Matsuura, 2015

Ichthyol Res (2015) 62:72–113
DOI 10.1007/s10228-014-0444-5
REVIEW FOR IPFC9 SPECIAL ISSUE Ichthyology and Indo-Pacific Fish Conferences
from the 1980s to the 2010s
Taxonomy and systematics of tetraodontiform fishes: a review

focusing primarily on progress in the period from 1980 to 2014
Keiichi Matsuura
Received: 2 October 2014 / Revised: 12 October 2014 / Accepted: 13 October 2014 / Published online: 11 November 2014
Ó The Author(s) 2014. This article is published with open access at Springerlink.com
Abstract When the first Indo-Pacific Fish Conference species in four genera, Balistidae 37 species in 12 genera,
(IPFC1) was held in Sydney in 1981, there were still many Monacanthidae 102 species in 27 genera, Aracanidae 13
problems in the generic- and species-level taxonomy of all species in six genera, Ostraciidae 22 species in five genera,
tetraodontiform families except for the recently reviewed Triodontidae monotypic, Tetraodontidae 184 species in 27
Triacanthodidae and Triacanthidae. The period from IPFC1 genera, Diodontidae 18 species in seven genera, and Mo-
to IPFC9 (1981-2013) was a time of great progress in the lidae five species in three genera. Phylogenetic relation-
taxonomy and systematics of the Tetraodontiformes: many ships of the families have been clarified by morphological
review and revisional papers have been published for and molecular analyses and have provided well-supported
various genera and species, with descriptions of many new sister relationships of the families: Triacanthodidae and
taxa occurring mainly on coral reefs and in tropical Triacanthidae, Balistidae and Monacanthidae, and Tetra-
freshwaters; and cladistic analyses of morphological char- odontidae and Diodontidae. However, there remain prob-
acters have been performed to clarify phylogenetic rela- lems with the phylogenetic positions of the Triodontidae
tionships of various families and molecular analyses have and Molidae due to conflicts of differing positions in
greatly progressed to provide detailed phylogenetic rela- morphological and molecular studies (e.g., Molidae has
tionships of families, genera, and even species. The pur- been placed differently among molecular studies).
pose of this paper is to provide a review on developments
in the taxonomy and systematics of the Tetraodontiformes, Keywords Tetraodontiformes Classification
focusing primarily on contributions since 1980 (when Morphology Molecular Phylogenetic relationships
James C. Tyler’s monumental work was published)
through the period of IPFCs, including pertinent publica-
tions before 1980. This paper recognizes 412 extant species Introduction
in the 10 families of living Tetraodontiformes, with the
allocation of species and genera as follows: Triacanthodi- Tetraodontiform fishes are distributed in tropical to tem-
dae including 23 species in 11 genera, Triacanthidae seven perate seas and freshwaters of the world. They show a
remarkable diversity in shape, size, and way of life
(Fig. 1). A small filefish of the genus Rudarius Jordan and
This article was registered in the Official Register of Zoological Fowler 1902 and a small pufferfish of the genus Carino-
Nomenclature (ZooBank) as C37A57BA-30FF-48AD-8AEC- tetraodon Benl 1957 mature at about 2 cm in total length
14C084BBB4BA.
This article was published as an Online First article on the online
(TL) (Lim and Kottelat 1995; Tyler 1970), whereas ocean
publication date shown on this page. The article should be cited by sunfishes of the genus Mola Koelreuter 1766 attain over
using the doi number. 300 cm TL (Yoshita et al. 2009). Tetraodontiform fishes
are characterized by a small mouth with either relatively
K. Matsuura (&)
few teeth that are often enlarged or massive beak-like tooth
Division of Fishes, National Museum of Nature and Science,
4-1-1 Amakuobo, Tsukuba, Ibaraki 305-0005, Japan plates, a small gill opening restricted to the side of the
e-mail: [email protected] body, scales usually modified as spines, enlarged plates, or
123
Taxonomy and systematics of tetraodontiforms 73
123
74 K. Matsuura
b Fig. 1 Representatives of the 10 extant families of Tetraodontifor- systematics of the Tetraodontiformes, focusing primarily
mes. a Triacanthodidae, Triacanthodes anomalus; b Triacanthidae, on contributions from 1980 (when James C. Tyler’s
Triacanthus biaculeatus; c Balistidae, Abalistes filamentosus; d Mon-
acanthidae, Thamnaconus hypargyreus; e Aracanidae, Kentrocapros monumental work was published) through the period of
aculeatus; f Ostraciidae, Ostracion immaculatus; g Triodontidae IPFCs, but including pertinent publications before 1980.
Triodon macropterus; h Tetraodonitidae, Arothron mappa; i Diodonti- This paper is composed of two parts, the first reviewing
dae, Diodon liturosus; j Molidae, Masturus lanceolatus. Photographs taxonomic studies and the second focusing of studies on
of a and e provided by BSKU; b, d, f, h, and i by KAUM; c and g by
NSMT; j by Hideki Sugiyama systematics. Institutional abbreviations follow Fricke and
Eschmeyer (2014).
a carapace, and pelvic fins that are reduced or absent (Tyler

1980; Nelson 2006). In addition, many pufferfishes are Triacanthodidae (Spikefishes, Fig. 1a; Table 1)
characterized by having a strong toxin in the viscera and
skin, and even in the musculature of some species of Spikefishes of the family Triacanthodidae are usually
Lagocephalus Swainson 1839 (Matsuura 1984). Because of found in depths of 100-600 m on continental shelves and
their peculiar morphological characters, tetraodontiforms slopes (Tyler 1968; Matsuura and Tyler 1997). They are
have long attracted the attention of ichthyologists and easily distinguished externally from other families of the
biologists. order Tetraodontiformes by the following combination of
Since Cuvier (1816) classified tetraodontiforms in the characters: body deep and slightly compressed, covered by
order Plectognathi, tetraodontiforms were usually consid- moderately thick skin with numerous small scales not
ered to form a monophyletic group among the advanced readily distinguishable to the unaided eye, with each scale
percomorph fishes (Nelson 2006). Although the taxonomy bearing upright spinules and having a roughly shagreen-
of tetraodontiform fishes progressed greatly in the nine- like appearance; two separate dorsal fins, six spines in the
teenth century (see Tyler 1980, for a history of the clas- first dorsal fin and 12-18 soft rays in the second dorsal fin;
sification of the order), information about tetraodontiform caudal fin rounded to almost truncate, not forked; most
taxonomy was scattered in many articles, making it diffi- dorsal-, anal-, and pectoral-fin rays branched; pelvic fins
cult to understand the taxonomic relationships of tetra- with a large spine and one or two inconspicuous and
odontiforms overall. Alec Fraser-Brunner published an rudimentary soft rays; mouth small and usually terminal;
important series of articles on various groups of tetra- teeth of moderate size, usually conical, 10 or more in an
odontiforms as reviews of genera and families during the outer series in each jaw; caudal peduncle compressed,
mid-1930s to early 1950s: however, many parts of his deeper than wide, not distinctly tapered (Matsuura 2001).
publications were based on cursory examinations of rela- Spikefishes are still relatively poorly known in terms of
tively few specimens. There were no comprehensive phy- taxonomy and biology among tetraodontiform families,
logenetic studies until Winterbottom (1974) and Tyler although Tyler (1968) published an excellent monograph
(1980) provided their interpretations of the phylogenetic on the superfamily Triacanthoidea including the Triacan-
relationships of tetraodontiforms based on myology and thodidae and Triacanthidae. He recognized 19 triacantho-
osteology, respectively. did species in 11 genera (Table 1): Atrophacanthus Fraser-
When the first Indo-Pacific Fish Conference (IPFC1) Brunner 1950 (one species), Bathyphylax Myers 1934 (two
was held in Sydney in 1981, many problems remained in species), Halimochirurgus Alcock 1899 (two species),
the generic- and species-level taxonomy of all tetra- Hollardia Poey 1861 (three species), Johnsonina Myers
odontiform families, except for the Triacanthodidae and 1934 (one species), Macrorhamphosodes Fowler 1934 (two
Triacanthidae for which Tyler (1968) had provided a species), Mephisto Tyler 1966b (one species), Parahol-
monographic revision. The period from IPFC1 to IPFC9 lardia Fraser-Brunner 1941b (two species), Paratri-
(1981-2013) was a time of great progress in the taxon- acanthodes Fowler 1934 (two species), Triacanthodes
omy and systematics of tetraodontiforms. Many review Bleeker 1857 (two species), and Tydemania Weber 1913
and revisional papers have been published for various (one species). Because all these genera and species were
genera and species, with descriptions of new taxa found described in detail by Tyler (1968), their taxonomic fea-
mainly in coral reefs and tropical freshwaters; and cla- tures are not repeated here, except for new information that
distic analyses of morphological characters have clarified provides a better understanding of taxa.
phylogenetic relationships of a number of families and Many species of the Triacanthodidae are known from
molecular analyses greatly assisted our understanding of the tropical and warm regions of the Indo-Pacific. How-
the detailed phylogenetic relationships of families, genera, ever, five species are distributed in the western Atlantic:
and even species. The purpose of this paper is to present a Hollardia hollardi Poey 1861, Hollardia meadi Tyler
review of the developments in the taxonomy and 1966a, Johnsonina eriomma Myers 1934, Parahollardia
123
Table 1 A list of 23 species of the Triacanthodidae in the world

Subfamily Species Distribution
Hollardiinae Hollardia goslinei Tyler 1968 Hawaii

Hollardia hollardi Poey 1861 Bermuda and Florida Keys southward to Gulf of Mexico and Caribbean Sea
Hollardia meadi Tyler 1966a Bahamas, Cuba, and Barbados
Paraholardia lineata (Longley 1935) Atlantic coast of the USA from Virginia to Florida; Gulf of Mexico from Florida to
Yucatan
Paraholardia schmidti Woods 1959 Honduras to Panama in the western Caribbean Sea
Triacanthodinae Atrophacanthus japonicus (Kamohara Off Tanzania, Maldive Islands, Japan, and Celebes Sea
1941)
Bathyphylax bombfrons Myers 1934 Off Kenya, China Sea (off Hong Kong), Australia, and New Caledonia
Bathyphylax omen Tyler 1966c Off Kenya
Bathyphylax pruvosti Santini 2006 Marquesas Islands
Halimochirurgus alcocki Weber 1913 Arabian Sea, Japan, Mariana Islands, Australia, and New Caledonia
Halimochirurgus centriscoides Alcock Arabian Sea and Australia
1899
Johnsonia eriomma Myers 1934 Bahamas to Greater and Lesser Antilles and western Caribbean Sea
Macrorhamphosodes platycheilus Bay of Bengal, Philippines, and Australia
Fowler 1934
Macrorhamphosodes uradoi (Kamohara South Africa, off Kenya, Japan, Korea, East China Sea, South China Sea,
1933) Australia, New Caledonia, and New Zealand
Mephisto fraserbrunneri Tyler 1966b Somalia, Arabian Sea, and Andaman Sea
Paratriacanthodes abei Tyler 1997 South China Sea
Paratriacanthodes herrei Myers 1934 Philippines
Paratriacanthodes retrospinis Fowler East Africa, Japan, Kyushu-Palau Ridge, South China Sea, Australia, and New
1934 Caledonia
Triacanthodes anomalus (Temminck Japan, Korea, East China Sea, South China Sea
and Schlegel 1850)
Triacanthodes ethiops Alcock 1894 East Africa, Maldive Islands, Japan, East China Sea, Philippines, Indonesia, New
Caledonia, and Australia
Triacanthodes indicus Matsuura 1982 Saya de Malha Bank (western Indian Ocean)
Triacanthodes intermedius Matsuura New Caledonia
and Fourmanoir 1984
Tydemania navigatoris Weber 1913 East Africa, Bay of Bengal, Japan, China, Philippines, Indonesia, and Australia
lineata (Longley 1935), and Parahollardia schmidti Massachusetts, whereas this species had previously only
Woods 1959. Hollardia goslinei Tyler 1968 are known been known from the Florida Keys, Bahamas, Bermuda,
only from nine specimens collected from the Hawaiian Gulf of Mexico, Caribbean, and south to Brazil, with the
Islands. new northern occurence perhaps associated with warming
Little information about the Atlantic and Hawaiian currents along the east coast of North America.
spikefishes has been published since Tyler’s (1968) In contrast to the Atlantic spikefishes, many papers on
monograph. Matsuura (1983) provided a brief morpho- Indo-Pacific spikefishes were published after 1980.
logical account of Hollardia hollardi with a color photo- Matsuura (1982) described Triacanthodes indicus based on
graph based on specimens collected from off Surinam. 13 specimens collected from the western Indian Ocean.
McEachran and Fechhelm (2005) also provided a brief This species is characterized by relatively large nasal
account and line drawing of Hollardia hollardi and Hol- organs compared to other species of Triacanthodes.
lardia meadi from the Gulf of Mexico. McEachran and Matsuura and Fourmanoir (1984) described Triacanthodes
Fechhelm (2005) and Hartel et al. (2008) reported briefly intermedius based on two specimens collected from New
Parahollardia lineata from the Gulf of Mexico and off Caledonia. Triacanthodes intermedius is unique among
greater New England, respectively. Tyler et al. (2013) spikefishes in having several intermediate characters
documented a large northern range extension for Hollardia between Paratriacanthodes and Triacanthodes. Tyler
hollardi to the northeast coast of the USA off (1997) described Paratriacanthodes abei based on a single
123
76 K. Matsuura
specimen collected in the South China Sea. Santini (2006) South Wales and Western Australia), Triacanthodes ethi-
described Bathyphylax pruvosti based on 25 specimens ops (Northern Territory, Queensland, Western Australia),
collected from the Marquesas Islands. In addition to these and Tydemania navigatoris (Western Australia). Shao et al.
contributions, papers and books on taxonomic and zoo- (2008) recorded seven species of spikefishes from off
geographical studies of spikefishes were published by southern Taiwan in the South China Sea: B. bombifrons,
authors from various countries including Australia, China, Halimochirurgus alcocki, Halimochirurgus centriscoides,
Japan, Korea, Russia, Taiwan, Ukraine, and the USA. Macrorhamphosodes uradoi (first record from the South
Amaoka (1982) reported Halimochirurgus alcocki China Sea), Paratriacanthodes retrospinis, Triacanthodes
Weber 1913, Macrorhamphosodes uradoi (Kamohara anomalus, and Tydemania navigatoris. All species records
1933), and Paratriacanthodes retrospinis Fowler 1934 of Shao et al. (2008) were based on specimens deposited at
with detailed descriptions and color photographs of fresh museums and institutions in Taiwan and the USA. Larson
specimens collected from the Kyushu-Palau Ridge. et al. (2013) published an annotated checklist of the fishes
Matsuura (1984) provided brief accounts and color pho- of the Northern Territory, Australia, in which they recorded
tographs of tetraodontiform fishes found in the Japanese Halimochirurgus centriscoides, Macrorhamphosodes
Archipelago, including Atrophacanthus japonicus Kamo- platycheilus, Triacanthodes ethiops, and Tydemania
hara 1941, Halimochirurgus alcocki, Macrorhamphosodes navigatoris.
uradoi, Triacanthodes anomalus (Temminck and Schlegel Tyler (1983) reported four spikefishes from off Kenya in
1850), and Triacanthodes ethiops Alcock 1894. In the the western Indian Ocean: B. bombifrons, Bathyphylax
following year, Matsuura (1985) provided detailed omen Tyler 1966c, Macrorhamphosodes uradoi, and
accounts and a color photograph of Triacanthodes anom- Macrorhamphosodes platycheilus. When Tyler (1968)
alus collected from the Okinawa Trough. Matsuura (1987) compared B. bombifrons and B. omen, the main differences
and Stewart and Clark (1988) recorded Macrorhampho- were that B. bombifrons has a narrower pelvis, a shorter
sodes uradoi from New Zealand, a species previously postorbital length, a less distinctly supraterminal mouth,
known only from Japan (Kamohara 1933; Tyler 1968; and a more concave snout profile. Because the holotype of
Matsuura 1984), Kenya (Tyler 1983) and South Africa B. omen (37.5 mm in standard length, SL) from the western
(Hulley 1972). Matsuura and Tyler (1997) reported Indian Ocean is much smaller than that of B. bombifrons
Bathyphylax bombifrons Myers 1934, Halimochirurgus (77.6 mm SL) from off Hong Kong in the South China Sea,
alcocki, Macrorhamphosodes uradoi, Paratriacanthodes some of these differences were considered to be due in part
retrospinis, Triacanthodes ethiops, and Triacanthodes to the size differences of the holotypes (Tyler 1968). Based
intermedius based on many specimens collected from New on four additional, newly collected specimens of B.
Caledonia. Except for Triacanthodes intermedius, these are bombifrons (80.9-84.0 mm SL) and two specimens of B.
the first records for the species from New Caledonia. Chen omen (67.0-93.5 mm SL), Tyler (1983) concluded that the
et al. (1997) recorded Triacanthodes anomalus from the most reliable character separating the two species is the
South China Sea. Matsuura (2000) compiled a checklist of degree of concavity of the snout. The depth of the snout in
Tetraodontiformes from the South China Sea including the middle of its length is 10.8-13.4 % SL (average 12.0)
eight species of spikefishes: A. japonicus, B. bombifrons, in B. bombifrons and 12.5-15.5 % SL (average 13.9) in B.
Halimochirurgus alcocki, Macrorhamphosodes platychei- omen. Matsuura and Tyler (1997) confirmed this difference
lus Fowler 1934, Paratriacanthodes retrospinis, Tri- in three specimens (86.2-93.0 mm SL) of B. bombifrons
acanthodes anomalus, Triacanthodes ethiops, and collected off New Caledonia.
Tydemania navigatoris Weber 1913. Tyler (1986) provided keys to the genera and species of
Myers and Donaldson (2003) published a checklist of spikefishes known from South and East Africa. He recog-
fishes from the Mariana Islands including a record of nized 11 species in this region: A. japonicus, B. bombi-
Halimochirurgus alcocki. Kim et al. (2005) provided brief frons, B. omen, Halimochirurgus alcocki, Halimochirurgus
accounts and color photographs of Triacanthodes anoma- centriscoides, Macrorhamphosodes platycheilus, Mac-
lus and Macrorhamphosodes uradoi from Korea. Hoese rorhamphosodes uradoi, Mephisto fraserbrunneri, Para-
et al. (2006) recorded eight species of spikefishes for the triacanthodes retrospinis, Triacanthodes ethiops, and
first time from Australia: B. bombifrons (New South Wales Tydemania navigatoris. Adam et al. (1998) recorded A.
and Queensland), Halimochirurgus alcocki (New South japonicus and Triacanthodes ethiops for the first time from
Wales, Queensland and Western Australia), Hali- the Maldive Islands in the central Indian Ocean. Atroph-
mochirurgus centriscoides (Western Australia), Macrorh- acanthus japonicus had previously been recorded only
amphosodes platycheilus (Northern Territory and Western from southern Japan, the Celebes Sea, and off Tanzania
Australia), Macrorhamphosodes uradoi (New South Wales (Kamohara 1941; Fraser-Brunner 1950; Tyler 1968;
and Queensland), Paratriacanthodes retrospinis (New Matsuura 1984). Although T. ethiops is relatively common
123
Table 2 A list of 7 species of the Triacanthidae in the world

Species Distribution
Pseudotriacanthus strigilifer (Cantor 1849) Persian Gulf, Gulf of Oman, Arabian Sea, Bay of Bengal, Philippines, South China
Sea including Gulf of Thailand, Indonesia, and northern Australia
Triacanthus biaculeatus (Bloch 1786) Persian Gulf, Gulf of Oman, Arabian Sea, Bay of Bengal, and Japan (north to central Honshu),
China, South China Sea including Gulf of Thailand, Indonesia, northern Australia
Triacanthus nieuhofii Bleeker 1852b Arabian Sea, Bay of Bengal, Andaman Sea, Indonesia, South China Sea, northern Australia
Tripodichthys angustifrons (Hollard 1854) Indonesia and northern Australia
Tripodichthys blochii (Bleeker 1852b) Arabian Sea, Japan (stray), China (stray), South China Sea including Gulf of Thailand, and Indonesia
Tripodichthys oxycephalus (Bleeker 1851c) Bay of Bengal, South China Sea including Gulf of Thailand, and Arafura Sea
Trixiphichthys weberi (Chaudhuri 1910) Bay of Bengal, South China Sea including Gulf of Thailand, and Indonesia
in the West Pacific and New Caledonia (Tyler 1968; agencies have recently implemented a joint survey along
Matsuura and Tyler 1997), it had previously been known the Indian Ocean coast of Sumatra and Java. Their surveys
only from East Africa in the Indian Ocean (Tyler 1968, have resulted in many interesting spikefishes, including
1986). Manilo and Bogorodsky (2003) studied coastal two undescribed species and several rarely collected spe-
fishes of the Arabian Sea based on specimens collected by cies, which will be published elsewhere (Matsuura and
research vessels of Russia and Ukraine from 1967 to 1991. Kawai in preparation).
They also examined specimens collected by the German
RV Meteor, which are deposited at the Zoological Museum
of Hamburg University. The two authors recorded Hali- Triacanthidae (Triplespines, Fig. 1b; Table 2)
mochirurgus centriscoides and Mephisto fraserbrunneri
Tyler 1966b (previously known only from the holotype, Triplespines of the family Triacanthidae are found on
52.2 mm SL from the Andaman Sea and a non-type continental shelves in the Indo-West Pacific, usually just
specimen, 66.0 mm SL from off Somalia in the western below the sea surface to 60 m depth (Tyler 1968). They
Indian Ocean), but did not provide descriptions of the two clearly differ from other families of the order Tetraodonti-
species. The specimens they studied are deposited at the formes in having the following combination of characters:
National Museum of Natural History, National Academy of moderately elongate, strongly compressed body; skin
Sciences, Ukraine, the Zoological Museum of the Russian moderately thick with numerous scales not easily discern-
Academy of Sciences, and the Zoological Museum of ible to the unaided eye, with each scale bearing upright
Moscow State University. In addition to these, Venkatar- spinules and having a rough, shagreen-like appearance; two
amanujam et al. (1993) erroneously described a new so- separate dorsal fins, six spines (usually only five spines
lenostomid species, Solenostomus tuticoriensis based on a visible, the sixth rudimentary) in the first dorsal fin and
specimen of Macrorhamphosodes platycheilus from 20-26 soft rays in the second dorsal fin; pelvic fin with a
southern India (see Orr and Fritzsche 1997). large spine and no visible soft rays; mouth small and
This brief historical review clearly shows that there are usually terminal; teeth in jaws with an outer series of about
many collection lacunae with respect to the distribution and 10 heavy incisors and an internal series of several molars,
diversity of spikefishes in the world oceans. New Caledo- usually four in the upper jaw and two in the lower jaw;
nia is one example of how collection efforts could provide caudal fin deeply forked; caudal peduncle distinctly
us with better understanding of spikefish diversity. When tapering to a narrow transversely indented region just in
Matsuura and Fourmanoir (1984) described Triacanthodes front of the caudal-fin base where the peduncle is wider
intermedius from New Caledonia, only two type specimens than deep (Matsuura 2001).
were available for their study. Long-term deep-water sur- In his monograph of the superfamily Triacanthoidea,
veys by ORSTOM around New Caledonia added six spe- Tyler (1968) recognized seven species (Table 2): Pseudo-
cies of spikefishes including 16 specimens of triacanthus strigilifer (Cantor 1849), Triacanthus biacule-
Triacanthodes intermedius to a taxonomic study on tetra- atus (Bloch 1786), Triacanthus nieuhofii Bleeker 1852b,
odontiform fishes of this region by Matsuura and Tyler Tripodichthys blochii (Bleeker 1852a), Tripodichthys an-
(1997). Many regions of the world oceans need to be sur- gustifrons (Hollard 1854), Tripodichthys oxycephalus
veyed, but the most promising areas would be waters in (Bleeker 1851c), and Trixiphichthys weberi (Chaudhuri
French Polynesia and the Indian Ocean coast of Sumatra 1910). He appeared to have resolved all taxonomic prob-
and Java in Indonesia. Japanese and Indonesian fisheries lems in the Triacanthidae, except for two unusual
123
78 K. Matsuura
Table 3 A list of 37 species of the Balistidae in the world

Abalistes filamentosus Matsuura and Yoshino Ryukyu Islands, North West Shelf of Australia, and Timor Sea
2004
Abalistes stellatus (Anonymous 1798) Red Sea, East Africa eastward through northern Australia to Fiji, northward to southern Japan
Balistapus undulates (Park 1797) Red Sea, East Africa eastward through northern Australia to Tuamotu Islands, northward to
southern Japan
Balistes capriscus Gmelin 1789 Tropical western and eastern Atlantic
Balistes polylepis Steindachner 1876 Eastern Pacific from northern California to Chile and Hawaiian Islands
Balistes punctatus Gmelin 1789 Eastern Atlantic from Madeira to Angola
Balistes vetula Linnaeus 1758 Tropical western and eastern Atlantic
Balistoides conspicillum (Bloch and East Africa eastward through northern Australia to Samoa, northward to southern Japan
Schneider 1801)
Balistoides viridescens (Bloch and Scneider Red Sea, East Africa eastward through northern Australia to Line and Tuamotu islands,
1801) northward to southern Japan
Canthidermis macrolepis (Boulenger 1888) Red Sea, Gulf of Oman, and Arabian Sea
Canthidermis maculata (Bloch 1786) Circumgloba in warm and tropical seas
Canthidermis sufflamen (Mitchill 1815) Western Atlantic from Massachusetts through Gulf of Mexico to Lesser Antilles, Bermuda
Melichthys indicus (Randall and Klausewitz Red Sea, East Africa eastward to western Thailand and Sumatra, southern Java, and Bali
1973)
Melichthys niger (Bloch 1786) Circumglobal in tropical seas
Melichthys vidua (Richardson 1845) East Africa eastward through northern Australia to Hawaiian and Tuamotu islands, northward to
southern Japan
Odonus niger (Rüppell 1836) Red Sea, East Africa eastward through northern Australia to Line, Marquesas and Society
islands, northward to southern Japan
Pseudobalistes flavimarginatus (Rüppell Red Sea, East Africa eastward through northern Australia to Tuamotu Islands, northward to
1829) southern Japan
Pseudobalistes fuscus (Bloch and Schneider Red Sea, East Africa eastward through northern Australia to Society Islands, northward to
1801) southern Japan
Pseudobaliste naufragium (Jordan and Starks Eastern Pacific from Baja California to Ecuador
1895)
Rhinecanthus abyssus Matsuura and Shiobara Known only from Kume-jima Island, Ryukyu Islands
1989
Rhinecanthus aculeatus (Linnaeus 1758) East Africa eastward through northern Australia to Hawaiian, Marquesas and Tuamotu islands,
northward to southern Japan
Rhinecanthus assasi (Forsskål 1775) Red Sea and Gulf of Oman
Rhinecanthus cinereus (Bonnaterre 1788) Mauritius and Reunion
Rhinecanthus lunula Randall and Steene Queensland to Pitcairn Islands
1983
Rhiencanthus rectangulus (Bloch and Red Sea, East Africa eastward through northern Australia to Hawaiian and Pitcairn islands,
Schneider 1801) northward to southern Japan
Rhinecanthus verrucosus (Linnaeus 1758) Seychelles, Chagos Archipelago eastward through northern Australia to Vanuatu, northward to
southern Japan
Sufflamen albicaudatum (Rüppell 1829) Red Sea, Gulf of Oman, and Arabian Sea
Sufflamen bursa (Bloch and Schneider 1801) East Africa eastward through northern Australia to Hawaiian, Marquesas and Pitcairn islands,
Sufflamen chrysopterum (Bloch and East Africa eastward through northern Australia to Samoa, northward to southern Japan
Schneider 1801)
Sufflamen fraenatum (Latreille 1804) Red Sea, East Africa eastward through northern Australia to Hawaiian, Marquesas and Tuamotu
islands, northward to southern Japan
Sufflamen verres (Gilbert and Starks 1904) Eastern pacific from Baja California to Ecuador
Xanthichthys auromarginatus (Bennett 1832) Mauritius eastward through northern Australia to Hawaiian and Society islands, northward to
southern Japan
Xanthichthys caeruleolineatus Randall, Angela Islands in western Indian Ocean eastward through Indonesia to Galapagos Islands,
Matsuura and Zama 1978 northward to southern Japan
123
Table 3 continued
Xanthichthys lineopunctatus (Hollard 1854) South Africa eastward through northern Australia to New Guinea, northward to southern Japan
Xanthichthys mento (Jordan and Gilbert Antitropical in Pacific, southern Japan, Hawaiian Islands, Clipperton Island, Revillagigedo
1882) Islands, southern California, Easter Island, and Pitcairn Islands
Xanthichthys ringens (Linnaeus 1758) Western Atlantic from South Carolina through Gulf of Mexico to Lesser Antilles
Xenobalistes punctatus Heemstra and Smith South Africa
1983
Xenobalistes tumidipectoris Matsuura 1981 Mariana Islands
specimens of Triacanthus biaculeatus collected from the distributions (e.g., Matsuura 1984, 2003a, 2009, 2011,
Gulf of Thailand. Tyler (1968) reported that four speci- 2013; Allen and Swainston 1988; Talwar and Jhingran
mens from the Gulf of Thailand near Paknam have dif- 1991; Kottelat et al. 1993; Randall 1995; Allen 1997; Chen
ferences in eye size and the width of the interorbital space. et al. 1997; Fricke 1999; Johnson 1999; Matsuura and
Two of these specimens (111.5 mm SL and 113.9 mm SL) Peristiwady 2000; Hutchins 2001b; Hoese et al. 2006;
have exceptionally large eyes (10.4 % SL) and the other Kottelat 2013; Larson et al 2013). Although most of these
two (105.6 mm SL and 107.9 mm SL) have a normal eye publications did not greatly extend the previously reported
size (7.8 and 8.5 % SL). The two large-eye specimens have distributions of triplespines, Fricke (1999) recorded Tri-
eyes that are also larger than those of any other specimens acanthus biaculeatus from the Mascarene Islands and
of equivalent size. The large-eye specimens have a wider Kottelat et al. (1993) reported a 60-mm SL specimen of
interorbital space (10.2-10.5 % SL vs. 6.3-10.0 % SL in this species from freshwater in Indonesia.
many other specimens). Tyler (1968) stated that they could
be an undescribed species, but tentatively considered them
as abnormal Tiracanthus biaculeatus. Although I have Balistidae (Triggerfishes, Fig. 1c; Table 3)
been to the Gulf of Thailand many times and examined
local collections in Thailand, I have not seen specimens of Triggerfishes of the family Balistidae occur in shallow
Triacanthus with such large eyes. waters, mainly on coral reefs around the world usually
Tyler (1968) provided color descriptions of P. strigilifer, from just below the sea surface to 50 m depth. They differ
Triacanthus biaculeatus, and Trixiphichthys weberi based externally from other families of the order Tetraodonti-
on color notes or color photographs of fresh specimens, but formes by the following combination of characters: body
neither was available for the other species. Matsuura deep, moderately compressed, encased in very thick, tough
(2009) included a color photograph of a juvenile specimen skin with large scales easily discernible as individual units;
(50 mm SL) of Triacanthus nieuhofii collected from Phu- scales above pectoral-fin base in many species enlarged,
ket in the Andaman Sea: body silvery white with light forming a flexible tympanum; mouth small and terminal, or
brown tinge on dorsal half of body; first dorsal spine sil- almost terminal; teeth strong, eight in the outer series of
very white with a black membrane between the first and both jaws; gill opening, a moderately short, vertical to
third dorsal spines; second dorsal, anal, and pectoral fins oblique slit in front of pectoral-fin base; two dorsal fins,
with light yellowish rays; caudal fin dark yellow. Matsuura first dorsal fin with three visible spines, the second spine
(2013) provided brief accounts and a color photograph of more than 1/2 length of first spine; first spine capable of
Tripodichthys blochii collected from the northern Gulf of being locked in an erect position by second spine; second
Thailand: body silvery white with several irregular golden dorsal and anal fin similar in shape; most dorsal-, anal-, and
brownish markings on head and body; basal half of first pectoral-fin rays branched; pelvic fins rudimentary, repre-
dorsal spine silvery white and proximal half black with sented by a series of four pairs of enlarged scales encasing
yellowish orange first dorsal fin membrane; second dorsal, posterior end of pelvis (Matsuura 2001).
anal, and pectoral fins with light yellowish rays; caudal fin Triggerfishes of the Balistidae have not yet been com-
dusky yellow. prehensively reviewed, but Matsuura (1979) described the
Distribution ranges of triplespines were well docu- osteology of family and its generic relationships, and
mented by Tyler (1968). After his monograph appeared, Matsuura (1980) diagnosed and illustrated the 20 species of
many publications on fish faunas, field guides and check- 10 genera found in Japanese waters. His latter paper cov-
lists have provided additional information on triplespine ered many of the 25 species that were known from the
123
80 K. Matsuura
Indo-Pacific before 1980. Prior to this publication two (Bloch and Schneider 1801) and Sufflamen chrysopterum
papers reviewed triggerfishes in the eastern Pacific (Berry (Bloch and Schneider 1801), were included and neither was
and Baldwin 1966) and those of the western Atlantic designated type species. Smith (1949b) subsequently des-
(Moore 1967a). In addition, Randall and Klausewitz (1973) ignated Balistes bursa Bloch and Schneider 1801 as the
reviewed Melichthys Swainson 1839, with a description of type species of the genus Hemibalistes. However, the
Melichthys indicus, and Randall et al. (1978) revised diagnostic character of Hemibalistes, all scales on cheek
Xanthichthys Kaup in Richardson 1856 with a description smaller than those on body, is not adequate to differentiate
of Xanthichthys caeruleolineatus. These were important it from other triggerfish genera because squamation on the
contributions about the taxonomy of triggerfishes up to and cheek of Sufflamen and Hemibalistes is variable (Matsuura
including 1980. A brief historical review of triggerfish 1980). As stated above, the papers of Berry and Baldwin
taxonomy follows. (1966), Moore (1967a), Randall and Klausewitz (1973),
Fraser-Brunner (1935a) published a review of the Bal- Randall et al. (1978), and Matsuura (1980) contributed to
istidae, one of his series of taxonomic studies of tetra- the proper recognition of 11 genera, Abalistes, Balistapus,
odontiform fishes. He recognized 13 genera in the Balistes, Balistoides, Canthidermis, Melichthys, Odonus,
Balistidae and provided a platform for triggerfish taxon- Pseudobalistes, Rhinecanthus, Sufflamen, and Xanthich-
omy. Although de Beaufort and Briggs (1962) lumped thys. Tyler (1980) also accepted these 11 genera in his
Fraser-Brunner’s 11 genera into the single genus Balistes phylogenetic study on tetraodontiform fishes. The oste-
Linnaeus 1758, they relegated all 11 genera to subgenera. ologically based phylogenies of balistoids given by
Berry and Baldwin (1966) reviewed triggerfishes of the Matsuura (1979) and Tyler (1980) were largely in agree-
eastern Pacific with comments on Fraser-Brunner’s clas- ment, as were the majority of their morphological
sification. They pointed out that Fraser-Brunner (1935a) description; the few conflicts between their respective
recognized Nematobalistes Fraser-Brunner 1935a and accounts were clarified by Tyler and Matsuura (1981) as an
Verrunculus Jordan 1924 on trivial differences of squa- aid to subsequent workers.
mation and the anterior few soft rays of the dorsal fin. Matsuura (1981) described Xenobalistes tumidipectoris
Berry and Baldwin (1966) examined many individuals of as a new genus and species based on a small juvenile
the eastern Pacific triggerfishes, which diminished differ- collected from stomach contents of a marlinfish, Makaira
ences among Balistes, Nematobalistes, and Verrunculus. It mazara (Jordan and Snyder 1901), captured in the Mariana
led Berry and Baldwin (1966) to conclude that the latter Islands. Although the holotype of Xenobalistes tumidi-
two are junior synonyms of Balistes. They partially pectoris was partially damaged by digestion, it differs
accepted Fraser-Brunner’s system by recognizing six gen- clearly from other triggerfishes by its possession of a
era: Balistes, Canthidermis Swainson 1839, Melichthys, number of unique characters, such as there is a large lat-
Pseudobalistes Bleeker 1865–1869, Sufflamen Jordan eral expansion protruding laterally just below the pectoral
1916, and Xanthichthys, but they left the other five genera, fin, the supraorbital ridge is well developed and convex
Abalistes Jordan and Seale 1906, Balistapus Tilesius 1820, dorsolaterally, and the mid-lateral portion of the coracoid
Balistoides Fraser-Brunner 1935a, Odonus Gistel 1848, is greatly expanded to form a disk-like bone. Two years
and Rhinecanthus Swainson 1839 to the deliberations of later, Heemstra and Smith (1983) described Xenobalistes
subsequent authors. Just after Berry and Baldwin’s (1966) punctatus as a new species, based on a small juvenile
paper appeared, Moore (1967a) published a review of the washed up on a beach at the mouth of the Van Stadens
triggerfishes of the western Atlantic and recognized four River, South Africa.
genera, Balistes, Canthidermis, Melichthys, and Xanthich- Randall and Steene (1983) described Rhinecanthus
thys (i.e., triggerfishes of other genera not occurring in the lunula and provided brief comments about the other five
western Atlantic). Just after his review of the western species, Rhinecanthus aculeatus (Linnaeus 1758), Rhine-
Atlantic triggerfishes, Moore (1967b) clarified the taxo- canthus assasi (Forsskål 1775), Rhinecanthus cinereus
nomic status of Balistes punctatus Gmelin 1789, stating (Bonnaterre 1788), Rhinecanthus rectangulus (Bloch and
that it occurs only in the eastern Atlantic. Schneider 1801), and Rhinecanthus verrucosus (Linnaeus
Matsuura (1980) provided keys to genera and species, 1758). Matsuura and Shiobara (1989) described Rhine-
and taxonomic accounts of 20 Japanese species that he canthus abyssus on the basis of two specimens collected
classified in 10 genera, Abalistes, Balistapus, Balistoides, off the northeast coast of Kume-jima in the Ryukyu
Canthidermis, Melichthys, Odonus, Pseudobalistes, Rhin- Islands, at depths of 120-150 m, which is quite deep for
ecanthus, Sufflamen, and Xanthichthys. He also addressed a triggerfishes. Matsuura and Yoshino (2004) studied the
taxonomic problem in the genus Hemibalistes. Hemibal- monotypic genus Abalistes and recognized two species,
istes as proposed by Fraser-Brunner (1935a) was not an Abalistes stellatus (Anonymous 1798) and a new species,
available name because two species, Sufflamen bursa Abalistes filamentosus.
123
Taxonomic studies of triggerfishes continued during the Xanthichthys to clarify the relationships of the two species
period from Berry and Baldwin (1966) to Matsuura and of Xenobalistes with the five species of Xanthichthys. This
Yoshino (2004), resulting in the classification shown in study will be published elsewhere to resolve the taxonomic
Table 3, which recognizes 37 species of triggerfishes in 11 status of Xenobalistes. Santini et al. (2013a) also men-
genera. In addition to the above contributions, many tioned that Balistoides and Pseudobalistes are not mono-
checklists and illustrated books on shallow-water fishes phyletic: Balistoides conspicillum (Bloch and Schneider
also provided taxonomic and zoogeographical information 1801) is sister to Melichthys, and Pseudobalistes flavi-
of the Balistidae and other tetraodontiforms (Böhlke and marginatus (Rüppell 1829) forms a clade with Balistoides
Chaplin 1968; Jones and Kumaran 1980; Kyushin et al. viridescens (Bloch and Schneider 1801), whereas Pseudo-
1982; Gloerfelt-Tarp and Kailola 1984; Matsuura 1984, balistes fuscus (Bloch and Schneider 1801) and Pseudo-
2000, 2001, 2002, 2003a, 2009, 2011, 2013; Sainsbury balistes naufragium (Jordan and Starks in Jordan 1895)
et al. 1984; Smith and Heemstra 1986; Tortonese 1986; represent a single clade. These species should be revisited
Allen and Swainston 1988; Winterbottom et al. 1989; with detailed morphological examinations.
Kuiter 1993; Allen and Robertson 1994; Randall 1995,
1996, 2005, 2010; Kyushin et al. 1977; Randall et al. 1997,
2004; Fricke 1999; Myers 1999; Manilo and Bogorodsky Monacanthidae (Filefishes, Fig. 1d; Table 4)
2003; Munday 2005; Shao et al. 2008; Fricke et al. 2009;
Allen and Erdman 2012). Still, taxonomic challenges Filefishes of the family Monacanthidae occur on coral and
remain. Sufflamen verres (Gilbert and Starks 1904) in the rocky reefs and in sea grass beds, shallower than 200 m,
eastern Pacific is said to be different from the Indo-West but some species are found at depths reaching almost
Pacific species, Sufflamen fraenatum (Latreille 1804), in 300 m (Matsuura and Tyler 1997; Hutchins 2001b). They
having higher counts of fin rays of the second dorsal and differ externally from other families of the order Tetra-
anal fins (Berry and Baldwin 1966). According to their odontiformes by the following combination of characters:
study, S. verres has 30-33 dorsal rays and 27-30 anal body usually deep, strongly compressed, body shape
rays, while S. fraenatum has 28-30 dorsal rays and 24-26 varying from oblong to almost circular; skin smooth to
anal rays. However, Berry and Baldwin (1966) also noted rough, shagreen-like, with minute to small scales armed
that a specimen of S. fraenatum from South Africa with 31 with one to many fine spinules, spinules enlarged in some
dorsal rays and 27 anal rays, overlapping the counts of S. species forming bristles or spines on the posterior portion
verres. Matsuura (1980) showed on the basis of 22 speci- of the body; scales on head of some species with strong
mens of S. fraenatum from Japan that dorsal rays range flattened spinules; mouth small, generally terminal, non-
from 27 to 30 and anal rays from 24 to 27. Randall (2010) protractile; teeth pointed and not fused together, central
stated that the Hawaiian population of S. fraenatum has pair usually the largest in each jaw; gill opening, a short
27-31 dorsal rays and 24-28 anal rays. These data vertical to oblique slit in front of, or above, pectoral-fin
strongly suggest that S. verres is a junior synonym of S. base; two dorsal fins, first dorsal fin consisting of a
fraenatum. Molecular comparisons of S. verres and S. prominent spine, which can be locked upright by a second
fraenatum may help clarify their relationships. Fricke very small spine, second dorsal fin with 22 to 52 simple
(1999) stated that Xanthichthys lineopunctatus (Hollard (unbranched) soft rays; anal fin with 20 to 62 simple soft
1854) is a synonym of Xanthichthys lima (Bennett 1832). rays; pelvic fin and spines rudimentary or absent, repre-
However, the original description of Balistes lima by sented by a series of three or fewer pairs of enlarged scales
Bennett (1832) comprised only 50 words with fin ray encasing the pelvic terminus, or segments of indeterminate
counts ‘‘D. 1?, A. 26, P. 13.’’ Because the poor original number, or entirely absent; pelvis usually capable of ver-
description fits several species of the Balistidae, it is tical movement giving rise to a ventral flap (Hutchins
impossible to verify that Balistes lima is conspecific with 2001b; Matsuura 2003b).
Xanthichthys lineopunctatus. As no type specimens are There are no comprehensive reviews of the Monacan-
known for Balistes lima, I conclude that Balistes lima thidae, although in 1988 Barry Hutchins provided taxo-
should be treated as a nomen dubium. nomic, morphological, and phylogenetic data for all
Santini et al. (2013a) showed Xenobalistes tumidipec- monacanthid species in the world in his doctoral disserta-
toris to be deeply nested among five species of Xanthich- tion. Some parts of his dissertation were published, how-
thys in a phylogenetic tree of the Balistidae based on ever, the taxonomic accounts of many species and genera
molecular analysis. Their molecular analysis suggests that and the phylogenetic analysis of the genera remain
Xenobalistes is a junior synonym of Xanthichthys. I am unpublished. Hutchins published a series of articles on the
currently comparing larger specimens of Xenobalistes Monacanthidae between 1977 and 2002. His first article on
tumidipectoris and Xenobalistes punctatus with those of Australian filefishes recognized 28 genera in Australia
123
82 K. Matsuura
Table 4 A list of 102 species of the Monacanthidae in the world

Acanthaluteres brownii (Richardson 1848) Southern Western Australia and South Australia
Acanthaluteres spilomelanurus (Quoy and Southern Western Australia, South Australia, Victoria, Tasmania, and southern New South Wales
Gaimard 1824)
Acanthaluteres vittiger (Castelnau 1873) Southern Western Australia, South Australia, Victoria, Tasmania and southern New South Wales
Acreichtys hajam (Bleeker 1851a) Indonesia to Papua New Guinea, northward to southern Japan
Acreichthys radiatus (Popta 1900) Indonesia to Papua New Guinea, northward to southern Japan
Acreichthys tomentosus (Linnaeus 1758) Sri Lanka eastward thorough Indonesia to New Caledonia, northward to southern Japan
Aluterus heudelotii Hollard 1855 Both sides of Atlantic: in the western Atlantic from Massachusetts and Bermuda to Brazil, in the
eastern Atlantic Mauritania to Angola
Aluterus monoceros (Linnaeus 1758) Circumglobal in warm and tropical seas
Aluterus schoepfii (Walbaum 1792) Nova Scotia and Bermuda to Brazil
Aluterus scriptus (Osbeck 1765) Circumglobal in warm and tropical seas
Amanses scopas (Cuvier 1829) Red Sea, East Africa eastward through northern Australia to Tuamotu Islands, northward to
southern Japan
Anacanthus barbatus Gray 1831 India eastward to northern Australia
Artrolepis filicauda (Günther 1880) Northern Western Australia, Northern Territory, Queensland, New South Wales, eastern
Victoria, northern Tasmania, and southern Papua New Guinea
Brachaluteres fahaqa Clark and Gohar 1953 Red Sea
Brachaluteres jacksonianus (Quoy and Southern Western Australia, South Australia, Victoria, Tasmania, and New South Wales
Gaimard 1824)
Brachaluteres taylori Woods in Schultz et al. Indonesia eastward to Papua New Guinea and eastern Australia, northward to Philippines
1966
Brachaluteres ulvarum Jordan and Fowler Japan
1902
Cantherhines cerinus Randall 2011 Philippines
Cantherhines dumerilii (Hollard 1854) East Africa, Mozambique Channel, Comores and Mascarenes, north to southern Japan,
Ogasawara Islands and Hawaiian Islands, south to New Caledonia, Lord Howe Island, Rapa
and Ducie (Pitcairn Group). East to Central America
Cantherhines fronticinctus (Günther in East Africa eastward through Indonesia to New Caledonia, northward to southern Japan
Playfair and Günther1867)
Cantherhines longicaudus Hutchins and Tahiti, Cook Islands, New Caledonia, and Tuamotu Islands
Randall 1982
Cantherhines macroceros (Hollard in Western Atlantic and St. Paul’s Rocks, Cape Verde Islands
Guichenot 1853)
Cantherhines multilineatus (Tanaka 1918) Eastern Indian Ocean, South China Sea, and Japan
Cantherhines nukuhiva Randall 2011 Marquesas Islands and Tuamotu Archipelago
Cantherhines pardalis (Rüppell 1837) Red Sea, East Africa eastward through northern Australia to Marquesas and Pitcairn islands,
Cantherhines pullus (Ranzani 1842) Massachusetts and Bermuda to Brazil, also recorded from West Africa
Cantherhines rapanui (de Buen 1963) Easter Island
Cantherhines sandwichiensis (Quoy and Hawaiian Islands, Johnston Island, and Cook Islands
Gaimard 1824)
Cantherhines verecundus Jordan 1925 Hawaiian Islands
Cantheschenia grandisquamis Hutchins Western Australia, New South Wales, and Lor Howe Island
1977
Cantheschenia longipinnis (Fraser-Brunner Southern Western Australia and northern New South Wales
1941c)
Chaetodermis penicilligerus (Cuvier 1816) Indonesia east ward to northern Australia, northward to southern Japan
Colurodontis paxmani Hutchins 1977 Northern Australia
Enigmacanthus filamentosus Hutchins 2002 Seychelles and Marshall Islands
Eubalichthys bucephalus (Whitley 1931) Southern Western Australia, South Australia, Tasmania, and southern New South Wales
123
Table 4 continued
Eubalichthys caeruleoguttatus Hutchins Northern Western Australia

1977
Eubalichthys cyanoura Hutchins 1987 Southern Western Australia and South Australia
Eubalichthys gunnii (Günther 1870) Eastern South Australia, Victoria, and Tasmania
Eubalichthys mosaicus (Ramsay and Ogilby Southern Western Australia, South Australia, Victoria, Tasmania, New South Wales, and
1886) southern Queensland
Eubalichthys quadrispinis Hutchins 1977 Southern Western Australia and South Australia
Lalmohania velutina Hutchins 1994b Southeastern India
Meuschenia australis (Donovan 1824) Eastern South Australia, Victoria, and Tasmania
Meuschenia flavolineata Hutchins 1977 Southern Western Australia, South Australia, Victoria, Tasmania, and southern New South Wales
Meuschenia freycineti (Quoy and Gaimard Southern Western Australia, South Australia, Tasmania, and southern New South Wales
1824)
Meuschenia galii (Waite 1905) Southern Western Australia, South Australia, and western Victoria
Meuschenia hippocrepis (Quoy and Gaimard Southern Western Australia, South Australia, Victoria, and northeastern Tasmania
1824)
Meuschenia scaber (Forster in Bloch and Southern Western Australia, South Australia, Tasmania, and southern New South Wales
Schneider1801)
Meuschenia trachylepis (Günther 1870) New South Wales and Queensland
Meuschenia venusta Hutchins 1977 Southern Western Australia, South Australia, Tasmania, and southern New South Wales
Monacanthus chinensis (Osbeck 1765) West Pacific northward to southern Japan and southward to Victoria
Monacanthus ciliates (Mitchill 1818) Newfoundland and Bermuda to the Antilles
Nelusetta ayraud (Quoy and Gaimard 1824) Southern Western Australia, South Australia, Victoria, northern Tasmania, New South Wale, and
southern Queensland
Oxymonacanthus halli Marshall 1952 Red Sea
Oxymonacanthus longirostris (Bloch and East Africa eastward through northern Australia to Samoa, northward to southern Japan
Schneider 1801)
Paraluteres arqat Clark and Gohar 1953 Red Sea
Paraluteres prionurus (Bleeker 1851b) East Africa eastward through Indonesia to Nieu, northward to southern Japan
Paramonacanthus arabicus Hutchins 1997 Arabian Gulf
Paramonacanthus choirocephalus (Bleeker Thailand eastward through Indonesia to Papua New Guinea
1853)
Paramonacanthus frenatus (Peters 1855) East Africa and Seychelles
Paramonacanthuys lowei Hutchins 1997 Eastern Australia
Paramonacanthus matsuurai Hutchins 1997 Ogasawara Islands
Paramonacanthus nematophorus (Günther Red Sea, East Africa, and Seychelles
1870)
Paramonacanthus oblongus (Temminck and Indonesia through northern Australia to New Caledonia, northward to southern Japan
Schlegel 1850)
Paramonacanthus otisensis Whitley 1931 Eastern Australia
Paramonacanthus pusillus (Rüppell 1828) Red Sea, East Africa eastward through Indonesia to northern Australia, northward to southern
Japan
Paramonacanthus sulcatus (Hollard 1854) Malaysia, Indonesia, Thailand, Taiwan, and China
Paramonacanthus tricuspis (Hollard 1854) India eastward to west coast of Thailand
Pervagor alternans (Ogilby 1899) Eastern Australia, Lord Howe Island, New Caledonia, and Marshall Islands
Pervagor aspricaudus (Hollard 1854) Mauritius eastward to Marshall Islands, Hawaiian Islands, and New Caledonia, northward to
southern Japan
Pervagor janthinosoma (Bleeker 1854) East Africa eastward through Indonesia to Fiji and Tonga, northward to southern Japan
Pervagor marginalis Hutchins 1986a Marquesas Islands and Line Islands
Pervagor melanocephalus (Bleeker 1853) Indonesia eastward through northern Australia to Samoa, northward to southern Japan
Pervagor nigrolineatus (Herre 1927) Indonesia eastward through Papua New Guinea to Solomon Islands, northward to southern Japan
Pervagor randalli Hutchins 1986a Red Sea
123
84 K. Matsuura
Table 4 continued
Pervagor spilosoma (Lay and Bennett 1839) Hawaiian Islands

Pseudalutarius nasicornis (Temminck and East Africa eastward through northern Australia to New Caledonia, northward to southern Japan
Schlegel 1850)
Pseudomonacanthus elongatus Fraser- Australia
Brunner 1940
Pseudomonacanthus macruru (Bleeker Singapore, Malaysia, Brunei, Indonesia, Philippines, and Japan
1856)
Pseudomonacanthus peroni (Hollard 1854) Australia and New Guinea
Pseudomonacanthus tweediei Fraser- Singapore
Brunner 1940
Rudarius ercodes Jordan and Fowler 1902 Japan, Korea, and China
Rudarius excelsus Hutchins 1977 Malaysia (Sabah), Papua New Guinea, Queensland, and Ryukyu Islands
Rudarius minutus Tyler 1970 Malaysia (Sabah), Indonesia, Palau, and Great Barrier Reef
Scobinichthys granulatus (Shaw in White Southern Western Australia, South Australia, Victoria, northern Tasmania, New South Wale, and
1790) southern Queensland
Stephanolepis auratus (Castelnau 1861) Eastern Western Australia, South Australia, Victoria, and Tasmania
Stephanolepis cirrhifer (Temminck and Japan, Korea, Taiwan, China, and northern Philippines
Schlegel 1850)
Stephanolepis diaspros Fraser-Brunner 1940 Red Sea, Gulf of Oman, and Persian Gulf
Stephanolepis hispidus (Linnaeus 1766) Both sides of Atlantic: western Atlantic from Nova Scotia and Bermuda to Brazil; eastern
Atlantic from Canary Islands to Angola
Stephanolepis setifer (Bennett 1831) North Carolina and Bermuda to northern South America
Thamnaconus analis (Waite 1904) Southwestern Pacific
Thamnaconus arenaceus (Barnard 1927) Western Indian Ocean
Thamnaconus degeni (Regan 1903) Tasmania, Victoria and South Australia
Thamnaconus fajardoi Smith 1953 Mozambique Channel, Madagascar, western Mascarenes
Thamnaconus fijiensis Hutchins and Fiji
Matsuura 1984
Thamnaconus garretti (Fowler 1928) Hawaiian Islands
Thamnaconus hypargyreus (Cope 1871) Western Pacific and southeastern Indian Ocean
Thamnaconus melanoproctes (Boulenger Muscat, Oman, Gulf of Oman, Arabian Sea, northwestern Indian Ocean
1889)
Thamnaconus modestoides (Barnard 1927) Red Sea, Indo-West Pacific: East Africa, South Africa, Seychelles, Madagascar and Réunion
(Mascarenes) east to Indonesia, north to southern Japan, south to northwestern Australia
Thamnaconus modestus (Günther 1877) Japan, Korea, China
Thamnaconus paschalis (Regan 1913) Easter Island and southeastern Pacific
Thamnaconus septentrionalis (Günther China, East China Sea, and Japan
1874)
Thamnaconus tessellatus (Günther 1880) Western Pacific and southeastern Indian Ocean
(Hutchins 1977) where the greatest diversity of filefishes is Psilocephalus Swainson 1839), Paraluteres Bleeker 1865,
found (Hutchins 2001a). and Pseudalutarius Bleeker 1865 as distinctive genera.
Before Hutchins’ (1977) review of Australian filefishes Berry and Vogele (1961) reviewed nine species of the
appeared, Fraser-Brunner (1941a) published a review paper Monacanthidae of the western North Atlantic with detailed
on the Monacanthidae (his Aluteridae), recognizing 22 accounts of Amanses pullus (Ranzani 1842), Aluterus
genera. Although de Beaufort and Briggs (1962) lumped monoceros (Linnaeus 1758), Aluterus heudelotii Hollard
Amanses Gray 1835, Chaetodermis Swainson 1839, Par- 1855, Aluterus schoepfi (Walbaum 1792), Aluterus scriptus
amonacanthus Bleeker 1865–1869, Pervagor Whitley (Osbeck 1765), Monacanthus ciliatus (Mitchill 1818),
1930a, and Pseudomonacanthus Bleeker 1865 into a single Monacanthus tuckeri Bean 1906, Stephanolepis hispidus
genus, Monacanthus Linnaeus 1758, they recognized (Linnaeus 1758), and Stephanolepis setifer (Bennett 1831).
Aluterus Cloquet 1816, Anacanthus Gray 1830 (their Although Berry and Vogele (1961) placed Monacanthus
123
pullus in the genus Amanses, they also pointed out that M. Based on the contributions shown above, I recognize
pullus should be classified in the subgenus Cantherhines 102 species in 27 genera worldwide (Table 4). I provi-
Swainson 1839 because M. pullus does not have a patch of sionally place Pseudomonacanthus multilineatus Tanaka
long spines on the mid-side of the body between the dorsal 1918 in the genus Cantherhines following previous
and anal fins, which is a diagnostic character of Amanses. authors (Matsuura 1984; Kyushin et al. 1977; Lindberg
When Randall (1964) reviewed the filefish genera Amanses et al. 1997). Although Randall (1964) placed Pseudo-
(monotypic) and Cantherhines, he recognized 11 species of monacanthus multilineatus in Cantherhines, Hutchins and
Cantherhines in the tropical seas of the world. He showed Randall (1982) said that Cantherhines multilineatus
that Amanses possesses a patch of long stout spines (male) should be classified in the genus Thamnaconus Smith
or a dense brushlike mass of long setae (female) on the 1949b. However, they did not provide characters to jus-
mid-side of the body, whereas all species of Cantherhines tify the transfer from Cantherhines to Thamnaconus. To
lack long spines or long setae on the mid-side of the body. resolve the taxonomic status of Cantherhines multilinea-
Randall (1964) described Cantherhines tiki as a new spe- tus, a detailed study of diagnostic characters, including
cies based on a single specimen collected from Easter encasing scales of the posterior end of the pelvis, is
Island. Cantherhines tiki was said to be distinguishable needed. Hutchins suggested in his dissertation that
from other species of Cantherhines by its strongly pro- Thamnaconus melanoproctes (Boulenger 1889) is a senior
duced snout (Randall 1964). However, Caldwell and synonym of Pseudomonacanthus multilineatus. However,
Randall (1967) showed Cantherhines tiki to be a junior an examination of specimens from various localities in
synonym of Cantherhines rapanui (de Buen 1963) because the Indo-West Pacific is necessary to properly access the
of the considerable variation in the snout shape of Can- relationships of the two nominal species. Although Barry
therhines rapanui. Hutchins made great contributions to the better under-
Hutchins (1977) described three new genera, Bigener, standing of taxonomy and diversity of filefishes, in par-
Cantheschenia and Colurodontis, and eight new species ticular Australian filefishes, taxonomic problems in
from Australia. However, Hutchins (1994a) subsequently several genera such as Acreichthys Fraser-Brunner 1941c,
placed Aleuterius brownii Richardson 1848, the type spe- Pseudomonacanthus, and Thamnaconus remain. As in the
cies of Bigener, in Acanthaluteres Bleeker 1865, making case of triggerfishes, many checklists and illustrated
Bigener a junior synonym of Acanthaluteres. Hutchins and guidebooks have provided much useful information on
Randall (1982) reviewed the Cantherhines fronticinctus the taxonomy, distribution, and biology of filefishes (see
complex and described Cantherhines longicaudus as a new the section on the Balistidae).
species from Tahiti and Rarotonga. Matsuura (1984) pro-
vided brief accounts and color photographs of 26 mon-
acanthids found in Japanese waters. Hutchins and Matsuura Aracanidae (Trunkfishes, Fig. 1e; Table 5)
(1984) described Thamnaconus fijiensis from Fiji. In the
following year, Hutchins (1985) reviewed the genus Trunkfishes of the family Aracanidae occur in shallow to
Brachaluteres Bleeker 1865 and recognized four species relatively deep waters in depths from 5 m to 300 m. The
with comments on their ability to inflate the abdomen. species of Aracana Gray 1833, Anoplocapros Kaup 1855,
Hutchins (1986a) published a revision of the genus Per- Caprichthys McCulloch and Waite 1915, Capropygia
vagor and recognized eight species including two new Kaup 1855 and Polyplacapros Fuji and Uyeno 1979 are
species, Pervagor randalli and Pervagor marginalis. found in temperate seas of southern Australia, whereas
Hutchins (1986b) provided a key and brief taxonomic species of Kentrocapros Kaup 1855 are known from deep
accounts for 16 species of filefishes found along South waters in the warm, tropical regions of the Indo-Pacific
Africa. Hutchins (1987) published Eubalichthys cyanoura (Matsuura and Yamakawa 1982; Matsuura and Tyler 1997;
as a new species from South Australia. Hutchins (1994b) Matsuura 2006, 2008).
described and illustrated Lalmohania velutina from India Trunkfishes differ externally from other families of
as a new genus and species. Hutchins (2002) reported Tetraodontiformes by the following combination of char-
another new genus and species, Enigmacanthus filamen- acters: body almost completely encased in a bony shell or
tosus, from the Seychelles and Marshall Islands. Enigm- carapace formed of enlarged, thickened scale plates, usu-
acanthus filamentosus is a very small filefish, 27-36 mm ally hexagonal in shape and firmly sutured to one another;
SL, resembling in size species of the genus Rudarius. isolated bony plates on caudal peduncle; carapace trian-
Randall (2011) again reviewed the genus Cantherhines, gular or hexagonal in cross section, with openings for
recognizing 11 species, including two new species, Can- mouth, eyes, gill slit, pectoral, dorsal, and anal fins, and for
therhines cerinus from the Philippines and Cantherhines the flexible caudal peduncle; scale plates often with surface
nukuhiva from the Marquesas Islands. granulations and prolonged in some species into prominent
123
86 K. Matsuura
Table 5 Distributions of 13 species of the Aracanidae in the world

Anoplocapros amygdaloides Fraser-Brunner 1941d Western Australia and South Australia

Anoplocapros inermis (Fraser-Brunner 1935b) Southern Queensland, New South Wales and Victoria
Anoplocapros lenticularis (Richardson 1841) Western Australia, South Australia and Victoria
Aracana aurtia (Shaw in Shaw and Nodder 1798) Western Australia, South Australia, Victoria, Tasmania, and New South Wales
Aracana ornata (Gray 1838) Western Australia, South Australia, Victoria, and Tasmania
Caprichthys gymnura McCulloch and Waite 1915 Western Australia and South Australia
Capropyguia unistriata (Kaup 1855) Western Australia, South Australia, and Victoria
Kentrocapros aculeatus (Houttuyn 1782) Japan, Korea, China, and Hawaiian Islands
Kentrocapros eco Phillipps 1932 New Zealand
Kentrocapros flavofasciatus (Kamohara 1938) Japan, New South Wales, Queensland, and New Caledonia
Kentrocapros rosapinto (Smith 1949a) Western Indian Ocean
Kentrocapros spilonota (Gilbert 1905) Hawaiian Islands
Polyplacapros tyleri Fujii and Uyeno 1979 New South Wales and Norfolk Ridge
carapace spines over the eyes or along ventrolateral or the trunkfish as ‘‘Many Spined Coffin Fish. Ostracion
dorsolateral angles of the body; mouth small, terminal, (Acerana) auritus’’; however, the legend of plate 98
with fleshy lips; teeth of moderate size, conical, usually spelled the name as ‘‘Ostracion (Acarana) auritus.’’
fewer than 15 in each jaw; gill opening moderately short, a Plate 98 provided a good illustration of the species
vertical to oblique slit in front of pectoral-fin base; spinous showing appropriate characters to identify the species. It is
dorsal fin absent; most dorsal-, anal-, and pectoral-fin rays the type species of the subgenus by monotypy. However,
branched; caudal fin with nine branched rays; pelvic fins in 1838 John E. Gray published another paper in which he
absent (Matsuura 2001). described three species, Ostracion (Aracana) ornata, Os-
The Aracanidae and Ostraciidae have been placed in tracion (Aracana) flavigaster, and Ostracion (Aracana)
separate families by some authors (e.g., Winterbottom lineata. The second species is a junior synonym of Ara-
1974; Tyler 1980; Matsuura 1984, 2001), but they have cana ornata and the third Aracana aurita (Shaw in Shaw
also been treated as two subfamilies of the family Ostrac- and Nodder 1798). Gray (1838) stated for his three species
iidae by others (e.g., Winterbottom and Tyler 1983; Klas- that ‘‘I have formed a subgenus under the name of Ara-
sen 1995; Allen et al. 2006; Nelson 2006; Matsuura 2008). cana’’ indicating his intention to establish the group at the
All the above authors recognized the two groups as closely subgeneric level. Although the name Aracana has long
related phylogenetic clades (sister groups), but placed them been used for the trunkfish genus, the two names Acerana
at different taxonomic ranks. Santini and Tyler (2003) and Acarana were published earlier than Aracana, thus
studied fossils and extant members of the Tetraodontifor- creating a nomenclatural problem. If one follows the
mes extensively and recognized familial groupings for the priority rule of the Code (ICZN 1999), the family name
Aracanidae and Ostraciidae. Recently, Santini et al. should be Aceranidae or Acaranidae. As Aracana has
(2013a) investigated the phylogenetic relationships of the been used almost universally in the literature, with Acer-
Aracanidae and Ostraciidae based on molecular analysis of ana or Acarana virtually ignored by taxonomists, Aracana
many taxa (nine species of five genera of Aracanidae and should be preserved under Article 33.3.1 of the Code,
17 species of six genera of Ostraciidae). They demon- making the spelling of the family group name Aracanidae
strated that the species examined in their studies (Santini preserved under Article 35.4.1.
and Tyler 2003; Santini et al. 2013a) have adequate char- Trunkfishes have not yet been comprehensively
acteristics to recognize two separate groups at the family reviewed. Although Fraser-Brunner (1935b, 1941d) pro-
level. vided brief reviews of the family, his accounts of genera
A problem exists with the spelling of the genus Ara- and species were cursory as they were based on few
cana. When Gray (1833) published his article on Indian specimens and did not provide adequate characteristics for
zoology, he used two different spellings for a species of the species treated. Kuiter (1994) and Matsuura (2008)
trunkfish. At the beginning of the article, a section titled diagnosed and illustrated seven species in four genera of
‘‘Directions for arranging the plates’’ provided the name Australian aracanids, covering many representatives of the
of animals in the plates where he presented the name of family except for the genera Kentrocapros and
123
Fig. 2 Kentrocapros eco

(NMNZ P.903, 102 mm TL,
from Pahia, Bay of Islands, New
Zealand; a lateral view; b dorsal
view) and an undescribed
species of Kentrocapros
(c NSMT-P 43344, 104 mm SL)
collected from New Zealand
Polyplacapros. Polyplacapros tyleri Fujii and Uyeno 1979 (NMNZ P. 903) revealed Matsuura’s (1990) Tasman Sea
is unique in having a fusiform body and the caudal material to be an undescribed species (Fig. 2). This unde-
peduncle nearly completely covered by discrete bony scribed species has a spine on the dorsolateral ridge mid-
plates. In addition to Australian genera of Aracanidae, way between the eye and the dorsal-fin origin, whereas the
detailed descriptions and illustrations of all four known holotype of O. eco lacks this spine. This new species will
species of Kentrocapros were provided by Matsuura and be described and published elsewhere.
Yamakawa (1982), Matsuura and Tyler (1997), and As discussed above, no major taxonomic problems
Matsuura (2006). Matsuura (1990) provided brief accounts remain in the Aracanidae. However, there still remains a
and a color photograph of a species of Kentrocapros col- need to clarify the nomenclature of some species of
lected from the Tasman Sea (32°430 S, 167°300 E) under the Anoplocapros as well as the intraspecific variation of
name Kentrocapros eco (Phillipps 1932). However, an morphological characters related to ontogenetic develop-
examination of the holotype (102 mm TL) of Ostracion eco ment of Kentrocapros.
123
88 K. Matsuura
Table 6 Distributions of 22 species of the Ostraciidae in the world

Acanthostracion guineensis Bleeker 1865 Eastern Atlantic on West Coast of Africa from Guinea to Gabon
Acanthostracion notacanthus Bleeker 1863 Eastern Atlantic from St. Helena, Ascension and Azores islands, Ghana and Angola
Acanthostracion polygonius Poey 1876 Western Atlantic from New Jersey and Bermuda to Brazil
Acanthostracion quadricornis (Linnaeus 1758) Western Atlantic from Massachusetts and Bermuda to Brazil
Lactophrys bicaudalis (Linnaeus 1758) Western Atlantic from Florida to Brazil
Lactophrys trigonus (Linnaeus 1758) Western Atlantic from Massachusetts and Bermuda to Brazil
Lactophrys triqueter (Linnaeus 1758) Western Atlantic from Massachusetts and Bermuda to Brazil
Lactoria cornuta (Linnaeus 1758) Red Sea, East Africa eastward through northern Australia to Marquesas Islands,
Lactoria diaphana (Bloch and Schneider 1801) East Africa eastward through northern Australia to Hawaiian and Marquesas islands,
Lactoria fornasini (Bianconi 1846) East Africa through Indonesia to Easter Island, southward to New South Wales,
Ostracion cubicus Linnaeus 1758 Red Sea, East Africa eastward through northern Australia to Tuamotu Islands (stray
found in Hawaiian Islands), northward to southern Japan
Ostracion cyanurus Rüppell 1828 Red Sea, Gulf of Oman, and Arabian Sea
Ostracion immaculatus Temminck and Schelgel 1850 Japan, Korea, China, and East China Sea
Ostracion meleagris Shaw in Shaw and Nodder 1796 East Africa eastward through Indonesia to Hawaiian and Pitcairn islands, northward to
southern Japan
Ostracion nasus Bloch 1785 India eastward through Indonesia to Papua New Guinea, northward to Philippines
Ostracion rhinorhynchos Bleeker 1851a Sri Lanka eastward through Indonesia to Queensland, northward to southern Japan
Ostracion solorensis Bleeker 1853 Philippines through Indonesia to Great Barrier Reef
Ostracion trachys Randall 1975 Réunion and Mauritius
Ostracion whitleyi Fowler 1931 Hawaiian, Society, Marquesas, and Tuamotu islands
Tetrosomus concatenatus (Bloch 1785) East Africa through Indonesia to New South Wales, northward to southern Japan
Tetrosomus gibbosus (Linnaeus 1758) Red Sea, East Africa eastward through northern Australia to New Caledonia, northward
to southern Japan
Tetrosomus reipublicae (Ogilby 1913) East Africa through Indonesia to New South Wales, northward to southern Japan
Ostraciidae (Boxfishes, Fig. 1f; Table 6) branched; caudal fin with eight branched rays; pelvic fins
absent (Matsuura 2001).
Boxfishes of the family Ostraciidae occur in shallow Fraser-Brunner (1935b) published a brief review of the
tropical and warm seas of the world. Boxfishes differ Aracanidae and Ostraciidae, recognizing six genera in the
externally from other families of Tetraodontiformes by the Ostraciidae: Acanthostracion Bleeker 1865, Lactophrys
following combination of characters: body almost com- Swainson 1839, Lactoria Jordan and Fowler 1902, Ostra-
pletely encased in a bony shell or carapace formed of cion Linnaeus 1758, Rhinesomus Swainson 1839, and
enlarged, thickened scale plates, usually hexagonal in Rhynchostracion Fraser-Brunner 1935b. Boxfishes found
shape and firmly sutured to one another; no isolated bony in the Indo-Australian Archipelago were studied by de
plates on caudal peduncle; carapace triangular, rectangular, Beaufort and Briggs (1962) who lumped Fraser-Brunner’s
or pentangular in cross section, with openings for mouth, (1935b) Indo-Pacific genera into the genus Ostracion.
eyes, gill slits, pectoral, dorsal, and anal fins, and for the However, they recognized five subgenera in Ostracion:
flexible caudal peduncle; scale-plates often with surface Lactoria, Ostracion, Rhynchostracion, Tetrosomus Swain-
granulations, which are prolonged in some species into son 1839, and Triorus Jordan and Hubbs 1925. Triorus is a
prominent carapace spines over the eye or along the ven- junior synonym of Tetrosomus because the type species of
trolateral or dorsal angles of the body; mouth small, ter- the former, Lactophrys tritropis Snyder 1911, is a junior
minal, with fleshy lips; teeth moderate, conical, usually synonym of Tetrosomus concatenatus (Bloch 1786) (see
fewer than 15 in each jaw; gill opening, a moderately short, Randall et al. 1997). Although Ostracion nasus Bloch 1785
vertical to oblique slit in front of pectoral-fin base; spinous and Ostracion rhinorhynchos Bleeker 1851a have been
dorsal fin absent; most dorsal-, anal-, and pectoral-fin rays placed in the genus Rhynchostracion by some authors
123
Böhlke and Chaplin (1968) provided keys and accounts

accompanied by illustrations to boxfishes found in the
western Atlantic. They recognized five species, A. polyg-
onus, A. quadricornis, Lactophrys bicaudalis (Linnaeus
1758), Lactophrys trigonus (Linnaeus 1758), Lactophrys
triqueter (Linnaeus 1758). Lactophrys bicaudalis and L.
triqueter were placed in the genus Rhinesomus by Matsu-
ura (2003b) and McEachran and Fechhelm (2005). How-
ever, Klassen (1995) argued that L. bicaudalis, L. triqueter,
and L. trigonus are nested in a single clade on the basis of
his extensive osteological analysis; I agree with this.
Acanthostracion quadricornis is common in the western
Atlantic, but strays were recorded from waters off South
Fig. 3 Holotype of Acanthostracion bucephalus (AMS IB.6355) Africa (Tyler 1965). Whitley (1965) described Acanthos-
collected from Queensland, Australia. Photograph provided by Mark tracion bucephalus on the basis of a specimen from
McGrouther
Queensland, Australia. However, there seems little doubt
that it is actually A. quadricornis (Fig. 3). The record of A.
(Kyushin et al. 1982; Gloerfelt-Tarp and Kailola 1984; quadricornis from Australia may suggest that the specimen
Sainsbury et al. 1984; Mohsin and Ambak 1996; Randall was transferred by ballast water from a ship or released
et al. 1997; Allen et al. 2006), they were classified in the from an aquarium. As with the Balistidae, many checklists
genus Ostracion by other authors (Klassen 1995; Matsuura and illustrated guidebooks have provided a great deal of
2001; Hayashi and Hagiwara 2013). When Fraser-Brunner information on the taxonomy, distribution, and biology of
(1935b) established Rhynchostracion, he differentiated it boxfishes (see the section on the Balistidae). On the basis
from other ostraciid genera by the degree of the develop- of the above overview, I recognize 22 species in the Ost-
ment of the dorsal ridge, the convexity of the dorsal surface raciidae (Table 6). However the relationships of T. con-
of the carapace, and the projection of the snout anterior to catenatus and T. reipublicae need to be examined in detail
the mouth. However, the two characters involving the based on a comparison of a large selection of specimens
carapace do not clearly differentiate Rhynchostracion from including the types from various localities of the Indo-West
other ostraciids and the snout of Ostracion nasus, the type Pacific to better understand the morphological variations
species of Rhynchostracion, protrudes anteriorly beyond observed in T. reipublicae (Fig. 4).
the mouth in adults but not juveniles. In addition, large
adults of Ostracion cubicus Linnaeus 1758 have a snout
protruding to some extent beyond the mouth. These factors Triodontidae (Threetooth puffer, Fig. 1g)
led Matsuura (2001) to place O. nasus and O. rhinorhyn-
chos in their own genus in his key and brief accounts of The family Triodontidae is represented by a single extant
ostraciids found in the western central Pacific. Klassen species, Triodon macropterus Lesson 1831, but several
(1995) studied osteological characters of many species of fossil taxa referred to this genus date back to the Eocene
the Ostraciidae extensively and provided evidence that (see Tyler and Patterson 1991). The family differs exter-
Rhynchostracion is a junior synonym of Ostracion. Klassen nally from others of the order Tetraodontiformes by the
(1995) also regarded Tetrosomus as a junior synonym of following combination of characters: body moderately
Lactoria because all species of Lactoria and Tetrosomus elongate and moderately compressed, with a very large
examined by him formed a single clade. If Klassen’s ventral flap that may be extended downward by a rotation
(1995) proposal is accepted, Lactoria should be a junior of the long, shaft-like pelvis; jaws modified to form a beak
synonym of Tetrosomus because the latter has priority over of three heavy, massive teeth, two on the upper jaw and one
the former. However, it seems premature to accept his on the lower jaw; two rudimentary dorsal-fin spines present
proposal because there are species-level taxonomic prob- or absent; and caudal fin deeply forked (Matsuura 2001).
lems in several species of Tetrosomus (Allen and Erdman Triodon macropterus is rare in museum collections and
2012). poorly known. It is a deep-water species collected at depths
Tyler (1965) revised the genus Acanthostracion in the of 50-377 m (Kyushin et al. 1977; Matsuura and Tyler
Atlantic and recognized four species, Acanthostracion 1997). When Tyler (1967) redescribed T. macropterus in
guineensis Bleeker 1865, Acanthostracion notacanthus detail, only 20 specimens were available for his study: five
(Bleeker 1863), Acanthostracion polygonus Poey 1876, from the Indian Ocean (Mauritius, Reunion and India) and
and Acanthostracion quadricornis (Linnaeus 1758). 15 from the West Pacific (Indonesia, Philippines and
123
90 K. Matsuura
Fig. 4 Two forms of

Tetrosomus reipublicae
collected from the Ryukyu
Islands (a, b) and Kochi, Japan
(c). a Deep body with no blue
lines (BSKU 29663); b,
c slender body with wavy blue
lines (b BSKU 29697; c BSKU
59588)
Japan). Due to the poor condition of two specimens, Tyler first dorsal fin (Table 7). However, he did not find other
(1967) used 18 specimens to determine whether the first differences between the Indian Ocean specimens and those
dorsal fin, composed of one or two spines, is present or from the West Pacific. This led him to recognize the two
absent. His examination revealed 12 specimens from the groups as being conspecific. I have recently had the
West Pacific to have a first dorsal fin, and five Indian Ocean opportunity of examining 28 specimens of T. macropterus,
specimens and a specimen from the West Pacific to lack a nine from the Indian Ocean and 19 from the West Pacific
(see Appendix). My examination revealed that the first
dorsal fin to be absent in the nine specimens from the
Table 7 Presence or absence of first dorsal fin (two spines) in Tri- Indian Ocean, and present in the 19 specimens from the
odon macropterus. Tyler (1967) showed that a specimen (448.3 mm West Pacific (Table 7). Tyler (1967) also provided fre-
SL) collected from the Volcano Islands, south of the Ogasawara
Islands, had a single spine
quency distributions of fin-ray counts. My examination of
28 specimens produced almost the same frequency distri-
First dorsal fin butions of fin-ray counts as those of Tyler (1967), except
Present Absent for a count of 17 fin rays on the right side pectoral fin in a
single specimen (Table 8).
Indian Ocean 9 ? 5* = 14
Since Tyler’s (1967) redescription was published, T.
West-Pacific 19 ? 12* = 31 1*
macropterus was reported (with color photographs) from
* Data from Tyler (1967) several areas in the Indo-West Pacific: the Chagos
123
Table 8 Frequency distributions of fin-ray counts in Triodon macro- Table 9 A list of 27 genera of the Tetraodontidae in the world
pterus. Counts of fin rays of damaged or abnormal fins are excluded.
Counts of pectoral-fin rays were obtained from fins on both sides Genus Number of species
Dorsal-fin rays Amblyrhynchote Bibron in Duméril 1855 1

Arothron Müller 1841 11
10 11 12
Auriglobus Kottelat 1999 5
This study 3 24 1 Canthigaster Swainson 1839 36
Tyler (1967) 2 14 1 Carinotetraodon Benl 1957 5
Total 5 38 2 Chelonodontops Smith 1958 2
Anal-fin rays Colomesus Gill 1884 3
Contusus Whitley 1947 2
9 10
Dichotomyctere Duméril 1855 6
This study 8 19 Ephippion Bibron in Duméril 1855 1
Tyler (1967) 3 12 Feroxodon Su, Hardy and Tyler 1986 1
Total 11 31 Guentheridia Gilbert and Starks 1904 1
Javichthys Hardy 1985 1
Pectoral-fin rays
Lagocephalus Swainson 1839 11
14 15 16 17 Leiodon Swainson 1839 1
This study 1 12 41 1 Marilyna Hardy 1982a 3
Tyler (1967) 2 7 17 Omegophora Whitley 1934 2
Total 3 19 58 1 Pao Kottelat 2013 14
Pelagocephalus Tyler and Paxton 1979 2
Polyspina Hardy 1983a 1
Reicheltia Hardy 1982a 1
Archipelago (Kyushin et al. 1977), the South China Sea Sphoeroides Anonymous 1798 20
(Kyushin et al. 1982), the Kyushu-Palau Ridge (Okamura Takifugu Abe 1949c 25
et al. 1985), the Timor Sea (Gloerfelt-Tarp and Kailola
Tetractenos Hardy 1983a 2
1984), and the Japanese Archipelago (Matsuura 1984).
Tetraodon Linnaeus 1758 6
Color photographs taken by research staff of the National
Torquigener Whitley 1930a 20
Museum of Nature and Science show T. macropterus to
Tylerius Hardy 1984a 1
have the following coloration: body yellowish brown dor-
Total number of species 184
sally, becoming white ventrally; ventral flap yellow in the
ventral four-fifths and white in the proximal one-fifth; a
large black ocellus with a narrow white edge in the prox- Tetraodontidae (Pufferfishes, Fig. 1h, Table 9)
imal part of the ventral flap; second dorsal, anal, pectoral,
and caudal fins yellow or yellowish brown; when present, Pufferfishes of the family Tetraodontidae usually occur in
membrane of first dorsal fin black (Fig. 1g). In addition to shallow warm, tropical seas, and freshwaters of the world,
the above reports, T. macropterus has been recorded in although some species may be found at depths over 350 m
checklists of fishes from several areas in the West Pacific: (Matsuura 1982; Tyler and Matsuura 1997). They differ
the Mariana Islands (Myers and Donaldson 2003), Tonga externally from other families of the order Tetraodonti-
(Randall et al. 2004), off southern Taiwan (Shao et al. formes by the following combination of characters: head
2008), Northern Territory of Australia (Larson et al. 2013), large and blunt; jaws modified to form a beak of four
and northern half of Australia (Hoese et al. 2006). Kim heavy, powerful teeth, two above and two below; eyes high
et al. (2005) provided a brief account and a color illustra- on head; gill opening, a simple slit in front of pectoral fins;
tion of T. macropterus from Korea. dorsal and anal fins located far posteriorly, containing
Johnson and Britz (2005b) reported the smallest known seven to 15 soft rays; caudal fin truncate, rounded, or
T. macropterus (20 mm SL), collected at Wallis and Futuna emarginate to somewhat lunate; pelvic fins absent; lateral
at a depth of 245-440 m. They provided detailed charac- line (when present) often indistinct, forming an intercon-
ters and the visceral anatomy of the early juvenile that nected pattern on sides of the head and body, but quite
differs from adults in having a huge head (45 % SL vs distinct in some genera (e.g., Lagocephalus and Torqui-
28.5-32.7 % SL in adults), no ventral flap, and different gener Whitley 1930c); typical scales absent, but many
scale structure. Juveniles also differ from adults in having a spinules often present on back and/or belly, and sometimes
very large distended stomach. on sides (Matsuura 2001).
123
92 K. Matsuura
The Tetraodontidae is the most speciose family in the 1970s to present. Allen and Randall (1977) reviewed Can-
Tetraodontiformes, including 184 species (Table 9). thigaster and described seven new species, bringing the total
Because pufferfishes possess very few external characters number of species to 22. Since Allen and Randall’s (1977)
useful for taxonomy and specimens are easily distorted revisionary study was published, six Indo-Pacific species
when they are fixed in formalin and preserved in ethanol, it have been described as new as follows: Canthigaster leop-
is often not easy for ichthyologists to recognize species arda Lubbock and Allen 1979 from the Indo-West Pacific,
limits and classify them into natural groups. Consequently, Canthigaster flavoreticulata Matsuura 1986 from the Tonga
they have been poorly studied and taxonomic confusion Submarine Ridge, South Pacific, Canthigaster cyanetron
remains. Although the Tetraodontidae has never been Randall and Cea-Egaña 1989 from Easter Island, Canthi-
comprehensively reviewed, the taxonomy of pufferfishes gaster punctata Matsuura 1992 from the Mascarene Sub-
has progressed since Fraser-Brunner’s (1943) brief review marine Ridge in the western Indian Ocean, Canthigaster
of tetraodontid genera. He classified pufferfishes into 11 cyanospilota Randall, Williams and Rocha 2008 from the
genera, Canthigaster Swainson 1839, Lagocephalus, Red Sea, and Canthigaster criobe Williams, Delrieu-Trottin,
Sphoeroides Anonymous 1798, Amblyrhynchote Bibron in Planes 2012 from the Gambier Archipelago in French
Duméril 1855, Torquigener, Colomesus Gill 1884, Ephip- Polynesia. In addition, Randall et al. (2008) reviewed Can-
pion Bibron in Duméril 1855, Tetraodon Linnaeus 1758, thigaster coronata (Vaillant and Sauvage 1875) and recog-
Arothron Müller 1841, Chonerhinos Bleeker 1854, Xe- nized Canthigaster axiologa Whitley 1931 as a separate
noptere Bibron in Duméril 1855, based on osteological species. Although Allen and Randall (1977) synonymized
characters of the skull, the lateral-line system on the head Canthigaster papua (Bleeker 1848) with Canthigaster sol-
and body, a skin fold on the ventrolateral side of the body, andri (Richardson 1845), Randall (1995) and Allen and
and spinule distributions on the body. Although Fraser- Erdman (2012) recognized it as a distinctive species. Moura
Brunner (1943) used the spellings Amblyrhynchotes and and Castro (2002) reviewed Canthigaster in the Atlantic
Xenopterus, I follow Kottelat’s (2001, 2013) Am- where many authors recognized only a single species Can-
blyrhynchote and Xenoptere because Kottelat (2001) thigaster rostrata (Bloch 1786). Moura and Castro (2002)
showed that Amblyrhynchotes and Xenopterus of Troschel extensively studied specimens collected from various
(1856) are incorrect spellings. localities from both sides of the Atlantic and recognized six
When Alec Fraser-Brunner studied pufferfishes during species, three of which they described as new: Canthigaster
World War II, Abe (1939, 1942, 1944, 1949a, b, c, 1950, figueiredoi from the east coast of South America, Canthi-
1952, 1954) independently studied pufferfishes found in gaster jamestyleri from the southeast coast of the United
Japan and adjacent regions. Abe (1949c) provided keys and States and the Gulf of Mexico, and Canthigaster supra-
accounts of pufferfishes in the seas around Japan. Subse- macula from the west coast of Africa. The above studies
quently, Abe (1952) proposed Fugu as a new genus to bring the total number of species of Canthigaster to 36,
include five subgenera Akamefugu, Higanfugu, Shosaifugu, making it the largest genus in the Tetraodontidae.
Takifugu, and Torafugu, which had been previously pro- Shipp (1974) reviewed pufferfishes of the Atlantic
posed and made available by Abe (1949c, 1950). However, belonging to Canthigaster, Colomesus Gill 1884, Ephippion
the genus Fugu is an objective synonym of the subgenus Bibron in Duméril 1855, Lagocephalus, and Sphoeroide.
Torafugu Abe 1950, because the type species of these Prior to Shipp’s (1974) publication, Tyler (1964) reviewed
genus group names is Tetraodon rubripes Temminck and two species of Colomesus, Colomesus asellus (Müller and
Schlegel 1850. Matsuura (1990) clarified the nomenclatural Troschel 1848), and Colomesus psittacus (Bloch and
problems concerning the genus group names proposed by Schnedier 1801) in detail. Amaral et al. (2013) described
Abe (1949c, 1950, 1952) and showed that Takifugu Abe Colomesus tocantinensis from the Tocantins River, Brazil.
1949c is the appropriate generic name for warmwater Shipp (1974) recognized 13 species of Sphoeroides in the
pufferfishes found mainly around Japan, Korea, and China. Atlantic and provided a key to the species, along with
As with other tetraodontiform families, de Beaufort and detailed accounts of all Atlantic species. Walker and Bussing
Briggs (1962) lumped Fraser-Brunner’s (1943) Indo-Paci- (1996) described the eastern Pacific Sphoeroides lispus
fic genera, Amblyrhynchote, Lagocephalus, and Torqui- Walker in Walker and Bussing 1996 and Sphoeroides ro-
gener into the genus Sphoeroides although they recognized senblatti Bussing in Walker and Bussing 1996 as new spe-
Canthigaster, Chelonodon Müller 1841, Chonerhinos, and cies. Prior to Shipp (1974) and Walker and Bussing (1996),
Tetraodon. As shown by subsequent authors, Am- five species of Sphoeroides had already been reported from
blyrhynchote, Lagocephalus, and Torquigener are distinc- the eastern Pacific. With 20 species, Sphoeroides is the third
tive genera. largest genus in the Tetraodontidae. Most species of Spho-
Marine pufferfishes occurring in warm and tropical eroides occur in shallow waters of the Atlantic and the
regions of the world were studied by many authors from the eastern Pacific, but Sphoeroides pachygaster (Müller and
123
Troschel in Schomburgk 1848) is found in deep waters Tyler and Paxton (1979) described Pelagocephalus
exceeding 350 m in all oceans. coheni as a new genus and species based on a single
Shipp (1974) also provided detailed accounts and pho- specimen collected at Norfolk Island. Pelagocephalus is
tographs of Lagocephalus laevigatus (Linnaeus 1766) and externally distinguished from other pufferfishes by its nasal
Lagocephalus lagocephalus (Linnaeus 1758). The former apparatus in the form of an open, flat, and relatively
is endemic to the Atlantic and the latter is distributed in unornamented disk. Heemstra and Smith (1981) described
tropical and other warm seas of the world. Abe and Tabeta a second species of Pelagocephalus, Pelagocephalus marki
(1983) described Lagocephalus gloveri as a new species from South Africa. Hardy (1982b) recorded P. marki from
from Japan. Abe et al. (1984) described another new spe- New Zealand.
cies, Lagocephalus wheeleri, from Japan, but Matsuura In the period from 1980 to 1989, Graham S. Hardy
(2010) showed it to be a junior synonym of Lagocephalus published a series of papers on the taxonomy of pufferf-
spadiceus (Richardson 1845). Matsuura et al. (2011) re- ishes that are distributed mainly in the Southern Hemi-
described Lagocephalus guentheri (Miranda Ribeiro 1915) sphere. Hardy (1980) reviewed the antitropical species
based on an examination of the holotype and many speci- Arothron firmamentum (Temminck and Schlegel 1850)
mens from the Red Sea. Although the genus Lagocephalus indicating that it should be classified in Arothron, whereas
has not been comprehensively reviewed, an ongoing study it had previously been classified in Boesemanichthys by
by the author has revealed 11 valid species in the genus. Abe (1952). The monotypic genus Contusus Whitley 1947
This study will be published elsewhere. was reviewed by Hardy (1981), who described a second
Although Abe (1949a, b, c, 1950, 1952, 1954) made species Contusus brevicaudus restricted to southern Aus-
progress on the taxonomy of Takifugu, many species found tralia. Hardy and Hutchins (1981) recognized the validity
along the coast of China were not included in his articles. of Omegophora Whitley 1934 with the description of
Cheng et al. (1975) reviewed pufferfishes found in Chinese Omegophora cyanopunctata from the southwestern part of
waters and described Takifugu flavidus (Li, Wang and Australia. Hardy (1982a) erected two new genera, Mari-
Wang in Cheng et al. 1975) and Takifugu reticularis (Tian, lyna and Reicheltia, based on detailed morphological
Cheng and Wang in Cheng et al. 1975). Subsequently, comparisons including skull osteology. Hardy (1983a)
Chinese authors described four species of Takifugu from reviewed the Australian species of Torquigener and
China: Takifugu orbimaculatus Kuang, Li and Liang 1984 established two new generic names, Polyspina with one
from a creek in Lianhuashan, Guangdong; Takifugu cor- species and Tetractenos with two species. He recognized
onoides Ni and Li 1992 from an estuary of Chang Jiang; 12 species of Torquigener, including five new species from
Takifugu plagiocellatus Li in Su and Li 2002 from the Australia. Hardy and Randall (1983) described Torqui-
coast of Hainan Island; and Takifugu variomaculatus Li in gener flavomaculosus as a new species from the Red Sea.
Su and Li 2002 from brackish waters of Modaomen Zhu- Hardy described Torquigener randalli Hardy 1983b from
hai, Guangdong. These Chinese and Japanese authors the Hawaiian Islands and Torquigener gloerfelti Hardy
increased the number of Takifugu species making it the 1984b from Indonesia. Hardy (1989) described Torqui-
second largest genus in the Tetraodontidae, with 25 spe- gener balteus as a new species from South Africa and
cies. However, taxonomic problems still remain in Taki- provided a key to 19 species of Torquigener, although he
fugu, such as Takifugu basilevskianus (Basilewsky 1855) overlooked Sphoeroides marleyi Fowler 1929, which is a
and Takifugu pseudommus (Chu 1935), which show sig- member of Torquigener. My examination of the holotypes
nificant variation in morphological features including of Torquigener balteus and S. marleyi from South Africa
ontogenetic color changes. revealed them to be conspecific, making the former a junior
Benl (1957) described Carinotetraodon chlupatyi as a synonym of the latter. Matsuura (2014) recently described
new genus and species from Thailand. However, Dekkers Torquigener albomaculosus as a new species from Amami-
(1975) synonymized Carinotetraodon chlupatyi with Cari- oshima Island, Ryukyu Islands. Torquigener albomaculo-
notetraodon lorteti (Tirant 1885), making that species the sus is unique in building large spawning nests, called
type of the genus. Carinotetraodon is similar to Canthigaster ‘‘mystery circles’’ by local SCUBA divers. Thus, the above
in having a laterally compressed body and several derived studies of Torquigener bring the total number of species of
osteological features, as well as a shared derived behavioral Torquigener to 20, the co-equal third largest genus in the
peculiarity in their skin ridge-lifting along the midline of the Tetraodontidae along with Sphoeroides. Hardy (1984a)
body (Tyler 1978, 1980). Additional new species of Cari- recognized Sphoeroides spinosissimus Regan 1908 as
notetraodon were described: Carinotetraodon salivator Lim being distinctive from other genera of the Tetraodontidae
and Kottelat 1995 from Sarawak, Carinotetraodon irrubesco and established a new genus Tylerius for it. Hardy (1985)
Tan 1999 from Sumatra, and Carinotetraodon imitator Britz described Javichthys kailolae as a new genus and species
and Kottelat 1999 from India. from Indonesia. Su et al. (1986) established a new genus,
123
94 K. Matsuura
Feroxodon, for Anchisomus multistriatus Richardson 1854 pleurospilus (Regan 1919) and congeneric with Tetraodon
from Australia. patoca (see Smith 1986). Thus, Tetraodon patoca is a
The taxonomy of freshwater pufferfishes in Asia was in a species of Chelonodontops (Kottelat 2013). Matsuura
confused state until Dekkers (1975) reviewed the genus (2002) reviewed two species of Chelonodon, Chelonodon
Tetraodon. Subsequently, Sontirat and Soonthornsatit laticeps Smith 1948, and Chelonodon patoca, both of
(1985) described Tetraodon suvattii as a new species from which belong in Chelonodontops. Kottelat (2013) showed
Thailand, and Roberts (1988) described Tetraodon abei as a that Gastrophysus Müller 1843 is an objective synonym of
new species from Laos. Roberts (1982) reviewed the genus Takifugu Abe 1949, as the type species of both genus group
Chonerhinos Bleeker 1854 and described three species from names is Tetrodon oblongus Bloch 1786. However, this is
Borneo, Chonerhinos amabilis, Chonerhinos nefastus, and likely to cause confusion not only in pufferfish taxonomy,
Chonerhinos silus. Still, complex nomenclatural problems but also in the fisheries of East Asia where pufferfishes are
involving Chonerhinos and Xenoptere remained until Ko- treated as important and expensive fishes (e.g., several
ttelat (1999) proposed the new genus, Auriglobus, to rectify species of Takifugu cost about US$100 per kilogram at fish
these problems. Kottelat (2013) discussed nomenclatural markets). In addition, because pufferfishes have fatal poi-
issues of the Tetraodontidae found in Southeast Asia in son in their viscera, scientific names are very important for
detail. He established the new genus Pao (type species, food security management in East Asia. As suggested by
Tetraodon leiurus Bleeker 1850a) that includes 13 species Kottelat (2013), Gastrophysus can be suppressed under the
previously placed in Tetraodon and Monotrete, but differ- International Code of Zoological Nomenclature (ICZN
entiated from other pufferfishes by their unique color pattern 1999). I will ask the International Commission on Zoo-
and a very elongate premaxillary pedicel that creates a logical Nomenclature to suppress Gastrophysus.
greatly enlarged open space between their dorsomedial Matsuura and Okuno (1991) redescribed a rare puffer-
edges. Thus, the generic name Tetraodon is now applied to fish, Arothron carduus (Cantor 1849), on the basis of the
only six species of African freshwater pufferfishes. Kottelat holotype from Penang and two additional specimens from
(2013) also recognized Leiodon Swainson 1839 as a valid the West Pacific. Matsuura (1994) described Arothron
genus that includes Tetrodon cutcutia Hamilton 1822 dis- caeruleopunctatus as a new species from the tropical
tributed in freshwaters of southern Asia. Kottelat (2013) region of the Indo-West Pacific. Matsuura (1999) briefly
recognized Dichotomyctere Duméril 1855 for six species: reviewed Arothron with a key to species and accounts of all
Dichotomyctere erythrotaenia (Bleeker 1853), Dichoto- members of the genus including two undescribed species.
myctere fluviatilis (Hamilton 1822), Dichotomyctere kre- As indicated above, there has been a great deal of pro-
tamensis (Inger 1953), Dichotomyctere nigroviridis (Marion gress in taxonomy of the Tetraodontidae during the past
de Procé 1822), Dichotomyctere ocellatus (Steindachner several decades, especially in the period from the mid-
1870), and Dichotomyctere sabahensis (Dekkers 1975). 1970s to the present. However, there remain taxonomic
Saenjudaeng et al. (2013) described Tetraodon palustris as a problems in genera such as Arothron, Chelonodontops,
new species from the Mekong basin of Thailand. Because of Lagocephalus, Pao, Takifugu, and Torquigener where
its publication date, July 2013, the authors were unaware of many species await description and detailed morphological
Kottelat’s (2013) article proposing the genus Pao. However, and molecular comparisons to classify them into appro-
it is clear from the original description of Tetraodon pa- priate groups. The taxonomy of pufferfishes in Southeast
lustris, that it is a species of Pao, bringing the total number Asia is important not only for the understanding of fish
of species in that genus to 14. diversity of the region, but also for the welfare and food
Kottelat (2013) also commented on genera of marine management for humans. Dao et al. (2012) reported that
pufferfishes. Although Tetrodon patoca Hamilton 1822 has the number of victims of food poisoning by eating puf-
long been placed in Chelonodon Müller 1841, Kottelat ferfishes reached 737, with 127 mortalities from 1999 to
(2013) pointed out that Tetrodon cutcutia Hamilton 1822, 2003. Due to the lack of knowledge about pufferfishes and
the type species of Chelonodon, had been placed in the their toxicity, pufferfishes are still found in Southeast Asian
same genus as Tetraodon fluviatilis Hamilton 1822 and fish markets (Fig. 5), although local and state governments
related species by Tyler (1980). Tyler (1980) called this in countries around the South China Sea have prohibited
group Chelonodon but Kottelat (2013) stated that the oldest the sale of pufferfishes for food.
available name for a genus including Tetraodon fluviatilis
is Dichotomyctere. Tetrodon patoca and Tetraodon flu-
viatilis have a sufficient number of character differences to Diodontidae (Porcupinefishes, Fig. 1i)
warrant being placed in separate genera. The type species
of Chelonodontops Smith 1958 is Chelonodontops pul- Many species of the family Diodontidae are benthic
chellus Smith 1958, a junior synonym of Chelonodon inhabitants around coral or rocky reefs, but some occur
123
Fig. 5 Pufferfishes found in a

fish landing place 60 km north
of Nha Trang in southern
Vietnam (a) and a fish market in
Sabah, Malaysia (b): a many
small pufferfishes of
Lagocephalus and a large
specimen of L. inermis in the
center; b large specimens of
Arothron hispidus, A.
reticularis, and A. stellatus
frequently in seagrass beds and on sand or mud bottom to form a strong, beak-like crushing structure without a
depths of 150 m, with one species being found in pelagic median suture dividing the upper and lower jaws into left
waters. They differ externally from other families of the and right halves; gill opening, a relatively small, vertical
order Tetraodontiformes by the following combination of slit immediately preceding the pectoral-fin base; short-
characters: body wide and capable of great inflation, based dorsal and anal fins with soft rays, set far back on
covered with massive spines, which may be quite long; body, and like caudal fin, generally rounded; most fin rays
spines with large bases or roots, which are beneath the branched; bases of fins often thick and fleshy; no pelvic
skin; long spines that are usually erectile and double- fins (Leis 2001).
rooted, or short spines fixed in an erect position by their Porcupinefishes were well studied by Leis (1978, 1986,
triple-rooted bases; head broad and blunt; nasal organ 2001, 2003, 2006). Leis (1978) reviewed Diodon Linnaeus
usually appearing as small tentacles located in front of the 1758 and provided detailed accounts of five species: Dio-
large eyes; mouth large, wide, and terminal, teeth fused to don eydouxii Brisout de Barneville 1846, Diodon
123
96 K. Matsuura
holocanthus Linnaeus 1758, Diodon hystrix Linnaeus how many species in the family, some believing a single
1758, Diodon liturosus Shaw 1804, and Diodon nicthe- species exists in each genus, and others recognizing two
merus Cuvier 1818. Leis (2006) provided a list of syn- species each in Mola and Masturus and one species in
onyms and keys for all species of the Diodontidae. Keys to Ranzania (Bray 2008).
regional species were also provided by Leis (1986, western Parenti (2003) published a list of nominal species in the
Indian Ocean; 2001, eastern Indian Ocean and western Molidae. Bass et al (2005) published a molecular analysis
central Pacific; 2003, western Atlantic). Accounts and of ocean sunfishes and recognized two clades in Mola, M.
color photographs for species of the eastern Pacific were mola and M. ramsayi. They demonstrated that M. ramsayi
provided by Allen and Robertson (1994). Australian spe- occurs in South Africa and Australia, whereas M. mola is
cies were diagnosed and illustrated by Gomon (2008). As found in all oceans. However, they were unable to differ-
pointed out by Leis (2006), taxonomic problems remain for entiate among specimens of Masturus from the West
some species, such as Chilomycterus reticulatus (Linnaeus Pacific (Taiwan) and the western Atlantic (Florida).
1758) and species of ‘‘Atlantic Chilomycterus.’’ Yoshita et al. (2009) studied ocean sunfishes within the
genus Mola around Japan. They took measurements and
counted 99 specimens of young and adults (maximum size
Molidae (Ocean sunfishes, Fig. 1j) 332 cm TL, Fig. 6). They showed convincingly that there
are two species with clear morphological differences: a
Ocean sunfishes of the family Molidae occur in tropical well-developed head bump (head bump height 12.1 % TL)
and other warm seas of the world. They are usually pelagic, in group A vs. with no distinct head bump (head bump
descending to a depth of over 200 m. The ocean sunfishes height 7.8 % TL) in group B, number of clavus fin rays
differ externally from other families of Tetraodontiformes 14-17 in group A vs 10-13 in group B, number of clavus
by the following combination of characters: body short and ossicles 8-15 in group A vs. 8-9 in group B, and edge of
deep or oblong, prominently compressed; caudal peduncle clavus not wavy in group A vs. wavy in group B. Yoshita
and typical caudal fin absent; eyes small; mouth terminal, et al. (2009) suggested strongly that their group A was M.
small; teeth united and beak-like in each jaw without a ramsayi and group B was M. mola. However, a clearer
median suture; no palatine teeth; gill opening small, pore- view of intra- and interspecific relationships of Mola must
like, located in front of pectoral-fin base; dorsal and anal await the availability of more specimens from other regions
fins of similar shape, generally triangular, dorsal fin located of the world’s oceans for study by both morphological and
opposite anal fin; dorsal and anal fins spineless, each with molecular methods.
15 to 21 soft rays; pectoral fins of small to moderate size,
located midlaterally, fitting into a shallow concavity in side
of body in some; pelvic fins absent; caudal fin replaced by a Systematics
leathery, rudder-like lobe known as a pseudocaudal fin or
clavus (supported mostly by fin-ray elements originally Cuvier (1816) pioneered the classification of tetraodonti-
belonging to dorsal and anal fins); skin leathery, with many form fishes by placing them in the order Plectognathi based
small scales (small juveniles may also have some larger on his detailed anatomical studies. Since Cuvier’s (1816)
scattered spiny scales) (Hutchins 2001b). work was published, there have been various arrangements
Fraser-Brunner (1951) reviewed ocean sunfishes and of tetraodontiform classification (see Tyler 1980 for a his-
recognized five species in the Molidae: Masturus lanceol- tory of tetraodontiform classification), but the Plectognathi
atus (Liénard 1840), Masturus oxyuropterus (Bleeker or Tetraodontiformes has nearly always been considered a
1873), Mola mola (Linnaeus 1758), Mola ramsayi (Giglioli monophyletic group, except for Le Danois (1955, 1959,
1883), and Ranzania laevis (Pennant 1776). Because spe- 1961). As appropriately criticized by Tyler (1963), Yseult
cies of Mola and Masturus reach more than 3 m in length Le Danois tried to destroy the order Tetraodontiformes. She
and two tons in weight, it is difficult to preserve specimens stated that the triacanthoids and balisoitds are of acanthurid
of adults in museums. It is also difficult for ichthyologists origin and that the other plectognaths (her Orbiculati) are
to obtain measurements and counts on large adult ocean not even of percoid derivation, being related to the iso-
sunfishes in the field, and specimens of Mola and Masturus spondylous fishes, and that Canthigaster is related to the
are not frequently collected and returned to museum col- ostracioids rather than the tetraodontids. However, her
lections. Ranzania laevis is relatively small, reaching statements were based on erroneous observations and
around 80 cm TL, but specimens of Ranzania have also interpretations of the osteological and myological charac-
been rarely collected. Since ichthyologists have few ters of tetraodontiforms (Tyler 1963; Winterbottom 1974).
opportunities to study an adequate number of adult speci- No comprehensive phylogenetic studies of tetraodontiforms
mens of ocean sunfishes, authors have been unable to agree appeared prior to studies by Winterbottom (1974) and Tyler
123
Fig. 6 A large specimen of

Mola ramsayi (330 cm TL)
captured with a set net along the
Pacific coast of northern
Honshu, Japan
(1980). Morphological synapomorphies of tetraodontiforms On the basis of a comprehensive myological study,

have been provided by Winterbottom (1974), Tyler (1980), Winterbottom (1974) analyzed relationships of Tetra-
Rosen (1984), Tyler and Sorbini (1996), and Santini and odontiformes using cladistic methods. He recognized three
Tyler (2003). In their review of the interrelationships of major clades: (1) Triacanthodidae ? Triacanthidae, (2)
actinopterygian fishes, Lauder and Liem (1983) supported (Balistidae ? Monacanthidae) ? (Aracanidae ? Ostracii-
the monophyly of Tetraodontiformes by adding to the dae), and (3) Triodontidae ? [(Tetraodontidae ? Dio-
already substantial list of synapomorphies. In addition, dontidae) ? Molidae] (Fig. 7). The first clade, including
Wiley and Johnson (2010) provided a list of 10 synapo- the Triacanthodidae and Triacanthidae, was considered to
morphies for fossil and extant tetraodontiforms. be the sister group of all other tetraodontiforms. Tyler
123
98 K. Matsuura
phylogenetic relationships of the Tetraodontiformes by an

evolutionary (traditional) method and his systematic
arrangements of families are similar to those of Winter-
bottom (1974), except for placing the superfamily Triac-
anthoidea (Triacanthodidae ? Triacanthidae) as the basal
sister group to the two superfamilies, the Balistoidea
(Balistidae and Monacanthidae) and the Ostracioidea
(Aracanidae and Ostraciidae).
Winterbottom (1974) and Tyler (1980) stated that tet-
raodontiforms were probably related to acanthuroid fishes
of the Perciformes, although Winterbottom (1974) sug-
gested that some zeiforms might also be related to tetra-
odontiforms. Rosen (1984) proposed on the basis of his
osteological analysis that zeoids have a sister-group rela-
tionship with tetraodontiforms and the two groups are sister
to the caproids. Leis (1984) used characters of eggs and
larvae to investigate tetraodontiform relationships.
Although no aracanid and triodontid larvae were available
for his study, Leis (1984) showed relationships of tetra-
odontiform families that were similar to those of Winter-
bottom (1974) and Tyler (1980), except for the
phylogenetic position of the Ostraciidae. Leis (1984)
placed the Ostraciidae in an unresolved trichotomy, Ost-
raciidae ? Diodontidae ? Molidae, and placed the three
families as a sister clade to the Tetraodontidae (Fig. 7b).
This was the first time that the Ostraciidae was placed in
the gymnodonts (Triodontidae, Tetraodontidae, Diodonti-
dae, and Molidae). Winterbottom and Tyler (1983) also
provided many synapomorphies involving osteological and
myological characters that supported a sister-group rela-
tionship of balistoids and ostracioids. Klassen (1995)
similarly supported the relationship of the two groups.
James C. Tyler and other authors published many papers
on fossil tetraodontiforms (e.g., Tyler and Patterson 1991;
Tyler and Banikov 1992, 1994, 2011, 2012; Tyler et al.
1992, 1993, 2000, 2003, 2006; Tyler and Sorbini 1996,
1998; Tyler and Winterbottom 1999; Tyler and Santini
2001, 2002; Sorbini and Tyler 2004; Bannikov and Tyler
2008a, b; Gregorova et al. 2009; Carnevale and Tyler 2010;
Tyler and Kriznar 2013; Miyajima et al. 2014).
Santini and Tyler (2003) used osteological data on fossil
and extant tetraodontiform fishes accumulated by previous
Fig. 7 Phylogenetic relationships of the extant families of Tetra-
odontiformes inferred from cladistic analyses of morphological contributions to generate a new classification of all known
characters. a Winterbottom 1974; b Leis 1984 (larvae of the families represented by fossil and extant forms. For extant
Aracanidae and Triodontidae not available for Leis); c Santini and families, Santini and Tyler (2003) placed the Triacan-
Tyler 2003 thodidae as a sister group to all other families of the Tet-
raodontiformes (Fig. 7). They arranged the other families
(1980) studied tetraodontiforms extensively and provided a into two suborders, the Balistoidei and Tetraodontoidei.
huge number of osteological descriptions, comparative The former is composed of the Triacanthidae ? [(Balisti-
diagnoses, and illustrations for all extant families, major dae ? Monacanthidae) ? (Aracanidae ? Ostraciidae)].
representatives of extant genera, and most of the known The latter suborder is composed of the Triodontidae ?
fossil taxa of the Tetraodontiformes. Tyler (1980) analyzed [(Tetraodontidae ? Diodontidae) ? Molidae]. This
123
classification is similar to those of Winterbottom (1974) suggestive of a close relationship between the Ostraciidae
and Tyler (1980). and Molidae. However, because they studied only one
Miya et al. (2003) analyzed molecular data for 100 species of Ostraciidae and two species of Molidae, it seems
species of higher teleosteans. Their study was the first to premature to hypothesize a close relationship of the two
hypothesize close relationships among lophiiforms, tetra- families until further support for it may be forthcoming. In
odontiforms, and caproids. They retrieved phylogenetic contrast, the Molidae was recovered as the sister group to a
topologies placing caproids as a sister group with tetra- clade comprising the Tetraodontidae and Diodontidae by
odontiforms and the two groups as sister to lophiiforms. Santini et al. (2013c) as previously indicated by morpho-
However, they used few taxa: one species of Caproidae, logical studies. The Triodontidae was considered to be
two species of Tetraodontiformes, and six species of Lo- close to the Tetraodontidae, Diodontidae, and Molidae by
phiiformes. Yamanoue et al. (2007) analyzed more species morphological studies (Winterbottom 1974; Tyler 1980;
of the three groups and provided a robust phylogenetic Santini and Tyler 2003), whereas molecular studies
topology that placed the Caproidei as sister to the Lophii- recovered close relationship of the Triodontidae with the
formes, and the two groups as a clade that is sister to the Aracanidae and Ostraciidae (Alfaro et al. 2007; Yamanoue
Tetraodontiformes. et al. 2008; Santini et al. 2013c).
Nakae and Sasaki (2010) studied the lateral-line system Despite differences in molecular trees of familial rela-
and its innervations of nine species of tetraodontiforms tionships among authors, several sister group relationships
(representing all families examined except for the Molidae) are generally supported by molecular studies (Holcroft
and a single species each from the Lophiidae, Zeidae, Cap- 2005; Alfaro et al. 2007; Yamanoue et al. 2007; Santini
roidae, and Siganidae. Their analysis supported a close et al. 2013c) as follows: Triacanthodidae ? Triacanthidae,
relationship of the Tetraodontiformes with the Lophiidae, Balistidae ? Monacanthidae, Aracanidae ? Ostraciidae,
but not with the Zeidae, Caproidae, or Siganidae. Recently, and Tetraodontidae ? Diodontidae, as previously seen in
Chanet et al. (2013) presented synapomorphies of tetra- morphological studies. On the basis of whole mitochon-
odontiforms and lophiiforms involving soft anatomical drial genome sequences, Yamanoue et al. (2008) docu-
characters: rounded and anteriorly disposed kidneys, a mented different phylogenetic relationships from those of
compact thyroid included in a blood sinus, an abbreviated other authors. Whereas the Triacanthidae was usually
spinal cord, an asymmetric liver, and clusters of suprame- placed close to the Triacanthodidae, Balistidae, and Mon-
dullary neurons in the rostral part of the spinal cord. Baldwin acanthidae in other studies (Winterbottom 1974; Tyler
(2013) also provided a putative synapomorphy of some tet- 1980; Leis 1984; Santini and Tyler 2003; Holcroft 2005;
raodontiforms and lophiiforms: they are strikingly similar in Alfaro et al. 2007; Santini et al. 2013c), Yamanoue et al.
having the trunk enclosed in an inflated sac covered with (2008) recovered a relationship of the Triacanthidae with
xanthophores. Although Nakae and Sasaki (2010), Chanet the Tetraodontidae and Diodontidae.
et al. (2013), and Baldwin (2013) did not study many species Phylogenetic relationships of genera and species were
of tetraodontiform and outgroup fishes, and the relationship studied by various authors in the Triacanthidae, Balistidae,
between the Tetraodontiformes and Lophiiformes is now Monacanthidae, Aracanidae, Ostraciidae, Tetraodontidae,
supported both by molecular and morphological characters. and Molidae, but the numbers of taxa studied differed
Molecular studies by various authors have generally greatly and resulted in different resolutions of phylogenetic
supported the monophyly of tetraodontiform families analyses. Santini and Tyler (2002b) analyzed phylogenetic
(Holcroft 2005; Alfaro et al. 2007; Yamanoue et al. 2007, relationships of species and genera of the Triacanthidae
2008), although conflicts exist between the constructed using morphological characters. They found two clades,
topologies of familial relationships by morphological and one composed of two species of Triacanthus Oken 1817,
molecular analyses (Figs. 7, 8). In the morphological and the other composed of three species of Tripodichthys
studies by Winterbottom (1974), Tyler (1980), and Santini Tyler 1968 and one species each of Pseudotriacanthus
and Tyler (2003), the Triodontidae, Tetraodontidae, Dio- Fraser-Brunner 1941b and Trixiphichthys Fraser-Brunner
dontidae, and Molidae form a monophyletic group, 1941b. Matsuura (1979, 1981) analyzed cladistic relation-
whereas Holcroft (2005) placed the Molidae with Araca- ships of all extant genera of the Balistidae and 22 genera of
nidae ? Ostraciidae, treating them as subfamilies. Alfaro the Monacanthidae using osteological characters. Although
et al. (2007) placed the Molidae as the basal sister group to many balistid genera showed no significant differences in
Triodontidae ? (Aracanidae ? Ostraciidae). Britz and osteological characters, Matsuura (1979) found that a
Johnson (2005) and Johnson and Britz (2005a) found the derived condition of the interhyal was shared by Rhine-
fusion of anterior vertebral centra in the occipital region canthus and Sufflamen. He also recovered Canthidermis as
and a thick band of cartilage on the side of the pterygio- sister to all other balistid genera based on a scale bone in
phores of the vertical fins in an ostraciid and molids the posterior part of the skull. However, Tyler and
123
100 K. Matsuura
Fig. 8 Phylogenetic relationships of the Tetraodontiformes inferred (2005) and Yamanoue et al. (2007) included two subfamilies the
from molecular analyses. a Holcroft 2005; b Alfaro et al. 2007; Aracaninae and Ostraciinae, which were classified as families in the
c Yamanoue et al. 2007; d Santini et al. 2013c. The Triodontidae was other papers
not included in Holcroft (2005), and the Ostraciidae of Holcroft
Matsuura (1981) revealed Matsuura’s (1979) ‘‘scale bone’’ In her molecular analyses of tetraodontiforms, Holcroft
to be the posterior part of the sphenotic that appears sep- (2005) also recovered the close relationship of Rhinecan-
arated from the major portion of the sphenotic. Matsuura thus and Sufflamen found by Matsuura (1979). She further
(1981) found derived conditions of the pectoral girdle and showed that three species of Balistes form a sister group to
the skull in Xenobalistes, making it a sister group to all Pseudobalistes fuscus (Bloch and Schneider 1801) and that
other balistid genera. Matsuura (1979) recognized two Xanthichthys auromarginatus (Bennett 1832) is sister to
clades in the Monacanthidae: the first composed of Aro- Balistoides viridescens. Her analysis revealed that Bal-
trolepis Fraser-Brunner 1941c, Paramonacanthus, Mon- istoides conspicillum is sister to other balistids, including
acanthus, Stephanolepis Gill 1861, Chaetodermis, Balistes, Pseudobalistes, Rhinecanthus, Sufflamen, Abal-
Acreichthys, and Pervagor; the second Cantherhines, Eu- istes, Canthidermis, Xanthichthys, and Balistoides virides-
balichthys Whitley 1930b, Thamnaconus (= Navodon cens. In addition, she found Melichthys niger (Bloch 1786)
Whitley 1930b of Matsuura 1979), Pseudomonacanthus, to be the first lineage to diverge in the Balistidae. Although
Scobinichthys Whitley 1931, Nelusetta Whitley 1939, Holcroft (2005) studied phylogenetic relationships of nine
Meuschenia Whitley 1929, Rudarius, Aluterus, Oxymon- genera of Monacanthidae, her study covered relatively
acanthus Bleeker 1865, Pseudalutarius, Brachaluteres, small numbers of monacanthid genera making it difficult to
Paraluteres, and Anacanthus. The second clade possesses compare her phylogenetic tree with that constructed by
more advanced characters than the first clade and is com- Matsuura (1979). Yamanoue et al. (2009) studied the
posed of three subclades: (1) Amanses and Cantherhines, phylogenetic relationships of 12 species of Balistidae and
(2) Eubalichthys, Thamnaconus, Pseudomonacanthus, 21 genera of Monacanthidae, involving one species each
Scobinichthys, Nelusetta, Meuschenia and Rudarius, and from 33 genera of the two families. The molecular analysis
(3) Aluterus, Oxymonacanthus, Pseudalutarius, Brachalu- placed Balistes as sister to all other balistid genera that
teres, Paraluteres, and Anacanthus. were separated into two clades, the first comprising
123
Canthidermis, Abalistes, Rhinecanthus, and Sufflamen, and (two species), Caprichthys (monotypic), Capropygia
the second Pseudobalistes, Xanthichthys, Xenobalistes, (monotypic), Aracana (two species), and Anoplocapros
Odonus, Balistapus, Balistoides, and Melichthys. In the (three species), with Polyplacapros (monotypic). The two
Monacanthidae they found the clade that included two monotypic genera Caprichthys and Capropygia were found
genera, Oxymonacanthus and Pseudalutarius, form a sister to represent a clade to other genera. Kentrocapros was
group to other genera that are divisible into two major placed as sister to the subclade composed of Aracana and
clades, the first composed of Rudarius, Brachaluteres, Anoplocapros.
Paraluteres, Pervagor, Stephanolepis, Acreichthys, Chae- Klassen (1995) studied the osteological characters of 19
todermis, Monacanthus, and Paramonacanthus, and the species of the Ostraciidae extensively. His phylogenetic
second Aluterus, Pseudomonacanthus, Amanses, Can- analysis found two clades in the Ostraciidae, the first
therhines, Eubalichthys, Thamnaconus, Nelusetta, Scobi- composed of Acanthostracion and Lactophrys Swainson
nichthys, Meuschenia, and Acanthaluteres. 1839, and the second Ostracion and Lactoria, having
Santini et al. (2013a) provided the largest molecular synonymized Rhynchostracion with Ostracion and Tetro-
dataset for the Balistidae and Monacanthidae based on two somus with Lactoria. Santini et al. (2013a) studied the
mitochondrial and three nuclear loci of 33 species of Bal- phylogenetic relationships of 17 species of Ostraciidae by
istidae and 53 species of Monacanthidae. The molecular molecular analysis. The resultant phylogenetic tree closely
analysis supported monophylies of the following genera: resembled that of Klassen (1995), but where Klassen
Odonus, Balistapus, Melichthys, Xanthichthys, Canthider- (1995) found Rhynchostracion nasus (Bloch 1785) deeply
mis, Balistes, Abalistes, Rhinecanthus, and Sufflamen. nested within Ostracion, Santini et al. (2013a) presented it
Species of Balistoides and Pseudobalistes were recovered as sister to Ostracion.
in separate clades with Balistoides conspicillum sister to On the basis of many osteological characters, nasal
Melichthys, and B. viridescens forming a clade with P. organ, and lateral-line system, Tyler (1980) inferred three
flavimarginatus. The latter two species share at least two groups of pufferfish genera: (1) Lagocephalus, Colomesus,
morphological characters, a small naked area on the cheek Guentheridia Gilbert and Starks 1904, and Sphoeroides;
just behind the mouth, and a large maximum size, reaching (2) Amblyrhynchote, Torquigener, and Fugu (= Takifugu);
over 60 cm TL. In the balistid tree Xenobalistes tumidi- and (3) all other genera. Holcroft (2005) and Alfaro et al.
pectoris was deeply nested in a clade composed of five (2007) analyzed the phylogenetic relationships of 19 spe-
species of Xanthichthys, strongly supporting Xenobalistes cies of 10 genera (Arothron, Canthigaster, Lagocephalus,
as a junior synonym of Xanthichthys (see Santini et al. Marilyna, Monotreta, Sphoeroides, Takifugu, Tetractenos,
2013b). Santini et al. (2013b) recovered virtually all of the Tetraodon, and Torquigener). Their results are similar,
subclades of Yamanoue et al. (2009), except for Chaeto- placing Lagocephalus at the base of the topologies.
dermis and Acreichthys relationships. In the phylogenetic Yamanoue et al. (2011) studied the phylogenetic relation-
topology of Santini et al. (2013b), Chaetodermis and ships of 50 species of Tetraodontidae by molecular ana-
Acreichthys are sequential sister taxa to the clade com- lysis. The study found four major clades, the first
posed of three species of Paramonacanthus and Mon- composed of species of Lagocephalus, the second Co-
acanthus chinensis, whereas Yamanoue et al. (2009) lomesus and three species of Sphoeroides, the third Mari-
presented sister-group relationships of Chaetodermis and lyna, Tetractenos, Tylerirus, Polyspina, Torquigener, and
Acreichthys. Santini et al. (2013b) supported monophylies Takifugu, and the fourth and the largest clade Carinotet-
or close relationships of most genera, but species of raodon, Tetraodon, Auriglobus, Pelagocephalus, Canthi-
Monacanthus and Paramonacanthus were separated into gaster, Chelonodon, Omegophora, and Arothron. The
different clades suggesting strongly that these genera phylogenetic tree of Yamanoue et al. (2011) recognized
require revision. many monophyletic lineages, but Colomesus and species of
A cladistic analysis of the external characters, osteology, Tetraodon fell into distantly related clades. A relaxed
and myology of all genera of Aracanidae led Winterbottom molecular-clock Bayesian divergence time estimation cal-
and Tyler (1983) to recognize two clades in the Aracani- culated three invasions to freshwater by pufferfishes during
dae, the first composed of Kentrocapros and Polyplaca- their evolution: the first 48-78 million years ago in
pros, and the second Aracana, Strophiurichthys Fraser- Southeast Asia, the second 17-38 million years ago in
Brunner 1935b, Anoplocapros, Caprichthys, and Capro- Africa, and the third 0-10 million years ago in South
pygia. In the second clade Aracana was sister to the America.
remaining genera, Strophiurichthys and Anoplocapros Igarashi et al. (2013) studied the phylogenetic rela-
diverged sequentially, and Caprichthys and Capropygia tionships of 17 species of Tetraodon by molecular ana-
were placed in a small terminal clade. Santini et al. (2013a) lysis. The study clearly showed that they did not form a
studied the phylogenetic relationships of Kentrocapros monophyletic group. Except for T. cutcutia, 16 species
123
102 K. Matsuura
were placed into three major clades: an Asian freshwater species in 27 genera, Diodontidae with 18 species in seven
group composed of T. leiurus, Tetraodon palembangensis genera, and Molidae with five species in three genera.
Bleeker 1850b, T. abei, Tetraodon baileyi Sontirat 1985, However, many taxonomic problems involving species and
Tetraodon cochinchinensis Steindachner 1866, T. suvattii, genera of various families still remain. In particular, tax-
and Tetraodon turgidus Mitchill 1815; an Asian brackish onomic anomalies exist in the Monacanthidae and Tetra-
water group of Tetraodon erythrotaenia Bleeker 1853, odontidae, which have the greatest number of species and
Tetraodon biocellatus Tirant 1885, T. fluviatilis, and Tet- diversity within the order. Studies of spikefishes of the
raodon nigroviridis Marion de Procé 1822; and an African Triacanthodidae reveal clarity on the number of genera and
freshwater group of Tetraodon lineatus Linnaeus 1758, species, but poorly surveyed deep areas of the world’s
Tetraodon pustulatus Murray 1857, Tetraodon mbu Bou- oceans exist where new species may occur. Several genera
lenger 1899, Tetraodon miurus Boulenger 1902, and Tet- of Balistidae, Aracanidae, and Ostraciidae require study
raodon duboisi Poll 1959. Two molecular studies by both by morphological and molecular analyses to clarify
Yamanoue et al. (2011) and Igarashi et al. (2013) pro- generic and species limits. The large size attained by ocean
duced essentially the same phylogenetic tree for fresh- sunfishes has made it difficult for us to have a clear
water pufferfishes. understanding of species limits, although a combination of
In his review of ocean sunfishes, Fraser-Brunner (1951) molecular and morphological approaches have clarified
found Masturus Gill 1884 and Mola to be more closely that the presence of several populations probably represent
related to each other than to Ranzania. Tyler’s (1980) species of Mola (see Bass et al. 2005; Yoshita et al. 2009).
extensive surveys of osteological characters supported the When the taxonomic levels of these populations are
phylogenetic tree of Fraser-Brunner (1951). Santini and established, difficult nomenclatural challenges still remain:
Tyler (2002a) performed the first cladistic analysis of the many nominal species of ocean sunfishes were published in
phylogenetic relationships of ocean sunfishes, using 48 the old days without type specimens.
morphological characters. Their study supported the results With regard to the systematics of the Tetraodontiformes
of the previous studies. Yamanoue et al. (2004) used overall, familial relationships have been clarified by mor-
mitochondrial genomes to analyze the phylogenetic rela- phological and molecular analyses, providing us with a
tionships of ocean sunfishes. Their study resulted in the firm understanding of the sister relationships of the fol-
same phylogenetic tree as recovered by previous morpho- lowing family groups: Triacanthodidae and Triacanthidae,
logical studies. Molecular studies on tetraodontiform rela- Balistidae and Monacanthidae, and Tetraodontidae and
tionships by Alfaro et al. (2007) also produced the same Diodontidae. However, the phylogenetic positions of the
result. Bass et al. (2005) examined many specimens of Triodontidae and Molidae remain unclear because of con-
ocean sunfishes captured from various localities world- flicts about their positions in morphological and molecular
wide. The molecular analysis found that M. ramsayi is studies (the position of Molidae differs even among
distributed in the Southern Hemisphere and M. mola in molecular studies). More taxon sampling is needed for
both the Northern and Southern hemispheres. Yoshita et al. molecular analyses to produce more robust phylogenetic
(2009) studied ocean sunfishes of Mola around Japan topologies. Although generic and specific phylogenetic
employing both morphological and molecular characters. relationships for the families Triacanthidae, Balistidae,
As discussed in the section of Molidae, the study clearly Monacanthidae, Aracanidae, Ostraciidae, and Molidae
showed two species of Mola around Japan, probably have been studied using morphological and/or molecular
identifiable as M. mola and M. ramsayi. analyses, the genera and species of the Triacanthodidae and
Diodontidae have never been studied cladistically and
await future studies.
Concluding remarks
Acknowledgments I am grateful to the members of the Organizing
The taxonomy of tetraodontiforms has progressed greatly Committee of the 9th Indo-Pacific Fish Conference for inviting me to
since the mid-1960s. On the basis of contributions to date, write this paper. I thank numerous people in museums and universi-
this paper identifies 412 extant species in the 10 living ties for supporting my research on tetraodontiform fishes. Ralf Britz
in the Natural History Museum kindly provided access to old litera-
families of Tetraodontiformes, with an allocation of species ture. Martin F. Gomon of the Museum Victoria and Edward O. Murdy
and genera as follows: Triacanthodidae including 23 spe- of the George Washington University kindly provided comments on
cies in 11 genera, Triacanthidae with seven species in four the draft manuscript. My thanks go to James C. Tyler and an anon-
genera, Balistidae with 37 species in 12 genera, Mon- ymous reviewer for providing helpful comments to the manuscript.
Mark McGrouther of the Australian Museum kindly photographed the
acanthidae with 102 species in 27 genera, Aracanidae with holotype of Acanthostracion bucephalus. I also thank Hiromitsu Endo
13 species in six genera, Ostraciidae with 22 species in five and Naohide Nakayama of BSKU, Hiroyuki Motomura and his stu-
genera, monotypic Triodontidae, Tetraodontidae with 184 dents of KAUM, Eri Katayama of NSMT, and Hideki Sugiyama, for
123
providing me with beautiful photographs of tetraodontiforms. This Abe T (1949b) Taxonomic studies on the puffers (Tetraodontidae,
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Researches, A (19208019) and B (24370041) from the Japan Society caudal fin. Bull Jpn Soc Sci Fish 15:19-27
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Science. pectoral fin (with some additions to the previous reports of the
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Open Access This article is distributed under the terms of the Abe T (1952) Taxonomic studies on the puffers (Tetraodontidae,
Creative Commons Attribution License which permits any use, dis- Teleostei) from Japan and adjacent regions—VII. Concluding
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author(s) and the source are credited. Boesemanichthys. Jpn J Ichthyol 1:35-44
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Ichthyol Res (2015) 62:72–113

Taxonomy and systematics of tetraodontiform fishes: a review

a carapace, and pelvic fins that are reduced or absent (Tyler

Table 1 A list of 23 species of the Triacanthodidae in the world

Hollardiinae Hollardia goslinei Tyler 1968 Hawaii

Table 2 A list of 7 species of the Triacanthidae in the world

Table 3 A list of 37 species of the Balistidae in the world

Table 4 A list of 102 species of the Monacanthidae in the world

Eubalichthys caeruleoguttatus Hutchins Northern Western Australia

Pervagor spilosoma (Lay and Bennett 1839) Hawaiian Islands

Table 5 Distributions of 13 species of the Aracanidae in the world

Anoplocapros amygdaloides Fraser-Brunner 1941d Western Australia and South Australia

Fig. 2 Kentrocapros eco

Table 6 Distributions of 22 species of the Ostraciidae in the world

Böhlke and Chaplin (1968) provided keys and accounts

Fig. 4 Two forms of

Dorsal-fin rays Amblyrhynchote Bibron in Duméril 1855 1

Fig. 5 Pufferfishes found in a

Fig. 6 A large specimen of

(1980). Morphological synapomorphies of tetraodontiforms On the basis of a comprehensive myological study,

phylogenetic relationships of the Tetraodontiformes by an

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