Identification Guide to
Pufferfishes (Tetraodontidae, Tetraodontiformes)of the South China Sea

Keiichi Matsuura
Hiroyuki Motomura

The Kagoshima University Museum2022

Identification Guide to

Pufferfishes(Tetraodontidae, Tetraodontiformes)ofof the South China Sea

Keiichi Matsuura and Hiroyuki Motomura

K. Matsuura & H. Motomura
Authors
Keiichi Matsuura, PhD
Curator Emeritus
National Museum of Nature and Science
4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan
E-mail: matsuura@kahaku.go.jp

Bibliographic info

Hiroyuki Motomura, PhD
Professor, Director
The Kagoshima University Museum
1-21-30 Korimoto, Kagoshima 890-0065, Japan
E-mail: motomura@kaum.kagoshima-u.ac.jp

Juveniles of Torquigener hypselogeneion,
Okinoerabu-jima Island, photo by K. Uehara

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Copyright © 2022 by the Kagoshima University Museum
All rights reserved. No part of this publication may be reproduced or transmitted in any form or by
any means without prior written permission from the publisher. Copyrights of the photographs areheld by the photographers.
For bibliographic purposes this book should be cited as follows:
Matsuura, K. and H. Motomura. 2022 (Feb.). Identification guide to pufferfishes (Tetraodontidae,
Tetraodontiformes) of the South China Sea. The Kagoshima University Museum, Kagoshima. 40 pp.,
70 figs.
ISBN 978-4-905464-20-4
Corresponding author: Keiichi Matsuura
Design & DTP by Hiroyuki Motomura
Printing by Asahi Co. Ltd. (Kagoshima City)
Published on 1 February 2022
Cover image: Eye of Canthigaster axiologus, off Yaku-shima Island, Kagoshima Prefecture, Japan,
photo by S. Harazaki.
Inside cover image: Arothron manilensis, off Okinoerabu-jima Island, Kagoshima Prefecture, Japan,
photo by K. Uehara.
Back cover image: same as Fig. 1b.

Identification Guide to Pufferfishes of the South China Sea

Introduction

Introduction

Pufferfishes of the family Tetraodontidae differ externally from other families of the order Tetraodontiformes by the following combination of characters: head large and blunt; jaws modified to form a beak offour heavy, powerful teeth, two above and two below; eyes high on head; gill opening, a simple slit in frontof pectoral fins; dorsal and anal fins located far posteriorly, containing seven to 15 soft rays; caudal fin truncate, rounded, or emarginate to somewhat lunate; pelvic fins absent; lateral line (when present) often indistinct, forming an interconnected pattern on sides of the head and body, but quite distinct in some genera(e.g., Lagocephalus and Torquigener); typical scales absent, but many spinules often present on back and/or belly, and sometimes on sides (Matsuura 2001).
The Tetraodontidae is the most speciose family in the Tetraodontiformes, including about 190 species.
Because pufferfishes possess very few external characters useful for taxonomy and specimens are easily
distorted when they are fixed in formalin and preserved in ethanol, it is often not easy for ichthyologists to
identify them to species. The taxonomy of pufferfishes in the South China Sea is important not only for theunderstanding of fish diversity of the region, but also for the welfare and food management for humans. Daoet al. (2012) reported that the number of victims of food poisoning from eating pufferfishes reached 737, with
127 mortalities from 1999 to 2003. Due to the lack of knowledge about pufferfishes and their toxicity, poisonous pufferfishes have been frequently found at fish landing places and fish markets in countries around theSouth China Sea (Fig. 1), although local and state governments in these countries have announced that eating pufferfishes are quite dangerous for humans. The main purpose of this identification guide is to provide
distinguishing characters of pufferfishes of the South China Sea, which would be helpful for local ichthyologists and food security people to identify pufferfishes and prevent food poisoning in their countries.

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Fig. 1. Pufferfishes found at a fish landing place 60 km north of NhaTrang in southern
Vietnam (a) and a fish market in Malaysia (b). Photos by K. Matsuura.

K. Matsuura & H. Motomura

Methods

Methods

Counts and measurements follow Dekkers (1975). Standard lengthand total length are abbreviated as SL and TL, respectively. Institutional
abbreviations are as follows: BPBM (Bernice Pauahi Bishop Museum,
Honolulu), BSKU (Department of Natural Science, Kochi University,
Kochi), CSIRO (Commonwealth Scientific and Industrial Research Organisation, Hobart), FAKU (Maizuru Fisheries Research Station, Kyoto
University, Maizuru), FRLM (Fisheries Research Laboratory, Mie University), HUMZ (Hokkaido University Museum, Hakodate), KAUM (The
Kagoshima University Museum, Kagoshima), KPM (Kanagawa Prefectural Museum of Natural History, Odawara), MUFS (Miyazaki University,
Miyazaki), NMMB (National Museum of Marine Biology and Aquarium,
Pingtung), NRF-SAIAB (National Research Foundation - South African
Institute for Aquatic Biodiversity), NSMT (National Museum of Natureand Science, Tsukuba), SMSM (Shimonoseki Marine Science Museum,
Shimonoseki), and USNM (National Museum of Natural History, Washington, D. C.). Credit of photographs taken by H. Senou belongs to the
Kanagawa Prefectural Museum of Natural History, Odawara. Illustrations of Amblyrhynchote honckenii drawn by E. Heemstra were provided by the South African Institute for Aquatic Biodiversity.

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Takifugu bimaculatus, 33 cm SL, Yellow Sea, drawn by K. Matsuura.

Identification Guide to Pufferfishes of the South China Sea

Key to Genera

Key to Genera

1a. Snout long, its length 55–77% of head length; nasal organ barely
visible without aid of magnification; dorsal surface posterior to theeye distinctly keeled (Fig. 2a, b) ......………………….... Canthigaster1b. Snout short, its length 32–55% of head length; nasal organ easily visible without magnification; dorsal surface posterior to the eyemore or less smooth, without a distinct keel (Fig. 2c, d) ………....... 2

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Fig. 2. Typical body shapes of pufferfishes. Top, Canthigaster (a, front view; b, lateral view); bottom, Arothron (c, front view; d,lateral view). Fins are not shown in front views.

2a. Nasal organ covered by a small sac with 2 nostrils (Fig. 3a) ...…... 3
2b. Nasal organ not covered by a small sac with 2 nostrils (Fig. 3b–d) ..
...................................................................................................….... 8

Fig. 3. Nasal organs of pufferfishes. a, Lagocephalus; b, Arothron; c, Chelonodontops; d, Canthigaster.

K. Matsuura & H. Motomura
3a. No raised skin fold along the ventral side of the caudal peduncle ... 4
3b. A raised skin fold along the ventral side of the caudal peduncle) ... 5

Key to Genera

4a. No spinules on head and body ..........…………............ Sphoeroides
4b. Spinules all over the head and body except for the caudal peduncle............................................................................................... Tylerius
5a. Mouth supraterminal (Fig. 4a) ....……………........ Amblyrhynchote5b. Mouth terminal (Fig. 4b–d) .…………………………………….......... 6
6a. Chin distinct (Fig. 4a, b) ..........................…………....... Torquigener
6b. Chin indistinct (Fig. 4c, d) ...………………...................................... 7

Fig. 4. Mouth positions and chin shapes of pufferfishes. a, Amblyrhynchote honckenii; b, Torquigener brevipinnis; c, Arothronstellatus; d, Takifugu flavipterus.

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7a. A longitudinal wide silver-white stripe running from chin to caudal-finbase on ventro-lateral part of head and body ....……. Lagocephalus
7b. Body variously colored, but no longitudinal silver-white stripe on
ventro-lateral part of body …................................................ Takifugu
8a. A single lateral line on the side of the body (Fig. 5a); nasal organwith 2 bifid tentacles (Fig. 3b) ….......................................... Arothron
8b. Two lateral lines, the upper joining the lower in the region above
or behind anal fin (Fig. 5b); nasal organ in the form of a depression
with slightly raised margin expanded before and behind into a pair ofelongate flaps (Fig. 5c) ….........……….....….......... Chelonodontops

Fig. 5. Lateral line system of Arothron (a) and Chelonodontops (b).

Identification Guide to Pufferfishes of the South China Sea

Genus Amblyrhynchote Bibron in Duméril 1855

Amblyrhynchote honckenii (Bloch 1785)
Evileye Pufferfish

Amblyrhynchote honckenii, BPBM 21755, 8.4 cm SL, Natal, South Africa, photo by J. E. Randall.

Amblyrhynchote honckenii, 19 cm TL (above, breeding female), 8 cm TL (below, juvenile), South Africa,
drawn by E. Heemstra, courtesy of NRF-SAIAB.

Dorsal-fin rays 9–10; anal-fin rays 8; pectoral-fin rays 14–16; caudal peduncle short, its
length 22–28 % of SL; bony interorbital width 13–26% of head length. Body somewhat squarish
in cross section. Dorsal surface of body blackish brown with yellowish white spots; lateral side
yellow; ventral surface whitish; dorsal, pectoral and caudal fins dusky with yellow tinge; anal
fin in breeding females yellow, in males white. Attains 30 cm TL. Collected at depths shallower
than 400 m. Indo-West Pacific from East Africa to China.

Genus Amblyrhynchote

Mouth supraterminal situated above dorsal end of pectoral-fin base; chin prominent; nasal organ covered by a small sac with 2 nostrils; ventrolateral skin foldprominent on area between caudal-fin base and anus; small spinules on back andbelly. Monotypic. Although previous authors used the spelling Amblyrhynchotes forthis genus (e.g., Hardy 1984, 1988), we follow Kottelat (2001) in using Amblyrhynchote because Kottelat (2001) showed that Amblyrhynchotes is an incorrect spelling.

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K. Matsuura & H. Motomura

Genus Arothron Müller 1841

Genus Arothron

Body ovoid, heavy; single lateral line on lateral side of body; no skinfold on lower side of body; nasal organ with 2 bifid tentacles; spinules
present on body. Most members of Arothron solitary but young of A. firmamentum form schools. Fourteen species in the world, 12 species inSouth China Sea.

Key to Species of Arothron
1a. Body lacking markings; caudal fin dusky yellow with prominent black margin …...............
................................................…………………………………………...... A. immaculatus
1b. Body with various color markings; caudal fin without black margin ................................ 22a. Dorsal-fin rays 13–15 (usually14); anal-fin rays 13–15 (usually14); dorsal and anal fins
bluntly pointed …………..............................................................……… A. firmamentum
2b. Dorsal-fin rays 9–13: anal-fin rays 9–13; dorsal and anal fins rounded …..................… 3
3a. Eye encircled by alternating dark brown and pale blue lines ....… A. caeruleopunctatus
3b. Eye not encircled by lines …………………………………………………....……………... 44a. Many dark brown lines radiating from eye ……………….………..…………….. A. mappa
4b. No radiating lines from eye ……………………………....…………………………………. 5

85a. Wide dark vertical bars on side of head and ventrolateral side of body; dorsal part ofbody greenish brown with many white spots ………..…………….....………… A. hispidus
5b. No dark vertical bars on head or body …………………….....……………………………...6
6a. Dorsal part of body white …………………………………………………….....…..……….. 7
6b. Dorsal part of body greenish brown, dark brown, dark gray, light gray, yellow or black .. 8
7a. Body white with many black spots, forming a network pattern; ventral surface of headand body white without black spots (in juveniles many dark bands on belly) ...................
....................................................................................................................... A. stellatus
7b. Dorsal and lateral sides of body white with many longitudinal black lines ….. A. carduus
8a. Many longitudinal lines on body ………......………………..……………………….……… 98b. No longitudinal lines on body …………………......………………………………….…… 10
9a. Several longitudinal brown lines on light gray or light brown body ……….. A. manilensis
9b. Many longitudinal white lines on dark brown or greenish-dark brown body…………….....
...............................…………………………………………………………..A. multilineatus10a. Several curved white lines on side of head ……….……………………….. A. reticularis
10b. No curved lines on side of head …………………………………….....………………… 1111a. Body dark brown or brownish gray with black spots ……………… A. nigropunctatus11b. Body and fins black with many white spots or body yellow with many black or darkbrown speckles ……....…………………………………..…….……….………. A. meleagris

Identification Guide to Pufferfishes of the South China Sea
Arothron caeruleopunctatus Matsuura 1994
Blue-spotted Pufferfish

Genus Arothron
Arothron caeruleopunctatus, NSMT-P 45451, 42.4 cm SL, Bikini Atoll, Marshall Islands, photo by H. Kishimoto.

Dorsal-fin rays 11–12, anal-fin rays 10–12, pectoral-fin rays 18–19. A large pufferfish attaining 80 cm TL; head and body brown dorsally, covered with many pale blue spots; ventral
regions of head and body pale, with or without black blotches just below pectoral fin; eye encircled by alternating dark brown and pale blue lines. Occurs solitarily on coral reefs. Indo-West
Pacific, from Réunion eastward through Indonesia to Coral Sea and northward to Japan.

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Arothron carduus (Cantor 1849)
Black-lined Pufferfish

Arothron carduus, HUMZ 35438, 19.5 cm SL, Ishigaki-jima Island, Ryukyu Islands, photo by K. Matsuura.

Dorsal fin ray 10–11, anal-fin rays 9–10, pectoral-fin rays 18–19. Head and body white withmany black lines, some encircling eye and gill opening; side of body behind dorsal fin with longitudinal black lines; ventral surface of body yellowish; dorsal and anal fins yellowish brown;
pectoral fin dusky yellow; caudal fin white with many longitudinal black lines. Attains 43 cm TL.
Collected at depths shallower than 30 m. Known only from Ryukyu Islands and Penang Island,
Malaysia.

K. Matsuura & H. Motomura
Arothron firmamentum (Temminck & Schlegel 1850)
Starry Pufferfish

Genus Arothron
Arothron firmamentum, KAUM–I. 30569, 9.4 cm SL, Iwo-jima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 13–15 (usually14), anal-fin rays 13–15 (usually14), pectoral-fin rays 15–17(usually 16). Body relatively oblong compared to other congeners; head and body bluish black,covered with many white spots. Attains 45 cm TL. Collected at depths of 100–140 m. Antitropical in West Pacific, Japan to Taiwan, south-eastern Australia and northern New Zealand; recorded also from South Africa and Argentina.

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Arothron hispidus (Linnaeus 1758)
White-spotted Pufferfish

Arothron hispidus, KPM-NI 31894, 26.8 cm SL, Iriomote-jima Island, Ryukyu Islands, photo (KPM-NR 107143A) by H. Senou.

Dorsal-fin rays 10–11, anal-fin rays 10–11, pectoral-fin rays 17–19. Head and body greenish
brown with small white spots dorsally, becoming white ventrally with curved dark stripes; wide
dark bars on side of head and body; pectoral-fin base and gill opening encircled with alternating
white and black lines. Attains 45 cm TL. Inhabits coral reefs, but juveniles frequently occur inestuaries. Indian and Pacific oceans, from East Africa eastward to tropical eastern Pacific.

Identification Guide to Pufferfishes of the South China Sea
Arothron immaculatus (Bloch & Schneider 1801)
Black-edged Pufferfish

Genus Arothron
Arothron immaculatus, KAUM–I. 43918, 16.3 cm SL, Uchinoura Bay, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 9–11 (usually 10), anal-fin rays 9–10 (usually 10), pectoral-fin rays 16–19.
Body light brown with olivaceous tinge; dorsal and anal fins yellowish, caudal fin yellow withbroad black edge, pectoral-fin rays yellow with hyaline membranes. Attains 30 cm TL. Occurs
in estuaries, and particularly common in areas with sea grasses and mangroves. Indo-West
Pacific from East Africa eastward to Indonesia and northward to Japan.

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Arothron manilensis (Marion de Procé 1852)
Striped Pufferfish

Arothron manilensis, HUMZ 41440, 24 cm SL, Iriomote-jima Island, Ryukyu Islands, photo by K. Matsuura.

Dorsal-fin rays 9–11 (usually 10), anal-fin rays 9–10 (usually10), pectoral-fin rays 16–19.
Body light gray or light brown with greenish tinge; ventral side of body whitish or light yellowish;
several longitudinal dark brown lines on body; caudal fin light yellow or brownish yellow withblack margins. Occurs in mangroves, sea grass beds, and sandy-muddy bottoms in depths
shallower than 20 m. Tropical western Pacific eastward to Samoa and northward to Japan.

K. Matsuura & H. Motomura
Arothron mappa (Linnaeus 1758)
Map Pufferfish

Genus Arothron
Arothron mappa, KAUM–I. 22222, 14 cm SL, Malaysia, photo courtesy of KAUM.

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Dorsal-fin rays 11–12, anal-fin rays 10–11 (usually11), pectoral-fin rays 17–19. A large pufferfish, attaining 60 cm TL; head and body brownish gray with many irregular dark lines dorsally,
appearing as numerous reticulations; ventral parts of head and body white, except for blackanus; reticulated lines developing with growth of fish, making color patterns of juveniles andadults different, but with dark lines radiating from the eye at every stage of growth. Coral reef
inhabitant but juveniles occasionally occur in estuaries. Indo-West Pacific from East Africa eastward to Marshall Islands and northward to Japan.
Arothron meleagris (Bloch & Schneider 1801)
Guineafowl Pufferfish

Arothron meleagris, KAUM–I. 37803, 25.8 cm SL, Iwo-jima Island, Kagoshima Prefecture.
Yellow phase (upper right), KAUM–I. 32360, 27.9 cm SL, Iwo-jima Island. Photos courtesy of KAUM

Dorsal-fin rays 10–13 (usually 11–12), anal-fin rays 11–13 (usually 12), pectoral-fin rays 18–
21 (usually 19–20). Entire body usually black with many white spots; fins also black with whitespots; also a yellow form with many dark brown spots on body; mouth not surrounded by darkarea. Attains 35 cm TL. Inhabits coral reefs. Tropical Indian and Pacific oceans from East Africa
eastward through Indonesia to tropical eastern Pacific.

Identification Guide to Pufferfishes of the South China Sea
Arothron multilineatus Matsuura 2016
Many-lined Pufferfish

Genus Arothron
Arothron multilineatus, KAUM–I. 13606, 34.3 cm SL, Kawajiri, Kagoshima Prefecture, photo courtesy of KAUM.

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Arothron multilineatus, MUFS 46817, 11.6 cm SL, Iorigawa, Miyazaki Prefecture. Photo by M. Wada.

Dorsal-fin rays 9–11, anal-fin rays 9–11, pectoral-fin rays 18–19. Lateral and dorsal surfaces of head and body dark brown or greenish-dark brown covered with many longitudinal whitelines; ventral surface of head and body white with light yellowish tinge; many spinules on headand body except on the lateral surface of the caudal peduncle, around mouth, eye, gill opening,and on dorsal- and anal-fin bases. Attains 25 cm TL. Occurs in coral reefs. Indo-West Pacificfrom East Africa eastward to Samoa and northward to Ryukyu Islands.

K. Matsuura & H. Motomura
Arothron nigropunctatus (Bloch & Schneider 1801)
Blackspotted Pufferfish

Genus Arothron
Arothron nigropunctatus, KAUM–I. 39888. 25.8 cm, Iwo-jima Island, Kagoshima Prefecture. Photo courtesy of KAUM.

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Arothron nigropunctatus, BSKU 108306, 16.9 cm SL, Yoron-jima Island, Kagoshima Prefecture. Photo courtesy of BSKU.

Dorsal-fin rays 9–11 (usually 10); anal-fin rays 10–11 (usually 11); pectoral-fin rays 17–20(usually 18–19). Body dark brown or dark gray dorsally, paler ventrally, with many dark brownor black spots; also a yellow form similar to A. meleagris but distinguished from it by the presence of a dark area around mouth. Attains 25 cm TL. Occurs in coral reefs; Indo-West Pacificfrom East Africa eastward to Samoa and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea
Arothron reticularis (Bloch & Schneider 1801)
Reticulated Pufferfish

Genus Arothron
Arothron reticularis, KPM-NI 31898, 34.1 cm SL, Iriomote-jima Island, Ryukyu Islands, photo (KPM-NR 107147A) by H. Senou.

Dorsal-fin rays 10–11; anal-fin rays 9–11; pectoral-fin rays 17–19. Body greenish brown dorsally, white ventrally; many white spots on side and back behind eye; several curved white lines
on side of head and around gill opening; dorsal, anal, and pectoral fins pale yellow; caudal fingreenish brown with many white spots. Attains 43 cm TL. Occurs around mangroves and sandyor muddy bottoms of sheltered bays at depths shallower than 20 m; India eastward to Fiji andnorthward to Japan. Similar to Arothron hispidus but distinguishable by having several curved
white lines on side of head and lacking vertical dark bars on ventral side of body.
Arothron stellatus (Bloch & Schneider 1801)
Star Pufferfish

Arothron stellatus, KAUM–I. 52638, 33.9 cm SL, Tanega-shima Island, Kagoshima Prefecture.
Juvenile (upper right), KAUM–I. 51417, 4.5 cm SL, Yoron-jima Island, Kagoshima Prefecture. Photos courtesy of KAUM.

Dorsal-fin rays 11–12, anal-fin rays 11, pectoral-fin rays 17–20 (usually 18). A large pufferfish, attaining 100 cm TL; body white with many black spots dorsally forming a network pattern;small black spots also covering vertical fins; pectoral-fin base and gill opening with black spots
larger than those elsewhere on of the body; ventral side of head and body white with blackanus; juveniles orange with many black spots dorsally and many irregular oblique lines on belly.
Occurs in coral reefs; Indo-West Pacific, from East Africa eastward through Indonesia to Tuamotu Islands and northward to Japan.

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K. Matsuura & H. Motomura

Genus Canthigaster Swainson 1839

Genus Canthigaster

Body and head somewhat compressed; a single inconspicuous nostril; lateral lines usually not apparent; back with a skin ridge on dorsal
midline; caudal fin truncate or slightly rounded; head and body with
scattered tiny spinules. Thirty-two species in the world, 13 species in
the South China Sea.

Key to Species of Canthigaster
1a. Dorsal-fin rays 11–12, anal-fin rays 10–11 ………………………………. C. amboinensis
1b. Dorsal-fin rays 8–10, anal-fin rays 8–10 …………………...……………………………… 2
2a. Dorsal part of head and body with 2 or 4 dark brown or black saddle-like bars ……..…. 3
2b. Dorsal part of head and body without dark saddle-like bars ……………………..………. 4
3a. Dorsal part of caudal peduncle covered with a dark saddle-like bar ….……….………… 5
3b. Dorsal part of caudal peduncle without a dark saddle-like bar ….……..… C. ocellicincta
4a. A rounded black blotch with a diameter subequal to eye on ventral side of body between
pectoral-fin base and anal-fin origin ……………………………...……..….. C. inframacula
4b. No rounded black blotch on ventral side of body ……………....…………………………. 6
5a. Second and third dark saddle-like bars extending ventrally to belly ………… C. valentini
5b. Second and third dark saddle-like bars extending ventrally to level of ventral end of
pectoral-fin origin, not extending to belly ………………………….........…….. C. axiologus

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6a. Side of body with 1 or 2 longitudinal stripes (sometimes fragmented) from pectoral-fin
region to caudal peduncle ………….......…………………………………..……………..…. 7
6b. Side of body without longitudinal stripes ……………....……………………….….....….... 8
7a. Side of body with 2 parallel, longitudinal stripes joining to form an arch in front of gill
opening ………………………………………….…..…..............………………….….......… 97b. Side of body with 1 dark longitudinal stripe, equal to, or slightly wider than eye diameter,
running from upper gill opening to caudal peduncle …....………………...…… C. bennetti
8a. Side of body pale brown with clusters of dark brown spots ……...………….. C. leoparda
8b. No clusters of dark brown spots on side of body …………………..…………………….. 10
9a. Dorsal part of body brown with many wavy dark blue lines; 2 parallel yellow stripes onbody much narrower than diameter of pupil; caudal fin yellow with longitudinal dark bluelines …………………………................…………………………………..…….… C. rivulata
9b. Dorsal part of body light brown with many blue spots; 2 parallel, narrow yellow stripes onbody; caudal fin light yellow with several transverse light blue lines ….......…. C. punctata
10a. Body dark brown dorsally and white ventrally without white spots; caudal fin yellow……………………..........………………………………………………….……. C. epilampra
10b. Body brown or reddish brown with many white spots ……….…………………………. 11
11a. No longitudinal pale blue to white lines on back ….……………………. C. janthinoptera
11b. Longitudinal blue to white lines on back ………………......……………………………. 1212a. Body brown with combination of pale blue to white wavy lines and white spots; paleblue-edged, black ocellus on dorsal-fin base; caudal fin reddish brown with manytransverse pale blue lines …............………………………………….……… C. compressa
12b. Body reddish brown with many small blue-white spots and short wavy lines on back;pale blue-edged, black ocellus on dorsal-fin base; several blue-white lines on snout;caudal fin orange with many small blue-white spots …………...………...………. C. papua

Identification Guide to Pufferfishes of the South China Sea
Canthigaster amboinensis (Bleeker 1864)
Spotted Toby

Genus Canthigaster

Canthigaster amboinensis, KPM-NI 30731, 7.7 cm SL, Yoron-jima Island, Kagoshima Prefecture,
photo (KPM-NR 106625B) by H. Senou.

Dorsal-fin rays 10–12 (usually 12); anal-fin rays 10–11 (usually 11); pectoral-fin rays 16–17(usually 17). Body brown with numerous white spots; several white lines radiating from eye;cheek with many pale blue spots or lines; snout with several narrow pale blue lines; dorsal,
anal, and pectoral fins pale with brown base; caudal fin brown with pale blue spots proximally.
Attains 15 cm TL. Inhabits very shallow water, frequently in lower reaches of surge zone; eats
mostly small invertebrates. Indo-Pacific from East Africa eastward through Indonesia to Fiji andnorthward to Japan.
Canthigaster axiologus (Whitley 1931)
Saddle Toby

Canthigaster axiologus, KAUM–I. 55127, 6.9 cm SL, Yoron-jima Island, Kagoshima Prefecture,
photo courtesy of KAUM.

Dorsal-fin rays 9–10 (usually 10); anal-fin rays 9–10 (usually 9); pectoral-fin rays 16–17(usually 17). Head and body with 4 black, saddle-like bars, first on top of head, second between
pectoral-fin bases, third on middle of body, and fourth on caudal peduncle, the two middle bars
extending ventrally to level of pectoral-fin base, not reaching belly; ventral half of body whitewith many orange spots; eye with several radiating pale blue lines; dorsal, anal, and pectoral
fins pale; caudal fin pale with a black spot at top and bottom of base. Attains 14 cm TL. Foundin depths of 10–80 m, but usually in shallow depths. Feeds primarily on invertebrates, especially mollusks. West Pacific southward to New South Wales in Australia and northward to Japan.

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K. Matsuura & H. Motomura
Canthigaster bennetti (Bleeker 1854)
Exquisite Toby

Genus Canthigaster
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Canthigaster bennetti, BPBM 15579, 4.0 cm SL, Guadalcanal Island, Solomon Islands, photo by J. E. Randall.

Dorsal-fin rays 9–11 (usually 9–10); anal-fin rays 8–10 (usually 9); pectoral-fin rays 14–16(usually 15–16). Greenish brown dorsally, with irregular orange lines and spots on back; whitespots on side and back posterior to pectoral fin; side of body with a dark brown longitudinal
stripe from gill opening to caudal peduncle; ventral half of body white with many pale blue
spots; eye with several radiating blue lines; several vertical blue lines just posterior to mouth;
dorsal-fin base with black ocellus and irregular blue lines; dorsal, anal, and pectoral fins pale;caudal fin greenish brown. Attains 10 cm TL. Inhabits inner reef flats and sheltered lagoons, often on algal or silty reefs and among attached Sargassum on shallow rubble flats. Feeds mainly on filamentous green algae, sponges and other benthic invertebrates. Indo-Pacific from East
Africa eastward through Indonesia to Tuamotu Islands and northward to Japan.

Canthigaster compressa (Marion de Procé 1822)
Compressed Toby

Canthigaster compressa, BPBM 16000, 5.7 cm SL, Guadalcanal Island, Solomon Islands, photo by J. E. Randall.

Dorsal-fin rays 8–10 (usually 9); anal-fin rays 8–9 (usually 9); pectoral-fin rays 15–18. Body
brown with combination of pale blue to white wavy lines and white spots; pale blue-edged black
ocellus on dorsal-fin base; caudal fin reddish brown with pale blue spots forming vertical pale
blue lines. Attains 11 cm TL. Usually found in depths shallower than 25 m at silty bays and harbors. Western Pacific from Indonesia eastward to Vanuatu and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea
Canthigaster epilampra (Jenkins 1903)
Lantern Toby

Genus Canthigaster
Canthigaster epilampra, BPBM 35231, 6.6 cm SL, Ani-jima Island, Ogasawara Islands, photo by J. E. Randall.

Dorsal-fin rays 9–10 (usually 10); anal-fin rays 9–10 (usually 9); pectoral-fin rays 16–17(usually 17). Ventral half of body creamy white covered with many blue-white spots and shortlines, dorsal half dark brown; several blue lines radiating from eye; several blue lines on snout;
dorsal, anal, and pectoral fins pale; caudal fin yellow with longitudinal blue lines. Attains 11
cm TL. Usually occurs on steep outer reef slopes below 25 m. Western Pacific from Indonesia
eastward to Society Islands and northward to Japan; also recorded from Christmas Island ineastern Indian Ocean.

19Canthigaster inframacula Allen & Randall 1977
Bellyspot Toby

Canthigaster inframacula, USNM 20843, 7.6 cm SL, Oahu, Hawaiian Islands, photo by J. E. Randall.

Dorsal-fin rays 10–11; anal-fin rays 10–11; pectoral-fin rays 17–18. Body light brown dorsally, becoming whitish ventrally; a longitudinal dark brown stripe running from eye to upper baseof caudal fin; a round dark brown spot with diameter about equal to eye, on lower side of body
between lower base of pectoral fin and origin of anal fin; dark brown lines radiating from eye;short, irregular, dark brown lines on dorsal half of body; dorsal, anal, and pectoral fins pale;caudal fin pale with several dark yellowish spots in transverse rows on rays. Attains 9.5 cm TL.
Collected at depths of 124–247 m. Known only from Hawaiian Islands, Tori-shima Island, Japan, and central Vietnam.

K. Matsuura & H. Motomura
Canthigaster janthinopter (Bleeker 1855)
Honeycomb Toby

Genus Canthigaster
Canthigaster janthinoptera, KAUM–I. 20296, 5.1 cm SL, Yaku-shima Island, Kagoshima Prefecture, photo courtesy of KAUM.

20

Dorsal-fin rays 8–10 (usually 9); anal-fin rays 8–10 (usually 9); pectoral-fin rays 15–18 (usually 16–17). Body reddish brown with many white spots on sides of head and body forming reddish brown reticulations; eye with several radiating reddish brown lines; a dark ocellus at dorsal-fin base sometimes present; dorsal, anal, and pectoral fins pale; caudal fin reddish brown orbrown. Attains 9 cm TL. Occurs in clear lagoon and on seaward reefs, solitary or paired. Feeds
on sponges, polychaetes, other invertebrates and filamentous algae. Indo-Pacific from East Africa eastward through Indonesia to Society Islands and northward to Japan.
Canthigaster leoparda Lubbock & Allen 1979
Leopard Toby

Canthigaster leoparda, BPBM 28498, 5.8 cm SL, Cebu, Philippines, photo by J. E. Randall.

Dorsal-fin rays 8–9; anal-fin rays 9; pectoral-fin rays 16. Body light brown with clusters of
irregular brown spots on side and blue lines across dorsal side of snout; many light blue spots
on ventral side of head and body; many alternating light blue and orange lines radiating fromeye; pectoral fin pale; dorsal and anal fins light bluish; caudal fin light bluish with yellow upperand lower rays. Attains 7 cm TL. Inhabits depths of 30–50 m; solitary or in pairs. Christmas Island (Indian Ocean), Indonesia, Philippines, and Mariana Islands.

Identification Guide to Pufferfishes of the South China Sea
Canthigaster ocellicincta Allen & Randall 1977
Shy Toby

Genus Canthigaster
Canthigaster ocellicincta, BPBM 15993, 5.0 cm SL, Florida Island, Solomon Islands, photo by J. E. Randall.

Dorsal-fin rays 9; anal-fin rays 9; pectoral-fin rays 16. Body light brown with gray lines onhead and light bluish gray spots on posterior part of body; two vertical dark brown bars between eye and dorsal-fin origin separated by white lines; a black spot on dorsal-fin base; finspale. Attains 9 cm TL. Inhabits seaward reef slopes at depths of 20–53 m. Western Pacific fromIndonesia eastward to New Caledonia and northward to Japan.
Canthigaster papua (Bleeker 1848)
Papuan Toby

Canthigaster papua, BPBM 9381, 6.7 cm SL, Palau Islands, photo by J. E. Randall.

Dorsal-fin rays 8–10 (usually 9); anal-fin rays 8–10 (usually 9); pectoral-fin rays 15–18 (usually 17). Body reddish brown with many small blue-white spots and short wavy lines on back;pale blue-edged black ocellus on dorsal-fin base; several blue-white lines on snout; caudal finorange with many small blue-white spots; dorsal, anal, and pectoral fins pale; caudal fin light
reddish brown with blue-white spots. Attains 9 cm TL. Occurs in depths shallower than 35 m
in coral reefs. Eastern Indian Ocean eastward through Indonesia to New Caledonia and northward to Japan.

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K. Matsuura & H. Motomura
Canthigaster punctata Matsuura 1992
Yellow-lined Toby

Genus Canthigaster
Canthigaster punctata, NMMBP 26276, 5.5 cm SL, Dong-gang, southern Taiwan, photo by K. Koeda.

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Dorsal-fin rays 10; anal-fin rays 9–10; pectoral-fin rays 16. Dorsal sides of head and bodylight brownish gray with many blue spots; side and belly white; two parallel longitudinal yellow
lines running from pectoral-fin base to caudal peduncle, joining each other just in front of gill
opening; alternating yellow and light blue lines radiating from eye; dorsal, anal, and pectoral
fins pale; caudal fin light yellow with wavy, light blue vertical lines. Attains 12 cm TL. Type specimens collected at a depth of 92 m. Known only from Mascarene Submarine Ridge, western
Indian Ocean and Taiwan.
Canthigaster rivulata Temminck & Schlegel 1850
Brown-lined Toby

Canthigaster rivulata, KAUM–I. 5676, 10.3 cm SL, Tanega-shima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 9–10 (usually 10); anal-fin rays 9–10 (usually 9); pectoral-fin rays 16–18 (usually 17). Dorsal half of head and body brownish gray with many wavy dark blue lines forming a
vermiculated pattern; ventral half of body white; side of body with 2 parallel, dark longitudinalbands, joining each other just in front of gill opening; a dark blotch at dorsal-fin base; dorsal,
anal, and pectoral fins pale; caudal fin with yellowish brown rays. Attains 20 cm TL. Collected
at depths around 100 m in Hawaii, though commonly found in shallow reefs. Indo-West Pacificfrom East Africa eastward to Western Australia and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea
Canthigaster valentini (Bleeker 1853)
Model Toby

Genus Canthigaster
Canthigaster valentini, KAUM–I. 37661, 5.2 cm SL, Iwo-jima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 9; anal-fin rays 9; pectoral-fin rays 16–17 (usually 17). Body white with 4
blackish saddle-like bars on back; first on top of head connecting eyes, second and third bars
extending ventrally to belly region, and fourth on caudal peduncle; belly white; side of body behind pectoral-fin base with dark yellow spots; interorbital region with several dark yellow longitudinal lines; 5–7 transverse dark yellow lines on snout; dorsal, anal, and pectoral fins pale withyellowish orange base; caudal fin yellowish orange. Attains 11 cm TL. Found in coastal reefs atvarious depths. Indo-Pacific from East Africa eastward through Indonesia to Tuamotu Islands
and northward to Japan.

Pair of Canthigaster valentini, spawning, Yaku-shima Island, Kagoshima Prefecture, photo by S. Harazaki.

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K. Matsuura & H. Motomura

Genus Chelonodontops Smith 1958

Genus Chelonodontops
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Mouth below a horizontal line through dorsal end of gill opening;
nasal organ in the form of a depression with slightly raised margin expanded before and behind into a pair of elongate flaps; 2 lateral lines
on body, upper lateral line joining lower behind anal fin, and lower extending forward above anal fin; no lateral skin fold along ventrolateralpart of body and caudal peduncle. Four species in the world, 1 species
in South China Sea.
Chelonodontops patoca (Hamilton 1822)
Milkspotted Pufferfish

Chelonodontops patoca, KAUM–I. 39848, 4 cm SL, Yoron-jima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Chelonodontops patoca, KAUM–I. 66503, 4.3 cm SL, Tokuno-shima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 9–11 (usually 10); anal-fin rays 8–10 (usually 8); pectoral-fin rays 16–19 (usually 17). Dorsal surface of body covered with spinules from posterior part of interorbital region
to just dorsal to pectoral fin; ventral surface of body covered with spinules from throat to justbefore anus; body brownish gray with many round white spots; 5–6 dark relatively wide bars on
body, first across eye and last on caudal peduncle; a longitudinal yellow line running along ventrolateral corner of body from below mouth to caudal peduncle; dorsal, anal, and pectoral fins
pale; caudal fin dusky yellow. Attains 33 cm TL. Occurs in shallow water at depths of 1–22 m,
frequently found in mangroves at depths of 1–5 m. Tropical eastern Indian Ocean eastward toeastern Australia and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea

Genus Lagocephalus Swainson 1839

Key to Species of Lagocephalus
1a. Dorsal surface of body smooth, not covered with spinules ………….……..…………….. 2
1b. Dorsal surface of body covered with spinules …….…………………....…………...…….. 3

Genus Lagocephalus

Body oblong; a longitudinal skin fold running along ventrolateral partof body; dorsal and anal fins pointed; caudal fin emarginate or truncate;
two lateral lines on head and body, ventral element on skin fold andlateral element on lateral midline of side from above gill opening to caudal-fin base with an anterior extension from below eye to the snout region; nasal organs covered by a small sac with 2 nostrils; ventraolateralside of body silver. Nine species in the world, 7 species in South China
Sea. Although muscles of L. inermis, L. cheesemanii, and L. spadiceus
in seas around Japan and East China Sea are edible, individuals of thesame species are poisonous in the South China Sea.

2a. Lower one-fourth to one-third of pectoral fin white, in contrast to the remaining black
upper section; lower caudal-fin lobe longer than the upper …………..… L. lagocephalus
2b. Lower section of pectoral fin not white; caudal fin truncate, slightly rounded or slightly
lunate; upper and lower lobes of caudal fin equal in length ………....…..……… L. inermis
3a. Body oblong; caudal peduncle depressed, wider than deep …………………………….. 43b. Body not oblong; caudal peduncle laterally compressed, deeper than wide …………. 5
4a. Dorsal surface of head and body brown with many black spots; dorsal-fin rays 12–13(usually 12), anal-fin rays 10–11 (usually 11), pectoral-fin rays 16–17 (usually 17) …......
……………….…………………..……..............……………………………… L. sceleratus
4b. Dorsal surface of head and body light brown with irregular dark brown markings andshort dark brown lines; dorsal-fin rays 10–12 (usually 11), anal-fin rays 8-10, pectoral-fin
rays 14–16 (usually 15) ………......................………………………………… L. suezensis
5a. Middle part of caudal fin slightly produced posteriorly; caudal fin black with upper andlower white tips …………….......…………………………………………….. L. cheesemanii
5b. Caudal fin slightly lunate or truncate; caudal fin not black, dusky yellow or light brown .. 6
6a. Spinules on dorsal surface reaching to or beyond dorsal-fin origin …………… L. lunaris
6b. Spinules on dorsal surface not reaching to dorsal-fin origin, ending only halfway frominterorbital region to dorsal-fin origin …….......……………………………….. L. spadiceus

Lagocephalus lunaris, juvenile, KAUM–I. 52335, 6.9 cm SL, Malaysia, photo courtesy of KAUM.

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K. Matsuura & H. Motomura
Lagocephalus cheesemanii (Clarke 1897)
Black-tail Pufferfish

Genus Lagocephalus
26

Lagocephalus cheesemanii, FRLM 33430, 15.5 cm SL, Ago Bay, Mie Prefecture, photo by S. Kimura.

Dorsal-fin rays 11–15 (usually 13–14), anal-fin rays 11–14 (usually 12–13), pectoral-fin rays
15–18 (usually 17). Spinules on back in a rhomboidal patch, starting anteriorly between nasalorgan and eye with widest expanse above midpoint between eye and gill opening, tapering
posteriorly and ending above posterior part of pectoral fin; caudal fin double emarginate withmiddle rays produced posteriorly; dorsal half of body dark brown to brownish black; side ofhead silver with deep yellowish tinge; ventral side of body between gill opening and caudal-finbase silver; dorsal and pectoral fins dark; caudal fin dark brown to black with dorsal and ventral
white tips. Attains 40 cm TL. Occurs at depths of 15–200 m. Eastern Indian Ocean eastward
through South China Sea to Japan and southward to eastern Australia and New Zealand. Individuals in South China Sea are toxic.

Lagocephalus inermis (Temminck & Schlegel 1850)
Smooth Pufferfish

Lagocephalus inermis, KAUM–I. 45420, 24.9 cm SL, Uchinoura Bay, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 12–13, anal-fin rays 11–13, pectoral-fin rays 15–17. A large pufferfish, attaining 56 cm. Body smooth without spinules; caudal fin slightly rounded or truncate; body greenish brown or dark brown above; silvery white on side; belly white; gill opening black internally.
Attains 63 cm TL. Occurs at depths of 35–130 m. Indo-West Pacific from East Africa eastward
through Indonesia to eastern Australia and northward to Japan. Individuals in South China Sea
are toxic.

Identification Guide to Pufferfishes of the South China Sea
Lagocephalus lagocephalus (Linnaeus 1758)
Oceanic Pufferfish

Genus Lagocephalus

Lagocephalus lagocephalus, KPM-NI 4372, 23.8 cm SL, Sagami Bay, Kanagawa Prefecture, photo (KPM-NR 53375A) by H. Senou.

27

Lagocephalus lagocephalus, KAUM–I. 77800, 18 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

Lagocephalus lagocephalus, KAUM–I. 9491, 5.3 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 13–15, anal-fin rays 12–13, pectoral-fin rays 13–16. Blackish-gray to dark
bluish-gray above, with transverse darker bands in juveniles; sides silver between level of upper margin of orbits and ventrolateral angles of body; belly white with dark brown or black spots
near pectoral-fin bases continuing onto underside of body; all fins dark, with lower one-third ofpectoral fin white. Attains 70 cm TL. Occurs at depths of 40–400 m. Circumglobal in warm waters.

K. Matsuura & H. Motomura
Lagocephalus lunaris (Bloch & Schneider 1801)
Lunartail Pufferfish

Genus Lagocephalus
Lagocephalus lunaris, KAUM–I. 19713, 28.9 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

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Dorsal-fin rays 16–17, anal-fin rays 11–12, pectoral-fin rays 16–17. Dorsal surface of bodycovered with spinules from posterior edge of nostrils to origin of dorsal fin; caudal fin deeply
concave; dorsal half of head and body brownish gray; silver on sides; belly white; pectoral, dorsal, and anal fins pale; dorsal half of caudal fin yellowish gray and lower half of caudal fin bluishwhite. Attains 30 cm TL. Occurs at depths of 1–150 m. Indo-West Pacific from East Africa eastward through Indonesia to eastern Australia, northward to Japan. Every part of body is strongly
toxic.
Lagocephalus sceleratus (Gmelin 1789)
Silver-cheeked Toadfish

Lagocephalus sceleratus, KAUM–I. 54093, 68.1 cm SL, Tanega-shima Island, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 12–13 (usually 12), anal-fin rays 10–11 (usually 11), pectoral-fin rays 16–17(usually 17). Dark greenish-gray above, with moderately dense array of irregularly scattered
black spots extending from snout to end of caudal peduncle; sides bluish-gray, each with broad
silver stripe extending from corner of mouth, becoming thinner and descending under eye,then broadening considerably and passing through pectoral-fin base, tapering along lateralsurface of body and terminating near posterior end of caudal peduncle; ventral side of bodygrayish-white; dorsal and pectoral fins grayish; caudal fin yellowish-gray; anal fin pale. Attains
97 cm TL. Occurs at depths of 10–180 m. Indo-Pacific from East Africa eastward through Indonesia to Samoa and northward to Japan. Every part of body is strongly toxic.

Identification Guide to Pufferfishes of the South China Sea
Lagocephalus spadiceus (Richardson 1845)
Half-smooth Golden Pufferfish

Genus Lagocephalus
Lagocephalus spadiceus, KAUM–I. 22558, 19.2 cm SL, Kagoshima Bay, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 11–14 (usually 12–13); anal-fin rays 10–12 (usually 11); pectoral-fin rays 1417 (usually 15–16). Dorsal surface of body covered with spinules from interorbital region to
above pectoral fin, but never reaching dorsal-fin origin; caudal fin truncate or slightly concave;dorsal half of head and body brownish gray; silver on sides; belly white; dorsal, anal, and pectoral fins pale; dorsal half of caudal fin yellowish gray and ventral half dusky white. Attains 30
cm TL. Occurs at depths of 4–190 m. Indo-West Pacific from East Africa eastward through Indonesia to eastern Australia. Many individuals in South China Sea are toxic.

29
Lagocephalus suezensis Clark & Gohar 1953
Suez Pufferfish

Lagocephalus suezensis, KAUM–I. 13613, 13.8 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 10–12 (usually 11); anal-fin rays 9–10; pectoral-fin rays 14-16 (usually 15).
Dark brownish-gray above, with many brown dots and spots; side with broad silver stripe extending from corner of mouth, becoming thinner and descending under eye, then broadening
considerably and passing through pectoral-fin base, tapering along lateral surface of body andreaching caudal-fin base; ventral side of body grayish-white; dorsal and pectoral fins grayish;caudal fin yellowish-gray; anal fin pale. Attains at least 18 cm TL but probably reaches morethan 20 cm TL. Occurs at depths of 2–60 m. Because this species has frequently been confused with L. sceleratus, much of the literature cannot be relied upon for distribution records.
The examination of specimens and photographs in publications revealed it to occur in the Indo-West Pacific from East Africa and Red Sea eastward to Australia, and northward to Japan. It
is probably toxic.

K. Matsuura & H. Motomura

Genus Sphoeroides Anonymous 1798

Genus Sphoeroides

Only one lateral line; no raised skin fold along ventrolateral part ofbody and caudal peduncle; nasal organ covered by a small sac with 2
nostrils; caudal fin truncate; anal-fin origin at or behind vertical from lastdorsal-fin ray; vertebrae 17–20. About 20 species in the world (mostspecies in Atlantic and eastern Pacific), 1 species in South China Sea.
Sphoeroides pachygaster (Müller & Troschel 1848)
Blunthead Pufferfish

Sphoeroides pachygaster, KPM-NI 40071, 30.5 cm SL, Suruga Bay, Shizuoka Prefecture, photo (KPM-NR 109141B) by H. Senou.

30

Sphoeroides pachygaster, KAUM–I. 22572, 24 cm SL, East China Sea, photo courtesy of KAUM.

Dorsal-fin rays 8–9; anal-fin rays 8–9; pectoral-fin rays
14–17. Body smooth without spinules; caudal fin truncate;greenish gray on dorsal half of body, white ventrally; several dark spots occasionally on flanks. Fins pale but caudal
fin dusky with a ventral white tip. Attains 48 cm. Inhabits
the continental shelf and slope at depths of 25–480 m andfrequently collected below 200 m. Circumglobal in warm waters.

Sphoeroides pachygaster, KAUM–I. 86292, 6.5 cm SL,
East China Sea, photo courtesy of KAUM.

Identification Guide to Pufferfishes of the South China Sea

Genus Takifugu Abe 1949

Key to Species of Takifugu
1a. Many alternating brown and white vertical bands on head and body, beginning fromsnout and ending on caudal peduncle ……………….……………...…………. T. oblongus
1b. No vertical bands on head and body ……………..……………………………………..…. 2

Genus Takifugu

Body oblong; a longitudinal skin fold running along ventrolateralpart of body; caudal fin rounded or truncate; nasal organs covered bya small sac with 2 nostrils; two lateral lines, lower along ventrolateralpart of caudal peduncle on skin fold; caudal fin truncate to rounded; nasal organs covered by a small sac with 2 nostrils. About 24 species in
the world; most species in seas around China and Japan; 5 species inSouth China Sea.

2a. Dorsal side of body light gray covered with 7–8 oblique dark blue bands; a large black
blotch behind gill opening ………………………….......………………….… T. bimaculatus
2b. No oblique dark bands on head and body ……………………………....…………………. 3
3a. Dorsal side of head and body light brown covered with many rounded white spots; nolarge black blotch on dorsal-fin base ……………….....……………………… T. flavipterus
3b. Dorsal side of body dark brown or greenish brown without many white spots; large black
blotch sometimes on dorsal-fin base ……………………………......………………………. 4
4a. Dorsal and ventral surfaces of body covered continuously with spinules, not interrupted
by naked space in front of gill opening; dorsal side of head and body dark brown; a large
rounded black blotch behind gill opening and another large rounded black blotch on
dorsal-fin base …………………………………………………………....……… T. obscurus
4b. Dorsal and ventral spinule covered area separated by naked area in front of gill opening;dorsal side of body greenish brown; a wide transverse black band edged with an orange
line on dorsal side of body just behind gill opening; another large black blotch edged
with an orange line on dorsal-fin base ……………………...………....…….… T. ocellatus

Takifugu bimaculatus (Richardson 1845)
Two-blotch Pufferfish

Takifugu bimaculatus, 12 cm SL, Hong Kong, photo by J. Mann.

Dorsal-fin rays 13–14; anal-fin rays 12–13; pectoral-fin rays 15–18. Dorsal surface of body covered byspinules from interorbital region to area in front of dorsal-fin origin; ventral surface of body also covered byspinules; dorsal and ventral spinule covered areas separated by naked space on sides; dorsal side of body
above ventral edge of gill opening light gray, covered with 7–8 oblique blackish blue bands; a large rounded
black blotch behind gill opening; a small black blotch on pectoral-fin base; ventral side of body white; dorsal,
anal, and pectoral fins dusky; caudal fin dusky yellow. Attains 36 cm TL. Collected on continental shelves but
no depth records available. From northern Vietnam northward through East China Sea to Japan.

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K. Matsuura & H. Motomura
Takifugu flavipterus Matsuura 2017
Fine-patterned Pufferfish

Genus Takifugu
Takifugu flavipterus, NSMT-P 124758, 10.4 cm SL, Yamaguchi Prefecture, Sea of Japan, photo courtesy of NSMT.

32

Dorsal-fin rays 11–15; anal-fin rays 10–13; pectoral-fin rays 14–17. Dorsal and ventral surfaces of body covered with spinules, spinules continuous across side in front of gill opening;dorsal side of head and body light brown, covered with many rounded white spots; a longitudinal yellow stripe running along fin fold from chin to caudal-fin base; ventral side of body white;dorsal, anal, and pectoral fins light yellow; caudal fin dusky yellow. Attains 25 cm TL. Occurs incoastal waters shallower than 100 m. Widely distributed along eastern coasts of Asian Continent from northern Vietnam northward through Korea and Japan to southern Sakhalin Island.
Takifugu oblongus (Bloch 1786)
Lattice Pufferfish

Takifugu oblongus, KAUM–I. 27758, 22.8 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 12–14; anal-fin rays 10–12; pectoral-fin rays 14–16. Dorsal and ventral surfaces of body covered with spinules, spinules continuous across side in front of gill opening;many alternating transverse brown and white bands on head and body, beginning on snout andending on caudal peduncle; ventral side of head and body white; dorsal and pectoral fins dusky
yellow; anal fin white; dorsal three-fourths of caudal fin dusky yellow, ventral one fourth white.
Attains 40 cm TL. Occurs at depths of 1–73 m. Indo-West Pacific from East Africa eastward
through Indonesia to Australia and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea
Takifugu obscurus (Abe 1949)
Obscure Pufferfish

Genus Takifugu
Takifugu obscurus, 30 cm TL, aquarium photograph, courtesy of SMSM.

Dorsal-fin rays 15–19; anal-fin rays 13–16; pectoral-fin rays 16–18. Dorsal and ventral
spinule covered areas separated by naked area on side in front of gill opening; dorsal sideof body dark brown without white spots; a large white-edged black rounded blotch behind gill
opening and another on dorsal-fin base; short irregular white lines frequently observed on
dorsal part of head; a longitudinal wide yellow band running just above fin fold from chin to
caudal-fin base; ventral side of body white. Attains 50 cm TL. Inhabits coastal areas, frequently
entering rivers. Northern South China Sea and East China Sea.
Takifugu ocellatus (Linnaeus 1758)
Ocellated Pufferfish

Takifugu ocellatus, 10 cm TL, aquarium photograph, courtesy of SMSM.

Dorsal-fin rays 13–15; anal-fin rays 12–13; pectoral-fin rays 16–18. Dorsal and ventral
spinule covered areas separated on side by naked area in front of gill opening; dorsal side ofbody greenish brown; a wide orange-edged black transverse band on dorsal side of body justbehind gill opening and another on dorsal-fin base; fins pale. Attains 15 cm TL. Inhabits coastal
areas, frequently entering rivers. From northern Vietnam northward through East China Sea to
Yellow Sea.

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K. Matsuura & H. Motomura

Genus Torquigener Whitley 1930

Genus Torquigener

Two lateral lines, lower on low skin ridge ventrolaterally on caudal
peduncle; anal-fin origin below middle of dorsal fin; caudal fin truncate
to rounded; nasal organs covered by a small sac with 2 nostrils; eyes
adnate dorsally only; chin strongly developed. Twenty species in theworld, 5 species in South China Sea.

Key to Species of Torquigener
1a. A solid, dark brown, longitudinal stripe or distinct longitudinal row of spots or blotches on
mid-laterally on body from pectoral-fin base to caudal-fin base ….............……...…..…... 2
1b. No solid longitudinal stripe nor longitudinal row of spots on side of body …….………… 4
2a. Dorsal-fin rays 9–11 (usually 10); anal-fin rays 8–9 (usually 8); pectoral-fin rays 16–17;
no vertical bands on side of head ………….....………………….…………….… T. florealis
2b. Dorsal-fin rays 8–9; anal-fin rays 7–8; pectoral-fin rays 13–16; several vertical bands onside of head below eye ………………………......…………………………………….…….. 3
3a. Vertical dark yellowish-brown bars on side of head; eye diameter smaller than 24% ofhead length; a solid, dark brown, longitudinal stripe on side of body …....... T. brevipinnis
3b. Vertical dark brown bars on side of head; eye diameter larger than 4.1 times in headlength; a distinct longitudinal row of yellowish-brown spots on side of body …...................
...........………………………………….…..………………...…………… T. hypselogeneion

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4a. Spinules on head and body well developed and distinct; dorsal side of head and body
above level through ventral end of gill opening light brown with many small white spots………………………………..............……………………………………..… T. purcaspinus
4b. Spinules on head and body small; dorsal side of head and body above level throughventral end of gill opening pale yellowish white covered with many small rounded brown
blotches and spots …………………...............……………………..…………… T. gloerfelti

Torquigener brevipinnis (Regan 1902)
Yellow-lined Pufferfish

Torquigener brevipinnis, KAUM–I. 162, 8.3 cm SL, Satsuma Peninsula, Kagoshima Prefecture, photo courtesy of KAUM.

Dorsal-fin rays 8–9; anal-fin rays 7–8; pectoral-fin rays 15–16. Dorsal side of body lightbrown with many round white spots; a solid distinct longitudinal row of yellowish-brown spots(frequently coalesced to form a solid stripe) running from pectoral-fin region to caudal-fin base;side of head white with 5 vertical yellowish-brown bars; ventral side of head and body white;dorsal, anal, and pectoral fins pale; caudal fin membrane pale but rays white with many small
brown spots, forming several vertical rows. Attains 11 cm TL. Inhabits shallow waters at depths
of 2–100 m. Eastern Indian Ocean eastward to New Caledonia and northward to Japan.

Identification Guide to Pufferfishes of the South China Sea
Torquigener florealis (Cope 1871)
Floral Pufferfish

Genus Torquigener
Torquigener florealis, BPBM 15488, 11.6 cm SL, Hawaii, photo by J. E. Randall.

Dorsal-fin rays 9–11 (usually 10); anal-fin rays 8–9; (usually 8); pectoral-fin rays 16–17. A
distinct longitudinal row of brownish yellow spots mid-laterally on side from pectoral-fin baseto caudal-fin base; dorsal side of body above level through ventral end of gill opening covered
with many small, irregularly sized, white spots, delineated by a rosette of smaller brown spots;
about 10 rounded yellowish brown spots just below a longitudinal row of yellowish brown spots;side of head white with many small brown spots; ventral side of body white; dorsal, anal, andpectoral fins pale; caudal fin with several vertical light brown bands. Attains 19 cm TL. Inhabits
shallow waters but collected from depths exceeding 100 m. South China Sea, East China Sea,and Hawaiian Islands.

Torquigener gloerfelti Hardy 1984
Gloerfelt's Pufferfish

Torquigener gloerfelti, KAUM–I. 17043, 17.2 cm SL, Malaysia, photo courtesy of KAUM.

Dorsal-fin rays 8–9; anal-fin rays 6–7; pectoral-fin rays 15–16. Dorsal side of head and body
above level through ventral end of gill opening pale yellowish white covered with many smallrounded brown blotches and spots; dorsal, anal, and pectoral fins pale; caudal-fin rays yellowish with dark brown margins. Attains 20 cm TL. Usually collected at depths of 50–60 m. JavaSea and South China Sea.

35

K. Matsuura & H. Motomura
Torquigener hypselogeneion (Bleeker 1852)
Dark-striped Pufferfish

Genus Torquigener

Torquigener hypselogeneion, 7.5 cm SL, KPM-NI 2468, Okinawa-jima Island, Ryukyu Islands, photo (KPM-NR 9777A) by K. Hanazaki (©).

Dorsal-fin rays 8–9; anal-fin rays 7–8; pectoral-fin rays 13–15. Dark brown reticulations on
dorsal surface of body; a solid, dark brown curved stripe on side of body angled from above gill
opening to mid-lateral center of body and continuing horizontally to caudal-fin base; many small
dark brown spots on cheek, forming broad vertical bands, separated by narrow, irregular whitebands; belly white. Attains 10 cm TL. Occurs at depths shallower than 22 m. Indo-West Pacific,
from East Africa eastward through Indonesia to Samoa and northward to Japan.

36Torquigener parcuspinus Hardy 1983
Yellow-eyed Toadfish

Torquigener parcuspinus, CSIRO H 8267-06, 7.5 cm SL, Western Australia, photo courtesy of CSIRO.

Dorsal-fin rays 9; anal-fin rays 6; pectoral-fin rays 16. Spinules on head and body well developed and distinct; dorsal side of head body above level through dorsal end of gill opening lightbrown with many small white spots; no markings on side of head; ventral side of head and bodywhite; fins pale. Attains 10 cm TL. Collected at depths shallower than 400 m. Tropical eastern
Indian Ocean eastward to western Australia and northward to Philippines.

Identification Guide to Pufferfishes of the South China Sea

Genus Tylerius Hardy 1984

Tylerius spinosissimus (Regan 1908)
Fine-spined Pufferfish

Genus Tylerius

Chin not prominent; well-developed spinules covering head andbody except for posterior part of caudal peduncle and fin bases; nasalorgan a short flattened papilla with two openings; no ventrolateral skinfold; body width at pectoral-fin base greater than body depth at end of
dorsal-fin base; lips papillose. Monotypic.

37
Tylerius spinosissimus, NMMB-P 28533, 7.5 cm SL, Dong-gang, southern Taiwan, photo by K. Koeda.

Dorsal-fin rays 8–9; anal-fin
rays 6–7; pectoral-fin rays 14–15.
Body short; body width greaterthan depth; caudal fin longer than
distance between anal-fin origin
and caudal-fin base; dorsal part ofbody above level through ventraledge of pectoral-fin base yellowish-brown with many white spots;
fins pale; posterior margin of
caudal fin whitish with dark brown
posterior margin; belly white,speckled with black. Attains 12 cm
TL. Inhabits shallow waters, frequently collected from depths lessthan 100 m. Indo-West Pacific,
from East Africa eastward throughSouth China Sea to western Australia and northward to Ryukyu
Islands.

Tylerius spinosissimus, 1 cm TL, Suruga Bay, Shizuoka Prefecture, Japan,
underwater photograph by I. Takaku.

K. Matsuura & H. Motomura

Acknowledgments

Acknowledgments
38

We are grateful to the following persons and institutions for facilities provided during our field surveys in Southeast Asia: Aziz Arshad(Universiti of Putra Malaysia, Selangor), Mazlan Abd Ghaffar, Seah
Ying Giat, Siti TafzilMeriam Sheikh Abdul Kadir, and other colleagues(Universiti Malaysia Terengganu, Terengganu), B. Mabel Manjaji-Matsumoto (Universiti Malaysia Sabah, Kota Kinabalu), Nguyen Van Quan
(Institute of Marine Environment and Resources, Haiphong), Teguh
Peristiwady (Indonesian Institute of Sciences, Bitung), Veera Vilasri(National Science Museum Thailand, Pathumthani), and Ricardo P.
Babaran, Wilfredo L. Campos, Ulysses B. Alama, and other colleagues(University of the Philippines Visayas, Miagao). Our thanks also go to
the following persons for providing us photographs and/or drawings of
pufferfishes: Richard Pyle and late John E. Randall (BPBM), Hiromitsu
Endo (BSKU), John Pogonoski (CSIRO), Yoshiaki Kai (FAKU), RachelAtanacio (FishBase), Seishi Kimura (FRLM), Hisashi Imamura (HUMZ),
Hiroshi Senou (KPM), Yukio Iwatsuki (MUFS), Keita Koeda and Hsuan-Ching Ho (NMMB), Masanori Nakae and Gento Shinohara (NSMT),
Elaine Heemstra (SAIAB), Takayuki Sonoyama (SMSM), Kazutoshi
Uehara (GT Divers Okinoerabu), Shigeru Harazaki (Yakushima Diving Service Mori-to-Umi), Itaru Takaku (Yakushima Diving Life), Jason
Mann (Hong Kong), Katsuji Hanazaki, Hirokazu Kishimoto, and Masaki
Wada. Martin F. Gomon of the Museum Victoria kindly provided helpfulcomments on the manuscript. We thank Hiroaki Saito (Atmosphere andOcean Research Institute, the University of Tokyo) and the JSPS Coreto-Core CREPSUM program fish team for opportunity of publishing thisidentification guide. Malaysian specimens were collected during the
JSPS Asian Core and Core-to-Core programs, supported by the Ministry of Higher Education (Government of Malaysia), University Putra
Malaysia, and University Malaysia Terengganu. Vietnamese specimens
were collected with the support of the Institute of Marine Environment
and Resources (Haiphone) and the Ha Long Bay Management Department (Ha Long), with permission for the use of specimens granted by
the Biodiversity Conservation Agency, Ministry of Natural Resources
and Environment (Hanoi). This study was supported in part by JSPS
KAKENHI Grant Numbers 20H03311 and 21H03651; the JSPS Coreto-Core CREPSUM JPJSCCB20200009; and the “Establishment of
Glocal Research and Education Network in the Amami Islands” projectof Kagoshima University adopted by the Ministry of Education, Culture,
Sports, Science and Technology, Japan.

Identification Guide to Pufferfishes of the South China Sea

References

References

Dao VH, Nguyen TD, Nguyen TH, Takata Y, Sato S, Kodama M, Fukuyo
Y (2012) High individual variation in the toxicity of three marine pufferin Vietnam. Coastal Mar Sci, 35:1–6.
Dekkers WJ (1975) Review of the Asiatic freshwater puffers of thegenus Tetraodon Linnaeus, 1758 (Pisces, Tetraodontiformes, Tetraodontidae). Bijdr Dierk, 45:87–142.
Hardy GS (1984) Tylerius, a new generic name for the Indo-Pacific pufferfish, Spheroides spinosissimus Regan, 1908 (Tetraodontiformes:
Tetraodontidae) and comparisons with Amblyrhynchotes (Bibron)
Duméril. Bull Mar Sci, 35:32–37.
Hardy, GS (1988) Unavailability of the name Amblyrhynchotes hypselogeneion rufopunctatus Anon., 1962, and notes on Torquigener gloerfelti (Pisces: Tetraodontidae). Uo, (38):1–2.
Kottelat M (2001) Nomenclatural status of names of tetraodontiform
fishes based on Bibron’s unpublished work. Zoosystema, 23:605–
618.
Matsuura K (2001) Triacanthodidae, Triacanthidae, Balistidae, Ostraciidae, Aracanidae, Triodontidae, Tetraodontidae. In: Carpenter
KE, Niem VH (eds) FAO species identification guide for fishery purposes. The living marine resources of the western central Pacific.
Vol 6. Bony fishes part 4 (Labridae to Latimeriidae). FAO, Rome, pp.
3902–3928, 3948–3957.

39

Arothron stellatus found at a fish market in Malaysia. Photo by K. Matsuura.

K. Matsuura & H. Motomura

Scientific & English name index

Index
40

AmblyrhynchoteAmblyrhynchote honckenii
Amblyrhynchotesamboinensis, Canthigaster
Arothron
Arothron caeruleopunctatus
Arothron carduus
Arothron firmamentum
Arothron hispidus
Arothron immaculatus
Arothron manilensis
Arothron mappa
Arothron meleagris
Arothron multilineatus
Arothron nigropunctatus
Arothron reticularis
Arothron stellatus
axiologus, Canthigaster
Bellyspot Toby
bennetti, Canthigasterbimaculatus, Takifugu
Black-edged Pufferfish
Black-lined Pufferfish
Blackspotted Pufferfish
Black-tail Pufferfish
Blue-spotted Pufferfish
Blunthead Pufferfish
brevipinnis, Torquigener
Brown-lined Toby
caeruleopunctatus, Arothron
Canthigaster
Canthigaster amboinensis
Canthigaster axiologus
Canthigaster bennetti
Canthigaster compressa
Canthigaster epilampra
Canthigaster inframacula
Canthigaster janthinopter
Canthigaster leoparda
Canthigaster ocellicincta
Canthigaster papua
Canthigaster punctata
Canthigaster rivulata
Canthigaster valentini
carduus, Arothroncheesemanii, Lagocephalus
Chelonodontops
Chelonodontops patoca
compressa, Canthigaster
Compressed Toby
Dark-striped Pufferfish
epilampra, Canthigaster
Evileye Pufferfish
Exquisite Toby
Fine-patterned Pufferfish
Fine-spined Pufferfish
firmamentum, Arothron
flavipterus, Takifugu
Floral Pufferfish
florealis, Torquigener
gloerfelti, Torquigener
Gloerfelt's Pufferfish
Guineafowl Pufferfish
Half-smooth Golden Pufferfish
hispidus, Arothronhonckenii, Amblyrhynchote
Honeycomb Toby
hypselogeneion, Torquigener
immaculatus, Arothron
inermis, Lagocephalus
inframacula, Canthigasterjanthinopter, Canthigaster
Lagocephalus
Lagocephalus cheesemanii

6, 7
4, 6, 7
7
16, 17
5, 6, 8
8, 9
8, 9
8, 10
8, 10, 158, 111, 8, 11
8, 12
8, 12
8, 13
8, 14
8, 15
6, 8, 15, 38, 39
16, 1716, 1916, 18
4, 31
8, 11
8, 9
8, 14
25, 26
8, 9
30
6, 34
16, 228, 9
5, 16
16, 1716, 1716, 1816, 1816, 1916, 1916, 2016, 2016, 21
16, 21
16, 2216, 2216, 238, 9
25, 265, 6, 24
24
16, 1816, 1834, 3616, 197
16, 1831, 3237
8, 10
6, 31, 32
34, 3534, 3534, 3534, 358, 12
25, 298, 10, 15
4, 6, 716, 2034, 368, 11
25, 2616, 1916, 203, 5, 6, 25
25, 26

Lagocephalus inermis
Lagocephalus lagocephalus
Lagocephalus lunaris
Lagocephalus sceleratus
Lagocephalus spadiceus
Lagocephalus suezensis
lagocephalus, Lagocephalus
Lantern Toby
Lattice Pufferfish
Leopard Toby
leoparda, Canthigasterlunaris, Lagocephalus
Lunartail Pufferfish
manilensis, Arothron
Many-lined Pufferfish
Map Pufferfish
mappa, Arothron
meleagris, Arothron
Milkspotted Pufferfish
Model Toby
multilineatus, Arothron
nigropunctatus, Arothron
oblongus, Takifugu
Obscure Pufferfish
obscurus, Takifugu
Oceanic Pufferfish
Ocellated Pufferfish
ocellatus, Takifugu
ocellicincta, Canthigaster
pachygaster, Sphoeroides
papua, Canthigaster
Papuan Toby
parcuspinus, Torquigener
patoca, Chelonodontops
punctata, Canthigasterreticularis, Arothron
Reticulated Pufferfish
rivulata, Canthigaster
Saddle Tobysceleratus, Lagocephalus
Shy Toby
Silver-cheeked Toadfish
Smooth Pufferfish
spadiceus, Lagocephalus
Sphoeroides
Sphoeroides pachygasterspinosissimus, Tylerius
Spotted Toby
Star Pufferfish
Starry Pufferfish
stellatus, Arothron
Striped Pufferfish
Suez Pufferfish
suezensis, Lagocephalus
Takifugu
Takifugu bimaculatus
Takifugu flavipterus
Takifugu oblongus
Takifugu obscurus
Takifugu ocellatus
Torquigener
Torquigener brevipinnis
Torquigener florealis
Torquigener gloerfelti
Torquigener hypselogeneion
Torquigener parcuspinus
Two-blotch Pufferfish
Tylerius
Tylerius spinosissimus
valentini, CanthigasterWhite-spotted Pufferfish
Yellow-eyed Toadfish
Yellow-lined Pufferfish
Yellow-lined Toby25, 2625, 2725, 2825, 28, 29
25, 2925, 2925, 2716, 1931, 3216, 2016, 2025, 2825, 281, 8, 11
8, 13
8, 12
8, 12
8, 12
24
16, 238, 13
8, 14
31, 3231, 3331, 3325, 2731, 3331, 3316, 2130
16, 21
16, 2134, 3624
16, 228, 15
8, 15
16, 22
16, 1725, 28, 29
16, 21
25, 2825, 2625, 296, 30
30
37
16, 178, 15
8, 10
6, 8, 15, 38, 39
8, 11
25, 2925, 296, 31
4, 31
6, 31, 32
31, 3231, 3331, 333, 6, 346, 34
34, 3534, 352, 34, 36
34, 3631
6, 37
37
16, 238, 10
34, 3634
16, 22

Ichthyological Publications by the Kagoshima University Museum2009 (March)
Motomura, H. (ed.). Fish collection building
and procedures manual. Japanese edition.
Kagoshima University Museum, Kagoshima. 70 pp.

2010 (March)
Motomura, H. and K. Matsuura (eds.). Fishes of Yaku-shima Island - A World Heritage
island in the Osumi Group, Kagoshima Prefecture, southern Japan. National Museumof Nature and Science, Tokyo. viii + 264
pp., 704 figs.

2011 (Nobember)
Matsunuma, M., H. Motomura, K. Matsuura, N. A. M. Shazili and M. A. Ambak (eds.).
Fishes of Terengganu - east coast of Malay
Peninsula, Malaysia. National Museumof Nature and Science, Tokyo, Universiti
Malaysia Terengganu, Terengganu, and Kagoshima University Museum, Kagoshima.
viii + 251 pp., 678 figs.

2013 (February)
Motomura, H. and S. Ishikawa (eds.). Fishcollection building and procedures manual.
English edition. Kagoshima University
Museum, Kagoshima and Research Institutefor Humanity and Nature, Kyoto. 70 pp.

2013 (March)
Motomura, H., S. Dewa, K. Furuta and K.
Matsuura (eds.). Fishes of Iou-jima andTake-shima islands, Mishima, Kagoshima,
Japan. Kagoshima University Museum, Kagoshima and National Museum of Natureand Science, Tsukuba. 390 pp., 883 figs.

2013 (September)
Yoshida, T., H. Motomura, P. Musikasin-thorn and K . Matsuura (eds.). Fishesof northern Gulf of Thailand. National
Museum of Nature and Science, Tsukuba,
Research Institute for Humanity and Nature,
Kyoto, and Kagoshima University Museum,Kagoshima. viii + 239 pp., 621 figs.

2014 (March)
Motomura, H. and K. Matsuura (eds.). Field
guide to fishes of Yoron Island in the middleof the Ryukyu Islands, Japan. Kagoshima
University Museum, Kagoshima andNational Museum of Nature and Science,
Tsukuba. 648 pp., 1808 figs.

2016 (July)
Iwatsubo, H., S. Kato and H. Motomura(eds.). Field guide to fishes of Ei in southern Kyushu, Japan. Kagoshima Museumof Aquatic Biodiversity, Makura-zaki, Kagoshima University Museum, Kagoshima,and Seahorse Ways, Minami-kyushu. 80
pp., 192 figs.

Most publications listed above can be downloaded fromhttp://www.museum.kagoshima-u.ac.jp/staff/motomura/dl.html

Ichthyological Publications by the Kagoshima University Museum2016 (August)Matsunuma, M., Y. Fukui and H. Motomura(eds.). Freshwater fishes of Kagoshima
City, southern Kyushu, Japan. Kagoshima
University Museum, Kagoshima. 86 pp.,
221 figs.

2016 (September)
Koeda, K., Y. Hibino, T. Yoshida, Y. Kimura, R. Miki, T. Kunishima, D. Sasaki, T. Furukawa, M. Sakurai, K. Eguchi, H. Suzuki,
T. Inaba, T. Uejo, S. Tanaka, M. Fujisawa, H.
Wada and T. Uchiyama. Annotated checklist of fishes of Yonaguni-jima island, thewesternmost island in Japan. Kagoshima
University Museum, Kagoshima. v + 120
pp., 523 figs.

2017 (January)
Motomura, H., U. B. Alama, N. Muto, R.
Babaran, and S. Ishikawa (eds.). Commer-cial and bycatch market fishes of Panay
Island, Republic of the Philippines. The
Kagoshima University Museum, Kagoshima, University of the Philippines Visayas,
Iloilo, and Research Institute for Humanity
and Nature, Kyoto. 246 pp., 911 figs.

2017 (February)
Motomura, H. and S. Harazaki. Annotatedchecklist of marine and freshwater fishesof Yaku-shima island in the Osumi Islands,
Kagoshima, southern Japan, with 129 newrecords. Bulletin of the Kagoshima University Museum, 9: 1–183.

2017 (October)
Iwatsubo, H. and H. Motomura (eds.). Field
guide to fishes of Kagoshima Bay in southern Kyushu, Japan. Kagoshima Museum of
Aquatic Biodiversity, Kagoshima and theKagoshima University Museum, Kagoshima. 302 pp., 618 figs.

2017 (December)
Kimura, Y., Y. Hibino, R. Miki, T. Minetoma and K. Koeda (eds.) Field guide tofishes of Kuchinoerabu-jima island in theOsumi Group, Kagoshima, southern Japan.
Kagoshima University Museum, Kagoshima. 200 pp.

2018 (March)
Koeda, K., H. Hata, M. Yamada and H.
Motomura (eds.). Field guide to fishes landed at Uchinoura Fishing Port, Kagoshima,
Japan. Kagoshima University Museum,Kagoshima. 520 pp., 1059 figs.

2018 (April)Motomura, H., K. Hagiwara, H. Senou
and M. Nakae (eds.). Identification guideto fishes of the Amami Islands, Japan. Kagoshima University Museum, Kagoshima,
Yokosuka City Museum, Yokosuka,Kanagawa Prefectural Museum of Natural History,
Odawara, and National Museumof Natureand Science, Tsukuba. 414 pp., 3290 figs.

Most publications listed above can be downloaded fromhttp://www.museum.kagoshima-u.ac.jp/staff/motomura/dl_en.html

Ichthyological Publications by the Kagoshima University Museum2018 (September)
Mochida, I. and H. Motomura. An annotated checklist of marine and freshwater
fishes of Tokunoshima island in the Amami
Islands, Kagoshima, southern Japan, with202 new records. Bulletin of the Kagoshima
University Museum, 10: 1–80.

2019 (February)
Motomura, H., K. Hagiwara, H. Senou and
M. Nakae (eds.). Identification guide tofishes of the Amami Islands in the Ryukyu
Archipelago, Japan. The Minaminippon
Shimbun Kaihatsu Center, Kagoshima. 436
pp., 3370 figs.

2020 (February)
Sonoyama, T., K. Ogimoto, S. Hori, Y.
Uchida and M. Kawano. An annotatedchecklist of marine fishes of the Sea of Japan off Yamaguchi Prefecture, Japan, with74 new records. Bulletin of the Kagoshima
University Museum, 11: 1–152.

2020 (March)
Motomura, H. and K. Uehara. An annotatedchecklist of marine and freshwater fishes of
Okinoerabu Island in the Amami Islands,
Kagoshima, southern Japan, with 361 newrecords. Bulletin of the Kagoshima University Museum, 12: 1–125.

2020 (May)
Motomura, H. List of Japan’s all fish species. Current standard Japanese and scientific names of all fish species recorded fromJapanese waters. The Kagoshima University
Museum, Kagoshima. 560 pp.

2020 (September)
Motomura, H., T. Yamamoto and S. Tagane(eds.). Field guide to wildlife of the Takaono River mouth and its vicinity areas, Izumi
City, northwestern Kagoshima Prefecture,
Japan. The Kagoshima University Museum,Kagoshima. 316 pp.

2020 (November)
Koeda, K., H. Hata, M. Yamada and H.
Motomura (eds.). Fishes from markets inOsumi Peninsula, Kagoshima, Japan. The
Kagoshima University Museum, Kagoshima. 634 pp.

2020 (December)
Fujiwara, K. and H. Motomura. An annotated checklist of marine and freshwater
fishes of Kikai Island in the Amami Islands,
Kagoshima, southern Japan, with 259 newrecords. Bulletin of the Kagoshima University Museum, 14: 1–73.

Most publications listed above can be downloaded fromhttp://www.museum.kagoshima-u.ac.jp/staff/motomura/dl_en.html

Ichthyological Publications by the Kagoshima University Museum

2021 (January)
Motomura, H., S. Kimura, Y. G. Seah, S.
T. Sheikh Abdul Kadir, and M. A. Ghaffar(eds). Reef and shore fishes of Bidong Island, off east coast of Malay Peninsula. The
Kagoshima University Museum, Kagoshima. 80 pp., 224 figs.

2021 (October)Jeong, B. and H. Motomura. An annotatedchecklist of marine and freshwater fishesof five islands of Mishima in the Osumi
Islands, Kagoshima, southern Japan, with
109 new records. Bulletin of the Kagoshima
University Museum 16: 1–116.

2022 (February)
Motomura, H. (ed.). ICHTHY, Natural
History of Fishes of Japan. Vols. 1–12
(2020.10–2021.9). The Kagoshima University Museum, Kagoshima. 614 pp.

2022 (February)
Iwatsubo, H., M. Itou, M. Yamada and H.
Motomura (eds.). Field guide to fishes of
the East China Sea side of Satsuma Peninsula in Kagoshima, southern Kyushu,
Japan. Kagoshima Museum of Aquatic Biodiversity, Kagoshima and the Kagoshima
University Museum, Kagoshima. 329 pp.,1833 figs.

Arothron stellatus (top two individuals) and A. hispidus (bottom) found at a fish market in Malaysia.
Photo taken by K. Matsuura on 23 May 2011.

THE
KAGOSHIMA
UNIVERSITYMUSEUM