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Actinopterygii: Perciformes

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Actinopterygii: Perciformes JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Species Diversity. 2004, 9, 1-36 Phylogenetic Relationships and New Classification of the Superfamily Scorpaenoidea (Actinopterygii: Perciformes) Hisashi Imamura lhe HOickaido University Museum, hacudy ofFisheries, HOkkaido Ultiversity, 3-1-1 Minato-cho. Htikodate, Hbkkaido, 041-8611 .Jtipan E-maii;imamura/(ti/museum.hokudai.ac.jp (Received 10 October 2e02; Accepted 10 August 2003) re- The phylogenetic relationships of the supeirlami!y Scorpaenoidea are constructed cladistically. based on speeimens belonging to 18 families, 59 genera, and 86species, by using osteological and myological characters rec- ognized in 111 transformation series. The tbllowing relationships are in- ferred; <1) the former Scerpaenoidei is paraphyletic; (2) the family Sebasti- dae is not monophyletic, most of the genera included having initially branched ofi' other ingroup taxa: (3) the family Setarchidae and Trachyscor- pia have a sister relationship, being nested within the paraphylatic Scor- paenidae; and <4) the family Neosebastidae is the sister group of the former Platycephaloidei, In conelusion, the Scorpaenoidea is reclassified into 20 families, accepting several redefined taxa, such as the Sebastidae, Sebastolo- bidae, and Seorpaenidae. Key Words: Scorpaenoidea, monophyly, phylogony, classification. Introduction The percifbrm superfamily Scorpaenoidea, including the former scorpaeni- fbrm suborders Scorpaenoidei and Platycephaloidei (sensu Imamura 1996; Esch- meyer 1998), was proposed by Imamura and Yabe (2002), who separated the poly- phyletic Scorpaenifbrmes into the perciform Scorpaenoidea and Cottoidei. Al- though Imamura (1996) had earlier recognized the monophyly of the super:lamily and inferred its phylogenetic relationshtps, his cladogram included many unre- solved relationships. Furthermore, although many ichthyologists have recognized the former Scorpaenoidei as a valid taxonomic unit (e.g., Greenwood et al, 1966; Nelson 1976, 1984, 1994; Ishida 1994; Shinohara 1994; Imamura 1996; Imamura and Shinohara 1998; Eschmeyer 1998; Mandrytsa 2001), neither Shinohara (1994) nor Imamura (1996) could find any synapomorphies supporting the monophyly of that taxon. Imamura and Yabe (2002) did not provide taxonomic names corresponding to the former suborders Scorpaenoidei and Platycephaloidei. Recently, although Mandrytsa (2001) initially reconstructed the phylogenetic relationships of Scorpaenoidei (excluding Platycephaloidei) using PAUP ver. 3.1.1, he did not accept the findings and reanalyzed by hand (but it is unclear what methodology he fo11owed) both the fami!ial relationships within his Scorpaenoidei and the intergeneric relationships within each family, He recognized the mono- phyly of his Scorpaenoidei. excluding the fetmilies Congiopodidae (sensu Nelson NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 2 Hisashi Imamura 1994) and Pataecidae, on the basis of two derived characters (presence of three neu- romasts on the first infraorbital and insertion of the first two dorsal proximal pterygiophores into the space between the second and third neural spines), using Percoidei as an outgroup. However, these characters are found not acceptable in the present study, as is explained below. In addition, Mandrytsa (2001) recognized several paraphyletic taxa (e.g., subfamilies Scerpaeninae and Neocentropogoni- nae). The purposes of the present study are to reconstruct the phylogenetic relation- ships of the Scorpaenoidea and to propose a new classification based on the rela- tionships. Methods and Materials The terminology fbllows Imamura (1996) for osteolog}i, except fbr infraorbitals and extrascapulars, which fo11ow Imamura (2000) and Imamura and Yabe (2002), respectively, and Winterbottom (1974) for myology, exeept for the swimbladder muscles, which fbllow Hallacher (1974). Institutional abbreviations fbllow Leviton et al. (1985). Although Serranidae (sensu Baldwin and Johnson 1993) was inferred to be closely related to Scorpaenoidea by Imamura and Yabe (2002), the most closely related serranid taxon to Scorpaenoidea has not been specMed; in this study, therefore, three subfamilies (Anthiinae, Epinephelinae, and Serraninae) were used as separate outgroups (Fig, 1). The data were analyzed using PAUP' 4.0blO (Swoflbrd 2002), including the heuristic search option. Character evolution "ordered" "unordered" was assumed as (Wagner parsimony) or (Fitch parsimony) "loss" when the transformation series (TS) contained a derived or character order was equivocal owing to many modifications. The initial familial classification of the scorpaenoids fbllowed Ishida (1994), except for the plectrogeniid PlectrQgenium, and Imamura (1996). Material examined Scorpaenoidea. Apistidae: Apistus carinatus (Bloch, 1801), HUMZ 37372 and one uncatalegued specimen (2, 93-102mm SL). Aploactinidae: Aploactis aspera (Richardson, 1845), HUMZ 37483 (55mm); Erisphex potti (Steindachner, 1896), HUMZ 64275, 108774 (2, 74-87mm). Bembridae: Bembras jqponica Cuvier, 1829, HUMZ 49409, 49411, 49412 (3, 160-182mm), Congiopodidae: Alertichthys btacki (Moreland, 1960), HUMZ 66621 (161 mm); Congiopodus coriaceus Paulin and More- land, 1979, HUMZ 91192 (142mm), Gnathanacanthidae: Gnathanacanthus goetzeei Bleeker, 1855, AMS I. 20188001 (47mm) and AMS IB. 668 (158mm, dissected by Ishida 1994). Hoplichthyidae: Hbplichthys gilberti Jordan and Richardson, 1908, HUMZ 51736 (152mm); U haswelli McCulloch, 1907, HUMZ 50268 (341mm); Il tangsdoi:fii Cuvier, 1829, HUMZ 75337 (156mm). Neosebastidae: Maxillicosta raoulensis Eschmeyer and Poss, 1976, HUMZ 110722 (88mm); AJeosebastes thetidis (Waite, 1899), HUMZ 21124 (196mm). Parabembridae: Parabembras curta (Tem- minck and Schlegel, 1843), HUMZ 108335, 108460, 108778 (3, 125-165mm). Pataecidae: Pataecus fronto Richardson, 1844, AMS I. 38479002 (110mm), Peristediidae: Periste- dion orientale Temminck and Schlegel, 1843, HUMZ 106621 (152mm); Satyrichtdys serrutatus (Alcock, 1898), HUMZ 108541 (200mm). Plectrogeniidae: Bembradium ro- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Phylogeny of Scorpaenoidea 3 seum Gilbert, 1905, HUMZ 75105, 79190 (2, 85-133mm); Ptectrogenium nanum Gilbert, 1905, HUMZ 37337 (52mm). Scorpaenidae: Dendrochirus zebra (Quoy and Gaimard, 1824), HUMZ 63987 (132mm); Ponttnus macrocephalus (Sauvage, 1882), HUMZ 99503 (121mm); Pterois volitans (Linnaeus, 1758), NSMT-P 54350 (93mm); Scorpaena izensis Jordan and Starks, 1904, HUMZ 79974 (134mm); Scorlpaenodes lit- toralis (Tanaka, 1917), HUMZ 101394 (74mm); ScorpaenQpsis neglecta Heckel, 1837, HUMZ (uncatalogued, 125mm); Taenianotus triacanthus Lacepede, 18e2, USNM 106917 (60 mm). Sebastidae: AdeZosebastes latens Eschmeyer, Abe and Nakano, 1979, HUMZ 72035, 75569 (2, 206-2e7mm); Helicolenus hiigendoi:t7 (Steindachner and DOderlein, 1884), HUMZ 39743 (136mm); H percoides (Richardson and Solander, 1842), HUMZ 91501 (164mm); Hbzukius gayotensis Barsukov and Fedrov, 1975, HUMZ 71936 (232mm); Sebastes matsubarae Hilgendorf 1880, HUMZ 90303 (114 mm); S. owstoni (Jordan and Thompson, 1914), HUMZ 42641 (183 mm); S. thompsoni (Jordan and Hubbs, 1925), HUMZ 92291 (156mm); Sebastiscus marmoratus (Cuvier, 1829), HUMZ 69895, 117714 (2, 125-157mm); Sebastolobus macrochir(GUnther, 1880), HUMZ 59268, 68329 (2, 112-169 mm); Tracbyscorpia eschnzeyeri Whitley. 1970, NSMT- P 41332 (235 mm). Setarchidae: Ectreposebastes imus Garman, 1899, HUMZ 751]2 and one uncatalogued specimen (2, 85-113mm); Setarches longimanus (Alcock, 1894), HUMZ 79505 (125 mm). Synanceiidae: ChoridactyIus multibarbus Richardsen, 1848, CAS 15067 (71mm); Erosa erosa (Cuvier, 1829), HUMZ 64250 (63mm); Inimicus japonicus (Cuvier, 1829), HUMZ 79103, 140587 (2, 104-143 mm); Minous monodactylus (Bloch and Schneider, 1801), HUMZ 101864 (95mm SL); S,nanceia horrida (Lin- naeus, 1766), CAS 15073 (106mm). Tetrarogidae: TetrarQge niger (Cuvier, 1829), URM-P 31819 (18mm, data from G. Shinohara). Triglidae: Bellotor egretta (Goode and Bean, 1896), HUMZ 69393 (87 mm); Chelidonichthys spinosus (McClelland, 1844), HUMZ 108541 (200mm); Lepidotrigla micrQptera GUnther, 1873, HUMZ 48942 (159 mm); Prionotus stearnsii Jordan and Swain, l884, HUMZ 32456 (98mm); Pter.v- gotrigla macrortrynchus Kamehara, 1936, HUMZ (uncatalogued, 95mm). In addition, platycephalid specimens, including l6 genera and 38 species, were also examined, such being listed in Imamura (1996). Comparative materials Serranidae. Anthiinae: Ptectranthias keltoggi azumonus (Jordan and Richard- son, 1910), HUMZ 79389 (104 mm). Epinephelinae: Diploprion bdesciatum Kuhl and van Hasselt, 1828, HUMZ 87031 (122mm); EZpinqphelus awoara (Temminck and Schlegel, 1843), HUMZ 97035 (125mm); Niphon sptnosus Cuvier, 1829, HUMZ (uncat- alogued, 126mm). Serraninae: ChelidQperca hirundinacea (Valenciennes, 1831), HUMZ 36481 (102 mm). A further 31 percoid species that were previously examined were listed in Ima- mura (2000), Phylogenetic Analysis of Scorpaenoidea Monophyly of Scorpaenoidea Imamura and Yabe (2002) recognized the monophyly of the Scorpaenoidea on the basis of three synapomorphies: presence of a suborbital stay, the parietal sen- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 4 Hisashi Imamura sory canal bony structure with spines, and an extrinsic swimbladder muscle de- rived from the obliquus superioris (Fig. 1). They also suggested that the single pos- tocular spine in the larval
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