Ichthyological Research

, Volume 62, Issue 1, pp 72–113 | Cite as

Taxonomy and systematics of tetraodontiform fishes: a review focusing primarily on progress in the period from 1980 to 2014

Open Access
REVIEW FOR IPFC9 SPECIAL ISSUE Ichthyology and Indo-Pacific Fish Conferences from the 1980s to the 2010s
Part of the following topical collections:
  1. Ichthyology and Indo-Pacific Fish Conferences from the 1980s to the 2010s

Abstract

When the first Indo-Pacific Fish Conference (IPFC1) was held in Sydney in 1981, there were still many problems in the generic- and species-level taxonomy of all tetraodontiform families except for the recently reviewed Triacanthodidae and Triacanthidae. The period from IPFC1 to IPFC9 (1981−2013) was a time of great progress in the taxonomy and systematics of the Tetraodontiformes: many review and revisional papers have been published for various genera and species, with descriptions of many new taxa occurring mainly on coral reefs and in tropical freshwaters; and cladistic analyses of morphological characters have been performed to clarify phylogenetic relationships of various families and molecular analyses have greatly progressed to provide detailed phylogenetic relationships of families, genera, and even species. The purpose of this paper is to provide a review on developments in the taxonomy and systematics of the Tetraodontiformes, focusing primarily on contributions since 1980 (when James C. Tyler’s monumental work was published) through the period of IPFCs, including pertinent publications before 1980. This paper recognizes 412 extant species in the 10 families of living Tetraodontiformes, with the allocation of species and genera as follows: Triacanthodidae including 23 species in 11 genera, Triacanthidae seven species in four genera, Balistidae 37 species in 12 genera, Monacanthidae 102 species in 27 genera, Aracanidae 13 species in six genera, Ostraciidae 22 species in five genera, Triodontidae monotypic, Tetraodontidae 184 species in 27 genera, Diodontidae 18 species in seven genera, and Molidae five species in three genera. Phylogenetic relationships of the families have been clarified by morphological and molecular analyses and have provided well-supported sister relationships of the families: Triacanthodidae and Triacanthidae, Balistidae and Monacanthidae, and Tetraodontidae and Diodontidae. However, there remain problems with the phylogenetic positions of the Triodontidae and Molidae due to conflicts of differing positions in morphological and molecular studies (e.g., Molidae has been placed differently among molecular studies).

Keywords

Tetraodontiformes Classification Morphology Molecular Phylogenetic relationships 

Introduction

Tetraodontiform fishes are distributed in tropical to temperate 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 Fowler 1902 and a small pufferfish of the genus Carinotetraodon Benl 1957 mature at about 2 cm in total length (TL) (Lim and Kottelat 1995; Tyler 1970), whereas ocean sunfishes of the genus Mola Koelreuter 1766 attain over 300 cm TL (Yoshita et al. 2009). Tetraodontiform fishes are characterized by a small mouth with either relatively few teeth that are often enlarged or massive beak-like tooth plates, a small gill opening restricted to the side of the body, scales usually modified as spines, enlarged plates, or a carapace, and pelvic fins that are reduced or absent (Tyler 1980; Nelson 2006). In addition, many pufferfishes are characterized by having a strong toxin in the viscera and skin, and even in the musculature of some species of Lagocephalus Swainson 1839 (Matsuura 1984). Because of their peculiar morphological characters, tetraodontiforms have long attracted the attention of ichthyologists and biologists.
Fig. 1

Representatives of the 10 extant families of Tetraodontiformes. a Triacanthodidae, Triacanthodes anomalus; b Triacanthidae, Triacanthus biaculeatus; c Balistidae, Abalistes filamentosus; d Monacanthidae, Thamnaconus hypargyreus; e Aracanidae, Kentrocapros aculeatus; f Ostraciidae, Ostracion immaculatus; g Triodontidae Triodon macropterus; h Tetraodonitidae, Arothron mappa; i Diodontidae, Diodon liturosus; j Molidae, Masturus lanceolatus. Photographs of a and e provided by BSKU; b, d, f, h, and i by KAUM; c and g by NSMT; j by Hideki Sugiyama

Since Cuvier (1816) classified tetraodontiforms in the order Plectognathi, tetraodontiforms were usually considered to form a monophyletic group among the advanced percomorph fishes (Nelson 2006). Although the taxonomy of tetraodontiform fishes progressed greatly in the nineteenth century (see Tyler 1980, for a history of the classification of the order), information about tetraodontiform taxonomy was scattered in many articles, making it difficult to understand the taxonomic relationships of tetraodontiforms overall. Alec Fraser-Brunner published an important series of articles on various groups of tetraodontiforms as reviews of genera and families during the mid-1930s to early 1950s: however, many parts of his publications were based on cursory examinations of relatively few specimens. There were no comprehensive phylogenetic studies until Winterbottom (1974) and Tyler (1980) provided their interpretations of the phylogenetic relationships of tetraodontiforms based on myology and osteology, respectively.

When the first Indo-Pacific Fish Conference (IPFC1) was held in Sydney in 1981, many problems remained in the generic- and species-level taxonomy of all tetraodontiform families, except for the Triacanthodidae and Triacanthidae for which Tyler (1968) had provided a monographic revision. The period from IPFC1 to IPFC9 (1981−2013) was a time of great progress in the taxonomy and systematics of tetraodontiforms. Many review and revisional papers have been published for various genera and species, with descriptions of new taxa found mainly in coral reefs and tropical freshwaters; and cladistic analyses of morphological characters have clarified phylogenetic relationships of a number of families and molecular analyses greatly assisted our understanding of the detailed phylogenetic relationships of families, genera, and even species. The purpose of this paper is to present a review of the developments in the taxonomy and systematics of the Tetraodontiformes, focusing primarily on contributions from 1980 (when James C. Tyler’s monumental work was published) through the period of IPFCs, but including pertinent publications before 1980. This paper is composed of two parts, the first reviewing taxonomic studies and the second focusing of studies on systematics. Institutional abbreviations follow Fricke and Eschmeyer (2014).

Triacanthodidae (Spikefishes, Fig. 1a; Table 1)

Spikefishes of the family Triacanthodidae are usually found in depths of 100−600 m on continental shelves and slopes (Tyler 1968; Matsuura and Tyler 1997). They are easily distinguished externally from other families of the order Tetraodontiformes by the following combination of characters: body deep and slightly compressed, covered by moderately thick skin with numerous small scales not readily distinguishable to the unaided eye, with each scale bearing upright spinules and having a roughly shagreen-like appearance; two separate dorsal fins, six spines in the first dorsal fin and 12−18 soft rays in the second dorsal fin; caudal fin rounded to almost truncate, not forked; most dorsal-, anal-, and pectoral-fin rays branched; pelvic fins with a large spine and one or two inconspicuous and rudimentary soft rays; mouth small and usually terminal; teeth of moderate size, usually conical, 10 or more in an outer series in each jaw; caudal peduncle compressed, deeper than wide, not distinctly tapered (Matsuura 2001).
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 1941)

Off Tanzania, Maldive Islands, Japan, and Celebes Sea

 

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 1899

Arabian Sea and Australia

 

Johnsonia eriomma Myers 1934

Bahamas to Greater and Lesser Antilles and western Caribbean Sea

 

Macrorhamphosodes platycheilus Fowler 1934

Bay of Bengal, Philippines, and Australia

 

Macrorhamphosodes uradoi (Kamohara 1933)

South Africa, off Kenya, Japan, Korea, East China Sea, South China Sea, 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 1934

East Africa, Japan, Kyushu-Palau Ridge, South China Sea, Australia, and New Caledonia

 

Triacanthodes anomalus (Temminck and Schlegel 1850)

Japan, Korea, East China Sea, South China Sea

 

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 and Fourmanoir 1984

New Caledonia

 

Tydemania navigatoris Weber 1913

East Africa, Bay of Bengal, Japan, China, Philippines, Indonesia, and Australia

Spikefishes are still relatively poorly known in terms of taxonomy and biology among tetraodontiform families, although Tyler (1968) published an excellent monograph on the superfamily Triacanthoidea including the Triacanthodidae and Triacanthidae. He recognized 19 triacanthodid species in 11 genera (Table 1): Atrophacanthus Fraser-Brunner 1950 (one species), Bathyphylax Myers 1934 (two species), Halimochirurgus Alcock 1899 (two species), Hollardia Poey 1861 (three species), Johnsonina Myers 1934 (one species), Macrorhamphosodes Fowler 1934 (two species), Mephisto Tyler 1966b (one species), Parahollardia Fraser-Brunner 1941b (two species), Paratriacanthodes Fowler 1934 (two species), Triacanthodes Bleeker 1857 (two species), and Tydemania Weber 1913 (one species). Because all these genera and species were described in detail by Tyler (1968), their taxonomic features are not repeated here, except for new information that provides a better understanding of taxa.

Many species of the Triacanthodidae are known from the tropical and warm regions of the Indo-Pacific. However, five species are distributed in the western Atlantic: Hollardia hollardi Poey 1861, Hollardia meadi Tyler 1966a, Johnsonina eriomma Myers 1934, Parahollardia lineata (Longley 1935), and Parahollardia schmidti Woods 1959. Hollardia goslinei Tyler 1968 are known only from nine specimens collected from the Hawaiian Islands.

Little information about the Atlantic and Hawaiian spikefishes has been published since Tyler’s (1968) monograph. Matsuura (1983) provided a brief morphological account of Hollardia hollardi with a color photograph based on specimens collected from off Surinam. McEachran and Fechhelm (2005) also provided a brief account and line drawing of Hollardia hollardi and Hollardia meadi from the Gulf of Mexico. McEachran and Fechhelm (2005) and Hartel et al. (2008) reported briefly Parahollardia lineata from the Gulf of Mexico and off greater New England, respectively. Tyler et al. (2013) documented a large northern range extension for Hollardia hollardi to the northeast coast of the USA off Massachusetts, whereas this species had previously only been known from the Florida Keys, Bahamas, Bermuda, Gulf of Mexico, Caribbean, and south to Brazil, with the new northern occurence perhaps associated with warming currents along the east coast of North America.

In contrast to the Atlantic spikefishes, many papers on Indo-Pacific spikefishes were published after 1980. Matsuura (1982) described Triacanthodes indicus based on 13 specimens collected from the western Indian Ocean. This species is characterized by relatively large nasal organs compared to other species of Triacanthodes. Matsuura and Fourmanoir (1984) described Triacanthodes intermedius based on two specimens collected from New Caledonia. Triacanthodesintermedius is unique among spikefishes in having several intermediate characters between Paratriacanthodes and Triacanthodes. Tyler (1997) described Paratriacanthodes abei based on a single specimen collected in the South China Sea. Santini (2006) described Bathyphylax pruvosti based on 25 specimens collected from the Marquesas Islands. In addition to these contributions, papers and books on taxonomic and zoogeographical studies of spikefishes were published by authors from various countries including Australia, China, Japan, Korea, Russia, Taiwan, Ukraine, and the USA.

Amaoka (1982) reported Halimochirurgus alcocki Weber 1913, Macrorhamphosodes uradoi (Kamohara 1933), and Paratriacanthodes retrospinis Fowler 1934 with detailed descriptions and color photographs of fresh specimens collected from the Kyushu-Palau Ridge. Matsuura (1984) provided brief accounts and color photographs of tetraodontiform fishes found in the Japanese Archipelago, including Atrophacanthus japonicus Kamohara 1941, Halimochirurgus alcocki, Macrorhamphosodes uradoi, Triacanthodes anomalus (Temminck and Schlegel 1850), and Triacanthodes ethiops Alcock 1894. In the following year, Matsuura (1985) provided detailed accounts and a color photograph of Triacanthodes anomalus collected from the Okinawa Trough. Matsuura (1987) and Stewart and Clark (1988) recorded Macrorhamphosodes uradoi from New Zealand, a species previously known only from Japan (Kamohara 1933; Tyler 1968; Matsuura 1984), Kenya (Tyler 1983) and South Africa (Hulley 1972). Matsuura and Tyler (1997) reported Bathyphylax bombifrons Myers 1934, Halimochirurgus alcocki, Macrorhamphosodes uradoi, Paratriacanthodes retrospinis, Triacanthodes ethiops, and Triacanthodes intermedius based on many specimens collected from New Caledonia. Except for Triacanthodes intermedius, these are the first records for the species from New Caledonia. Chen et al. (1997) recorded Triacanthodes anomalus from the South China Sea. Matsuura (2000) compiled a checklist of Tetraodontiformes from the South China Sea including eight species of spikefishes: A. japonicus, B. bombifrons, Halimochirurgus alcocki, Macrorhamphosodes platycheilus Fowler 1934, Paratriacanthodes retrospinis, Triacanthodes anomalus, Triacanthodes ethiops, and Tydemania navigatoris Weber 1913.

Myers and Donaldson (2003) published a checklist of fishes from the Mariana Islands including a record of Halimochirurgus alcocki. Kim et al. (2005) provided brief accounts and color photographs of Triacanthodes anomalus and Macrorhamphosodes uradoi from Korea. Hoese et al. (2006) recorded eight species of spikefishes for the first time from Australia: B. bombifrons (New South Wales and Queensland), Halimochirurgus alcocki (New South Wales, Queensland and Western Australia), Halimochirurgus centriscoides (Western Australia), Macrorhamphosodes platycheilus (Northern Territory and Western Australia), Macrorhamphosodes uradoi (New South Wales and Queensland), Paratriacanthodes retrospinis (New South Wales and Western Australia), Triacanthodes ethiops (Northern Territory, Queensland, Western Australia), and Tydemania navigatoris (Western Australia). Shao et al. (2008) recorded seven species of spikefishes from off southern Taiwan in the South China Sea: B. bombifrons, Halimochirurgus alcocki, Halimochirurgus centriscoides, Macrorhamphosodes uradoi (first record from the South China Sea), Paratriacanthodes retrospinis, Triacanthodes anomalus, and Tydemania navigatoris. All species records of Shao et al. (2008) were based on specimens deposited at museums and institutions in Taiwan and the USA. Larson et al. (2013) published an annotated checklist of the fishes of the Northern Territory, Australia, in which they recorded Halimochirurgus centriscoides, Macrorhamphosodes platycheilus, Triacanthodes ethiops, and Tydemania navigatoris.

Tyler (1983) reported four spikefishes from off Kenya in the western Indian Ocean: B. bombifrons, Bathyphylax omen Tyler 1966c, Macrorhamphosodes uradoi, and Macrorhamphosodes platycheilus. When Tyler (1968) compared B. bombifrons and B. omen, the main differences were that B. bombifrons has a narrower pelvis, a shorter postorbital length, a less distinctly supraterminal mouth, and a more concave snout profile. Because the holotype of B. omen (37.5 mm in standard length, SL) from the western Indian Ocean is much smaller than that of B. bombifrons (77.6 mm SL) from off Hong Kong in the South China Sea, some of these differences were considered to be due in part to the size differences of the holotypes (Tyler 1968). Based on four additional, newly collected specimens of B. bombifrons (80.9−84.0 mm SL) and two specimens of B. omen (67.0−93.5 mm SL), Tyler (1983) concluded that the most reliable character separating the two species is the degree of concavity of the snout. The depth of the snout in the middle of its length is 10.8−13.4 % SL (average 12.0) in B. bombifrons and 12.5−15.5 % SL (average 13.9) in B. omen. Matsuura and Tyler (1997) confirmed this difference in three specimens (86.2−93.0 mm SL) of B. bombifrons collected off New Caledonia.

Tyler (1986) provided keys to the genera and species of spikefishes known from South and East Africa. He recognized 11 species in this region: A. japonicus, B. bombifrons, B. omen, Halimochirurgus alcocki, Halimochirurgus centriscoides, Macrorhamphosodes platycheilus, Macrorhamphosodes uradoi, Mephisto fraserbrunneri, Paratriacanthodes retrospinis, Triacanthodes ethiops, and Tydemania navigatoris. Adam et al. (1998) recorded A. japonicus and Triacanthodes ethiops for the first time from the Maldive Islands in the central Indian Ocean. Atrophacanthus japonicus had previously been recorded only from southern Japan, the Celebes Sea, and off Tanzania (Kamohara 1941; Fraser-Brunner 1950; Tyler 1968; Matsuura 1984). Although T. ethiops is relatively common in the West Pacific and New Caledonia (Tyler 1968; Matsuura and Tyler 1997), it had previously been known only from East Africa in the Indian Ocean (Tyler 1968, 1986). Manilo and Bogorodsky (2003) studied coastal fishes of the Arabian Sea based on specimens collected by research vessels of Russia and Ukraine from 1967 to 1991. They also examined specimens collected by the German RV Meteor, which are deposited at the Zoological Museum of Hamburg University. The two authors recorded Halimochirurgus centriscoides and Mephisto fraserbrunneri Tyler 1966b (previously known only from the holotype, 52.2 mm SL from the Andaman Sea and a non-type specimen, 66.0 mm SL from off Somalia in the western Indian Ocean), but did not provide descriptions of the two species. The specimens they studied are deposited at the National Museum of Natural History, National Academy of Sciences, Ukraine, the Zoological Museum of the Russian Academy of Sciences, and the Zoological Museum of Moscow State University. In addition to these, Venkataramanujam et al. (1993) erroneously described a new solenostomid species, Solenostomus tuticoriensis based on a specimen of Macrorhamphosodes platycheilus from southern India (see Orr and Fritzsche 1997).

This brief historical review clearly shows that there are many collection lacunae with respect to the distribution and diversity of spikefishes in the world oceans. New Caledonia is one example of how collection efforts could provide us with better understanding of spikefish diversity. When Matsuura and Fourmanoir (1984) described Triacanthodes intermedius from New Caledonia, only two type specimens were available for their study. Long-term deep-water surveys by ORSTOM around New Caledonia added six species of spikefishes including 16 specimens of Triacanthodes intermedius to a taxonomic study on tetraodontiform fishes of this region by Matsuura and Tyler (1997). Many regions of the world oceans need to be surveyed, but the most promising areas would be waters in French Polynesia and the Indian Ocean coast of Sumatra and Java in Indonesia. Japanese and Indonesian fisheries agencies have recently implemented a joint survey along the Indian Ocean coast of Sumatra and Java. Their surveys have resulted in many interesting spikefishes, including two undescribed species and several rarely collected species, which will be published elsewhere (Matsuura and Kawai in preparation).

Triacanthidae (Triplespines, Fig. 1b; Table 2)

Triplespines of the family Triacanthidae are found on continental shelves in the Indo-West Pacific, usually just below the sea surface to 60 m depth (Tyler 1968). They clearly differ from other families of the order Tetraodontiformes in having the following combination of characters: moderately elongate, strongly compressed body; skin moderately thick with numerous scales not easily discernible to the unaided eye, with each scale bearing upright spinules and having a rough, shagreen-like appearance; two separate dorsal fins, six spines (usually only five spines visible, the sixth rudimentary) in the first dorsal fin and 20−26 soft rays in the second dorsal fin; pelvic fin with a large spine and no visible soft rays; mouth small and usually terminal; teeth in jaws with an outer series of about 10 heavy incisors and an internal series of several molars, usually four in the upper jaw and two in the lower jaw; caudal fin deeply forked; caudal peduncle distinctly tapering to a narrow transversely indented region just in front of the caudal-fin base where the peduncle is wider than deep (Matsuura 2001).
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 his monograph of the superfamily Triacanthoidea, Tyler (1968) recognized seven species (Table 2): Pseudotriacanthus strigilifer (Cantor 1849), Triacanthus biaculeatus (Bloch 1786), Triacanthus nieuhofii Bleeker 1852b, Tripodichthys blochii (Bleeker 1852a), Tripodichthys angustifrons (Hollard 1854), Tripodichthys oxycephalus (Bleeker 1851c), and Trixiphichthys weberi (Chaudhuri 1910). He appeared to have resolved all taxonomic problems in the Triacanthidae, except for two unusual specimens of Triacanthus biaculeatus collected from the Gulf of Thailand. Tyler (1968) reported that four specimens from the Gulf of Thailand near Paknam have differences in eye size and the width of the interorbital space. Two of these specimens (111.5 mm SL and 113.9 mm SL) have exceptionally large eyes (10.4 % SL) and the other two (105.6 mm SL and 107.9 mm SL) have a normal eye size (7.8 and 8.5 % SL). The two large-eye specimens have eyes that are also larger than those of any other specimens of equivalent size. The large-eye specimens have a wider interorbital space (10.2−10.5 % SL vs. 6.3−10.0 % SL in many other specimens). Tyler (1968) stated that they could be an undescribed species, but tentatively considered them as abnormal Tiracanthus biaculeatus. Although I have been to the Gulf of Thailand many times and examined local collections in Thailand, I have not seen specimens of Triacanthus with such large eyes.

Tyler (1968) provided color descriptions of P. strigilifer, Triacanthus biaculeatus, and Trixiphichthys weberi based on color notes or color photographs of fresh specimens, but neither was available for the other species. Matsuura (2009) included a color photograph of a juvenile specimen (50 mm SL) of Triacanthus nieuhofii collected from Phuket in the Andaman Sea: body silvery white with light brown tinge on dorsal half of body; first dorsal spine silvery white with a black membrane between the first and third dorsal spines; second dorsal, anal, and pectoral fins with light yellowish rays; caudal fin dark yellow. Matsuura (2013) provided brief accounts and a color photograph of Tripodichthys blochii collected from the northern Gulf of Thailand: body silvery white with several irregular golden brownish markings on head and body; basal half of first dorsal spine silvery white and proximal half black with yellowish orange first dorsal fin membrane; second dorsal, anal, and pectoral fins with light yellowish rays; caudal fin dusky yellow.

Distribution ranges of triplespines were well documented by Tyler (1968). After his monograph appeared, many publications on fish faunas, field guides and checklists have provided additional information on triplespine distributions (e.g., Matsuura 1984, 2003a, 2009, 2011, 2013; Allen and Swainston 1988; Talwar and Jhingran 1991; Kottelat et al. 1993; Randall 1995; Allen 1997; Chen et al. 1997; Fricke 1999; Johnson 1999; Matsuura and Peristiwady 2000; Hutchins 2001b; Hoese et al. 2006; Kottelat 2013; Larson et al 2013). Although most of these publications did not greatly extend the previously reported distributions of triplespines, Fricke (1999) recorded Triacanthus biaculeatus from the Mascarene Islands and Kottelat et al. (1993) reported a 60-mm SL specimen of this species from freshwater in Indonesia.

Balistidae (Triggerfishes, Fig. 1c; Table 3)

Triggerfishes of the family Balistidae occur in shallow waters, mainly on coral reefs around the world usually from just below the sea surface to 50 m depth. They differ externally from other families of the order Tetraodontiformes by the following combination of characters: body deep, moderately compressed, encased in very thick, tough skin with large scales easily discernible as individual units; scales above pectoral-fin base in many species enlarged, forming a flexible tympanum; mouth small and terminal, or almost terminal; teeth strong, eight in the outer series of both jaws; gill opening, a moderately short, vertical to oblique slit in front of pectoral-fin base; two dorsal fins, first dorsal fin with three visible spines, the second spine more than 1/2 length of first spine; first spine capable of being locked in an erect position by second spine; second dorsal and anal fin similar in shape; most dorsal-, anal-, and pectoral-fin rays branched; pelvic fins rudimentary, represented by a series of four pairs of enlarged scales encasing posterior end of pelvis (Matsuura 2001).
Table 3

A list of 37 species of the Balistidae in the world

Species

Distribution

Abalistes filamentosus Matsuura and Yoshino 2004

Ryukyu Islands, North West Shelf of Australia, and Timor Sea

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 Schneider 1801)

East Africa eastward through northern Australia to Samoa, northward to southern Japan

Balistoides viridescens (Bloch and Scneider 1801)

Red Sea, East Africa eastward through northern Australia to Line and Tuamotu islands, 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 1973)

Red Sea, East Africa eastward to western Thailand and Sumatra, southern Java, and Bali

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 1829)

Red Sea, East Africa eastward through northern Australia to Tuamotu Islands, northward to southern Japan

Pseudobalistes fuscus (Bloch and Schneider 1801)

Red Sea, East Africa eastward through northern Australia to Society Islands, northward to southern Japan

Pseudobaliste naufragium (Jordan and Starks 1895)

Eastern Pacific from Baja California to Ecuador

Rhinecanthus abyssus Matsuura and Shiobara 1989

Known only from Kume-jima Island, Ryukyu Islands

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 1983

Queensland to Pitcairn Islands

Rhiencanthus rectangulus (Bloch and Schneider 1801)

Red Sea, East Africa eastward through northern Australia to Hawaiian and Pitcairn islands, 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, northward to southern Japan

Sufflamen chrysopterum (Bloch and Schneider 1801)

East Africa eastward through northern Australia to Samoa, northward to southern Japan

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, Matsuura and Zama 1978

Angela Islands in western Indian Ocean eastward through Indonesia to Galapagos Islands, northward to southern Japan

Xanthichthys lineopunctatus (Hollard 1854)

South Africa eastward through northern Australia to New Guinea, northward to southern Japan

Xanthichthys mento (Jordan and Gilbert 1882)

Antitropical in Pacific, southern Japan, Hawaiian Islands, Clipperton Island, Revillagigedo 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 1983

South Africa

Xenobalistes tumidipectoris Matsuura 1981

Mariana Islands

Triggerfishes of the Balistidae have not yet been comprehensively reviewed, but Matsuura (1979) described the osteology of family and its generic relationships, and Matsuura (1980) diagnosed and illustrated the 20 species of 10 genera found in Japanese waters. His latter paper covered many of the 25 species that were known from the Indo-Pacific before 1980. Prior to this publication two papers reviewed triggerfishes in the eastern Pacific (Berry and Baldwin 1966) and those of the western Atlantic (Moore 1967a). In addition, Randall and Klausewitz (1973) reviewed Melichthys Swainson 1839, with a description of Melichthys indicus, and Randall et al. (1978) revised Xanthichthys Kaup in Richardson 1856 with a description of Xanthichthys caeruleolineatus. These were important contributions about the taxonomy of triggerfishes up to and including 1980. A brief historical review of triggerfish taxonomy follows.

Fraser-Brunner (1935a) published a review of the Balistidae, one of his series of taxonomic studies of tetraodontiform fishes. He recognized 13 genera in the Balistidae and provided a platform for triggerfish taxonomy. Although de Beaufort and Briggs (1962) lumped Fraser-Brunner’s 11 genera into the single genus Balistes Linnaeus 1758, they relegated all 11 genera to subgenera. Berry and Baldwin (1966) reviewed triggerfishes of the eastern Pacific with comments on Fraser-Brunner’s classification. They pointed out that Fraser-Brunner (1935a) recognized Nematobalistes Fraser-Brunner 1935a and Verrunculus Jordan 1924 on trivial differences of squamation and the anterior few soft rays of the dorsal fin. Berry and Baldwin (1966) examined many individuals of the eastern Pacific triggerfishes, which diminished differences among Balistes, Nematobalistes, and Verrunculus. It led Berry and Baldwin (1966) to conclude that the latter two are junior synonyms of Balistes. They partially accepted Fraser-Brunner’s system by recognizing six genera: Balistes, Canthidermis Swainson 1839, Melichthys, Pseudobalistes Bleeker 1865–1869, Sufflamen Jordan 1916, and Xanthichthys, but they left the other five genera, Abalistes Jordan and Seale 1906, Balistapus Tilesius 1820, Balistoides Fraser-Brunner 1935a, Odonus Gistel 1848, and Rhinecanthus Swainson 1839 to the deliberations of subsequent authors. Just after Berry and Baldwin’s (1966) paper appeared, Moore (1967a) published a review of the triggerfishes of the western Atlantic and recognized four genera, Balistes, Canthidermis, Melichthys, and Xanthichthys (i.e., triggerfishes of other genera not occurring in the western Atlantic). Just after his review of the western Atlantic triggerfishes, Moore (1967b) clarified the taxonomic status of Balistes punctatus Gmelin 1789, stating that it occurs only in the eastern Atlantic.

Matsuura (1980) provided keys to genera and species, and taxonomic accounts of 20 Japanese species that he classified in 10 genera, Abalistes, Balistapus, Balistoides, Canthidermis, Melichthys, Odonus, Pseudobalistes, Rhinecanthus, Sufflamen, and Xanthichthys. He also addressed a taxonomic problem in the genus Hemibalistes. Hemibalistes as proposed by Fraser-Brunner (1935a) was not an available name because two species, Sufflamen bursa (Bloch and Schneider 1801) and Sufflamen chrysopterum (Bloch and Schneider 1801), were included and neither was designated type species. Smith (1949b) subsequently designated Balistes bursa Bloch and Schneider 1801 as the type species of the genus Hemibalistes. However, the diagnostic character of Hemibalistes, all scales on cheek smaller than those on body, is not adequate to differentiate it from other triggerfish genera because squamation on the cheek of Sufflamen and Hemibalistes is variable (Matsuura 1980). As stated above, the papers of Berry and Baldwin (1966), Moore (1967a), Randall and Klausewitz (1973), Randall et al. (1978), and Matsuura (1980) contributed to the proper recognition of 11 genera, Abalistes, Balistapus, Balistes, Balistoides, Canthidermis, Melichthys, Odonus, Pseudobalistes, Rhinecanthus, Sufflamen, and Xanthichthys. Tyler (1980) also accepted these 11 genera in his phylogenetic study on tetraodontiform fishes. The osteologically based phylogenies of balistoids given by Matsuura (1979) and Tyler (1980) were largely in agreement, as were the majority of their morphological description; the few conflicts between their respective accounts were clarified by Tyler and Matsuura (1981) as an aid to subsequent workers.

Matsuura (1981) described Xenobalistes tumidipectoris as a new genus and species based on a small juvenile collected from stomach contents of a marlinfish, Makaira mazara (Jordan and Snyder 1901), captured in the Mariana Islands. Although the holotype of Xenobalistes tumidipectoris was partially damaged by digestion, it differs clearly from other triggerfishes by its possession of a number of unique characters, such as there is a large lateral expansion protruding laterally just below the pectoral fin, the supraorbital ridge is well developed and convex dorsolaterally, and the mid-lateral portion of the coracoid is greatly expanded to form a disk-like bone. Two years later, Heemstra and Smith (1983) described Xenobalistes punctatus as a new species, based on a small juvenile washed up on a beach at the mouth of the Van Stadens River, South Africa.

Randall and Steene (1983) described Rhinecanthus lunula and provided brief comments about the other five species, Rhinecanthus aculeatus (Linnaeus 1758), Rhinecanthus assasi (Forsskål 1775), Rhinecanthus cinereus (Bonnaterre 1788), Rhinecanthus rectangulus (Bloch and Schneider 1801), and Rhinecanthus verrucosus (Linnaeus 1758). Matsuura and Shiobara (1989) described Rhinecanthus abyssus on the basis of two specimens collected off the northeast coast of Kume-jima in the Ryukyu Islands, at depths of 120−150 m, which is quite deep for triggerfishes. Matsuura and Yoshino (2004) studied the monotypic genus Abalistes and recognized two species, Abalistes stellatus (Anonymous 1798) and a new species, Abalistes filamentosus.

Taxonomic studies of triggerfishes continued during the period from Berry and Baldwin (1966) to Matsuura and Yoshino (2004), resulting in the classification shown in Table 3, which recognizes 37 species of triggerfishes in 11 genera. In addition to the above contributions, many checklists and illustrated books on shallow-water fishes also provided taxonomic and zoogeographical information of the Balistidae and other tetraodontiforms (Böhlke and Chaplin 1968; Jones and Kumaran 1980; Kyushin et al. 1982; Gloerfelt-Tarp and Kailola 1984; Matsuura 1984, 2000, 2001, 2002, 2003a, 2009, 2011, 2013; Sainsbury et al. 1984; Smith and Heemstra 1986; Tortonese 1986; Allen and Swainston 1988; Winterbottom et al. 1989; 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 2003; Munday 2005; Shao et al. 2008; Fricke et al. 2009; Allen and Erdman 2012). Still, taxonomic challenges remain. Sufflamen verres (Gilbert and Starks 1904) in the eastern Pacific is said to be different from the Indo-West Pacific species, Sufflamen fraenatum (Latreille 1804), in having higher counts of fin rays of the second dorsal and anal fins (Berry and Baldwin 1966). According to their study, S. verres has 30−33 dorsal rays and 27−30 anal rays, while S. fraenatum has 28−30 dorsal rays and 24−26 anal rays. However, Berry and Baldwin (1966) also noted that a specimen of S. fraenatum from South Africa with 31 dorsal rays and 27 anal rays, overlapping the counts of S. verres. Matsuura (1980) showed on the basis of 22 specimens of S. fraenatum from Japan that dorsal rays range from 27 to 30 and anal rays from 24 to 27. Randall (2010) stated that the Hawaiian population of S. fraenatum has 27−31 dorsal rays and 24−28 anal rays. These data strongly suggest that S. verres is a junior synonym of S. fraenatum. Molecular comparisons of S. verres and S. fraenatum may help clarify their relationships. Fricke (1999) stated that Xanthichthys lineopunctatus (Hollard 1854) is a synonym of Xanthichthys lima (Bennett 1832). However, the original description of Balistes lima by Bennett (1832) comprised only 50 words with fin ray counts “D. 1?, A. 26, P. 13.” Because the poor original description fits several species of the Balistidae, it is impossible to verify that Balistes lima is conspecific with Xanthichthys lineopunctatus. As no type specimens are known for Balistes lima, I conclude that Balistes lima should be treated as a nomen dubium.

Santini et al. (2013a) showed Xenobalistes tumidipectoris to be deeply nested among five species of Xanthichthys in a phylogenetic tree of the Balistidae based on molecular analysis. Their molecular analysis suggests that Xenobalistes is a junior synonym of Xanthichthys. I am currently comparing larger specimens of Xenobalistes tumidipectoris and Xenobalistes punctatus with those of Xanthichthys to clarify the relationships of the two species of Xenobalistes with the five species of Xanthichthys. This study will be published elsewhere to resolve the taxonomic status of Xenobalistes. Santini et al. (2013a) also mentioned that Balistoides and Pseudobalistes are not monophyletic: Balistoides conspicillum (Bloch and Schneider 1801) is sister to Melichthys, and Pseudobalistes flavimarginatus (Rüppell 1829) forms a clade with Balistoides viridescens (Bloch and Schneider 1801), whereas Pseudobalistes fuscus (Bloch and Schneider 1801) and Pseudobalistes naufragium (Jordan and Starks in Jordan 1895) represent a single clade. These species should be revisited with detailed morphological examinations.

Monacanthidae (Filefishes, Fig. 1d; Table 4)

Filefishes of the family Monacanthidae occur on coral and rocky reefs and in sea grass beds, shallower than 200 m, but some species are found at depths reaching almost 300 m (Matsuura and Tyler 1997; Hutchins 2001b). They differ externally from other families of the order Tetraodontiformes by the following combination of characters: body usually deep, strongly compressed, body shape varying from oblong to almost circular; skin smooth to rough, shagreen-like, with minute to small scales armed with one to many fine spinules, spinules enlarged in some species forming bristles or spines on the posterior portion of the body; scales on head of some species with strong flattened spinules; mouth small, generally terminal, non-protractile; teeth pointed and not fused together, central pair usually the largest in each jaw; gill opening, a short vertical to oblique slit in front of, or above, pectoral-fin base; two dorsal fins, first dorsal fin consisting of a prominent spine, which can be locked upright by a second very small spine, second dorsal fin with 22 to 52 simple (unbranched) soft rays; anal fin with 20 to 62 simple soft rays; pelvic fin and spines rudimentary or absent, represented by a series of three or fewer pairs of enlarged scales encasing the pelvic terminus, or segments of indeterminate number, or entirely absent; pelvis usually capable of vertical movement giving rise to a ventral flap (Hutchins 2001b; Matsuura 2003b).
Table 4

A list of 102 species of the Monacanthidae in the world

Species

Distribution

Acanthaluteres brownii (Richardson 1848)

Southern Western Australia and South Australia

Acanthaluteres spilomelanurus (Quoy and Gaimard 1824)

Southern Western Australia, South Australia, Victoria, Tasmania, and southern New South Wales

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 Gaimard 1824)

Southern Western Australia, South Australia, Victoria, Tasmania, and New South Wales

Brachaluteres taylori Woods in Schultz et al. 1966

Indonesia eastward to Papua New Guinea and eastern Australia, northward to Philippines

Brachaluteres ulvarum Jordan and Fowler 1902

Japan

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 Playfair and Günther1867)

East Africa eastward through Indonesia to New Caledonia, northward to southern Japan

Cantherhines longicaudus Hutchins and Randall 1982

Tahiti, Cook Islands, New Caledonia, and Tuamotu Islands

Cantherhines macroceros (Hollard in Guichenot 1853)

Western Atlantic and St. Paul’s Rocks, Cape Verde Islands

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, northward to southern Japan

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 Gaimard 1824)

Hawaiian Islands, Johnston Island, and Cook Islands

Cantherhines verecundus Jordan 1925

Hawaiian Islands

Cantheschenia grandisquamis Hutchins 1977

Western Australia, New South Wales, and Lor Howe Island

Cantheschenia longipinnis (Fraser-Brunner 1941c)

Southern Western Australia and northern New South Wales

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

Eubalichthys caeruleoguttatus Hutchins 1977

Northern Western Australia

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 1886)

Southern Western Australia, South Australia, Victoria, Tasmania, New South Wales, and 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 1824)

Southern Western Australia, South Australia, Tasmania, and southern New South Wales

Meuschenia galii (Waite 1905)

Southern Western Australia, South Australia, and western Victoria

Meuschenia hippocrepis (Quoy and Gaimard 1824)

Southern Western Australia, South Australia, Victoria, and northeastern Tasmania

Meuschenia scaber (Forster in Bloch and Schneider1801)

Southern Western Australia, South Australia, Tasmania, and southern New South Wales

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 Schneider 1801)

East Africa eastward through northern Australia to Samoa, northward to southern Japan

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 1853)

Thailand eastward through Indonesia to Papua New Guinea

Paramonacanthus frenatus (Peters 1855)

East Africa and Seychelles

Paramonacanthuys lowei Hutchins 1997

Eastern Australia

Paramonacanthus matsuurai Hutchins 1997

Ogasawara Islands

Paramonacanthus nematophorus (Günther 1870)

Red Sea, East Africa, and Seychelles

Paramonacanthus oblongus (Temminck and Schlegel 1850)

Indonesia through northern Australia to New Caledonia, northward to southern Japan

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

Pervagor spilosoma (Lay and Bennett 1839)

Hawaiian Islands

Pseudalutarius nasicornis (Temminck and Schlegel 1850)

East Africa eastward through northern Australia to New Caledonia, northward to southern Japan

Pseudomonacanthus elongatus Fraser-Brunner 1940

Australia

Pseudomonacanthus macruru (Bleeker 1856)

Singapore, Malaysia, Brunei, Indonesia, Philippines, and Japan

Pseudomonacanthus peroni (Hollard 1854)

Australia and New Guinea

Pseudomonacanthus tweediei Fraser-Brunner 1940

Singapore

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 1790)

Southern Western Australia, South Australia, Victoria, northern Tasmania, New South Wale, and southern Queensland

Stephanolepis auratus (Castelnau 1861)

Eastern Western Australia, South Australia, Victoria, and Tasmania

Stephanolepis cirrhifer (Temminck and Schlegel 1850)

Japan, Korea, Taiwan, China, and northern Philippines

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 Matsuura 1984

Fiji

Thamnaconus garretti (Fowler 1928)

Hawaiian Islands

Thamnaconus hypargyreus (Cope 1871)

Western Pacific and southeastern Indian Ocean

Thamnaconus melanoproctes (Boulenger 1889)

Muscat, Oman, Gulf of Oman, Arabian Sea, northwestern Indian Ocean

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 1874)

China, East China Sea, and Japan

Thamnaconus tessellatus (Günther 1880)

Western Pacific and southeastern Indian Ocean

There are no comprehensive reviews of the Monacanthidae, although in 1988 Barry Hutchins provided taxonomic, morphological, and phylogenetic data for all monacanthid species in the world in his doctoral dissertation. Some parts of his dissertation were published, however, the taxonomic accounts of many species and genera and the phylogenetic analysis of the genera remain unpublished. Hutchins published a series of articles on the Monacanthidae between 1977 and 2002. His first article on Australian filefishes recognized 28 genera in Australia (Hutchins 1977) where the greatest diversity of filefishes is found (Hutchins 2001a).

Before Hutchins’ (1977) review of Australian filefishes appeared, Fraser-Brunner (1941a) published a review paper on the Monacanthidae (his Aluteridae), recognizing 22 genera. Although de Beaufort and Briggs (1962) lumped Amanses Gray 1835, Chaetodermis Swainson 1839, Paramonacanthus Bleeker 1865–1869, Pervagor Whitley 1930a, and Pseudomonacanthus Bleeker 1865 into a single genus, Monacanthus Linnaeus 1758, they recognized Aluterus Cloquet 1816, Anacanthus Gray 1830 (their Psilocephalus Swainson 1839), Paraluteres Bleeker 1865, and Pseudalutarius Bleeker 1865 as distinctive genera. Berry and Vogele (1961) reviewed nine species of the Monacanthidae of the western North Atlantic with detailed accounts of Amanses pullus (Ranzani 1842), Aluterus monoceros (Linnaeus 1758), Aluterus heudelotii Hollard 1855, Aluterus schoepfi (Walbaum 1792), Aluterus scriptus (Osbeck 1765), Monacanthus ciliatus (Mitchill 1818), Monacanthus tuckeri Bean 1906, Stephanolepis hispidus (Linnaeus 1758), and Stephanolepis setifer (Bennett 1831). Although Berry and Vogele (1961) placed Monacanthus pullus in the genus Amanses, they also pointed out that M.pullus should be classified in the subgenus Cantherhines Swainson 1839 because M.pullus does not have a patch of long spines on the mid-side of the body between the dorsal and anal fins, which is a diagnostic character of Amanses. When Randall (1964) reviewed the filefish genera Amanses (monotypic) and Cantherhines, he recognized 11 species of Cantherhines in the tropical seas of the world. He showed that Amanses possesses a patch of long stout spines (male) or a dense brushlike mass of long setae (female) on the mid-side of the body, whereas all species of Cantherhines lack long spines or long setae on the mid-side of the body. Randall (1964) described Cantherhines tiki as a new species based on a single specimen collected from Easter Island. Cantherhinestiki was said to be distinguishable from other species of Cantherhines by its strongly produced snout (Randall 1964). However, Caldwell and Randall (1967) showed Cantherhines tiki to be a junior synonym of Cantherhines rapanui (de Buen 1963) because of the considerable variation in the snout shape of Cantherhines rapanui.

Hutchins (1977) described three new genera, Bigener, Cantheschenia and Colurodontis, and eight new species from Australia. However, Hutchins (1994a) subsequently placed Aleuterius brownii Richardson 1848, the type species of Bigener, in Acanthaluteres Bleeker 1865, making Bigener a junior synonym of Acanthaluteres. Hutchins and Randall (1982) reviewed the Cantherhines fronticinctus complex and described Cantherhines longicaudus as a new species from Tahiti and Rarotonga. Matsuura (1984) provided brief accounts and color photographs of 26 monacanthids found in Japanese waters. Hutchins and Matsuura (1984) described Thamnaconus fijiensis from Fiji. In the following year, Hutchins (1985) reviewed the genus Brachaluteres Bleeker 1865 and recognized four species with comments on their ability to inflate the abdomen. Hutchins (1986a) published a revision of the genus Pervagor and recognized eight species including two new species, Pervagor randalli and Pervagormarginalis. Hutchins (1986b) provided a key and brief taxonomic accounts for 16 species of filefishes found along South Africa. Hutchins (1987) published Eubalichthys cyanoura as a new species from South Australia. Hutchins (1994b) described and illustrated Lalmohania velutina from India as a new genus and species. Hutchins (2002) reported another new genus and species, Enigmacanthus filamentosus, from the Seychelles and Marshall Islands. Enigmacanthus filamentosus is a very small filefish, 27−36 mm SL, resembling in size species of the genus Rudarius. Randall (2011) again reviewed the genus Cantherhines, recognizing 11 species, including two new species, Cantherhines cerinus from the Philippines and Cantherhines nukuhiva from the Marquesas Islands.

Based on the contributions shown above, I recognize 102 species in 27 genera worldwide (Table 4). I provisionally place Pseudomonacanthus multilineatus Tanaka 1918 in the genus Cantherhines following previous authors (Matsuura 1984; Kyushin et al. 1977; Lindberg et al. 1997). Although Randall (1964) placed Pseudomonacanthus multilineatus in Cantherhines, Hutchins and Randall (1982) said that Cantherhines multilineatus should be classified in the genus Thamnaconus Smith 1949b. However, they did not provide characters to justify the transfer from Cantherhines to Thamnaconus. To resolve the taxonomic status of Cantherhines multilineatus, a detailed study of diagnostic characters, including encasing scales of the posterior end of the pelvis, is needed. Hutchins suggested in his dissertation that Thamnaconus melanoproctes (Boulenger 1889) is a senior synonym of Pseudomonacanthus multilineatus. However, an examination of specimens from various localities in the Indo-West Pacific is necessary to properly access the relationships of the two nominal species. Although Barry Hutchins made great contributions to the better understanding of taxonomy and diversity of filefishes, in particular Australian filefishes, taxonomic problems in several genera such as Acreichthys Fraser-Brunner 1941c, Pseudomonacanthus, and Thamnaconus remain. As in the case of triggerfishes, many checklists and illustrated guidebooks have provided much useful information on the taxonomy, distribution, and biology of filefishes (see the section on the Balistidae).

Aracanidae (Trunkfishes, Fig. 1e; Table 5)

Trunkfishes of the family Aracanidae occur in shallow to relatively deep waters in depths from 5 m to 300 m. The species of Aracana Gray 1833, Anoplocapros Kaup 1855, Caprichthys McCulloch and Waite 1915, Capropygia Kaup 1855 and Polyplacapros Fuji and Uyeno 1979 are found in temperate seas of southern Australia, whereas species of Kentrocapros Kaup 1855 are known from deep waters in the warm, tropical regions of the Indo-Pacific (Matsuura and Yamakawa 1982; Matsuura and Tyler 1997; Matsuura 2006, 2008).
Table 5

Distributions of 13 species of the Aracanidae in the world

Species

Distribution

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

Trunkfishes differ externally from other families of Tetraodontiformes by the following combination of characters: body almost completely encased in a bony shell or carapace formed of enlarged, thickened scale plates, usually hexagonal in shape and firmly sutured to one another; isolated bony plates on caudal peduncle; carapace triangular or hexagonal in cross section, with openings for mouth, eyes, gill slit, pectoral, dorsal, and anal fins, and for the flexible caudal peduncle; scale plates often with surface granulations and prolonged in some species into prominent carapace spines over the eyes or along ventrolateral or dorsolateral angles of the body; mouth small, terminal, with fleshy lips; teeth of moderate size, conical, usually fewer than 15 in each jaw; gill opening moderately short, a vertical to oblique slit in front of pectoral-fin base; spinous dorsal fin absent; most dorsal-, anal-, and pectoral-fin rays branched; caudal fin with nine branched rays; pelvic fins absent (Matsuura 2001).

The Aracanidae and Ostraciidae have been placed in separate families by some authors (e.g., Winterbottom 1974; Tyler 1980; Matsuura 1984, 2001), but they have also been treated as two subfamilies of the family Ostraciidae by others (e.g., Winterbottom and Tyler 1983; Klassen 1995; Allen et al. 2006; Nelson 2006; Matsuura 2008). All the above authors recognized the two groups as closely related phylogenetic clades (sister groups), but placed them at different taxonomic ranks. Santini and Tyler (2003) studied fossils and extant members of the Tetraodontiformes extensively and recognized familial groupings for the Aracanidae and Ostraciidae. Recently, Santini et al. (2013a) investigated the phylogenetic relationships of the Aracanidae and Ostraciidae based on molecular analysis of many taxa (nine species of five genera of Aracanidae and 17 species of six genera of Ostraciidae). They demonstrated that the species examined in their studies (Santini and Tyler 2003; Santini et al. 2013a) have adequate characteristics to recognize two separate groups at the family level.

A problem exists with the spelling of the genus Aracana. When Gray (1833) published his article on Indian zoology, he used two different spellings for a species of trunkfish. At the beginning of the article, a section titled “Directions for arranging the plates” provided the name of animals in the plates where he presented the name of the trunkfish as “Many Spined Coffin Fish. Ostracion (Acerana) auritus”; however, the legend of plate 98 spelled the name as “Ostracion (Acarana) auritus.” Plate 98 provided a good illustration of the species showing appropriate characters to identify the species. It is the type species of the subgenus by monotypy. However, in 1838 John E. Gray published another paper in which he described three species, Ostracion (Aracana) ornata, Ostracion (Aracana) flavigaster, and Ostracion (Aracana) lineata. The second species is a junior synonym of Aracana ornata and the third Aracana aurita (Shaw in Shaw and Nodder 1798). Gray (1838) stated for his three species that “I have formed a subgenus under the name of Aracana” indicating his intention to establish the group at the subgeneric level. Although the name Aracana has long been used for the trunkfish genus, the two names Acerana and Acarana were published earlier than Aracana, thus creating a nomenclatural problem. If one follows the priority rule of the Code (ICZN 1999), the family name should be Aceranidae or Acaranidae. As Aracana has been used almost universally in the literature, with Acerana or Acarana virtually ignored by taxonomists, Aracana should be preserved under Article 33.3.1 of the Code, making the spelling of the family group name Aracanidae preserved under Article 35.4.1.

Trunkfishes have not yet been comprehensively reviewed. Although Fraser-Brunner (1935b, 1941d) provided brief reviews of the family, his accounts of genera and species were cursory as they were based on few specimens and did not provide adequate characteristics for the species treated. Kuiter (1994) and Matsuura (2008) diagnosed and illustrated seven species in four genera of Australian aracanids, covering many representatives of the family except for the genera Kentrocapros and Polyplacapros. Polyplacapros tyleri Fujii and Uyeno 1979 is unique in having a fusiform body and the caudal peduncle nearly completely covered by discrete bony plates. In addition to Australian genera of Aracanidae, detailed descriptions and illustrations of all four known species of Kentrocapros were provided by Matsuura and Yamakawa (1982), Matsuura and Tyler (1997), and Matsuura (2006). Matsuura (1990) provided brief accounts and a color photograph of a species of Kentrocapros collected from the Tasman Sea (32°43′S, 167°30′E) under the name Kentrocapros eco (Phillipps 1932). However, an examination of the holotype (102 mm TL) of Ostracion eco (NMNZ P. 903) revealed Matsuura’s (1990) Tasman Sea material to be an undescribed species (Fig. 2). This undescribed species has a spine on the dorsolateral ridge midway between the eye and the dorsal-fin origin, whereas the holotype of O. eco lacks this spine. This new species will be described and published elsewhere.
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

As discussed above, no major taxonomic problems remain in the Aracanidae. However, there still remains a need to clarify the nomenclature of some species of Anoplocapros as well as the intraspecific variation of morphological characters related to ontogenetic development of Kentrocapros.

Ostraciidae (Boxfishes, Fig. 1f; Table 6)

Boxfishes of the family Ostraciidae occur in shallow tropical and warm seas of the world. Boxfishes differ externally from other families of Tetraodontiformes by the following combination of characters: body almost completely encased in a bony shell or carapace formed of enlarged, thickened scale plates, usually hexagonal in shape and firmly sutured to one another; no isolated bony plates on caudal peduncle; carapace triangular, rectangular, or pentangular in cross section, with openings for mouth, eyes, gill slits, pectoral, dorsal, and anal fins, and for the flexible caudal peduncle; scale-plates often with surface granulations, which are prolonged in some species into prominent carapace spines over the eye or along the ventrolateral or dorsal angles of the body; mouth small, terminal, with fleshy lips; teeth moderate, conical, usually fewer than 15 in each jaw; gill opening, a moderately short, vertical to oblique slit in front of pectoral-fin base; spinous dorsal fin absent; most dorsal-, anal-, and pectoral-fin rays branched; caudal fin with eight branched rays; pelvic fins absent (Matsuura 2001).
Table 6

Distributions of 22 species of the Ostraciidae in the world

Species

Distribution

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, northward to southern Japan

Lactoria diaphana (Bloch and Schneider 1801)

East Africa eastward through northern Australia to Hawaiian and Marquesas islands, northward to southern Japan

Lactoria fornasini (Bianconi 1846)

East Africa through Indonesia to Easter Island, southward to New South Wales, northward to southern Japan

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

Fraser-Brunner (1935b) published a brief review of the Aracanidae and Ostraciidae, recognizing six genera in the Ostraciidae: Acanthostracion Bleeker 1865, Lactophrys Swainson 1839, Lactoria Jordan and Fowler 1902, Ostracion Linnaeus 1758, Rhinesomus Swainson 1839, and Rhynchostracion Fraser-Brunner 1935b. Boxfishes found in the Indo-Australian Archipelago were studied by de Beaufort and Briggs (1962) who lumped Fraser-Brunner’s (1935b) Indo-Pacific genera into the genus Ostracion. However, they recognized five subgenera in Ostracion: Lactoria, Ostracion, Rhynchostracion, Tetrosomus Swainson 1839, and Triorus Jordan and Hubbs 1925. Triorus is a junior synonym of Tetrosomus because the type species of the former, Lactophrys tritropis Snyder 1911, is a junior synonym of Tetrosomusconcatenatus (Bloch 1786) (see Randall et al. 1997). Although Ostracion nasus Bloch 1785 and Ostracion rhinorhynchos Bleeker 1851a have been placed in the genus Rhynchostracion by some authors (Kyushin et al. 1982; Gloerfelt-Tarp and Kailola 1984; Sainsbury et al. 1984; Mohsin and Ambak 1996; Randall et al. 1997; Allen et al. 2006), they were classified in the genus Ostracion by other authors (Klassen 1995; Matsuura 2001; Hayashi and Hagiwara 2013). When Fraser-Brunner (1935b) established Rhynchostracion, he differentiated it from other ostraciid genera by the degree of the development of the dorsal ridge, the convexity of the dorsal surface of the carapace, and the projection of the snout anterior to the mouth. However, the two characters involving the carapace do not clearly differentiate Rhynchostracion from other ostraciids and the snout of Ostracion nasus, the type species of Rhynchostracion, protrudes anteriorly beyond 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 led Matsuura (2001) to place O. nasus and O. rhinorhynchos in their own genus in his key and brief accounts of ostraciids found in the western central Pacific. Klassen (1995) studied osteological characters of many species of the Ostraciidae extensively and provided evidence that Rhynchostracion is a junior synonym of Ostracion. Klassen (1995) also regarded Tetrosomus as a junior synonym of Lactoria because all species of Lactoria and Tetrosomus examined by him formed a single clade. If Klassen’s (1995) proposal is accepted, Lactoria should be a junior synonym of Tetrosomus because the latter has priority over the former. However, it seems premature to accept his proposal because there are species-level taxonomic problems in several species of Tetrosomus (Allen and Erdman 2012).

Tyler (1965) revised the genus Acanthostracion in the Atlantic and recognized four species, Acanthostracion guineensis Bleeker 1865, Acanthostracion notacanthus (Bleeker 1863), Acanthostracion polygonus Poey 1876, and Acanthostracionquadricornis (Linnaeus 1758). Böhlke and Chaplin (1968) provided keys and accounts accompanied by illustrations to boxfishes found in the western Atlantic. They recognized five species, A. polygonus, A. quadricornis, Lactophrysbicaudalis (Linnaeus 1758), Lactophrys trigonus (Linnaeus 1758), Lactophrys triqueter (Linnaeus 1758). Lactophrysbicaudalis and L. triqueter were placed in the genus Rhinesomus by Matsuura (2003b) and McEachran and Fechhelm (2005). However, 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 Africa (Tyler 1965). Whitley (1965) described Acanthostracion bucephalus on the basis of a specimen from Queensland, Australia. However, there seems little doubt that it is actually A. quadricornis (Fig. 3). The record of A. quadricornis from Australia may suggest that the specimen was transferred by ballast water from a ship or released from an aquarium. As with the Balistidae, many checklists and illustrated guidebooks have provided a great deal of information on the taxonomy, distribution, and biology of boxfishes (see the section on the Balistidae). On the basis of the above overview, I recognize 22 species in the Ostraciidae (Table 6). However the relationships of T. concatenatus and T. reipublicae need to be examined in detail based on a comparison of a large selection of specimens including the types from various localities of the Indo-West Pacific to better understand the morphological variations observed in T. reipublicae (Fig. 4).
Fig. 3

Holotype of Acanthostracion bucephalus (AMS IB.6355) collected from Queensland, Australia. Photograph provided by Mark McGrouther

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)

Triodontidae (Threetooth puffer, Fig. 1g)

The family Triodontidae is represented by a single extant species, Triodon macropterus Lesson 1831, but several fossil taxa referred to this genus date back to the Eocene (see Tyler and Patterson 1991). The family differs externally from others of the order Tetraodontiformes by the following combination of characters: body moderately elongate and moderately compressed, with a very large ventral flap that may be extended downward by a rotation of the long, shaft-like pelvis; jaws modified to form a beak of three heavy, massive teeth, two on the upper jaw and one on the lower jaw; two rudimentary dorsal-fin spines present or absent; and caudal fin deeply forked (Matsuura 2001).

Triodon macropterus is rare in museum collections and poorly known. It is a deep-water species collected at depths of 50−377 m (Kyushin et al. 1977; Matsuura and Tyler 1997). When Tyler (1967) redescribed T. macropterus in detail, only 20 specimens were available for his study: five from the Indian Ocean (Mauritius, Reunion and India) and 15 from the West Pacific (Indonesia, Philippines and Japan). Due to the poor condition of two specimens, Tyler (1967) used 18 specimens to determine whether the first dorsal fin, composed of one or two spines, is present or absent. His examination revealed 12 specimens from the West Pacific to have a first dorsal fin, and five Indian Ocean specimens and a specimen from the West Pacific to lack a first dorsal fin (Table 7). However, he did not find other differences between the Indian Ocean specimens and those from the West Pacific. This led him to recognize the two groups as being conspecific. I have recently had the opportunity of examining 28 specimens of T. macropterus, 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 Indian Ocean, and present in the 19 specimens from the West Pacific (Table 7). Tyler (1967) also provided frequency distributions of fin-ray counts. My examination of 28 specimens produced almost the same frequency distributions of fin-ray counts as those of Tyler (1967), except for a count of 17 fin rays on the right side pectoral fin in a single specimen (Table 8).
Table 7

Presence or absence of first dorsal fin (two spines) in Triodon macropterus. Tyler (1967) showed that a specimen (448.3 mm SL) collected from the Volcano Islands, south of the Ogasawara Islands, had a single spine

First dorsal fin

 

Present

Absent

Indian Ocean

 

9 + 5* = 14

West-Pacific

19 + 12* = 31

1*

* Data from Tyler (1967)

Table 8

Frequency distributions of fin-ray counts in Triodon macropterus. Counts of fin rays of damaged or abnormal fins are excluded. Counts of pectoral-fin rays were obtained from fins on both sides

Dorsal-fin rays

 

10

11

12

 

This study

3

24

1

 

Tyler (1967)

2

14

1

 

Total

5

38

2

 

Anal-fin rays

 

9

10

  

This study

8

19

  

Tyler (1967)

3

12

  

Total

11

31

  

Pectoral-fin rays

 

14

15

16

17

This study

1

12

41

1

Tyler (1967)

2

7

17

 

Total

3

19

58

1

Since Tyler’s (1967) redescription was published, T. macropterus was reported (with color photographs) from several areas in the Indo-West Pacific: the Chagos Archipelago (Kyushin et al. 1977), the South China Sea (Kyushin et al. 1982), the Kyushu-Palau Ridge (Okamura et al. 1985), the Timor Sea (Gloerfelt-Tarp and Kailola 1984), and the Japanese Archipelago (Matsuura 1984). Color photographs taken by research staff of the National Museum of Nature and Science show T. macropterus to have the following coloration: body yellowish brown dorsally, 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 proximal part of the ventral flap; second dorsal, anal, pectoral, and caudal fins yellow or yellowish brown; when present, membrane of first dorsal fin black (Fig. 1g). In addition to the above reports, T. macropterus has been recorded in checklists of fishes from several areas in the West Pacific: the Mariana Islands (Myers and Donaldson 2003), Tonga (Randall et al. 2004), off southern Taiwan (Shao et al. 2008), Northern Territory of Australia (Larson et al. 2013), and northern half of Australia (Hoese et al. 2006). Kim et al. (2005) provided a brief account and a color illustration of T. macropterus from Korea.

Johnson and Britz (2005b) reported the smallest known T. macropterus (20 mm SL), collected at Wallis and Futuna at a depth of 245−440 m. They provided detailed characters and the visceral anatomy of the early juvenile that differs from adults in having a huge head (45 % SL vs 28.5−32.7 % SL in adults), no ventral flap, and different scale structure. Juveniles also differ from adults in having a very large distended stomach.

Tetraodontidae (Pufferfishes, Fig. 1h, Table 9)

Pufferfishes of the family Tetraodontidae usually occur in shallow warm, tropical seas, and freshwaters of the world, although some species may be found at depths over 350 m (Matsuura 1982; Tyler and Matsuura 1997). They differ externally from other families of the order Tetraodontiformes by the following combination of characters: head large and blunt; jaws modified to form a beak of four heavy, powerful teeth, two above and two below; eyes high on head; gill opening, a simple slit in front of pectoral fins; dorsal and anal fins located far posteriorly, containing seven to 15 soft rays; caudal fin truncate, rounded, or emarginate to somewhat lunate; pelvic fins absent; lateral line (when present) often indistinct, forming an interconnected pattern on sides of the head and body, but quite distinct in some genera (e.g., Lagocephalus and Torquigener Whitley 1930c); typical scales absent, but many spinules often present on back and/or belly, and sometimes on sides (Matsuura 2001).
Table 9

A list of 27 genera of the Tetraodontidae in the world

Genus

Number of species

Amblyrhynchote Bibron in Duméril 1855

1

Arothron Müller 1841

11

Auriglobus Kottelat 1999

5

Canthigaster Swainson 1839

36

Carinotetraodon Benl 1957

5

Chelonodontops Smith 1958

2

Colomesus Gill 1884

3

Contusus Whitley 1947

2

Dichotomyctere Duméril 1855

6

Ephippion Bibron in Duméril 1855

1

Feroxodon Su, Hardy and Tyler 1986

1

Guentheridia Gilbert and Starks 1904

1

Javichthys Hardy 1985

1

Lagocephalus Swainson 1839

11

Leiodon Swainson 1839

1

Marilyna Hardy 1982a

3

Omegophora Whitley 1934

2

Pao Kottelat 2013

14

Pelagocephalus Tyler and Paxton 1979

2

Polyspina Hardy 1983a

1

Reicheltia Hardy 1982a

1

Sphoeroides Anonymous 1798

20

Takifugu Abe 1949c

25

Tetractenos Hardy 1983a

2

Tetraodon Linnaeus 1758

6

Torquigener Whitley 1930a

20

Tylerius Hardy 1984a

1

Total number of species

184

The Tetraodontidae is the most speciose family in the Tetraodontiformes, including 184 species (Table 9). Because pufferfishes possess very few external characters useful for taxonomy and specimens are easily distorted when they are fixed in formalin and preserved in ethanol, it is often not easy for ichthyologists to recognize species limits and classify them into natural groups. Consequently, they have been poorly studied and taxonomic confusion remains. Although the Tetraodontidae has never been comprehensively reviewed, the taxonomy of pufferfishes has progressed since Fraser-Brunner’s (1943) brief review of tetraodontid genera. He classified pufferfishes into 11 genera, Canthigaster Swainson 1839, Lagocephalus, Sphoeroides Anonymous 1798, Amblyrhynchote Bibron in Duméril 1855, Torquigener, Colomesus Gill 1884, Ephippion Bibron in Duméril 1855, Tetraodon Linnaeus 1758, Arothron Müller 1841, Chonerhinos Bleeker 1854, Xenoptere Bibron in Duméril 1855, based on osteological characters of the skull, the lateral-line system on the head and body, a skin fold on the ventrolateral side of the body, and spinule distributions on the body. Although Fraser-Brunner (1943) used the spellings Amblyrhynchotes and Xenopterus, I follow Kottelat’s (2001, 2013) Amblyrhynchote and Xenoptere because Kottelat (2001) showed that Amblyrhynchotes and Xenopterus of Troschel (1856) are incorrect spellings.

When Alec Fraser-Brunner studied pufferfishes during World War II, Abe (1939, 1942, 1944, 1949a, b, c, 1950, 1952, 1954) independently studied pufferfishes found in Japan and adjacent regions. Abe (1949c) provided keys and accounts of pufferfishes in the seas around Japan. Subsequently, Abe (1952) proposed Fugu as a new genus to include five subgenera Akamefugu, Higanfugu, Shosaifugu, Takifugu, and Torafugu, which had been previously proposed and made available by Abe (1949c, 1950). However, the genus Fugu is an objective synonym of the subgenus Torafugu Abe 1950, because the type species of these genus group names is Tetraodon rubripes Temminck and Schlegel 1850. Matsuura (1990) clarified the nomenclatural problems concerning the genus group names proposed by Abe (1949c, 1950, 1952) and showed that Takifugu Abe 1949c is the appropriate generic name for warmwater pufferfishes found mainly around Japan, Korea, and China. As with other tetraodontiform families, de Beaufort and Briggs (1962) lumped Fraser-Brunner’s (1943) Indo-Pacific genera, Amblyrhynchote, Lagocephalus, and Torquigener into the genus Sphoeroides although they recognized Canthigaster, Chelonodon Müller 1841, Chonerhinos, and Tetraodon. As shown by subsequent authors, Amblyrhynchote, Lagocephalus, and Torquigener are distinctive genera.

Marine pufferfishes occurring in warm and tropical regions of the world were studied by many authors from the 1970s to present. Allen and Randall (1977) reviewed Canthigaster and described seven new species, bringing the total number of species to 22. Since Allen and Randall’s (1977) revisionary study was published, six Indo-Pacific species have been described as new as follows: Canthigaster leoparda Lubbock and Allen 1979 from the Indo-West Pacific, Canthigaster flavoreticulata Matsuura 1986 from the Tonga Submarine Ridge, South Pacific, Canthigastercyanetron Randall and Cea-Egaña 1989 from Easter Island, Canthigaster punctata Matsuura 1992 from the Mascarene Submarine Ridge in the western Indian Ocean, Canthigastercyanospilota Randall, Williams and Rocha 2008 from the Red Sea, and Canthigaster criobe Williams, Delrieu-Trottin, Planes 2012 from the Gambier Archipelago in French Polynesia. In addition, Randall et al. (2008) reviewed Canthigaster coronata (Vaillant and Sauvage 1875) and recognized Canthigaster axiologa Whitley 1931 as a separate species. Although Allen and Randall (1977) synonymized Canthigaster papua (Bleeker 1848) with Canthigaster solandri (Richardson 1845), Randall (1995) and Allen and Erdman (2012) recognized it as a distinctive species. Moura and Castro (2002) reviewed Canthigaster in the Atlantic where many authors recognized only a single species Canthigaster rostrata (Bloch 1786). Moura and Castro (2002) extensively studied specimens collected from various localities from both sides of the Atlantic and recognized six species, three of which they described as new: Canthigaster figueiredoi from the east coast of South America, Canthigaster jamestyleri from the southeast coast of the United States and the Gulf of Mexico, and Canthigaster supramacula from the west coast of Africa. The above studies bring the total number of species of Canthigaster to 36, making it the largest genus in the Tetraodontidae.

Shipp (1974) reviewed pufferfishes of the Atlantic belonging to Canthigaster, Colomesus Gill 1884, Ephippion Bibron in Duméril 1855, Lagocephalus, and Sphoeroide. Prior to Shipp’s (1974) publication, Tyler (1964) reviewed two species of Colomesus, Colomesus asellus (Müller and Troschel 1848), and Colomesus psittacus (Bloch and Schnedier 1801) in detail. Amaral et al. (2013) described Colomesus tocantinensis from the Tocantins River, Brazil. Shipp (1974) recognized 13 species of Sphoeroides in the Atlantic and provided a key to the species, along with detailed accounts of all Atlantic species. Walker and Bussing (1996) described the eastern Pacific Sphoeroides lispus Walker in Walker and Bussing 1996 and Sphoeroides rosenblatti Bussing in Walker and Bussing 1996 as new species. Prior to Shipp (1974) and Walker and Bussing (1996), five species of Sphoeroides had already been reported from the eastern Pacific. With 20 species, Sphoeroides is the third largest genus in the Tetraodontidae. Most species of Sphoeroides occur in shallow waters of the Atlantic and the eastern Pacific, but Sphoeroides pachygaster (Müller and Troschel in Schomburgk 1848) is found in deep waters exceeding 350 m in all oceans.

Shipp (1974) also provided detailed accounts and photographs of Lagocephalus laevigatus (Linnaeus 1766) and Lagocephalus lagocephalus (Linnaeus 1758). The former is endemic to the Atlantic and the latter is distributed in tropical and other warm seas of the world. Abe and Tabeta (1983) described Lagocephalus gloveri as a new species from Japan. Abe et al. (1984) described another new species, Lagocephalus wheeleri, from Japan, but Matsuura (2010) showed it to be a junior synonym of Lagocephalus spadiceus (Richardson 1845). Matsuura et al. (2011) redescribed Lagocephalus guentheri (Miranda Ribeiro 1915) based on an examination of the holotype and many specimens from the Red Sea. Although the genus Lagocephalus has not been comprehensively reviewed, an ongoing study by the author has revealed 11 valid species in the genus. This study will be published elsewhere.

Although Abe (1949a, b, c, 1950, 1952, 1954) made progress on the taxonomy of Takifugu, many species found along the coast of China were not included in his articles. Cheng et al. (1975) reviewed pufferfishes found in Chinese waters and described Takifugu flavidus (Li, Wang and Wang in Cheng et al. 1975) and Takifugu reticularis (Tian, Cheng and Wang in Cheng et al. 1975). Subsequently, Chinese authors described four species of Takifugu from China: Takifugu orbimaculatus Kuang, Li and Liang 1984 from a creek in Lianhuashan, Guangdong; Takifugu coronoides Ni and Li 1992 from an estuary of Chang Jiang; Takifuguplagiocellatus Li in Su and Li 2002 from the coast of Hainan Island; and Takifugu variomaculatus Li in Su and Li 2002 from brackish waters of Modaomen Zhuhai, Guangdong. These Chinese and Japanese authors increased the number of Takifugu species making it the second largest genus in the Tetraodontidae, with 25 species. However, taxonomic problems still remain in Takifugu, such as Takifugu basilevskianus (Basilewsky 1855) and Takifugu pseudommus (Chu 1935), which show significant variation in morphological features including ontogenetic color changes.

Benl (1957) described Carinotetraodon chlupatyi as a new genus and species from Thailand. However, Dekkers (1975) synonymized Carinotetraodon chlupatyi with Carinotetraodon lorteti (Tirant 1885), making that species the type of the genus. Carinotetraodon is similar to Canthigaster in having a laterally compressed body and several derived osteological features, as well as a shared derived behavioral peculiarity in their skin ridge-lifting along the midline of the body (Tyler 1978, 1980). Additional new species of Carinotetraodon were described: Carinotetraodon salivator Lim and Kottelat 1995 from Sarawak, Carinotetraodon irrubesco Tan 1999 from Sumatra, and Carinotetraodon imitator Britz and Kottelat 1999 from India.

Tyler and Paxton (1979) described Pelagocephalus coheni as a new genus and species based on a single specimen collected at Norfolk Island. Pelagocephalus is externally distinguished from other pufferfishes by its nasal apparatus in the form of an open, flat, and relatively unornamented disk. Heemstra and Smith (1981) described a second species of Pelagocephalus, Pelagocephalus marki from South Africa. Hardy (1982b) recorded P.marki from New Zealand.

In the period from 1980 to 1989, Graham S. Hardy published a series of papers on the taxonomy of pufferfishes that are distributed mainly in the Southern Hemisphere. Hardy (1980) reviewed the antitropical species Arothron firmamentum (Temminck and Schlegel 1850) indicating that it should be classified in Arothron, whereas it had previously been classified in Boesemanichthys by Abe (1952). The monotypic genus Contusus Whitley 1947 was reviewed by Hardy (1981), who described a second species Contusus brevicaudus restricted to southern Australia. Hardy and Hutchins (1981) recognized the validity of Omegophora Whitley 1934 with the description of Omegophora cyanopunctata from the southwestern part of Australia. Hardy (1982a) erected two new genera, Marilyna and Reicheltia, based on detailed morphological comparisons including skull osteology. Hardy (1983a) reviewed the Australian species of Torquigener and established two new generic names, Polyspina with one species and Tetractenos with two species. He recognized 12 species of Torquigener, including five new species from Australia. Hardy and Randall (1983) described Torquigener flavomaculosus as a new species from the Red Sea. Hardy described Torquigener randalli Hardy 1983b from the Hawaiian Islands and Torquigener gloerfelti Hardy 1984b from Indonesia. Hardy (1989) described Torquigener balteus as a new species from South Africa and provided a key to 19 species of Torquigener, although he overlooked Sphoeroides marleyi Fowler 1929, which is a member of Torquigener. My examination of the holotypes of Torquigener balteus and S. marleyi from South Africa revealed them to be conspecific, making the former a junior synonym of the latter. Matsuura (2014) recently described Torquigener albomaculosus as a new species from Amami-oshima Island, Ryukyu Islands. Torquigener albomaculosus is unique in building large spawning nests, called “mystery circles” by local SCUBA divers. Thus, the above studies of Torquigener bring the total number of species of Torquigener to 20, the co-equal third largest genus in the Tetraodontidae along with Sphoeroides. Hardy (1984a) recognized Sphoeroides spinosissimus Regan 1908 as being distinctive from other genera of the Tetraodontidae and established a new genus Tylerius for it. Hardy (1985) described Javichthys kailolae as a new genus and species from Indonesia. Su et al. (1986) established a new genus, Feroxodon, for Anchisomus multistriatus Richardson 1854 from Australia.

The taxonomy of freshwater pufferfishes in Asia was in a confused state until Dekkers (1975) reviewed the genus Tetraodon. Subsequently, Sontirat and Soonthornsatit (1985) described Tetraodon suvattii as a new species from Thailand, and Roberts (1988) described Tetraodon abei as a new species from Laos. Roberts (1982) reviewed the genus Chonerhinos Bleeker 1854 and described three species from Borneo, Chonerhinos amabilis, Chonerhinos nefastus, and Chonerhinos silus. Still, complex nomenclatural problems involving Chonerhinos and Xenoptere remained until Kottelat (1999) proposed the new genus, Auriglobus, to rectify these problems. Kottelat (2013) discussed nomenclatural issues of the Tetraodontidae found in Southeast Asia in detail. He established the new genus Pao (type species, Tetraodon leiurus Bleeker 1850a) that includes 13 species previously placed in Tetraodon and Monotrete, but differentiated from other pufferfishes by their unique color pattern and a very elongate premaxillary pedicel that creates a greatly enlarged open space between their dorsomedial edges. Thus, the generic name Tetraodon is now applied to only six species of African freshwater pufferfishes. Kottelat (2013) also recognized Leiodon Swainson 1839 as a valid genus that includes Tetrodon cutcutia Hamilton 1822 distributed in freshwaters of southern Asia. Kottelat (2013) recognized Dichotomyctere Duméril 1855 for six species: Dichotomyctere erythrotaenia (Bleeker 1853), Dichotomyctere fluviatilis (Hamilton 1822), Dichotomyctere kretamensis (Inger 1953), Dichotomyctere nigroviridis (Marion de Procé 1822), Dichotomyctere ocellatus (Steindachner 1870), and Dichotomyctere sabahensis (Dekkers 1975). Saenjudaeng et al. (2013) described Tetraodon palustris as a new species from the Mekong basin of Thailand. Because of its publication date, July 2013, the authors were unaware of Kottelat’s (2013) article proposing the genus Pao. However, it is clear from the original description of Tetraodon palustris, that it is a species of Pao, bringing the total number of species in that genus to 14.

Kottelat (2013) also commented on genera of marine pufferfishes. Although Tetrodon patoca Hamilton 1822 has long been placed in Chelonodon Müller 1841, Kottelat (2013) pointed out that Tetrodon cutcutia Hamilton 1822, the type species of Chelonodon, had been placed in the same genus as Tetraodon fluviatilis Hamilton 1822 and related species by Tyler (1980). Tyler (1980) called this group Chelonodon but Kottelat (2013) stated that the oldest available name for a genus including Tetraodon fluviatilis is Dichotomyctere. Tetrodon patoca and Tetraodon fluviatilis have a sufficient number of character differences to warrant being placed in separate genera. The type species of Chelonodontops Smith 1958 is Chelonodontops pulchellus Smith 1958, a junior synonym of Chelonodon pleurospilus (Regan 1919) and congeneric with Tetraodon patoca (see Smith 1986). Thus, Tetraodon patoca is a species of Chelonodontops (Kottelat 2013). Matsuura (2002) reviewed two species of Chelonodon, Chelonodon laticeps Smith 1948, and Chelonodon patoca, both of which belong in Chelonodontops. Kottelat (2013) showed that Gastrophysus Müller 1843 is an objective synonym of Takifugu Abe 1949, as the type species of both genus group names is Tetrodon oblongus Bloch 1786. However, this is likely to cause confusion not only in pufferfish taxonomy, but also in the fisheries of East Asia where pufferfishes are treated as important and expensive fishes (e.g., several species of Takifugu cost about US$100 per kilogram at fish markets). In addition, because pufferfishes have fatal poison in their viscera, scientific names are very important for food security management in East Asia. As suggested by Kottelat (2013), Gastrophysus can be suppressed under the International Code of Zoological Nomenclature (ICZN 1999). I will ask the International Commission on Zoological Nomenclature to suppress Gastrophysus.

Matsuura and Okuno (1991) redescribed a rare pufferfish, Arothron carduus (Cantor 1849), on the basis of the holotype from Penang and two additional specimens from the West Pacific. Matsuura (1994) described Arothron caeruleopunctatus as a new species from the tropical region of the Indo-West Pacific. Matsuura (1999) briefly reviewed Arothron with a key to species and accounts of all members of the genus including two undescribed species.

As indicated above, there has been a great deal of progress in taxonomy of the Tetraodontidae during the past several decades, especially in the period from the mid-1970s to the present. However, there remain taxonomic problems in genera such as Arothron, Chelonodontops, Lagocephalus, Pao, Takifugu, and Torquigener where many species await description and detailed morphological and molecular comparisons to classify them into appropriate groups. The taxonomy of pufferfishes in Southeast Asia is important not only for the understanding of fish diversity of the region, but also for the welfare and food management for humans. Dao et al. (2012) reported that the number of victims of food poisoning by eating pufferfishes reached 737, with 127 mortalities from 1999 to 2003. Due to the lack of knowledge about pufferfishes and their toxicity, pufferfishes are still found in Southeast Asian fish markets (Fig. 5), although local and state governments in countries around the South China Sea have prohibited the sale of pufferfishes for food.
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

Diodontidae (Porcupinefishes, Fig. 1i)

Many species of the family Diodontidae are benthic inhabitants around coral or rocky reefs, but some occur frequently in seagrass beds and on sand or mud bottom to depths of 150 m, with one species being found in pelagic waters. They differ externally from other families of the order Tetraodontiformes by the following combination of characters: body wide and capable of great inflation, covered with massive spines, which may be quite long; spines with large bases or roots, which are beneath the skin; long spines that are usually erectile and double-rooted, or short spines fixed in an erect position by their triple-rooted bases; head broad and blunt; nasal organ usually appearing as small tentacles located in front of the large eyes; mouth large, wide, and terminal, teeth fused to form a strong, beak-like crushing structure without a median suture dividing the upper and lower jaws into left and right halves; gill opening, a relatively small, vertical slit immediately preceding the pectoral-fin base; short-based dorsal and anal fins with soft rays, set far back on body, and like caudal fin, generally rounded; most fin rays branched; bases of fins often thick and fleshy; no pelvic fins (Leis 2001).

Porcupinefishes were well studied by Leis (1978, 1986, 2001, 2003, 2006). Leis (1978) reviewed Diodon Linnaeus 1758 and provided detailed accounts of five species: Diodon eydouxii Brisout de Barneville 1846, Diodon holocanthus Linnaeus 1758, Diodon hystrix Linnaeus 1758, Diodon liturosus Shaw 1804, and Diodon nicthemerus Cuvier 1818. Leis (2006) provided a list of synonyms and keys for all species of the Diodontidae. Keys to regional species were also provided by Leis (1986, western Indian Ocean; 2001, eastern Indian Ocean and western central Pacific; 2003, western Atlantic). Accounts and color photographs for species of the eastern Pacific were provided by Allen and Robertson (1994). Australian species were diagnosed and illustrated by Gomon (2008). As pointed out by Leis (2006), taxonomic problems remain for some species, such as Chilomycterus reticulatus (Linnaeus 1758) and species of “Atlantic Chilomycterus.”

Molidae (Ocean sunfishes, Fig. 1j)

Ocean sunfishes of the family Molidae occur in tropical and other warm seas of the world. They are usually pelagic, descending to a depth of over 200 m. The ocean sunfishes differ externally from other families of Tetraodontiformes by the following combination of characters: body short and deep or oblong, prominently compressed; caudal peduncle and typical caudal fin absent; eyes small; mouth terminal, small; teeth united and beak-like in each jaw without a median suture; no palatine teeth; gill opening small, pore-like, located in front of pectoral-fin base; dorsal and anal fins of similar shape, generally triangular, dorsal fin located opposite anal fin; dorsal and anal fins spineless, each with 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 leathery, rudder-like lobe known as a pseudocaudal fin or clavus (supported mostly by fin-ray elements originally belonging to dorsal and anal fins); skin leathery, with many small scales (small juveniles may also have some larger scattered spiny scales) (Hutchins 2001b).

Fraser-Brunner (1951) reviewed ocean sunfishes and recognized five species in the Molidae: Masturus lanceolatus (Liénard 1840), Masturus oxyuropterus (Bleeker 1873), Mola mola (Linnaeus 1758), Mola ramsayi (Giglioli 1883), and Ranzania laevis (Pennant 1776). Because species of Mola and Masturus reach more than 3 m in length and two tons in weight, it is difficult to preserve specimens of adults in museums. It is also difficult for ichthyologists to obtain measurements and counts on large adult ocean sunfishes in the field, and specimens of Mola and Masturus are not frequently collected and returned to museum collections. Ranzania laevis is relatively small, reaching around 80 cm TL, but specimens of Ranzania have also been rarely collected. Since ichthyologists have few opportunities to study an adequate number of adult specimens of ocean sunfishes, authors have been unable to agree how many species in the family, some believing a single species exists in each genus, and others recognizing two species each in Mola and Masturus and one species in Ranzania (Bray 2008).

Parenti (2003) published a list of nominal species in the Molidae. Bass et al (2005) published a molecular analysis of ocean sunfishes and recognized two clades in Mola, M. mola and M. ramsayi. They demonstrated that M. ramsayi occurs in South Africa and Australia, whereas M. mola is found in all oceans. However, they were unable to differentiate among specimens of Masturus from the West Pacific (Taiwan) and the western Atlantic (Florida). 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 332 cm TL, Fig. 6). They showed convincingly that there are two species with clear morphological differences: a well-developed head bump (head bump height 12.1 % TL) in group A vs. with no distinct head bump (head bump height 7.8 % TL) in group B, number of clavus fin rays 14−17 in group A vs 10−13 in group B, number of clavus ossicles 8−15 in group A vs. 8−9 in group B, and edge of clavus not wavy in group A vs. wavy in group B. Yoshita et al. (2009) suggested strongly that their group A was M. ramsayi and group B was M. mola. However, a clearer view of intra- and interspecific relationships of Mola must await the availability of more specimens from other regions of the world’s oceans for study by both morphological and molecular methods.
Fig. 6

A large specimen of Mola ramsayi (330 cm TL) captured with a set net along the Pacific coast of northern Honshu, Japan

Systematics

Cuvier (1816) pioneered the classification of tetraodontiform fishes by placing them in the order Plectognathi based on his detailed anatomical studies. Since Cuvier’s (1816) work was published, there have been various arrangements of tetraodontiform classification (see Tyler 1980 for a history of tetraodontiform classification), but the Plectognathi or Tetraodontiformes has nearly always been considered a monophyletic group, except for Le Danois (1955, 1959, 1961). As appropriately criticized by Tyler (1963), Yseult Le Danois tried to destroy the order Tetraodontiformes. She stated that the triacanthoids and balisoitds are of acanthurid origin and that the other plectognaths (her Orbiculati) are not even of percoid derivation, being related to the isospondylous fishes, and that Canthigaster is related to the ostracioids rather than the tetraodontids. However, her statements were based on erroneous observations and interpretations of the osteological and myological characters of tetraodontiforms (Tyler 1963; Winterbottom 1974). No comprehensive phylogenetic studies of tetraodontiforms appeared prior to studies by Winterbottom (1974) and Tyler (1980). Morphological synapomorphies of tetraodontiforms have been provided by Winterbottom (1974), Tyler (1980), Rosen (1984), Tyler and Sorbini (1996), and Santini and Tyler (2003). In their review of the interrelationships of actinopterygian fishes, Lauder and Liem (1983) supported the monophyly of Tetraodontiformes by adding to the already substantial list of synapomorphies. In addition, Wiley and Johnson (2010) provided a list of 10 synapomorphies for fossil and extant tetraodontiforms.

On the basis of a comprehensive myological study, Winterbottom (1974) analyzed relationships of Tetraodontiformes using cladistic methods. He recognized three major clades: (1) Triacanthodidae + Triacanthidae, (2) (Balistidae + Monacanthidae) + (Aracanidae + Ostraciidae), and (3) Triodontidae + [(Tetraodontidae + Diodontidae) + Molidae] (Fig. 7). The first clade, including the Triacanthodidae and Triacanthidae, was considered to be the sister group of all other tetraodontiforms. Tyler (1980) studied tetraodontiforms extensively and provided a huge number of osteological descriptions, comparative diagnoses, and illustrations for all extant families, major representatives of extant genera, and most of the known fossil taxa of the Tetraodontiformes. Tyler (1980) analyzed phylogenetic relationships of the Tetraodontiformes by an evolutionary (traditional) method and his systematic arrangements of families are similar to those of Winterbottom (1974), except for placing the superfamily Triacanthoidea (Triacanthodidae + Triacanthidae) as the basal sister group to the two superfamilies, the Balistoidea (Balistidae and Monacanthidae) and the Ostracioidea (Aracanidae and Ostraciidae).
Fig. 7

Phylogenetic relationships of the extant families of Tetraodontiformes inferred from cladistic analyses of morphological characters. a Winterbottom 1974; b Leis 1984 (larvae of the Aracanidae and Triodontidae not available for Leis); c Santini and Tyler 2003

Winterbottom (1974) and Tyler (1980) stated that tetraodontiforms were probably related to acanthuroid fishes of the Perciformes, although Winterbottom (1974) suggested that some zeiforms might also be related to tetraodontiforms. Rosen (1984) proposed on the basis of his osteological analysis that zeoids have a sister-group relationship 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 tetraodontiform families that were similar to those of Winterbottom (1974) and Tyler (1980), except for the phylogenetic position of the Ostraciidae. Leis (1984) placed the Ostraciidae in an unresolved trichotomy, Ostraciidae + 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, Diodontidae, and Molidae). Winterbottom and Tyler (1983) also provided many synapomorphies involving osteological and myological characters that supported a sister-group relationship 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 contributions to generate a new classification of all known families represented by fossil and extant forms. For extant families, Santini and Tyler (2003) placed the Triacanthodidae as a sister group to all other families of the Tetraodontiformes (Fig. 7). They arranged the other families into two suborders, the Balistoidei and Tetraodontoidei. The former is composed of the Triacanthidae + [(Balistidae + Monacanthidae) + (Aracanidae + Ostraciidae)]. The latter suborder is composed of the Triodontidae + [(Tetraodontidae + Diodontidae) + Molidae]. This classification is similar to those of Winterbottom (1974) and Tyler (1980).

Miya et al. (2003) analyzed molecular data for 100 species of higher teleosteans. Their study was the first to hypothesize close relationships among lophiiforms, tetraodontiforms, and caproids. They retrieved phylogenetic topologies placing caproids as a sister group with tetraodontiforms and the two groups as sister to lophiiforms. However, they used few taxa: one species of Caproidae, two species of Tetraodontiformes, and six species of Lophiiformes. Yamanoue et al. (2007) analyzed more species of the three groups and provided a robust phylogenetic topology that placed the Caproidei as sister to the Lophiiformes, and the two groups as a clade that is sister to the Tetraodontiformes.

Nakae and Sasaki (2010) studied the lateral-line system and its innervations of nine species of tetraodontiforms (representing all families examined except for the Molidae) and a single species each from the Lophiidae, Zeidae, Caproidae, and Siganidae. Their analysis supported a close relationship of the Tetraodontiformes with the Lophiidae, but not with the Zeidae, Caproidae, or Siganidae. Recently, Chanet et al. (2013) presented synapomorphies of tetraodontiforms and lophiiforms involving soft anatomical characters: rounded and anteriorly disposed kidneys, a compact thyroid included in a blood sinus, an abbreviated spinal cord, an asymmetric liver, and clusters of supramedullary neurons in the rostral part of the spinal cord. Baldwin (2013) also provided a putative synapomorphy of some tetraodontiforms and lophiiforms: they are strikingly similar in having the trunk enclosed in an inflated sac covered with xanthophores. Although Nakae and Sasaki (2010), Chanet et al. (2013), and Baldwin (2013) did not study many species of tetraodontiform and outgroup fishes, and the relationship between the Tetraodontiformes and Lophiiformes is now supported both by molecular and morphological characters.

Molecular studies by various authors have generally supported the monophyly of tetraodontiform families (Holcroft 2005; Alfaro et al. 2007; Yamanoue et al. 2007, 2008), although conflicts exist between the constructed topologies of familial relationships by morphological and molecular analyses (Figs. 7, 8). In the morphological studies by Winterbottom (1974), Tyler (1980), and Santini and Tyler (2003), the Triodontidae, Tetraodontidae, Diodontidae, and Molidae form a monophyletic group, whereas Holcroft (2005) placed the Molidae with Aracanidae + Ostraciidae, treating them as subfamilies. Alfaro et al. (2007) placed the Molidae as the basal sister group to Triodontidae + (Aracanidae + Ostraciidae). Britz and Johnson (2005) and Johnson and Britz (2005a) found the fusion of anterior vertebral centra in the occipital region and a thick band of cartilage on the side of the pterygiophores of the vertical fins in an ostraciid and molids suggestive of a close relationship between the Ostraciidae and Molidae. However, because they studied only one species of Ostraciidae and two species of Molidae, it seems premature to hypothesize a close relationship of the two families until further support for it may be forthcoming. In contrast, the Molidae was recovered as the sister group to a clade comprising the Tetraodontidae and Diodontidae by Santini et al. (2013c) as previously indicated by morphological studies. The Triodontidae was considered to be close to the Tetraodontidae, Diodontidae, and Molidae by morphological studies (Winterbottom 1974; Tyler 1980; Santini and Tyler 2003), whereas molecular studies recovered close relationship of the Triodontidae with the Aracanidae and Ostraciidae (Alfaro et al. 2007; Yamanoue et al. 2008; Santini et al. 2013c).
Fig. 8

Phylogenetic relationships of the Tetraodontiformes inferred from molecular analyses. a Holcroft 2005; b Alfaro et al. 2007; c Yamanoue et al. 2007; d Santini et al. 2013c. The Triodontidae was not included in Holcroft (2005), and the Ostraciidae of Holcroft (2005) and Yamanoue et al. (2007) included two subfamilies the Aracaninae and Ostraciinae, which were classified as families in the other papers

Despite differences in molecular trees of familial relationships among authors, several sister group relationships are generally supported by molecular studies (Holcroft 2005; Alfaro et al. 2007; Yamanoue et al. 2007; Santini et al. 2013c) as follows: Triacanthodidae + Triacanthidae, Balistidae + Monacanthidae, Aracanidae + Ostraciidae, and Tetraodontidae + Diodontidae, as previously seen in morphological studies. On the basis of whole mitochondrial genome sequences, Yamanoue et al. (2008) documented different phylogenetic relationships from those of other authors. Whereas the Triacanthidae was usually placed close to the Triacanthodidae, Balistidae, and Monacanthidae in other studies (Winterbottom 1974; Tyler 1980; Leis 1984; Santini and Tyler 2003; Holcroft 2005; Alfaro et al. 2007; Santini et al. 2013c), Yamanoue et al. (2008) recovered a relationship of the Triacanthidae with the Tetraodontidae and Diodontidae.

Phylogenetic relationships of genera and species were studied by various authors in the Triacanthidae, Balistidae, Monacanthidae, Aracanidae, Ostraciidae, Tetraodontidae, and Molidae, but the numbers of taxa studied differed greatly and resulted in different resolutions of phylogenetic analyses. Santini and Tyler (2002b) analyzed phylogenetic relationships of species and genera of the Triacanthidae using morphological characters. They found two clades, one composed of two species of Triacanthus Oken 1817, and the other composed of three species of Tripodichthys Tyler 1968 and one species each of Pseudotriacanthus Fraser-Brunner 1941b and Trixiphichthys Fraser-Brunner 1941b. Matsuura (1979, 1981) analyzed cladistic relationships of all extant genera of the Balistidae and 22 genera of the Monacanthidae using osteological characters. Although many balistid genera showed no significant differences in osteological characters, Matsuura (1979) found that a derived condition of the interhyal was shared by Rhinecanthus and Sufflamen. He also recovered Canthidermis as sister to all other balistid genera based on a scale bone in the posterior part of the skull. However, Tyler and Matsuura (1981) revealed Matsuura’s (1979) “scale bone” to be the posterior part of the sphenotic that appears separated from the major portion of the sphenotic. Matsuura (1981) found derived conditions of the pectoral girdle and the skull in Xenobalistes, making it a sister group to all other balistid genera. Matsuura (1979) recognized two clades in the Monacanthidae: the first composed of Arotrolepis Fraser-Brunner 1941c, Paramonacanthus, Monacanthus, Stephanolepis Gill 1861, Chaetodermis, Acreichthys, and Pervagor; the second Cantherhines, Eubalichthys Whitley 1930b, Thamnaconus (= Navodon Whitley 1930b of Matsuura 1979), Pseudomonacanthus, Scobinichthys Whitley 1931, Nelusetta Whitley 1939, Meuschenia Whitley 1929, Rudarius, Aluterus, Oxymonacanthus Bleeker 1865, Pseudalutarius, Brachaluteres, Paraluteres, and Anacanthus. The second clade possesses more advanced characters than the first clade and is composed of three subclades: (1) Amanses and Cantherhines, (2) Eubalichthys, Thamnaconus, Pseudomonacanthus, Scobinichthys, Nelusetta, Meuschenia and Rudarius, and (3) Aluterus, Oxymonacanthus, Pseudalutarius, Brachaluteres, Paraluteres, and Anacanthus.

In her molecular analyses of tetraodontiforms, Holcroft (2005) also recovered the close relationship of Rhinecanthus and Sufflamen found by Matsuura (1979). She further showed that three species of Balistes form a sister group to Pseudobalistes fuscus (Bloch and Schneider 1801) and that Xanthichthys auromarginatus (Bennett 1832) is sister to Balistoides viridescens. Her analysis revealed that Balistoides conspicillum is sister to other balistids, including Balistes, Pseudobalistes, Rhinecanthus, Sufflamen, Abalistes, Canthidermis, Xanthichthys, and Balistoides viridescens. In addition, she found Melichthys niger (Bloch 1786) to be the first lineage to diverge in the Balistidae. Although Holcroft (2005) studied phylogenetic relationships of nine genera of Monacanthidae, her study covered relatively small numbers of monacanthid genera making it difficult to compare her phylogenetic tree with that constructed by Matsuura (1979). Yamanoue et al. (2009) studied the phylogenetic relationships of 12 species of Balistidae and 21 genera of Monacanthidae, involving one species each from 33 genera of the two families. The molecular analysis placed Balistes as sister to all other balistid genera that were separated into two clades, the first comprising Canthidermis, Abalistes, Rhinecanthus, and Sufflamen, and the second Pseudobalistes, Xanthichthys, Xenobalistes, Odonus, Balistapus, Balistoides, and Melichthys. In the Monacanthidae they found the clade that included two genera, Oxymonacanthus and Pseudalutarius, form a sister group to other genera that are divisible into two major clades, the first composed of Rudarius, Brachaluteres, Paraluteres, Pervagor, Stephanolepis, Acreichthys, Chaetodermis, Monacanthus, and Paramonacanthus, and the second Aluterus, Pseudomonacanthus, Amanses, Cantherhines, Eubalichthys, Thamnaconus, Nelusetta, Scobinichthys, Meuschenia, and Acanthaluteres.

Santini et al. (2013a) provided the largest molecular dataset for the Balistidae and Monacanthidae based on two mitochondrial and three nuclear loci of 33 species of Balistidae and 53 species of Monacanthidae. The molecular analysis supported monophylies of the following genera: Odonus, Balistapus, Melichthys, Xanthichthys, Canthidermis, Balistes, Abalistes, Rhinecanthus, and Sufflamen. Species of Balistoides and Pseudobalistes were recovered in separate clades with Balistoides conspicillum sister to Melichthys, and B. viridescens forming a clade with P. flavimarginatus. The latter two species share at least two morphological characters, a small naked area on the cheek just behind the mouth, and a large maximum size, reaching over 60 cm TL. In the balistid tree Xenobalistes tumidipectoris was deeply nested in a clade composed of five species of Xanthichthys, strongly supporting Xenobalistes as a junior synonym of Xanthichthys (see Santini et al. 2013b). Santini et al. (2013b) recovered virtually all of the subclades of Yamanoue et al. (2009), except for Chaetodermis and Acreichthys relationships. In the phylogenetic topology of Santini et al. (2013b), Chaetodermis and Acreichthys are sequential sister taxa to the clade composed of three species of Paramonacanthus and Monacanthus chinensis, whereas Yamanoue et al. (2009) presented sister-group relationships of Chaetodermis and Acreichthys. Santini et al. (2013b) supported monophylies or close relationships of most genera, but species of Monacanthus and Paramonacanthus were separated into different clades suggesting strongly that these genera require revision.

A cladistic analysis of the external characters, osteology, and myology of all genera of Aracanidae led Winterbottom and Tyler (1983) to recognize two clades in the Aracanidae, the first composed of Kentrocapros and Polyplacapros, and the second Aracana, Strophiurichthys Fraser-Brunner 1935b, Anoplocapros, Caprichthys, and Capropygia. In the second clade Aracana was sister to the remaining genera, Strophiurichthys and Anoplocapros diverged sequentially, and Caprichthys and Capropygia were placed in a small terminal clade. Santini et al. (2013a) studied the phylogenetic relationships of Kentrocapros (two species), Caprichthys (monotypic), Capropygia (monotypic), Aracana (two species), and Anoplocapros (three species), with Polyplacapros (monotypic). The two monotypic genera Caprichthys and Capropygia were found to represent a clade to other genera. Kentrocapros was placed as sister to the subclade composed of Aracana and Anoplocapros.

Klassen (1995) studied the osteological characters of 19 species of the Ostraciidae extensively. His phylogenetic analysis found two clades in the Ostraciidae, the first composed of Acanthostracion and Lactophrys Swainson 1839, and the second Ostracion and Lactoria, having synonymized Rhynchostracion with Ostracion and Tetrosomus with Lactoria. Santini et al. (2013a) studied the phylogenetic relationships of 17 species of Ostraciidae by molecular analysis. The resultant phylogenetic tree closely resembled that of Klassen (1995), but where Klassen (1995) found Rhynchostracion nasus (Bloch 1785) deeply nested within Ostracion, Santini et al. (2013a) presented it as sister to Ostracion.

On the basis of many osteological characters, nasal organ, and lateral-line system, Tyler (1980) inferred three groups of pufferfish genera: (1) Lagocephalus, Colomesus, Guentheridia Gilbert and Starks 1904, and Sphoeroides; (2) Amblyrhynchote, Torquigener, and Fugu (= Takifugu); and (3) all other genera. Holcroft (2005) and Alfaro et al. (2007) analyzed the phylogenetic relationships of 19 species of 10 genera (Arothron, Canthigaster, Lagocephalus, Marilyna, Monotreta, Sphoeroides, Takifugu, Tetractenos, Tetraodon, and Torquigener). Their results are similar, placing Lagocephalus at the base of the topologies. Yamanoue et al. (2011) studied the phylogenetic relationships of 50 species of Tetraodontidae by molecular analysis. The study found four major clades, the first composed of species of Lagocephalus, the second Colomesus and three species of Sphoeroides, the third Marilyna, Tetractenos, Tylerirus, Polyspina, Torquigener, and Takifugu, and the fourth and the largest clade Carinotetraodon, Tetraodon, Auriglobus, Pelagocephalus, Canthigaster, Chelonodon, Omegophora, and Arothron. The phylogenetic tree of Yamanoue et al. (2011) recognized many monophyletic lineages, but Colomesus and species of Tetraodon fell into distantly related clades. A relaxed molecular-clock Bayesian divergence time estimation calculated three invasions to freshwater by pufferfishes during their evolution: the first 48−78 million years ago in Southeast Asia, the second 17−38 million years ago in Africa, and the third 0−10 million years ago in South America.

Igarashi et al. (2013) studied the phylogenetic relationships of 17 species of Tetraodon by molecular analysis. The study clearly showed that they did not form a monophyletic group. Except for T. cutcutia, 16 species were placed into three major clades: an Asian freshwater group composed of T. leiurus, Tetraodon palembangensis Bleeker 1850b, T. abei, Tetraodon baileyi Sontirat 1985, Tetraodon cochinchinensis Steindachner 1866, T. suvattii, and Tetraodon turgidus Mitchill 1815; an Asian brackish water group of Tetraodon erythrotaenia Bleeker 1853, Tetraodon biocellatus Tirant 1885, T. fluviatilis, and Tetraodon nigroviridis Marion de Procé 1822; and an African freshwater group of Tetraodon lineatus Linnaeus 1758, Tetraodon pustulatus Murray 1857, Tetraodon mbu Boulenger 1899, Tetraodon miurus Boulenger 1902, and Tetraodon duboisi Poll 1959. Two molecular studies by Yamanoue et al. (2011) and Igarashi et al. (2013) produced essentially the same phylogenetic tree for freshwater pufferfishes.

In his review of ocean sunfishes, Fraser-Brunner (1951) found Masturus Gill 1884 and Mola to be more closely related to each other than to Ranzania. Tyler’s (1980) extensive surveys of osteological characters supported the phylogenetic tree of Fraser-Brunner (1951). Santini and Tyler (2002a) performed the first cladistic analysis of the phylogenetic relationships of ocean sunfishes, using 48 morphological characters. Their study supported the results of the previous studies. Yamanoue et al. (2004) used mitochondrial genomes to analyze the phylogenetic relationships of ocean sunfishes. Their study resulted in the same phylogenetic tree as recovered by previous morphological studies. Molecular studies on tetraodontiform relationships by Alfaro et al. (2007) also produced the same result. Bass et al. (2005) examined many specimens of ocean sunfishes captured from various localities worldwide. The molecular analysis found that M. ramsayi is distributed in the Southern Hemisphere and M. mola in both the Northern and Southern hemispheres. Yoshita et al. (2009) studied ocean sunfishes of Mola around Japan employing both morphological and molecular characters. As discussed in the section of Molidae, the study clearly showed two species of Mola around Japan, probably identifiable as M. mola and M. ramsayi.

Concluding remarks

The taxonomy of tetraodontiforms has progressed greatly since the mid-1960s. On the basis of contributions to date, this paper identifies 412 extant species in the 10 living families of Tetraodontiformes, with an allocation of species and genera as follows: Triacanthodidae including 23 species in 11 genera, Triacanthidae with seven species in four genera, Balistidae with 37 species in 12 genera, Monacanthidae with 102 species in 27 genera, Aracanidae with 13 species in six genera, Ostraciidae with 22 species in five genera, monotypic Triodontidae, Tetraodontidae with 184 species in 27 genera, Diodontidae with 18 species in seven genera, and Molidae with five species in three genera. However, many taxonomic problems involving species and genera of various families still remain. In particular, taxonomic anomalies exist in the Monacanthidae and Tetraodontidae, which have the greatest number of species and diversity within the order. Studies of spikefishes of the Triacanthodidae reveal clarity on the number of genera and species, but poorly surveyed deep areas of the world’s oceans exist where new species may occur. Several genera of Balistidae, Aracanidae, and Ostraciidae require study both by morphological and molecular analyses to clarify generic and species limits. The large size attained by ocean sunfishes has made it difficult for us to have a clear understanding of species limits, although a combination of molecular and morphological approaches have clarified that the presence of several populations probably represent species of Mola (see Bass et al. 2005; Yoshita et al. 2009). When the taxonomic levels of these populations are established, difficult nomenclatural challenges still remain: many nominal species of ocean sunfishes were published in the old days without type specimens.

With regard to the systematics of the Tetraodontiformes overall, familial relationships have been clarified by morphological and molecular analyses, providing us with a firm understanding of the sister relationships of the following family groups: Triacanthodidae and Triacanthidae, Balistidae and Monacanthidae, and Tetraodontidae and Diodontidae. However, the phylogenetic positions of the Triodontidae and Molidae remain unclear because of conflicts about their positions in morphological and molecular studies (the position of Molidae differs even among molecular studies). More taxon sampling is needed for molecular analyses to produce more robust phylogenetic topologies. Although generic and specific phylogenetic relationships for the families Triacanthidae, Balistidae, Monacanthidae, Aracanidae, Ostraciidae, and Molidae have been studied using morphological and/or molecular analyses, the genera and species of the Triacanthodidae and Diodontidae have never been studied cladistically and await future studies.

Notes

Acknowledgments

I am grateful to the members of the Organizing Committee of the 9th Indo-Pacific Fish Conference for inviting me to write this paper. I thank numerous people in museums and universities for supporting my research on tetraodontiform fishes. Ralf Britz in the Natural History Museum kindly provided access to old literature. Martin F. Gomon of the Museum Victoria and Edward O. Murdy of the George Washington University kindly provided comments on the draft manuscript. My thanks go to James C. Tyler and an anonymous reviewer for providing helpful comments to the manuscript. Mark McGrouther of the Australian Museum kindly photographed the holotype of Acanthostracion bucephalus. I also thank Hiromitsu Endo and Naohide Nakayama of BSKU, Hiroyuki Motomura and his students of KAUM, Eri Katayama of NSMT, and Hideki Sugiyama, for providing me with beautiful photographs of tetraodontiforms. This study was partially supported by Grant-in-Aids for Scientific Researches, A (19208019) and B (24370041) from the Japan Society for the Promotion of Science; Grant-in-Aid for Scientific Research on Innovative Areas of the Ministry of Education, Culture, Sports, Science and Technology (MEXT), 24120001; Health Labour Sciences Research Grant (FY2013-2014) of the Ministry of Health, Labour and Welfare, Japan; and the “Biological Properties of Biodiversity Hotspots in Japan” Project of the National Museum of Nature and Science.

References

  1. Abe T (1939) Notes on Sphoeroides xanthopterus (Temminck et Schlegel) (Tetraodontidae, Teleostei). Zool Mag 51:334−337Google Scholar
  2. Abe T (1942) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—I. Vertebral variation. Palao Tropical Biol Stn Stud 2:477−496Google Scholar
  3. Abe T (1944) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—II. Variation of dorsal fin. Ann Zool Japon 22:200−234Google Scholar
  4. Abe T (1949a) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—III. Variation of anal fin. Bull Jpn Soc Sci Fish 14:123−144Google Scholar
  5. Abe T (1949b) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—IV. Variation of caudal fin. Bull Jpn Soc Sci Fish 15:19−27Google Scholar
  6. Abe T (1949c) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—V. Synopsis of the puffers from Japan and adjacent regions. Bull Biogeogr Soc Japan 14:89−140, pls 1−2Google Scholar
  7. Abe T (1950) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—VI. Variation of pectoral fin (with some additions to the previous reports of the present series). Jpn J Ichthyol 1:198−206Google Scholar
  8. Abe T (1952) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions—VII. Concluding remarks, with the introduction of two new genera, Fugu and Boesemanichthys. Jpn J Ichthyol 1:35−44Google Scholar
  9. Abe T (1954) Taxonomic studies on the puffers (Tetraodontidae, Teleostei) from Japan and adjacent regions–Corrigenda and addenda. Jpn J Ichthyol 1:121−128, pl 1Google Scholar
  10. Abe T, Tabeta O (1983) Description of a new swellfish of the genus Lagocephalus (Tetraodontidae, Teleostei) from Japanese waters and the East China Sea. Uo 32:1−8, pls 1−3Google Scholar
  11. Abe T, Tabeta O, Kitahama K (1984) Notes on some swellfishes of the genus Lagocephalus (Tetraodontidae, Teleostei) with description of a new species from Japan. Uo 34:1−10, pls 1−3Google Scholar
  12. Adam MS, Merrett NR, Anderson RC (1998) Additions to the fish fauna of the Maldive Islands. Part 1: An annotated checklist of the deep demersal fishes of the Maldive Islands. Ichthyol Bull JLB Smith Inst Ichthyol 67:1−19Google Scholar
  13. Alcock AW (1894) Natural history notes from H. M. Indian marine survey steamer `Investigator,’ Commander C. F. Oldham, R. N., commanding. Series II, No. 11. An account of a recent collection of bathybial fishes from the Bay of Bengal and from the Laccadive Sea. J Asiatic Soc Bengal 63:115−137, pls 6−7Google Scholar
  14. Alcock AW (1899) Halimochirurgus centriscoides, a new deep-sea fish from the Gulf of Manár. Proc Asiatic Soc Bengal 1899:78Google Scholar
  15. Alfaro ME, Santini, Brock CD (2007) Do reefs drive diversification in marine teleosts? Evidence from the pufferfish and their allies (Order Tetraodontiformes). Evolution 61:2104–2126Google Scholar
  16. Allen GR (1997) Marine fishes of tropical Australia and south-east Asia. Western Australian Museum, PerthGoogle Scholar
  17. Allen GR, Erdman MV (2012) Reef fishes of the East Indies. Volumes I−III. Tropical Reef Research, PerthGoogle Scholar
  18. Allen GR, Randall JE (1977) Review of the sharpnose pufferfish (Subfamily Canthigasterinae) of the Indo-Pacific. Rec Aust Mus 30:475−517 Google Scholar
  19. Allen GR, Robertson DR (1994) Fishes of the tropical eastern Pacific. University of Hawai’i Press, HonoluluGoogle Scholar
  20. Allen GR, Swainston R (1988) The marine fishes of north-western Australia. A field guide for anglers and divers. Western Australian Museum, PerthGoogle Scholar
  21. Allen GR, Cross NJ, Allen CJ (2006) Ostraciidae. In: Beesley PL, Wells A (eds) Zoological Catalogue of Australia. Volume 35, Part 3. Fishes. ARBS & CSIRO Publishing, Collingwood, pp 1901−1909Google Scholar
  22. Amaoka (1982) Triacanthodidae. In: Okamura O, Amaoka K, Mitani F (eds) Fishes of the Kyushu-Palau Ridge and Tosa Bay. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp 302−304, 409Google Scholar
  23. Amaral CRL, Brito PM, Silva DA, Carvalho EF (2013) A new cryptic species of South American freshwater pufferfish of the genus Colomesus (Tetraodontidae), based on both morphology and DNA data. PLoS ONE 8:1−15Google Scholar
  24. Anonymous (1798) ‘Histoire naturelle des poissons’ by Lacepède (1798). Allgemeine Literatur-Zeitung 3(287): columns 673−680Google Scholar
  25. Baldwin CC (2013) The phylogenetic significance of colour patterns in marine teleost larvae. Zool J Linn Soc 168:496−563Google Scholar
  26. Bannikov AF, Tyler JC (2008a) A new genus and species of triggerfish from the middle Eocene of the northern Caucasus, the earliest member of the Balistidae (Tetraodontiformes). Paleontol J (Moscow) 42:615−620Google Scholar
  27. Bannikov AF, Tyler JC (2008b) A new species of the pufferfish Eotetraodon from the Eocene of the northern Caucasus (Tetraodontiformes, Tetraodontidae). Paleontol J (Moscow) 42:526−530Google Scholar
  28. Barnard KH (1927) Diagnoses of new genera and species of South African marine fishes. Ann Mag Natr Hist (Ser 9) 20:66–79Google Scholar
  29. Basilewsky S (1855) Ichthyographia Chinae borealis. Nouveaux mémoires de la Société impériale des naturalistes de Moscou 10:215−263, pls 1−9Google Scholar
  30. Bass AL, Dewar H, Thys T, Streelman JT, Karl SA (2005) Evolutionary divergence among lineages of the ocean sunfish family, Molidae (Tetraodontiformes). Mar Biol 148:405–414Google Scholar
  31. Bean TH (1906) Descriptions of new Bermudan fishes. Proc Biol Soc Wash 19:29−33Google Scholar
  32. Beaufort LFde, Briggs J C (1962) The fishes of the Indo-Australian Archipelago XI. Scleroparei, Hypostomides, Pediculati, Plectognathi, Opisthomi, Discocephali, Xenopterygii. EJ Brill, LeidenGoogle Scholar
  33. Benl G (1957) Carinotetraodon chlupatyi nov. gen., nov. spec., ein Kugelfisch mit Kamm und Kiel [Pisces, Fam. Tetraodontidae] (vorläufige Mitteilung). Opscula Zool 5:1−4Google Scholar
  34. Bennett ET (1831) A small collection of fishes, formed during the voyage of H.M.S. Chanticleer, and presented to the Society by the Lords Commissioners of the Admiralty, together with numerous other Zoological specimens obtained during the same voyage, was laid upon the table. It contained among others a young individual of the Scyllium cirratum, in the statė in which it is described by Schneider as the Sąualus punctatus: a specimen of the Blennius pilicornis, Cuv., described originally by Marcgrave, and remarkable for the long acicular tooth at the back of the lower jaw on each side, a peculiarity which may hereafter cause it to be regarded as the type of a distinct genus: a specimen of the Antennarius scaber, Chironectes scaber, Cuv., also described by Marcgrave: and two species which appeared to be new to science, and which were thus characterized by Mr. Bennett. Proc Commit Sci Correspond Zool Soc London 1830−31:112Google Scholar
  35. Bennett ET (1832) Observations on a collection of fishes from the Mauritius, presented by Mr. Telfair, with characters of new genera and species. Proc Commit Sci Correspond Zool Soc London 1830−31:165−169Google Scholar
  36. Berry FH, Baldwin WJ (1966) Triggerfishes (Balistidae) of the eastern Pacific. Proc Calif Acad Sci (Ser 4) 34:429–474Google Scholar
  37. Berry FH, Vogele LE (1961) Filefishes (Monacanthidae) of the western North Atlantic. Fish Bull 61:61–109Google Scholar
  38. Bianconi GG (1846) Lettera [sul Ostracion fornasini, n. sp. de pesce del Mosambico]. Nuovi Annali delle Scienze naturali Bologna (Ser 2) 5:113–115, pl 1Google Scholar
  39. Bleeker, P. (1848) A contribution to the knowledge of the ichthyological fauna of Sumbawa. J Ind Archipel Eastern Asia (Singapore) 2:632–639Google Scholar
  40. Bleeker P (1850a) Over drie nieuwe soorten van Tetraödon van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië 1:96−97Google Scholar
  41. Bleeker P (1850b) Bijdrage tot de kennis der Blootkakige visschen van den Soenda-Molukschen Archipel. Verhandelingen van het Bataviaasch Genootschap van Kunsten en Wetenschappen. 24:1–20 + 21–26Google Scholar
  42. Bleeker P (1851a) Bijdrage tot de kennis der Balistini en Ostraciones van den Indischen Archipel. Verhandelingen van het Bataviaasch Genootschap van Kunsten en Wetenschappen 24:1−38, pls 1−7Google Scholar
  43. Bleeker P (1851b) Bijdrage tot de kennis der ichthyologische fauna van de Banda-eilanden. Natuurkundig Tijdschrift voor Nederlandsch Indië 2:225−261Google Scholar
  44. Bleeker P (1851c) Bijdrage tot de kennis der ichthyologische fauna van Riouw. Natuurkundig Tijdschrift voor Nederlandsch Indië 2: 469−497Google Scholar
  45. Bleeker P (1852a) Bijdrage tot de kennis der ichthyologische fauna van Singapore. Natuurkundig Tijdschrift voor Nederlandsch Indië 3:51−86Google Scholar
  46. Bleeker P (1852b) Bijdrage tot de kennis der ichthyologische fauna van het eiland Banka. Natuurkundig Tijdschrift voor Nederlandsch Indië 3: 443−460Google Scholar
  47. Bleeker P (1853) Vierde bijdrage tot de kennis der ichthyologische fauna van Celebes. Natuurkundig Tijdschrift voor Nederlandsch Indië 5:153−174Google Scholar
  48. Bleeker P (1854) Vijfde bijdrage tot de kennis der ichthyologische fauna van Celebes. Natuurkundig Tijdschrift voor Nederlandsch Indië 7:225−260 Google Scholar
  49. Bleeker P (1856) Bijdrage tot de kennis der ichthyologische fauna van het eiland Nias. Natuurkundig Tijdschrift voor Nederlandsch Indië 12:211–228Google Scholar
  50. Bleeker P (1857) Vierde bijdrage tot de kennis der ichthyologische fauna van Japan. Acta Soc Sci Indo-Neerlandicae 3:1−46, pls 1−4Google Scholar
  51. Bleeker P (1863) Mémoire sur les poissons de la côte de Guinée. Natuurkundige Verhandelingen van de Hollandsche Maatschappij der Wetenschappen te Haarlem (Ser 2) 18:1–136, pls 1–28Google Scholar
  52. Bleeker P (1865) Systema Balistidorum, Ostracionidorum, Gymnodontidorumque revisum. Nederlandsch Tijdschrift voor de Dierkunde 3:8–19Google Scholar
  53. Bleeker P (1865−1869) Atlas ichthyologique des Indes Orientales Néêrlandaises, publié sous les auspices du Gouvernement colonial néêrlandais. Tome V. Baudroies, Ostracions, Gymnodontes, Balistes. Frédemic Muller, Éditeur, AmsterdamGoogle Scholar
  54. Bleeker P (1873) Description et figure d’une espèce insulindienne d’Orthagoriscus. Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen. Afdeeling Natuurkunde (Ser 2) 7:151−153Google Scholar
  55. Bloch ME (1785) Naturgeschichte der ausländischen Fische. Vol 1. Schlesinger, BerlinGoogle Scholar
  56. Bloch ME (1786) Naturgeschichte der ausländischen Fische. Vol 2. Schlesinger, BerlinGoogle Scholar
  57. Bloch ME, Schneider JG (1801) M. E. Blochii, Systema Ichthyologiae Iconibus cx Ilustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo. Berolini. Sumtibus Auctoris Impressum et Bibliopolio Sanderiano Commissum, Berlin.Google Scholar
  58. Böhlke JE, Chaplin CG (1968) Fishes of the Bahamas and adjacent tropical waters. University of Texas Press, AustinGoogle Scholar
  59. Bonnaterre JP (1788) Tableau encyclopédique et methodique des trois règnes de la nature. Ichthyologie. Panckoucke, ParisGoogle Scholar
  60. Boulenger GA (1888) An account of the fishes obtained by Surgeon-Major A.S.G. Jayakar at Muscat, east coast of Arabia. Proc Zool Soc London 1887: 653−667, pl 54Google Scholar
  61. Boulenger GA (1889) Second account of the fishes obtained by Surgeon-Major A.S.G. Jayakar at Muscat, east coast of Arabia. Proc Zool Soc London 1889:236−246, pls 25−28Google Scholar
  62. Boulenger G A (1899) Matériaux pour la faune du Congo. Poissons nouveaux du Congo. Troisième Partie. Silures, Acanthoptérygiens, Mastacembles, Plectognathes. Annales du Musee du Congo (Ser Zool) 1:39–58, pls 20–29Google Scholar
  63. Boulenger GA (1902) Additions à la faune ichthyologique de bassin du Congo. Matériaux pour la faune du Congo. Annales du Musee du Congo (Ser Zool) 2:19–57, pls 7–16Google Scholar
  64. Bray D (2008) Family Molidae. In: Gomon M, Bray D, Kuiter R (eds) Fishes of Australia’s southern coast. New Holland Publishers Pty Ltd, Sydney, pp 858–861Google Scholar
  65. Brisout de Barneville CNF (1846) Note sur les Diodoniens. Revue Zoologique par la Société Cuvierienne (Paris) 1846:136−143Google Scholar
  66. Britz R, Johnson GD (2005) Occipto-vertebral fusion in ocean sunfishes (Teleostei: Tetraodontiformes: Molidae) and its phylogenetic implications. J Morphol 266:74−79Google Scholar
  67. Britz R, Kottelat M (1999) Carinotetraodon imitator, a new freshwater pufferfish from India (Teleostei: Tetraodontiformes). J South Asian Natr Hist 4:39−47Google Scholar
  68. Caldwell DK, Randall JE (1967) Cantherhines tiki, a junior synonym of the Easter Island filefish, Cantherhines rapanui. Copeia 1967:857−858Google Scholar
  69. Cantor TE (1849) Catalogue of Malayan fishes. J Asiatic Soc Bengal 18:i–xii + 983–1443, pls 1–14Google Scholar
  70. Carnevale G, Tyler JC (2010) Review of the fossil pufferfish genus Archaeotetraodon (Teleostei, Tetraodontidae), with description of three new taxa from the Miocene of Italy. Giobios 43:283–304Google Scholar
  71. Castelnau FL (1861) Mémoire sur les poissons de l’Afrique australe. J.-B. Bailliere, ParisGoogle Scholar
  72. Castelnau FL (1873) Contribution to the ichthyology of Australia. Proc Zool Acclim Soc Vic Melbourne 2:37−158Google Scholar
  73. Chanet B, Guintar C, Betti E, Gallut C, Dettaï, Lecointre G (2013) Evidence for a close phylogenetic relationship between the teleost orders Tetraodontiformes and Lophiiformes based on an analysis of soft anatomy. Cybium 37:179−198Google Scholar
  74. Chaudhuri BL (1910) Triacanthus weberi, sp. nov. J Proc Asiatic Soc Bengal (New Ser) 6:497−501Google Scholar
  75. Chen, QC, Cai YZand Ma XM (eds) (1997) Fishes from Nansha Islands to South China coastal waters 1. Science Press, BeijingGoogle Scholar
  76. Cheng QT, Wang CX, Tian MC, Li CS, Wang YG, Wang Q (1975) Studies on the Chinese tetraodonoid fishes of the genus Fugu. Acta Zool Sinica 21:359−378Google Scholar
  77. Chu YT (1935) Description of a new species of Lagocephalus from Chusan, China. China J 22:87 Google Scholar
  78. Clark E, Gohar HAF (1953) The fishes of the Red Sea: Order Plectognathi. Publ Mar Biol Stat Al Ghardaqa 8:1−80, pls 1−5Google Scholar
  79. Cloquet H (1816) Dictionnaire des sciences naturelles, vol. 1 (Pisces accounts). F.G. Levrault, StrasbourgGoogle Scholar
  80. Cope ED (1871) Contribution to the ichthyology of the Lesser Antilles. Trans Amer Phil Soc (New Ser) 14:445–483Google Scholar
  81. Cuvier G (1816) Le Règne Animal distribué d’après son organisation pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Les reptiles, les poissons, les mollusques et les annélides. Edition 1. Déterville, ParisGoogle Scholar
  82. Cuvier G (1818) Sur les Diodons, vulgairement orbes-épineux. Mémoires du Muséum National d’Histoire Naturelle, Paris 4:121−138, pls 6−7Google Scholar
  83. Cuvier G (1829) Le Règne Animal, distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Edition 2. Déterville, ParisGoogle Scholar
  84. Dao VH, Nguyen TD, Nguyen TH, Takata Y, Sato S, Kodama M, Fukuyo Y (2012) High individual variation in the toxicity of three marine puffer in Vietnam. Coastal mar Sci 35:1−6Google Scholar
  85. de Buen F (1963) Los peces de la Isla de Pascua. Catálogo descriptivo e ilustrado. Boletin de la Sociedad de Biologia de Concepción 35/36:3−80Google Scholar
  86. Dekkers WJ (1975) Review of the Asiatic freshwater puffers of the genus Tetraodon Linnaeus, 1758 (Pisces, Tetraodontiformes, Tetraodontidae). Bijdragen tot de Dierkunde 45:87−142Google Scholar
  87. Donovan E (1823−27) The naturalist's repository; or Monthly miscellany ... 5 vols. LondonGoogle Scholar
  88. Duméril C (1855) Note sur un travail inédit de Bibron relative aux poisons plectognathes gymnodontees (diodons et tétrodons). Rev Mag Zool (Sér 2) 8:274−282Google Scholar
  89. Forsskål PS (1775) Descriptiones animalium, avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit Petrus Forskål. Post mortem auctoris edidit Carsten Niebuhr. Adjuncta est materia medica kahirina atque tabula maris Rubri geographica. ex officina Mölleri, HauniaeGoogle Scholar
  90. Fowler HW (1928) The fishes of Oceania. Mem B P Bishop Mus 10:i–iii + 1–540, pls 1–49Google Scholar
  91. Fowler HW (1929) New and little-known fishes from the Natal coast. Ann Natal Mus 6:245−264Google Scholar
  92. Fowler HW (1931) The fishes of Oceania—Supplement I. Mem Bernice P. Bishop Mus 11:313–381Google Scholar
  93. Fowler HW (1934) Descriptions of new fishes obtained 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Proc Acad Natl Sci Philad 85:233−367Google Scholar
  94. Fraser-Brunner A (1935a) Notes on the Plectognath fishes—I. A synopsis of the genera of the family Balistidae. Ann Mag Natr Hist Ser 10, 15:658−663Google Scholar
  95. Fraser-Brunner A (1935b) Notes on the Plectognath fishes—II. A synopsis of the genera of the family Ostraciontidae. Ann Mag Natr Hist Ser 10, 16:313−320Google Scholar
  96. Fraser-Brunner A (1940) The fishes of the genus Pseudomonacanthus with descriptions of two new species. Bull Raffles Mus 16:62–67, pls 21–23Google Scholar
  97. Fraser-Brunner A (1941a) Notes on the plectognath fishes —IV. A synopsis of the genera of the family Aluteridae, and descritptions of seven new species. Ann Mag Natr Hist Ser 11 8:176−199Google Scholar
  98. Fraser-Brunner A (1941b) Notes on the plectognath fishes —V. The families of triacanthiform fishes, with a synopsis of the genera and description of a new species. Ann Mag Natr Hist Ser 11 7:420−430Google Scholar
  99. Fraser-Brunner A (1941c) Notes on the plectognath fishes—VI. A synopsis of the genera of the family Aluteridae, and descriptions of seven new species. Ann Mag Natr Hist Ser 11 8:176−199Google Scholar
  100. Fraser-Brunner A (1941d) Note on the Plectognath fishes—VII. The Aracanidae, a distinct family ostraciontoid fishes, with descriptions of two new species. Ann Mag Natr Hist Ser 18 8: 306−313Google Scholar
  101. Fraser-Brunner A (1943) Note on the plectognath fishes. —VIII. The classification of the suborder Tetraodontoidea, with a synopsis of the genera. Ann Mag Natr Hist Ser 11 10:1−18Google Scholar
  102. Fraser-Brunner A (1950) Studies in plectognath fishes from the “Dana”-Expeditions. I. An interesting new genus of triacanthodid fishes from the Celebes Sea. Dana Rep 35:1−8Google Scholar
  103. Fraser-Brunner A (1951) The ocean sunfishes (Family Molidae). Bull Brit Mus (Natr Hist) Zool 1:89−121Google Scholar
  104. Fricke R (1999) Fishes of the Mascarene Islands (Réunion, Mauritius, Rodriguez). An annotated checklist with descriptions of new species. Koeltz Scientific Books, KönigsteinGoogle Scholar
  105. Fricke R, Mulochau T, Durville P, Chabanet P, Tessier E, Letourneur Y (2009) Annotated checklist of the fish species (Pisces) of La Réunion, including a Red List of threatened and declining species. Stuttgarter Beiträge zur Naturkunde A (Neue Serie) 2:1−168Google Scholar
  106. Fricke R, Eschmeyer WN (2014) A guide to fish collections in the Catalog of fishes. Online version, updated 27 August 2014. http://research.calacademy.org/redirect?url=http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. Accssessed 1 September 2014
  107. Fujii E, Uyeno T (1979) Polyplacapros tyleri, a new genus and species of ostraciid trunkfish from off eastern Australia and Norfolk Ridge. Jpn J Ichthyo 26:1−10Google Scholar
  108. Giglioli EH (1883) Zoology at the Fisheries Exhibition. II—Notes on the Vertebrata. Nature 28:313−316Google Scholar
  109. Gilbert CH (1905) Section II. The deep-sea fishes of the Hawaiian Islands. In: Jordan DS, Evermann BW (eds) The aquatic resources of the Hawaiian Islands. Bull US Fish Comm 23: 577−713, pls 66−101Google Scholar
  110. Gilbert CH, Starks EC (1904) The fishes of Panama Bay. Mem Calif Acad Sci 4:1−304, pls. 1−33Google Scholar
  111. Gill TN (1861) On several new generic types of fishes contained in the museum of the Smithsonian Institution. Proc Acad Natl Sci Phil 13:77–78Google Scholar
  112. Gill TN (1884) Synopsis of the plectognath fishes. Proc US Natl Mus 7:411–427Google Scholar
  113. Gistel J (1848) Naturgeschichte des Thierreichs, für höhere Schulen. Stuttgart.Google Scholar
  114. Gloerfelt-Tarp T, Kailola PJ (1984) Trawled fishes of southern Indonesia and northwestern Australia. Australian Development Assistance Bureau, Canberra, Directorate General of Fisheries, Jakarta and German Agency for Technical Cooperation, EschbornGoogle Scholar
  115. Gmelin JF (1789) Caroli a Linné. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species; cum characteribus, differentiis, synonymis, locis. Editio decimo tertia, aucta, reformata 1:1033−1516Google Scholar
  116. Gomon F (2008) Family Diodontidae. In: Gomon M, Bray D, Kuiter R (eds) Fishes of Australia’s southern coast. New Holland Publishers Pty Ltd, Sydney, pp 857−858Google Scholar
  117. Gray JE (1830−35) Illustrations of Indian zoology; chiefly selected from the collection of Major-General Hardwicke, F.R.S., Vols 1–2. Adolphus Richter and Co., LondonGoogle Scholar
  118. Gray JE (1831) Description of twelve new genera of fish, discovered by Gen. Hardwicke, in India, the greater part in the British Museum. Zool Misc 1831:7−9Google Scholar
  119. Gray JE (1838) Notes on the fish. Annals Mag Natr Hist New Ser 1:109−111Google Scholar
  120. Gregorova R, Schultz O, Harzhauser M, Kroh A, Coric S (2009) A giant Miocene sunfish from the North Alpine Foreland Basin (Austria) and its implication for molid phylogeny. J Vert Paleo 29:359–371Google Scholar
  121. Guichenot A (1853) Poissons. In: Sagra MR de la (ed) Histoire physique, politique et naturelle de l’Ile de Cuba. Vol 2. Arthus Bertrand, ParisGoogle Scholar
  122. Günther A (1870) Catalogue of the fishes in the British Museum. Vol 8. Catalogue of the Physostomi, containing the families Gymnotidae, Symbranchidae, Muraenidae, Pegasidae, and of the Lophobranchii, Plectognathi, Dipnoi, Ganoidei, Chondropterygii, Cyclostomata, Leptocardii, in the British Museum. LondonGoogle Scholar
  123. Günther A (1874) Third notice of a collection of fishes made by Mr. Swinhow in chian. Ann Mag Natr Hist (Ser 4) 13:154–159Google Scholar
  124. Günther A (1877) Preliminary notes on new fishes collected in Japan during the expedition of H. M. S. `Challenger.’ Ann Mag Natr Hist (Ser 4) 20:433–446Google Scholar
  125. Günther A (1880) Report on the shore fishes procured during the voyage of H. M. S. Challenger in the years 1873-1876. Report on the scientific results of the voyage of H. M. S. Challenger during the years 1873-76, zoolgy. 1:1−82, pls. 1−32Google Scholar
  126. Hamilton F (1822) An account of the fishes found in the river Ganges and its branches. A. Constable, Edinburgh and LondonGoogle Scholar
  127. Hardy GS (1980) A redescription of the antitropical pufferfish Arothron firmamentum (Plectognathi: Tetraodontidae). NZ J Zool 7:115−125Google Scholar
  128. Hardy GS (1981) A redescription of the pufferfish Contusus richei (Tetraodontiformes: Tetraodontidae), and description of a second species of Contusus. NZ J Zool 8:11−23Google Scholar
  129. Hardy GS (1982a) Two new generic names for some Australian pufferfishes (Tetraodontiformes: Tetraodontidae), with species’ redescriptions and osteological comparisons. Aust Zool 21:1−26Google Scholar
  130. Hardy GS (1982b) First Pacific records of Pelagocephalus marki Heemstra & Smith (Tetraodontiformes: Tetraodontidae), and first male specimen of the genus. NZ J Zool 9:377−380Google Scholar
  131. Hardy GS (1983a) Revision of Australian species of Torquigener Whitley (Tetraodontiformes: Tetraodontidae), and two new generic names for Australian puffer fishes. J Roy Soc NZ 13:1−48Google Scholar
  132. Hardy GS (1983b) The status of Torquigener hypselogeneion (Bleeker) (Tetraodontiformes: Tetraodontidae) and some related species, including a new species from Hawaii. Pacif Sci 37:65−74Google Scholar
  133. Hardy GS (1984a) Tylerius, a new generic name for the Indo-Pacific pufferfish, Spheroides spinosissimus Regan, 1908 (Tetraodontiformes: Tetraodontidae) and comparisons with Amblyrhynchotes (Bibron) Duméril. Bull Mar Sci 35:32−37Google Scholar
  134. Hardy GS (1984b) Redescription of the pufferfish Torquigener brevipinnis (Regan) (Tetraodontiformes: Tetraodontidae), with description of a new species of Torquigener from Indonesia. Pacif Sci 38:127−133Google Scholar
  135. Hardy GS (1985) A new genus and species of pufferfish (Tetraodontidae) from Java. Bull Mar Sci 36:145−149Google Scholar
  136. Hardy GS (1989) Description of a new species of Torquigener Whitley (Pisces: Tetraodontidae) from South Africa, with a key to the genus. Natl Mus NZ Rec 3:119−123Google Scholar
  137. Hardy GS, Hutchins JB (1981) On the validity of the pufferfish genus Omegophora Whitley (Tetraodontiformes: Tetraodontidae) with the description of a new species. Rec West Aust Mus 9:187−201Google Scholar
  138. Hardy, GS, Randall JE (1983) Description of a new species of pufferfish (Tetraodontiformes: Tetraodontidae) from the Red Sea and adjacent waters. Israel J Zool 32:13−20Google Scholar
  139. Hartel KE, Kenaley CP, Galbraith JK, Sutton TT (2008) Additional records of deep-sea fishes from off greater New England. Northeastern Naturalist 15:317−334Google Scholar
  140. Hayashi M, Hagiwara K (2013) Ostraciidae. In: Nakabo T (ed) Fishes of Japan with pictorial keys to the species, third edition. Tokai University Press, Hadano, pp 1723−1726, 2238−2239Google Scholar
  141. Heemstra PC, Smith MM (1981) Pelagocephalus marki, a new species of puffer fish (Tetraodontidae) from South Africa. Bull Mar Sci 31:911−915Google Scholar
  142. Heemstra PC, Smith MM (1983) A new species of the triggerfish genus Xenobalistes Matsuura (Tetraodontiformes: Balistidae) from South Africa. JLB Smith Inst Ichthyol Spec Publ 26:1−5Google Scholar
  143. Herre AWCT (1927) A new genus and three new species of Philippine fishes. Philipp J Sci 32:413–419, pls 1–2Google Scholar
  144. Hoese DF, Bray DJ, and Paxton JR (2006) Triacanthodidae and Triacanthidae. In: Beesley PL, Wells A (eds) Zoological Catalogue of Australia. Volume 35, Part 3. Fishes. ARBS & CSIRO Publishing, Collingwood, pp 1861−1867Google Scholar
  145. Holcroft NI (2005) A molecular analysis of the interrelationships of tetraodontiform fishes (Acanthomorpha: Tetraodontiformes). Mol Phylogenet Evol 34:525−544Google Scholar
  146. Hollard HLGM (1854) Monographie de la famille des Balistides. Suite 2. Annales des Sciences Naturelles, Paris (Zoologie) (Sér 4) 1:39−72, 303−339, pls 2−3Google Scholar
  147. Hollard HLGM (1855) Monographie de la famille des balistides. (Suite et fin). Annales des Sciences Naturelles, Paris (Zoologie) (Sér 4) 4:5−27, pl 1Google Scholar
  148. Houttuyn M (1782) Beschryving van eenige Japanese visschen, en andere zee-schepzelen. Verhandelingen der Hollandsche Maatschappij der Wetenschappen, Haarlem 20:311−350Google Scholar
  149. Hulley PA (1972) The rare plectognath fish, Macrorhamphosodes uradoi (Kamohara) (Triacanthodidae) in South African waters. Ann S Afr Mus 60:191−195Google Scholar
  150. Hutchins JB (1977) Descriptions of three new genera and eight new species of monacanthid fishes from Australia. Rec West Aust Mus 5:3−58Google Scholar
  151. Hutchins JB (1986a) Review of the monacanthid fish genus Pervagor, with descriptions of two new species. Indo-Pacific Fishes (12):1–35, pls 1–2Google Scholar
  152. Hutchins JB (1986b) Family No. 264: Balistidae. In: Smith MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan South Africa, Johannesburg, pp 882−887, pls 137−140Google Scholar
  153. Hutchins JB (1987) New Australian fishes. Part 12. A new species of Eubalichthys (Monacanthidae). Mem Mus Vic 48:51−52Google Scholar
  154. Hutchins JB (1994a) Monacanthidae. In: Gomon, MF, Glover JCM, Kuiter RH (eds) The fishes of Australia’s south coast. State Print, Delaide, pp 866−891Google Scholar
  155. Hutchins JB (1994b) Description of a new genus and species of monacanthid fish from India. Rec West Aust Mus 16:567−574Google Scholar
  156. Hutchins JB (1997) Review of the monacanthid fish genus Paramonacanthus, with descriptions of three new species. Rec West Aust Mus Suppl 54:1–57Google Scholar
  157. Hutchins JB (2001a) Checklist of the fishes of Western Australia. Rec West Aust Mus Suppl No. 63: 9−50Google Scholar
  158. Hutchins JB (2001b) Monacanthidae and Molidae. In: Carpenter K, Niem (eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 6. Bony fishes part 4 (Labridae to Latimeriidae). FAO, Rome, pp 3929−3947, 3966−3968Google Scholar
  159. Hutchins JB (2002) Description of a new genus and species of miniature monacanthid fish from the Seychelles and Marshall Islands. Rec West Aust Mus 21:213−219Google Scholar
  160. Hutchins JB, Matsuura K (1984) Description of a new monacanthid fish of the genus Thamnaconus from Fiji. Rec West Aust Mus 11:387–391 Google Scholar
  161. Hutchins JB, Randall JE (1982) Cantherhines longicaudus, a new filefish from Oceania, with a review of the species of the C. fronticinctus complex. Pacif Sci 36:175−185Google Scholar
  162. Hutchins JB, Swainston R (1985) Revision of the monacanthid fish genus Brachaluteres. Rec West Aust Mus 12:57−78Google Scholar
  163. ICZN (Internationa Commision on Zoological Nomenclature) (1999) International code of zoological nomenclature, 4th ed. The International Trust for Zoological Nomenclature, LondonGoogle Scholar
  164. Igarashi Y, Doi H, Yamanoue Y, Kinoshita S, Ishibashi T, Ushio H, Asakawa S, Nishida M, Watabe S (2013) Molecular phylogenetic relationship of Tetraodon pufferfish based on mitochondrial DNA analysis. Fish Sci 79:243−250Google Scholar
  165. Inger RF (1953) A new fish from north Borneo. Fieldiana Zool 34:149−152Google Scholar
  166. Johnson GD, Britz R (2005a) Leis’ conundrum: homology of the clavus of the ocean sunfishes. 2. Ontogeny of the median fins and axial skeleton of Ranzania laevis (Teleostei, Tetraodontiformes, Molidae). J Morphol 266:11−21Google Scholar
  167. Johnson GD, Britz R (2005b) A description of the smallest Triodon on record (Teleostei: Tetraodontiformes: Triodontidae). Ichthyol Res 52:176−181Google Scholar
  168. Johnson JW (1999) Annotated checklist of the fishes of Moreton Bay, Queensland, Australia. Mem Qld Mus 43:709−762Google Scholar
  169. Jones S, Kumaran M (1980) Fishes of the Laccadive Archipelago. Nature Conservation and Aquatic Science Service, SantinivasGoogle Scholar
  170. Jordan DS (1895) The fishes of Sinaloa. Proc Calif Acad Sci (Ser 2) 5:377–514, pls 26–55Google Scholar
  171. Jordan DS (1916) The nomenclature of American fishes as affected by the opinions of the International Commission on Zoological Nomenclature. Copeia 29:25−28Google Scholar
  172. Jordan DS (1924) Rare species of fishes from the coast of southern California. Copeia 134: 81−82Google Scholar
  173. Jordan DS, Fowler HW (1902) A review of the trigger-fishes, file-fishes, and trunk-fishes of Japan. Proc US Natl Mus 25:251−286Google Scholar
  174. Jordan DS, Gilbert CH (1882) Notes on a collection of fishes made by Lieut. Henry E. Nichols, U. S. N., on the west coast of Mexico, with descriptions of new species. Proc US Natl Mus 4:225−233Google Scholar
  175. Jordan DS, Hubbs CL (1925) Record of fishes obtained by David Starr Jordan in Japan, 1922. Mem Carnegie Mus 10:93−346, pls 5−12Google Scholar
  176. Jordan DS, Seale A (1906) The fishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the fishes of Oceania. Bull Bur Fish 25 (for 1905):173−455 + 457−488, pls 33−53Google Scholar
  177. Jordan DS, Snyder JO (1901) Descriptions of nine new species of fishes contained in museums of Japan. J Col Sci Imp Univ Tokyo 15:301–311, pls 15–17Google Scholar
  178. Jordan EK (1925) Notes on the fishes of Hawaii, with descriptions of six new species. Proc US Natl Mus 66:1−43, pls. 1−2Google Scholar
  179. Kamohara T (1933) On a new fish from Japan. Zool Mag 45:389−393Google Scholar
  180. Kamohara T (1938) On the offshore bottom-fishes of Prov. Tosa, Shikoku, Japan. Maruzen Kabushiki Kaisha, TokyoGoogle Scholar
  181. Kamohara T (1941) Descriptions of one new and two rare fishes from Japan. Ann Zool Japon 20:166−168Google Scholar
  182. Kaup JJ (1855) Uebersicht über die Species einiger Familien der Sclerodermen. Archiv für Naturgeschichte 21: 215−233Google Scholar
  183. Kim IS, Choi Y, Lee CL, Lee YJ, Kim BJ, Kim JH (2005) Illustrated book of Korean fishes. Kyo−Hak Publishing Co., Ltd., SeoulGoogle Scholar
  184. Klassen GJ (1995) Phylogeny and biogeography of the Ostraciinae (Tetraodontiformes: Ostraciidae). Bull Mar Sci 57:393−441Google Scholar
  185. Koelreuter IT (1766) iscium rariorum e museo Petropolitano exceptorum descriptiones continuatae. Novi Commentarii Academiae Scientiarum Imperialis Petropolitanae 10:329–351, pl 8Google Scholar
  186. Kottelat M (1999) Nomenclature of the genera Barbodes, Cyclocheilichthys, Rasbora and Chonerhinos (Teleostei: Cyprinidae and Tetraodontidae), with comments on the definition of the first reviser. Raffles Bull Zool 47:591−600Google Scholar
  187. Kottelat M (2001) Fishes of Laos. WHT Publications, ColomboGoogle Scholar
  188. Kottelat M (2013) The fishes of the inland waters of southeast Asia: a catalogue and core bibiography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles Bull Zool Suppl 27:1−663Google Scholar
  189. Kottelat M, Whitten AJ, Kartikasari SN, Wirjoatmodjo S (1993) Freshwater fishes of western Indonesia and Sulawesi. Periplus Edition, Hong KongGoogle Scholar
  190. Kuang YD, Li CS, Liang SH (1984) A new species of the genus Takifugu (Tetraodontiformes)—Takifugu orbimaculatus. Trans Oceanol Limnol 4:58−61Google Scholar
  191. Kuiter RH (1993) Coastal fishes of south-eastern Australia. Universeity of Hawai’i Press, HonoluluGoogle Scholar
  192. Kuiter RH (1994) Aracanidae. In: Gomon, MF, Glover JCM, Kuiter RH (eds) The fishes of Australia’s south coast. State Print, Delaide, pp 892−902Google Scholar
  193. Kyushin K, Amaoka K, Nakaya K, Ida H (1977) Fishes of Indian Ocean. Japan Marine Fishery Resource Research Center, TokyoGoogle Scholar
  194. Kyushin K, Amaoka K, Nakaya K, Ida H, Tanino Y, Senta T (1982) Fishes of the South China Sea. Japan Marine Fishery Resource Research Center, TokyoGoogle Scholar
  195. Larson HK, Williams RS, Hammer MP (2013) An annotated checklist of the fishes of the Northern Territory, Australia. Zootaxa 3696:1−293Google Scholar
  196. Latreille PA (1804) Tableau méthodique des poissons. In: Nouveaux dictionnaire d’histoire naturelle, 1re éd. Déterville, Paris, pp 71−105Google Scholar
  197. Lauder GV, Liem KF (1983) The evolution and interrelationships of the actinopterygian fishes. Bull Comp Zool 150:95−197Google Scholar
  198. Lay GT, Bennett ET (1839) Fishes. In: Richardson J, Vigors NA, Lay GT, Bennett ET, Owen R, Gray JE, Burkland W, Sowerby GB (eds) The zoology of Captain Beechey’s voyage, compiled from the collections and notes made by Captain Beech EV, the officers and naturalist of the expedition, during a voyage to the Pacific and Behring’s Straits performed in his Majesty’s Ship Blossom, under the command of Captain F. W. Beechey, R.N., F.R.S., in the years 1825, 26, 27, and 28. Henty G. Bohn, London, pp 41−75, pls 15−23Google Scholar
  199. Le Danois Y (1955) Sur le remaniement du sous-ordre des poissons Plectognaths et la définition d’un nouveau sous-ordre: les Orbiculates. C R Acad Sci (Paris) 240:1933−1934Google Scholar
  200. Le Danois Y (1959) Etude ostéologique, myologique et systématique des poissons du sous-ordre des Orbiculates. Ann Inst Océanog 36:1−273Google Scholar
  201. Le Danois Y (1961) Remarques sur les poissons Orbiculates du sous-ordre des Ostracioniformes. Mém Mus Natl Hist Nat (Paris) (New Ser A) 19:207−338Google Scholar
  202. Leis JM (1978) Systematics and zeegeography of the porcupinefishes (Diodon, Diodontidae, Tetraodontiformes), with comments on egg and larval development. Fish Bull 76:535−567Google Scholar
  203. Leis JM (1984) Tetraodontiformes: relationships. In: Moser HG, Richards WJ, Cohen DM, Fahay MP, Kendall Jr AW, Richardson SL (eds) Ontogeny and systematics of fishes. Am Soc Ichthyol Herpetol Spec Publ 1, pp 459−463Google Scholar
  204. Leis JM (1986) Family No. 269: Diodontidae. In: Smith, MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan South Africa, Johannesburg, pp. 903−906, pl 144Google Scholar
  205. Leis JM (2001) Family Diodontidae. In: Carpenter K, Niem (eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 6. Bony fishes part 4 (Labridae to Latimeriidae). FAO, Rome, pp 3958−3965Google Scholar
  206. Leis JM (2003) Family Diodontidae. In: Carpenter KE (ed) The living marine resources of the Western Central Atlantic. Volume 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5. FAO, Rome, pp 2007−2010Google Scholar
  207. Leis JM (2006) Nomenclature and distribution of the species of the porcupinefish family Diodontidae (Pisces, Teleostei). Mem Mus Vic 63:77−90Google Scholar
  208. Lesson RP (1831) Poissons. In: Duperrey LI (ed) Voyage autour du monde : exécuté par ordre du roi, sur la corvette de Sa Majesté, la Coquille, pendant les années 1822, 1823, 1824, et 1825. Tome 2, zoologie. Arthus Bertrand, ParisGoogle Scholar
  209. Liénard E (1840) Description d’une nouvelle espèce du genre mole (Orthagoriscus, Schn.) découverte à l’île Maurice. Revue Zoologique par la Société Cuvierienne (Paris) 3:291−292Google Scholar
  210. Lim KKP, Kottelat M (1995) Carinotetraodon salivator, a new species of pufferfish from Sarawak, Malaysia (Teleostei: Tatraodontidae). Jpn J Ichthyol 41:359−365Google Scholar
  211. Lindberg GU, Fedorov VV, Krasyukova ZV (1997) Fishes of the Sea of Japan and the adjacent parts of the Sea of Okhotsk and Yellow Sea. Part 7. Teleostomi. Actinopterygii. Osteichthyes. XXXII. Dactylopteriformes—XXXVII. Pegasiformes. (CCII. Fam Dactylopteridae —CCXIX. Fam. Pegasidae). Handbook on the Identification of Animals, Zoological Institute of the Russian Academy of Sciences, St. PetersbergGoogle Scholar
  212. Linnaeus C (1758) Systema Naturae, 10th ed. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. HolmiaeGoogle Scholar
  213. Linnaeus C (1766) Systema naturae sive regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Laurentii Salvii. 12th ed. HolmiaeGoogle Scholar
  214. Longley WH (1935) Osteological notes and descriptions of new species of fishes. Carnegie Inst Wash Year Book 34: 86−89Google Scholar
  215. Lubbock R, Allen GR (1979) Canthigaster leopard a new sharpnose pufferfish (Teleostei: Tetraodontidae) from the central Indo-Pacifc. Rev Fr Aquariol 6:87−90 Google Scholar
  216. Manilo LG, Bogorodsky SV (2003) Taxonomic composition, diversity and distribution of coastal fishes of the Arabian Sea. J Ichthyol 43:S75−S149Google Scholar
  217. Marion de Procé PM (1822) Sur plusieurs espèces nouvelles de poissons et de crustacés observées. Bulletin de la Société Philomathique de Paris 1822:129−134Google Scholar
  218. Marshall NB (1952) The `Manihine’ expedition to the Gulf of Aqaba 1948–1949. IX. Fishes. Bull Brit Mus (Natr Hist) Zool 1:221–252Google Scholar
  219. Matsuura K (1979) Phylogeny of the superfamily Balistoidea (Pisces: Tetraodontiformes). Mem Fac Fish Hokkaido Univ 26:49−169Google Scholar
  220. Matsuura K (1980) A revision of Japanese balistoid fishes. I. Family Balistidae. Bull Natl Sci Mus (Ser A) 6:27–69Google Scholar
  221. Matsuura K (1981) Xenobalistes tumidipectoris, a new genus and species of triggerfish (Tetraodontiformes, Balistidae) from the Marianas Islands. Bull Natl Sci Mus (Ser A) 7:191−200Google Scholar
  222. Matsuura K (1982) A new triacanthodid fish, Triacanthodes indicus from the Indian Ocean. Jpn J Ichthyol 28:385−392Google Scholar
  223. Matsuura K (1983) Triacanthodidae. In: Uyeno TK, Matsuura K, Fujii E (eds) Fishes trawled off Suriname and French Guiana. Japan Marine Fishery Resource Research Center, Tokyo, p 464Google Scholar
  224. Matsuura K (1984) Tetraodontiformes. In: Masuda H, Amaoka K, Araga C, Uyeno T, Yoshino T (1984) The fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp 356−366, pls 321−334, 370Google Scholar
  225. Matsuura K (1985) Triacanthodidae to Tetraodontidae. In: Okamura O, Machida Y, Yamakawa T, Matsuura K, Yatou T (eds) Fishes of the Okinawa Trough and the adjacent waters. Vol. 2. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp 624−641, 742−750Google Scholar
  226. Matsuura K (1986) A new sharpnose pufferfish, Canthigaster flavoreticulata, collected from the South Pacific. Jpn J Ichthyol 33:223−224Google Scholar
  227. Matsuura K (1987) First record of a triacanthodid fish, Macrorhamphosodes uradoi from New Zealand. Jpn J Ichthyol 34:105−107Google Scholar
  228. Matsuura K (1990) The pufferfish genus Fugu Abe, 1952, a junior subjective synonym of Takifugu Abe, 1949. Bull Natl Sci Mus (Ser A) 16:15−20Google Scholar
  229. Matsuura, K (1992) A new sharpnose puffer, Canthigaster punctata (Teleostei: Tetraodontidae) from the Mascaren Submarine Ridge, western Indian Ocean. Bull Natl Sci Mus (Ser A) 18:127−130Google Scholar
  230. Matsuura K (1994) Arothron caeruleopunctatus, a new puffer from the Indo-West Pacific. Jpn J Ichthyol 41:29−33Google Scholar
  231. Matsuura K (1999) Taxonomic review of the puffers of the genus Arothron (Tetraodontiformes: Tetraodontidae) with a key to genera of the Indo-West Pacific puffers. Proc 9th Joint Sem Mar Fish Sci:125−140Google Scholar
  232. Matsuura, K (2000) Tetraodontiformes. In: Randall JE, Lim KKP (eds) A checklist of the fishes of the South China Sea. Raffles Bull Zool Suppl 8: 647−649Google Scholar
  233. Matsuura K (2001) Triacanthodidae, Triacanthidae, Balistidae, Ostraciidae, Aracanidae, Triodontidae, Tetraodontidae. In: Carpenter K, Niem (eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Vol 6. Bony fishes part 4 (Labridae to Latimeriidae). FAO, Rome, pp 3902−3928, 3948−3957Google Scholar
  234. Matsuura K (2002) A review of two morphologically similar puffers, Chelonodon laticeps and C. patoca. Natl Sci Mus Monogr 22:173−178Google Scholar
  235. Matsuura K (2003a) Balistidae, Monacanthidae, Ostraciidae, Tetraodontidae, and Diodontidae. In: Kimura S, Matsuura K (eds) Fishes of Bitung, northern tip of Sulawesi, Indonesia. Ocean Research Institute, University of Tokyo, Tokyo, pp 218−230Google Scholar
  236. Matsuura K (2003b) Balistidae, Monacanthidae. In: Carpenter KE (ed) The living marine resources of the Western Central Atlantic. Vol 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5. FAO, Rome, pp 1963−1979Google Scholar
  237. Matsuura K (2006) Redescription of the rare boxfish, Aracana spilonota, with comments on its taxonomic position and a record of another rare boxfish, Kentrocapros flavofasciatus, from southeastern Australia. Natl Sci Mus Monogr 34:273−277Google Scholar
  238. Matsuura K (2008) Family Ostraciidae and family Tetraodontidae. In: Gomon M, Bray D, Kuiter R (eds) Fishes of Australia’s southern coast. New Holland Publishers Pty Ltd, Sydney, pp 842−856Google Scholar
  239. Matsuura K (2009) Triacanthidae, Balistidae, Monacanthidae, Ostraciidae, Tetraodontidae, and Diodontidae. In: Kimura S, Satapoomin U, Matsuura K (eds) Fishes of Andaman Sea. National Museum of Nature and Science, Tokyo, pp 324−337Google Scholar
  240. Matsuura K (2010) Lagocephalus wheeleri Abe, Tabeta & Kitahama, 1984, a junior synonym of Tetrodon spadiceus Richardson, 1845 (Actinopterygii, Tetraodontiformes, Tetraodontidae). Mem Natl Mus Nat Sci 46:39−46Google Scholar
  241. Matsuura K (2011) Triacanthidae, Balistidae, Tetraodontidae, and Diodontidae. In: Matsunuma M, Motomura H, Matsuura K, Shazili NAMS, Ambak MA (eds) Fishes of Terengganu—east coast of Malay Peninsula, Malaysia. National Museum of Nature and Science, Tsukuba, Universiti Malaysia Terengganu, Kuala Terengganu, and Kagoshima University Museum, Kagoshima, pp 237−246Google Scholar
  242. Matsuura K (2013) Triacanthidae, Balistidae, Monacanthidae, Ostraciidae, Tetraodotnidae, and Diodontidae. In: Yoshida T, Motomura H, Mushikasinthorn P, Matsuura K (eds) Fishes of northern Gulf of Thailand. National Museum of Nature and Science, Tsukuba, Research Institute for Humanity and Nature, Kyoto, and Kagoshima University Museum, Kagoshima, pp 224−234Google Scholar
  243. Matsuura K (2014) A new pufferfish of the genus Torquigener that builds ‘‘mystery circles’’ on sandy bottoms in the Ryukyu Islands, Japan (Actinopterygii: Tetraodontiformes: Tetraodontidae) Ichthyol Res doi 10.1007/s10228-014-0428-5Google Scholar
  244. Matsuura K, Fourmanoir P (1984) A new triacanthodid fish, Triacanthodes intermedius, from New Caledonia. Bull Natl Sci Mus (Ser A) 10:31−35Google Scholar
  245. Matsuura K, Golani D, Bogorodsky SV (2011) The first record of Lagocephalus guentheri Miranda Ribeiro, 1915 from the Red Sea with notes on previous records of L. lunaris (Actinopterygii, Tetraodontiformes, Tetraodontidae). Bull Natl Mus Nat Sci (Ser. A) 37:163−169Google Scholar
  246. Matsuura K, Okuno J (1991) Redescription of a rare pufferfish, Arothron carduus (Cantor, 1849) (Teleostei: Tetraodontidae). Bull Natl Sci Mus (Ser A) 17:157−164Google Scholar
  247. Matsuura K, Shiobara Y (1989) A new triggerfish, Rhinecanthus abyssus, from the Ryukyu Islands. Jpn J Ichthyol 36:315−317Google Scholar
  248. Matsuura K, Peristiwady T (2000) Fishes Ikan. In: Matsuura K, Sumadhiharga OK, Tsukamoto K (eds) Field Guide to Lombok Island: Identification guide to marine organisms in seagrass beds of Lombok Island, Indonesia. Ocean Research Institute, University of Tokyo, Tokyo, pp 99−334Google Scholar
  249. Matsuura, K, Tyler JC (1997) Tetraodontiform fishes, mostly from deep waters, of New Caledonia. No. 9. In: B. Séret (ed) Résultats des Campagnes MUSORSTOM, 17. Mémoires du Muséum National d’Histoire Naturelle, Paris (N. S.) (Sér A) Zool 174:173−208Google Scholar
  250. Matsuura K, Yamakawa (1982) Rare boxfishes, Kentrocapros flavofasciatus and K. rosapinto, with notes on their relationships. Jpn J Ichthyol 29:31−42Google Scholar
  251. Matsuura K, Yoshino T (2004) A new triggerfish of the genus Abalistes (Tetraodontiformes: Balistidae) from the western Pacific. Rec Aust Mus 56:189−194Google Scholar
  252. McCulloch AR, Waite ER (1915) A revision of the genus Aracana and its allies. Trans Roy Soc S Aust 39:477−493, pls 16−25Google Scholar
  253. McEachran JD, Fechhelm JD (2005) Fishes of the Gulf of Mexico. Volume 2: Scorpaeniformes to Tetraodontiformes. University of Texas Press, Austin.Google Scholar
  254. Miranda Ribeiro A de (1913−15) Fauna brasiliense. Peixes. Tomo V. Museu Nacional, Rio de JaneiroGoogle Scholar
  255. Mitchill SL (1815) The fishes of New-York, described and arranged. Trans Lit Phil Soc NY 1:355−492, pls 1−6Google Scholar
  256. Mitchill SL (1818) Memoir on ichthyology. The fishes of New York, described and arranged. In a supplement to the Memoir on the same subject, printed in the New-York Literary and Philosophical Transactions. Amer Month Mag Crit Rev 2:321−328Google Scholar
  257. Miya M, Takeshima H, Endo H, Ishiguro NB, Inoue JG, Mukai T, Satoh TP, Yamaguchi M, Kawaguchi A, Mabuchi K, Shirai SM, Nishida M (2003) Major patterns of higher teleostean phylogenies: a new perspective based on 100 complete mitochondrial DNA sequences. Mol Phylogenet Evol 26:121–138Google Scholar
  258. Miyajima Y, Ohe F, Koike H, Matsuoka H (2014) First occurrence of a filefish (Tetraodontiformes; Monacanthidae) in Asia, from the Middle Miocene in Nagano Prefecture, central Japan. Zootaxa 3786:382–400Google Scholar
  259. Mohsin AKM, Ambak MA (1996) Marine fishes and fisheries of Malaysia and neighbouring countries. Universiti Pertanian Malaysia Press, SerdangGoogle Scholar
  260. Moore D (1967a) Triggerfishes (Balistidae) of the western Atlantic. Bull Mar Sci 17:689−722Google Scholar
  261. Moore D (1967b) Nomenclature of the spotted triggerfish, Balistes punctatus, of the eastern Atlantic. Copeia 1967:858−861Google Scholar
  262. Moura RL, Castro MC (2002) Revision of Atlantic sharpnose pufferfishes (Tetraodontiformes: Tetraodontidae: Canthigaster), with description of three new species. Proc Biol Soc Wash 115:32−50Google Scholar
  263. Müller J (1841) Vergleichende Anatomie der Myxinoiden. Dritte Fortsetzung. Über das Gefässystem. Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin 1839:175−304, pls 1−5Google Scholar
  264. Müller J (1843) Beiträge zur Kenntniss der natürlichen Familien der Fische. Archiv für Naturgeschichte 9:292–330Google Scholar
  265. Müller J, Troschel FH (1848) Fische. In: Schomburgk R (ed) Reisen in Britisch-Guiana in den Jahren 1840−44, im Auftrag Sr. Mäjestat des Königs von Preussen ausgeführt von Richard Schomburgk. Vol 3. J.J. Waber, Leipzig, pp. 618−644Google Scholar
  266. Munday BC (2005) Checklist of the fishes of the Hawaiian Archipelago. Bishop Museum Press, HonoluluGoogle Scholar
  267. Murray A (1857) Notice of a Tetraodon (believed to be new) from Old Calibar. Proc Royal Physical Soc Edinburgh 1:252–253Google Scholar
  268. Myers GS (1934) Three new deep-water fishes from the West Indies. Smithson Misc Coll 91:1−12, pl 1Google Scholar
  269. Myers RF (1999) Micronesian reef fishes. A comprehensive guide to the coral reef fishes of Micronesia. 3rd revised ed. Coral Graphics, GuamGoogle Scholar
  270. Myers RF, Donaldson TJ (2003) The fishes of the Mariana Islands. Micronesica 35/36:598−652Google Scholar
  271. Nakae M, Sasaki K (2010) Lateral line system and its innervations in Tetraodontiformes with outgroup comparisons: descriptions and phylogenetic implications. J Morphol 271:559−579Google Scholar
  272. Nelson JS (2006) Fishes of the world, fourth edition. John Wiley & Sons, Inc., HobokenGoogle Scholar
  273. Ni Y, Li CS (1992) Takifugu coronoidus, nov. sp. (Tetraodontiformes) from Chinese waters. Oceanol Limnol Sinica 23:527−532Google Scholar
  274. Ogilby JD (1899) Additions to the fauna of Lord Howe Island. Proc Linn Soc NSW 23:730–745Google Scholar
  275. Ogilby JD (1913) On six new or rare Queensland fishes. Mem Qld Mus 2:81–89Google Scholar
  276. Okamura O, Machida Y, Yamakawa T, Matsuura K, Yatou T (1985) Fishes of the Okinawa Trough and the adjacent waters. Vol. 2. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, TokyoGoogle Scholar
  277. Oken L (1817) V. Kl. Fische. Isis (Oken) 8:1779−1782Google Scholar
  278. Orr JW, Fritzsche RA (1997) Solenostomus tuticoriensis Venkataramanujam, Venkataramani, and Devaraj, 1993, a junior synonym of Macrorhamphosodes platycheilus Fowler, 1934 (Tetraodontiformes: Triacanthodidae). Copeia 1997:888−889Google Scholar
  279. Osbeck P (1765) Reise nach Ostindien und China. Nebst O. Toreens Reise nach Suratte und C. G. Ekebergs Nachricht von den Landwirthschaft der Chineser. J. C. Koppe, RostockGoogle Scholar
  280. Park M (1797) Descriptions of eight new fishes from Sumatra. Trans Linn Soc London 3:33−38, pl 6Google Scholar
  281. Parenti P (2003) Family Molidae Bonaparte 1832—molas or ocean sunfishes. Calif Acad Sci Annotated Checklists of Fishes 18:1−9Google Scholar
  282. Pennant T (1776) British zoology. 4th edition. Vol. 3: Class III. Reptiles. Class IV. Fish. Benjamin White, London.Google Scholar
  283. Peters W (1855) Übersicht der in Mossambique beobachteten Seefische. Monatsberichte der Königlichen Preuss [ischen] Akademie der Wissenschaften zu Berlin 1855:428–466Google Scholar
  284. Phillipps WJ (1932) Notes on new fishes from New Zealand. N Z J Sci Technol 13:26−234Google Scholar
  285. Playfair RL, Günther A (1867) The fishes of Zanzibar, with a list of the fishes of the whole east coast of Africa. Voorst, LondonGoogle Scholar
  286. Poey F (1861) Memorias sobra la historia natural de la Isla de Cuba, acompañadas de sumarios Latinos y extractos en Francés. Tomo 2. La Habana 2:337−442, pls 13, 15−19Google Scholar
  287. Poey F (1876) Enumeratio piscium cubensium (Parte Segunda). Anales de la Sociedad Española de Historia Natural Madrid 5:131–218, pls 4–9Google Scholar
  288. Poll M (1959) Résultats scientifiques des missions zoologiques au Stanley Pool subsidiées par le Cemubac (Université Libre de Bruxelles) et la Musée Royal du Congo (1957-1958). III. Récherches sur la faune ichthyologique de la région du Stanley Pool 71:75–174, pls 12–26Google Scholar
  289. Popta, CML (1900) On a small Monacanthus. Notes from the Leyden Museum 22:126−128Google Scholar
  290. Quoy, JRC, Gaimard JP (1824−25) Description des Poissons. Chapter IX. In: Freycinet LCD de (ed) Voyage autour du monde, entrepris par ordre du Roi sous le ministére et conformément aux instructions de S. Exc. M. le Vicomte du Bouchage, Secrétaire d’État au Département de la Marine, exécuté sur les corvettes de S.M. l’Uranie et la Physicienne, pendant les années 1817, 1818, 1819 et 1820; publié sous les auspices de S.E.M. le Comte Corbiére, Secrétaire d’État de l’Intérieur, pouv la partie historique et kes sciences naturelles, et de S.E.M. le Marquis de Clemont-Tonnerre, Secrétaire d’État de la Marine et des Colonies, pouv la partie nautique. “1824” Pillet, ParisGoogle Scholar
  291. Ramsay EP, Ogilby JD (1886) Descriptions of some new Australian fishes. Proc Linn Soc NSW (Ser 2) 1:4−7Google Scholar
  292. Randall JE (1964) A revision of the filefish genera Amanses and Cantherhines. Copeia 1964:331−361Google Scholar
  293. Randall JE (1975) Ostracion trachys, a new species of trunkfish from Mauritius (Ostraciontidae). Matsya 1:59–62Google Scholar
  294. Randall JE (1995) Coastal fishes of Oman. Crowford House Publishing Pty Ltd, BathurstGoogle Scholar
  295. Randall JE (1996) Caribbean reef fishes, 3rd edition. TFH Publications Inc.Neptune CityGoogle Scholar
  296. Randall JE (2005) Reef and shore fishes of the South Pacific. University of Hawai’i Press, HonoluluGoogle Scholar
  297. Randall JE (2010) Shore fishes of Hawai’i. University of Hawai’i Press, HonoluluGoogle Scholar
  298. Randall JE (2011) Review of the circumtropical monacanthid fish genus Cantherhines, with descriptions of two new species. Indo-Pacific Fishes (40):1−30, pls 1−7Google Scholar
  299. Randall JE, Cea-Egaña A (1989) Canthigaster cyanetron a new toby (Teleostei: Tetraodontidae) from Easter Island. Rev Fr Aquariol 15:93−96Google Scholar
  300. Randall, JE, Klausewitz, W (1973) A review of the triggerfish genus Melichthys, with description of a new species from the Indian Ocean. Senckenbergiana Biologica 54:57−69Google Scholar
  301. Randall JE, Steene RC (1983) Rhinecanthus lunula a new species of triggerfish from the South Pacific. Freshw Mar Aquar 6:45−51Google Scholar
  302. Randall JE, Matsuura K, Zama A (1978) A revision of the triggerfish genus Xanthichthys, with description of a new species. Bull Mar Sci 28:688–706Google Scholar
  303. Randall JE, Williams JT, Rocha LA (2008) The Indo-Pacific tetraodontid fish Canthigaster coronata, a complex of three species. Smithiana, Publ Aquatic Biodiv, Bull 9:3−13Google Scholar
  304. Randall JE, Allen GR, Steene RC (1997) Fishes of the Great Barrier Reef and Coral Sea. 2nd, revised ed. Crawford House Press, BathurstGoogle Scholar
  305. Randall JE, Williams JT, Smith DG, Kulbicki M, Mou Tham G, Labrosse P, Kronen M, Clua E, Mann BS (2004) Checklist of the shore and epipelagic fishes of Tonga. Atoll Res Bull (502):i−ii + 1−35Google Scholar
  306. Ranzani C (1842) De novis speciebus piscium. Dissertatio Secunda. Novi Commentarii Academiae Scientiarum Instituti Bononiensis 5:3−21, pls 1−7Google Scholar
  307. Regan CT (1903) On the classification of the fishes of the suborder Plectognathi; with notes and descriptions of new species from specimens in the British Museum Collection. Proc Zool Soc London 1902, 2:284–303, pls 24–25Google Scholar
  308. Regan CT (1908) Report on the marine fishes collected by Mr. J. Stanley Gardiner in the Indian Ocean. Trans Linn Soc London Sec Ser Zool 12:217−255, pls 23−32Google Scholar
  309. Regan CT (1913) A collection of fishes made by Professor Francisco Fuentes at Easter Island. Proc Zool Soc London 1913:368–374, pls 55–60Google Scholar
  310. Regan CT (1919) Fishes from Durban, Natal, collected by Messrs. H. W. Bell Marley and Romer Robinson. Ann Durban Mus 2:197−204Google Scholar
  311. Richardson J (1841) On some new or little known fishes from the Australian seas. Proc Zool Soc London 1841:21−22Google Scholar
  312. Richardson J (1844−48) Ichthyology of the voyage of H.M.S. Erebus & Terror, under the command of Sir James Clark Ross R.N., F.R.S. In: Richardson J and Gray JE (eds) The zoology of the voyage of H.M.S. Erebus & Terror, under the command of Captain Sir James Clark Ross R.N., F.R.S. during the years 1839 to 43. Longman & Brown, London, pp i–viii + 1–139, pls 1–60Google Scholar
  313. Richardson J (1845) Ichthyology.—Part 3. In: Hinds RB (ed) The zoology of the voyage of H.M.S. Sulphur, under the command of Captain Sir Edward Belcher, R.N., C.B., F.R.G.S., etc., during the years 1836−42, No. 10. Smith, Elder & Co, London, pp 71–98, pls 55–64Google Scholar
  314. Richardson J (1854) Vertebrals, including fossil mammals. Fish. In: Forbes E (ed) The zoology of the voyage of H.M.S. Herald, under the command of Captain Henry Kellett, R.N., C.B., during the years 1845–51. Lovell Reeve, London, pp 156–171, pls 30–33Google Scholar
  315. Richardson J (1856) Ichthyology. In: Brewster D (ed) The Encyclopedia Britannica, or Dictionary of Arts, Sciences, and General Literature. 8th Edition. Vol. 12. Encyclopedia Britannica, Edinburgh, pp 204−331Google Scholar
  316. Roberts TR (1982) The southeast Asian freshwater pufferfish genus Chonerhinos (Tetraodontidae), with descriptions of new species. Proc Calif Acad Sci (Ser 4) 43:1−16Google Scholar
  317. Roberts TR (1998) Freshwater fugu or pufferfishes of the genus Tetraodon from the Mekong basin, with descriptions of two new species. Ichthyol Res 45:225−234Google Scholar
  318. Rosen DE (1984) Zeiforms as primitive plectognath fishes. Am Mus Nov 2782:1−45Google Scholar
  319. Rüppell WPES (1828−30) Atlas zu der Reise im nördlichen Afrika. Fische des Rothen Meers. Heinrich Ludwig Brönner, Frankfurt am MainGoogle Scholar
  320. Rüppell WPES (1835−38) Neue Wirbelthiere zu der Fauna von Abyssinien gehörig. Fische des Rothen Meeres. Siegmund Schmerber, Frankfurt am MainGoogle Scholar
  321. Saenjudaeng P, Vidthayanon C, Grudpun C (2013) Tetraodon palustris, a new freshwater pufferfish (Tetraodontiformes: Tetraodontidae) from the Mekong Basin of Thailand. Zootaxa 3686:77−84Google Scholar
  322. Sainsbury KJ, Kailola PJ, Leyland GG (1984) Continental shelf fishes of northern and north-western Australia. Clouston & Hall and Peter Pownall Fisheries Information Service, CanberraGoogle Scholar
  323. Santini F (2006) A new species of Triacanthodidae (Tetraodontiformes, Acanthomorpha) from the central Pacific. Cybium 30:195−198Google Scholar
  324. Santini F, Tyler JC (2002a) Phylogeny of the ocean sunfishes (Molidae, Tetraodontiformes), a highly derived group of teleost fishes. Italian J Zool 69:37–43Google Scholar
  325. Santini F, Tyler JC (2002b) Phylogeny and biogeography of the extant species of triplespine fishes (Triacanthidae, Tetraodontiformes). Zool Scripta 31:321–330Google Scholar
  326. Santini F, Tyler JC (2003) A phylogeny of the families of fossil and extant tetraodontiform fishes (Acanthomorpha, Tetraodontiformes), upper Cretaceous to Recent. Zool J Linn Soc 139:565−617Google Scholar
  327. Santini F, Soresen L, Marcroft T, Dornburg A, AlfaroME (2013a) A multilocus molecular phylogeny of boxfishes (Aracanidae, Ostraciidae; Tetraodontiformes). Mol Phylogenet Evol 66:153−160Google Scholar
  328. Santini F, Sorenson L, Alfaro ME (2013b) A new multi-locus timescale reveals the evolutionary basis of diversity patterns in triggerfishes and filefishes (Balistidae, Monacanthidae; Tetraodontiformes). Mol Phylogenet Evol 69:165−176Google Scholar
  329. Santini F, Sorenson L, Alfaro ME (2013c) A new phylogeny of tetraodontiform fishes (Tetraodontiformes, Acanthomorpha) based on 22 loci. Mol Phylogenet Evol 69:177−187Google Scholar
  330. Schomburgk RH (1848) The history of Barbados; comprising a geographical description of the island and an account of its geology and natural productions. Longman, Brown, Green and Longmans, LondonGoogle Scholar
  331. Schultz LP, Woods LP, Lachner EA (1966) Fishes of the Marshall and Marianas islands. Vol. 3. Families Kraemeriidae through Antennariidae. Bull US Natl Mus 202, 3:i−vii + 1−176, pls 124−148Google Scholar
  332. Shao, KT, Ho HC, Lin PL, Lee PF, Lee MY, Tsai CY, Liao YC, Lin YC (2008) A checklist of the fishes of southern Taiwan, northern South China Sea. Raffles Bull Zool Suppl 19:233−271 Google Scholar
  333. Shaw G (1804) General zoology or systematic natural history. Vol. 5. George Kearsley, LondonGoogle Scholar
  334. Shaw G, Nodder FP (1789–1813) The Naturalist’s Miscellany, or coloured figures of natural objects; drawn and described from nature. Nodder & C, LondonGoogle Scholar
  335. Shipp, RL (1974) The pufferfishes (Tetraodontidae) of the Atlantic Ocean. Publ Gulf Coast Res Lab Mus 4:1−164Google Scholar
  336. Smith JLB (1948) Brief revisions and new records of South African marine fishes. Ann Mag Natr Hist (Ser 11) 14:335−346Google Scholar
  337. Smith JLB (1949a) A new aracanid fish from South Africa. Annals Mag Natr Hist (Ser 12) 2:354−359Google Scholar
  338. Smith JLB (1949b) The sea fishes of southern Africa. Central News Agency, Ltd., Cape TownGoogle Scholar
  339. Smith JLB (1953) Fishes taken in the Moçambique Channel by Mussolini P. Fajardo. Memorias do Museu Dr. Alvaro de Castro 2:3–20, pl 1Google Scholar
  340. Smith JLB (1958) Tetraodont fishes from from South and East Africa. Ann Mag Natr Hist (Ser 13) 1:156−160, pl 2Google Scholar
  341. Smith MM (1986) Family No. 267: Tetraodontidae. In: Smith MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan South Africa, Johannesburg, pp 894−903, pls 139−140, 142−145Google Scholar
  342. Smith MM, Heemstra PC (1986) Family No. 263: Balistidae. In: Smith MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan South Africa, Johannesburg, pp 876−882, pls 136−139Google Scholar
  343. Snyder JO (1911) Descriptions of new genera and species of fishes from Japan and the Riu Kiu Islands. Proc US Natl Mus 40:525–549Google Scholar
  344. Sontirat S (1985) Three new species of the freshwater fishes from Thailand. Thai Fish Gazette 38:41–49Google Scholar
  345. Sontirat S, Soonthornsatit S (1985) A new puffer species of Thailand: Tetraodon suvattii n. sp. Proc 23rd Conf Fish Sec Kasetsart Univ Bangkok 49−53Google Scholar
  346. Sorbini C, Tyler JC (2004) Review of the fossil file fishes of the family Monacanthhidae (Tetraodontiformes), Pliocene and Pleistocene of Europe, with a new genus, Frigocanthus, and two new species related to the Recent Aluterus. Bollettino del Museo Civico di Storia Naturale di Verona 26:41–76Google Scholar
  347. Steindachner F (1866) Ichthyologische Mittheilungen. (VIII) Verhandlungen der K.-K. zoologisch-botanischen Gesellschaft in Wien 16:75–484, pls 5–6Google Scholar
  348. Steindachner F (1870) Ichthyologische Notizen (X). (Schluss). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe 61:623–642, pls 1–5Google Scholar
  349. Steindachner F (1876) Ichthyologische Beiträge (V). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe 74:49−240, pls 1−15Google Scholar
  350. Stewart AL, Clark MR (1988) Records of three families and four species of fish new to New Zealand fauna. NZ J Zool 15:577−583Google Scholar
  351. Su JX, Li CS (2002) Fauna Sinica. Osteichthyes. Tetraodontiformes, Pegasiformes, Gobiesociformes, Lophioformes. Science Press, BeijingGoogle Scholar
  352. Su JX, Hardy GS, Tyler JC (1986) A new generic name for Anchisomus multistriatus Richardson 1854 (Tetraodontidae), with notes on its toxicity and pufferfish biting behavior. Rec West Aust Mus 13:101−120Google Scholar
  353. Swainson W (1839) On the natural history and classification of fishes, amphibians, & reptiles, or monocardian animals. Spottiswoode & Co., LondonGoogle Scholar
  354. Talwar PK, Jhingran AG (1991) Inland fishes of India and adjacent countries (2 vols.) Oxford & IBH Publishing Co., New DelhiGoogle Scholar
  355. Tan HH (1999) A new species of Carinotetraodon from Sumatra and Borneo and validity of C. borneensis (Teleostei: Tetraodontidae). Ichthyol Exploration Freshw10:345−354Google Scholar
  356. Tanaka S (1918) Figures and descriptions of the fishes of Japan including Riukiu Islands, Bonin Islands, Formosa, Kurile Islands, Korea and southern Sakhalin (27):475−494, pls 131−135Google Scholar
  357. Temminck CJ, Schlegel H (1850) Pisces, Fauna Japonica. Parts 7–9:113–172, pls 1–143Google Scholar
  358. Tilesius WG von (1820) De piscium Australium novo genere icone illustrato. Mém Acad Imp Sci St Petersbourg 7:301−310, second pl 9Google Scholar
  359. Tirant G (1885) Notes sur les poissons de la Basse-Cochinchine et du Cambodge. Excursions et reconnaissances. 10:91−198Google Scholar
  360. Tortonese E (1986) Balistidae, Monacanthidae, Ostraciontidae, Tetraodontidae, Diodontidae, and Molidae. In: Whitehead, PJP, Bauchot ML, Hureau JC, Nielsen J, Tortonese (eds) Fishes of the north-eastern Atlantic and the Mediterranean. Vol 3. UESCO, Paris, pp 1335−1348Google Scholar
  361. Troschel FH (1856) Bericht über die Leistungen in der Ichthyologie wärend des Jahres 1855. Archiv für Naturgeschichte 22:67−89Google Scholar
  362. Tyler JC (1963) A critique of Y. Le Danois’ work on the classification of the fishes of the oprder Plectognathi. Copeia 1963:203206Google Scholar
  363. Tyler JC (1964) A diagnosis of the two species of South American puffer fish (Tetraodontidfae, Plectognathi) of the genus Colomesus. Proc Acad Sci Philad 116:119−148Google Scholar
  364. Tyler JC (1965) A synopsis of the four species of cowfishes (Acanthostracion, Plectognathi) in the Atlantic Ocean. Proce Acad Nat Sci Philad 117:8261–287Google Scholar
  365. Tyler JC (1966a) A new species of triacanthodid Plectognath fish from the Caribbean, Hollardia meadi. Not Nat (Philad) 382:1−8Google Scholar
  366. Tyler JC (1966b) A new genus and species of triacanthodid fish (Plectognathi) from the Indian Ocean. Not Nat (Philad) 385:1−5Google Scholar
  367. Tyler JC (1966c) Bathyphylax omen, a new species of triacanthodid plectognath fish from the Indian Ocean. Not Nat (Philad) 395:1−5Google Scholar
  368. Tyler JC (1967) A redescription of Triodon macropterus Lesson, a phyletically important plectognath fish. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen. Biol Med Sci (Ser C) 70:84–96Google Scholar
  369. Tyler JC (1968) A monograph on plectognath fishes of the superfamily Triacanthoidea. Monogr Acad Natl Sci Philad 16:1−364Google Scholar
  370. Tyler JC (1970) An especially small, sexually dimorphic new species of filefish (Monacanthidae) from Australasian reefs. Proc Acad Natl Sci Philad 122:273−290Google Scholar
  371. Tyler JC (1978) Der Rotaugen-Kammkugelfisch, Carinotetraodon lorteti, aus Südostasien. Die Aquarien-und Terrarien Zeitschrift 31:118–121Google Scholar
  372. Tyler JC (1980) Osteology, phylogeny, and higher classification of the fishes of the order Plectognathi (Tetraodontiformes). NOAA Tech Rep NMFS Circular 434:1–422Google Scholar
  373. Tyler JC (1983) Records of fishes of the family Triacanthodidae (Tetraodontiformes) from the western Indian Ocean off east Africa. JLB Smith Inst Ichthyol Spec Publ 31:1−13Google Scholar
  374. Tyler JC (1986) Family No. 265: Triacanthodidae. In: Smith MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan South Africa, Johannesburg, pp 887−890Google Scholar
  375. Tyler JC (1997) New species of Paratriacanthodes spikefish (Triacanthodidae: Tetraodontiformes) from the South China Sea. Proc Biol Soc Wash 110: 310−313Google Scholar
  376. Tyler JC, Bannikov AF (1992) New genus of primitive ocean sunfish with separate premaxillae from the Eocene of southwest Russia (Molidae, Tetraodontiformes). Copeia 1992:1014−1023Google Scholar
  377. Tyler JC, Bannikov AF (1994) A new genus of fossil pufferfish (Tetraodontidae: Tetraodontiformes) based on a new species from the Oligocene of Russia and a referred species from the Miocene of Ukraine. Proc Biol soc Wash 107:97−108Google Scholar
  378. Tyler JC, Bannikov AF (2011) New specimen of rare fossil triacanthid fish genus Protacanthodes from the Eocene of Monte Bolca, Italy (Triacanthidae, Tetraodontiformes). Studi e Ricerche sui Giacimenti Terziari di Bolca, Museo Civico di Storia Naturale di Verona 13:29–35Google Scholar
  379. Tyler JC, Bannikov AF (2012) A new species of puffer fish, Eotetraodon tavernei, from the Eocene of Monte Bolca, Italy (Tetraodontidae, Tetraodontiformes) Studi e Ricerche sui Giacimenti Terziari di Bolca, Museo Civico di Storia Naturale di Verona 14:51–58Google Scholar
  380. Tyler JC, Kriznar M (2013) A new genus and species, Slovenitriacanthus saksidai, from southwestern Slovenia, of the Upper Cretaceous basal tetraodontiform family Cretatriacanthidae (Plectocretacicoidea). Bollettino del Museo Civico di Storia Naturale di Verona 37:45–56Google Scholar
  381. Tyler JC, Matsuura K (1981) Comments on the osteology of balistoid fishes (Tetraodontiformes), with notes on the triodontid pelvis. Proc Biol Soc Wash 94:52−66Google Scholar
  382. Tyler JC, Patterson C (1991) The skull of the Eocene Triodon antiquus (Triodontidae; Tetraodontiformes): similar to that of the Recent threetooth pufferfish T. macropterus. Proc Biol Soc Wash 104:878−891Google Scholar
  383. Tyler JC, Paxton JR (1979) New genus and species of pufferfish (Tetraodontidae) from Norfolk Island, southwest Pacific. Bull Mar Sci 29:202−215Google Scholar
  384. Tyler JC, Santini F (2001) A new species of triacanthid fish of the genus Protacanthodes from the Eocene of Monte Bolca, Italy (Tetraodontiformes). Bollettino del Museo Civico di Storia Naturale di Verona 25:3−9Google Scholar
  385. Tyler JC, Santini F (2002) Review and reconstructions of the tetraodontiform fishes from the Eocene of Monte Bolca, Italy, with comments on related Tertiary taxa. Museo Civico di Stria Naturale di Verona, Studie e Ricerche sui Giacimenti Terziari di Bolca 9:47−119Google Scholar
  386. Tyler JC, Sorbini L (1996) New superfamily and three new families of tetraodontiform fishes from the Upper Cretaceous: the earlies and most morphologically primitive plectognaths. Smithson Contr Paleobiol 82:1−259Google Scholar
  387. Tyler JC, Sorbini L (1998) A new genus and species of primitive triggerfishes from the Eocene of Monte Bolca, Italy; the earliest known balistoid (Tetraodontiformes). Museo Civico di Stria Naturale di Verona, Studie e Ricerche sui Giacimenti Terziari di Bolca (Misc Plaeontol) 7:43−265Google Scholar
  388. Tyler JC, Winterbottom R (1999) A review of the morphology and relationships of the Oligocene spikefish genera Acanthopleurus Agassiz 1844 and Cryptobalistes Tyler 1968 (Tetraodontiformes: Triacanthidae). Paleontologische Zeitschrift 73:351−367Google Scholar
  389. Tyler JC, Purdy RW, Oliver KH (1992) A new species of Sphoeorides pufferfish (Teleostei: Tetraodontidae) with extensive hyperostosis from the Pliocene of North Carolina. Proc Biol Soc Wash 105:462−482Google Scholar
  390. Tyler JC, Jerzmanska A, Bannikov AF, Swidnicki J (1993) Two new genera and species of Oligocene spikefishes (Tetraodontiformes: Tiracanthodidae), the first fossils of the Hollardinae and Tiracanthodinae. Smithoson Contr Paleobiol 75:1−27Google Scholar
  391. Tyler JC, Bronzi P, Ghiandoni A (2000) The Cretaceous fishes of Nardoi 11°. A new genus and species of Zeiformes, Cretazeus rinaldii, the earliest record for the order. Bollettino del Museo Civico di Storia Naturale di Verona 24:11−28Google Scholar
  392. Tyler JC, O’Toole B, Winterbottom R (2003) Phylogeny of the genera and families of zeiform fishes, with comments on their relationships with tetraodontiforms and caproids. Smithson Contr Zool 618:1−110Google Scholar
  393. Tyler JC, Mirzaie M, Nazemi A (2006) New genus and species of basal tetraodontoid puffer fish from the Oligocene of Iran, related to the Zignoichthyidae (Tetraodontiformes). Bollettino del Museo Civico di Storia Naturale di Verona 30:49–58Google Scholar
  394. Tyler JC, Collette BB, Broughton EA (2013) Northern range extension to Georges Bank for Hollardia hollardi (Reticulate Spikefish) (Triacanthodidae, Tetraodontiformes). Northeastern Naturalist 20:33–36Google Scholar
  395. Vaillant LL, Sauvage HE (1875). Note sur quelques espèces nouvelles de poissons des îles Sandwich. Rev Mag Zool (Ser 3) 3: 278−287Google Scholar
  396. Venkataramanujam K, Venkataramani VK, and Deveraj M (1993) A new solenostomid fish Solenostomus tuticoriensis sp. nov. from Tuticorin Bay, South India. J Mar Biol Assoc India 35:201−204Google Scholar
  397. Waite ER (1904) Additions to the fish fauna of Lord Howe Island, No. 4. Rec Aust Mus 5:135–186, pls 17–24Google Scholar
  398. Waite ER (1905) Notes on fishes from Western Australia.—No. 3. Rec Aust Mus 6:55−82, pls 8−17Google Scholar
  399. Walbaum JJ (1792) Petri Artedi sueci genera piscium. In quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Redactis speciebus 242 ad genera 52. Ichthyologiae pars III. Ant. Ferdin. Rose, GrypeswaldiaeGoogle Scholar
  400. Walker H. J. Jr., Bussing WA (1996) Two new pufferfishes of the genus Sphoeroides from the Eastern Pacific. Copeia 1996:677−684Google Scholar
  401. Weber M (1913) Die Fische der Siboga−Expedition. E. J. Brill, LeidenGoogle Scholar
  402. White J (1790) Journal of a voyage to New South Wales with sixty-five plates of non descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. J. D. Piccadilly, LondonGoogle Scholar
  403. Whitley GP (1929) Studies in ichthyology. No. 3. Rec Aust Mus17:101–143, pls 30–34Google Scholar
  404. Whitley GP (1930a) Additions to the checklist of the fishes of New South Wales, no. 3. Aust Zool 6:117−124, pl 14Google Scholar
  405. Whitley GP (1930b) Leatherjacket genera. Aust Zool 6:179Google Scholar
  406. Whitley GP (1930c) Ichthyological miscellanea. Mem Queensland Mus 10:8–31, pl 1Google Scholar
  407. Whitley GP (1931) New names for Australian fishes. Aust Zool 6:310−334, pls 25−27Google Scholar
  408. Whitley GP (1934) Studies in ichthyology. No 8. Rec Aust Mus 19:153–163Google Scholar
  409. Whitley GP (1939) Studies in ichthyology. No 12. Rec Aust Mus 20:264–277Google Scholar
  410. Whitley GP (1947) New sharks and fishes from Western Australia. Part 3. Aust Zool 11:129−150, pl 11Google Scholar
  411. Whitley GP (1965) Illustrations and records of fishes. Aust Zool 13:103−120 Google Scholar
  412. Wiley EO, Johnson GD (2010) A teleost classification based on monophyletic groups. In: Nelson JS, Schultze H, Wilson VH (eds) Origin and phylogenetic interrelationships of teleosts. Verlag Dr. Friedrich Pfeil, München, pp 123−182Google Scholar
  413. Williams JT, Delrieu-Trottin E, Planes S (2012) A new species of Indo-Pacific fish, Canthigaster criobe, with comments on other Canthigaster (Tetraodontiformes: Tetraodontidae) at the Gambier Archipelago. Zootaxa 3523:80−88Google Scholar
  414. Winterbottom R (1974) The familial phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as evidenced by their comparative myology. Smithson Contr Zool 155:1−201Google Scholar
  415. Winterbottom R. Tyler JC (1983) Phylogenetic relationships of aracanin genera of boxfishes (Ostraciidae: Tetraodontiformes). Copeia 1983:902−917Google Scholar
  416. Winterbottom, R, Emery AR, Holm E (1989) An annotated checklist of the fishes of the Chagos Archipelago, central Indian Ocean. Roy Ontario Mus Life Sci Contr 145:1−226, pls 1−8Google Scholar
  417. Woods LP (1959) Parahollardia schmidti, a new triacanthodid fish from the western Caribbean. Copeia 1959:222−225Google Scholar
  418. Yamanoue Y, Miya M, Matsuura K, Katoh M, Sakai H, Nishida M (2004) Mitochondrial genomes and phylogeny of the ocean sunfishes (Tetraodontiformes: Molidae). Ichthyol Res 51:269−273Google Scholar
  419. Yamanoue Y, Miya M, Matsuura K, Yagishita N, Mabuchi K, Sakai H, Katoh M, Nishida M (2007) Phylogenetic position of tetraodontiform fishes within the higher teleosts: Baysian inferences based on 44 whole mitochondrial genome sequence. Mol Phylogenet Evol 45:89−101Google Scholar
  420. Yamanoue Y, Miya M, Matsuura K, Katoh M, Sakai H, Nishida M (2008) A new perspective on phylogeny and evolution of tetraodontiform fishes (Pisces: Acanthopterygii) based on whole mitochondrial genome sequences: basal ecological diversification? BMC Evol Biol:212 doi 10.1186/471-2148-8-212Google Scholar
  421. Yamanoue Y, Miya M, Matsuura K, Sakai H, Katoh M, Nishida M (2009) Unique patterns of pelvic fin evolution: a case study of balistoid fishes (Pisces: Tetraodontiformes) based on whole mitochondrial genome sequences. Mol Phylogenet Evol 50:179−189Google Scholar
  422. Yamanoue Y, Miya M, Doi H, Mabuchi K, Sakai H, Nishida M (2011) Multiple invasion into freshwater by a pufferfishes (Teleostei: Tetraodontidae): A mitogenomic perspective. PLos ONE 6:e17410 doi 10.1371/journal.pone.0017410Google Scholar
  423. Yoshita Y, Yamanoue Y, Sagara K, Nishibori M, Kuniyoshi H, Umino T, Sakai Y, Hashimoto H, Gushima K (2009) Phylogenetic relationships of two Mola sunfishes (Tetraodontiformes: Molidae) occurring around the coasts of Japan, with notes on their geographical distribution and morphological characteristics. Ichthyol Res 56:232−244Google Scholar

Copyright information

© The Author(s) 2014

Open AccessThis article is distributed under the terms of the Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are credited.

Authors and Affiliations

  1. 1.Division of FishesNational Museum of Nature and ScienceTsukubaJapan

Personalised recommendations