FAO Fisheries Synopsis No. 125, Volume 5 FIR/S125 Vol.

FAO SPECIES CATALOGUE

VOL. 5. BILLFISHES OF THE WORLD

AN ANNOTATED AND ILLUSTRATED CATALOGUE

OF MARLINS, SAILFISHES, SPEARFISHES
AND SWORDFISHES KNOWN TO DATE

UNITED NATIONS DEVELOPMENT PROGRAMME

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
FAO Fisheries Synopsis No. 125, Volume 5 FIR/S125 Vol.5

FAO SPECIES CATALOGUE

VOL. 5 BILLFISHES OF THE WORLD

An Annotated and Illustrated Catalogue

of Marlins, Sailfishes, Spearfishes and Swordfishes Known to date

MarIins,

prepared by

Izumi Nakamura
Fisheries Research Station
Kyoto University
Maizuru
Kyoto 625, Japan

Prepared with the support from the

United Nations Development Programme (UNDP)

UNITED NATIONS DEVELOPMENT PROGRAMME

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS

Rome 1985
 The designations employed and the presentation
of material in this publication do not imply the
expression of any opinion whatsoever on the
part of the Food and Agriculture Organization
of the United Nations concerning the legal
status of any country, territory. city or area or
of its authorities, or concerning the delimitation
of its frontiers or boundaries.

M-42
ISBN 92-5-102232-1

All rights reserved . No part of this publicatlon may be reproduced.

stored in a retriewal system, or transmitted in any form or by any means,
electronic, mechanical, photocopying or otherwase, wthout the prior
permission of the copyright owner. Applications for such permission,
with a statement of the purpose and extent of the reproduction should
be addressed to the Director, Publications Division, Food and Agriculture
Organization of the United Nations Via delle Terme di Caracalla, 00100
Rome, Italy.

FAO 1985
 - - iii -

"Then the fish came alive, with his dead in

him, and rose high out of the water showing all his
power and his beauty. He seemed to hang in the
air above the old man in the skiff. Then he fell
into the water with a crash that sent spray over
the old man and over all of the skiff” (from The
Old Man and the Sea, by Ernest Hemingway). I
do hope for the eternal prosperity of these beauti-
ful and powerful animals in our water planet.

PREPARATION OF THIS DOCUMENT

The present publication was prepared under the FAO Regular Programme with the support of the United
Nations Development Programme as part of the UNDP/FAO Global Project on Survey and Identification of the
World’s Marine Fishery Resources (GLO/82/001). It is the fifth worldwide species catalogue issued under the
FAO Fisheries Synopses series.

The author is one of the foremost authorities on billfish taxonomy (see Nakamura, I., 1983) and has also had
a long field experience on the biology and fisheries of these species. He has prepared, in the course of the past
seven years, all FAO species identification sheets so far published on this group (Western Centra1 Atlantic,
Eastern Central Atlantic and Western Indian Ocean).

The index of scientific and vernacular names was prepared in collaboration with FAO's Fishery Information,
Data and Statistics Service.

Most illustrations were redrawn from draft material provided by the author.

Technical Editor: W. Fischer, Fishery Resources and Environment Division, FAO

Illustrators: P. Lastrico (principal illustrator) and O. Lidonnici, FAO, Rome

ABSTRACT

This is the fifth in the FAO series of worldwide annotated and illustrated
catalogues of major groups of organisms that enter marine fisheries. The present
volume includes the two families and all 12 species of billfishes known so far. It
provides a comprehensive and illustrated key to families and species, with a glossary of
technical terms and measurements. Within each family are given individua1 accounts
of species which include drawings, scientific and vernacular names, information on
habitat, biology and fisheries, and a distribution map. The section including family and
species accounts is followed by a table indicating the distribution of each species by
major marine fishing areas. The work is fully indexed and there is ample reference to
pertinent literature.

Distribution: For bibliographic purposes this document should

be cited as follows:
Author
FAO Fisheries Department N a k a m u r a , I . , F A O s p e c i e s c a t a l o g u e . Vo1.5.
FAO Regional Fisheries Officers 1985 Billfishes of the World. An anno-
Regional Fisheries Councils tated and illustrated catalogue of
and Commissions marlins, sailfishes, spearfishes and
Selector SM swordfishes known to date. FAO
Fish.Synop., (125)Vo1.5:65 p.

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TABLE OF CONTENTS

Code Page

1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.1 Pl an of the Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.2 General Remarks on Billfis hes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

1.3 Il l us trated Glos s ary of Technica l T e rm s a nd M e a s ure m e nts . . . . . . . . . . . . . . . . 10

2. SYSTEMATIC CATALOGUE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

2.1 Il l ustrated Key to Genera and S pe c ie s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

2.2 Additional Aids to Identificatio n of G e ne ra a nd S pe c ie s . . . . . . . . . . . . . . . . . . 19

2.3 FAM ILY ISTIOP HORIDAE . . . . . . . . . . . . . . . . . . . ISTIO . . . . . . . . . . . 20

Istiophorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ISTIO Istio . . . . . . . . 20

I. albicans (Atlantic sailfish) . . . . . . . . . . . . . . . . . ISTIO Istio 1 . . . . . . . 21

I. platypterus (Indo-P acific sailfis h) . . . . . . . . . . . . . . ISTIO Istio 2 . . . . . . . 23

Makaira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ISTIO Mak . . . . . . . . 27

M. indica (Black marlin) . . . . . . . . . . . . . . . . . . . . ISTIO Mak 2 . . . . . . . 27

M. mazara (Indo-P acific blue ma rlin) . . . . . . . . . . . . . ISTIO Mak 3 . . . . . . . 30
M. nigricans (Atlantic blue marlin) . . . . . . . . . . . . . . ISTIO Mak 1 . . . . . . . 33

Tetrapturus . . . . . . . . . . . . . . . . . . . . . . . . . . . . ISTIO Tetra . . . . . . . . 35

T. albidus (Atlantic white marlin) . . . . . . . . . . . . . . . ISTIO Tetra 1 . . . . . . . 35

T. angustiros tris (Shortbill s pea rfis h) . . . . . . . . . . . . . ISTIO Tetra 3 . . . . . . . 38
T. audax (Striped marlin) . . . . . . . . . . . . . . . . . . . . ISTIO Tetra 5 . . . . . . . 40
T. belone (Mediterranean s pearfis h) . . . . . . . . . . . . . . ISTIO Tetra 6 . . . . . . . 43
T. georgei (Rounds cale s pearfi s h) . . . . . . . . . . . . . . . ISTIO Tetra 4 . . . . . . . 45
T. pfluegeri (Longbill s pearfish) . . . . . . . . . . . . . . . . ISTIO Tetra 2 . . . . . . . 46

2.4 FAMILY XIPHIIDAE . . . . . . . . . . . . . . . . . . . . . . . XIPH . . . . . . . . . . . 48

Xiphias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XIPH Xiph . . . . . . . . 48

X. gladius (Swordfish) . . . . . . . . . . . . . . . . . . . . . XIPH Xiph 1 . . . . . . . . 48

3. LIST OF S PECI ES BY M AJOR FIS H I N G A R EA S . . . . . . . . . . . . . . . . . . . . . . . . 52

4. BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

5. INDEX OF SCIENTIFI C AND VERN A C U LA R N A M ES . . . . . . . . . . . . . . . . . . . . . 59

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1. INTRODUCTION
This review covers the 12 species of billfishes recognized to date, even though the taxonomic status of one
of them (Tetrapturus georgei) is still somewhat uncertain. A thirteenth form, the so-called “Hatchet marlin”,
often mentioned in recent literature, is also briefly presented (see page 5), but cannot be recognized as a valid
species on the basis of the presently available data.
The systematic part of the present catalogue is based primarily on Nakamura, Iwai and Matsubara (1968),
Nakamura (1974) and Nakamura (1983), while the remaining information, particularly that on biology and fisheries
of the various species, stems from the author’s personal experience and pertinent literature available to him. In
order to avoid overcharging the text with literature citations, every effort was made to restrict these to papers
considered to be of specific relevance to the biology and fisheries of the species in question. Many other
important papers, particularly on systematics, anatomy, distribution and the more general aspects of biology and
fisheries, although omitted in the text, have been included in the bibliography. Although great care was paid in
evaluating the published and unpublished information used in this review, some misjudgements and incorrect
interpretations will undoubtedly have occurred.
For more detailed information on billfish taxonomy, stocks, biology and fisheries, the reader is referred to
specialized periodical publications such as the Bulletin of the Far Seas Fisheries Research Laboratory (Shimizu,
Japan), the Fishery Bulletin of NMFS, NOAA, US Department of Commerce (Seattle, USA), and the more
comprehensive papers on billfish taxonomy and biology, such as the Proceedings of the International Billfish
Symposium, Kailua-Kona, Hawaii, 9 to 12 August 1972, edited by R.S. Shomura and F. Williams (NOAA Technical
Report NMFS SSRF-675), “The sailfish, the swashbuckler of the open sea”, by J.B. Tinsley (1964), “Billfish, marlin,
broadbill, sailfish”, by C.O. Mather (1976) and “Systematics of the Billfishes (Xiphiidae and Istiophoridae)” by
I. Nakamura (1983).
Acknowledgements

The author wishes to convey his deepest appreciation and thanks to Dr Walter Fischer (FAO, Rome) for his
initiative, guidance, encouragement and enthusiastic support; to Dr Shiro Chikuni (FAO, Rome) for his critical
revision of the fisheries information included in this catalogue; to Mr Paolo Lastrico for his neat work in
redrawing the rough sketches presented with the draft manuscript; to Mrs Michèle Kautenberger-Longo (FAO,
Rome) for typing of the final manuscript and to Ms Gloria Soave (FAO, Rome) for the revision of the bibliography.
The author also wishes to express his deep gratitude to:
Mr Sho Okano of the Fishermen’s Cooperative of Kyoto Prefecture who granted him permission to examine
billfishes landed at the Nishi-Maizuru Fish Market; to Captain Yoshio Kurohiji and the crew of the R/V SHUNYO
MARU and to Captain Sadamu Tanabe and the crew of the R/V SHOYO MARU for their valuable help during his
participation in several longline fisheries research cruises in the Pacific, Indian and Atlantic oceans.
The fishermen and fisheries technicians who have helped the author in the course of his billfish studies and
have given him invaluable information on fisheries, behaviour and biology of these fishes, in particular
Mr Guillermo M. Adachi (Manzanillo Game Fishing Club, Mexico); Mr Peter Goadby (Game Fishing Association of
Australia); Messrs Jack Izatt, John Covacevich, David Hopton, Peter Nielsen and Joe Bethune and Ms Daphnie
Nielson (Cairns Game Fishing Club, Australia); Messrs Toshiro Sato, Wataru Ichikawa, Shoji Nakano and Michio
Kuroiwa (Japan Marine Fisheries Resources Research Center); Mr Elwood K. Harry (President of the International
Game Fish Association); Messrs Shojiro Shimura and Soroku Togo (Fisheries Agency, Government of Japan);
Messrs Seki Araki, Ryoichi Shugyo and Tsuneo Okabe (Nishidomari Fishermen’s Cooperative, Tsushima, Japan).
The scientists and research institutions who have provided useful information, or accommodation to the
author and offered him their facilities during his visits, in particular: the late Dr Hiroshi Nakamura, Dr Shooji
Ueyanagi, Dr Shoji Kikawa, the late Mr Hisaya Watanabe, Dr Keiichiro Mori, Dr Hajime Yamanaka, Mr Yasuo
Nishikawa, Dr Toshio Shiohama and Mr Ikuo Warashina (formerly Nankai Regional Fisheries Research Laboratory,
Kochi now Far Seas Fisheries Research Laboratory, Shimizu, Japan); the late Prof. Kiyomatsu Matsubara and
Prof. Tamotsu Iwai (Kyoto University, Kyoto, Japan); Dr Jiro Isa and Mr Akio Tomori (Okinawa Prefectural
Fisheries Experimental Laboratory, Okinawa, Japan); Dr John R. Paxton and Dr Frank Talbot (Australian Museum,
Sydney, Australia); Dr Julian G. Pepperell, Mr John Matthews and Mr Robert Williams (New South Wales State
Fisheries, Sydney, Australia); Dr Robert H. Gibbs, Jr. (US National Museum of Natural History Washington, D.C.,
USA); Dr Bruce B. Collette and Dr Daniel Cohen (Systematics Laboratory, NMFS, NOAA, Washington, D.C., USA);
Dr W. Eschmeyer and Dr T. Iwamoto (California Academy of Sciences, San Francisco, USA); Dr Robert J.
Lavenberg (Los Angeles County Museum, Los Angeles, USA); the late Prof. Carl L. Hubbs and Prof. Richard H.
Rosenblatt (Scripps Institution of Oceanography, La Jolla, USA); Mr Withold L. Klawe (Inter-American Tropical
Tuna Commission, La Jolla, USA); Prof. C. Richard Robins and Prof. Donald P. de Sylva (University of Miami,
Miami, USA); Dr Francis G. Carey (Woods Hole Oceanographic Institution, Woods Hole, USA); Dr William Smith-
Vaniz and the late Dr John Böhlke (Academy of Natural Sciences of Philadelphia, Philadelphia, USA); Dr Donn E.
Rosen and Dr C.L. Smith (American Museum of Natural History, New York, USA); Dr Peter J.P. Whitehead, Mr
Alwyn Wheeler and Mr Oliver Crimmen (British Museum, Natural History, London, UK); Dr Marie-Louise Bauchot
and Ms M. Desoutter (Museum National d’Histoire Naturelle, Paris, France); Dr Alfred Post (Institut für
Seefischerei, Hamburg, Germany, Fed. Rep.); Dr Jørgen Nielsen (Universitetets Zoologiske Museum, Copenhagen,
Denmark); Dr E.G. Silas (Central Marine Fisheries Research Institute, Cochin, India); the late Dr H.T. Teng
(Taiwan Fisheries Research Institute, Keelung, China); Dr Thosaporn Wongratana (Department of Biology,
Chulalongkorn University, Bangkok, Thailand).
Finally, the author wishes to heartily acknowledge the technical assistance provided by Mrs Reiko
Nakamura.
 - 2 -

1.1 Plan of the Catalogue

This catalogue is arranged alphabetically by genera and species. Each of the multispecies genera is
introduced with general descriptive remarks, illustrations of diagnostic features, highlights on the biology, and
relevance to fisheries. The information pertaining to each species is arranged by paragraphs, as follows:
(1)scientific name, (2) synonymy, (3) FAO species names, (4) field marks, (5) diagnostic features,
( 6 ) g e o g r a p h i c a l d i s t r i b u t i o n , (7) habitat and biology, (8) size, (9) interest to fisheries, (10) local species
n a m e s , (11) literature, and (12) remarks.

(1((1) (1) Scientific name : : Reference is given to the original description of each species so no confusion will
arise as to precise identification.

(2) Synonymy : Synonims and different name combinations are listed (misidentifications and other
nomenclatorial problems are discussed under (11) remarks).
(3) (3) FAO species names : English, French and Spanish names for each species, to be used primarily within
FAO, were selected on the basis of the following
: criteria: (i) each name must apply to one species
only, in a worldwide context; (ii) the name should not lead to confusion with other groups. Wherever
possible, the names selected were based on vernacular names (or parts of names) already in existence
within the areas where the species is fished. FAO species names are, of course, not intended to replace
local species names, but they are considered necessary to overcome the considerable confusion caused
by the use of a single name for many different species, or several names for the same species.
(4) (4) Field marks : A few obvious field characters of use in field identification extracted from “Diagnostic
Features” at various levels.
(5) (5) Diagnostic features : Distinctive characters of the species are given as an aid for identification,
accompanied by pertinent illustrations. Species identifications should be attempted only after
consultation of the illustrated key to genera and species. Reference to FAOFSpecies Identification
Sheets is given wherever relevant.

(6) (6) Geographical distribution : : The entire known geographic range of the species, including areas of
seasonal occurrence, is given in the text and shown on a small map. In cases where only scattered
records of occurrence are available, interrogation marks have been used to indicate areas of
suspected distribution.
(7) (7) Habitat and biology : : The known depth range of the species,and information on salinity and
temperature of its habitat are given where known. Information on biological aspects, such as
migrations, spawning seasons and areas, food, predators and longevity is also included.
(8) (8) Size : The maximum known, as well as the common body length and weight (if available) are given.
Body length is measured from the tip of the lower jaw to the tip of the caudal rays in the middle of
the fork of the tail. The all-tackle angling record and length at first maturity are given where
known.
(9) (9) Interest to fisheries : This paragraph gives an account of the areas where the species is fished and
of the nature of the fishery; its importance is either qualitatively estimated or actual figures of
annual landings are provided. Data on utilization (fresh, dried, frozen, canned, etc.) are also given
where available. Here too, the quality and quantity of the information available vary considerably
with the species.
(11)
(10) (10) Local species names : These are the names used locally for the various species. The present
compilation is necessarily incomplete, since only a fraction of the local names used throughout the
world is actually published. In many cases, local names are available only for species supporting
documented fisheries. Apart from possible omissions due to limitations of literature available, some
of the names included may be somewhat artificial (i.e. through transliteration of indigenous words
into English). The local species name is preceeded by the name of the country concerned (in capital
letters) and, where necessary, by geographical specifications (in lower case). Whenever possible, the
language of the transcribed vernacular name is added in parenthesis. When more than one name is
used within a country, the official name, if available, is underlined.

(11) Literature : This includes references to the most important publications relevant to the species, the
emphasis being on biology and fisheries. Additional references are included in the bibliography. In
the case of a few uncommon species, only systematic papers are available.
(12)
(12) Remarks : Important information concerning the species and not fitting in any of the previous
paragraphs is given here. For instance, in some cases the scientific name used in the present
catalogue, although nomenclaturally correct, is not the best known. Other nomenclatural problems,
such as the use of subspecies, are discussed.
 - 3 -

1.2 General Remarks on Billfishes

The term “Billfishes” has been widely accepted by both, commercial and sports fishermen as well as
scientists, to apply to the large fishes of the families Xiphiidae and Istiophoridae, characterize d by the
prolongation of the upper jaw, much beyond the lower jaw into a long rostrum which is flat and swordlike
(swordfish) or rounded and spearlike (sailfishes, spearfishes and marlins). Needlefishes, (Belonidae ) are also
sometimes referred to as billfishes, but they are easily distinguished from the true billfishes by having both jaws
prolonged, the dorsal and anal fins both single and similar in size and shape, and the pelvic fins inserted far behind
the pectorals.

The billfishes include 12 species arranged in four genera and two families as follows (see also Fig.1):

Phylum Chordata

Superclass Gnathostomata

Class Osteichthyes

Subclass Actinopterygii

Infraclass Teleostei

Division Euteleostei

Superorder Acanthopterygii

Order Perciformes

Suborder Xiphioidei

Family Xiphiidae

Genus Xiphias
Xiphias gladius - Swordfish (worldwide)

Family Istiophoridae
Genus Istiophorus

Istiophorus albicans - Atlantic sailfish (Atlantic)

Istiophorus platypters - Indo-Pacific sailfish (Indian and Pacific oceans)

Genus Tetrapturus

Tetrapturus albidus - White marlin (Atlantic)

Tetrapturus angustirostris - Shortbill spearfish (Indian and Pacific oceans)

Tetrapturus audax - Striped marlin (Indian and Pacific oceans)

Tetrapturus belone - Mediterranean spearfish (Mediterranean Sea)

Tetrapturus georgei - Roundscale spearfish (Atlantic)

Tetrapturus pfluegeri - Longbill spearfish (Atlantic)

Genus Makaira

Makaira indica - Black marlin (Indian and Pacific oceans)

Makaira mazara - Indo-Pacific blue marlin (Indian and Pacific oceans)

Makaira nigricans - Atlantic blue marlin (Atlantic)

 - 4 -

Fig.1 Classification of billfishes

Sizes of drawings correspond roughly to maximum lengths of species; screened oval areas show
cross-sections of bodies at level of pectoral fin bases (arrows show position of anus) anus)
 - 5 -

Another billfish has recently been reported by

several authors (Robins, 1974; de Sylva, 1974;
McClane, 1974; Pristas, 1980) from the western
Atlantic Ocean, particularly in the Gulf of Mexico.
This might be either a new species or an aberrant
form of Tetrapturus albidus. Venezuelan and Cuban
fishermen have since long referred to this fish as
t h e “ H a t c h e t m a r l i n ” o r “Axe marlin”. It is
characterized chiefly by the truncate shape of the
anterior lobes of the first dorsal and anal fins
(Fig.2) However,in the absence of conclusive
taxonomic evidence, a final decision on the status of
the Hatchet marlin cannot be made at this time and “Hatchet marlin” Fig.2
further study on this problem is urgently needed. redrawn from McClane (1974)

Xiphias gladius (Family Xiphiidae) differs from all species of the Family Istiophoridae in both external and
internal features. Some superficial phenetic similarities, especially between Makaira and Xiphias seem to be
due to convergent evolution. As regards the relationship between the three istiophorid genera Tetrapturus,
Istiophorus and Makaira, various different opinions have been advanced. Generally, the sailfishes are included
in the genus Istiophorus, the small spearfishes in Tetrapturus and the large (blue and black) marlins in Makaira.
The smaller marlins striped and white), have been placed by many authors in either of the genera Tetrapturus
and Makaira. On the basis of the present author’s recent studies (Nakamura, 1983) it seems clear that these two
species, T. albidus and T. audax should be included in the genus Tetrapturus together with the small spearfishes,
T. angustirostris, T. belone,
belone T. pfulegeri and T. georgei (fig.1). (Fig.1).

It is likely that enlargement of the body as well as acquirement of the capability for fast swimming have
been an evolutionary trend in billfishes (see maximum body sizes of living billfishes in Fig.3).

Fig.3 Comparison of approximate maximum recorded body length and maximum recorded body weight (both are
not always coincidental with each other) in billfishes. The cross-sections of body and bill of each species
are shown schematically on left and right, respectively (modified after Nakamura, 1983, Fig.54)
 -6-

Furthermore, it has been shown that as billfishes grow larger, their bodies become much more robust,
particularly in the case of Makaira indica (see Fig.4).

O 1OO 2OO 3OO

BODY LENGTH cm.

Fig.4 Length-weight relationships of billfishes caught by setnets and landed at Nishi-Maizuru Fish Market, Kyoto
Pref., Japan between August 1976 and November 1978. Smaller I. platypterus (less than 110 cm body
length) were weighed whole, the other large fishes of all species were weighed without gills or guts, and
sometimes without the tip of the bill
 -7-
-

Billfishes are primarily oceanic, epipelagic species inhabiting tropical and temperate waters, and seasonally
also the cold waters of all oceans. They are usually confined to the water layers above the thermocline, but
some may occur at greater depths (to below 800 m). The general distribution patterns of billfishes are
summarized in Fig.5. While Xiphias gladius is cosmopolitan, the istiophorids have more restricted distributions.
Although some billfish experts recognize only one cosmopolitan species of sailfish Istiophorus platypterus, the
present author prefers, for the time being, to retain the traditional distinction between I. platypterus for the
Indo-Pacific sailfish and I. albicans for the Atlantic sailfish, because there is evidence of some morphological
differences between these two forms (see pages 16, 21, 23 and 24). If we consider the presence of spawning
grounds to be a major element in the delimitation of principal areas of distribution, we may conclude that five of
the species of Istiophoridae are confined to the Pacific and Indian oceans, six (if we recognize Tetrapturus
georgei as valid) to the eastern atlantic and five (if we recognize the Hatchet marlin as valid) to the western
central Atlantic. Furthermore, three species, Makaira indica, Tetrapturus angustirostris and Tetrapturus audax
have been shown to occur incidentally as "invaders" int the eastern Atlantic, which brings to 10 the number of
species found at least occasionally in the Atlantic Ocean.

Fig.5 Schematic representation of the distribution of billfishes

W: Western; E: Eastern; N: Northern Hemisphere; S: Southern Wemisphere;

Principal distribution areas (including spawning grounds);

@$$$$$ Areas of occasiona1 distribution, or invasion, (no

(no spawning)
(modified after Nakamura, 1983, Fig.61)

Istiophorus albicans, I. platypterus, and Makaira indica migrate seasonally towards coastal waters, while T.
angustirostris, M. mazara and M. nigricans are the most typically oceanic species. Roughly speaking, the
migration pattern of all billfishes involves seasonal movements into temperate or cold waters for feeding and
back to subtropical or tropical waters for overwintering and spawning. Although being among the largest and
fastest bony fishes known from the sea, they seldom perform transoceanic migrations as is typical for the large
tunas.
 -- 8 - -

All species are dioecious (the sexes are separate) and their mating display is not completely known. None of
the species are known to show sexual dimorphism in morphological features or colour pattern, but females of
many species attain larger sizes than males. Batch-spawning of most species takes place in tropical and
subtropical waters. The eggs are very small (about 1 mm in diameter) and pelagic, hatching out into planktonic
larvae.

Billfishes are active and voracious predators, using their long rostrum for attacking their prey. Most
likely there are no true, regular predators of billfishes, although they are preyed upon occasionally by many large
oceanic fishes, such as tunas, wahoo, skipjack and dolfinfishes, particularly during their younger stages. The
young are also taken sometimes by adult billfishes.

Being excellent foodfish, all species are of some importance to fisheries. Their commerical value is
particularly high in Japanese markets. Most of them are exploited commercially by longliners and all are
regarded as excellent and exciting targets by sportsfishermen. The recent world production of billfishes fluctuates
around 95,000 metric tons per year (Table I) of which more than 90% is taken as bycatch in tuna longline
fisheries. T h i s b y c a t c h r e p r e s e n t s i n w e i g h t a b o u t 1 0 % o f t h e a c t u a l t u n a c a t c h e s ( g e n u s T h u n n u s ) . J a p a n
produced about 70% of the world's catch and is the principal consumer country for these fishes. The species
predominating in the catches are Xiphias glaudius, Makaira mazara, and Tetrapturus audax, accounting for about
40%, 23% and 17%, respectively, of the toal world catch.
-

TABLE I

Estimated world catch of billfishes and tunas in metric tons (source : FAO, 1983, 1984)

Systematic Category 1978 1979 1980 1981 1982

Istiophorus albicans 267 2 823 1 320 1 091 920

Istiophorus platypterus 10 516 7 961 7 767 6 438 7 214
Makaira indica 2 909 3 440 2 465 2 453 2 373
Makaira mazara 18 193 18 654 21 109 21 413 20 727
Makaira nigricans 1 842 2 160 2 328 2 218 2 448
Tetrapturus albidus 204 93 119 121 131
Tetrapturus audax 15 426 15 988 18 429 15 664 15 460
Istiophoridae n.e.i.* 5 920 5 339 6 062 6 350 7 320

Subtotal Istiophoridae 55 277 56 483 59 599 55 748 56 593

Xiphias gladius 40 279 37 922 36 402 37 726 40 321

Subtotal Xiphiidae 40 279 37 922 36 402 37 726 40 321

1 Total Billfishes (Xiphioidei) 95 556 94 405 96 001 93 474 96 914

Thunnus alalunga 222 775 191 014 182 084 186 125 183 481
albacares
Thunnus albacares 521 466 537 987 522 412 560 274 535 725
Thunnus atlanticus 78 105 300 845 807
Thunnus maccoyii 32 415 35 475 39 186 37 226 29 005
Thunnus obesus 210 358 195 635 201 021 175 047 182 940
Thunnus thynnus 35 601 35 485 32 479 44 478 41 365
Thunnus tonggol 34 48 9 368 4 020
Total Thunnus 1 022 727 995 749 977 491 1 004 363 977 343

Ratio: Xiphioidei/Thunnus 0.093 0.095 0.098 0.093 0.097

*n.e.i.= not elsewhere identified

 - 9 -

Billfishes are caught with a variety of fishing methods, such as longlining, trolling, harpooning, drift-netting,
set-netting and others. The most important fishing method used in commercial fisheries is surface-longlining
which was gradually developed by Japanese, Chinese (fishermen from Taiwan Province) and Korean fleets
originally operating in the northwestern Pacific and then extending to other Pacific areas and to the Indian and
Atlantic oceans. This method is also extensively used by other fishing nations, particularly the USSR and Cuba.
On the other hand, sportsfishing is exclusively effected by trolling. Figure 6 represents the general distribution
of commercial and sportsfishing areas for billfishes. The commercial longline fisheries cover nearly the entire
natural distribution areas of the species, while all other fisheries are more restricted. The areas where
commercial longline fisheries overlap with sportsfishing have recently been subjected to conflicts of interests in
relation to the extension of Exclusive Economic Zones, and hence, fishing rights, to 200 miles offshore.

.............
k Surface longline fishertes Principal sportfishing grounds iiHarpoon fishing grounds
------ Approximate natural distribution of billfishes based on literature and personal communications from fishermen

Fig.6 Schematic illustration of the world’s commercial and sport fisheries for billfishes
(modified after Nakamura, 1983, fig.62)

Virtually all billfishes are highly appreciated for the excellent quality of their flesh. Fish over 20 kg are
generally used for “sashimi” (sliced raw flesh with soy sauce and green mustard) and materials for “sushi” (sliced
raw flesh with green mustard on vinegar-boiled rice balls). The colour of billfish flesh is usually paler than that
of tuna. One of its advantages is that it does not change colour as easily as tuna flesh, thus withstanding longer
periods of transportation and having a longer market shelf-life than tuna meat.

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1.3 Illustrated Glossary of Technical Terms and Measurements

Fig.7 Schematic illustration of Indo-Pacific sailfish, Istiophorus platypterus (above) and a striped marlin,
Tetrapturus audax (middle and below)
All measurements are straight line distances. For detail of measurements, see Rives (1956) and Nakamura (1983)
 - 11 -

Air-bladder - Membranous sac filled with air Body width - G r e a t e s t w i d t h o f b o d y

or other gases lying in the abdomen just beneath the (Fig.7). Body width at origin of pectoral fins, pelvic
vertebrae. The swordfish’s air-bladder consists of a fins and first anal fin is sometimes also used.
single chamber (Fig.8a), while that of istiophorids is
made up of many bubble-shaped, small chambers Branchiostegal (rays) - Strut- or ray-like
(Fig.8b). Also known as gas-bladder or swim- bones attached to the hyoid arch, connected by the
bladder. branchiostegal membrane (Figs15,16,22).
caudal keel Branchiostegal membrane - The membrane
connecting the branchiostegals and enclosing the
gill chamber ventrally (Figs 15,16,22).
Caudal fin - Median fin situated at the pos-
terior end of the body, consisting of an upper and a
lower lobe (Fig.7).
air bladder Caudal keel - Xiphiidae have a large median
a. Xiphias glaudius caudal keel (Fig.8a)( and Istiophoridae have a pair of
a pair of caudal keels (Fig.8b)(on the middle of the caudal
caudal keels peduncle.
Caudal notch - An elongate, small pit on
dorsal and ventral margins of the caudal peduncle
(Fig.7). The caudal notch is shallow and small in
istiophorids, and deep and rather large in Xiphias.
Caudal peduncle - The narrow part of the
air bladder body between the posterior ends of the dorsal and
b. Makaira indica anal fins and the base of the caudal fin (Fig.7).
Schematic illustration of Caudal vertebrae - Vertebrae that bear a
air-bladder and caudal keels Fig.8 haemal spine ventral to the vertebral centrum
(Fig.10b). Caudal vertebrae lack pleural ribs. The
Anal fin(s) - The fin(s) on the ventral median number of caudal vertebrae is 15 or 16 in Xiphias
line of the body behind the anus (Fig.7). 12 in Istiophorus and Tetrapturus, and 13 in
Makaira.
A n u s - External opening of the intestine,
situated on the ventral midline of the body (Fig.7). anterior neural
The relative position of the anus to the first anal fin zygapophyses neural
is important in istiophorid taxonomy. Also known as spine
posterior neural arch
vent. neural neural canal
zygapophyses
Bill - Long, slender upper jaw. Both jaws are
elongate, but the upper is longer than the lower in
billfishes. The bill is flat in the swordfish and
round in istiophorids (Figs 7,9).
nasal maxilla
cross section
transverse
process
(parapophyses)
ethmoid X. glaudius frontal
premaxilla pleural
cross section rib

a. precaudal vertebrae
nasal neural spine
M. indica
Bill structure in dorsal view (schematic) Fig.9

Bill length - Meausred from the tip of the neural arch

neural canal
bill to the anteriormost point on the fleshy margin
of the orbit (Fig.7).
centrum
Body depth - Greatest depth of body (Fig.7).
Body depth at origin of pectoral fins, pelvic fins,
and first anal fin is sometimes also used. haemal canal
haemal arch
(fused transverse
Body length - Measured from the tip of the haemal process)
lower jaw (with the jaws closed) to the posterior spine
margin of the middle caudal rays (Fig.7). In other
fishes the measurement, “body length” is usually
“standard length”. In billfishbi taxonomy, this dimen- b. caudal vertebrae
sion is used following Rivas (1956).
Schematic illustration of vertebrae
of Xiphias glaudius
Fig.10
 - 12 -

Dorsal fin(s) - Fin(s) on the back of a fish Fork length - Measured from the tip of the
(Fig.7). Billfishes have two dorsal fins. bill (upper jaw) to the posterior margin of the
m i d d l e caudal rays (Fig.7). Usually used for
Ethmoid - Unpaired skull bone on the anterior scombroid (particularly tunas) studies, because the
part of the neurocranium (Fig.9). caudal fork area is very strong, in tunas as well as
in billfishes and the snout is not prolonged in tunas.

Eye diameter - Measured as the greatest dis-

tance across the cornea, that is, between the mar- Frontals - Paired skull bones in middle part
gins of the cartilaginous eyeball (Fig.7). of the neurocranium (Figs 9,16).

Eye-fork length - Measured from the poste- Gill arch - The J-shaped structure under
rior edge of the orbit to the posterior margin of the the gill cover that bears the gill filaments and
middle caudal rays (Fig.7). normally the gillrakers, but the billfishes lack
This dimension is
frequently used by Japanese fishery biologists gillrakers (Fig. 12). There are 4 gill arches on
because it is useful in specimens with bills cut at each side of billfishes.
fish markets or on fishing boats.
upper lobe

Fin grooves - The first dorsal, first anal and

pelvic fins fold down into grooves in all istiophorids
gill arch
when they are swimming rapidly (Fig.11). Fin
grooves are not developed in Xiphias gladius. arch angle
lower lobe

gill filaments ("gill rays")

first gill arch of Xiphias glaudius
(schematic)
Fig.12

slow-swimming Gill filaments - Organ for aquatic respira-

tion. In billfishes the gill filaments are ossified as
“gill rays” as in the case of tunas (Fig.12); see also
Iwai & Nakamura (1964).

Haemal arch - T h e a r c h t h a t i s f o r m e d
above the fused distal ends of the haemal spines of
caudal vertebrae (Fig.l0b).
fast-swimming
Haemal canal - The canal for the blood ves-
Schematic illustration of fin grooves sel, formed by the haemal arch of caudal vertebrae
(shaded areas) in Istiophorus platypterus (Fig.l0b).
Fig.11
Haemal spines - The spines that extend
Fin membranes - The thin membranes between ventrally from the centra of a caudal vertebra
the rays of the fins (Fig.7). (Fig.l0b). The first vertebra with a haemal spine is
the first caudal vertebra.
Fin rays - General term for the soft rays and
spines (spinous rays) that support the fins (Fig.7). Head length - Measured from the tip of the
Soft rays are branched, segmented or paired (left mandible (lower jaw) to the most distant point on
and right elements united). Spines are unsegmented the opercular membrane (Fig.7).
fin supports, unbranched, unpaired and usually stiff
and sharply pointed.

Fin spines - Sharp, pungent and pointed

structures (Fig.7). Usually called merely spines.
 - 13 -

Hypural p l a t e - The expanded ends of the hypu- Interneural bones - The bones situated
ral bones form a wide, fan-like plate onto which the between the neural spines of the vertebrae and the
caudal fin rays insert distally. Like tunas, billfishes spines or rays of the dorsal fin (Fig.14).
differ from most other fishes in having the caudal fin
rays so deeply divided that they completely cover the Interorbital width - Measured as the shortest
hy pural plate; the hypural plate consists of four distance between the fleshy margins of the orbits
hypural bones in Xiphias (Fig.13a) and five in Istio- (Fig.7) .
phoridae (fig. 13b).
hypural 5 Isthmus - Ventral fleshy area on the throat
between the gills (Fig.15).
eye
hypural plate
( hypurals 1-4 )
caudal
fin rays isthmus

brachiostegal
bill mandible membrane
(rostrum) brachiostegal ray
Head of Xiphias glaudius in ventral view
(schematic) Fig.15
parahypural Lacrymal bone - The largest of the infra-
a. Xiphias gladius orbital series of bones, located ventral and slightly
(Caudal fin rays shown only on lower half) anterior to the eye (Fig.16). Also known as pre-
orbital bone or first infraorbital bone.
uroneural hypural plate
epural (hypurals 1-5) preopercle
frontal opercle
autogenous neural spine nasal
maxilla
premaxilla

preural notch subopercle

centrum
predentary
dentary angular
lacrymal
parhypurapophysis retroarticular branchiostregals
interopercle

autogenous haemal spine Lateral view of head skeleton of

b. Istiophorus platypterus Istiophorus platypterus (Schematic) Fig.16
Schematic drawing of hypural plate Fig.13
Interhaemal bones - The bones situated between Laminae of the olfactory rosette - -Fleshy folds
the haemal spines of the vertebrae and the spines or (nasal laminae) containing cells that can detect
rays of the anal fin (Fig.14). odours, arranged i n a r a d i a t e p a t t e r n ( r o s e t t e )
beneath the skin between the anterior and posterior
nostril openings (Fig.17).
upturned skin
anterior
nostril
dorsal fin spine
interneural bone nasal
neural spine
laminae posterior nostril
olfactory
vertebral centrum rosette

pleural rib haemal spine

interhaemal bone
anal fin spine

accessory nasal sac sac nasal cavity

Position of interneural and interhaemal
bones in Istiophorus platypterus Nasal cavity of Istiophorus platypterus
(schematic) Fig.14 after dissection (schematic) Fig.17
 - 14 -

Lateral apophyses - The flanges that extend Lateral line - -A series of sense organs
laterally from the anterior part of each vertebral enclosed in tubular scales along the sides of the
centrum (Fig.18). Also known as the transverse body (Fig.7). The lateral line is looped in Makaira
flanges. mazara and reticulate in Makaira nigricans. O t h e r
precaudal vertebrae caudal vertebrae istiophorids have a single lateral line. Xiphias
gladius has a single lateral line in immature stages
which disappears in the adult.

Mandible - Known as the lower jaw (Figs 7,

15), consisting of predentary (Istiophoridae only),
dentary, angular and retroarticular bones (Fig.16).

X. glaudius Maxilla - The supporting bone of the pre-

maxilla which bears teeth in the upper jaw (Figs
9,16). The maxilla itself also bears teeth in
istiophorids, unlike most other fishes.

Nape - D o r s u m o f t h e n e c k a r e a i m m e -
I. platypterus diately posterior to the head (Fig.7).

Nasals - Paired bones in the ethmoid region

of the neurocranium (Figs 9,16).

I. albicans Neural arch - T h e a r c h t h a t i s f o r m e d

below the fused basal part of the neural spine of
the vertebrae (Fig.10).

Neural canal - The canal for the nerve cord

T. angustirostris formed by the neural arch (Fig.10).

Neural spines - The spines that extend dor-

sally from the centra of a vertebra (Fig.10).

Nostrils - External openings of the nasal

T. pfluegeri cavity. There are two (anterior and posterior)
nostrils in billfishes (Figs 7,17).

O p e r c u l u m - Gill cover, supported by four

bones: opercle, preopercle, interopercle and sub-
T. albidus opercle (Figs 7,16).

Orbit - The eye socket (Fig.16). The sclera

is ossified in billfishes.

T. audax Palatine - A pair of plow-shaped bones, the

ventral margin of which lies in the roof of the
mouth. The palatine bones may be toothed in
istiophorids (Fig. 19), but are edentate in Xiphias.

vomer palatine teeth

parapophysis (edentate)
M. mazara
lateral apophysis

M. nigricans

Roof of mouth in Tetratrapturus albidus

M. indica (schematic)
Fig.19

Lateral apophyses of billfish bvertebrae (schematic). Parapophyses - (Transverse processes). Pro-

White areas: lateral view; black areas: ventral view jections from the ventral centra of the vertebrae
Fig.18 (Fig.10).
 - 15 -

Pectoral fins - Lateral paired fins behind the Viscera - Internal organs of the body which
head (Fig.7). are well developed in all the species of billfishes
(Fig.20). The intestine is coiled, the spleen is not
Pelvic fins - Paired fins on the ventral edge visible in ventral view, and the gonads are symmet-
of anterior body (Fig.7). Also known as ventral rical in Xiphias (Fig.20a). The spleen is visible in
fins. ventral view, and the intestine is undulated in
Istiophoridae (Fig.20b,c). The gonads are symmetri-
Precaudal vertebrae - The anterior verte- cal in Istiophorus (Fig20b), in Makaira, and appa-
brae without haemal spines (Figs l0a,18). Also known rently in Tetrapturus except T. angustirostris and
as abdominal vertebrae. The number of precaudal T. pfluegeri where they are asymmetrical and Y-
vertebrae is 10 or 11 in Xiphias, 12 in Istiophorus shaped (Fig.20c) (possibly also in T. belone).
and Tetrapturus and 11 in Makaira.

Premaxillae - Paired bones of the upper jaw,

usually bearing teeth in higher teleosts and asso-
ciated with the maxillae (Figs 9,16).
liver
Predentary - Unpaired bone anterior to the
dentary. Present in Istiophorids (Figs 16,22), but
absent in Xiphias. pyloric
caeca
Rostrum - Projecting snout; or bill (Figs 7,9).

Scales - Thin, flat, bony plates covering the gall

body, usually cycloid or ctenoid. The scales of bladder
istiophorids do not fit into these categories, they are
intestine spleen
elongate and pungent, with sharp posterior points
(Fig.7). The arrangement and shape of the scales
are useful characters for the identification of bill- rectum stomach
fishes. Xiphias has no scales in the adult stage.

Snout - Forward part of the head, anterior to gonad

the eyes and above the mouth (Fig.7).

Snout length - Measured from the tip of the

mandibule (lower jaw) to the anteriormost point on
the fleshy margin of the orbit (Fig.7) in billfishes. a. Xiphias b. Istiophorus c. Tetrapturus
gladius platypterus angustirostris
Standard length - In systematic studies, the
standard length is the distance from the anterior- Viscera of billfishes in ventral view (schematic)
Fig.20
most part of the snout to the caudal fin base
(theoretically to the end of the terminal vertebra,
but this is not practical in general use) (Fig.7). In
Vomer - A median skull bone, the ventral
fishery studies the fork length, e.g., the distance
surface of which lies in the roof of the mouth. The
from the anteriormost point of the head to the
vomer is edentate in billfishes (Fig.19), but many
posterior margin of the middle caudal rays is used.
other fishes have vomerine teeth.
Total lengt h - Straight-line measurement from
the anteriormost to the posteriormost point of the
fish (Fig.7).

Trunk length - Measured from the posterior

edge of the orbit to the anterior insertion of the
caudal keels (Fig.7). Used in sport-fishing.

Vertebra - One of the bony or more or less

cartilaginous (in primitive fishes or young fishes)
segments composing the spinal column or backbone
(Figs 10,18). Number of vertebrae = number of pre-
caudal vertebrae plus number of caudal vertebrae:
26 in Xiphiidae (Xiphias) and 24 in Istiophoridae
(Istiophorus, Tetrapturus and Makaira).

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2. SYSTEMATIC CATALOGUE

dorsal fins
2.1 Illustrated Key to Genera and Species
well separated
cross section
l a . No pelvic fins; a large median keel on
each side of caudal peduncle region;
snout extremely long, forming a sword-
like bill, depressed in cross-section;
first dorsal fin short-based, well sepa- depressed large
rated from second dorsal fin in adults no scales median
bill no teeth
(Fig. 21); no scales on body nor teeth in keel
no pelvic fins
jaws in adults; right and left branchio- Xiphias glaudius Fig. 21
stegal membrane separated distally
(Fig.22a); vertebrae 26 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xiphias gladius
Worldwide in tropical eye
and temperate waters I

l b . Rigid, tapering pelvic fin present; a isthmus

pair of caudal keels on each side of
caudal peduncle region; snout long to
branchiotegal branchiostegal
somewhat shorter, round in cross-sec- mandible
bill (rostrum) ray membrane
tion; first dorsal fin long-based, close
to second dorsal fin (Fig.23); body
covered with small, elongate bony
scales; rasp-like small jaw teeth pre-
sent in adults; right and left branchio-
stegal membranes united broadly (Fig. I
22b); vertebrae 24 predentary eye
ventral view of head Fig. 22

2a. First dorsal fin (X) sail-like and elongate

remarkably higher than body depth bony dorsal fins
(Y) at level of midbody; pelvic fin scales close together
rays very long, nearly reaching to
anal fin origin, with well developed cross section
membrane (Fig.24)

paired
3a. Pectoral fins and caudal fin rasp-like teeth
keels
short in immature specimens
up to about 90 cm body length rigid tapering
(Fig.25); attains greater, size T. audax Fig.23
(about 100 kg maximum body
weight) than Atlantic sail-
fish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Istiophorus platypters
Pacific and Indian oceans

3b. Pectoral fins and caudal fin

long in immature specimens to
about 90 cm body length (Fig.
26); attains smaller size (about
60 kg maximum body weight)
than Indo-Pacific sailfish . . . . . . . Istiophorus albicans
anus Fig.24
Atlantic Ocean pelvic fin

I. platypterus I. albicans
(immature specimen) Fig.26
(immature specimen) Fig.25
 -- 17 -

22b. First dorsal fin (X) lower than body

depth (Y) at level of midbody, not sail-
like; pelvicfin rays short, well sepa-
r a t e f r o m origin of anal fin, with
moderately developed membrane (Fig.
27)

Pelvic fin anus

Fig. 27
4a. Height of anterior lobe of first
dorsal f i n (X) s l i g h t l y g r e a t e r
than, or nearly equal to, body
d e p t h ( Y ) ; nape slightly elevated
or not elevated (Fig.28); body well
compressed laterally; vertebrae
12+12=24

5a. Anterior lobe of first dorsal

fin slightly higher than rest
of fin which remains of about
equal height almost to the
end (Figs 27,30,31,32); a n u s
situated far anterior to first
anal fin origin (Fig.29), t h e
d i s t a n c e b e t w e e n t h e m (X) a. T. angustirostris
greater than height of first
anal fin (Y)
b. T. albidus
Fig. 28
6a. Bill short, its length (X)
usually equal to, or
shorter than, head length
(Y); pectoral fins narrow
and short, less than 18%
of body length (Figs 30,
31) anus second anal fin

7a. Bill very short, less first anal fin Fig. 29

than 15% of body
length (Fig. 30) . . . .Tetrapturus angustirostris
Pacific and Indian oceans

7b. Bill moderately

short, less than 18%
of body length (Fig.
31)) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrapturus belone
T. angustirostris Fig. 30
Mediterranean Sea

6b. Bill long, its length (X)

usually equal to, or
longer than, head length
(Y); pectoral fins wide
and long, more than 18%
of body length (Fig. 32). . . . . . . Tetrapturus pfluegeri
Fig. 31
Atlantic Ocean T. belone

T. pluegeri Fig.32
 - 18 -

5b. Anterior lobe of first dorsal fin

somewhat higher than remainder of second anal fin
t h e f i n , the height decreasing
gradually backward (Figs 34,35,36); anus
anus situated near origin of first first anal fin Fig. 33
a n a l f i n (Fig.33), t h e d i s t a n c e
b e t w e e n t h e m (X) s m a l l e r t h a n
height of first anal fin (Y) scales
soft

8a. Tip of first dorsal fin, pectoral

fins and first anal fin rounded
(Figs 34,35)

9a. First dorsal fin unspotted; T. georgei Fig. 34

s c a l e s o n midbody s o f t
and rounded (Fig.34).........Tetrapturus georgei
Eastern Atlantic
and Mediterranean

9b. First dorsal fin spotted;

s c a l e s o n midbody p u n -
gent, each scale elongate
with 1 or 2 spines (Fig.
35) . . . . . . . . . . . . . . . . . . . Tetrapturus albidus
T. albidus Fig. 35
Atlantic Ocean

8b. Tip of first dorsal fin, pectoral

fins and first anal fin pointed
(Fig.36) . . . . . . . . . . . . . . . . . . Tetrapturus audax
Pacific and Indian oceans

4b. Height of anterior lobe of first dorsal

fin (X) lower than body depth (Y); nape
highly elevated (Fig.37); body not com-
pressed laterally; vertebrae 11+13=24 T. audax Fig. 36

10a. Pectoral fins can be folded back

against sides of body

11a. Lateral line system with

simple loops (Fig.38) . . . . . . . . . . . Makaira mazara
Pacific and Indian oceans Fig. 37

l111b. Lateral line system r e t i -

culated (Fig.39) . . . . . . . . . .. Makaira nigricans
Atlantic Ocean

10b. Pectoral fins rigid, c a n n o t b e

folded back against sides of body
(Fig.40 ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Makaira indica
(Fig.40)
M. indica Fig. 38
Pacific and Indian oceans
not able to be folded back
to this position

pectoral fin rigid

M. nigricans Fig. 39 M. indica Fig. 40
 -- 19 -

2.2 Additional Aids to Identification of Genera and Species

The shape of vertebrae of billfishes shows characteristic variations by species or genus and may be used as
an additional aid to species identification, especially in the case of specimens that are damaged or cut (Fig.41)
anterior neural zigapophysis posterior neural zigapophysis
neural spine

hypural
plate

X. gladius haemal spine

anterior neural zigapophysis posterior neural zigapophysis

neural spine

I. platypterus
anterior haemal zigapophysis
haemal spine

I. albicans

T. angustirostris
precaudal vertebrae caudal vertebrae

T. pfluegeri

T. albidus

T. audax

M. maxara

M. nigricans

M. indica

Vertebrae of billfishes (modified after Nakamura, 1983, Fig. 10) Fig.41

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20

2.3 FAMILY ISTIOPHORIDAE ISTIO

Synonymy : Histiophoridae; Makairidae; Tetrapturidae.

FA0 Names: En - Marlins, Sailfishes, Spearfishes; Fr - Makaires, Marlins, Voiliers; Sp -- Agujas,

Marlines, Peces vela.

Field Marks : Bill long, round in cross-section;

double keels on each side of caudal peduncle; scales
ossified and elongate, each with one to several points
posteriorly; pelvic fins rigid, string-like, depressible into
a ventral groove; vertebrae 24, with plate-like spines.

Diagnostic Features : Premaxilla elongate, forming

a long, pointed rostrum, round in cross-section; jaws with
file-like teeth in adults; branchiostegal membranes
entirely united to each other and free from isthmus;
gillrakers absent. Caudal peduncle fairly depressed and
slightly compressed, with 2 keels on each side and a
shallow notch on dorsal and ventral profiles. Pelvic fins
with one spine and 2 soft rays fused together, depressible
into a groove. Body covered with elongate scales, each
with one to several points. Vertebrae 24, with plate-like
neural and haemal spines (triangular to square).

Habitat, Distribution and Biology : Istiophorids are

primarily inhabitants of warm seas, usually swimming in
the upper layers of water above the thermocline. They
often migrate into temperate or cold waters to feed on
forage organisms during the warmer season and back to
warm waters for spawning or over-wintering during the
colder season. Being among the largest and swiftest
teleosts of the oceans, they perform considerably long
migrations but only few records of transoceanic migra-
tions are known for this group.

Interest to Fisheries : All istiophorid species have

some commercial value throughout the world, but they
are most highly priced in Japan which is the major
consumer nation for this group of fishes. Most of the
species are exploited commercially by longline fleets and
all are regarded as excellent game fish by sports fisher-
men.

Remarks :The species of this family undergo dras-

tic morphological changes during growth, as is shown in
the illustrated example of Makaira mazara.

Body changes of Makaira mazara with growth

(schematic)

ISTIO Istio
Istiophorus Lacepède, 1801

Genus : Istiophorus Lacepède, 1801, Histoire naturelle des poissons, 3:374-5

Type Species : Istiophorus gladifer Lacepède, based on “Le voilier” of Broussonet, Mem.Acad.Sci., 1786:
450-5, pl. 10.
Synonymy : Nostidium Hermann, 1804; Histiophorus Cuvier, in Cuvier and Valenciennes, 1832; Skeponopodus
Nardo, 1833; Zanclurus Swainson, 1839.
Remarks: There are many research workers who consider that this genus is monotypic, including only
Istiophorus platypterus. Others recognize two species (or subspecies).
 -- 21 -

ISTIO Istio 1
Istiophorus albicans (Latreille, 1804)

Makaira albicans Latreille in Bose & Latreille, 1804, Nouveau Dictionnaire d’Histoire Naturelle, 1(24):104
(Brasil).
F
Synonymy: Histiophorus americanus Cuvier in Cuvier & Valenciennes, 1832; Histiophorus pulchellus Cuvier
in Cuvier & Valenciennes,1832; Makaira velifera Cuvier, 1832; Skeponopodus guebucu Nardo, 1833; Histiophorus
granulifer Castelnau, 1861; Xiphias velifer Rochebrune, 1882; Istiophorus americanus-Jordan, 1885; Istiophorus
wrigthi Jordan & Evermann, 1926; Istiophorus maguirei Jordan & Evermann, 1926; Istiophorus volador Jordan &
Evermann, 1926; Histiophorus albicans-Whitley, 1936; Istiophorus albicans-Whitley, 1937.
Fao Name :: En - Atlantic sailfish; Fr - Voilier de Il'Atlantique; Sp - Pez vela del Atlántico.

Field Marks: First dorsal fin sail-like and remarkably higher than greatest body depth; pelvic fins very
long, nearly reaching to anus, with a well developed membrane; pectoral and caudal fins long compared with those
of Istiophorus platypterus in young fish up to about 90 cm body length.

Diagnostic Features : Body fairly com-

pressed. Bill long, slender and round in cross sec-
tion; jaws and palatines with small, file-like teeth
in adults; no gillrakers; right and left branchio-
stegal membranes united to each other, free from
isthmus. Two dorsal fins, the first large, with 42 to
46 rays, the second small with 6 or 7 rays; first
dorsal fin sail-like, remarkably higher than body
depth at level of midbody, its base long and close to
that of second dorsal fin; two anal fins, the first Immature of I. albicans ca. 90 cm. body length
with 11 to 14, the second with 6 or 7 rays; position ( screened areas show characters of I. platypterus )
of second anal fin slightly more forward than that of
second dorsal fin; pectoral fins with 17 to 20 rays, relatively longer than in I.
platypterus in immature specimens (to about 90 cm body length); pelvic fins extremely
long, almost reaching to anus, depressible into a groove, with one spine and several
soft rays fused tightly together and with a well developed membrane. Caudal peduncle
with double keels on each side and a shallow caudal notch on both, the dorsal and the
ventral surfaces; anus situated near to first anal fin origin. Lateral line single and
well visible. Scales vary in shape with growth (see I. platypterus); in adults they are
somewhat sparse, imbedded in the skin, each with a single, rather blunt point or two
posterior points. Vertebrae 24 (12 precaudal and 12 caudal). Colour: body dark blue
dorsally, light blue splattered with brown laterally, and silvery white ventrally; about
20 rows of longitudinal stripes on sides, each stripe composed of many light blue round
dots. Bases of first and second anal fins often tinged with silvery white; membrane of
scales
first dorsal fin dark blue or blackish blue, with scattered small round black dots;
remaining fins blackish blue, sometimes tinged with dark brown.
 -- 22 -

Geographical Distribution : I. albicans is widely

distributed in the tropical and temperate waters of the
Atlantic Ocean. The latitudinal range, based on data
from longline catches, is approximately 40°N in the
western North Atlantic, 50°N in the eastern North Atlan-
tic, 40°S in the western South Atlantic, and 32°S in the
eastern South Atlantic. However, the southern range limit
of this species in the eastern South Atlantic near the
Cape of Good Hope is uncertain because of possible joint
occurrence of both sailfish species in that area, and the
difficulty of separating these accurately in the field.
The Atlantic sailfish is also known to migrate into the
Mediterranean Sea, although the records from this area
are few and mostly based on juvenile specimens. Further-
more, in at least one case (Ben-Tuvia, 1966) a specimen
recorded from the Mediterranean as sailfish has been
subsequently identified as Tetrapturus belone.

Habitat and Biology : This is an epipelagic and

oceanic species, usually found in the upper layers of warm
water above the thermocline (temperature range between
21° and 28°C), but also capable of descending to rather
deep water. This is the least oceanic of the Atlantic
billfishes, and it often migrates into near-shore waters.
In the western Atlantic, it concentrates particularly in
the Caribbean Sea, the Gulf of Mexico and around the
West Indies and Florida. The distribution along the east coast of the USA appears to be influenced by meteoro-
logical conditions (wind) and by water temperatures; during summer, the fish move northward along with the
extension of warm water, and with the beginning of cold weather and northerly winds they are driven back south-
ward to congregate in schools off the Florida coast. In the eastern Atlantic, the shifts of the frontal zone of the
Canaries Current and the Equatorial Countercurrent are responsible for the aggregation of the populations off
West Africa; in spring the population in that area moves northward along the coast, and apparently returns toward
the south in autumn, following the 28°C isotherm; the period of increased abundance off Ivory Coast coincides
with the period of maximum surface water temperture, around 28°C.
I. albicans occasionally forms schools or smaller groups of 3 to 30 individuals, but more often occurs in loose
aggregation over a wide area. Observations on the die1 activity of this species off Florida have shown scattering
of the population in the early morning, but by 09.00 hours schools of up to 30 individuals begin to form and feed on
concentrations of small forage fish.

Mather et al. (1974) reported on the results of 18 years of billfish tagging under the Woods Hole
Oceanographic Institution’s Cooperative Gamefish Tagging Program; 12 525 sailfish were tagged and 97 recoveries
recorded; the longest distance between the points of release and recovery was about 3 070 km, from off Cape
Hatteras, North Carolina (USA) to off Guianas; the longest time at liberty of tagged fish was about 4
years.

The Atlantic sailfish is heterosexual, but there are no external morphological characters or colour to
distinguish males and females; large fish are usually females. Around Florida, this species often moves inshore
into shallow waters where the females, swimming sluggishly with their dorsal fins extended and accompanied each
by one or more males, may spawn near the surface in the warm season (Voss, 1956; Jolley, 1977). However,
Gehringer (1956), stated that spawning occurs in offshore waters beyond the 100 fathom isobath and appears to
extend from April to September from south of Cuba to Carolina. In the eastern Atlantic, spawning has been
observed in West African shelf waters throughout the year with a peak intensity during the summer months in the
Conakry-St. Luis region, and from February to April in the Conakry-Freetown region (Ovchinnikov, 1970). Off
southeast Florida, the presence of three distinct groups of maturing ovocytes in ripe ovaries reveals that ovocyte
development is asynchronous, this resulting in fractional or multiple spawning; a 33.4 kg female may shed up to
4.8 million eggs in three batches during one spawning season (Jolley, 1977).
Although I. albicans probably competes for food with many other large pelagic fishes, such as other
billfishes, tunas, sharks and dolphinfishes, it is most likely less affected by food competition than the other
species. In the Atlantic it overlaps in geographical range and hence, competes for food during certain seasons of
the year, particularly with Tetrapturus albidus and Makaira nigricans.
The food of larval Atlantic sailfish consists primarily of copepods, but the diet of the larvae changes rapidly
to include predominantly fishes after an increase in size of only a few millimetres. Around Florida, adult I.
albicans have been shown to feed mainly on pelagic fishes such as Euthynnus alletteratus, Hemiramphus spp.,
Trichiurus lepturus, Strongylura notatus, Caranx ruber, Lagodon rhomboides and squids, e.g., Argonauta argo,
Ommastrephes bartrami, etc. They also often feed on bottom-dwelling organisms including sea robins (Triglidae),
cephalopods and gastropods. This shows that I. albicans does not feed only in surface waters but also near the sea
bottom down to considerable depths.
 -- 23 -

Size : This species reaches a maximum size beyond 3.15 m in total length and 58 kg in weight. The all-
tackle angling record is a 58.10 kg fish taken off Luanda (Angola) on 27 March 1974. Other large records include
individuals of 58 kg in 1975 and 52.75 kg in 1972 (both off Louanda) and 58 kg in 1979 from Cancun, Quintana Roo,
Mexico (IGFA, 1981). The all-line-class world records recognized by IGFA are from West Africa (Angola and
Senegal) and Mexico (Cancun, Quintana Roo). The largest fish recorded by sportsfishermen measured 3.15 m in
total length and 55.8 kg weight (Walker Bay, Bahamas, 25 April 1950).

The majority of sportsfishing catches in southern Florida range from about 102 to 140 cm body length (or
173 to 229 cm total length) with considerable differences in weight (from 6.0 to 49.4 kg). The second largest size
group ranges from 61 to 94 cm body length, averaging less than 6 kg in weight. The small fish begin to appear in
the catches in late summer, becoming prominent in November, December and January. The body length of
individuals caught by commercial tuna longliners in the Atlantic ranges from about 125 to 210 cm (mostly between
150 and 195 cm).

Interest to Fisheries : Catches of I. albicans in the period from 1978 to 1982 have been reported from five
FA0 major marine fishing areas (21, 31, 34, 41 and 47), mostly by Ghana, Japan, Brazil and the USSR. The total
world catch was 267 metric tons in 1978, 2 823 t in 1979, 1 320 t in 1980, 1 091 t in 1981 and 920 t in 1982. More
than 88% of the total catch from 1979 to 1982 was taken in Fishing Area 34 (eastern central Atlantic),
predominantly by Ghana (FAO, 1984).
The principal commercial fishing gear for I. albicans is the tuna longline used primarily for tunas and
marlins, and hence the Atlantic sailfish is a byproduct of the fisheries for these species. Trolling is the primary
method used by sportsfishermen, mainly by towing baited hooks and lures through the surface water to stimulate
swimming fish.

The flesh is good in summer, but usually not as good as that of marlins. It is marketed mostly frozen and
often fresh in local markets.

Local Names: B R A Z I L : Agulhão, Agulhão bandeira, Agulhão de vela, Agulhão vela, Bicudo, Guebuçù;
CANADA: Sailfish; CUBA: Abanico, Aguja de abanico, Aguja voladora, Bicuda, Prieta, Voladeira; FRANCE:
Voilier, Voilier de 1’Atlantique; GHANA: Adzietekwesi, Fetiso, Onyankle; GOLD COAST: Fetiso; JAPAN:
Nishibashoo, Nishibashookajiki (names for Atlantic sailfish landed in Japan); MEXICO: Pez vela, Volador;
MOROCCO: Espadon; NETHERLANDS: Zegal-fisch; PORTUGAL: Bicuda, Espardarte veleiro, Peixe de vela,
Veleiro, Veleiro do Atlantico; SENEGAL: Espadon, Oumbajhe; SPAIN: Pez vela, Pez vela de1 Atlantico; SOUTH
AFRICA: Sailfish, Seilvis; UNITED KINGDOM: Sailfish; USA: Atlantic sailfish, Sail, Sailfish; USSR: Parusnik,
Parusnik-ryba; VENEZUELA: Aguja vela; WEST INDIES: Balahoo, Billfish, Mère, Ocean gar, Sailfish, Squadron.

Literature : Voss (1953); de Sylva (1957); Cadenat (1961); Tinsley (1964); Wise & Davis (1973); Jolley (1974,
1977); Beardsley, Merrett & Richards (1975).

Remarks : Adults of I. albicans and I. platypterus are extremely similar and very difficult to distinguish. In
the immature stage (to about 90 cm body length), this species has longer pectoral and caudal fins than I.
platypterus; this seems to be due to the more rapid growth of I. albicans which attains a much smaller maximum
size than I. platypterus. Further study is required, to clarify the speciation problem of the two sailfish species.
For the time being, the author prefers to retain the use of Istiophorus albicans (Latreille, 1804) for the Atlantic
sailfish and Istiophorus platypterus (Shaw & Nodder, 1792) for the Indo-Pacific sailfish, but other research workers
recognize a single species under the name Istiophorus platypterus.

Istiophorus platypterus (Shaw & Nodder, 1792) 1 ISTIO Istio 2

Xiphias platypterus Shaw & Nodder, 1792, Nat.Misc., (28):no pagination, pl. 88 (Indian Ocean).
Synonymy : Scomber gladius Bloch, 1793; Istiophorus gladifer Lacepède, 1801; Xiphias velifer Schneider, in
Bloch & Schneider, 1801; Histiophorus indicus Cuvier, in Cuvier & Valenciennes, 1832; Histiophorus immaculatus
Rüppel 1835; Histiophorus orientalis Schlegel in Temminck & Schlegel, 1842-50; Histiophorus gladius-Günther,
1860; Istiophorus dubius Bleeker, 1873; Istiophorus triactis Hemprich & Ehrenberg, 1899; Istiophorus japonicus
Jordan & Thompson; Istiophorus gladius-McCulloch, 1921; Istiophorus orientalis-Jordan & Snyder, 1901; Istiophorus
eriquius Jordan & Ball in Jordan & Evermann, 1926; Istiophorus Greyi Jordan & Evermann, 1926; Istiophorus greyi-
Jordan, Evermann & Clark, 1930; Istiophorus brookei Fowler, 1933; Istiophorus ludibundus Whitley, 1933;
Istiophorus immaculatus-La Monte & Marcy, 1941; Istiophorus amarui Curtis, 1944; Istiophorus gladius greyi-
Nichols & Murphy, 1944; Istiophorus greyii-de Buen, 1958; Istiophorus platypterus-Whitehead, 1964.

FAO Name : En - Indo-Pacific sailfish; Fr - Voilier de l’Indo-Pacifique; Sp - Pez vela del Indo-Pacífico.
 -- 24 -

Field Marks : First dorsal fin sail-like and remarkably higher than greatest body depth; pelvic fins very
long, nearly reaching to anus, with a well developed membrane. Pectoral and caudal fins short compared with
those of Istiophorus albicans in young up to about 90 cm body length.

Diagnostic Features: : Body fairly compressed.

Bill long, slender and round in cross section; jaws and
palatines with small, file-like teeth; no gillrakers;
right and left branchiostegal membranes united to
each other, free from isthmus. Two dorsal fins, the
first large with 42 to 49 rays, the second small with 6
or 7 rays; first dorsal fin sail-like, with the middle
rays longest, remarkably higher than body depth at
level of midbody, its base long and close to that of Immature of I. platypterus ca. 90 cm. body length
second dorsal fin; two anal fins, the first with 12 to ( screened areas show characters of I.-___
albicans )
17, the second with 6 or 7 rays; position of second
anal fin slightly more forward than that of second
dorsal fin; pectoral fins with 18 to 20 rays, relatively
shorter than in I. albicans in immature specimens (to
about 90 cm body length pelvic fins extremely long,
almost reaching to anus, depressible into a groove,
with one spine and several soft rays fused tightly
together and with a well developed membrane. Caudal
peduncle with double keels on each side and a shallow
notch on both, the dorsal and the ventral surfaces;
anus situated near to first anal fin origin. Lateral line
single and well visible. Scales varying in shape with ca. 15 cm ca. 21 cm
growth; in adults they are somewhat sparse, imbedded body length
body lenght
in the skin, each with a single, rather blunt point, or
with two posterior points. Vertebrae 24 (12 precaudal
and 12 caudal). Colour: body dark blue dorsally, light
blue splattered with brown laterally, and silvery white
ventrally; about 20 rows of longitudinal stripes on
sides, each stripe composed of many light blue round
dots. Bases of first and second anal fins often tinged
with silvery white; membrane of first dorsal fin dark
blue or blackish blue, with scattered small, round
black dots; remaining fins blackish brown or dark blue.
ca. 90 cm ca. 200 cm
body length body length

Schematic drawings of scales

(not same size)
 -- 25 -

Geographical Distribution : I. pla- I.

typterus is widely distributed in the tro-
pical and temperate waters of the Paci-
fic and Indian oceans. Its latitudinal
range, based on data from longline
catches, is approximately 45° to 50° N in
the western North Pacific, 35° N in the
eastern North Pacific, 40° to 35°S in the
western South Pacific, 35°S in the eas- -
tern South Pacific, 45°S in the western
Indian Ocean, and 35°S in the eastern
Indian Ocean. This species shows a
strong tendency to come close to the
shore, even though a few individuals
have been caught in the central parts of
the oceans. Large numbers of Indo- Indo-
Pacific sailfish are found in waters off
Papua New Guinea, around the Solomon
Islands, in the warm Kuroshio Current and its branch, the Tsushima Current, in the Sea of Japan, in the East China
Sea, around the Philippines, in the Banda Sea, in the central South Pacific from Tahiti to the Marquesas, and off
the Pacific coast of Mexico. They are also fairly abundant off the northern coast of Australia and around the
Hawaiian Islands, India and Sri Lanka, but have been seldom seen off the Pacific coast of South America from
Peru to Chile. This species has also entered the Mediterranean Sea from the Red Sea through the Suez Canal.

Habitat and Biology

: : The Indo-Pacific sailfish is an epipelagic and oceanic species, usually found above
the thermocline. It shows a strong tendency to approach continental coasts, islands and reefs.

In the western Pacific, the distribution of postlarvae and adults appears to be closely related to the Kuroshio
Current, and the densest concentrations seem to coincide with the spawning season. Individuals over 160 cm eye-
fork length (=(84 to 88.7 % of body length) migrate southward out of the East China Sea, presumably for spawning.
In the eastern Pacific, the seasonal north-south displacements of this species off the coast of Mexico appear to
coincide with the seasonal movements of the 28’C 28°C water isotherm. In the Indian Ocean, off East Africa, the
abundance and distribution of I. platypterus is positively correlated with the months of the northeast monsoons
when the East African coastal current reaches its maximum temperature (28° to 30° C) and minimum salinity
(35.2 to 35,3% 0 ). This also the time of highest biological productivity in the surface waters caused by a
mixing of waters resulting from the junction of the southward-flowing Somalia Current and the northward-flowing
East African Coastal Current. In the Sea of Japan, considerable numbers of individuals of this species migrate in
schools northward with the warm Tsushima Current (a branch of the Kuroshio) during summer (peak in later
summer), and southward against the current during autumn (peak in early autumn), and are caught in these periods
by the coastal setnets. These schools consist of young (55 to 110 cm body length) and adults (145 to 235 cm body
length) with no fish of intermediate sizes caught by the setnets. Sometimes the young and the adults are mixed in
the catches, but more often they are landed separately. Therefore, I. platypterus most likely schools by size.
Usually, the young form more dense schools than the adults. In the East China Sea, the Indo-Pacific sailfish
migrates northward in summer and returns southward in autumn where it overwinters in the southernmost area;
fish under 160 cm eye-fork length move into the area between May and July while those over 160 cm migrate
southward out of the area, probably for spawning.

Spawning of this species occurs with males and females swimming in pairs or with two or three males
chasing a single female (probably mating behaviour). I_
I. platypterus seems to spawn throughout the year in tropical
and subtropical waters of the Pacific with peak spawning occurring in the respective local summer seasons. The
ripe ovarian eggs are about 0.85 mm in diameter and have a single oil globule surrounded by a pale yellow
indefinite nimbus; there are no structures on the vitelline
vitelline membrane and the egg as a whole is transparent or
translucent. Eggs shed from a captured female in the Indian Ocean averaged 1.304 mm in diameter.

The feeding behaviour of I. platypterus has been observed by fishermen as follows: when one or several
sailfish found a school of prey fishes sardines,
(sardines, anchovies, mackerels, or jack mackerels), they began to pursue it
at half speed with their fins half-folded back into the grooves. They then drove at the prey at full speed with
their fins completely folded back and once they had caught up with it, they suddenly made sharp turns with their
fins expanded to confront a part of the school and then hit the prey with the bill. Subsequently they ate the killed
and stunned fish, usually head first. Surprisingly, several individuals showed a kind of team-behaviour in capturing
the prey .
I. platypterus and Makaira indica are the billfish
billfish species dominantly migrating into inshore waters and they
undoubtedly compete with each other for food and habitat, although M. indica probably feeds on larger forage
organisms, and swims a little deeper and farther offshore than thesailfish.
the sailfish. The habitat of this species is
strikingly different from that of the closely related Tetrapturus angustirostris which does not usually occur within
500 km off the coast, while M. indica is mostly distributed within this range.
 -- 26 -

The major forage items of the lndo-Pacific sailfish are fishes and squids, but the adults are fairly
opportunistic feeders and take almost any food they come across. During several longline cruises of Japanese
research vessels in the eastern North Pacific Ocean, adult I. platypterus have been shown to feed mainly on
cephalopods and fishes (Bramidae, Stromateidae, Carangidae, Ostracion spp., Gempylidae, Auxis spp., Trachip-
terus spp., Belonidae, Balistidae, Coryphaena spp., Lagocephalus spp., etc.).

Large pelagic sharks, the killer whale and other related species attack billfishes and tunas hooked by
longlines, but predation on free-swimming tunas and billfishes is thought to be very rare.

Size : This species reaches a maximum size beyond 340 cm in total length and 100 kg in weight. In the
sports fishery (all tackle angling record) corresponds to a fish of 327.7 cm total length and 100.24 kg (221 lb)
weight, taken at Santa Cruz Island, Galapagos, Ecuador on 12 February 1947. Another very large fish (340.4 cm
total length and 89.81 kg (198 lb) weight) was taken at La Paz, Baja California, Mexico on 23 August 1957. Other
records over 85 kg are the following: 90.26 kg (199 lb) at Pinas Bay, Panama on 17 January 1968; 89.81 kg (198 lb)
at Mazatlan, Mexico on 10 November 1954; 87.54 kg (193 lb) at Acapulco, Mexico on 8 January 1978; 87.28 kg
(192.7 lb) at Acapulco, Mexico on 4 October 1961; 87.09 kg (192 lb) at La Paz, Baja California, Mexico on 6
September 1950; and 85.72 kg (189 lb) at Yanuca, Fiji on 7 December 1967 (IGFA, 1981). All-line-class world
records up to 1981 for both men and women recognized by IGFA are from the eastern Pacific, except the one for
the women’s 130-lb line class of 189 lb already mentioned from Yanuca, Fiji.

In sportsfishing at Malindi, Kenya, the majority of individuals caught ranged from 203 to 254 cm fork length
(224 to 279 cm total length) and from 18.1 to 47.2 kg in weight. Length frequency data for sailfishes caught by
longlines in the East China Sea, give size range from 105 to 240 cm body length (60% between 165 and 190 cm
body length); the size distribution is essentially unimodal, except in June when a group of 125 to 150 cm fish
suddenly appeared in the catch. Koto, Furukawa & Kodama (1959) believed that these small fish enter the East
China Sea from other areas during this month. The average size of the individuals caught by longlines is unimodal,
about 140 to 240 cm body length, while it is bimodal in those caught by driftnets and setnets, about 50 to 110 cm
and 140 to 240 cm body length, respectively.

Interest to Fisheries : In the period from 1978 and 1982 catches of I. platypterus have been reported from
seven FAO Fishing Areas (51, 57, 61, 71, 77, 81 and 87), predominantly by Japan and the Republic of Korea. The
total world catch was 10 516 t in 1978, 7 916 t in 1979, 7 767 t in 1980, 6 438 t in 1981 and 7 214 t in 1982. Only
4% (285 t) of the 1982 total world catch were taken in the Indian Ocean, and 96% (6 929 t) in the Pacific Ocean
particularly in Fishing Areas 61 and 77 (northwest Pacific and eastern central Pacific) of which 6 218 t were taken
by Japan, China (Taiwan Province) and the Republic of Korea. In Fishing Area 77 (eastern central Pacific), the
Korean and Japanese landings have decreased remarkably in recent years: 5 425 t (1978), 4 351 t (1979), 1 525 t
( (1980), 539 t (1981) and 2 483 t (1982) (FAO, 1984). It should be noted that the catch statistics given for this
species by Japanese longliners include Tetrapturus angustirostris, but that the share of the latter species in these
catches is quite negligible.
lndo-Pacific sailfish are often taken as bycatch by the commercial surface tuna longliners. They are also
caught by commercial fishermen with surface driftnets, and by trolling, harpooning and setnetting. In sportsfishing
this species is caught by surface trolling.

The flesh is dark red, and not as good as that of marlins. In summer it is good for sashimi (sliced flesh with
soy-sauce and horse radish) or sushi (vinegared boiled rice with sliced flesh and horse radish, dipped in soy-sauce
during the meal).

Local Names : A U S T R A L I A : Bayonet fish, Pacific sailfish, Sailfish; CHINA: Ho-soan-ki-hi, Tong-fang-
chiyii, Yu-san-chi-yu, Yu-san-yu; COMOROS: Mbassi kouri; INDIA: Fung-hibaru, Mylmeen, Ola-meen, Peacock
fish, Sailfish, Tadmasa, Yemungolah; INDONESIA: Djangilus, Geulang pajang, Hohoo malays, lkan jegan, lkan laya,
lkan layar, Lijarang, Landjareng, Lkajar plajaren, Panombuk, Stuji stuji; JAPAN: Akitaroo, Atsutaro, Banba,
Baren, Barin, Bashoo, Bashookajiki, Byoobu, Byoobusashi, Haio, Haioshibi, Haou, Hauo, Kanga, Kannushi, Koomori,
Minokajiki, Nourage, Oba, Suginairage, Suginairagi, Sugiyama, Tobihira; KENYA: Mbassi, Nsulinsuli;
MADAGASCAR: Ndwaro; MALAYSIA: Mersuji, Layeran; MEXICO: Pez vela, Volador; NEW CALEDONIA:
Empereur éventail, Voilier; PHILIPPINES: Dogso, Dugso, Dumosok, Kandelan, Kandayan, Liplipan, Malasugi,
Sailfin, Sailfish; REPUBLIC OF KOREA: Dot-sae-chi; SRI LANKA: Mylmeen, Thalapatha; TAHITI: Haurepe;
TANZANIA: Mbassi, Nsuli nsuli; USA: Pacific sailfish, Sailfish; USSR: Parusnik, Parusnik-ryba; VIET NAM: C á co.

Literature : Nakamura, H. (1937, 1938, 1940, 1942, 1949, 1951); Koto, Furukawa & Kodama (1959); Koto &
Kodama (1962); Ueyanagi (1963, 1963a, 1964); Tinsley (1964); Williams (1964, 1970); Kume & Joseph (1969, 1969a);
Nakamura, I. (1974, 1983); Beardsley, Merrett & Richards (1975).

Remarks: See remarks for lstiophorus albicans.

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-- 27 -

Makaira Lacepède, 1802 ISTIO Mak

Genus : Makaira Lacepède, 1802. Histoire naturelle des poissons, 1802, 4:688-95, pl.13 (fig.3).

Type Species : Makaira nigricans Lacepède, 1802.

Synonymy : Machaera Cuvier, 1832 (amended spelling); Macaira Nardo, 1833 (amended spelling); Istiompax
Whitley, 1931; Marlina Hirasaka & Nakamura, 1947; Eumakaira Hirasaka & Nakamura, 1947; Orthocraeros Smith,
1956; Istiomax Abe, 1963 (? missprint).
Eumakairazka
Remarks: There have been different opinions regarding the species composition of this genus (see discussion
on page 3.5).

Makaira indica (Cuvier, 1832) ISTIO Mak 2

T e t r a p t u r u s i n d i c u s Cuvier, in Cuvier & Valenciennes, 1832, Histoire Naturelle des Poissons, 8:209-10
(Sumatra, Indonesia). Indonesiar

Synonymy : Tetrapturus australis Macleay, 1854; Histiophorus brevirostris Playfair, in Playfair & Günther,
1866; Tetrapturus brevirostris-Goode, 1882; Makaira marlina Jordan & Hill, in Jordan & Evermann, 1926; Makaira
indica-Jordan & Evermann, 1926; Istiompax australis-Whitley, 1931; Makaira indicus-Deraniyagala, 1933; Makaira
australis-Fowler, 1934; Makaira nigricans marlina-Nichols & LaMonte, 1935; Makaira nigricans tahitiensis Nichols
& LaMonte, l935; Makaira brevirostris-LaMonte &Marcy, 1941; Makaira ampla marlina-LaMonte & Marcy, 1941;
M a k a i r a a m p l a t a h i t i e n s i s - L a M o n t e & M a r t y , 1 9 4 1 ; M a r l i n a marlina-Hirasaka & Nakamura, 1947; Malina malina-
Chen, 1951; Makaira marhina-Mori, 1952; Istiompax dombraini Whitley,, 1954; IstioMakaira marhina-Chyung, 1954;;
M a k a i r a m a z a r a t a h i t i e n s i s - L a M o n t e , 1 9 5 5 ; M a k a i r a x a n t h o l i n e a t u sDeraniyagv56;
D e r a n i y a g a l a , 1 9 5 6 ; Makaira herscheri-
Fournmanoir, 1957; Makaira marlina marlina-Morrow, 1957; Makaira marlina tahitiensis-Morrow, 1 9 5 7 ; I s t i o m p a x
marlina-Royce, 1957; Istiompax brevirostris-Morrow, 1958; Istiompax indicus-Morrow, 1959; Makaira (Istiompax)
indica-Robins & de Sylva, 1961; Istiomax indicus-Abe, 1963.

FAO Names : En - Black marlin; Fr - Makaire noir; Sp - Aguja negra.

Field Marks : Body not very compressed; nape highly elevated; height of anterior lobe of first dorsal fin
smaller than greatest body depth; second dorsal fin slightly forward of second anal fin; pectoral fins rigid, not
adpressible against sides of body.

Diagnostic Features : Body not strongly compressed. Bill long, extremely

stout, and round in cross section; nape conspicuously elevated; right and left
branchiostegal membranes completely united to each other, but free from isthmus; no
gillrakers; both jaws and palatines (roof of mouth) with small, file-like teeth. Two
dorsal fins, the first with 34 to 43 rays, low (anterior lobe lower than body depth); with
a long base ending close to second dorsal fin origin; second dorsal fin with 5 to 7 rays,
its position slightly forward with respect to that of second anal fin; two anal fins, the
first with 10 to 14 and the second with 6 or 7 rays; pectoral fins with 12 to 20 rays,
rigid, not adpressible to sides of body; pelvic fins with a poorly developed membrane,
shorter than the pectorals and depressible into ventral grooves. Caudal peduncle fairly
scales
 -- 28 -

compressed (laterally) and slightly depressed (dorso-ventrally), with strong double keels on each side and a poorly
developed notch on both, the dorsal and ventral surfaces; anus situated near first anal fin origin. Lateral line
single but obscured, especially in larger fish. Body densely covered with thick, elongate bony scales, each with 1
or 2 (mostly 1) sharp posterior points. Vertebrae 24 (11 precaudal and 13 caudal). Colour: body dark blue dorsally
and silvery white ventrally; usually no blotches or dark stripes on body in adults, although light blue vertical
stripes may occur in a few fish. First dorsal fin blackish to dark blue; other fins usually dark brown, sometimes
tinged with dark blue.

Geographical Distribution :
The black marlin is distributed
throughout the tropical and sub-
tropical waters of the Pacific
and Indian oceans where the
main population and the spawn-
ing grounds occur, but it occa-
sionally also enters temperate
waters. Stray individuals have
been found to migrate into the
Atlantic Ocean by way of the
Cape of Good Hope, but the
existence of Atlantic breeding
stocks is unlikely. The latitudi-
nal range of this species, based
on Japanese commercial long-
liners’ catches extends north- Area of occasional distribution, or invasion, (no spawning)
ward to 35°-40°N in the North
Pacific and southward to 45°S 4in the western Pacific and 30°-35’S
30°-35°S in the eastern Pacific, and northward to 25°N
and southward to 45°S in the Indian Ocean.

Habitat and Biology: : This is an epipelagic and oceanic species usually found in surface waters above the
thermocline at temperatures ranging from 15° to 30°C,
30°C often in nearshore waters close to land masses, islands,
and coral reef areas. In tropical oceanic areas its distribution is continuous, although rather scattered, whereas
its presence in temperate waters is occasional.

M. indica usually occurs nearer to the surface than most other billfishes (except the sailfish), but this is by
no means a general rule. The commercial longline fishery generally operates in waters deeper than 100 m, and in the
equatorial western Indian Ocean, peak catch rates of black marlin have been reported to occur in the upper water
layers (0 to 200 m) over depths of 365 to 915 m (200 to 500 fathoms).

Like other billfishes, the black marlin effects seasonal migrations. In the East China Sea, schools of this
species move northward during spring and summer and southward during autumn and winter. In the Sea of Japan,
the north-bound migration occurs in late summer and early autumn following the core of the Tsushima Current (a
warm branch of the Kuroshio Current), and the return to southern waters takes place in autumn, with the fish
swimming against the current.

The sexes are separate in M. indica, but males and females are indistinguishable by external features. In the
waters around Taiwan Island, these-ratio was found to be 53/414 males in fish ranging from 20 to 200 kg weight.
As in other marlins, females become larger than males. M. indica is densely distributed in the northwestern part
of the Coral Sea between October and December. Almost all fish Acaught at that time in this area have well-
developed gonads, and are therefore believed to belong to spawning schools. The skewed sex ratio (541/615
females) may also be indicative of this possibility. Sportsfishermen from Cairns, Queensland (Australia)
eyewitnessed what might be the courtship behaviour of the black marlin: a group composed of one large fish
(supposed to be female) followed by several smaller fish (supposed to be males) was seen swimming on an
undulating course, sometimes back and forth, the fish being closer together than is usual in the shallow coral reef
areas near Cairns.

On the basis of records of the presence of larvae and mature females, spawning is believed to occur in the
vicinity of Hainan Island and in the South China Sea in May and June, around Taiwan Island from August to
October, in the northwestern part of the Coral Sea between October and December, and off Cairns (Australia)
from August to November. Black marlin are believed to prefer water temperatures around 27° to 28’C 28°C during
spawning. Egg counts of ripe roe totalled about 40 million per female.

Black marlins use their long and stout bill for feeding. Analyses of stomach contents show that the prey is
usually swallowed head first and bears slashes that had obviously been inflicted by the predator’s bill. The prey
includes skipjack (Katsuwonus pelamis), yellowfin tuna (Thunnus albacares), bigeye
bigeye tuna (Thunnus obesus), frigate
tunas (Auxis species) and other tuna species. The feeding of M. indica vary from area to area and by
seasons. In Pacific equatorial waters, food items include mainly Scombridae, Gempylidae, Coryphaenidae,
Xiphiidae and Carangidae, squids and cuttlefishes, while other groups, i.e., Sternoptychidae, Paralepididae,
Alepisauridae, Chiasmodontidae, Chaetodontidae, Balistidae, Ostraciidae and Tetraodontidae, octopods and large
decapod crustaceans are of lesser importance.
 -- 29 -

The larvae and juveniles of the black marlin are predated upon by pelagic carnivorous fishes such as large
sharks, scombrids, carangids, dolphinfishes and other billfishes. There are no true predators of the adults except
for the killer whale (Orcinus orca) and related species (only a few cases known). The main food competitors f o r
the black marlin are thought to be large sharks, swordfish, large tunas and other marlins.

Size : This species reaches sizes beyond 448 cm in total length and 700 kg in weight. The heaviest record
of black marlin in sportsfishing (the all tackle angling record) is a specimen of 442 cm (body length) and 707.61 kg
weight, caught at Cabo Blanco, Peru, on 4 August 1953. Considerable numbers of black marlin weighing in excess
of 500 kg are often caught off Cairns, Queensland, Australia. The measurements taken of the two female black
marlins in excess of 1 000 lb caught during the First International Black Marlin Tournament held at Cairns in
October 1973, were 448 cm total length (481.4 kg) and 437.2 cm total length (484.3 kg). Other examples of
gamefishing records are specimens of 426.7 cm total length (509.8 kg), 436.9 cm total length (691.7 kg) and
447 cm total length (552.5 kg).

The size range of black marlin taken by commercial longline fisheries is 150 to 310 cm (mostly 170 to
210 cm) body length in the western Indian Ocean and 170 to 310 cm (mostly 185 to 240 cm) body length in the
Coral Sea.

Interest to Fisheries : There are important longlining, harpooning, trolling and setnet fisheries for M.
indica in the Pacific as well as in the Indian Ocean. Catches are reported from five FAO Fishing Areas (51, 57,
61, 77 and 81) by various nations. The world catch was 2 909 t in 1978, 3 440 metric tons in 1979, 2 465 t in 1980
2 453 t in 1981 and 2 373 t in 1982. About 80 to 90% of this catch was taken by Chinese (Taiwan Province) vessels
in Fishing Area 61 (northwestern Pacific), amounting to 2 591 t (89%) in 1978, 3 183 t (93%) in 1979, 1 991 t (81%)
in 1980, 1 981 t (81%) in 1981 and 2 416 t (91%) in 1982 (FAO, 1984). In addition, it is estimated that Japan and
the Republic of Korea, though excluded from FAO statistics, are currently catching about 2 000 t of this species
in Fishing Area 61.

M. indica is mostly caught by ordinary surface tuna longliners. These vessels vary considerably in size, the
largest being about 2 000 GRT mother ships with several small catch boats on either side of the deck (this type of
ship stopped operating recently), but the majority are between 250 and 350 GRT, as this size appears to be the
most economical and efficient. Longlining vessels larger than 100 GRT are usually made of steel, while the
smaller boats are mostly wooden.

Harpoon-fishing is carried out on the fishing grounds off Taiwan Island and southern Japan. Small (about
10 m long) wooden, engine; powered boats are used in this activity. Optimum temperatures for harpoon-fishing for
this species range from 23° to 25° C (westward of Uotsurijima, in the East China Sea, between October and April).

Sportfishering off northeastern Australia, Peru and Ecuador is usually carried out from ordinary trolling
boats (36 to 42 feet long with inboard engines). In these areas, black marlins are more dominant among big game
fishes than on other sportsfishing grounds.
Finally, black marlins are also often taken by setnets displayed to catch the “yellowtail", (Seriola
quinqueradiata) in southern and western Japan during summer and autumn.

Good fishing grounds for black marlin (shallower than those of other billfishes) are found in the East China
Sea in the area where the Kuroshio and Tsushima currents mix with the waters of the Yellow Sea. The fishing
season extends from July to January (peak between August and October) between 30° and 34°N; from May to July
and from October to April (peaks in June and July and October and January, respectively), between 25° and 30°N,
and throughout the year around Taiwan Island (peak between October and December in the South China Sea and in
February and March off eastern Taiwan Island). Juvenile and young black marlin migrate in schools in July and
August, and in the coral reef areas of northern Queensland, adults come near the shore between September and
November.

Data on fishing effort and intensity are insufficient in all of the above areas. However, it is known that
most of the fishing effort and intensity are displayed in the East China Sea, the waters around Taiwan Island, off
northwestern Australia, in the Coral Sea, Arafura Sea, Sulu Sea, Celebes Sea and northwestern Arabian Sea.

Local Names : AUSTRALIA: Black marlin; CHINA: Kyau-shit-á, Lih-ch’ih-Ch’i-yü; JAPAN: Genba,
Katahari, Shiro, Shiroka, Shirokajiki, Shirokawa, Shirokawajiki, Shiromazaara, Shiruachi; NEW ZEALAND: Black
marlin; PAPUA NEW GUINEA: Black marlin; REPUBLIC OF KOREA: Baek-sae-chi; SRI LANKA: Ahin
koppara, Kopparaikulla, Kopparan, Kopparava, Makara, Marlin, Saparava; USA: Black marlin, Giant black marlin,
Pacific black marlin, Silver marlin; USSR: Sere bristyi marlin.

Literature : Howard & Ueyanagi (1965); Ueyanagi (1960); Merrett (1971); Nakamura (1975, 1983); Goadby
(1972); Howar d & Starck (1975); Mather (1976).
Remarks : The English common name “black marlin” had not been universally accepted for Makaira indica
until the midsixties. Prior to this agreement, many authors, particularly Japanese, called this species "white
marlin” which is a direct translation from the Japanese common name, “Shirokajiki” (shiro = white, kajiki = marlin)
The general colour of this species is black or blueblack while alive, but after death it changes to white and this
seems to be the cause of the above-mentioned nomenclatorial discrepancy.
 -- 30 -

Makaira mazara (Jordan & Snyder, 1901) ISTIO Mak 3

Tetrapturus mazara Jordan & Snyder, 1901, J.Coll.Sci.Imp.Univ.Tokyo , 15(2):305 (Japan).

Synonymy : Makaira mazara Jordan & Evermann, 1926; Makaira ampla mazara-LaMonte, 1941; Makaira
nigricans-Fowler, 1944 (many authors erroneously use the name Makaira nigricans for this species); Eumakaira
nigra Hirasaka & Nakamura, 1947; Makaira nigricans mazara-Rosa, name1950; Istiompax howardi Whitley, 1954;
Istiompax mazara-Whitley, 1968; Makaira nigra-Abe, 1957; Makaira (Makaira) nigricans-Robins & de Sylva, 1961.

FA0 Names : En - Indo-Pacific blue marlin; Fr -Makaire bleu de l’Indo-Pacifique; Sp - Aguja azul de1
Indo-Pacifico

Field Marks : Body not very compressed; nape

highly elevated; height of anterior lobe of first dorsal fin
less than greatest body depth; lateral line system with
simple loops.

Diagnostic Features : Body not strongly com- ca. 17 cm body length ca. 170 cm. body length
pressed. Bill long, extremely stout and round in cross Schematic drawings of scales (not same size)
section; nape conspicuously elevated; right and left bran-
chiostegal membranes completely united to each other,
but free from isthmus; no gillrakers; both jaws and
palatines (roof of mouth) with small, file-like teeth. Two
dorsal fins, the first with 40 to 45 rays, lower than body
depth throughout its length, with a pointed anterior lobe head . tail
and a long base, originating above the posterior margin of
preopercle and ending close to the second dorsal fin
origin; second dorsal fin with 6 or 7 rays, its position
slightly backward with respect to that of second anal fin;
two anal fins, the first with 12 to 17 rays and the second
with 6 or 7 rays; pectoral fins long and narrow, adpres-
sible to sides of body, with 20 to 23 rays; pelvic fins
shorter than the pectorals, with a poorly developed mem-
brane and depressible into deep ventral grooves. Caudal
peduncle fairly compressed (laterally) and slightly depres-
sed (dorsoventrally), with strong double keels on each side
and a shallow notch on both the dorsal and ventral
surfaces; anus situated near first anal fin origin. Lateral
line single, following a single loop pattern, obvious in
juveniles and immature fish, but obscure in adults, as it
becomes progressively imbedded in the skin with growth
(however, the line becomes always clearly visible when
the epidermis is removed). Body densely covered with
elongate, thick, bony scales, each with usually 1 or 2,
sometimes with 3 posterior points. Vertebrae 24 (11
precaudal and 13 caudal). Colour: body blue-black dor-
sally and silvery white ventrally, with about 15 rows of
Lateral line systems of left side in size growth
( schematic )
 -- 31 -

pale, cobalt-coloured stripes, each consisting of round dots and/or narrow bars (these stripes may not always be
visible, especially in long-preserved specimens). First dorsal fin blackish or dark blue, other fins blackish brown,
sometimes tinged with dark blue; bases of first and second anal fins tinged with silvery white.

Geographical Distribution : : The

In d o -Pac i fic b lu e ma rl i n i s f o u n d p r i -
marily in the tropical and subtropical
waters of the Pacific and Indian oceans.
It is the most tropical billfish species
and is frequent in equatorial waters. Its
latitudinal range, based on data from the
commercial longline fishery, extends to
about 45°N in the western North Pacific
O c e a n , 35°N in the eastern North Paci-
f i c 35°S i n t h e western South Pacific,
25°S in the eastern South Pacific 40 to
45°S in the southwestern Indian Ocean
a n d 35°S in the southeastern Indian
Ocean.

Habitat and Biology : This is an epipelagic and oceanic species mostly confined to the waters on the
warmer-side of the 24°C surface isotherm and known to effect seasonal north-south migrations. Pacific blue
marlins are not usually seen close to land masses or islands, unless there is a deep drop-off of the shelf (900 to
1 800 m as in the waters off Kailna-Kona, Hawaii Island).

The monthly distribution of catches by Japanese longliners shows two main seasonal concentrations of M.
mazara: one from December through March in the western and central South Pacific between 8° and 26°S, and
t h e other from May through October in the western and central North Pacific between 2° and 24°N. In the
remaining two months (April and November) the fish tend to concentrate in the equatorial Pacific between 10°N
and 10°S. This species becomes less abundant toward the eastern Pacific; in the Indian Ocean, it is known to be
relatively abundant around Sri Lanka and Mauritius; off the east coast of Africa, it is apparently abundant
between the equator and 13°S during the southeast monsoon period (from April to October).

In Hawaiian waters, an ecological interaction is known between M. mazara and the striped marlin,
Tetrapturus audax, with the two species responding in different, respectively; exclusive ways, to certain environ-
mental factors such as temperature or food.

Of 170 individuals of M. mazara tagged from 1963 through 1970 in the Pacific Ocean, no recoveries were
made up to 1972. Five specimens were tagged with ultrasonic transmitters and tracked off Hawaii in 1971 and
1972. A fish of 270 kg was successfully tagged and tracked on 14 and 15 July 1971, 3.1 miles west of Keauhou,
Hawaii Island (Yuen, Dizon & Uchiyama, 1974). The tag was inserted on 14 July at 09:35 hours and the fish was
tracked by the R.V. CHARLES H. GILBERT until 08.00 hours of the next morning. A temperature-sensitive tag
was used on this occasion for the purpose of obtaining information on depth. During the tracking period, the fish
moved to about 25 miles north of the point of release on an erratic course cruising between the 183 m and 1 830 m
(100 and 1 000 fathom) isobaths. The calculated speed of the fish ranged from 0.6 to 4.4 knots (0.09 to 0.62 body
length/sec) with an average of 1.6 knots (0.23 body length/sec). Swimming depth varied from the surface to 73 m,
but the fish remained mostly within the upper 37 m.

Larvae of M. mazara have been extensively collected in the tropical and subtropical waters of the western
and central Pacific, and south of the Maldive Islands, around the Mascalene Islands, and off the south
coasts of Java and Sumatra in the Indian Ocean. Ripe eggs in the ovary are transparent with a yellow oil globule,
and measure about 0.8 to 0.9 mm in diameter.

This species is believed by commercial fishermen to form small-scale schools consisting of at most ten
individuals. Larger fish tend to swim solitary.
M. mazara is known to feed in and near surface waters, but sometimes takes food in relatively deep waters
as is suggested by the finding of the deep-dwelling squirrel fish (Holocentrus laeteoguttatus) in the stomachs of
this species off Hawaii. The feeding behaviour has been observed by a fishery biologist off Baja California: after
a fish of about 3 m body length had found a school of squids (Dosidicus gigas measuring about 40 cm mantle
length) that was gathering under the night-light of a squid-fishing boat, it approached the school at almost full
speed with its fins completely held back in the grooves, then suddenly hit the squids with its bill, subsequently
nudging the stunned prey and eating it head first. This species has also been observed to swallow big tunas like
skipjack (Katsuwonus pelamis), yellowfin tuna (Thunnus albacares) and bigeye tuna (Thunnus obesus) head first, and
the fishes found in stomachs of M. mazara often showed deep slashes on their bodies, presumably caused by the
bill of the marlin. These observations show that this species seems to use its bill quite often for feeding.
 - 32 -

Stomachs of M. mazara contained mostly squids (Philippine Sea), and tuna-like fishes (off New Zealand and
in the central Pacific) In Hawaiian waters, tuna-like fishes make up more than 85% in volume of the Indo-Pacific
blue marlin’s diet. A large individual caught off Hawaii had a 29 kg bigeye tuna in its stomach and its weight,
including the bigeye was 340 kg. In the eastern North Pacific, M. mazara has been shown (from the results of
Japanese longline research cruises) to feed primarily on squids and fishes, in particular Bramidae, Carangidae,
Gempylidae, Auxis spp., Xiphias gladius, etc.

Dr Radtke estimated the age of this species based on otolith readings (Pacific Gamefish Research News,
1982) as follows: (1) Males: 52.1 kg=6 years; 68.3 to 71.2 kg=7 to 8 years; 82 kg=9 years; 96.9 to 114.2 kg=11 to
12 years; (2) Females: 135.4 to 147.4 kg=8 years; 209.2 to 228.8 kg=13 to 15 years; 286.3 kg=16 years,
336.5 kg=17 years.

Large pelagic sharks like Isurus, Prionace, Lamna and Carcharhinus, as well as the killer whale (Orcinus
orca) and related species have often been observed to attack tunas and billfishes (including this species) hooked on
longlines. The author believes that such attacks are highly improbable under natural conditions.

Size : M. mazara attains sizes over 906 kg (2 000 lb) in commercial longline fisheries and to about 820 kg in
sportsfishing activities. The heaviest record in sportsfishing (unofficial data, since more than one person was on
the fishing rod) is the 818 kg (1 805 lb) “Choys monster” caught by Captain Cornelius Choy and his party off
Waikiki, Howaii; the second-largest is a specimen of 447 cm total length, 523 kg (1 153 lb) weight, and 185 cm of
girth caught at Ritidian Point in Guam (this is the all tackle angling record). The heaviest record listed in “World
Record Game Fishes, 1982” published by the International Game Fish Association is a fish of 498.95 kg (1 100 lb)
weight, 420.4 cm total length, and 200 cm girth width from Le Morne, Mauritius. The size range of M. mazara
caught by commercial longliners averages approximately 200 to 285 cm body length in Pacific equatorial waters
and 215 to 300 cm body length in the Indian Ocean.

In the Pacific, size at first maturity of males is thought to range from 130 to 140 cm eye-fork length (86.8
to 87.8% of body length). Females attain larger sizes than males; around the Bonin-Islands, fish over 200 cm
eye-fork length are all females; around Taiwan Island males attain less than 120 kg weight, while females grow to
over 300 kg.

Interest to Fisheries : Catches of M. mazara have been reported by about 10 countries from seven FAO
Fishing Areas (51, 57, 61, 71, 77, 81 and 87 ). The major fishing nations in the period from 1978 to 1982 were Japan
and the Republic of Korea. The total world catch was 18 193 t in 1978, 18 654 t in 1979, 21 109 t in 1980, 21 413 t
in 1981 and 20 727 t in 1982. Only 8.2% (1 761 t) of the 1981 catch came from the Indian Ocean, while 91.8%
(19 652 t) were taken in the Pacific Ocean, particularly in Fishing Area 61, northwest Pacific (7 543 t), by
Japanese and Chinese (Taiwan Province) vessels (about equal shares); in Fishing Area 71, western central Pacific
(6 048 t) by Japan, Malaysia, the Philippines and the Republic of Korea; and in Fishing Area 77, eastern central
Pacific (4 979 t) by Japan and the Republic of Korea. In 1982, 8.3% of the total catch were taken in the Indian
Ocean, while 91.7% came from the Pacific Ocean, particularly from Fishing Areas 61, 71 and 77 (FAO, 1984).

The gear most commonly used for fishing M. mazara is the Japanese type of longline and its derivates, but
this species is most often caught incidental to fisheries directed at other marlins (M. indica and Tetrapturus
audax) and tunas (Thunnus thynnus, Thunnus obesus and Thunnus albacares). Most Japanese tuna longline vessels
range in size from 240 to 340 GRT. Recently many of them have been equipped with freezing facilities capable
of preserving the fish in excellent condition at very low temperatures, between -40° and -50°C or at even lower
temperatures in the most modern vessels. M. mazara is also taken incidentally by harpooning boats aiming at
Makaira indica, Tetrapturus audax or Xiphias glaudius southern Japan and Taiwan Island.

The quality of the flesh is excellent for sashimi (sliced raw fresh flesh with soy-sauce and horse radish).
Most of it is marketed frozen.

Local Names : AUSTRALIA: Blue marlin; CHILE: Pez zuncho; CHINA: Lan fu yii; JAPAN: Aburakajiki,
Genba, Katokui, Katsuokui, Kudamaki, Kuro, Kuroka, Kurokajiki, Kurokawa, Kurokawakajiki, Kuromazaara,
Mazaara, Njiachi, Njiara, Tsun; MALAYSIA: Mersudji; MEXICO: Maríln azul, Maríln negro; NEW CALEDONIA:
Empéreur, Marlin bleu; NEW ZEALAND: Marlin, Taketonga; REPUBLIC OF KOREA: Nok-sae-chi; TAHITI: Haura;
USA: Blue marlin, Cuban black marlin; USSR: Ch’joernij marlin; VIET NAM: cá cè' den.

Literature : Nakamura, H. (1938, 1942); Royce (1957); Ueyanagi (1964); Howard & Ueyanagi (1965);
Nakamura, Iwai & Matsubara (1968); Strasburg (1969, 1970); Merrett (1971); Nakamura, I. (1974, 1983); Howard &
Starck (1975).

Remarks : The English name “black marlin” was often used among Japanese scientists for Makaira mazara
until the mid-sixties. “Black marlin” Is a direct translation of the Japanese common name, "Kurokajiki" (kuro=
black, kajiki = marlin) for this species. See also “Remarks” under Makaira nigricans.
 -- 33 -

Makaira nigricans Lacepède, 1802 ISTIO Mak 1

Makaira nigricans Lacepède, 1802, Histoire naturelle des Poissons, 4:688-91, pl. 12 (Fig. 3). (Ile de Re, Bay of
Biscay).
Synonymy : Tetrapturus herschelii Gray, 1838; Histiophorus herschelii-Günther, 1860; Tetrapturus amplus
Poey, 1860; Tetrapturus H e r s c h e l i i - L ü t k e n, 1880; Makaira herscheli-Jordan & Evermann, 1896; Makaira ampla-
Jordan & Evermann, 1926; Makaira ensis Jordan & Evermann, 1926; Makaira bermudae Mowbray, 1931; Makaira
nigricans nigricans-Nichols & LaMonte, 1935; Makaira nigricans ampla-Nichols & LaMonte, 1935; Makaira ampla
ampla -LaMonte & Marcy, 1 9 4 1 ; M a k a i r a p e r e z i d e B u e n , 1950; Orthocraeros bermudae-Smith, 1956; Makaira
herschelii-Smith, 1956; Makaira (Makaira) nigricans-Robins & de Sylva.

FAO Names : En - Atlantic blue marlin; Fr - Makaire bleu de l’Atlantique; Sp - Aguja azul de1 Atlántico

Field Marks : Body not very compressed; nape highly

elevated; height of anterior lobe of first dorsal fin less than
greatest body depth; lateral line system chickenwire-shaped.

Diagnostic Features : Body not strongly compressed.

Bill long, very stout and round in cross section; nape conspi-
cuously elevated; right and left branchiostegal membranes
completely united to each other, but free from isthmus; no
gillrakers; both jaws and palatines (roof of mouth) with small, scales
file-like teeth. Two dorsal fins, the first with 39 to 43 rays,
lower than maximum body depth throughout its length, with a
pointed anterior lobe and a long base, originating above the
posterior margin of preopercle and ending near second dorsal
fin origin; second dorsal fin with 6 or 7 rays, its position slightly
backward with respect to that of second anal fin; two anal fins,
the first with 13 to 16, the second with 6 or 7 rays, and very
similar in size and shape to the second dorsal; pectoral fins long
and narrow, adpressible to sides of body, with 19 to 22 rays;
pelvic fins shorter than the pectorals, with a poorly developed
membrane, and depressible into deep ventral grooves. Caudal
peduncle fairly compressed (laterally) and slightly depressed
(dorsoventrally), with strong double keels on each side and a
poorly developed notch on both the dorsal and ventral surfaces;
anus situated just in front of first anal fin origin. Lateral line
forming a complicated network pattern, obvious in immature
fish but obscure in adults, as it becomes progressively imbedded
in the skin (however, the line becomes always clearly visible
when the epidermis is removed). Body densely covered with
elongate, thick, bony scales, each with usually 1 or 2 (mostly 1),
sometimes with 3 posterior points. Vertebrae 24(((((11 precaudal
a n d 13 caudal). Colour: body blue-black dorsally and silvery
white ventrally, with about 15 rows of pale cobalt-coloured
stripes, each consisting of round dots and/or narrow bars. First
dorsal fin membrane blackish or dark blue, without any dots or
marks, other fins usually brown black, sometimes tinged with
dark blue; bases of first and second anal fins tinged with silvery
white.
Lateral line systems of left side in size growth
( schematic )
 -- 34 -

Geographical Distribution : : This species is distributed

mainly in the tropical and temperate waters of the Atlantic. It
is the most tropical of all Atlantic billfishes and probably more
abundant in the western than in the eastern Atlantic, judging
from the hook rates attained by commercial fisheries in both
areas. Its latitudinal range, based on data from the commercial
longline fishery, extends from about 40°40° to 45°N in the North
Atlantic to 40°S in the western South Atlantic, 30°S in the
central South Atlantic, and 35°S in the eastern South Atlantic.

Habitat and Biology : This is an epipelagic and oceanic

species usually found in waters with surface temperatures
r a n g i n g f r o m 22° to 31°C. Water colour affects the occur-
rence of M. nigricans, at least in the northern Gulf of Mexico,
where the fish show preference for blue water.

In the western Atlantic, the monthly distribution of

Japanese longliner catches shows two main seasonal concentra-
tions; one from January through April in the western South
Atlantic, between 5° and 30°S, and the other from June through
October, in the western North Atlantic, between 10° and 35°N.
May, November and December appear to be transitional
months. In the eastern Atlantic, where the blue marlin is less
abundant, it occurs mostly off the east coast of Africa between
25°N and 25°S. There are indications of a periodical fluctua-
tion in abundance of this species which is probably due to ecological interaction with other species of marlin. For
example, during the fishing season in the northern Gulf of Mexico (April to November), anglers tend to catch more
M. nigricans at times when white marlin (Tetrapturus albidus) catches are lowest and vice versa.

In the northwestern Atlantic, 561 individuals of M. nigricans were tagged from 1955 to 1971, but only 4 have
been recaptured, all near their respective release points.. Although these tag returns are inconclusive, they
suggest that this technique might be successful if applied on a larger scale. Unlike Istiophorus albicans, M.
nigricans does not form concentrations in coastal waters. In the open ocean, it rarely gathers in schools and is
usually found as scattered single individuals.

Little is known about the spawning grounds and seasons. In the western central Atlantic, three larvae have
been recorded off Georgia (30°5’N/79°37’W), two off Cat Cay, Bahamas, one at 32°06’N/72°00’W, and one at about
40 miles northeast of Fort Pierce, Florida. Several juveniles were recorded from off Jamaica. In the
southwestern Atlantic, 85 larvae were found from off Brazil, between Cabo de Sao Roque and 26 S. Subripe ova
are opaque, white to yellow, and 0.3 to 0.5 mm in diameter. Transparent spherical eggs flowing out of a ripe
ovary measured 1 mm in diameter.

The Atlantic blue marlin feeds mostly in near-surface waters but sometimes makes trips to relatively deep
water for feeding, as is shown by the presence of deep sea fishes such as Pseudoscopelus in the stomachs of
specimens caught off Puerto Rico. However, the Atlantic blue marlin is believed not to feed on the surface at
night; surface-trolled baits are taken by this species especially in the morning between 1O:OO and 11:OO hours.
Ovchinnikov (1970) and Rivas (1975) both concluded that the marlin’s bill does not play an important role in the
capture of food, based on observations that marlins without bills or with broken or malformed bills are as healthy
as normal fish. The present author disagrees with these conclusions.
Stomachs of M. nigricans contained dolphinfishes (Coryphaena) and tuna-like fishes, especially frigate
mackerel (Auxis) around the Bahamas; tuna-like fishes, predominant in both number and volume, off Puerto Rico
and Jamaica, and mainly dolphinfishes and scombrids in the northern Gulf of Mexico.

The size range of the prey taken by this species is relatively wide, including fishes from about 20 to 102 cm
total length and octopods from at least 15 cm to about 61 cm (Bimini, Bahamas). Off Puerto Rico, a 135 kg
Atlantic blue marlin was found to have swallowed a 38 mm long postlarval surgeonfish while another fish of
similar size had ingested a 11 kg squid. In the Gulf of Guinea, a 290 kg M. nigricans had swallowed a bigeye tuna,
(Thunnus obesus) weighing about 50 kg.

Size : M. nigricans is smaller than M. mazara, averaging between 136 kg (300 lb) and 181 kg (400 lb). The
maximum size of this species exceeds 375 cm body length and 580 kg in weight. However, many anglers report
having seen this species reaching sizes up to 680 kg (1 500 lb). The largest specimen recorded by the International
Game Fish Association (the all tackle angling record) was from St. Thomas, Virgin Islands and weighed 581.51 kg
(1 282 lb). The second-largest was a fish of 461.98 kg (1 018 lb 8 oz) from South Pass, Louisiana, USA. Generally,
M. nigricans heavier than 136 kg (300 lb) are females. Throughout the Atlantic, the size of fish caught by
commercial longliners ranges from about 230 cm tc 345 cm total length (200 to 275 cm body length).

Interest to Fisheries : From 1978 to 1982, catches of M. nigricans have been recorded by several countries
from five FAO Fishing Areas (21, 31, 34, 41 and 47). The total world catch was 1 842 t in 1978, 2 160 t in 1979,
2 328 t in 1980, 2 218 t in 1981 and 2 448 t in 1982. Of the above-mentioned five Fishing Areas, the only one that

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yielded catches over 1 000 t during this period was the eastern central Atlantic (Fishing Area 34) in 1980 (1 054 t
taken predominantly by Cuba). In 1981, 1% (22 t) of the total catch was taken in Fishing Area 21 (northwest
Atlantic) by Japan exclusively, 38.1% (846 t) in Fishing Area 31 (western central Atlantic) by Cuba, Venezuela,
Japan and others, 24% (532 t) in Fishing Area 34 (eastern central Atlantic) predominantly by Cuba, 14.3% (318 t)
in Fishing Area 41 (southwest Atlantic) by several countries, and 22.5% (500 t) in Fishing Area 47 (southeastern
Atlantic) by Japan and other several countries (FAO, 1983). In 1982, 0.12% (3 t) of the total catch was taken in
Fishing Area 21 exclusively by Japan, 34.3% (839 t) in Fishing Area 31 by Cuba, Japan, Venezuela, Republic of
Korea and others, 29.7% (726 t) in Fishing Area 34 by Cuba, Japan, Republic of Korea and others, 9.5% (232 t) in
Fishing Area 41 by Japan and other countries, and 26.5% (648 t) in Fishing Area 47 by Japan and China (Taiwan
Province)(FAO, 1984).

In the commercial fisheries, the fishing gear currently in use is the conventional or modified Japanese
longline. Around Cuba, M. nigricans is caught commercially with “palangres” a type of gear similar to the
ordinary longline, but with only a few hooks used. As regards vessels, Japanese-type longliners are used
extensively in the commercial fisheries, with certain modifications in size and equipment according to purposes
and localities. Small boats powered by sail and/or motors (inboard and outboard) are used for operating the Cuban
palangres.

In the sportsfisheries, M. nigricans is taken by ordinary rod and reel methods. Natural (mostly) or artificial
baits are trolled along the surface at speeds varying from 4 to 8 knots and usually 2 to 5 lines are used
simultaneously.

The quality of the flesh is excellent. It is marketed fresh or frozen.

Local Names : BRAZIL: Agulhao preto; CANADA: Blue marlin, Makaire bleu; CUBA: Abanico, Aguja,
Aguja casta, Castero, Prieta, Voladora; FRANCE: Makaire bleu; JAPAN: Nishikuro, Nishikurokajiki; MEXICO:
Marlín azul, Marlín negro; MOROCCO: Espadon; PORTUGAL: Espadium azul, Espadium azul do Atlantico, Peito,
Peixe agulha; REPUBLIC OF KOREA: Nog-Sae-chi; SOUTH AFRICA: Blue marlin, Blou marlyn; SPAIN: Aguja
azul; USA: Blue marlin, Cuban black marlin; USSR: Chernyi marlin, Goluboi marlin; VENEZUELA: Marlín azul;
WEST INDIES: Blue marlin, Squadron.

Literature : Krumholz & deSyIva (1958); Erdman (1962, 1968); deSylva (1963); Ovchinnikov (1970); Ueyanagi
et al. (1970); Mather, Jones and Beardsley (1972); Rivas (1975).

Remarks : The present author believes that the Indo-Pacific blue marlin Makaira , mazara and the Atlantic
blue marlin, Makaira nigricans, are distinct species chiefly because of differences in the pattern of the lateral line
system (simple-looped in M. mazara and reticulated in M..nigricans). Many scientistis, however, do not recognize
this character as specifically diagnostic and consider M. nigricans as a single pantropical species occurring in the
Atlantic, Pacific, and Indian oceans (Rivas, 1956, 1975; Royce, 1957; Briggs, 1960; Robins & de Sylva, 1960;
Jones & Silas, 1964; Morrow, 1964).

ISTIO Tetra
Tetrapturus Rafinesque, 1810

Genus : Tetrapturus Rafinesque, 1810, Caratteri di alcuni nuovi generi e nuove specie di animali e piante
della Sicilia, con varie osservazioni sopra i medesimi. Palermo, 1810, 105 pp., 20 pl. (ref.p. 54-5, pl.1, fig.1).

Type Species : Tetrapturus belone Rafinesque, 1810.

Synonymy : Skeponopodus Nardo, 1833; Tetraptururus Bonnaterre, 1841 (amended spelling); Tetrapterus
Agassiz, 1841 (amended spelling); Tetraplurus Verany, 1847 (? misprint); Scheponopodus Canestrini, 1872 (amended
spelling); Marlina Grey; 1928; Kajikia Hirasaka and Nakamura, 1947; Pseudohistiophorus de Buen , 1950, Lamontella
Smith, 1956.

Remarks : This genus includes six species of which one (Tetrapturus georgei) still requires proof with regard
to its validity. The possibility of existence of a seventh species, the so-called hatchet marlin (Tetrapturus sp.) is
even more doubtful.

Tetrapturus albidus Poey, 1860

ISTIO Tetra 1

Tetrapturus albidus Poey, Memorias sobrela historia natural de la isla de Cuba , 2:237-244, 258-60, pl. 151
(fig. 1), pl. 16 (figs 2-13), pl. 17 (figs 1,5,6-10,11,26) (Cuba ).
Synonimy: Tetrapturus le ssonae Canestr ini, 1861; Makaira lessonae-Jordan & Evermann, 1926; Makaira
albida-Jordan & Evermann, 1926; Lamontella albida-Smith, 1956.
 -- 36 -

FAO Names : --Atlantic white marlin; Fr - M a k a i r e blanc de l'Atlantique; Sp - Aguja blanca de1
Atlántico.

Field Marks : Anterior lobe of first dorsal fin rounded and higher than
remainder of fin, the height decreasing gradually backward; anus situated near origin
of first anal fin, the distance between them smaller than half of first anal fin height.

Diagnostic Features : Body elongate and fairly compressed. Bill stout and long,
round in cross section; nape fairly elevated; right and left branchiostegal membranes
completely united to each other, but free from isthmus; no gillrakers; both jaws and
palatines (roof of mouth) with small, file-like teeth. Two dorsal fins, the first with 38
to 46 rays, usually with a rounded anterior lobe, higher than body depth anteriorly,
then abruptly decreasing in height to about the 12th dorsal fin ray and gently
decreasing further backward; first dorsal fin base long, extending from above posterior
margin of preopercle to near second dorsal fin origin; second dorsal fin with 5 or 6 scales
rays, its position slightly backward with respect to the second anal fin; two anal fins,
the first with 12 to 17 rays, the second with 5 or 6 rays and very similar in size and shape to the second dorsal;
pectoral fins long and wide, round-tipped, adpressible against sides of body and with 18 to 21 rays; pelvic fins
slender and almost equal to, or slightly shorter than the pectorals. Caudal peduncle well compressed (laterally)
and slightly depressed (dorsoventrally), with strong double keels on each side and a shallow notch on both, the
dorsal and ventral surfaces; anus situated just in front of first anal fin origin. Lateral line single and obvious,
curving above base of pectoral fin and then continuing in a straight line toward the caudal fin base. Body densely
covered with elongate bony scales, each with 1 or 2 posterior points. Vertebrae 24 (12 precaudal and 12 caudal).
Colour: body blue-black dorsally, silvery white splattered with brown laterally, and silvery white ventrally;
usually no blotches or marks on body, but sometimes more than 15 rows of obscure whitish stripes. First dorsal
fin dark blue with many black dots; second drosal fin dark blue; pectoral fins blackish brown, sometimes tinged
with silvery white; pelvic fins blue-black with a black fin membrane; caudal fin blackish brown.

Geographical Distribution : Based on Japanese com-

mercial longliner’s catches, the distribution of T. albidus
extends over most of the Atlantic, roughly from 45°N in the
North Atlantic to 45°S in the western South Atlantic and
35°S in the eastern South Atlantic. T. albidus is also
known from the Mediterranean Sea, and from Bretagne,
France, though these records seem to correspond to a few
straying individuals.

Habitat and Biology : This is a pelagic and oceanic

species, usually swimming above the thermocline. Its distri-
bution varies seasonally, reaching the higher latitudes in
both the northern and southern hemispheres only during the
respective warm seasons. It is usually found in deep (over
100 m) blue water with surface temperatures over 22°C and
salinities of 35 to 37°/00. Average air temperatures of
regions where it occurs are usually moderate to warm (15°
to 25°C). Currents of 0.5 to 2 knots occur over much of its
habitat. Angling success for white marlin in the Gulf of
Mexico is greater in proportion to the blueness of the water,
and poorer in proportion to its greenness. In some areas, T.
albidus is found to concentrate near rips (usually occurring
at interfaces between different masses of water) or weed
lines. Its differential distribution is also influenced by
 -- 37 -

bottom topography. Steep dropoffs, submarine canyons and shoals, when located in areas with suitable water
conditions, are often the scene of important feeding concentrations of this species and exceptionally productive
fishing.

The recovery of tagged individuals shows that this species may accomplish fairly long journeys, but not the
extensive transoceanic migrations achieved by the bluefin tuna (Thunnus thynnus) and the albacore (Thunnus
alalunga). Tagging experiments carried out the Cooperative Game fish Tagging Program of the Woods Hole
Oceanogrphic Institution have produced considerable insights into the movements of T. albidus in the western
North Atlantic. As of January 1973,1some 9 000 of these fish had been marked in that area,zan and 144 tags had
been returned (Mather, Clark and Mason, 1975).

The information presently available indicates that T. albidus spawns once a year. Knowledge of spawning
seasons, areas, and mating behaviour is incomplete, because of difficulties in identifying the eggs and larvae and
the lack of continuous and comprehensive gonad studies. Ueyanagi et al. (1970) concluded that this species
migrates into subtropical waters to spawn, with peak spawning occurring in early summer. The spawning areas are
found in deep and blue oceanic waters, generally at high surface temperatures (20 to 29°C, except in the southern
Atlantic gyrals) and high surface salinities (over 35°/00). Except off Cabo Frio, Brazil, the productivity of these
waters is considered to be low.

T. albidus is not generally considered a schooling fish, and is most often found as single individuals or in
pairs "tailing" with only the dorsal lobe of their caudal fins showing. Small schools (5 to 12 fish), however, are
occasionally seen feeding on schools of bait, or tailing, but loose aggregations of numerous fish scattered over
fairly large areas are most typical. They may school according to size or sex at various seasons of the year.

Hemingway (1935) described “white marlins” breeding off Cuba in May: “they breed in the same way the
groupers do, except that as current-dwelling fishes, they do this in the current instead of on the reef. The female
marlin heads into the current while the male heads in the opposite direction, and while they are side by side, the
female expells the eggs and the male the milt;the male then catches the eggs in the basket-like opening of his
gill covers and lets them pass out through his mouth”. The feasibility of the latter action seems questionable, but
the observation of paired spawning may be true.

Atlantic white marlin are known to kill or stun their food by spearing it or hitting it with their bill. This
may not always be true, however, as whole specimens found in the stomachs appeared to be unscathed. In such
cases the marlin may have simply overtaken the prey. Squids seem to play a most important part as food of T.
albidus in the different areas of its abundance. In the Gulf of Mexico, the most consistently important food items
observed from 1966 to 1971 were squids, dolphinfish (Coryphaena hippurus), and hardtail jack, (Caranx crysos).
Mackerels were next in importance and flying fishes and bonitos also played a big part. Other food items found
were cutlassfishes, swellfishes, herrings, barracudas, moonfishes, triggerfishes, remoras, hammerhead sharks, and
crabs, but to a much lesser and more inconsistent degree. Along the central Atlantic coasts, the favorite food
items appear to be round herring (Etrumeus teres) and squid (Loligo pealei), but carangids are also well
represented in addition to several other species.

Size : This species reaches a maximum size of over 280 cm in total length and over 82 kg in weight. The all
tackle angling record given by the International Game Fish Association is a fish caught off Victoria, Brazil, on 8
December 1979, weighing 82.50 kg (181 lb 14 oz). Other large records (over 70 kg) are: 79 kg (174 lb 3 oz) off
Victoria, Brazil, on 1 November 1975; 77.40 kg (170 lb 10 oz) off Gurapari, Espirito Santo, Brazil on 2 December
1978; and 73.2 kg/274.3 cm total length at Pompano Beach, Florida on 25 April 1953. The size of Atlantic white
marlins caught by commercial longliners ranges from 130 to 210 cm body length (mostly around 165 cm body
length).

Interest to Fisheries : In the period from 1978 to 1981, catches of T. albidus have been reported from five
FAO Fishing Areas (21, 31, 34, 41 and 47) by Japan and the Republic of Korea. The world total catch was very
small throughout these areas: 204 t in 1981, 93 t in 1979, 119 t in 1980, 121 t in 1981 and 131 t in 1982 (FAO,
1984).

The types of fishing gear used for billfishes and tunas, including T. albidus, vary somewhat from one area to
another but all employ the basic hook and line technique.The major gear classifications are rod and reel, handline
and longline. The fishing gear is operated from various types of boats, ranging from large ocean-going longliners
and very luxurious sportsfishing crafts down to outboard wooden or FRP motorboats and small 4.5 to 6 m wooden
sailboats and row boats in the Caribbean area.

The quality of the flesh is excellent. It is mostly marketed frozen in Japan, and fresh locally.

Local Names : BRAZIL: Agulhao, Agulhao branca, Bicuda, Espadarte meca; CANADA: Makaire blanc,
White marlin; CUBA: Aguja blanca, Aguja de paladar, Blanca, Cabezona; FRANCE: Makaire blanc; ITALY:
Marlin bianco; JAPAN: Nishimaka, Nishimakajiki (name for white marlin landed in Japan); MORROCO: Espadon;
PORTUGAL: Agulha, Espadium branco, Espadium pequenho, Espadon branco do Atlantico; REPUBLIC OF
KOREA: Bag-sae-chi; SOUTH AFRICA: White marlin, Wit marlyn; SPAIN: Aguja blanca, Alfiler, Alton, Cometa,
Pez aguja; USA: Skilligalee, White marlin;USSR: Belyi marlin; VENEZUELA: Aguja blanca; WEST INDIES: White
marlin.
 -- 38 -

Literature :: Wallace && Wallace (1942); Gibbs (1957); de Sylva (1963); Stephens (1965); Ueyanagi et al.
(1970); Nakamura (1971); Nakamura & Rivas (1972); Robins (1974); Mather, Clark & Mason (1975).

Tetrapturus angustirostris Tanaka, 1915

ISTIO tetra 3

tTetrapturus angustirostris Tanaka, 1914 to 1915, figures and description of the fishes of Japan, 18:p1.88
(fig.285 1914), 19:324 (1915) (Sagami Bay, Japan).

Synonymy :: Tetrapturus illingworthi Jordan & Evermann, 1926; Tetrapturus kraussi Jordan & Evermann,
1926; Pseudohistiophorus angustirostris-de Buen, 1950; Pseudohistiophorus illingworthi-de Buen, 1950.

FA0 Names : E n - S h o r t b i l l s p e a r f i s h ; F r -M a k a i r e à r o s t r e c o u r t ; S p - M a r l i n t r o m p a c o r t a

Field Marks : Bill very short, usually less than 15%

of body length; pectoral fins narrow and short, less than
15% of body length; distance between anus and anal fin
oriyin nearly equal to anal fin height.

Diagnostic Features : Body elongate and fairly

compressed. Bill short and slender, round in cross
section; lower jaw shorter than upper jaw, but still
projecting; nape nearly straight; right and left branchio-
stegal membranes completely united to each other, but
free from isthmus; no gillrakers; both jaws and palatines
(roof of mouth) with small, file-shaped teeth. Two dorsal ca . 55 cm. ca. 160 cm.
fins, the first with 45 to 50 rays and with a pointed body length body length
anterior lobe, higher than body depth anteriorly, the fin
then abruptly decreasing in height to about the 19th Schematic drawings of scales ( not same size )
dorsal fin ray and gradually increasing thereafter, but
maintaining a uniform height posteriorly; first dorsal fin base long, extending from above posterior margin of
preopercle to near second dorsal fin origin; second dorsal fin with 6 or 7 rays, its position slightly backward with
respect to the second anal fin; two anal fins, the first with 12 to 15 rays, and the second with 6 to 8 rays and very
similar in size and shape to the second dorsal; pectoral fins with 17 to 19 rays; pelvic fins slender, about twice
the length of the pectorals. Caudal peduncle fairly compressed (laterally) and slightly depressed (dorsoventrally)
with double keels on each side and a shallow notch on both dorsal and ventral surfaces; anus situated far anterior
to first anal fin origin, at a distance usually longer than the height of first anal fin. Lateral line single and well
visible. Body densely covered with elongate bony scales, each scale with 3 to 5 posterior points. Vertebrae 24
(12 precaudal and 1 2 caudal). Gonad Y-shaped. Colour: body dark blue dorsally, blue splattered with brown
laterally, and silvery white ventrally, without dots or stripes. First dorsal fin dark blue, without dots or blotches;
remaining fins brown or dark brown; bases of first and second anal fins often tinged with silvery white.
 -- 39 -

Geographical Distribution :
T.angustirostris is distributed
throughout the tropical and tem-
perate waters of the Pacific and
Indian oceans and is thought to
be strongly oceanic, rarely ente-
ring coastal waters. Its latitu-
dinal range based on longline
c a t c h e s , extends roughly from
40°N to 35°S in the Pacific
Ocean and from 20°N to 35°-
45°S in the Indian Ocean.

Habitat and Biology : : T.

angustirostris is an oceanic
pelagic fish which does not gene-
rally occur in coastal or enclosed Area of occasional distribution, or invasion, (no spawning)
waters but is found well off-
shore. Longline fisheries in the
equatorial Indian Ocean take relatively few individuals in the upper water layers (0 to 200 m) over depths
shallower than 914 m (500 fathoms) while the highest catch rates are obtained above the 915 to 1830 m (501 to
1000 fathoms) isobaths. The catch rate decreases over deeper areas. The population density, based on catch per
unit of effort data of the longline fishery appears to be usually low, except in the northwestern Pacific between
15° and 30°N where it is high from about November through February. This species is apparently more abundant
during the southeast monsoon period in the western Indian Ocean, when the maturity of female fish is more
advanced and the surface temperatures are at the lowest (mean 25.5°C). Although some stray individuals of this
fish are found in the Atlantic Ocean, this species is believed to have its spawning grounds and principal popu-
lations only in the Pacific and Indian oceans.

Spawning is believed to occur mainly during the winter months, especially in warm offshore currents with
surface temperatures of about 25°C. Thus fish caught in waters around Taiwan Island were found to release ripe
eggs in November. Females with ripe ovaries were also reported to occur during the winter months in the
western Indian Ocean, and in March in the central Pacific Ocean. From the occurrence of larvae and mature fish,
spawning seems to be also more active in winter than in summer in the tropical and subtropical waters of the
Pacific and Indian oceans between 25°N and 25°S. The frequency distribution of the diameters of eggs shed by
the shortbill spearfish ranges roughly from 1.3 to 1.6 mm, with a mean of 1.442 mm in the equatorial western
Indian Ocean. The eggs released from the ovaries around Taiwan Island and preserved in alcohol are spherical and
about 1 mm in diameter. The ovarian eggs of nearly mature (not fully ripe) females are almost colourless and
semitransparent, with slightly yellowish brown oil globules which later unite into two fairly large globules. At this
stage, the diameter of the ovarian eggs is about 0.8 m m .

Like in other billfishes, stomach contents of T. angustirostris differ from place to place and from season to
season. Data from Japanese longline research cruises show that (i) in the eastern Pacific Ocean this species
feeds mainly on cephalopods and fishes, such as Gempylidae, Scombridae, Exocoetidae, Bramidae, Stromateidae,
Alepisaurus spp., Auxis spp., Katsuwonus pelamis etc.; (ii) in the central South Pacific Ocean, the number of fish
species preyed upon is more limited than in Tetrapturus audax and Makaira mazara, but the crustacean and
cephalopod species found in stomachs are nearly the same as those taken by other billfishes. On the other hand,
deepwater fishes such as Myctophidae, Triacanthidae and Polyipnus are lacking, which suggests that T.
angustirostris swims in shallower waters than T. audax and M. mazara. A comparative analysis of the relative
volume of stomach contents of tunas and billfishes from the central South Pacific shows that T. angustirostris and
the yellowfin tuna (Thunnus albacares) have their stomachs filled with food more often than the albacore,
(Thunnus alalunga), the striped marlin (Tetrapturus audax) or the blue marlin (Makaira mazara) and that T.
angustirostris and T. alalunga tend to eat smaller food items than other billfishes and tunas.

Size : The maximum known size of this species is about 2 m in total length and 52 kg in weight. The
average length of fish caught by the longline fishery is about 135 cm eye-fork length in the central South Pacific
and approximately 150 cm in the eastern Pacific, and the average weight is about 18 kg.

Interest to Fisheries : There are no special fisheries for T. angustirostris, but this species is caught
incidentally by tuna longlines and very rarely by trolling or sportfishing. The catch statistics by Japanese
longliners for the Indo-Pacific sailfish (Istiophorus platypterus) includes a negligible proportion of T.
angustirostris. Usually, nearshore records roughly apply to I. platipterus and offshore records to T.
angustirostris. All in all, the annual total catch of this species is estimated at several hundred metric tons.
The shortbill spearfish is marketed mostly frozen in Japan. The flesh is scanty and not of high value,
compared with that of other billfishes. It is mainly used for fish cakes and sausages.

Local Names : JAPAN: Fuurai, Fuuraikajiki, Sanmakajiki, Sugiyama; SOUTH AMERICA: Pez aguja corta;
USA: Japanese spearfish, Shortbill spearfish, Shortnose spearfish, Shortnosed spearfish, Slender spearfish; USSR:
Kop’jenosjets; VIET NAM: Cá cò'Nhâtban.

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Literature : Nakamura (1937, 1938); Royce (1957); Ueyanagi (1962); Watanabe & Ueyanagi (1963); Howard &
Ueyanagi (1965); Koga (1967); Merrett (1970, 1971); Howard & Starck (1975); Kikawa (1975).

Tetrapturus audax (Philippi, 1887) ISTIO Tetra 5

Histiophorus audax Philippi, 1887, Anal.Univ.Chile, 71:35-8, p1.8 (figs 2 to 3) (Iquique, Chile).

Synonymy : Istiophorus audax-Abbott, 1899; Tetrapturus mitsukurii Jordan & Snyder, 1901; Tetrapturus
ectenes Jordan & Evermann, 1926; Makaira grammatica Jordan & Evermann, 1926; Makaira holei Jordan &
Evermann, 1926; Makaira zelandica Jordan & Evermann, 1926; Makaira audax-Jordan & Evermann, 1926; Marlina
mitsukurii-Grey, 1928; Marlina zelandica-Whitley, 1937; Kajikia mitsukurii-Hirasaka & Nakamura, 1947; Kajikia
formosana Hiraska & N a k a m u r a , 1 9 4 7 ; Tetrapturus tenuirostratus Deraniyagala, 1951; Tetrapturus
acutirostratus Deraniyagara, 1952; Makaira formosana-Matsubara, 1955; Marlina audax-Smith, 1956; Tet rapt urus
audax-Robins & de Sylva, 1961; Makaira audax zelandica-Whitley, 1962.
FAO Names : En - Striped marlin; Fr - Marlin rayé; Sp - Marlin rayado.

Field Marks : Anterior lobe of first dorsal fin pointed and higher than remainder of the fin, the height
decreasing gradually backward; anus situated near origin of first anal fin, the distance between them smaller than
half of anal fin height; tips of pectoral and first anal fins pointed.

Diagnostic Features : Body elon-

gate and fairly compressed. Bill stout
and long,round in cross section; nape
fairly elevated; right and left branchio-
stegal membranes completely united to
each other, but free from isthmus; no
gillrakers; both jaws and palatines (roof
of mouth) with small, file-like teeth.
Two dorsal fins, the first with 37 to 42
rays, usually with a pointed anterior
lobe, higher than (or occasionally equal
to) body depth anteriorly, the fin then ca. 12 cm ca.100 cm. ca.180 cm.
abruptly decreasing in height to about body lengh body length body length
the 10th dorsal fin ray and gently
decreasing further backward; first dor- Schematic drawings of scales (not same size)
sal fin base long, extending from above
posterior margin of preopercle to just in
front of second dorsal fin origin; second dorsal fin with 5 or 6 rays, its position slightly backward in respect to the
second anal fin; two anal fins, the first with 13 to 18 rays, the second with 5 or 6 rays and very similar in size and
shape to the second dorsal; pectoral fins long and narrow, with pointed tips, adpressible against sides of body and
with 18 to 22 rays; pelvic fins slender and almost equal to, or slightly shorter than the pectorals in large
specimens, and slightly longer than pectorals in smaller individuals. Caudal peduncle well compressed (laterally)
and slightly depressed (dorsoventrally), with a pair of keels on each side and a shallow notch on both, the dorsal
and ventral surfaces; anus situated just in front of first anal fin origin. Lateral line single and obvious, curving
above base of pectoral fin and then continuing in a straight line toward the caudal fin base. Body densely covered
with elongate bony scales, each with 1 or 2 posterior points. Vertebrae 24 (12 precaudal and 12 caudal). Colour:
body blue-black dorsally and silvery white ventrally, with about 15 rows of cobalt-coloured stripes, each consisting
of round dots and/or narrow bands. First dorsal fin dark blue; other fins usually dark brown, sometimes tinged
with dark blue; bases of first and second anal fins tinged with silvery white.
 -- 41 -

Geographical Distribution :
T. audax occurs mainly in the
tropical, subtropical and tempe-
rate waters of the Pacific and
Indian oceans. J a p a n e s e long-
line catch data collected over
many years show that the distri-
bution pattern of this species
within the Pacific Ocean is
horseshoe-shaped, with the base
of the horseshoe located along
the Central American coast. T.
audax is occasionally found on
the Atlantic side of the Cape of
Good Hope (Talbot & P e n r i t h ,
1962) and one individual was
caught off Angola, West Africa Area of occasional distribution, or invasion, (no spawning)
in October 1976, having probably
strayed from the Indian Ocean as in a few cases of Tetrapturus angustirostris and Makaira indica. The
latitudinal range limits of T. audax, based on data from the commercial longline fishery, extend from about 45°N
in the North Pacific to 30° S in the eastern South Pacific and to 45°S in the western South Pacific, as far south as
45°S in the southwestern Indian Ocean and 35°S in the southeastern Indian Ocean.

Habitat and Biology : This is an epipelagic and oceanic species, usually swimming above the thermocline.
Parin (1968) included it among the holoepipelagic species which inhabit the isothermic, surface-pelagic layer of
the ocean at all stages of their life cycle, and are usually confined to tropical and subtropical waters where
permanent thermoclines exist, but penetrate higher latitudes in the local warm seasons. However, the striped
marlin has a somewhat atypical distribution as compared to most other billfishes and tunas, and seems to prefer
more temperate waters. In the Pacific, its distribution resembles that of the albacore (Thunnus alalunga) and the
bluefin tuna (Thunnus thynnus), in contrast to that of the other billfishes and tunas. In the Indian Ocean,
however, it is found in warmer waters. The total distributional range of this species, is generally bounded by the
20° and 25°C isotherms, at least in the western Pacific Ocean. This is the most dominant and widely distributed
of all billfishes, especially in the eastern and northcentral Pacific, where it is much more abundant than in the
western Pacific. In the Indian Ocean, the striped marlin is abundant in the western Arabian Sea.

Larvae of striped marlin have been recorded from the western North Pacific (west of 180° long.) between
10° to 30°N, and from the central South Pacific (west of 13O°W) between 10° and 30°S. They are most abundant
in the respective local early summers, with peak occurrences during May through June in the western North
Pacific, and in November and December in the central South Pacific. The seasonal occurrence of mature females
coincides with that of the larvae. While the distribution of larvae for the eastern Pacific (east of 12O°W) is not
known, mature fish are reported to occur there between 5°’ and 20°N, largely in May and June. Larvae have also
been reported to occur in the Banda and Timor seas in January and February,in the eastern Indian Ocean in
O c t o b e r a n d N o v e m b e r b e t w e e n 6°N and 6°S, and in the western Indian Ocean between 10°S and 18°S in
December and January. Mature females are found in March and May in the Bay of Bengal, although larval
occurrence is not yet known there. The lower temperature limit in the distribution of larvae is approximately
24°C, both in the Indian and Pacific oceans. However, larval distributions in the two oceans differ in that, in the
Pacific, the larvae of this species are scarcely found in equatorial waters. It has been noted that larvae of T.
audax are not likely to appear in the Kuroshio Current area, while those of the Indo-Pacific sailfish, Istiophorus
platypterus occur there extensively. Two juveniles of striped marlin (12.2 and 14.5 cm body length) were found
in stomachs of a yellowfin tuna (Thunnus albacares), and of a dolphinfish (Coryphaena hippurus) taken by longlines
on 13 January 1955 at 23°52’S/175°49’W and on 21 December 1964 at 17°5'S/67°29'E. These two occurrences
coincide with larval distributions of the species in the South Pacific and Indian oceans, respectively.

The ovarian eggs of striped marlin from New Zealand average about 0.85 mm in diameter shortly before
spawning. The size of ovulated eggs is presumed to exceed 1 mm in diameter, considering that the mean
diameter of the eggs of the shortbill spearfish (Tetrapturus angustirostris) is 1.442 mm and that the eggs of the
Indo-Pacific sailfish (Istiophorus platypterus), measure 1.304 mm in diameter.

T. audax, like the other marlins, does not form dense schools like the tunas, and the individuals are usually
dispersed at considerably wide distances. Several fish, however, are often seen together, sometimes following
one another, especially during the spawning season. Surfacing is apparently more common with strong wind and
high waves. When wind and current are moving in the same direction, the water surface is rather smooth, but
when the wind runs against the current, high waves develop and this is when striped marlin are most often seen at
the surface around Taiwan Island, usually swimming in the direction of the wind. When surfaced, striped marlins
usually swim very slowly, with the upper caudal fin lobe above the surface and the dorsal fin retracted and not
showing, a characteristic which reportedly distinguishes them from swordfishes which are unable to depress the
dorsal fin and show both the dorsal and caudal fins when surfaced. Striped marlins swim faster and are less easily
approached when surfaced than the swordfish. Like other billfishes and tunas, they tend to school by size. For
example, in the eastern Pacific, fish on the southern spawning grounds (forming a single size mode at 180 to
200 cm eye-fork length = 83.9 to 86.1% of body length) are larger than those on the northern spawning grounds
 -- 42 -

(two size modes, one at 140 cm and one at 180 cm). Smaller striped marlins occur in equatorial waters of the
Pacific, but these small fish are absent between 5° and 16°S; in midlatitudes (15° to 30°S) of the central South
Pacific, a longitudinal stratification is apparent, the larger fish (over 180 cm eye-fork length) occurring in
the western Pacific; harpooned fish tend to be larger than longlined fish in the East China Sea, and the harpooned
fish are also fatter at a given length.

Dolphinfishes (Coryphaena spp.), wahoo (Acanthocybium solandri) and the pelagic large sharks (Prionace,
Isurus, Lamna, Carcharhinus and Alopias) feed on many of the same forage organisms as the striped marlin. Its
closest competitors for food are possibly the other billfishes and larger tunas. The striped marlin, however, tends
to feed more on epipelagic organisms and less on mesopelagic ones than the swordfish and the oceanic tunas. Food
habits do not appear to vary significantly with sex or size, at least in adults. Considerable variation in species
composition of the diet occurs, however, with seasons and geographic localities. T. audax, like other billfishes and
tunas, is thought to be carnivorous and a non-selective feeder. Some of its reported major forage species by
localities are: Fistularia sp., Auxis, sp., squid (East Africa); Scomberoesox saurus, Arripis trutta, Loligo sp.,
Omnastrephes sloani, Caranx lutescens, Scomber japonicus (New Zealand); Alespisauridae, Clupeidae (Tasman
Sea; Gempylus serpens, Cololabis saida, Engraulis mordax, Sardinops caerulea, Trachurus symmetricus (Califor-
nia); Etrumeus teres, Fistularia sp., Argonauta sp., squid (Mazatlan, Mexico); Etrumeus teres, Scomber japonicus,
Fistularia sp., squid (Baja California,+Mexico); Auxis spp., Bramidae, Gempylidae, squid (eastern North Pacific)
Alepisaurus spp., squid (eastern South Pacific); squid (Peru-Chile); Engraulis ringens, Trachurus symmetricus, squid
7rkI-
(Chile).

Predators of adults of this species are probably extremely rare or almost inexistent, the only likely
candidates being some of the large pelagic sharks and the toothed whales, although there are many predators to
the earlier life-stages of the striped marlin.

Size : The maximum size attained by this species exceeds 350 cm in total length and 200 kg in weight. The
all tackle angling record is a fish caught off the Cavalli Islands, New Zealand, on 14 January 1977, weighing
189.37 kg (417 lb 8 oz). Other records of large specimens (over 180 kg) are the following: 183.47 kg (404 lb
8 oz), Bay of Islands, New Zealand, on 12 March 1980; 181.89 kg (401 lb), Cavalli Islands, New Zealand, 24
February 1970; 180.53 kg (398 lb), Mayor Island, New Zealand, 30 December 1974 (IGFA, 1981). All world records
for both men and women recognized by IGFA are from New Zealand, except a record from Botany Bay, Sydney, on
24 October 1976 (161.93 kg=357 lb for men’s 30 lb line class).

Size at first capture (longline fisheries) of T. audax is approximately 80 cm eye-fork length (=83.9 to 86.1%
of body length). Around Taiwan Island, size at first maturity generally estimated between 140 cm and 160 cm
eye-fork length, and the biological minimum size of males at about 137 cm eye-fork length. The maximum size
in commercial fisheries is probably about 290 cm eye-fork length or 258.6 kg (570 lb). The sizes of fish taken by
commercial longliners range mainly from 205 to 225 cm body length in the northern part of the western North
Pacific, between 145 and 185 cm body length in the southern part of the western North Pacific, between 235 and
255 cm body length in the central North Pacific, and about 280 cm body length in the western South Pacific.

Interest to Fisheries: In the period from 1978 to 1982, catches of T. audax have been reported from seven
FAO Fishing Areas (51, 57, 61, 71, 77, 81 and 87),87),mostly by Japan and the Republic of Korea. The total world
catch was 15 426 t in 1978, 15 988 t in 1979, 18 429 t in 1980, 15 664 t in 1981 and 15 460 t in 1982. Only 13.6%
(2 132 t) of the 1981 total catch was taken in the Indian Ocean, and 86.4% (13 532 t) in the Pacific Ocean,
particularly in Fishing Area 61 (northwest Pacific) with 7 229 t predominantly by Japan, in Fishing Area 77
(eastern central Pacific), with 2 414 t predominantly by Japan, and in Fishing Area 87 (southeast Pacific) with
2 217 t by Japan and the Republic of Korea (FAO, 1983); of the 1982 catch, 12.6% (1 949 t) was taken in the
Indian Ocean and 87.6% (13 511 t) in the Pacific Ocean, predominantly by Japan, China (Taiwan Province) and the
Republic of Korea (FAO, 1984).

The commercial catch of T. audax is taken mostly by surface longlining, while harpooning may be
responsible for less than 1% of the total catch in recent years. The longliners aim chiefly at tunas and marlins
which are frequently swimming at depths between 100 and 150 m. Longline gear consists of a mainline, float
lines, branch lines, hooks, bouys including several radio-bouys, and flags. Several hundred of these units (each
unit is referred to as a “basket”) are joined in a series to make up a set of longline. The longline is retrieved with
a line hauler. Japanese longliners in the eastern Pacific use about 2 000 hooks (about 400 baskets) per set. Due to
recent manpower problems, considerable effort has been directed toward developing labour-saving devices in
longlining. The reel-type and the tub-type of longlining are two examples of this development. In the reel-type
the mainline (usually wire) is continuous and reeled onto a large drum, while in the tub-type, the retrieved line is
coiled into a large tub. In harpooning, the electric harpoon has been recently introduced in Japanese fisheries to
speed up the killing of the fish.

The quality of the flesh is the best among billfishes for sashimi and sushi. It is marketed mostly frozen,
sometimes fresh.

Local Names : : AUSTRALIA: Striped marlin; CHILE: Pez aguja;CHINA: Chi zuo fo yii, Hung ju chi yii,
Hung ju ting pan; JAPAN: Achinoiyo, Achinoiyu, Achinuigu, Akinoio, Akinoiyo, Amenashi, Bai, Boke, Chiruguwa,
Dainanbo, Haihage, Hainouo, Haise, Mage, Maka, Makajiki, Masashi, Masasu, Naeragi, Naidonbo, Nairage, Nairagi,
Nairanbo, Neeranbo, Nooragi, Nouragi, Oiragi, Oiraki, Okajiki, Sashi, Sasu, Shitore, Shiutome, Tenguzawara,
 -- 43 -

Tsukinbo, Unjiachi; KENYA: Nduaro; MEXICO: Agujón, Marlin, Marlin rayado, Pez puerco; NEW CALEDONIA:
Empéreur, Marlin rayé, Empéreux; NEW ZEALAND: New Zealand marlin, Striped marlin; PHILIPPINES: Dugso,
Liplipan, Marasugi, Spearfish; REPUBLIC OF KOREA: Cheong-sae-chi; SRI LANKA: Haura; USA: Barred
marlin, Pacific striped marlin, Spearfish, Spikefish, Striped marlin, Striped swordfish; USSR: Polosatii marlin;
VIET NAM: C á cò mitsukurii.

Literature : Nakamura, H. (1938); Ueyanagi (1959, 1964); Jones & Kumaran (1964); Howard & Ueyanagi
(1965); Nakamura, I. (1968); Parin (1968); Kume & Joseph (1969, 1969a); Ueyanagi & Wares (1975); and Silas &
Pillai (1982).

Remarks : Honma & Kamimura (1958) and Kamimura & Honma (1958) advanced the hypothesis that North
and South Pacific populations of this species, being quite separate from each other, may represent different
species, based on differences in pectoral fin length, and ecology. Further study is needed on this problem.

Tetrapturus belone Rafinesque, 1810 / ISTIO Tetra 6

Tetrapturus belone Rafinesque, 1810, Caratteri di alcuni nuovi generi e nuove specie di animali e piante della
Sicilia, con varie osservazioni sopri i messimi. Palermo, 1810, 105 pp., 20pl. (ref.p. 54-5, p1.1, fig. 1).

Synonymy : Skeponopodus typus Nardo, 1833; Tetrapterurus belone-Bonaparte, 1841; Tetrapterus belone-
Agassiz, 1843; Tetraplurus belone-Verany, 1847; Histiophorus belone Günther 1860; Scheponopodus prototypus
Canestrini, 1872; Histiophorus (Tetrapturus) belone-Lütken, 11376; Makaira belone-Tortonese, 1958.

FA0 Names : En - Mediterranean spearfish; Fr - Marlin de la Méditerranée; Sp - Marlin de1 Mediterráneo.

Field Marks : Bill very short, about 18% of body length; pectoral fins narrow and short, less than 15% of
body length; distance between anus and anal fin origin nearly equal to anal fin height.

Diagnostic Features : Body elongate and fairly compressed. Bill rather short
and slender, round in cross section; nape almost straight; right and left branchio-
stegal membranes completely united to each other, but free from isthmus; no
gillrakers; both jaws and palatines (roof of mouth) with small, file-like teeth. Two
dorsal fins, the first with 39 to 46 rays and a rounded anterior lobe higher than body
depth anteriorly, the fin then abruptly decreasing to about the 10th dorsal fin ray and
keeping the same height further backward; first dorsal fin base long, extending from
above posterior margin of preopercle to just in front of second dorsal fin origin;
second dorsal fin with 5 to 7 rays, its position backward with respect to the second
anal fin by half the length of the anal fin base; two anal fins, the first with 11 to 15, scales
rays, the second with 6 or 7 rays and very similar in size and shape to the second dorsal; pectoral fins short (10 to
13% of body length), adpressible against sides of body, their upper margins curved, lower margins nearly straight
and tips pointed, with 16 to 20 rays; pelvic fins long and slender, slightly shorter than twice the pectoral fin
length and depressible into deep ventral grooves. Caudal peduncle well compressed (laterally) and slightly
depressed (dorsoventrally), with strong double keels on each side and a shallow notch on both, the dorsal and
ventral surfaces; anus situated far anterior to first anal fin origin. Lateral line single and obvious, its arch
ending between midpoint and tip of pectoral fin. Body densely covered with elongate bony scales, each with 3 to 5
posterior points. Vertebrae 24 (12 precaudal and 12 caudal). Colour: body dark bluish grey to nearly black
dorsally and silvery white ventrally; usually no blotches or marks on body or fins.
 - 44

Geographical Distribution : T. belone is limited in

its distribution to the Mediterranean Sea, being conside-
rably abundant around Italy. Its eastern distributional
limits have not been clearly defined and there have been
no records of its occurrence in the Black or the Aegean
seas. Although juveniles are known from off Lebanon and
Haifa, Israel, no adults have been reported east of the
Ionean Sea. T. belone is the most common istiophorid in
the central basin of the Mediterranean and its life cycle is
completed inside this sea as far as is known to date.
Reliable literature records of this species are from around
Sicily, the Straits of Messina, Palermo, Taranto, the Gulf
of Naples, Venice, the Ligurian Sea, Malta, Mallorca, the
Adriatic Sea, and Split, Yugoslavia (adults); Haifa and
Lebanon (juveniles); the Straits of Messina (larvae).

Habitat and Biology : Like other billfishes, this

species probably swims in the upper 200 m water layer
(epipelagic), generally above or within the thermocline. It
is frequently reported to travel in pairs, this being known
to occur in other istiophorids, possibly corresponding to a
feeding behaviour. In the Straits of Messina, the area of most heavy fishing, adults occur in August and
September, and occasionally in October and November. They seem to prefer the upper water layers of the Straits,
a fact that may well be associated with upwelling and the consequent concentration of food in these waters.

Winter or spring might not be an unreasonable hypothesis for the spawning season of this species. Its
pelagic eggs have been found in plankton collections from the Straits of Messina; they average 1.48 mm in
diameter and the oil globule is yellow-green; the eggs are incubated for several days and their development has
been described (Sparta, 1953, 1961).

T. belone probably feeds on pelagic fishes, such as sardines (Clupeidae), flyingfishes (Exocoetidae),
carangidis, scombridis, dolphinfishes (Coryphaenidae) and others. Around Sicily, it pursues dolphinfishes, Atlantic
sauries, sardine-like fishes, needlefish (Belone belone) and pilotfish (Naucrates ductor).

T. belone is ecologically similar to Xiphias gladius, Tetrapturus albidus, Thunnus thynnus and some large
sharks, with which it competes for food. Because of its relatively large size, it is probabily not preyed upon
extensively in the adult phase, although the larval and juvenile stages are taken by large pelagic fishes.

Size : The maximum size reached by this species exceeds 240 cm in body length and 70 kg in weight. The
usual size composition in commercial catches ranges from 10 to 30 kg (mostly 14 to 18 kg) and averages about
200 cm in body length.

Interest to Fisheries : T. belone is usually taken at the surface by harpoons, longlines, driftnets and
setnets.. In the Gulf of Castellammare, Sicily, and near the towns of Torretta, Granitola and Marinetta, a few
individuals are taken by tuna traps. It is also occasionally caught by flag lines and drifting handlines.

The fishery of this species is incidental to those for swordfish (Xiphias gladius), bluefin tuna (Thunnus
thynnus) and albacore (Thunnus alalunga). In the Straits of Messina, the gear most often used is the harpoon
(fiocine) although this species is also occasionally caught by nets (revastina) normally used for the Atlantic saury,
locally called “costardella”, on which T. belone feeds. To the south of the Straits of Messina, T. belone is taken
at night with vertical nets (palamideras) normally used to catch albacore.

No quantitative data are available on the annual or seasonal catch.

Local Names : ALGERIA: Auggia imbriale; FRANCE: Marlin, Poisson-pique; ITALY: Acura ‘mperiale
(regional variations of spelling and pronounciation based on Aguglia imperiale), Aguggha imperiali, Agugghia
‘mpiriale, Aguglia imperiale, Aguglia pelerana, Ugghia ‘mpriali, Ugulia imperiali; JAPAN: Chichukaifuurai,
Nishifuuraikajiki (even though this species does not occur around Japan, the Japanese have a keen interest to have
names for all billfishes); MALTA: Pastardella; MONACO: Aguglia imperiale; MOROCCO: Bumkhiat; SPAIN:
Marlin; USA: Mediterranean shortbill spearfish, Mediterranean spearfish; YUGOSLAVIA: Iglan, Iglokljun.

Literature : Sparta (1953, 1961); Robins & de SyIva (1960, 1963); Cavaliere (1962); Rodriguez-Roda &
Howard (1962); de Sylva (1973, 1975).

Remarks : Because of difficulties in identification of juvenile and adult billfish, it is possible that this
species is more widely distributed in the Mediterranean Sea than has been reported in the literature, and that
Mediterranean spearfish may have been identified by sports- and commercial fishermen as white marlin,
Tetrapturus albidus, a species which also occurs in the western and central Mediterranean Sea.

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-- 45 -

Tetrapturus georgei Lowe, 1840 ISTIO tetra 4

Tetrapturus georgii Lowe, 1840, Proc.Zool.Soc.London, 8:36-7 (Madeira).

Synonymy : Tetrapturus georgii-de Sylva, 1973; Tetrapturus georgei-Robins, 1874.

FA0 Names : En - Roundscale spearfish; Fr - Makaire épée; Sp - Marlín peto

Field Marks : First dorsal fin unspotted; tips of first dorsal and anal fins
rounded; distance between anus and anal fin origin nearly equal to half of anal fin
height; scales on mid-body soft and round.

Diagnostic Features : Body fairly robust and compressed. Bill long and
slender, round in cross section; nape moderately humped; right and left branchiostegal
membranes completely united to each other, but free from isthmus; no gillrakers; both
jaws and palatines (roof of mouth) with small, file-like teeth. Two dorsal fins, the
first with 43 to 48 rays, higher than the maximum body depth anteriorly, and lower
posteriorly, with a rounded anterior lobe and a long base extending from above
posterior margin of preopercle to just in front of second dorsal fin origin; second
dorsal fin with 6 or 7 rays, its position slightly backward with respect to the second
scales
anal fin; two anal fins, the first high and broadly rounded, with 14 to 16 rays, the
second with 5 to 7 rays and very similar in size to the second dorsal; pectoral fins long, subequal to pelvic fins,
reaching beyond the curve of the lateral line, adpressible against sides of body and with 19 or 20 rays; pelvic fins
long and slender. Caudal peduncle with a pair of keels on each side; anus moderately far from first anal fin
origin, at a distance equal to about half the height of first anal fin. Lateral line single and simple. Scales on
sides of body rounded anteriorly, only slightly imbricated and soft; scales on dorsal and ventral parts of body
elongate, imbricated and stiff. Vertebrae 24 (12 precaudal and 12 caudal). Colour: most probably there are no
bars on body, but this is still uncertain: first dorsal fin completely unspotted. Flesh distinctly redder than in T.
belone, more similar to T. albidus.

Geographical Distribution : This species is posi-

tively known only from the type locality, Madeira, and
from the specimens studied by Robins (1974) from
Sicily, the Straits of Gibraltar, and the adjacent Atlan-
tic Ocean off southern Portugal. Obviously it can be
expected to range widely in the eastern and perhaps
central North Atlantic, and clarification of the central
and eastern Atlantic records of spearfish from the
Japanese longliner’s data is of vital importance. The
larvae and juveniles and their areas of occurrence are
unknown.

Habitat and Biology : The available data are too

few to permit a discussion of the seasonal or annual
variations in occurrence of this species. It is probably
epipelagic and oceanic.

All three of the known females were in a refrac-

tary state with no developed ova. They were collected
on 27 May, 9 August and 5 October 1961. The only
known male, collected on 2 August, had fairly large
testes, but was not in spawning condition. Nothing
else is known of the bionomics and life history of this
species.
 -- 46 -

Size : Male: 160 cm body length, 21.5 kg body weight; females: 157 cm body length and 20 kg body weight;
more or less 1.5 m body length and 23.5 kg body weight; 154 cm body length and 23.5 kg body weight (Robins,
1974).

Interest to Fisheries : No data are available.

Local Names : “Roundscale spearfish” was proposed by Robins (1974) as the English common name for the
species in recognition of its peculiar lateral scales. No other names are available to the author.

Literature : de Sylva (1973); Robins (1974).

Remarks : T. georgei resembles most closely the white marlin, T. albidus, especially in the somewhat
humped nape and the broadly rounded anterior lobes of the first dorsal and first anal fin. The possibility that the
known specimens of T. georgei represent hybrids between other istiophorids has been discussed and rejected by
Robins (1974). Further study is strongly needed to clarify the validity of this species.

Tetrapturus pfluegeri Robins & de Sylva, 1963 ISTIO tetra 2

Tetrapturus pfluegeri Robins & de Sylva, 1963, Bull.Mar.Sci.Gulf Caribb., 3(1):86-69,

3(1):86-69, figs 1-2 (off San Juan,
Puerto Rico).

Synonymy: : Tetrapturus belone: Fowler, 1936 (and many other authors who had not distinguished this species
from Tetrapturus belone Rafinsque, 1810, until Tetrapturus pfluegeri was described by Robins & de Sylva, 1963);
Tetrapterus belone-La Monte, 1940; Tetrapturus beloni -(sic) sic Briggs, 1958.

FAO Names : En - Longbill spearfish; Fr - Makaire bécune; Sp - Aguja picuda.

Field Marks : Bill long, its length usually equal to or slightly longer than head
length; pectoral fins wide, long and rounded, longer than 18% of body length; anus
situated far anterior to first anal fin origin, the distance between anus and anal fin
origin nearly equal to anal fin height.

Diagnostic Features : Body elongate and remarkably compressed, its depth very
low. Bill slender and rather long, round in cross section; nape nearly straight; right
and left branchiostegal membranes completely united to each other, but free from
isthmus; no gillrakers; both jaws and palatines (roof of mouth) with small, file-like
teeth. Two dorsal fins, the first with 44 to 50 rays, and a rounded anterior lobe higher
than body depth anteriorly, the fin then abruptly decreasing in height to about the 9th
dorsal fin ray and maintaining almost the same height further backward, except at scales
posterior end; first dorsal fin base long, extending from above posterior margin of
preopercle to just in front of second dorsal fin origin; second dorsal fin small, with 6 or 7 rays, its position
backward with respect to second anal fin by one third of second anal fin base; two anal fins, the first with 12 to
17 rays, the second with 6 or 7 rays and very similar in size and shape to the second dorsal; pectoral fins long and
wide, round-tipped, adpressible against sides of body and with 18 to 21 rays; pelvic fins slender and almost equal
to, or slightly longer than the pectorals, and depressible into deep ventral grooves. Caudal peduncle compressed
(laterally) and slightly depressed (dorsoventrally), with strong double keels on each side and a shallow notch on
both, the dorsal and ventral surfaces; anus situated far anterior of first anal fin origin. Lateral line single and
obvious, curving above base of pectoral fin and then continuing in a straight line toward the caudal fin base. Body
densely covered with elongate bony scales, each with 2 to 5 posterior points. Vertebrae 24 (12 precaudal and 12
caudal). Gonad y-shaped. Colour: body blue-black dorsally, silvery white, splattered with brown, laterally, and
silvery white ventrally. First dorsal fin dark blue without dots or blotches;second dorsal fin dark blue; pectoral
fins blackish brown, sometimes tinged with greyish white; pelvic fins blue-black with a black fin membrane; first
anal fin dark blue, its base tinged with silvery white; second anal fin blackish brown.
 - 47

Geographical Distribution :: T. pfluegeri was reported

only rather recently as a new species by Robins & de Sylva
(1963). At that time, this species was known certainly only
from the western North Atlantic where it occurs from Georges
Bank to Puerto Rico and from the Gulf of Mexico to the
Caribbean Sea. Recent surveys of research vessels have clearly
shown that this species is also widely distributed in Atlantic
offshore waters, much more densely so in the western than in
the eastern Atlantic. Its latitudinal range, based on catches
by surface longlines of research vessels and commercial boats,
extends from approximately 40°N to 35°S.

Habitat and Biology : An epipelagic and oceanic species,

chiefly found in offshore waters, usually above the thermocline.

Based on occurrence of larvae and mature fish, spawning

of T. pfluegeri t a k e s p l a c e t h r o u g h o u t w i d e a r e a s o f t h e
tropical and subtropical Atlantic: mature individuals are
caught only in the January-March quadrant (mostly) and in the
April-June quadrant (less frequently), with the exception of the
areas around the Cape Verde Islands and the Caribbean Sea,
where some mature individuals have also been recorded in the
October-December quadrant; maure individuals have not been
recorded from north of 20°N in the western North Atlantic,
north of 30°N in the eastern North Atlantic, south of 10°S in
the western South Atlantic, or south of 30°S in the eastern South Atlantic. Maturation seems to occur at the same
time rather than in the same season both in the northern and southern hemispheres, which could be suggestive of
homogeneity of the population of this species. Like the other billfishes, T. pfluegeri feeds chiefly on pelagic fishes
and squids. For example, off northeastern Brazil, forage organisms for this species are Alepisaurus spp., Vomer
spp., Cephalacanthus spp., anchovies (Engraulidae), dolphinfishes (Coryphaena spp.) and squids. Within any region,
billfishes show negligible food selectivity and to some extent are food competitors to each other.

The term “double header” used by sportsfishermen applies to a situation where two longbill spearfish cross
the stern and take both trolled baits. Such pair-formation is known for other billfishes and the dolphinfish
Coryphaena hippurus. Sexes of the paired fish are unknown and frequently only one of them is boated. The pair-
formation, however, is most likely related to hunting procedure and sometimes to mating behaviour. Whether
spawning is done in pairs or larger groups is unknown. No information on the reproductive behaviour nor the
fecundity is available. Fertilization is external, the eggs have not been identified, but the larvae have been
described.

Size : The maximum size of this species exceeds 200 cm in body length and 45 kg in weight. The weight
range of T. pfluegeri taken by sportsfishing is from 9 to 36.5 kg, with rare specimens over 45 kg. The most
common size caught by surface longlines is about 165 cm body length throughout the Atlantic fishing grounds.

Interest to Fisheries : The longbill spearfish is fished by anglers with the same method they use for other
billfishes. No one specifically fishes for this species only. Gear and tackle preferences vary with fishermen but
all are well described in the sportsfishing literature, such as Mygdalski (1958), Tinsley (1964), Rybovich (1965),
Goadby (1970, 1972), etc. As many as 10 longbill spearfish per year are sent to each of the two principal
taxidermy shops in southeastern Florida. Catches everywhere are low; probably fewer than 100 specimens are
caught per year by sportsfishermen in the western Atlantic.

Commercial longliners take T. pfluegeri along with tunas, swordfish, other billfishes, wahoo (Acanthocybium
solandri), dolphinfishes (Coryphaena spp.), pelagic sharks and miscellaneous other larger pelagic fishes. Catches
of this species are incidental and the gear used is the ordinary surface tuna longline. Japanese statistical data
lump this species with the Atlantic sailfish, Istiophorus albicans as “spearfish and sailfish”; usually, nearshore
records roughly apply to I. albicans and offshore records to T. pfluegeri. In this category, 67 000, 51 000,
118 000, 118 000, 65 000, 59 000, 52 000, 28 000, 39 000, 23 000, 11 000, 8 000, 7 000, 10 000, 7 000, 2 000, 1 000,
2 000 and 3 000 fishes were caught in the years from 1962 to 1980, with an effort ranging from 22 million to 97.5
million hooks per year. The number of fish caught is apparently decreasing recently.

Local Names : JAPAN: Kuchinaga, Kuchinagafuurai (names for this species landed in Japan, see: Howard &
Ueyanagi, 1963); USA: Atlantic longbill spearfish, Longbill spearfish; USSR: Malyi kopénosets.

Literature : Robins & de Sylva (1963); Ovchinikov (1970); Ueyanagi et al. (1970); Robins (1975).

Remarks : Outline drawings of five specimens of this species (Nakamura, Matsubara & Iwai, 1968, p. 64, fig.
18) based on field notes, were thought by de Sylva (1973) to correspond to T. georgei rather than to T. pfluegeri,
or even to an undescribed species. The present author believes that those five specimens were in fact T.
pfluegeri, since very wide individual variations are recognized in external appearance of this species and none of
the above-mentioned specimens had the round scales typical of T. georgei; they also showed a wider distance
between anus and first anal fin origin (usually greater than height of first anal fin) than in T. georgei (about half
the height of first anal fin).

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2.4 FAMILY XIPHIDAE 1 XIPH

Synonymy : : None.

Remarks : : This family includes a single species, Xiphias gladius, which is easily distinguished from the
istiophorids by its flat bill, the absence of jaw teeth and scales in adults, the short-based dorsal fin which is well
separated from the second dorsal in adults, the absence of pelvic fins, and the presence of a single median keel on
each side of the caudal peduncle.

Xiphias Linnaeus, 1758 1 XIPH Xiph 1

Genus : Xiphias Linnaeus, 1758, Systema naturae, ed.10:248.

Type Species : Xiphias gladius Linnaeus, 1975 by monotypy.

Synonymy : Ziphius Hector, 1875 (?misspelling); Ziphias Cheeseman, 1876 (?misspelling); Phaethonichthys
Nichols, 1923 (based on incomplete juvenile specimen from stomach of a red-tailed tropic bird).
Remarks : This genus is monotypic.

Xiphias gladius XIPH Xiph 1

Linnaeus, 1758

Xiphias Gladius Linnaeus, 1758, Syst.Nat., 10:248 (Habitat in Oceano Europae).

Synonymy : Xiphias gladius-Bloch, 1786; Xiphias imperator Bloch & Schneider, 1801; Xiphias rondeletti
Leach, 1818; Phaethonichthys tuberculatus Nichols, 1923; Xiphias estara Phillips, 1932; Terapterus imperator-
Rohl, 1942; Xiphias thermaicus Serbetis, 1951; Xiphias gladius estara- Whitley, 1964.

FAO Names: En - Swordfish; Fr - Espadon; Sp - Pez espada

Field Marks : Bill extremely long, its cross-section flat; no teeth in jaws, in adults a large single median
caudal keel on each side; no pelvic fins; body without scales.
 -- 49 -

Schemattc drawings of scales (not same size)

Diagnostic Features : : Body
elongate and cylindrical. Upper
jaw prolonged into a long bill,
flat-oval in cross-section (but both
jaws prolonged into long bills in
immature individuals); eyes large;
mouth not protrusible; fine, file-
like teeth present in specimens of
about 1 m (body length), disappear- ca. 6 cm. ca. 25 cm. ca. 90 cm. ca. 200 cm.
ta
ing with growth; gill openings body length body length body length body length
wide , gill membranes united only (no scales)
basally and free from isthmus; no gillrakers. Two
widely separate dorsal fins in adults (continuous in
immature specimens), the first much larger than the
second; first dorsal with 34 to 49, second dorsal with 4
to 6 rays; two separate anal fins in adults (continuous in
immature specimens) the first much larger than the
second; first anal with 13 or 14, second anal with 3 or 4
rays; position of second anal fin slightly more forward
than that of second dorsal fin; pectoral fins falcate, a
little rigid and situated low on body sides, with 16 to 18
rays; pelvic fins and pelvic girdle absent; caudal fin
large and lunate. Caudal peduncle with a large keel
present on each side and a deep notch on both the
dorsal and ventral surfaces; anus situated near first
anal fin origin. Lateral line absent in adults, but
recognizable in specimens to about 1 m body length as
wavy line, disappearing with growth. Adults scaleless
but scales with small spines present in specimens to
about 1 m body length. Vertebrae 26 (15 or 16 precaudal
and 10 or 11 caudal). Colour: back and sides of body
blackish-brown, gradually fading to light-brown on ven-
tral side; fin membrane of first dorsal fin dark blackish-
brown; other fins brown or blackish-brown.

In its preadult stage, the swordfish undergoes

drastic morphological changes with growth, which
affect the body shape, the bill and particularly the
dorsal, anal and caudal fins.
Body changes of swordfish with growth ( schematic )
Geographical Distribution :
Cosmopolitan in tropical, tempe-
rate and sometimes cold waters
o f a l l o c e a n s , including the
Mediterranean Sea, the Sea of
Marmara, the Black Sea, and the
Sea of Azov. Based on data
from commercial longliners’
catches, the latitudinal range o f
this species extends from 50°N
to 45°S In the western Pacific,
from 50°N to 35°S in the eastern
P a c i f i c , f r o m 25°N to 45°S in
the Indian Ocean, from 50°N to
40°-45°S in the western Atlan-
tic, and from 60°N to 45°-50°S
in the eastern Atlantic.

Habitat and Biology : This is an epi-and mesopelagic, oceanic species, usually found in surface waters
warmer than 13°C, the optimum temperature range being 18° to 22°C in the northwestern Pacific Ocean. The
swordfish has the greatest temperature tolerance among billfishes, ranging from 5° to 27°C. Based on records of
forage organisms taken by this species, its depth distribution in the northwestern Pacific ranges normally from the
surface to about 550 m depth. It is believed, however, to descend occasionally into waters of 5° to 10°C and to
depths of at least 650 m.

The swordfish is primarily a warm-water s p e c i e s a n d , g e n e r a l l y s p e a k i n g , i t s m i g r a t i o n s c o n s i s t o f

movements toward temperate or cold waters for feeding in summer and back to warm waters in autumn for
spawning and overwintering. There are two hypotheses on the migration of the swordfish in the northwestern
Atlantic: (1) they migrate to the north and east along the edge of the continental shelf during summer and return
to the south and west in autumn, or (2) there are different groups of swordfish migrating from deep waters toward
the continental shelf in summer and moving back to deep waters in autumn.
 -- 50 -

Larvae are more frequently encountered at temperatures above 24°C. In the Atlantic, spawning apparently
occurs throughout the year in the Caribbean, the Gulf of Mexico, and in the waters off Florida, with the peak of
the spawning season from April through September. In the Pacific Ocean, it occurs in spring and summer (March
through July) in the central part, in spring (September to December) in the western South Pacific, and alI the year
round in equatorial Pacific waters. The best known spawning grounds of the swordfish are found in the
Mediterranean Sea, off the southern part of the Italian Peninsula and Sicily, the main spawning concentrations
occurring in the Straits of Messina. Adults are present on these spawning grounds in all months except January and
February, and spawning is most intensive from the end of June to August, when males are often observed pursuing
females. Eggs of this species have been found from June to September, and young swordfish up to 5 kg have been
recorded from October to December. Large numbers of juveniles occur throughout the Mediterranean from
November to March. Female gonads contain 2 to 5 million eggs. In the Atlantic Ocean swordfish spawn in the
upperwater layer at depths between 0 and 75 m, at temperatures around 23°C, and salinity of 33.8 to 37.4°/00.

Adult swordfish are opportunistic feeders, known to forage for their food from the surface to the bottom
over a wide depth range. Over deep water, they feed primarily on pelagic fishes, including tunas (Thunnus),
dolphinfishes (Coryphaena), Alepisaurus, Gempylus, flyingfishes (Exocoetidae), barracudas (Sphyraenidae) and
others, and pelagic squids (Ommastrephes, Loligo, Illex and others), while in relatively shallow waters they take
chiefly neritic pelagic fishes (mackerels, herrings, anchovies, sardines, sauries, needlefishes, etc.). Large adults
often make feeding trips to the bottom where the temperatures may be 5° to 10°C and feed on demerslal fishes
(hakes, Bramidae, trichiurids, gempylids, redfish, lanternfishes (Myctophydae), Gonostomatidae, Sternoptychidae
etc.). Based on stomach contents from X . gladius, it is most likely true that the swordfish uses its sword to kill
some of its prey, particularly squids and cuttlefishes, as is shown by the slashes on the bodies of prey found in
swordfish stomachs.

Size : This species reaches a maximum size of 445 cm total length and about 540 kg weight. The size range
of fish taken by the commercial swordfish longliners is 120 to 190 cm body length in the northwestern Pacific; the
average weight in the Mediterranean Sea ranges from 115 to 160 kg. Usually females are larger than males, and
most swordfish over 140 kg are females. Adults grow over 230 kg (rarely) in the Mediterranean, up to 320 kg in
the western Atlantic, and up to 537 kg in the southeastern Pacific. The all-tackle-angling record for this species
is a 536.15 kg (1182 lb) fish caught off Iquique, Chile in 1953.

There is little information on biological minimum size and age and some of the data are contradictory. X.
gladius first spawns at 5 to 6 years of age and 150 to 170 cm eye-fork length (which is 85 to 88% of body lenght) in
the Pacific and Indian oceans (Yabe et al., 1959). Males reach sexual maturity at a length of around 100 cm and
females at a length of 70 cm in the Atlantic (Ovchinnikov, 1970). However, recent research conducted on
swordfish off the southeast coast of the United States indicates that males mature at a smaller size than females
(at about 21 kg for males and 74 kg for females) (E. Houde, pers.comm., cited from Palko, Beardsley & Richards,
1981). Kume and Joseph (1969) regarded swordfish of less than 130 cm eye-fork length as immature.
Interest to Fisheries : There are important fisheries for X. gladius in the Atlantic, Indian and Pacific
oceans. Catches have been reported from 14 FAO Fishing Areas (21, 27, 31, 34, 41, 47, 51, 57, 61, 67, 70, 77, 81
and 87) by about 30 countries (major fishing nations: Japan, USA, Italy, Spain, Canada, Republic of Korea, China
(Taiwan Province), the Philippines and Mexico) in the period from 1978 to 1982. The world catch was 40 279 t in
1978, 37 992 t in 1971, 36 402 t in 1980, 37 726 t in 1981 and 40 321 t in 1982. Only 1 439 t of the 1981 catches
were taken in the Indian Ocean, while the rest were distributed in halves between the Pacific and Atlantic oceans
(including the Mediterranean Sea), particularly in Fishing Area 61 (northwestern Pacific) with 8 085 t predomi-
nantly by Japan, and secondly by China (Taiwan Province), Fishing Area 37 (Mediterranean) with 6 584 t
predominantly by Italy and Spain, and Fishing Area 77 (eastern central Pacific) with 5 210 t predominantly by
Mexico and Japan. More than 2 000 t were reported in 1981 from 5 other Fishing Areas, i.e., Fishing Area 21
(northwestern Atlantic) with 2 315 t predominantaly by USA, Fishing Area 27 (northeastern Atlantic) with 2 163 t
predominantly by Spain, Fishing Area 31 (western central Atlantic) with 2 548 t predominantly by USA, Fishing
Area 34 (eastern central Atlantic) with 2 117 t by some 10 countries, and Fishing Area 71 (western central
Pacific) with 2 940 t predominantly by the Philippines (FAO, 1983). Of the 1982 world catch (40 321 t), only 3.7%
(1 500 t) were taken in the Indian Ocean, 43.9% (17 705 t) in the Pacific Ocean, and 52.4% (21 116 t) in the
Atlantic Ocean including the Mediterranean Sea (FAO, 1984).

Catch records from the high seas tuna longline fishery indicate that swordfish are taken almost throughout
the range of that fishery. For the most part, however, swordfish catches are incidental to the tuna longline
fishery, except for the Japanese swordfish longlines (nocturnal longlines) called in Japanese “Mekanawa”
(=swordfish longline) or “Yonawa” (=night longline) which operate in the northwestern Pacific, from Japan
eastward almost to 14O°W. Other important commercial fisheries directed at the swordfish are located in the
western North Atlantic from the Grand and Georges Banks to the Gulf of Mexico (harpooning and longlining), in
the eastern Atlantic, especially in the Gulf of Guinea and the Mediterranean Sea (longlining, harpooning and
various kinds of trapping or setnetting), and in the South Atlantic off the coasts of Brazil and Uruguay (longlining).

Major sportingfishing areas for trolling and drifting baited lines are located off the east coast of the USA from
New York to Texas, from off the coast of California to Ecuador, Peru and northern Chile, off the east coast of
Australia and around New Zealand.
 -- 51 -

The quality of the flesh is excellent for steaks, canning or “Teriyaki” (grilled meat with sugar; soy-sauce and
rice wine in the Japanese way). Marketed mostly fresh or frozen.

Local Names : ALGERIA: Pez espada; ARGENTINA: Pez espada ; AUSTRALIA: Broadbill swordfish,
Swordfish; BRAZIL: Espadarte, Peixe espada; BELGIUM: Espadon ; CANADA: Broadbill swordfish, Espadon,
Swordfish; CHILE: Albacora ,, Pez espada ; CHINA: Chien-chi-yu, Ki-hi-khu, Tinmankhu; CUBA: Pez espada;
DENMARK: Svaerdfisk; FRANCE: Espadon G E R M; A N Y (FED.REP.) Schwertfisch;
: GREECE: Xiphias; INDIA:
Kuthira-meen
n (Lacadive eArchipelago?), Mas-hibaru ; IRELAND: Luinniasc ; ITALY: Pesc Pescee spada; JAPAN:
Andaachi, Dakuda, Ginzasu, Goto , Hirakucha, Hyu,, Io, Izasu, Kudamaki, Medara, Meka, Mekajiki, Mesara,
Okizaara, Rakuda, Shiutome, Shutome, Suzu, Teppo, Tsun ; MALTA: Piscispat, Pixxi spad; MEXICO: Pez espada;
NETHERLANDS: Zwaardvisch; NEW ZEALAND : Brodbill; NORWAY: Sverdfisk ;PERU: Pez espada;
PHILIPPINES: Dugso , Doguso, Lumod, Malasagi, Malasugi, Manumbuk, Mayaspus; PORTUGAL: Agulha, ,Agulhao,
Espada, Espadarte, Peixe agulha, Peixe espada; REPUBLIC OF KOREA: Whang-Sae-chi; ROMANIA: Reste
cuspada; SOUTH
; AFRICA: Broadbill, Swaardvis, Swordfish; SPAIN: Ajai para, Chichi
, is pada, Emperador, Espada,
Espardarte, Pez espada ; SRI LANKA: Kadu kpooara ; SWEDEN: Swardfisk; TUNISIA: Bou : sif ; UNITED
KINGDOM: Swordfish ; USA: Broadbill, Broadbill swordfish, Swordfish ; USSR: Mechenos, Mech-ryba,
Meshvenosouiye; VENEZUELA: Pez espada; VIET NAM: Ho cá mui i kiem ; YUGOSLAVIA: Babljan, Iglun, Igo,
Jaglun, Macokljun, Sablijck.
Literature : Nakamura et al., 1951; Yabe et al., 1959; Tibbo, Day and Doucet (1961); Ovchinnikov, 1970;
Palko, Beardsley and Richards (1981); R
Nakamura,
;i c h a r d
1983.
s

Remarks : The local name “Albacore” is used for Xiphias gladius in Chile, while “albacore” is commonly used
for the longfinn tuna, Thunnus alalunga in many English speaking countries. In the French speaking countries, the
yellowfin tuna, Thunnus albacares iss called “Albacore”.

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3. LIST OF SPECIES BY MAJOR FISHING AREAS

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areas of residence
areas of invasion (occasional presence)
MAJOR MARINE FISHING AREAS FOR STATISTICAL PURPOSES

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B I B L I O 4. BIBLIOGRAPHY

Beardsley, G.L., Jr., N.R. Merrett and W.J. Richards, Synopsis of the biology of the sailfish, Istiophorus
1975 platypterus (Shaw and Nodder, 1791). NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part
3:95-120

Ben-Tuvia, A., Red Sea fishes recently found in the Mediterranean. Copeia, 1966(2):254-75
1966

Briggs, J.C., Fishes of worldwide (circumtropical) distribution. Copeia, 1960(3):171-80

1960

Cadenat, J., Notes d’ichthyologie ouest-africaine. 3 3 . N o t e sur les poisson-épée de côte d’Ivoire et p l u s
1961 spécialement sur les ‘Voiliers’. Bull.Inst.Fondam.Afr.Noire (A Sci.Nat.), 23(1):120-30

Cavaliere, A., Notizie su biologia e pesca di Tetrapturus belone Raf. cenni sul’adulto e descrizione di un suo
1962 rarissimo stadio giovanile. Boll.Pesca Piscic.Idrobiol., 15(2):171-6

de Sylva, D.P., Studies on the age and growth of the Atlantic sailfish, Istiophorus americanus (Cuvier), using
1957 length-frequency curves. Bull.Mar.Sci.Gulf Caribb., 7(2):1-20

, Postlarva of the white marlin, Tetrapturus albidus, from the Florida current off the Carolinas.
1963 Bull.Mar.Sci.Gulf Caribb., 13(1):123-32

, Istiophoridae. In Clofnam I. Check-list of the fishes of the North-Atlantic and of the

1973 Mediterranean, edited by J.C. Hureau and Th. Monod. Paris, Unesco, vol.l:477-81

, A review of the world sport fishery for billfishes (Istiophoridae and Xiphiidae). NOAA Tech.
1974 Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 2:12-23

, Synopsis of biological data on the Mediterranean spearfish, Tetrapturus belone Rafinesque.

1975 NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:121-31
Erdman, D.S., The sport fishery for the blue marlin off Puerto Rico. Trans.Am.Fish.Soc., 91:225-7
1962

, Spawning cycle, sex ratio and weights of blue marlin off Puerto Rico and the Virgin Islands.
1968 Trans.Am.Fish.Soc., 97:131-7

FAO, Yearbook of fishery statistics. Annuaire statistique des pêches. Anuario estadistico de pesca, 1981.
1983 Catches and landings. Captures et quantités débarquées. Capturas y desembarques. Yearb.Fish.
Stat./Annu.Stat.Pêches/Anu.Estad.Pesca, (52):357 p.

, Yearbook of fishery statistics. Annuaire statistique des pêches. Anuario estadistico de pesca,
1984 1982. Catches and landings. Captures et quantités debarquées. debarquees. Capturas y desembarques.
Yearb.Fish.Stat./Annu.Stat.Pêches/Anu.Estad.Pesca, (54).393 p.
Gehringer, G.W., Observations on the development of the Atlantic sailfish Istiophorus americanus (Cuvier),
1956 with notes on an unidentified species of istiophorid. Fish.Bull.U.S.Fish.Wild.Serv, 57 (110):139-71
~-
Gibbs, R.H., Jr., Preliminary analysis of the distribution of white marlin, Makaira albida (Poey ) in the Gulf of
1957 Mexico. Bull.Mar.Sci.Gulf.Caribb., 7(4):360-9

Goadby P., Big fish and blue water: gamefishing in the Pacific. Sydney, Angus and Robertson, 334 p. 2nd
1972 rev.ed.

Hemingway, E., Marlin off Cuba. In American big game fishing, edited by E.V. Connet.
Connet. New York, Derrydale
1935 Press, pp. 55-82

Honma, M. and T. Kamimura, A population study of the so-called makajiki (striped marlin) of both northern and
1958 southern hemispheres of the Pacific. 2. Fishing conditions in the southern hemisphere.
8:12-21 (in Japanese). English translation by G.Y. Beard, National
Rep.Nankai Reg.Fish.Res.Lab., i3:12-21
Marine Fisheries Service Southwest Fisheries Center, Honolulu Laboratory, 11 p.

Howard, J.K. and W.A. Starck II, Distribution and relative abundance of billfishes (Istiophoridae) of the Indian
1975 Ocean.Stud.Trop.Oceanogr., (13):1-31
 -- 55 -

Howard, J.K. and S. Ueyanagi, Japanese standard common names of Istiophoridae. Occas.Pap.IFFC, 63(12):1-3
1963

, Distribution and relative abundance of billfishes (Istiophoridae) of the Pacific Ocean. Stud.
1965 Trop.Oceanogr., (2):1-134

IGFA (International Game Fish Association), World record game fishes, 1981. Florida, International Game Fish
1981 Association, 308 p.

Iwai, T. and I. Nakamura, Branchial skeleton of the bluefin tuna, with special reference to the gill rays.
1964 BulI.Misaki Mar.Biol.Inst., (6):21-5

Jolley, J.W., Jr., On the biology of Florida east coast Atlantic sailfish (Istiophorus platypterus). NOAA Tech.
1974 Rep.NMFS (Spec.Sci.Rep.-Fish.Ser. , (675)Part 2:81-8

, The biology and fishery of Atlantic sailfish Istiophorus platypterus, from southeastern Florida.
1977 Fla.Mar.Res.PubI., (28):1-31

Jones, S. and M. Kumaran, Distribution of larval billfishes (Xiphiidae and Istiophoridae) in the Indo-Pacific with
1964 special reference to the collection made by the Danish Dana Expedition. Symp.Ser.Mar.BioI.
Assoc.India, (1)Pt 1:483-98

Jones, S. and E.G. Silas, A systematic review of the scombroid fishes of India. Symp.Ser.Mar.Biol.Assoc.India,
1964 (1)Pt l:l-105

Kamimura, T. and M. Honma, A population study of the so-called Makajiki (striped marlin) of both northern and
1958 southern hemispheres of the Pacific. 1. Comparison of external characters. Rep.Nankai Reg.
Fish.Res.Lab., (8):1-11 (in Japanese). English transl. by National Marine Fisheries Service,
Southwest Fisheries Center, Honolulu Laboratory

Kikawa, S., Synopsis of biological data on the shortbill spearfish, Tetrapturus angustirostris Tanaka, 1914 in the
1975 Indo-Pacific areas. NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:39-54

Koga, S., Studies on the fishery biology of the tuna and marlin in the Indian Ocean and the South Pacific
1967 Ocean. Contrib.Shimonoseki Univ.Fish. , (490):1-208 (in Japanese)

Koto., T., I. Furukawa and K. Kodama, Studies on the tuna longline fishery in the East China Sea. 3. Ecological
1959 studies on the sailfish. Rep.Nankai Reg.Fish.Res.Lab., 10:88-106 (in Japanese, English summary)

Koto, T. and K. Kodama, Some considerations on the growth of marlins, using size-frequencies in commercial
1962 catches. 1. Attempts to estimate the growth of sailfish. Rep.Nankai Reg.Fish.Res.Lab., 15:97-107
(in Japanese, English summary)

Krumholtz, L.A. and D.P. de Sylva, Some foods of marlins near Bimini, Bahamas. BulI.Am.Mus.Nat.Hist.,
1958 114(5):406-11

Kume, S. and J. Joseph, The Japanese longline fishery for tunas and billfishes in the eastern Pacific Ocean east
1969 of 130°W, 1964-1966. Bull.I-ATTC , 13:277-418 (in English and Spanish)

, Size composition and sexual maturity of billfishes caught by the Japanese Iongline fishery in the
1969a Pacific Ocean east of 13O°W. Bull.Far Seas Fish.Res.Lab., 2:115-62

Mather, C.O., Billfish, marlin, broadbill, sailfish. Sidney, B.C., Canada, Saltaire Publishing Ltd., 272 p.
1976

Mather, F.J., III, H.L. Clark and J.M. Mason, Jr., Synopsis of the biology of the white marlin, Tetrapturus
1975 albidus Poey (1861). NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:55-94

Mather, F.J., III, A.C. Jones and G.L. Beardsley, Jr., Migration and distribution of white marlin and blue marlin
1972 in the Atlantic Ocean. Fish.Bull.NOAA/NMFS, 70:283-98

Mather, F.J., III, J.M. Mason, Jr. and H.L. Clark, Migration of white marlin and blue marlin in the western
1975 North Atlantic Ocean - tagging results since May 1970. NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-
Fish.Ser.), (675)Part 2:211-22

Mather, F.J. et al., Results of sailfish tagging in the western North Atlantic Ocean. NOAA Tech.Rep.NMFS
1974 spec.Sci.Rep.-Fish.Ser.), (675)Part 2:194-210
 - 56 -

McClane, A.J. (ed.) McClane’s field guide to salt water fishes of North America. New York, Holt, Rinehart
1974 and Winston, 283 p.

Merrett, N.R., Gonad development in billfish (Istiophoridae) from the Indian Ocean. J.Zool., Lond., 160:355-70
1970

, Aspects of the biology of billfish (Istiophoridae) from the equatorial western Indian Ocean.
1971 J.Zool., Lond., 163:351-95

Migdalski, E.C., Anglers guide to the salt water game fishes: Atlantic and Pacific. New York, Ronald Press,
1958 506 p.

Morrow, J.E., Marlins, sailfish and spearfish of the Indian Ocean. Symp.Ser.Mar.Biol.Assoc.India, (1)Pt 1:429-
1964 40

Nakamura, E.L., An analysis of the catches and biology of big game fishes caught by the New Orleans Big
1971 Game Fishing Club, 1966-1970. Rep.East Gulf Sport Fish.Mar.Lab., 38 p.

Nakamura, C.L. and L.R. Rivas, Big game fishing in the northeastern Gulf of Mexico during 1971. Panama
1972 City, Florida, National Marine Fisheries Service, 20 p.

Nakamura, H., On the habits of some istiophorid fishes of Taiwan waters, especially the short-nosed spearfish.
1937 Zool.Mag./Dobutsugaku Zassi, 49(6):233-8 (in Japanese)

, Report of an investigation of the spearfishes of Formosan waters. Rep.Taiwan Gov.Gen.Fish.

1738 E x p . S t n . , 1 0 : 1 - 3 4 ( i n J a p a n e s e ) . English transl. in Spec.Sci.Rep.U.S.FishWild.Serv.(Fish),
(153):1-46 (1955)
, On the spawning habits of the sailfish. Zool.Mag./Dobutsugaku Zassi, 5 2 ( 8 ) : 2 9 6 - 7 (in Japanese)
1940 ’

, On the ecology of the istiophorid fishes of Taiwan waters. Bull.Sci.Fish.Assoc.Fac.Agric.Tokyo

1942 Imp.Uni./Suisan Gakkai Ho, 9(1):45-51
9(1):45-51 (in Japanese)
, Tunas and their fisheries. Tokyo, Takeuchishobo, 118 p. (in Japanese). English transl. in
1949 Spec.Sci.Rep.U.S.FishWild.Serv.(Fish), (82):1-115 (1952)

, Tuna longline fishery and fishing grounds. Tokyo, Association of Japanese Tuna Fisheries
1951 Cooperatives (in Japanese). English transl. in Spec.Sci.Rep.U.S.FishWild.Serv.(Fish), (112):1-168
(1954)

Nakamura, H. et al. Notes on the life-history of the swordfish, Xiphias gladius Linnaeus. Jap.J.Ichthyol.,
1951 1(4):264-71

Nakamura, I., Juveniles of the striped marlin, Tetrapturus audax (Philippi). Mem.Coll.Agric.Kyoto Univ.,
1968 (94):17-29

, Some aspects of the systematics and distribution of billfishes. NOAA Yech.Rep.NMFS(Spec.

1974 (Sci.Rep.-Fish.Ser.), (675)Part 2:45-53

, Synopsis of biological data on the black marlin, Makaira indica (Cuvier) 1831. NOAA Tech.
1975 3:17-27-
Rep.NMFS(Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:17-27

, Systematics of the billfishes (Xiphiidae and Istiophoridae). Publ.Seto Mar.Biol.Lab., 28(5/6):

1983 ;55-396
255-396

Nakamura, I., T. Iwai and K. Matsubara, A review of the sailfish, spearfish, marlin and swordfish of the world.
1968 Spec.Rep.Misaki Mar.Biol.Inst., (4):1-95
(4):1-95 (in Japanese)

Ovchinnikov, V.V., Swordfishes and billfishes in the Atlantic Ocean. Ecology and functional morphology.
1970 Kaliningrad, Atlantic Scientific Research Institute for Fisheries Oceanography. Translated into
English by Israel Program for Scientific Translations, Jerusalem, 77 p. (1971)

Pacific Gamefish Research News, Published by Pacific Gamefish Foundation (cited indirectly from Iwami, T.,
1982 1984. All about billfish. Game Fish Blue Water, (2):76-80, in Japanese)

Palko, B.J., G.L. Beardsley and W.J. Richards, Synopsis of the biology of the swordfish, Xiphias gladius
1981 Linnaeus. NOAA Tech.Rep.NMFS Circ., (441):21p.
 - 57 -

Parin,
n N.V., Ikhtiofauna okeanoskoi epipelagiali (Ichthyofauna of the Epipelagic zone) Moscow, Nauka .
1968 Publishing House. Translated into English by Israel Program for Scientific Translations, Jerusalem,
IPST Catalog No.5528:210 No.5528:21
p.0 (1970)

Pristas, P., A possible hatchet marlin (Tetrapturus sp.) from the Gulf of Mexico. Northeast Gulf Sci., 4(1):51-6 .
1980

Rivas, L.R., Definitions and methods of measuring and counting in the billfishes (Istiophoridae, Xiphiidae).
1956 BulI.Mar.Sci.Gulf Caribb., 6(1):18-27 f

, Synopsis of biological data on blue marlin, Makaira nigricans Lacepède, 1802. NOAA Tech.
1975 Rep.NMFS(Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:1-16

Robins, CR., The validity and status of the roundscale spearfish, Tetrapturus georgei. NOAA Tech.Rep.
1974 NMFS (Spec.Sci Rep.-Fish.Ser.), (675)Part 2:54-61

l
, Synopsis of biological data on the longbill spearfish, Tetrapturus pfluegeri Robins and de Sylva.
1975 NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:28-38

Robins, C.R. and D.P. de Sylva, Description and relationships of the longbill spearfish,
l Tetrapturus belone
1960 based on western North Atlantic specimens. BulI.Mar.Sci.GuIf Caribb.,
f 10(4):383-413

, A new western Atlantic spearfish, Tetrapturus pfluegeri with a redescription of the Mediter-
1963 ranean spearfish Tetrapturus belone. BuII.Mar.Sci.Gulf Caribb., 13(1):84-122

Rodriguez-Roda, J. and J.K. Howard, Presence of Istiophoridae along the south Atlantic and Mediterranean.
1962 Nature, Lond., 196:495-6

Royce, W.C., O b s e r v a t i o n s o n t h e spearfishes of the central Pacific. Fish.Bull.U.S.Fish.Wild.Serv., h

1957 57(124):497-554

Rybovich, J.., Sportfisherman (boat). in McClane’s sstandard fishing encyclopedia edited

, by A.J. McClane.
1965 New York, Holt , Rinehart and Winston, pp. 851-62

Silas ,E.G. and P.P. Pillai, ,Resources of tunas and related species and their fisheries in the Indian Ocean.
1982 BulI.Cent.Mar.Fish.Res.Inst., Cochin, (32):174 4p.

Sparta, A., Uova e larve di Tetrapturus belone eRaf. (agulia aimperiale). BolI.Pesca Piscic.Idrobiol., 8(1):58-62
1953

, Biologia e pesca di Tetrapturus belonee Raf. e sueform

formeepostlarvali
postlarvali. Boll.Pesca
. Piscic.Idrobiol.,
1961 ;5(
15(1):20-4

Stephens, W.M., Summer cruise to the Sargasso Sea. Sea Front., 11:108-23
1965

Strasburg, D.W., Billfishes of the central Pacific Ocean. Circ.U.S.FishWild.Serv., (311):1-11

1969
, A report on the billfishes of the central Pacific Ocean. Bull.Mar.Sci., 20(3):575-604
1970 ’
Talbot, F.H. and M.J. Penrith , Tunnies and marlins of South Africa. Nature, Lond., 193:558-9
1962

Tibbo, S.N., L.R.Day and W.F. Doucet, The swordfish (Xiphias gladius L.), its life history and economic
1961 .
importance in the northwest Atlantic Bull.Fish.Res.Board Can., (130):1-47

Tinsley ,J.B. , The sailfish, swashbukler of the

Tinsley, r open sea. Gainesville, Florida, University of Florida Press,
1964 216 p.

Ueyanagi, S. , Larvae of the striped marlin, Makaira mitsukurii (Jordan et Snyder). Rep.Nankai Reg.Fish.Res.
1959 Lab., (11):130-46 (in
6 Japanese)

, On the larvae and the spawning areas of the shirokajiki , Marlina marlina
a (Jordan and Hill).
1960 Rep.Nankai Reg.Fish.Res.Lab., (12):

, On the larvae of the shortnosed spearfish, Tetrapturus angustirostris Tanaka. Rep.Nankai Reg.
1962 Fish.Res.Lab. (16):173-89
, 9
(in Japanese)
 -- 58 -

Ueyanagi, S., Methods for identification and discrimination of the larvae of five istiophorid species distributing
1963 in the Indo-Pacific. Rep.Nankai Reg.Fish.Res.Lab., (17):137-50 (in Japanese)

, A study of the relationships of the Indo-Pacific istiophorids. Rep.Nankai Reg.Fish.Res.Lab.,

1963a (17):151-65 (in Japanese)

, Description and distribution of larvae of five istiophorid species in the Indo-Pacific. Symp.Ser.
1964 Mar.Biol.Assoc.India, (1)Pt 1:499-528

Ueyanagi, S. and P.G. Wares, Synopsis of biological data on striped marlin, Tetrapturus audax (Philippi), 1887.
1975 NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 3:132-59

Ueyanagi, S. et al., Distribution,spawning, and relative abundance of billfishes in the Atlantic Ocean.
1970 BullFar Seas Fish.Res.Lab., (3):15-55 (in Japanese)

Voss, G.L., A contribution to the life history and biology of the sailfish, Istiophorus americanus Cuv. and Val.,
1953 in Florida waters. Bull.Mar.Sci.Gulf Caribb., 3(3):206-40

Solving life secrets of the sailfish. Natl.Geogr.Mag., 109:859-72

1956 ’
Wallace, D.H. and E.M. Wallace, Observations on the feeding habits of the white marlin, Tetrapturus albidus
1942 Poey. Rep.Md.Dep.Resour.Educ., (50):1-10

Watanabe, H. and S. Ueyanagi, Young of the shortbill spearfish, Tetrapturus angustirostris Tanaka. Rep.
1963 Nankai Reg.Fish.Res.Lab., (17):1-8 (in Japanese)

Williams, F., The scombroid fishes of East Africa. Symp.Ser.Mar.Biol.Assoc.India, (1)Pt 1:107-64
1964

, The sport fishery for billfish at Malindi, K e n y a , 1 9 5 8 - 1 9 6 8 w i t h s o m e b i o l o g i c a l n o t e s .

1970 Bull.Mar.Sci., 20(4):830-52

Wise, J.P. and C.W. Davis, Seasonal distribution of tunas and billfishes in the Atlantic. NOAA Tech.Rep.NMFS
1973 (Spec.Sci.Rep.-Fish.Ser.), (622):1-24

Yabe, H. et al., Study on the life history of the swordfish, Xiphias gladius Linnaeus. Rep.Nankai Reg.Fish.
1959 Res.Lab., (10):107-50 (in Japanese)

Yuen, H.S.H., A.E. Dizon and J.H. Uchiyama, Notes on the tracking of the Pacific blue marlin, Makaira
1974 nigricans. NOAA Tech.Rep.NMFS (Spec.Sci.Rep.-Fish.Ser.), (675)Part 2:265-8

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5. INDEX OF SCIENTIFIC AND VERNACULAR NAMES

EXPLANATION OF THE SYSTEM

The index applies exclusively to family and species accounts

under Section 2 (Systematic Catalogue)

Type faces used:

Italics : Valid scientific names (genera and species)

: Synonyms (preceded by an asterisk)

ROMAN (caps) : Family Names

Roman : International (FAO) species names

: Local names (preceded by an asterisk)
 - 60 -

PAGE PAGE

*australis, Tetrapturus
A
*Abanico 23,35 27
*Aburakajiki 32
*Achinoiyo
*Achinoiyu
*Achinuigu
42
42
B
*Acura 'mperiale 44
Tetrapturus *Babljan
*acutirostratus, 40 *Baek-sae-chi
*Adzietekwesi 23 *Bag-sae-chi
*Aguggha 'mperiali 44 *Bai
*Agugghia 'mperiale 44 *Balahoo
*Aguglia imperiale 44 *Banba 51
*Aguglia pelerana 44 *Baren 29
*Aguja 35 *Barin 37
Atlantico *Barred marlin 42
Aguja azul del 33 *Bashoo 23
Pacífico *Bashookajiki 26
Aguja azul del Indo- 30 *Bayonet fish 26
*Aguja blanca 37 *belone, Histiophorus 26
Atlántico (Tetrapturus) 43
Aguja blanca del 36 * belone, Histiophorus 26
*Aguja casta 35 *belone, Makaira 26
*Aguja de abanico 23 belone, Tetrapturus 26
*Aguja de paladar 37 * Belyi marlin 43
Aguja negra 27 *bermudae, Hakaira
Aguja picuda 46 *bermudae, Orthocraeros 43
*Aguja vela 23 *Bicuda 43
*Aguja voladora 23 *Bicudo 43
Agujas 20 *Billfish 37
*Agujón 43 Black marlin 33
*Agulha 37,51 *Blanca 33
*Agulhao 23,37,51 *Blou marlyn 23,37
*Agulhao bandeira 23 *Blue marlin 23
*Agulhao branca 37 *Boke 23
*Agulhao de vela 23 *Bou sif 27,29
*Agulhao preto 35 *brevirostris, Histiophorus 37
*Agulhao vela 23 *brevirostris, Istiompax 35
*Ahin koppara 29 *brevirostris, Makaira 32,35
*Aja para 51 *b revirostris, Tetrapturus 42
*Akinoio 42 *Broadbill 51
*Akinoiyo 42 *Broadbill swordfish 27
*Akitaroo 26 *brookei, Istiophorus 27
*Albacora 51 *Bumkhiat 27
*albicans, Histiophorus 21 *Byoobu 27
albicans, Istiophorus 21 *Byoobusashi 51
*albicans, Makaira 21 51
*albida, Lamontella 35 23
*albida, Makaira 35 44
albidus, Tetrapturus 35 26
*Alfiler 37 26
*Altón 37
*amarui, Istiophorus 23
*Amenashi
*americanus, Histiophorus
42
21
C
*americanus, Istiophorus 21
*ampla, Makaira 33 *Cá cè' den
*ampla, Makairampla 33 *Cá co
*ampla, Makaira nigricans 33 *Cá co mitsukurii
*amplus, Tetrapturus 33 *Cá cò ' Nhâtban
*Andaachi 51 *Cabezona
Pseudohistiophorus *Castero 32
*angustirostris, 35 *Ch'joernij marlin 26
Tetrapturus *Gheong-sae-chi 43
angustirostris. 38 *Chernyi marlin 39
Atlantic blue marlin 33 *Chi zuo fo yii 37
spearfish *Chichi spada 35
*Atlantic longbill 47 *Chichukai fuurai 32
Atlantic sailfish 21 *Chien-chi-yu 43
Atlantic white marlin 36 *Chiruguwa 35
*Atsutaro 26 *Cometa 42
*audax, Histiophorus 40 *Cuban black marlin 51
*audax, Hakaira 40 44
*audax, Marlina 40 51
audax, Tetrapturus 40 42
* Auggia imbriale 44 37
*australis, Istiompax 27 32,35
*australis, Makaira 27
 - 61 -

PAGE PAGE

*gladius, Scomber 23
gladius, Xiphias 48
D *Goluboi marlin
*Goto
35
51
*grammatica, Makaira 40
*Dainanbo 42 *granulifer, Histiophorus 21
*Dakuda 51 *greyii, Istiophorus 23
*Djangilus 26 *greyi, Istiophorus 23
*Dogso 26 *greyi, Istiophogladius 23
*Doguso 51 *Guebuε ú 23
*dombraini, Istionpax 27 *guebucu, Skeponopodus 21
*Dot-sae-chi 26
*dubius, Istiophorus 23
*Dugso 26,43,51
*Dumosok 26

H
*Haihage 42
E *Hainouo
*Haio
42
26
*Haioshibi 26
*ectenes, Tetrapturus 40 *Haise 42
*Emperador 51 *Haou 26
*Empéreur 32,43 *Hauo 26
*Empéreur éventail 26 *Haura 32,43
*Empéreux 43 *Haurepe 26
*ensis, Makaira 33 *herschelii, Histiophorus 33
*eriquius, Istiophorus 23 *herschelii, Tetrapturus 33
*Espada 51 *herscheli, Makaira 33
*Espadarte 51 *herscheri, Makaira 27
*Espadarte meca 37 *Hirakucha 51
*Espadium azul 35 belone
Atlantico *Histiophorus (Tetrapturus) 43
*Espadium azul do 35 *Histiophorus albicans 21
*Espadium branco 37 *Histiophorus americanus 21
*Espadium pequenho 37 *Histiophorus audax 40
*Espadon 23,35,37,48 *Histiophorus belone 43
Atlantico *Histiophorus brevirostris 27
*Espadon branco do 37 *Histiophorus gladius 23
*Espardarte veleiro 23 *Histiophorus granulifer 21
*estara, Xiphias 48 *Histiophorus herschelii 33
*estara, Xiphias gladius 48 *Histiophorus immaculatus 23
*Eumakaira nigra 30 *Histiophorus indicus 23
*Histiophorus orientalis 23
*Histiophorus pulchellus 21
*Ho cá mui kiem 51
*Ho-soan-ki-hi 26
*Hohoo malays 26
*holei, Makaira 40
*howardi, Istiompax 30
F *Hung ju chi yii
*Hung ju ting pan
42
42
*Hyu 51
*Fetiso 23
*formosana, Kajikia 40
*formosana, Makaira 40
*Fung-hibaru 26
*Fuurai 39
*Fuuraikajiki 39
I
*Iglan 44
*Iglokljun 44
*Iglun 51
G *Igo
*Ikan jegan
51
26
*Ikan laya 26
*Genba 29,32 *Ikan layar 26
georgei, Tetrapturus 45 Pseudohistiophorus
*georgii, Tetrapturus 45 *illingworthi, 38
*Geulang pajang 26 *illingworthi, Tetrapturus 38
*Giant black marlin 29 *immaculatus, Histiophorus 23
*Ginzasu 51 *immaculatus, Istiophorus 23
*gladifer, Istiophorus 23 *imperator, Tetrapterus 48
*gladius, Histiophorus 23 *imperator, Xiphias 48
*gladius, Istiophorus 23 indica, Makaira 27
 - 62 -

PAGE PAGE

(Istiompax) *Kuchinagafuurai 47
*indica, Makaira 27 *Kudamaki 32,51
*indicus, Histiophorus 23 *Kuro 32
*indicus, Istiomax 27 *Kuroka 32
*indicus, Istiompax 27 *Kurokajiki 32
*indicus, Makaira 27 *Kurokawa 32
*indicus, Tetrapturus 27 *Kurokawakajiki 32
marlin *Kuromazaara 32
Indo-Pacific blue 30 *Kuthira-meen 51
Indo-Pacific sailfish 23 *Kyau-shit-á 29
*Io 51
*Istiomax indicus 27
*Istiompax australis 27
*Istionpax brevirostris 27
*Istiompax dombraini 27
*Istiompax howardi 30
*Istiompax indicus 27
*Istiompax marlina
*Istiompax mazara
27
30
L
ISTIOPHORIDAE 20
Istiophorus albicans 21 *Lamontella albida 35
*Istiophorus amarui 23 *Lan fu yii 32
*Istiophorus americanus 21 *Landjareng 26
*Istiophorus brookei 23 *Layeran 26
*Istiophorus dubius 23 *lessonae, Makaira 35
*Istiophorus eriquius 23 *lessonae, Tetrapturus 35
*Istiophorus gladifer 23 *Lih-ch'ih-Ch'i-yü 29
*Istiophorus gladius 23 *Lijarang 26
*Istiophorus gladius greyi 23 *Liplipan 26,43
*Istiophorus greyi 23 *Lkajar plajaren 26
*Istiophorus greyii 23 Longbill spearfish 46
*Istiophorus immaculatus 23 *ludibundus, Istiophorus 23
*Istiophorus japonicus 23 *Luinniasc 51
*Istjophorus ludibundus 23 *Lumod 51
*Istiophorus maguirei 21
*Istiophorus orientalis 23
Istiophorus platypterus 23
*Istiophorus triactis 23
*Istiophorus volador 21
*Istiophorus wrighti 21
*Izasu 51
M
*Macokljun 51
*Mage 42
*maguirei, Istiophorus 21
*Maka 42
J *Makaira (Istiompax) indica
nigricans
27
*Makaira (Makaira) 30,33
*Jaglun 51 *Makaira albicans 21
*Japanese spearfish 39 *Makaira albida 35
*japonicus, Istiophorus 23 *Makaira ampla 33
*Makaira ampla ampla 33
*Makaira ampla marlina 27
*Makaira ampla mazara 30
*Makaira ampla tahitiensis 27
*Makaira audax 40
*Makaira audax zelandica 40
*Makaira australis 27
K *Makaira belone
*Makaira bermudae
43
33
*Makaira brevirostris 27
*Kadu kpooara 51 *Makaira ensis 33
*Kajikia formosana 40 *Makaira formosana 40
*Kajikia mitsukurii 40 *Makaira grammatica 40
*Kandayan 26 *Makaira herscheli 33
*Kandelan 26 *Makaira herscheri 27
*Kanga 26 *Makaira holei 40
*Kannushi 26 Makaira indica 27
*Katahari 29 *Makaira indicus 27
*Katokui 32 *Makaira lessonae 35
*Katsuokui 32 *Makaira marlina 27
*Ki-hi-khu 51 *Makaira marlina marlina 27
*Koomori 26 tahitiensis
*Kop'jenosjets 39 *Makaira marlina 27
*Kopparaikulla 29 Makaira mazara 30
*Kopparan 29 *Makaira mazara tahitiensis 27
*Kopparava 29 *Makaira nigra 30
*kraussi, Tetrapturus 38 Makaira nigricans 33
*Kuchinaga 47 *Makaira nigricans ampla 33
 - 63 -

PAGE PAGE

*Makaira nigricans marlina 27 *Mersuji 26

*Makaira nigricans mazara 50 *Mesara 51
nigricans *Meshvenosouiye 51
*Makaira nigricans 33 *Minokajiki 26
tahitiensis *mitsukurii, Kajikia 40
*Makaira nigricans 27 *mitsukurii, Marlina 40
*Makaira perezi 33 *mitsukurii, Tetrapturus 40
*Makaira velifera 21 *Mylmeen 26
*Makaira xantholineatus 27
*Makaira zelandica 40
Makaire à rostre court 38
Makaire bécune 46
*Makaire blanc 37
1'Atlantique
Makaire blanc de 36
*Makaire bleu
1'Atlantique
35 N
Makaire bleu de 33
Pacifique *Naeragi 42
Makaire bleu de l'Indo- 30 *Naidonbo 42
Makaire épée 45 *Nairage 42
Makaire noir 27 *Nairagi 42
Makaires 20 *Nairanbo 42
*Makajiki 42 *Nduaro 43
*Makara 29 *Ndwaro 26
*Malasagi 51 *Neeranbo 42
*Malasugi 26,51 *New Zealand m a r l i n 43
*malina, Malina 27 *nigra, Eumakaira 30
*Malina malina 27 *nigra, Makaira 30
*Malyi kopénosets 47 nigricans, Makaira 33
*Manumbuk 51 (Makaira)
*Marasugi 43 *nigricans, Makaira 30,33
*Marlin 29,32 nigricans
*Marlin 43-44 *nigricans, Makaira 33
*Marlin 44 *Nishibashoo 23
*Marlin azul 32,35 *Nishibashookajiki 23
*Marlin bianco 37 *Nishifuuraikajiki 44
*Marlin bleu 32 *Nishikuro 35
Mediterranée *Nishikurokajiki 35
Marlin de la 43 *Nishimaka 37
Marlín del Mediterráneo 43 *Nishimakajiki 37
*Marlín negro 32,35 *Njiachi 32
Marlín peto 45 *Njiara 32
Marlín rayado 40 *Nog-sae-chi 35
Marlin rayé 40 *Nok-sae-chi 32
Marlín trompa corta 38 *Nooragi 42
*Marlina audax 40 *Nourage 26
*marlina, Istiompax 27 *Nouragi 42
*marlina, Makaira 27 *Nsuli nsuli 26
*marlina, Makairampla 27 *Nsulinsuli 26
*marlina, Makaira marlina 27
ns
*marlina, Makaira nigrica 27
*Marlina marlina 27
*marlina, Marlina 27
*Marlina mitsukurii 40
*Marlina zelandica 40
Marlines
Marlins
20
20
O
*Mas-hibaru 51
*Masashi 42 *Oba 26
*Masasu 42 *Ocean gar 23
*Mayaspus 51 *Oiragi 42
*Mazaara 32 *Oiraki 42
*mazara, Istiompax 30 *Okajiki 42
mazara, Makaira 30 *Okizaara 51
*mazara, Makairanpla 30 *Ola-meen 26
s *Onyankle 23
*mazara, Makaira nigrican 30 *orientalis, Histiophorus 23
*mazara, Tetrapturus 30 *orientalis, Istiophorus 23
*Mbassi 26 *Orthocraeros bermudae 33
*Mbassi kouri 26 *Oumbajhe 23
*Mech-ryba 51
*Mechenos 51
*Medara 51
spearfish
*Mediterranean shortbill 44
Mediterranean spearfish 43
*Meka 51
*Mekajiki 51
*Mère 23
*Mersudji 32
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*Sere bristyi marlin 29

*Shiro 29
P *Shiroka
*Shirokajiki
29
29
*Shirokawa 29
*Pacific black marlin 29 *Shirokawajiki 29
*Pacific sailfish 26 *Shiromazaara 29
*Pacific striped marlin *Shiruachi 29
*Panombuk 26 *Shitore 42
*Parusnik 23,26 *Shiutome 42,51
*Parusnik-ryba 23,26 Shortbill spearfish 38
*Pastardella 44 *Shortnose spearfish 39
*Peacock fish 26 *Shortnosed spearfish 39
Peces vela 20 *Shutome 51
*Peito 35 *Silver marlin 29
*Peixe agulha 35,51 *Skeponopodus guebucu 21
*Peixe de vela 23 *Skeponopodus typus 43
*Peixe espada 51 *Skilligalee 37
*perezi, Makaira 33 *Slender spearfish 39
* Pesce spada 51 *Spearfish 43
*Pez aguja 37,42 Spearfishes 20
*Pez aguja corta 39 *S pikefish 43
Pez espada 48 *Squadron 23,35
*Pez puerco 48 Striped marlin 40
*Pez vela 23,26 *Striped swordfish 43
Pez vela del Atlántico 21 *S tuji stuji 26
Pac ífico *Suginairage 26
Pez vela del Indo- 23 *Suginairagi 26
* Pez zuncho 32 *Sugiyama 26,39
pfluegeri, Tetrapturus 46 *Suzu 51
tuberculatus *Svaerdfisk 51
*Phaethonichthys 48 *Sverdfisk 51
*Piscispat 51 *Swaardvis 51
* Pixxi spad 51 *Swardfisk 51
platypterus, Istiophorus 23 Swordfish 48
*platypterus, Xiphias 23
* Poisson-pique 44
*Polosatii marlin 43
*Prieta 23,35
*prototypus, Scheponopodus 43
angustirostris
*Pseudohistiophorus 38
illingworthi
* Pseudohistiophorus 38
T
*pulchellus, Histiophorus 21
* Tadmasa 26
*tahi tiensis, Makairampla 27
marlina
*tahitiensis, Makaira 27
mazara
*tahitiensis, Makaira 27
nigricans
R *tahitiensis, Makaira
* Taketonga
27
32
*Tenguzawara 42
*Rakuda 51 Tetrapturus
*Reste cuspada 51 *tenuirostratus, 40
*rondeletti, Xiphias 48 *Teppo 51
Roundscale spearfish 45 *Tetrapterus imperator 48
*Tetrapturus acutirostratus 40
Tetrapturus albidus 35
*Tetrapturus amplus 33
Tetrapturus angustirostris 38
Tetrapturus audax 40
*Tetrapturus australis 27
Tetrapturus belone 43
S *Tetrapturus brevirostris
*Tetrapturus ectenes
27
40
Tetrapturus georgei 45
*Sablijck 51 *Tetrapturus georgii 45
*Sail 23 *Tetrapturus herschelii 33
*Sailfin 26 *Tetrapturus illingworthi 38
*Sailfish 23,26 *Tetrapturus indicus 27
Sailfishes 20 *Tetrapturus kraussi 38
*Sanmakajiki 39 *Tetrapturus lessonae 35
*Saparava 29 *Tetrapturus mazara 30
*Sashi 42 *Tetrapturus mitsukurii 40
*Sasu 42 Tetrapturus pfluegeri 46
*Scheponopodus prototypus 43 *Tetrapturus tenuirostratus 40
*Schwertfisch 51 * Thalapatha 26
*Scomber gladius 23 *thermaicus, Xiphias 48
* Seilvis 23 *Tinmankhu 51
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*Tobihira 26
*Tong-fang-chiyii 26
*triactis, Istiophorus
*Tsukinbo
23
43
Y
*Tsun 32,51
Phaethonichthys *Yemungolah 26
*tuberculatus, 48 *Yu-san-chi-yu 26
*typus, Skeponopodus 43 *Yu-san-yu 26

U Z
*Ugghia 'mpriali 44 *Zegal-fisch 23
*Ugulia imperiali 44 *zelandica, M akaira 40
*Unjiachi 43 *zelandica, Makairaudax 40
*zelandica, Marlina 40
* Zwaardvisch 51

V
*Veleiro 23
*Veleiro do Atlantico 23
*velifera, Makaira 21
*velifer, Xiphias 21,23
*Voilier 23,26
Voilier de l'Atlantique 21
Pacifique
Voilier de l'Indo- 23
Voiliers 20
*Voladeira 23
*Volador 23,26
*Voladora 35
*volador, Istiophorus 21

W
*Whang-Sae-chi 51
*White marlin 37
*Wit marlyn 37
*wrighti, Istiophorus 21

X
*xantholineatus, Makaira 27
*Xiphias 51
*Xiphias estara 48
Xiphias gladius 48
*Xiphias gladius estara 48
*Xiphias imperator 48
*Xiphias platypterus 23
*Xiphias rondeletti 48
*Xiphias thermaicus 48
*Xiphias velifer 21,23
XIPHIIDAE 48