JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology SpeciesDiversity,2008, 13, 1･ 34 Phylogenetic Systematics of the Family Peristediidae (Teleostei: Actinopterygii) Toshio Kawai Ciollection Center, IVUtional Mttseum ofAJUture and Science, 3-23-1 Flyakunin-cho, Shinjuku-ku, Tokyo, 169-OOrs Jtrpan E-mail: kawai@,kahaku..uo.jp (Received 3 November 2006; Accepted 10 January 2008) The family Peristediidae, comprising about 36 species of armered sea robins in five genera, inhabit the bottoms of the tropical and temperate wa- ters of the world oeeans in depths of about 50 to 800m. The aim of this studs, is to infer Telationships among the species in the family based on morpho- logical characters and to revise the genus-level classification en the basis of the inferred pattern of phylogeny. Twenty-four peristediid and 16 outgroup taxa, i.e., 13 triglid and three hoplichthyid species, were used for the phylo- genetic analysis. Monophyly, of the Peristediidae is highly corroborated and two major clades are recognized in the family. The first clade ineludes Gar- gariscus, Heminodus, Satyrichtbys, and Paraheminodus, and the second in- cludes only Peristedion. It beeame clear that the genus SatyrichthJ)s is a non- monophyletic group, In conclusion, six monophyletic genera are recogriized in the Peristediidae: Oargariscus, Hentinodus, Paraheminodus, Peristedion, Satyrichtltys, and Scalicus, Accordingly, I propose seven new combinations with Scalicus as follows: Scaticus engyceros, S. gilberti, S. hians, S. investiga- toris, S. orientalis, S, quadratorostratus, and S. serrutatus, Key Words: Teleostei, Scorpaeniformes, Peristediidae, phylogenetic sys- tematics. Introduction Armored sea robins, the family Peristediidae (sensu Nelson 2006), comprise about 36 benthic species that are currently classMed tn four or five genera (Miller 1974; Kawai et al, 2004a, 2004b; Nelson 2006). Fishes of this family inhabit tropical and temperate waters of the world oceans in depths ranging from about 50 to 800m. The family is characterized by having the body encased in bony plates as well as by pessession of a rostral projection, the lower two pectoral fin rays free, and bar- bels on the lower jaw (Nelsen 2006), Systematics of the Peristediidae have been studied by many ichthyologists. Most of them have considered that the Peristediidae are closely related to the Triglidae (e.g., Gill 1888; Matsubara 1943; Greenwood et at, 1966; Washington et al. 1984). Recognition of genera has been subject to disagreement, Nelson (1976, 1984), Eschmeyer (1998), and Yamada (2e02) listed fbur genera in this family, viz., Periste- dion Lac6pede, 1801, Salyrichthys Kaup, 1873, Gargariscus Smith, 1917, and Henzi- nodus Smith, 1917, while treating Paraheminodus Kamohara, 1957 as a junior syn- onym of Satyrichthys without comment. On the contrary, Miller (1974), del Cerro NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 2 Toshio Kawai and Lloris (1997), Richards (1999), and Kawai et aL (2004a, 2004b) regarded Para- heminodus as valid. Imamura (1996) inferred the phylogenetic relationships of the Platycephalidae and related taxa using anatomical and external morphological characters. He fbund nine synapomorphies supporting the monophyly of the Peristediidae, Re- cently, the phylogenetic relationships of the superfamily Scorpaenoidea were in- ferred cladistically by Imamura (2004), and 14 synapomorphies supported again the monophyly of the Peristediidae. The sister group of the Peristediidae was the fam- ily Hoplichthyidae in both studies, Smith and Wheeler (2004) investigated the mo- lecular phylogeny of 69 scorpaenifbrm species and they also concluded that the Peristediidae are a monophyletie group, but that the sister group is the family Triglidae, not the Hoplichthyidae. Interrelationships within the Peristediidae have never been fu11y examined. The aim of this study is to infer relationships among species of the Peristediidae, and to propose a revised genus-level classification re- flecting the inferred phylogeny. Materials and Methods Data acquisition Osteological and myological examinations were made on specimens stained with Alizarin Red-S and Alcian Blue, dissected under microseopes (Leica MZ 8 and MZ 12), and drawn using a camera lucida. The teuminology fo11ows Imamura (1996, 2004) for the osteology, Winterbottom (1974) and Imamura (1996) for myology, and Teague (1961) and Kawai et al, (2004a) for external morphology, Measurement of standard length (SL) follows Hubbs and Lagler (1958). The institutional codes fo1- low Leviton et al. (1985), with the addition of Miyazaki University, Fisheries Sci- ence, Japan (MUFS). Terminal taxa I dissected specimens of 24 peristediid, 13 triglid, and three hoplichthyid species (see Appendix), and used primarily these 40 species fbr the phylogenetic analysis. The fbllowing 11 species were a]so examined, but dissection of representative specimens was not allewed (see Appendix): Paraheminodus kamoharai Kawai, Imamura, and Nakaya, 2004, P. Iaticephalus <Kamohara, 1952), Peristedion am- blygeays Fowler, 1938, Satyrichtdys clavilopis b]owler, 1938, S. hians (Gilbert and Cramer, 1897), S. tsokawae Yatou and Okamura, 1985, S. Iingi (Whitley, 1933), S. Iongiceps (Fowler, 1943), S. nza.crnus Yatou, 1985, S. rugosus (Fowler, 1943), and S. weZchi (Herre, 1925). The examinatien enabled me to assess only the external fea- tures of these species accurately. Although no specimens were available for examination of Peristedion altipin- nis Regan, 1903, P. crustosum (Garman, 1899), P, imberbe Poey, 1861, P. paucibarbi- ger Castro-Aguirre and Garcia-Dominguez, 1984, P. riversandersoni (Alcock, 1894), P. unicuspis Miller, 1967, Satyrichthys engyceros (GUnther, 1872), S. investigatoris (Alcock, 1898), S. moluccensis (Bleeker, 1851), and S, quadratorostratus (Four- manoir and Rivaton, 1979), data on the external characters of these species were taken from previous descriptions. I tried to assign these species to the clades recog- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Peristediid systematics 3 nized in this study based on the available information, although the assignment should be considered provisional for some species. Phylogenetic methods The present analysis was based primarily on an examination of 24 peristediid, 13 triglid, and three hoplichthyid species, as mentioned above, A data matrix ef 26 taxa (two outgroups and 24 ingroups) and 23 transfbrmation series (TS) was con- structed (Table 1). Characters only fbund among outgroup taxa were not included in the analysis, because the aim of this study was to infer relationships among in- group species. If the 16 outgroup species were coded independently, relationships among the taxa were not fully resolved because of much homoplasy. Therefore, the 13 triglid and three hoplichthyid species were united as two family-level terminal taxa in the construction ef the data matrix. This procedure is justified because the monophyly of these two families is well established (Imamura 1996, 2004). Charac- ters for the outgroup taxa were coded as a single state when all members of the family shared the same character state. When character states differed among sub- taxa, the state was coded as polymorphic. The multistate transformation series "unordered". was treated as All characters were unweighted fbr the sake of objec- tivity. Characters showing intraspecific variation were not considered. The data were analyzed using PAUP*4.0blO (Swofford 2002>, with ACCTRAN fbr character optimization, 1000 heuristic searches involving random addition sequenees, and TBR (tree bisection and reconnection) branch swapping under the optimality crite- rion of parsimony, Results Characters used for phylogenetic analysis Osteology. CrlS J, Fourth infraorbital and hyomandibular (O: separated; 1: su- tured) (Figs 1, 2). The fourth infraorbital is separated flrom the hyomandibular in Peristedion, Triglidae, and Hoplichthyidae (TS 1-O), whereas these bones are su- tured in Gangariscus, Sa4yrichtbys adeni (Lloyd, 1907), and S. riqffeli (Kaup, 1859) (TS 1-1). In Henzinodus, Paraheminodus, Satyrichthys amiscus (Jordan and Starks, 1904), S. gilberti (Jordan, 1921), S. orientalis (Fowler, 1938), and S. serrulatus (AI- cock, 1898), the relation between the fourth inflraorbital and the hyomandibular is undeterminable (coded as ?), because the hyomandibular is fused with the fifth in- firaorbital (Figs 1, 2C). 71S 2. Fifth imbaorbital and hyomandibular (O: sutured; 1: fused) (Figs 1, 2). The fifth inhraorbital and hyomandibular are clearly sutured in Gailgariscus, Periste- dion, Satyrichth{vs adeni, S. riofeli, Triglidae, and Hoplichthyidae (TS 2-O), whereas they are fused in Heminodus, Paraheminodus, Satyrichthys amiscus, S. gilberti, S. orientalis, and S. serrulatus (TS 2-1). 71S 3. Sixth infraorbital and sphenotic (O: sutured; 1: fused) (Fig. 3), The sixth infiraorbital is sutured to the sphenotic in Garlgariscus, Peristedion, SaCyrichthys adeni and S. riqtlTeli, Chetidonichtbys, Lepidotrigla, and Pter3,gotrigla (TS 3-O), whereas these bones are fused in Heminodus, Paraheminodus, Satyrichtdys amis- cus, S. gilberti, S orientalis, S. serrulatus, BeZlator, Prionotus, and Hoplichthyidae (TS 3-1). NII-Electronic Library Service Japanese SooietySociety of SystematicSystematio Zoology 4 Toshio Kawai 卜 ・ひ ・ OOOHHHHHHHHHHHHHHHoooooo の NIH o 卜・ ooor 図HHHHHHHHHH 阿 囲倒 剛例 HOOHri oo HHH 問 問 個 闘 囚 嗣 闘 囚 咽 H 国 刳 囚 HHHHH 一 ＜ H 州 州 剛 回 HHHHHHHH