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Taxonomy and Systematics of Tetraodontiform Fishes

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Taxonomy and Systematics of Tetraodontiform Fishes Ichthyol Res DOI 10.1007/s10228-014-0444-5 REVIEW FOR IPFC9 SPECIAL ISSUE Ichthyology and Indo-Pacific Fish Conferences from the 1980s to the 2010s Taxonomy and systematics of tetraodontiform fishes: a review focusing primarily on progress in the period from 1980 to 2014 Keiichi Matsuura Received: 2 October 2014 / Revised: 12 October 2014 / Accepted: 13 October 2014 Ó The Author(s) 2014. This article is published with open access at Springerlink.com Abstract When the first Indo-Pacific Fish Conference species in four genera, Balistidae 37 species in 12 genera, (IPFC1) was held in Sydney in 1981, there were still many Monacanthidae 102 species in 27 genera, Aracanidae 13 problems in the generic- and species-level taxonomy of all species in six genera, Ostraciidae 22 species in five genera, tetraodontiform families except for the recently reviewed Triodontidae monotypic, Tetraodontidae 184 species in 27 Triacanthodidae and Triacanthidae. The period from IPFC1 genera, Diodontidae 18 species in seven genera, and Mo- to IPFC9 (1981-2013) was a time of great progress in the lidae five species in three genera. Phylogenetic relation- taxonomy and systematics of the Tetraodontiformes: many ships of the families have been clarified by morphological review and revisional papers have been published for and molecular analyses and have provided well-supported various genera and species, with descriptions of many new sister relationships of the families: Triacanthodidae and taxa occurring mainly on coral reefs and in tropical Triacanthidae, Balistidae and Monacanthidae, and Tetra- freshwaters; and cladistic analyses of morphological char- odontidae and Diodontidae. However, there remain prob- acters have been performed to clarify phylogenetic rela- lems with the phylogenetic positions of the Triodontidae tionships of various families and molecular analyses have and Molidae due to conflicts of differing positions in greatly progressed to provide detailed phylogenetic rela- morphological and molecular studies (e.g., Molidae has tionships of families, genera, and even species. The pur- been placed differently among molecular studies). pose of this paper is to provide a review on developments in the taxonomy and systematics of the Tetraodontiformes, Keywords Tetraodontiformes Á Classification Á focusing primarily on contributions since 1980 (when Morphology Á Molecular Á Phylogenetic relationships James C. Tyler’s monumental work was published) through the period of IPFCs, including pertinent publica- tions before 1980. This paper recognizes 412 extant species Introduction in the 10 families of living Tetraodontiformes, with the allocation of species and genera as follows: Triacanthodi- Tetraodontiform fishes are distributed in tropical to tem- dae including 23 species in 11 genera, Triacanthidae seven perate seas and freshwaters of the world. They show a remarkable diversity in shape, size, and way of life (Fig. 1). A small filefish of the genus Rudarius Jordan and This article was registered in the Official Register of Zoological Fowler 1902 and a small pufferfish of the genus Carino- Nomenclature (ZooBank) as C37A57BA-30FF-48AD-8AEC- tetraodon Benl 1957 mature at about 2 cm in total length 14C084BBB4BA. This article was published as an Online First article on the online (TL) (Lim and Kottelat 1995; Tyler 1970), whereas ocean publication date shown on this page. The article should be cited by sunfishes of the genus Mola Koelreuter 1766 attain over using the doi number. 300 cm TL (Yoshita et al. 2009). Tetraodontiform fishes are characterized by a small mouth with either relatively & K. Matsuura ( ) few teeth that are often enlarged or massive beak-like tooth Division of Fishes, National Museum of Nature and Science, 4-1-1 Amakuobo, Tsukuba, Ibaraki 305-0005, Japan plates, a small gill opening restricted to the side of the e-mail: [email protected] body, scales usually modified as spines, enlarged plates, or 123 K. Matsuura 123 Taxonomy and systematics of tetraodontiforms b Fig. 1 Representatives of the 10 extant families of Tetraodontifor- systematics of the Tetraodontiformes, focusing primarily mes. a Triacanthodidae, Triacanthodes anomalus; b Triacanthidae, on contributions from 1980 (when James C. Tyler’s Triacanthus biaculeatus; c Balistidae, Abalistes filamentosus; d Mon- acanthidae, Thamnaconus hypargyreus; e Aracanidae, Kentrocapros monumental work was published) through the period of aculeatus; f Ostraciidae, Ostracion immaculatus; g Triodontidae IPFCs, but including pertinent publications before 1980. Triodon macropterus; h Tetraodonitidae, Arothron mappa; i Diodonti- This paper is composed of two parts, the first reviewing dae, Diodon liturosus; j Molidae, Masturus lanceolatus. Photographs taxonomic studies and the second focusing of studies on of a and e provided by BSKU; b, d, f, h, and i by KAUM; c and g by NSMT; j by Hideki Sugiyama systematics. Institutional abbreviations follow Fricke and Eschmeyer (2014). a carapace, and pelvic fins that are reduced or absent (Tyler 1980; Nelson 2006). In addition, many pufferfishes are Triacanthodidae (Spikefishes, Fig. 1a; Table 1) characterized by having a strong toxin in the viscera and skin, and even in the musculature of some species of Spikefishes of the family Triacanthodidae are usually Lagocephalus Swainson 1839 (Matsuura 1984). Because of found in depths of 100-600 m on continental shelves and their peculiar morphological characters, tetraodontiforms slopes (Tyler 1968; Matsuura and Tyler 1997). They are have long attracted the attention of ichthyologists and easily distinguished externally from other families of the biologists. order Tetraodontiformes by the following combination of Since Cuvier (1816) classified tetraodontiforms in the characters: body deep and slightly compressed, covered by order Plectognathi, tetraodontiforms were usually consid- moderately thick skin with numerous small scales not ered to form a monophyletic group among the advanced readily distinguishable to the unaided eye, with each scale percomorph fishes (Nelson 2006). Although the taxonomy bearing upright spinules and having a roughly shagreen- of tetraodontiform fishes progressed greatly in the nine- like appearance; two separate dorsal fins, six spines in the teenth century (see Tyler 1980, for a history of the clas- first dorsal fin and 12-18 soft rays in the second dorsal fin; sification of the order), information about tetraodontiform caudal fin rounded to almost truncate, not forked; most taxonomy was scattered in many articles, making it diffi- dorsal-, anal-, and pectoral-fin rays branched; pelvic fins cult to understand the taxonomic relationships of tetra- with a large spine and one or two inconspicuous and odontiforms overall. Alec Fraser-Brunner published an rudimentary soft rays; mouth small and usually terminal; important series of articles on various groups of tetra- teeth of moderate size, usually conical, 10 or more in an odontiforms as reviews of genera and families during the outer series in each jaw; caudal peduncle compressed, mid-1930s to early 1950s: however, many parts of his deeper than wide, not distinctly tapered (Matsuura 2001). publications were based on cursory examinations of rela- Spikefishes are still relatively poorly known in terms of tively few specimens. There were no comprehensive phy- taxonomy and biology among tetraodontiform families, logenetic studies until Winterbottom (1974) and Tyler although Tyler (1968) published an excellent monograph (1980) provided their interpretations of the phylogenetic on the superfamily Triacanthoidea including the Triacan- relationships of tetraodontiforms based on myology and thodidae and Triacanthidae. He recognized 19 triacantho- osteology, respectively. did species in 11 genera (Table 1): Atrophacanthus Fraser- When the first Indo-Pacific Fish Conference (IPFC1) Brunner 1950 (one species), Bathyphylax Myers 1934 (two was held in Sydney in 1981, many problems remained in species), Halimochirurgus Alcock 1899 (two species), the generic- and species-level taxonomy of all tetra- Hollardia Poey 1861 (three species), Johnsonina Myers odontiform families, except for the Triacanthodidae and 1934 (one species), Macrorhamphosodes Fowler 1934 (two Triacanthidae for which Tyler (1968) had provided a species), Mephisto Tyler 1966b (one species), Parahol- monographic revision. The period from IPFC1 to IPFC9 lardia Fraser-Brunner 1941b (two species), Paratri- (1981-2013) was a time of great progress in the taxon- acanthodes Fowler 1934 (two species), Triacanthodes omy and systematics of tetraodontiforms. Many review Bleeker 1857 (two species), and Tydemania Weber 1913 and revisional papers have been published for various (one species). Because all these genera and species were genera and species, with descriptions of new taxa found described in detail by Tyler (1968), their taxonomic fea- mainly in coral reefs and tropical freshwaters; and cla- tures are not repeated here, except for new information that distic analyses of morphological characters have clarified provides a better understanding of taxa. phylogenetic relationships of a number of families and Many species of the Triacanthodidae are known from molecular analyses greatly assisted our understanding of the tropical and warm regions of the Indo-Pacific. How- the detailed phylogenetic relationships of families, genera, ever, five species are distributed in the western Atlantic: and even species. The purpose of this paper is to present a Hollardia hollardi Poey 1861, Hollardia meadi Tyler review of the developments
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