contained residual eggs in the lumen; their diameters

varied between 0.75 and 1.30 mm and averaged 1.08 mm. No free residual eggs were found in A. rochei. Concerning primordial eggs, two size groups—one ranging from 0.07 to 0.10 mm and the other from 0.17 to 0.22 mm in diameter—were noted in ovaries of A. thazard. In A. rochei, there was only one group of primordial eggs ranging from 0.07 to 0.08 mm in diameter. The development of eggs and larvae of Auxis has been

described for the two species from the Atlantic and

Pacific Oceans. Mayo (1973) successfully hatched eggs

of two species of Auxis collected from the Straits of

Florida and described the growth, behavior, ecology, and development of the larvae. In the Pacific, artificial fertilization of eggs and subsequent rearing of larvae were conducted by Japanese scientists and their results have been briefly discussed in section 2.41. Harada, Murata, and Miyashita (1973) and Ueyanagi et al. (1973) published the results of these tuna-rearing ex- periments in which fish in advanced stages of sexual maturity were used. Of three conventional fishing methods attempted, only purse seining proved effective

in capturing mature yellowfin and skipjack tunas and

set nets for collecting mature Auxis. The "dry method" was used to obtain mature eggs and sperm. Mature eggs from ripe female Auxis generally oozed out when pressure was applied to the abdomen;

therefore, no incision was necessary (Ueyanagi et al. 1973). The investigators then obtained mature males and applied pressure on the abdomen to force the sperm-bearing milt over the eggs. The mixture was gently stirred before any water was added.

Collected and fertilized at sea, the eggs were then

transported to various laboratories to be hatched

(Ueyanagi et al. 1973). Fertilized eggs of Auxis were

transported to laboratories in 2-4 h on eight different oc- casions. The mortality of the eggs, which were trans- ported in plastic bags filled with oxygen-saturated

seawater, was negligible. The details of the success obtained in fertilizing and rearing both A. thazard and

A. rochei follow.

Auxis thazard

Mature eggs of A. thazard caught at Mera, Shizuoka

Prefecture, were collected, fertilized, and successfully hatched on five occasions between 15 July and 15 August 1971 (Ueyanagi et al. 1973). Larval rearing was

most successful on the first trial when survival lasted up

to 10 days. Newly hatched larvae emerged 25 h after fer- tilization and measured 2.59 mm TL. Heavy mortality

usually occurred 4-5 days after hatching. At Kushimoto, Wakayama Prefecture, mature eggs were collected, fertilized, and transported to the Marine

Laboratory of Kinki University on three occasions in 1972 (Harada, Murata, and Miyashita 1973; Ueyanagi

et al. 1973). The spherical, fertilized eggs were buoyant and varied from 0.93 to 0.98 mm in diameter (Fig. 18). Emerging 34-62 h after fertilization at temperature

ranging between 21.4° and 23.5°C, the newly hatched

larvae measured from 3.26 to 3.60 mm TL. On the third day after hatching, the larvae were fed rotifers, which

were replaced by marine plankton, mainly copepods, on

the seventh day. Twenty days after hatching, the larvae were fed several types of fish flesh. The larvae were

placed in various-sized aquaria to determine the effects of tank capacity on growth and survival; the investi- gators found that growth was rapid in the 0.5-ton (500

liter) aquarium during the first 30 days after which time

the rate slowed appreciably (Table 11, Fig. 19). Max- imum survival was 36 days. Larvae reared in aquaria which had capacities of 3 tons (3,000 liters) and 70 tons

22
1 cnili/cil c.sj; ol Ii/v/v ihuzurj (Iiu.ix.hI.i) m

morula slaee.(1.95iiini in dianictcf.

20 hours .illor rotlili/jlion. o

I'ostl.irv.i alniiil 5 days old.

Posll.irv.i .iIhhiI 10 d.iVN old. t:5V-5*

Jusi before halchlnfi, 40 hours

'n-jinnitiJiiiiiHT

> oung fisli about I 7 days old. (i4iimi in loi.ii IcTiijlh.

o »

ii,;iii;ii«iTM. I) ii| i'V''iji"i|"'|i«|iiipi|iiiijiiii|ii.f .|.iii|i*#ii|iiii|wippjj^,.^..,,,,,,L,.. :,",

I arva about I day old. .V4iiim in total loiii^lh. Youns; fish ab-nii 4o d.i\ s old, I :ihnin in total Icnuth.

Figure 18.— Various stages of development of egg and larvae of Auxin thazard (Harada, Murata. and Miyashita 197.3).

(70,000 liters) grew faster than those reared in the 0.5- ton aquarium, reaching a length of 12 cm in 33 days

(Table 12, Fig. 19). Growth decreased significantly after that and the last larvae died on the 41st day.

Auxis rochei

In 1972, the collection and fertilization of eggs from mature A. rochei succeeded in seven trials at Mera,

23
Table II. —Measurements on Auxis thazard larvae reared in a small

(500 liter) aquarium (Harada, Murata. and Miyashita 1973). Days

after hatching

Total

length Fork

length Body

depth (mm)

10
n Fertilized egga of Auxls rochei

(marusoda) In morula stage, 0.98 mm

In diameter.

l'osll;irv;i aboiii 5 il.ivs old.

2(1 hours iillor rcrtili/;ilioii

l'o<ll.irv;i ;ilioiil 10 J.ivs old.

2S hours al'lor lerlilizalion.

N ouii!: fish .iboul I') d.iys old, 4')tiimi in lol.il lcii-;ih.

Jusi bclorc lutchin!:. 38 hours.

\J -.-. \

*

Lurvj ahoul I d.i> old. ,V5iliiii in loljl Icnplh. iinni.ilun- llsli .iboiil 52 d.i>s oUI. I.S7nini in lolal lonmh

Figure 20.—Various stages of development of egg and larva of Auxia rochei (Harada, Murata, and Furutani 1973).

Experiments on artificial fertilization were continued

in 1973. On 15 July 1973, at 0500, ripe eggs were col- lected from A. rochei caught in a set net off Kashino, Oshima (Kushimoto, Wakayama Prefecture), and artificially fertilized (Kinki University Fisheries Experimental Station 1974). The water temperature at the time of fertilization was 25°C. The fertilized eggs, which

averaged 0.903 mm in diameter, were transported to the experimental station at Shirahama. The following day, an estimated 7,000 newly hatched larvae were placed in 25
• • TOTAL LENGTH ^^ ^^^^^ j^^^^ O O WATER TEMP ^ " LAnt.lL iur«r3

4 A WATER TEMP _i " 5"i"ll lAiMr

DAYS AFTER HATCHING

Figure 21. —Growth curves of Auxia rochei larvae reared in two

large (3,000 and 30,000 liter) and one small (500 liter) aquaria. Length measurements are based on a single fish. (Harada, Mura- U, and Furutani 1973.)

a 3-ton aquarium and at age 2 days were fed rotifers and

marine plankton. By age 18 days, the larvae were 3 cm

long. From the 24th day, the larvae were fed minced fish flesh, and they grew rapidly, reaching 5-6 cm by the 27th

day; however, heavy mortality reduced the number of larvae to 36. By the 35th day, the survivors were 7.7-7.9 cm long. Only one larva survived for 50 days. The

Japanese investigators observed that this rate of growth was slower than in previous experiments and believed that it was due to lack of fish larvae as food. They emphasized, however, that the experiments succeeded in

artificially producing "seedlings" of A. rochei which

were several centimeters in length.

In general, then, these experiments on frigate and bullet tunas have shown that:

1. the eggs are buoyant, spherical, and measure

between 0.93 and 0.99 mm in diameter;

2. the eggs hatch from 34 to 62 h after fertilization in water temperatures ranging between 21.4° and

23.5°C;

3. the newly hatched larvae measure 3.26 and 3.60 mm

TL;

4. the larvae can be fed rotifers, copepods, and minced

fish flesh, in that order, after hatching;

5. the growth and survival rates of larvae are better in large rather than in small aquaria; and that

6. under the best conditions, A. thazard larvae grew 120 mm in 33 days whereas A. rochei larvae grew 157 mm

in 52 days.

3.2 Preadult phase

3.21 Embryonic phase

See section 3.17 above.

3.22 Larval phase

Studies of anatomical development of larval Auxis show that gross nucleation of the central nervous system

of young Auxis (3.6-23.0 mm SL) is similar to young Thunnus albacares but heavier than in Katsuwonus

pelamis, T. obesus, T. thynnus, T. alalunga, T. atlanticus, and Euthynnus alletteratus (Richards and Dove

1971). Using sectioned organs and tissues, Richards and Dove determined that the swim bladder in katsuwonus and Auxis, which, among scombrids is similar in the

very early stages of development, starts to degenerate when the larvae reach the size of 9 mm, and is nearly completely degenerated in specimens 24 mm long at which time only the gas gland remains. Richards and Dove also determined that in Auxis the kidney is short

until the larvae reach 11 mm after which it lengthens

significantly and that the postcardinal vein is larger in Auxis and Euthynnus than in other species. The criteria for specific identification and separation

of Auxis larvae from those of other tuna and tunalike

fishes appear to be quite well established. Ueyanagi

(1964) has provided a key to identifying tuna larvae and

this is given below: Key to the larvae of tunas from

the western Pacific'

a Melanophores visible on the forebrain

bj One conspicuous melanophore present on

the caudal peduncle. No chromatophores on the isthmus or directly £inte- rior to the anus.

.... skipjack tuna, Katsuwonus pelamis

bj There is a dotted line of melanophores

along the midlateral line on the posterior part of the body. Chromatophores

appear on the isthmus and directly anterior to the anus. kawakawa, Euthynnus affinis a No melanophores visible on the forebrain.

c Melanophores appear on the sides of the body, d Chromatophores appear on the isth- mus and directly anterior to the anus. Melanophores appetir as dotted

lines along the middorsal, midlateral, and midventral lines of the caudal

region (usually in three lines). .... frigate and bullet tunas, Auxis spp.

dj Chromatophores do not appear on the

Applicable to larvae under about 8 mm TL. 26
isthmus nor directly anterior to the anus,

f One or several melanophores ap- pear along the dorsal and ventral

sides of the body,

g The most anterior of the chro- matophores on the dorsal

side is located posterior to the origin of the second dorsal fin. bluefin tuna, Thunnus thynnus

g The most anterior of the chro2 matophores on the dorsal

side is located anterior to the

origin of the second dorsal fm.

long-tailed tuna, Thunnus tonggol

f One of several melanophores

appear along the ventral side

of the body. bigeye tuna, Thunnus obesus

c No melanophores appear on the sides of the body. h Chromatophores appear at tip of lower

jaw. In the profile of the head, the

center of the eye is clearly located higher than the tip of the snout.

yellowfin tuna, Thunnus albacares

hj No chromatophores appear on tip of lower jaw. In the profile of the head,

the center of the eye is about on the same level as the tip of the snout or

slightly higher. albacore, Thunnus alalunga

Briefly, larval Auxis can be distinguished from other

larvae of tuna and tunalike fishes by the presence of a dotted line of melanophores along the dorsal, lateral, and ventral center lines of the caudal peduncle

(Ueyanagi 1964). The standard pattern is three lines, but sometimes only two—the dorsal and ventral—are

seen. In small specimens, however, the line on the

ventral side is not confined to the vicinity of the caudal peduncle but extends farther forward. Other characteristics include the appearance of melanophores on the isthmus and just anterior to the anus, late appearance of chromatophores on the forebrain (they do not appear

until the fish attains a body length of about 8 mm), and

poorly developed chromatophores on the first dorsal fin (they do not appear until the fish becomes about 10 mm

long). The outline of the anterior surface of the head

gives an impression of roundness and the separation betweeen the first and second dorsal fins becomes quite apparant at lengths of 10 mm or greater. Additional observations on the morphology of larval Auxis showed that the snout is shorter than in other tuna and tunalike larvae of comparable size (Jones

1963). Jones also noted that larval Auxis has 39 myotomes, a large chromatophore at the symphysis of the pectoral girdle, and that fin ray development is later

than in other tuna and tunalike larvae. Jones and Kumaran (1964) added that compared with larval Euthynnus affinis. larval Auxis exhibits later develop- ment of the first dorsal fin and has relatively few chro- matophores on the first dorsal fin membrane, and its fin membrane is relatively narrow. The variations in pigmentation among larval Auxis

has been used as a basis for separating the larvae into two possible types. Matsumoto (1959), who described

the morphology of larval and postlarval Auxis on a worldwide basis, pointed out the possibility that this variation may well be due to the presence of two species

of frigate tuna; therefore, he provided two separate descriptions calling them Type I and Type EI, as shown in Figure 22. Type I and Type II larvae show only minor differ- ences and the most obvious one is the variation in pig- mentation near the caudal peduncle (Matsumoto 1959). Type I, which resembles the form described as A. thazard (Matsumoto 1958), has three equally developed rows of pigment whereas in Type 11 the middorsal edge

of the caudal peduncle usually contains only one or two

chromatophores and the midlateral line has none. Subsequently, Type I larvae, which are stouter than Type n. were provisionally identified as A. thazard and Type II, which are relatively elongate, were identified as

A. rochei (Jones 1963). Further observations revealed

that compared to A. thazard, larval A. rochei has a

relatively shallow body, shows later development of the spinous dorsal, and has less intense pigmentation on the caudal peduncle (Jones 1963; Gorbunova 1969). A later study confirmed Jones' observation; Yabe and Ueyanagi

(1962) published the results of a study on larval tuna in which they positively identified specimens of A. rochei. A controversy over the identification of larval Auxis

erupted in 1963 when Jones (1963) questioned the re- sults obtained by Mito (1961), who described the egg and larval development oi Auxis (Fig. 23). Jones pointed

out that because A. rochei is the most common frigate mackerel in Japanese waters, the eggs and larval stages described by Mito may be this species; but he also noted

that there were striking differences between larvae described by Mito and those described by Matsumoto

(1959) and Jones (1961). These differences, he believed, should be explained by Mito. The breakthrough in artificial rearing accomplished by the Japanese should make it possible to comptire specimens of artificially reared larvae and those obtaind from net tows and night lighting to obtaln positive identification.

3.23 Adolescent phase

Juvenile Auxis can be distinguished from other tunas by certain morphological features (Schaefer and Marr

1948; Wade 1949; Mead 1951; Matsumoto 1962). They

can be distinguished from other tunas by the wide gap

between the first and second dorsal fins and the virtually colorless appearance of the first dorsal. Compared to Katsuwonus, however, the first dorsals are somewhat

similar, both having a lightly pigmented or colorless fin.

27
Figure 22.—Auxis larvae Type I. 7.05 mm (A) and T>pe II, 7.2 mm (B) (Matsumoto 1959). Katsuwonus, nevertheless, possess more pigment in the

first dorsal, which is also more or less brown or tan distally, whereas in Auxis, a few scattered chromatophores

are found along several of the anterior rays in the distal portion of the first dorsal. Compared with Euthynnus,

Auxis are rounder in cross section, relatively less compre.ssed laterally, and the length of the head and the caudal region shorter in comparison to total length. Juvenile Auxis also pKJssess a black sptot at the isthmus which is seen only in one other genus, Euthynnus. In young Euthynnus, however, the first and second dorsals are continuous. The large interspace between the dorsals, the low ray count of the first dorsal, and the absence of an elaborate "trellis" of the vertebrae also help to separate Auxis from Euthynnus and Katsuwonus. The pattern of pigmentation is also a good identify- ing character. Schaefer and Marr (1948) wrote "In the

smaller specimens the prominent areas of pigmentation

are on the upper and lower jaws, above the snout, around the postero-ventral margin of the orbit, on the upper operculum, between the orbits, along the mid-line

of the body, along the bases of the dorsal and anal fins including the finlets, and around the posterior end of the

urostyle. Large chromatophores in the peritoneum show

through the body wall along the upper half of the body

cavity. None of the fins or finlets bears pigment spMsts, with the exception of the first dorsal. The first dorsal bears a few scattered chromatophores, largely distrib- uted along the spines." Several features of the axial skeleton also are useful in separating the juveniles of partially digested tunas. Potthoff and Richards (1970), who studied regurgitated

food of terns and noddies from the Dry Tortugas, Fla., observed that damaged or partly digested Auxis and

those <30 mm could easily be separated from Euthynnus if judged on fin position and gill raker counts. Potthoff and Richards revealed that extreme care must be

used to examine Auxis specimens because the spines of the first dorsal in some juveniles are subcutaneous and come to the surface in the fin interspace. Furthermore,

they noted that the number of gill rakers is evenly dis- tributed over the range for Auxis whereas they are more

normally distributed in other species. Counts were as

follows: 19 in 38% of the ceratobranchials, 20 in 25%, 21

in 25' c, and 22 in 12%. Potthoff and Richards suggested

that this kind of distribution may be attributable to the presence of two types of Auxis £md their intermediates.

28
Figure 23.—Various stages of development of egg and larvae of Auxis (Mito 1961); 1) Pelagic egg, 7'j h after

collecting. 1.04 mm in diameter, oil lobule 0.26 mm; 2) 29-myotome stage, 94 h after; .3) 10'2 h after; 4) 14'? hr after, shortly before hatching (24'-27"C); 5) empty egg capsule; 6) larva just hatched, 2.70 mm TL, myotomes 11 + 32 = 43; 7) larva 16'; h after hatching, 3.68 mm TL, myotomes 9 + 31 = 40; 8) larva 30 h

after. 3.58 mm TL, myotomes 9 + 31 = 40; 9) larva 42'2 h after, 3.92 mm TL, myotomes 9 + 32 = 41; 10) larva .Vi h after. 3.75 mm TL, myotomes 8 + 31 = 39.

In addition to the gill raker counts (38-43 for A. thazard and 43-49 for A. rochei), the form of the free parapophyses is a useful feature of the axial skeleton for separating Auxis to species. Watanabe (1964) deter- mined that in A. thazard, the free parapophyses are

long, extend downward and almost touch the haemal

spine of the preceding vertebra whereas in ^4. rochei they

are short. This difference in length of the parapophyses

IS particularly evident on specimens of 111 mm or more

(Figs. 24, 25). Four other characters used by Watanabe (1964) in- clude: 1) the number of the vertebra on which the first free parapophysis occurs, 2) the number of the vertebra on which the first inferior foramen occurs, 3) the ratio of the length of the caudal to the precaudal vertebra, and

4) the ratio of the body height at the origin of the anal

29
Auxis thazard Auxis rochei

Specimen No. 7 Standard length 88.5 mm. Specimen No. 28 Standard length 94.0 mm.

(free parapophysis) Specimen No. 23 Standard length 124.6 mm. Specimen No. 37 Standard length 123.6 mm.

Figure 24. —Caudal vertebrae o( Auxis thazard (left) and A. rochei (Watanabe I964I.

fin to the distance between the origin of the first and sec- ond dorsal fins. Watanabe found that in A. thazard, the

first free parapophysis occurred most frequently on the

r~l Auxis rocl<ei F1 Auxis tliazord

nil

nn n n n H

-, nn n n n m

nn n n nO n

„ nn n n mH

riEi n D n nn

90 100 no 120 130 140 STANDARD LEIWTH(mm)

23d vertebra, whereas in A. rochei, it was on the 21st or 22d vertebra on SO'^ of the specimens; however, there were some in which it occurred on the 23d vertebra. Furthermore, the inferior foramen in young specimens

occurs on the 27th or 28th vertebra in A. thazard and

usually on the 29th and 30th vertebra in A. rochei. Watanabe also observed that the ratio of the length of the caudal to the precaudal vertebra for young speci- mens was 0.973 or less in A. rochei and 0.985 or more for A. thazard. This character also changes as growth occurs; therefore, the stage of growth needs to be consid- ered in making a diagnosis. Figure 26 shows a plot of the ratios—body height at the origin of the anal fin to the distance between the origins of the first and second

dorsal fins —on standard lengths and except for a single specimen which showed an intermediate value, those for A. thazard tend to be higher than for A. rochei.
Table IS. —Standard length and gill raker counts of juvenile Auxi»

thazard and A. roehei (Wade 1949).

Species
Table 16. —Measurements of Indian Ocean juvenile ^uxm thazard (specimen Nos. 1-12, 14) and A. rochei (specimen Nob. 17, 19-26) in millimeters and the gill raker counts in the upper and lower limbs (Jones 1961).
Bluefin tuna, T. thynnus (Yokota et al. 1961; de Jager

et al. 1963; Matthews et al. 1977) Kawakawa, Euthynnus affinis (Ronquillo 1954; Philippine Islands)

Little tunny, E. alletteratus (Dragovich 1969) Skipjack tuna, Katsuwonus pelamis (Kishinouye

1917, 1923, Japan; Ronquillo 1954; Yokota et al. 1961; Batts 1972, North Carolina) Yellowfin tuna, T. albacares (Kishinouye 1917; Naka- mura 1936, Celebes Sea; Mead 1951, eastern tropical Pacific; Ronquillo 1954; King and Ikehara

19.56; Yokota et al. 1961; Alverson 1963, eastern

tropical Pacific; de Jager et al. 1963; Watanabe

1964; Dragovich 1969; Matthews et al. 1977)

Billfishes

Black marlin, Makaira indica (Nakamura 1942, Taiwan; Watanabe 1964) Atlantic blue marlin, M. nigricans (Krumholz and de

Sylva 19.58, Bahamas; Erdnian 1962, Puerto Rico)

Perhaps because of their abundance, Auxis are considered forage elements which occupy an important

position in the food chain. In the eastern Pacific, A. thazard are of no commercial importance but they contribute to the tropical skipjack and yellowfin tuna

fishery indirectly, because they constitute a significant part of the food of adult yellowfin tuna. In the western

Pacific, Auxis, because of their distribution and abun- dance close to land masses, form a relatively high

proportion of the forage of tunas and billfishes that are

similarly distributed (Watanable 1964). The size oi Auxis consumed by large predators varies considerably. Klawe (1963) noted that specimens of A. thazard found in the stomachs of skipjack tuna, yellowfin tuna, and kawakawa varied from 60 to 125 mm, but Watanabe (1964) has shown that Auxis up to 320 mm

may fall prey to large tunas and billfishes (Table 18). In the Atlantic. Krumholz and de Sylva (1958) reported

that Auxis ranging in size from 200 to 250 mm have been

recovered from stomachs of blue marlin caught near Bimini in the Bahamas.

Table 18.—Size-frequency distribution ot AuxU found in the stomachs of tunas and marlins (Watanabe 1964). Body length (cm)

Period

•Ian. 19.53 Feb. 19.53 Nov.-Dec. 19.56 Dec. 1951- Mar. 19.52

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32

18 1 7 —

A - Banda Sea. north of Timor Island. Indonesia. B Sawu [Saru] Sea. .south of Flores Island. Indonesia. C - Off the coast of Queensland. Australia. n - Off the southern coast of Shikoku, .Japan. 3.35 Parasites, diseases, injuries, and abnormalities

Indo-Pacific sailfish, Istiophorus platypterus (Jones

19.58. India; Idyll and de Sylva 1963, western

Atlantic)

.Striped marlin, Tetrapturus audax (Royce 1957, central Pacific) White marlin, T. alhidus (Idyll and de Sylva 1963; Davies and Bortone 1976, northeast Gulf of Mexico)

Other Species

Barracuda, Sphyraena sp. (Idyll and de Sylva 1963) Lancetfish, Alepisaurus sp. (Matthews et al. 1977) Mahimahi, Coryphaena hippurus (Ronquillo 1954;

Idyll and de Sylva 1963; Rose and Hassler 1974, North Carolina)

Porpoise, Stenella sp. (Perrin et al. 1973, eastern tropical Pacific) Thresher shark, Alopias yulpinus (Whitley 1964, Australia) Tiger flat head, Neoplatycephalus richardsoni (Whitley 1964) Wahoo, Acanthocyhium solandri (Idvll and de Sylva

1963)

Table 19, extracted from MacCallum and MacCal- lum (1916), Linton (1940), Manter (1940, 1947, 1954), Vervoort (1965), Lewis (1967), Pillai (1967), Silas (1967a), Silas and Ummerkutty (1967), and Mamaev

(1968), lists the monogenetic and digenetic trematodes,

cestodes, and copepods that are parasitic on Auxis.

Table 19. —Monogenetic and digenetic trematodes, cestodes, and

parasitic copepods on Auxis (Mact^allum and MacC^allum 19iri; Linton 1940; Manter 1940. 1947, 19.54; Vervoort I9B5; Lewis 19fi7; Pillai 1967; Silas 1967a; Silas and Ummerkutty 1967; Mamaev 1968). Type of parasite Family Sp'>cies

Monogenetic trematode

Digenetic trematode

Parasitic copepod

Hexostomatidae

Bucephalidae

Fellodistomidae Didymozoidae

Gorgoderidae Dasyrhynchidae

(^aligidae

Bomolochidae

Hexoxtoma auxidi Rhipidocotyle capitatum

Tergextia laticollis Didymozoon auxis I'hacclotrema claviforme Opepherotrema planum

Colocyntotrema auxis Phyllodislomum lancea

Callitetrarhynchus

gracilis Caligus macarnvi

C. productus

C. auxisi Bomolochus mycterobius

33
3.4 Nutrition and growth

3.41 Feeding

Observations on the feeding habits of larval A. rochei

indicate that they are particulate feeders. Yokota et al. (1961) observed that larval A. rochei between 3.0 and 5.4 mm in length consumed nauplii ranging in size from

0.10 to 10.30 mm and copepodites of about 0.55 mm.

Auxis have been observed to use their caudal fin to hold their position in swift currents to capture drifting food and to migrate away from swift currents in search

for food. They seek food not only by sight but also by us- ing their lateral line sensory system (Imamura 1949; Uchihashi 1953). The maximum size of fish eaten and also the total weight of food consumed by different sizes of A. rochei caught in the pole-and-line and set net fisheries have

been discussed by Morita (1972). For the relationship between length of prey and body length of the predator, he calculated the following formula: Y = -0.015X2 + 0.358X + 9.075

where X = length of predator in centimeters y = length of prey in centimeters. The curvilinear relationship can be seen in Figure 28. The linear regression describing the relationship between weight of stomach contents and body length is shown in Figure 29. The equation is as follows: Y = 2.33X - 40.78

where X = length of A. rochei Y = weight of the stomach contents in grams.

The selection of food organisms above a minimum size apparently is determined by the magnitude of gill raker gaps among mackerels, tunas, and dolphins. Magnuson

and Heitz (1971), who calculated mean gill raker gaps of 0.74 mm for A. thazard and 0.51 mm for A. rochei, con- cluded that among scombrids Auxis and Katsuwonus

have small gill raker gaps. The filtering areas of the two

species of Auxis, which measured 570 mm^ for A. thazard and 55 mm^ for A. rochei, were intermediate when compared to other scombrids. Magnuson and

Heitz suggested that gill raker gap and the maximum

distensibility of the esophagus would set limits on the

size ot toed eaten. Usually, however, selectivity in lead- ing among scombrids is masked because of the diversity

of food organisms in the size range consumed by them. Competition for food appears to be of fundamental importance in inducing young scombrids to feed (Clemens 1956). In the eastern Pacific, juvenile Auxis

kept in shipboard aquaria learned to eat unnatural food

that was offered them by watching other species

mahimahi, Coryphaena sp.; jacks, Caranx sp.; pompano, Trachinotus sp.; threadfins, Polydactylus sp.; and

pomacentrids, Chromis sp. —splashing and moving

FORK LENGTH (cm)

Figure 28.— Relationship between fork length of Auxis rochei and

total length of prey consumed, off Japan (Morita 1972).
stomachs of yellowfin and skipjack tunas, had larval copepods and decajxxls in their stomachs (Kishinouye

1924). Auxis feed on a large variety of fishes, crustaceans, and molluscs. In Auxis captured in Hawaiian waters,

fishes comprised the greatest volume with crustaceans

next in rank (Tester and Nakamura 1957). Those cap- tured in Japanese waters fed on plankton as well as a wide variety of herringlike fishes such as silverside, Atherina sp.; anchovy, Stolephorus sp.; small round

herring, Spratelloides sp.; and immature Engraulis sp. (Kishinouye 1923). Okada (1955) observed that A.

rochei fed on small pelagic organisms, anchovies, silver- sides, and other small fishes. Of A. rochei taken in Australian waters, Whitley (1964) found the stomach cram- med with anchovies and young mullet. Auxis also feed on their own young. Yokota et al. (1961) examined A. thazard caught by trolling in Japanese waters in June 1959 and discovered that their diet included young Auxis as well as skipjack tuna. Other items in their diet included jack mackerel, Trachurus japonicus; flyingfishes, Exocoetidae; file- fishes, Monacanthidae; Mene maculata; round herring, Spratelloides japonicus; anchovy; and squids (Table

20).Troll-caught A. rochei consumed anchovy as the principal food and other items only infrequently whereas

those taken in set nets fed predominantly on jack mackerel with anchovy next in imjxirtance (Yokota et

al, 1961). Spotted mackerel. Scomber australasicus. and

Table 20. —The number of identifiable orgamisms found in stomachs

of 30 AuxU thazard caught by trolling gear in Japanese waters (Tokara fishing ground), June 1959 (Yokota et al. 1961).

Prey No. Prey No. Prey No.

Skipjack tuna 9

Frigate tuna 1 Jack mackerel 2

Flyingfishes

Filefishes Ginkagami^

1 Kibinago'

1 Squid

1 Anchovy

'Kibinago — Spratelloides japonicus (Houttuyn). 'Ginkagami — Mene maculata (Bloch).

lizardfish, Saurida undosquamis, were also found in fair numbers in stomachs of fish caught by set net but were

usually absent in stomachs of fish taken bv trolling (Table 21). In Indian waters, Thomas and Kumaran (1963) and Kumaran (1964), reporting on the stomach contents of

11 juvenile A. thazard (49-132 mm), observed that in small individuals <75 mm, fishes constituted 88"^ and

crustaceans 12''c by volume with squids entirely absent

(Fig. 30). In individuals >75 mm, fishes formed 39% and

Crustacea 42''r by volume. Squids occurred very infre- quently. Anchouiella sp. and Leiognathus sp. were the

AUXIS THAZARD 49-75mm

AUXIS THAZARD

76- 132 mm

I [ FISH

j I CRUSTACEA

j

j CEPHALOPODA ^H MISCELLANEOUS

AUXIS ROCHEI

170-252 mm

Figure 30.—Diagrams illustrating the composition, by volume, of the stomach contents of i4uxM thazard and A. Rochei caught in the Indian Ocean (Thomas and Kumaran 1963).

Table 21.—The stomach contents ot Auxis rochei caught by trolling gear and set net in Japanese waters. 19,'>8-61 (Yokota et al. 1961).

Date

Fishing ground Num- Spotted Jack Nezumi Kurotachi Sagi- Lizard- Hataber mackerel mackerel Saury gisu' kamasu" fue^ fish aji' Matouishi- mochi Mishimaokoze'' Squid Anchovy

Trolling line: Dec. 19.t8 Kumanonada

Jan. 19.59 Kumanonada

Feb. 1959 Kumanonada

Mar. 19.59 Kumanonada

Dec. 1960 Kumanonada

Jan. 1961 Kumanonada

Mar. 1961 Kumanonada

Apr. 1961 Kumanonada

Set or fixed net: Apr. 1961 Tanegashima

187
most common among the fish species consumed. Among

31 preadult A. rochei (170-252 mm), fishes constituted

42'~f by volume and were found in 80% of the samples

(Fig. 31). Those that were important were Sardinella

spp., Anchouiella sp., Leiognathus sp., and carangids (Table 22). Crustaceans were next in importance ac- counting for 24% by volume and found in 77% of the samples. Rabindra Nath (1962) reported that among the most common crustaceans consumed hy Auxis in Indian waters were Rhopolophthalmus sp., Hyperia

bengalensis, Oxycephalus clausi, Pseudophausia lati- frons, Acetes erythreus, and Squilla larvae. Cephalopods formed 22% of the food consumed

(Kumaran 1964), but pteropods were relatively unimportant in the diet (Rabindra Nath 1962). Kumaran

(1964) also noted that larval stomatofX)ds and Lucifer constituted a major portion of the diet of some

specimens captured near Quilandy on the west coast of

India. Other items of food occasionally seen were

chaetognaths, Halobates, and polychaetes.

-List of food items of preadult specimens of Auxu rochei from the Indian Ocean (Kumaran 1964).

Figure 31. —Percentages, by volume, of the types of food con- sumed by preadult specimens of Auxis rochei in the Indian Ocean (Kumaran 1964).

3.43 Growth rate

Observations on jKjstlarval growth indicate that captive Auxis grow faster than Euthynnus lineatus (Clemens 1956). Of five postlarval Auxis placed in an aquarium for observation, three died several hours later from injuries received in handling. Of the two remaining

Auxis, one measuring 20 mm TL grew to 40 mm in 6

days; the larger 30 mm specimen reached 46 mm during

the same period. The rate of growth of A. rochei estimated from modal

progression of length-frequency data appears to be

rather slow compared to that observed in larvae that were hatched from artificially fertilized eggs. Hotta
increases with age because older fish tend to gain pro- portionately more in weight than in length. The formula

for K is K = aW

L3

in which a = constant W = weight in grams L = standard length in millimeters (Rounsefell and Everhart 1953).

At La Linea, Spain, the average values of the condition factor for each length group of A. rochei were usually larger for males than for females (Rodriguez-Roda

1966). Furthermore, the values of X decreased from May

to August, but increased in September which coincided with the spawning period. Comparison of K among the

size groups showed no significant differences. Ishida (1971), who also calculated condition factor of Auxis caught off Japan, found that of the two species, A. thazard caught at Mikomoto and Shionomisaki had

higher indices than A. rochei. Between areas, however,

there appeared to be no appreciable difference. The length-weight relationship has been described for A. thazard and A. rochei caught in the world's oceans. The expression for the relationship is W = af

in which W = weight in grams L = length in centimeters

a and b = constants.

Table 23 gives the constant for the expression of the

predictive length-weight relationship calculated by

Ishida (1971) for both species of Auxis and by Yasui

(1975) for A. rochei caught in Japanese waters, by

Lenarz (1974) for A. rochei captured in the Atlantic Ocean, by Rodriguez-Roda (1966) for A. rochei caught in Spanish waters near the Strait of Gibraltar, and by Sivasubramaniam (1966) for both species caught around

Sri Lanka in the Indian Ocean. Figure 33 depicts the length-weight curves constructed by Rodriguez-Roda

(1966) and Yasui (1975). For A. rochei caught in Spanish

waters, Rodriguez-Roda has also provided the average

and calculated weights for the length groups studied (Table 24). The maximum size reported is 66 cm for A. rochei in the eastern Atlantic Ocean (Collignon see footnote 2).

Table 24. —Length groups, average weights, and calculated average

weights of Auxis rochei caught at Barbate, Tarifa, and La Linea, Spain, in 1958, 1961, 1963, and 1974 combined (Rodriguez-Roda 1966).

Size
W-l 549XI0"'L'

FORK LENGTH (cm) FORK LENGTH (cm)

Figure 33. —Length-weight relationship of Aiuu rorhei caught in (A) Japanese waters (Yasui 1973) and in (B) Mediterranean waters (Rod- riguez-Roda 1%6).

contents of molybdenum and nickel in the liver show no

relationship to sexual maturity, rate of growth, and area

of catch. The blood of Pacific A. rochei has been tested and

found to be relatively high in hemoglobin concentration (Klawe et al. 1963). Barrett and Williams (1965) showed

that compared with other scombrids, the mean level found for A. rochei was among the highest, reaching 19.2 g/100 ml and varying from 16.5 to 22.8 g/100 ml. Blood smears of A. thazard caught in Puerto Rican waters showed a blood count of 1 large hemoblast, 1 small hemoblast, 1 large lymphocyte, 30 small lymphocytes,

45 thrombocytes, and 22 granulocytes (neutrophils) (Saunders 1966). Mature erythrocytes from A. thazard

blood averaged 10 /i in length and 7.5 n in width.

3.5 Behavior

3.51 Migrations and local movements

Not much is known about the migration of Auxis in the world's oceans. Most of what is known comes from

results of studies conducted in Japanese waters. Hotta

(1955), after examining the fluctuations in landings of A. rochei along the coast of Japan in 1952, determined

their seasonal movement up and down the coast of Japan. He observed that they occur in the Satsunan Sea

region and off Shikoku early in the year, but as the year

progresses, larger catches are made farther north and by

June they are landed in the Izu Archipelago region (Fig. 35). The schools reach their most northern point around Hokkaido in September, then move southward until in December they appear to be concentrated mostly off the

Izu Archipelago and Shikoku.

Detailed studies of long-distance movement of A.

rochei in Japanese waters showed that fish caught, tag- ged, and released in August, September, and November

had a general tendency to move southward (Table 25, Fig. .36). Yamashige (1974) determined from his tagging

studies that within the population of A. rochei off Japan, differential migration occurs according to size. The large influx of small fish into the fishery, as shown

by the length-frequency distribution (Fig. 37), is the consequence of small fish moving south before the move- ment of large fish begins. Short-distance movement of tagged A. rochei in Japanese waters shows that they tend to move south- ward late in the year although some northward move- ment is also seen, but early in the year there is a definite northward movement (Table 25) (Morita 1972; Hamada, Ishida, Morita, Takezawa, Okabayashi, and

Ishii 1973; Hamada et al. 1974).

3.52 Schooling The schooling instinct, according to a number of

sources, is very strong and orderly in Auxis. Jones (1963) noted that the tendency of A. rochei to form large 38
K««Tl/!£ EARLV .«E
-Partial data on release and recovery otAuxu rochei in the 1965 and 1970-72 (iahing Hamada et al.
seasons ofr Japan (adapted from Morita 1972:
KAN (8 27 POUNDS)
.^"i^
eastern), off Tohoku Prefecture, and off Taiwan, deter- mined that the size range was usually narrowest among

fish caught in October-November whereas it varied widely among the fish caught in May-June (Fig. 40). Comparison of size data collected in Japan with those from the Philippines shows that the former's commercial catch constitutes smaller fish than the latter's. The

size of Auxis taken by the purse seiners Royal Venture and Southward Ho in the Philippines, shown in Figure

41, ranged from about 25 to 55 cm with the majority of the fish in the 40-4.5 cm range Data from the Indian Ocean show that the size of Auxis in the commercial catch also varies widely. Silas (1969) showed that A. thazard taken by drift net ranged from about 20 to 51 cm, but a large proportion of the

catch fell between 38 and 43 cm (Fig. 42). Silas also showed that compared with A. thazard, A. rochei are much smaller, ranging from about 19 to 29 cm and the mode at about 20 to 21 cm. The purse seine apparently

catches Auxis of more uniform sizes. Figure 43 shows

that A. thazard caught by purse seine in Indian waters had a relatively wide range from about 32 to 46 cm, but a

large proportion of the fish fell between 40 and 42 cm

(Silas 1969). Off Sri Lanka, A. rochei appearing in the commercial

catch are relatively small and their sizes vary from 16 to 34 cm; however, most are between 22 and 30 cm. Auxis

thazard, on the other hand, are much larger and range from 20 to 58 cm, but a large proportion of them falls be- tween 22 and 40 cm (Sivasubramaniam 1968). Separat20 25 30 FORK LENGTH (cm)

20 25 30 35 FORK LENGTH (cm)

Figure 40. —Percentage

area of capture. frequency distribution of lengths of Auxis rochei, by date and Japan (Hamada, Merita, Ishida, and Takezawa 197.3).

44

VengurloEXPLORATORY FISHING BY

R.V. VARUNA AND M.V.TUNA

FEBRUARY -APRIL 1966

1 POSITIVE STATIONS ( WITH TUNAS)

i NEGATIVE STATIONS ( NET FOULED

OR NIL CATCH OF TUNAS ) NUMBERS INDICATE TUNA CATCH

IN kg PER OPERATION

Auxis thazard

FORK LENGTH

TOTAL NO OF OPERATION : 38 NO OF POSITIVE STATIONS WITH TUNAS: 10(39-50 HOURS)

TOTAL CATCH OF TUNAS : 19438 kg AVERAGE DURATION OF EACH OPERATION : 3, 95 HOURS

AVERAGE CATCH OF TUNAS / OPERATION ; 1943-8 kg 72° 73» 74° 75° 76°

Figure 43.—Location of exploratory purse seine sets and catches

for tunas by the RV Varuna and MV Tuna off southwestern In- dia and the Laccadive Sea, and the length-frequency distribution and average weight of Aujcis thazard in the catches (Silas

1969). (N, = number of fishing operations; >', = number of specimens.

Figure 44.—The percentage frequencies of the number of Auxis

thazard caught per operation, by type of gear, and of the fork lengths

of the catches made off the southern and eastern coasts of Sri Lanka

(Sivasubramaniam 1968).

* 109 ISO 200 X 100 CATCH NUMBER PER OPERATION

34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 FORK LENGTH (cm)

Figure 45. —Length-frequency distributions of .4uJti8 rochei caught by traps at Barbate,

Tarifa, and La Linea. by month and year

(1961, 1963, 1964) (Rodriguez-Roda 1966).

28 32 36 40 FORK LENGTH (cm) 46
FORK LENGTH (cm)

Figure 46. —Length-frequency distribution of Auxis rochei caught by traps at Barbate, Tarifa, and La Linea. by month

and year (1961, 1964) ( Rodriguez- Roda 1966).

Taking a vessel-day as a unit of fishing effort, Ishida

(1972a) calculated the monthly landings per day and

found that this index fluctuated in a similar fashion as the number of vessels in operation per day (Fig. 47). I- shida also constructed a frequency distribution of the

landings per day and found that the primary mode fell at 30 t/day (t = metric tons) (Fig. 48). The annual mean

was 52.4 t/day but the seasonal mean, presumably cal- culated from selected data on landings per day for the peak months, reached 56.5 t/day. A plot of landings per day on number of vessels operating ijer day showed that em increase in fishing intensity was usually accompanied by a similar increase in catch per day. Off the coasts of Sri Lanka, Auxis are distributed in

all waters surrounding the island but their densities vary widely (Sivasubramaniam 1973). Using the number of fish caught per standard trolling unit (a vessel with a fixed number of lines) per day, Sivasubrama- niam found that A. thazard was most abundant along the southwest coast as catch rates reached 21.7

fish/boat-day. Almost equally high were the catch rates

JAN FEB M4R APR MAY JUNE JULY AUG SEPT OCT NOV DEC

Figure 47. —Landings of AuxU spp. per day in the fishery off Ashizurimisaki, Japan, calculated from data collected from October

1964 to June 1967 and combined by months (Ishida 1972a).

LANDING (METRIC TONS)

Figure 48.—Frequency distribution of landings per day in the fishery

for Auxi» spp. off Ashizurimisaki, Japan (Ishida 1972a).

along the southern coast which peaked at 21.3 fish/boat- day. Fish appeared to be fewer along the east and west

coasts with catch rates of 18.9 and 14.3 fish/boat-day, re- spectively. The lowest catch rate occurred off the north

coast with only 4.2 fish/boat-day. For A. rochei, the

variation in catch rates followed a similar pattern. Si- vasubramaniam (1973) estimated that usually a fishing

vessel will encounter two to three mixed schools per day

in waters around Sri Lanka.

47
4.3 Natality and recruitment

4.31 Reproduction rates An A. thazard of moderate size (44.2 cm) produces about 280,000 eggs/spawning and about 1.37 million eggs are extruded over the entire spawning season (Rao

1964). From Auxis caught in drift nets in Indian waters,

Silas (1969) also estimated their fecundity and con- cluded that A. thazard {n = 9) spawn from 197,000 to 1,056 million eggs and average 601,000 eggs/spawning;

for A. rochei (n = 4), he found fewer eggs per spawning.

Silas estimated that females spawn from 31,000 to 103,000 eggs and average 52,000 eggs/spawning.

4.33 Recruitment

Off Sri Lanka, major recruitment to the exploitable stock of A. thazard occurs annually along the south and southwest coasts between March and August

(Sivasubramaniam 1973). There is also evidence that sporadic recruitment occurs during the rest of the year. Off the east coast, recruitment has been observed in July-September, but no clear trend in recruitment has been found in catches along the north coast. For A.

rochei, Sivasubramaniam conjectured that recruitment probably takes place at about the same time as that for A. thazard. Therefore, the exploitation of new recruits

starts around the middle of the year then shifts to large

size classes until the following year's recruitment. The time of recruitment influences the annual pro- duction of Auxis in Sri Lanka (Sivasubramaniam 1973). When recruitment occurs during the southwest monsoon, the schools that enter the fishery are available at a time when the relative abundance of skipjack tuna, yel- lowfin tuna, and kawakawa is low and frigate and bullet tunas are fished until the beginning of the northeast monsoon. But when recruitment occurs late in the year, the fishing effort expended on these two sp>ecies is less intense because of the presence of more desirable species such as skipjack and yellowfin tunas. Thus, fishing emphasis shifts from trolling to pole and line, longline, and

drift netting. November-March marks the peak fishing season for the skipjack tuna pole-and-line fishery in the south and southwest coasts of Sri Lanka. It is at this time that large Auxis (40-50 cm), usually mixed with

skipjack and yellowfin tunas, show up frequently in the

catch.

4.4 Mortality and morbidity

4.42 Factors causing or affecting mortalty An instance of natural mass mortality of larval Auxis

in Hawaiian waters has been reported by Strasburg

(1959). Noting the presence of two distinct conditions of

larval Auxis in the preserved plankton samples collect- ed in the vicinity of Lanai (an island in the Hawaiian

chain), Strasburg hyjxithesized that these conditions represented fish that were alive and dead just prior to

capture. In a simple experiment, he placed several fresh, dead specimens in dishes of seawater and allowed them

to decompose. Other fresh, dead specimens were pre- served in Formalin. The results showed that fresh pre- served specimens were clean cut and normal in ap- pearance whereas dead, deteriorated specimens were

imperfect. Strasburg concluded that large numbers of

larval Auxis in the samples were dead before capture, but no cause of death could be isolated. One pKtssible cause of death is the passage of the larvae through an

area of marked temperature discontinuity in the surface layer.

5 EXPLOITATION

5.1 Fishing equipment

5.11 Gears

Pole and line is the most commonly used gear to catch

Auxis. In Japan, pole-and-line gear designed exclusively for fishing A. rochei differs from that used for skipjack tuna fishing. According to Yasui the pwle-andline gear for bullet tuna consists of a bamboo pole, nylon monofilament, trolling board, and jig, and is attached at the butt end of a tag line to the vessel so that the entire gear is "trolled" at a speed of 2-3 nmi/h while searching

for bullet tuna schools (Fig. 49). Two to three fishermen manage the "trolled" poles at the stem. When a fish

M. Yasui, Shizuoka Prefectural Fisheries Experimental Station, Shizuoka. Japan, pers. commun. December 1975.

NYLON MONOFILAMENT NO. 10 2 5m

-) TROLLING BOARD

Figure 49. —Diagram of the pole-and-iine gear used and the method of fishing in the fisher}' for Auxia spp. in Japanese water

(lahida 197!). 48
strikes the lure, the fisherman pulls on the tag line, re- trieves the pole, and flips the hooked fish aboard. A mixture of dead, salted "shirasu" (larval fish) and boiled noodles (somen) is chummed over the side while the boat circles the school counterclockwise, thus keeping

the boat close to the dead shirasu and boiled noodles which are pushed toward the school by the rotation of the propeller. When biting is fast, the poles are no longer

"trolled"; rather, each fisherman grasps one pole and

flips the hooked fish aboard. The use of other commercial gear depends on availability of Auxis and the efficiency of the gear. Among

them are trolling line, handline, small-scale longline, and a wide assortment of nets including traps (set net, pound net), gill or drift nets, bag net, ring net, beach

seine, otter trawl, and purse seine. In some countries where these gears are in use, Auxis are not the principal

species sought by the fishermen; rather, they are caught when other species of tuna such as yellowfin, bluefin, and skipjack tunas either become unavailable or are

relatively low in abundance. Some of these gears are discussed in the section that follows. In the Philippines, purse seines appear to be effective

in taking Auxis. To accelerate commercial develop- ment of pelagic fisheries, chartered commercial purse

seiners, and research and survey boats under the South China Sea Fisheries Development and Coordinating

Programme have been doing exploratory fishing includ- ing the detection and estimation of the abundance of the

pelagic resources. Rosenberg and Simpson, reporting on

voyage 3 of the purse seiners Royal Venture and South- ward Ho, stated that during the period when both worked the southeasterly portion of Moro Gulf, some of the floating logs seen had good signs of tuna. About 26,0- 00 kg of "Tulingan" (probably Auxis) and kawakawa

were landed, composing roughly 21'~r of the landings of the two seiners. Rosenberg and Simpson noted that

large quantities of kawakawa and Auxis are consumed

fresh in the Philippines, but there is no market for them when frozen. The total landings of tuna by the two seiners after voyage 4 reached 31,000 kg of which Auxis represented about ll'f. Southward Ho usually caught less Auxis

than the Royal Venture because the former used a

slightly larger mesh in the bunt section of the seine (Rosenberg et al.'). The result was that Auxts, which are smaller than either skipjack or yellowfin tunas, tended

to gill and had to be discarded.

In Indian waters, A. rochei occasionally occur in very

large schools close to shore and large quantities are usually taken by beach seines (Jones and Silas 1964).

'Rosenberg. K. J., and A. C. Simpson. 197.5. Pelagic fisheries devel- opment—Trip reports, chartered purse seine vessels. (Trip reports of chartered purse seine vessels Royal Venture and Southward Ho, covering Voyage .3. 9 February' to 26 March 1975. ( South China Sea Fish. Dev.

Coord. Programme. Manila. .June 197.5. SCS/7,5AVP/10, 28 p. 'Rosenberg, K. •!.. A. C. Simpson, and C. M. Rcnwick. 197,5. Pelagic

fisheries development —Trip reports, chartered purse seine vessels. (Trip reports of chartered purse seine vessels Royal Venture and Southward Ho, covering V'oyage 4. 9 April to 24 May 197,5.) South China

Dev. Coord. Programme. .Manila. -June 1975. SCS^5AVP12, 36 p.

Silas (1967b) rejaorted that Auxis are very rare in the

multiple troll catches made along the Tinnevelly coast

in the Gulf of Mannar but are caught in small numbers

together with skipjack tuna on pole-and-line and troll gear in the Laccadive Islands.

India, like the Philippines, has also been active in upgrading her tuna fishing capability. In January 1962, the Indo-Norwegian Project made the RV Varuna available

to the Central Marine Fisheries Research Institute of In- dia for exploratory fishing and oceanographic surveys in the Indian Ocean (Silas 1969). Two types of gear—drift net and purse seine—were used. Each unit of nylon drift net was 25.85 m long and 6.10 m wide. Five mesh sizes were used—2.5, 5.5, 10.0, 12.5, and 17.0 cm. The smaller mesh sizes captured juvenile pelagic fishes whereas the

larger meshes captured A. t hazard tmd A. rochei in both

the continental shelf and oceanic areas. Figure 42 shows

the number of drift net sets, the number of specimens

collected, the average weight, and the length-frequency

distribution of Auxis in the tuna catch.

Initial attempts at exploratory purse seining in the Laccadive Sea by the RV Varuna were unsuccessful, but

subsequent attempts by the MV Tuna were reasonably

successful (Silas 1969). The purse seine used in the ex- ploratory work was 540 m long and 67 m wide with a mesh size of 10 cm. Out of 38 purse seine sets, made with

a net 540 m long, 67 m wide, and mesh size of 10 cm, 10 were successful and the maximum catch exceeded 10 t in one instance. The catch was composed mainly of schools of E. affinis and A. thazard. Figure 43 shows the

locations of the purse seine sets, catch per set, and sizes caught. Silas noted that the average weight of A. rochei was only 0.168 kg compared with an average of 0.750 kg

for A. thazard. Of these early attempts at tuna purse

seining in Indian waters, Sileis remarked that they were

not highly successful because the sinking rate of the net was too slow thus allowing the schools to escape. Also, he noted that the crew lacked experience in purse seining, but the failures, he felt, should not deter further at- tempts at purse seining.

In Sri Lanka, five types of gear are used to exploit tuna—longline, po\e and line, trolling line, drift net, and

beach seine. The longline is used for Ifirge, deep- swimming tunas and will not be discussed here. The re- maining four are used in varying degrees to catch Auxis. The beach seine is not a selective gear for Auxis;

therefore, the catch by this gear is insignificant (Sivasubramaniam 1965, 1973). Beach seining, requir- ing about 30 men to conduct successfully, is possible in places like Tangalle, Udapu, and Trincomalee, because

schools of Auxis occasionally migrate close (within 0.9 km) to shore. The most important method of capturing Auxis off Sri Lanka, however, is by trolling. The gear is adapted from salmon trolling gear and is unlike the standard tuna

trolling gear used extensively by Japanese and United

States fishermen (Sivasubramaniam 1965). Consisting

of three mainlines each with 30 branch lines which end

in jigs, the trolling gear is well suited for Sri Lanka's

waters because of the denseness of the schools of Auxis

49
and kawakawa (Sivasubramaniam 1973). Trolling, usually limited to 3-5 h in the morning when Auxis

schools are at the surface, is effective during periods of monsoon when the water is turbid. The gear was formerly operated from "orus" (outrigger canoes with sail), but

catches were poor because of difficulty in maneuvering

the vessel. Nowadays, 3.5-gross ton motorized boats, manned by crews of at least three fishermen, are used and their efficiency have made them popular for fishing the various species of tunas. Trolling contributes over 50% of the total Auxis production.

Pole-and-line fishing boats in Sri Lanka concentrate

primeirily on skipjack tuna but also catch significant quantities of other tunas including kawakawa, yellowfin tuna, and Auxis (Fig. 50). Sivasubramaniam (1965)

jxjinted out that Sri Lanka's pole-and-line boats operate

gear similar to that used by fishermen of Minicoy,

Laccadive, and Maldive Islands. The boats, manned by

at least five fishermen, carry live bait in cane baskets

that are partially submerged alongside the craft. Wooden spades are used for spraying water. Originally,

p)ole-and-line fishing was restricted to waters off the southwest coast of the island, but mechanization of the boats has expanded this type of fishing to the east coast. Like other pole-and-line fisheries operating in the In- dian Ocean, however, the availability of live bait has

limited the popularity of this fishing method. Although pole-and-line fishing is effective for skipPOLE AND LINE

a n

TROLL n

r^

LONGLINE

(

J V////A BEACH SEINE

jack tuna, it is not equally effective for Auxis

(Sivasubramaniam 1973). The reason is that Auxis re- spond poorly to chumming probably because the live bait used are usually larger than the most common food items in their stomachs. Furthermore, this type of fish- ing is not effective during periods of southwest monsoon, because water turbidity makes baiting difficult. The best season for pole-and-line fishing is in No- vember-March off the southwest coast and in July-Sep- tember off the east coast. Tuna production depends

heavily on the availability of bait during these periods.

Drift nets are operated around the entire island of Sri Lanka (Sivasubramaniam 1973). A 3.5-gTOss ton boat

usually carried 15 pieces of netting whereas as 11-gross ton boat carried 60 pieces. With mesh sizes varying from

10.2 to 14.0 cm, the synthetic drift net has proven effec- tive for skipjack and large kawakawa. For Auxis, however, fish <30 cm in length are too small to be enmeshed. The catch of Auxis, therefore, amounts to only about

18 kg/day for a 3.5-gross ton boat and about 54 kg/day

for an 11-gross ton boat. Auxis rochei, being smaller, are very rare in drift net catches. Auxis are usually taken by

drift net from about September to March.

5.12 Boats

As with gear, a wide variety of fishing boats, ranging from modern high-seas purse seiners and pole-and-line boats to primitive sailing crafts, is used in fishing for Auxis.

In Japanese waters, the pole-and-line boats that are engaged in theAuxis fishery are small, usually <15 gross tons (Ishida 1972a). These boats are smaller than the usual pole-and-line boats of >30 gross tons which fish in the coastal waters and high seas for surface schools of tunas such as skipjack tuna, yellowfin tuna, and albacore. Japanese pole-and-line boats that fish in the Atlantic for yellowfin and skipjack tunas and occasionally

for Auxis and bigeye tuna vary from 151 to 239 gross

tons, whereas the seiners, both single and double boats, range from 50 up to >400 gross tons (Borgstrom 1964; Hayasi 1973, 1974). Among the more primitive types of crafts used in Auxis fishing are dugout canoes and catamarans (Silas 1967b). In the Maldive Islands, for example, sailing boats called vadu dhony, which are about 6.1 m long and 1.8 m wide, and fish three to four baited trolling

lines, are used as trollers (Sivasubramaniam'"). Slightly

larger sailing crafts, called the mas dhony, are used in pole-and-line fishing; they vary in length from 10.7 to

12.2 m, have a beam of about 3.4 m and a draft of nearly

0.8 m. The Maldivian boats are constructed of coconut wood, beautifully streamlined, and keeled for wind- ward sailing. Compartments for carrying live bait have

continuous water circulation through holes along the bottom. The fishermen fish at the stem atop a U-shaped

removable wooden platform.

Figure 50.—Percentage composition of tuna species caught by

various types of gear from the coastal waters of Sri Lanka (Siva- subramaniam 1%5).

'"K. Sivasubramaniam. UNDP-Sri Lanka Skipjack Fishery Develop- ment Project. FAO, Colombo, Sri Lanka, pers, commun. August 1975.

50
In 1972, the Maldivian fishing fleet had an estimated

5,100 fishing boats which included 2,980 trollere and

2,100 pole-and-line tuna boats ojjerating within a 46 km

radius of the islands. Skipjack tuna, yellowfin tuna, kawakawa, and Auxis are the main species caught by

the pole-and-line boats. Hiebert and Alverson (1971)

p)ointed out that one of the factors hindering the development of the Maldivian tuna fishery is the

limited access of the boats to more distant fishing grounds. Lack of refrigeration also precludes long trips; therefore, the tuna boats leave port for the fishing grounds at about 0300 and return late in the evening. The introduction of motorized, refrigerated boats can

probably increase the present fish catch 3-4 times.

In waters around Sri Lanka, trolling gear was formerly operated from "orus" or outrigger canoes with sails. Nowadays, however, 3- to 5-gross ton motorized boats, manned by a crew of at least three fishermen, are used (Sivasubramaniam 1973). In the Canary Islands where

Auxis are landed along with other species of tuna, there

are large boats (average 400 gross tons), which have re- frigerated holds and carry about 20 men, and smaller,

artisanal fishing boats (average 10 gross tons), which are crewed by 4-5 fishermen, are about 12 m long and have open decks and bait tanks. Propelled by engines that vary in size from 50 to 120 hp, these small boats usually engage in pole-and-line and occasionally purse seine

fishing. Off Angola, yellowfin and skipjack tunas are the

principal species caught by the small, pole-and-line

boats, but Auxis are also landed in smaller quantities (De Campos Rosado 1972). These boats, which operate

within a narrow strip about 75 km wide and 370 km long along the coast, vary in size from 12 to 20 m long. The Cuban pole-and-line fishery, which lands small amounts of Auxis, operates motorized sailing boats, which have been described as modified sloops with a

gaff-rigged mainsail and usually with a flying jib (Rawlings 1953; Wise and Jones 1971).

5.2 Fishing areas

5.21 General geographic distribution

Almost all the Auxis caught in the Japan Sea are A.

rochei (Okachi 1958). Percentagewise, Okachi deter- mined that of the fishing areas established for Auxis in Japanese waters, the South Pacific region had 62% of the catch, the middle Pacific region 22%, and the Japan

Sea and East China Sea regions combined only 12%. Catches from west Japan Sea and East China Sea

regions were usually similar whereas catches from the north Japan Sea region were smaller than either one.

In other areas of the world, Auxis are usually caught

as incidental species. In the Pacific, for example, Auxis

are caught incidentally in waters around Taiwan, the

Philippines, Thailand, West Malaysia, Hawaii, and

Australia (Serventy 1941; (josline and Brock 1960; Kume 1973; Philippine Bureau of Fisheries 1973). Small numbers of Auxis are also taken in the Indian Ocean.

Jones (1967) reported that Auxis are captured along the

coasts of India and in waters surrounding the Maldive

Islands (Sivasubramaniam footnote 10). According to Sivasubramaniam (1973), Auxis are caught in waters around Sri Lanka on a commercial scale throughout the year in the south and southwest coast of Sri Lanka.

In the Mediterranean region, the waters around

Cyprus, Greece, Italy, Malta, Morocco, Spain, and

Yugoslavia are fishing grounds for Auxis. Spanish trap- net fishermen land commercial quantities of Auxis at Barbate on the Atlantic Spanish coast near the Strait of

Gibralter, at Tarifa in the middle of the Strait, and at La Linea on the Mediterranean Spanish coast near the entrance of the Strait. Some fishing is also carried on around the Canary Islands (Bas 1967; Cendrero and

Garcia-Cabrera 1972). Auxis are taken in Moroccan waters mostly by trap fishermen. Traps located at Larache and at Cape

Spartel are set primarily to catch bluefin tuna that make their spawning migration toward the Mediterra- nean Sea along the Moroccan coast in April-July. Traps

at M'diq in the Mediterranean also capture tuna as they

exit after spawning. In traps located along the Atlantic Moroccan coast, Auxis account for only 3% of the tuna

landed, but in those situated along the Mediterranean Moroccan coast, they represent 98% of the tuna catch (Lamboeuf 1972). Around Portugal, the tuna fishing grounds are along

the north and west coasts where small pole-and-line vessels operate. Other grounds fished by Portuguese fisher- men are located off the islands of Azores and Madeira

where small- to medium-sized pole-and-line vessels operate, and off Angola (De Campos Rosado 1972; Dias and Barraca 1972). Catches of little tunny, bonito, and

Auxis represent roughly 60% of the total tuna landings from Angola. Off Ghana, in the Gulf of Guinea, there is another

fishing ground for Auxis. Although most of the tuna

landings at Tema are made by foreign longline, pole- and-line, and purse seine vessels, Ghana's canoes and

motorized fishing crafts also account for some of the landings of tuna and tunalike fishes (Di Palma 1968). In the western Atlantic, Auxis were previously taken

along the east coast of the United States. Pound nets

fished along the Middle Atlantic States during the

1940's and early 1950's frequently caught Auxis along with bluefin tuna, little tunny, and bonito (Anderson et

al. 1953). Southward, the waters around Cuba, off Venezuela, and off Brazil also yield small quantities of Auxis.

5.22 Geographic ranges

See section 5.21.

5.23 Depth ranges

Adult A. thazard has been reported to distribute themselves vertically from the surface down to a depth

of about 45 m (KisKinouye 1923).

51
5.3 Fishing seasons

5.31 General pattern of seasons The average monthly landings of Auxis at Tosashimizu (Kochi Prefecture), at Hachijo (Izu Archipelago), and at Mera (Shizuoka Prefecture) are shown in Figure 51. At Tosashimizu, where a pole-andline fishery operates, Auxis are caught throughout the

year, but peak fishing usually extends from October

through May (Yasui 1975). During the remainder of the year (June-September), some of the pole-and-line boats concentrate on other species; therefore, there is a reduction in fishing effort for Auxis.

In the region of the South China Sea, the fishing season for Auxis varies considerably. For example, in the Philippines, the fishing season varies for different parts of the island group beginning at any time between November and January and extending into May

(Philippine Bureau of Fisheries 1973). In Thailand and West Malaysia, however, the landings of /luxjs, Euthynnus affinis, and Thunnus tonggol are usually better during the latter half of the year (Kume 1973). Fishing for A. thazard in Indian waters begins about August and extends to about December (Nair et al. 1970). Sivasubramaniam (1973), who examined the

l-l
Figure 52. —Distribution of surface temperatures in July- September in years of good (A) and poor (B) fishing (or A axis rochei at Tosashimizu. Kochi Prefecture, Japan (Y'asui 1975).

fluenced to a large extent by a seasonal in-migration of

fish from the west and south. At Hachijo, where Auxis are landed by surface troUers, catches are made throughout the year except in October (Fig. 51). Yasui (1975) observed that there are two

seasons for Auxis at Hachijo—one in April-August and

another in November-January. The appearance of A uxjs

in April-May at Hachijo coincides with the end of the

gill net fishery for flyingfish, which are preyed on by

Auxis. Yasui also noted that sexually mature fish appear

in the catches in April-May followed by larger, older fish

in .July. At Ashizurimisaki in Kochi Prefecture, Ishida (1972b) noted considerable fluctuations in landings of Auxis. To

determine the possibility of predicting the degree of success of a fishing season, Ishida, using data for 1952-67., obtained the following:

C, = catch in October-March in year;

Cj+, = catch in October-March in year; + 1 ACjti = difference between Cj and C,»i.

Plotting C, against ACj»,, Ishida found that when C, was small, AC, + , tended to be positive and vice versa. Ishida also noted that larger catches of Auxis tended to be associated with cooler temperatures. Yamashige

(1974), on the other hand, found a positive relatonship between catches of A. rochei in August-July and the dif- ference in the mean surface temperature between August and October of a given year, and he described

the relationship between these two variables as follows: y = 1.42 + 1.15 ((g - t,o)

where Y = catch, in 1,000 t, from August {n year) to July (« + 1 year)

tf^ = mean surface temperature in August at Ashizurimisaki (°C).

t^^ = mean surface temi>erature in October at Ashizurimisaki (°C).

In Sri Lanka, the mechanization of fishing boats made fishing jjossible even during periods of monsoons and has been primarily responsible for a shift in the fish- ing season. Originally, Williams (1963) reported that the main fishing season for Auxis in Sri Lanka extended

from October or November to May. Sivasubramaniam

(1973), however, found that Williams defined the fish- ing season by examining catches made during the northeast and intermonsoons when beach seines are usually in operation. Excellent fishing occurs in all fishing areas around Sri Lanka particularly during the end of the southwest monsoon. Sivasubramaniam observed that

this period is associated with heavy recruitment and

vulnerability of the recruits to trolling gear. The result is that the number of fish caught per day rises abruptly, but the net tonnage landed, does not rise proportionately.

5.4 Fishing operation and results

5.41 Effort and intensity

In the Auxis fishery off Ashizurimisaki in Kochi Prefecture, Japan, fishing intensity (the number of small,

3-gross ton pole-and-line boats operating per day) varies widely from as few as 10 to as many as 260 boats/day

(Ishida 1972a). Fishing intensity usually peaks once in March and again in November to about 200 boats/day. During the balance of the fishing season, which extends

usually from October to May, however, the average number operating is about 170 boats/day.

5.42 Selectivity

Although pole-and-line fishing is effective for skip- jack and yellowfin tunas, it is not equally effective for Auxis. Sivasubramaniam (1973) observed that in Sri Lanka, Auxis respond poorly to chumming probably

because the live bait used is larger than the most com- mon food items in their stomachs. The drift net, which is in operation around the entire island of Sri Lanka, is also ineffective at times particularly for Auxis <30 cm. It is, however, effective for skipjack and large kawakawa. Catches of Auxis by drift nets, therefore, amount to only about 18 kg/day for a 3.5- 53
gross ton boat and about 54 kg/day for an 11 -gross ton

boat. Auxis rochei, being smaller, are very rare in drift net catches.

5.43 Catches

Total catches of frigate and bullet tunas by various countries in the Atlantic (including the Mediterranean

Sea), Pacific, and Indian Oceans in 1953-76 are given in Table 26. By far, the total catch in the Pacific was the

largest among the three oceans, averaging 40,700 t annually. The Atlantic and Indian Ocean catches, on the other hand, averaged about 13.8 1 in each ocean or about

a third of the Pacific catch.

In the Report of the Ad Hoc Committee Meeting of

Specialists to Review the Biology and Status of Stocks

of Small Tunas, it was pointed out that the catches of Auxis reported by the various countries do not truly re- flect the abundance of the two species in the world's ocean. For example, purse seine fishermen avoid

catching Auxis because they are a nuisance as many

become gilled in the net and require considerable time and manpower to remove. Furthermore, the catches of Auxis by tuna seiners are often discarded at sea. Other

difficulties noted with Auxis catch statistics are that ar- tisanal catches are usually not fully reported and that

there is confusion and inclusion of Auxis with other

spwcies.

Pacifc Ocean

Japan lands, by far, more Auxis than any other country and is, in fact, the only country that has a well- established commercial fishery. In 1953-76, the catch of Auxis averaged 24,300 t, varying from 14,900 1 in 1975 to 48,300 t in 1963 (Table 26). In Taiwan, the catches of frigate and bullet tunas in 1961-73 varied between 600 and 1,200 t and averaged

1,000 t (Table 26). Perhaps it should be pointed out that

there is a discrepancy in the catch statistics for frigate

Table 2(i. —Estimated catches of Auxis sp. by countries, l!).5.!-76 (in units of 1.(1(10 metric tons). (Data compiled from FAO 1959, I9()4. 19fi9, 1974.

1977; Lamboeuf 1972; Miyake and Tibbo'; Miyake et al.": Sivasubramaniam'; Hagborg.') — = data not available; 0.0 = magnitude negligible or

insignificant. Atlantic Ocean (including Mediterranean Sea) Pacific Ocean Indian Ocean

Z coo

O ii •? S S £ 23

1953
and bullet tunas from Taiwan for 1961-68. Examination

of the catch statistics from Taiwan for 1958-63 (FAO

1964) showed catches to be rather high. In 1961-63, for example, they were as follows:

Year Catch (1,000 metric tons)

1961 11.9 1962 18.2 1963 15.1

But in FAO (1967), the frigate and bullet tuna catches

in l%l-63 were given as follows:

Year Catch (1,000 metric tons)

1961 0.7 1962 0.8 1963 0.6

Further research into the method of compiling catch

statistics indicated that data from FAO (1964) included

other species under frigate and bullet tunas. Kume

(1973) revealed that Taiwan uses a categroy called "bonito" in which are included Auxis as well as kawa- kawa and perhaps other small tunas. Kume gave the "bonito" catch for 1970 as 15,500 t but estimated that of

this total, only 778 t or roughly 5% were Auxis. Further- more, the frigate and bullet tuna catches in 1961-63, as given in FAO (1967), amounts to only 4-6% of the

catches as given in FAO (1964). Therefore, it can be presumed that roughly 5% of the catches of frigate and

bullet tunas in Taiwan, as given in FAO (1964), are ac- tually of this species with the remainder constituting catches of other small tunas. Southward, in the South China Sea region, the Philippines, Sabah, Sarawak, Thailand, and West Malaysia

also harvest Auxis. In the Philippines, catches of Auxis

in 1964-76 varied from 5,000 t in 1974 to 19,500 t in 1968 and averaged 11,700 t (Table 26). The Thailand

landings of Auxis, Euthynnus affinis, and Thunnus

tonggol are combined and called "bonito" or "pla o"

locally in the catch statistics (Kume 1973). In 1971, Thai and Chinese seiners landed 5,090 t or 78% of the 6,548 t of "bonito" reportedly caught in Thailand. In West

Malaysia, landings of Auxis are also combined with

those of E. affinis and T. tonggol. The 1958-71 production of these three species combined varied between 789

t in 1959 and 5,578 t in 1967 and averaged 3,131 t annually. Table 26 shows that the catches of Auxis from China mainland varied from 2,400 to 5,300 t in 1964-71 and averaged 3,700 t.

Indian Ocean

In addition to E. affinis and T. Tonggol, which are the two most common tuna species landed from Indian

waters, small numbers of Auxis are also captured in the tuna fishery along the coast of India (Jones 1967). Of the two species, A. thazard are more common whereas A.

rochei are rarely seen in the commercial catch. Thomas

(1967) reported that Auxis are taken in the Laccadive

Islands, but catches are usually recorded in the

category, "other fishes," which includes E. affinis, Acanthocybium solandri, Istiophorus gladius, Elagatis

bipinnulatus. Caranx sp., Chorinemus sp., and sharks. Sivasubramaniam" estimated that Auxi.s catches in In- dia account for about 5% of the tuna and tunalike

catches. It is also apparent from the data that the west

coast of India usually yields Auxis catches that are about 5 times more than those from the east coast. In 1964-73, India's Auxis catches varied between 200 and

7,900 t and averaged 2,100 t annually (Table 26). The tuna fishery in the Maldives is carried out by sail- ing boats; therefore, fluctuations in the catches are asso- ciated with various environmental conditions such as wind direction and velocity and ocean surface currents.

In 1964-76, the catches of Auxis from the Maldives fluc- tuated between 1,500 t in 1964 and 6,200 t in 1973 and

averaged 3,200 t annually (Table 26.) There are, however, serious discrepancies in the catch statistics. For

1970 and 1971, for example, Sivasubramaniam (see foot- note 11) gave the catch of Auxis from the Maldives as

1,700 and 1,800 t, respectively. Data from a tuna can- ning plant project, however, indicate catches of 3,094 t in 1970 and 26,871 t in 1971. Based on landings in 1965- 69 and in 1972-73, it appears that the 1971 catch of 26,871 t is unreasonably high. Furthermore, FAO (1974) estimated the 1970 frigate and bullet tuna catch to be

20,000 t and the 1971 catch as 26,900 t, also unreasonably high; therefore, until these discrepancies can

be resolved, data from Sivasubramaniam (see footnote

11) will be accepted provisionally.

In waters around Sri Lanka, Auxis are caught on a commercial scale from all the main fishing grounds

(Sivasubramaniam 1973). Being the smallest member of the tuna and tunalike fishes, Auxis contribute only 15- 20'r to the total catch, by weight, but are the most abundant of all the tuna varieties in the waters around

Sri Lanka. Sivasubramaniam reported that although both species appear in the commercial catches, A. thazard contributes 92% and A. rochei 8% to the total annual production. In 1964-73, Sri Lanka's production

of frigate and bullet tunas ranged between 3,200 and

5,.500 t and averaged 4,500 t annually (Table 26). Per- centagewise, they constituted more than a third of the catch of tuna and tunalike fishes from Sri Lanka in 1964-67. The proportion fell to about a fourth in 1968-69, one fifth in 1970, and has been about one-sixth of the

total production since 1971. Other countries that harvest Auxis in the Indian Ocean include Bangladesh, Pakistan, Reunion Island, and Tanzania. Table 26 shows that in 1964-72, Auxis

catches in Bangladesh fluctuated between 100 and 800 t and averaged 400 t. In Pakistan, catches in 1964-73

"Sivasubramaniam, K. 1974. More recent information on the tuna

fisher>' in Sri Lanka, India and the Maldive Islands. Indo-Pacific Fishery Council/Indian Ocean Fishery Commission Ad Hoc Working Group,

.\antes. France, 16-18 September 1974. (Il'FC/IOFCAVPU4/12, 4 p.

55
varied from 1,000 to 5,600 1 and averaged 2,900 1 annually. Catches from the Reunion Island, recorded for 1970-

72, are relatively small, varying between 100 and 200 t and an annual average of about 200 t. Tanzania landed

400 t in 1971 and 600 t in 1972 and averaged 500 t for these 2 years.

Atlantic Ocean

In the Atlantic, Spain, which annually lands about

3,500 t of Auxis, exploits mackerel. Scomber scombrus;

bluefin tuna; bonito, Sarda sarda; and occasionally albacore along the Mediterranian Spanish coast (Bas

1967). Mackerel are caught by ring net and trawl; bluefin tuna, bonito, and Auxis are caught occasionally by

very large ring nets. The annual production of Auxis by

large ring nets is small, ranging between 274 and 2,500 1. The most important gear is the trap. At Barbate, A.

rochei appear in the traps in May-July and catches average 15.6 t but larger catches usually occur in MayJune (Rodriguez-Roda 1966). Little tunny and bonito

are also taken at Barbate, but A. rochei predominate,

accounting for about 53% of the production. The traps

at La Linea make the largest catches of these small tunas with A. rochei constituting about 97'~c of the catches (216 t). At Tarifa, 95'~'c of the catch is A. rochei and production averages about 67.1 t. The annual

catches of A. rochei by traps throughout Spain vary

widely. Catches at Barbate in some years are very low and fall below 2,000 fish/yr but in other years they may

be very high. Table 26 gives the annual production of Auxis in Spain and Table 27 gives a partial breakdown

of the catches by type ot gear. In 1953-76, the annual

landings varied from 500 to 10,000 1 and averaged 3,100

(Rt>driguez-Roda 1966, 1967). Auxis are taken in Moroccan waters mostly by trap

fishermen, but surface gear is also employed (Table 27). Lamboeuf (1972) stated that both bait boats and seiners operate as a team in fishing. The bait boat, which is usually an old, low tonnage sardine boat converted to carry bait, chums fish to the surface and keeps them

there long enough for the seiner to set the net around the

school. Lacking such a bait boat frequently results in an

unsuccessful purse seine set. Along the Atlantic Moroccan coast, boats landed 69% of the Auxis whereas traps contributed less than half or 31% (Lamboeuf 1973). Along the Mediterranean Moroccan coast, however,

traps contributed 91% of the Auxis landed with the remaining 9% produced by boats. In 1953-76, the annual

catches of Auxis fluctuated between 100 and 4,000 1 and

averaged 1,400 t (Table 26). Auxis landings in Morocco

rose from about 15% to 20%; of the total tuna landings in 1963-65, fell to about 12% in 1966, then rose steadily to about 27^0 of the total catch in 1969 (Lamboeuf 1972). The change in the relative importance of Auxis in the

total tuna landings resulted from a very sharp decline in bluefin tuna landings in Morocco. It should be noted

that slight discrepancies exist in the catch data for Auxis; only 200 t in Table 26 versus a 1969 catch of 588 1, according to Lamboeuf (1972).

In Portugal, traps, which are the principal tuna-fishing gear, are usually fished off the southern coast whereas pole-and-line boats operate along the north and west

coasts and off the islands of Azores and Madeira (Dias and Barraca 1972). In the Portuguese overseas province

of Angola where the yield of tuna is much higher, the

fishery is carried on by small, pole-and-line boats (De

Table 27.—Annual catches o( Auxis (in units of 1.000 metric tons) In the Atlantic Ocean, by country and type of gear. 1960-74
Campos Rosado 1972). Landings of Aujris from Angola

in 195.i-76 varied from 500 to 7,300 t and averaged 2,000

t (Table 26). Ghana's modern fishing fleet and industry, which

developed only since the early 1960's, are probably the

best among African nations (Di Palma 1968). Di Palma

observed that confusion exists in the catch statistics on

mackerels and scads because some boats do not differentiate between the various species. For example, several species may be reported together including mackerel.

Scorn her japonicus, various species of horse mackerels of the family Carangidae, Auxis, and mackerel scad, Decapterus rhuncus. Miyake et al.'" reported that Ghana's landings of Auxis by unclassified gear varied between 1,800 and 5,300 t and averaged 3,160 t in 1968-

72. In 1965-76, Ghana's annual catch of Auxis varied from 1,600 to 6,300 t and averaged 3,400 t (Table 26). Japan's resurgence as a fishing power after World War

II was not restricted to the Pacific and Indian Oceans. Japanese tuna longliners first made their appearance in the Atlantic in 1957 and by 1961 at least 60 vessels were

fishing there for yellowfin tuna and albacore (Borgstrom

1964). In 1962, the Japanese placed pole-and-line fishing boats of 239 gross tons each in Ghana to fish for surface

schools of tuna. Yellowfin and skipjack tunas are the main species landed by these Japanese bait boats but Auxis and bigeye tuna are also taken in limited quantities (Shomura 1966; Hayasi 1973). Three or four Japanese purse seiners and five to seven pole-and-line

(14 in 1972) boats operated in the Atlantic Ocean in 1962-63 and in 1967-72. Tema, with excellent shore facilities, including cold

storage, serves as a base for Japanese purse seiners, pole-and-line boats, and longliners (Di Palma 1968). Auxis as well as yellowfin, skipjack, and bigeye tunas

are caught along the coast of Africa and in coastal waters of the offshore islands in the eastern Atlantic.

In 1967-73, Japanese seiners landed from 177 to 1,256

t of Auxis and averaged 670 t annually (Hayasi 1973,

1974). The proportion of the purse seine tuna catch consisting of Auxis, however, was small varying from 3'r to

15^f in 1969-72. Auxis catches by Japanese pole-and-line boats also fluctuated widely in 1967-70 varying between 675 and

3,200 t and averaging 1,375 t annually (Hayasi 1973,

1974). The proportion of the pole-and-line tuna catch which constitutes /I uxi.s varied widely from an insignifi- cant part of the total pole-and-line tuna landings in 1972

to nearly one-third in the 1969 catch. Japanese boats fish not only in the eastern but also in the western Atlantic Ocean. The Caribbean region is an

important fishing ground for tuna and tunalike fislies including Auxis. Wise and Jones (1971) pointed out that

the catches of several species of tunas and related fishes occurring in the Caribbean region are shared by several

Miyake. M. P.. A. Ue Boisset, and .1. .Manning (compilers). 1974.

Statistical bulletin, international Commission for the Conservation of Atlantic Tunas, Vol. 4. ST/Total/74/2 - 13-15 Rev. |No pagination.!

countries. Japan lands about two-thirds, Cuba and

Venezuela each take about one-tenth, and the remaining countries, each with <2,000 t annually, land the balance of the catch. Among the species landed, yellowfin tuna is the most important, contributing slightly over a third of the total landings. Second most important is the "bonitos" but this category includes few true bonitos and is made up largely of skipjack tuna and

blackfin tuna, Thunnus atlanticus, with small amounts

of other species such as little tunny and Auxis. Bigeye tuna is third in importance followed by bluefin tuna and

albacore. The total landings of Auxis by all Japanese

fishing boats operating in the Atlantic varied from 400

to 3,200 t and averaged 1,200 t (Table 26). In Venezuela, scombrids of commercial interest are Sarda sarda, Auxis, Scomberomorus maculatus (= S. brasiliensis) , S. caualla, and Scomber japonicus (Griffiths 1971). The catches of Sarda and Auxis, primarily by baited hooks ("cordel") and gill nets along eastern Venezuela, are grouped in the official statistics into "cabanas." Unfortunately, the proportion of each

species in this category is unknown. Griffiths noted that the proportions may be variable among the fishing areas and also with time. "Cabanas" are usually sold fresh

locally or salted and sometimes canned as "tuna" or "bonito." In 1953-76, the landings of Auxis fluctuated between 400 and 1,800 t and averaged 1,000 t (Table

26).A number of other countries, given in Table 26, lists Auxis as among the species they land. Cyprus, Malta, and Yugoslavia are among them but their landings ap- pear to be insignificant. Greece also lands Auiis, but its annual catches are small and appear to be erratic, varying from 500 to 1,200 t and averaging 700 t. Italy's landings are also quite small fluctuating between 400 and 1,700 t and averaging 1,000 t annually.

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1975. Systematics and morphology of the bonitos (iJarcfa) and their

relatives (Scombridae, Sardmil. Fish. Bull., U.S. 73:516-625. COLLETTE, B. B.. and R. N. GIBBS, JR.

1963a. Preliminary field guide to the mackerel- and tuna-like fishes of the Indian Ocean (Scombridae). Smithson. Inst., 48 p. 1963b. A preliminary review of the fishes of the family Scombridae. FAG Fish. Rep. 6:23-32. CUVIER. G.

1829. Le regne animal. Nouvelle ed. torn. 11. Deterville. Paris. 40(i P- CUVIER, G.. and A VALENCIENNES.

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• U.S. a'J\/6I!NMEMr PQINriNS OCSICE; 1931-796-785/72 REGION lO 63

FISHERIES SYNOPSES

This series of documents, Issued by FAO. CSIRO, INP, and NMFS, contains comprehensive reviews of present knowledge on

species and stocl<s 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 In- formation 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- cerned or of related ones, as a means of exchange of knowledge among those already working on the species, and as the basis

for comparative study of fisheries resources. They will be brought up to date from time to time as further Information becomes

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The documents of this series are Issued under the following titles:

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Synopses In this series are compiled according to a standard outline described in Flb/S1 Rev. 1 (1965). FAO, CSIRO,

INP, and NMFS are working to secure the cooperation of other organizations and of individual scientists in drafting synopses on

species about which they have knowledge, and welcome offers of help In this task. Additions and corrections to synopsM

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

of the outline to FAO:

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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. Requests for copies of synopses should be addressed to the issuing organization; except for NMFS/S copies, these can be pur- chased from National Technical Information Service, U.S. Department of Commerce, 5285 Port Royal Road, Springfield, VA

22151.

The following synopses in this series have been issued since January 1978: 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, Pseudopleuronecles americanus (Waibaum) NMFS/S 123 Synopsis of biological data on the rock crab. Cancer irroratus Say NMFS/S 118 Synopsis of biological data on bonltos of the genus Sarda NMFS/S 122 Synopsisof biological data on tunas of the genus £ufhynnus NMFS/S 121 Synopsis of biological data on striped bass, Morone sexatilis (Waibaum)

May 1978

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October 1979 June 1980
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