Teleostei: Actinopterygii

Home , Peristediidae, Peristedion, Peristedion greyae, Satyrichthys

JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

SpeciesDiversity,2008, 13, 1･ 34

Phylogenetic Systematics of the Family Peristediidae

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 waters 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 phylogenetic 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. investigatoris, S. orientalis, S, quadratorostratus, and S. serrutatus, Key Words: Teleostei, Scorpaeniformes, Peristediidae, phylogenetic systematics.

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 barbels 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 synonym 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 inferred cladistically by Imamura (2004), and 14 synapomorphies supported again the monophyly of the Peristediidae. The sister group of the Peristediidae was the family Hoplichthyidae in both studies, Smith and Wheeler (2004) investigated the molecular 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

tures of these species accurately. Although no specimens were available for examination of Peristedion altipinnis Regan, 1903, P. crustosum (Garman, 1899), P, imberbe Poey, 1861, P. paucibarbiger 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: sutured) (Figs 1, 2). The fourth infraorbital is separated flrom the hyomandibular in Peristedion, Triglidae, and Hoplichthyidae (TS 1-O), whereas these bones are sutured 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 infiraorbital (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 amiscus, 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 − HHHHHH 回州回

＜ OH OHoooooooooOoooooooooooo O 1 Φ ooOOOH OoOOOOOOOOOOOOOOOOOHHHHO 回

OOOOOHHH − HHHHHHHHH − HHHOOOOOO

eL ・O 例國剛一一倒一倒一倒州倒 oooooo

囲＜ OHHHHHH − HH 目 HHOOOOHOHr 咽H 祠 o の

帽旨 H O o 剛一鬥 HHOOOHOOOOOOOOOHHHHO Φ 鴎 00 HI

目． ooo OHHHH 州倒 HHHHHHHHHHHHHHHH 一一〇口邸 ooHHHOOOOOOOOOOOOOOOHHHHHH宀 HooooooooooooooooHHHHO 日旨 Q oo 噛の口邸」h ［ ooH OOOH 目 HHHHHHHHOOOHHoooooo ．の ooH 阿一〇〇〇〇〇〇〇〇〇〇〇〇〇〇〇 HHH 州祠囲一の〇あ一一 IO ooHH 囚一一一一 HH 円倒 HHHHHHH 同 σ『 H 邸自弼 oo − ooooooooooooooo 例肖 H 円 HH h 一ρ OH 州 ooo 祠目 H 鬥 HHHHH 州 OOHHHOHHOH つ， Φ の〈二 o ◎ にの Φ 噂一肖 oooo 鬥鬥 HOOOOOOOOOOOOOOOHHHHH 同旨 ObOO

oD O O 囲 HHHHHHHHHHHHH − HH 例剛 HH 口 O Φ H ．』冨冖＜ OHHOOOOOOOOOOOOOOOOHHH − o 而邸の ← H 聶眉』 O O OHHOOOOOOOOOOOOOOOOH − Hrfo 口 ε 邸の〇自 oo H び・ひ．・OOOOOOOOOOOOOOOH ひ・ひ・卜・ひ・一応一二と〇 αつ ⊇ 国 bDq 口 97 祠謡一丶唱 gD 鵠』弓コ趙汽一 Φ の〇一織遣 qkk 9 魁口陛一甸 OO 幅 ON 』離心の邸鬻轄旨ミ N 』唱 ONk§ Φ 謎一 O O 篭物唱 § ミ Nk ⇔ × 蒿 NkO o 鶏』 cd ミっ越 O 邸邸織偽 NgO 鋤 O ℃ ℃ 鴇識ミ軸鞍朝熊嵩ミ k 的的』り O N ぺ N ミ建的邸 × のの勉 k 識鴇ミ摘越 h 嵩匿廻 9 ピ嵩 O の轟一凡魁鼠 k NO 劬 N 冖‘ 9 鳶描む穐の謡き角 OO ON k 齪 O N 燭 OkN 楠輪遣鳶り剃の ← の旭劬ミ努 q9 邑楠ミ ”隔ミ隔 ℃ 幅乱摘丶篇 ◎ 鳶 k 9 O 邸 ‘ O 魁り賠 § 丶国 q50 Qロ k 勉き Qk QOk 鋤霞物謡肩り燭 kq9 む § 賄ミ、急 9Nk 鴇ミ ◎ 窓虻 N 」 Q §穐匿 “ 鋤 kk 匿帽 q N 謡 NO9 能訂 O 鋤摘嚇効霜一肖防隔 O む嵩ミ嵐＝ dk 職 9 O 」ミ魅 kO ミ勉魅 q ．角 ⇔Ω 艶ミ墫 O Q 冒 qO 州 O ミ k 嚇 S 隔謎穐穐 6 劬ミ魯如 G り嚀角 bQ 』轄弓勉 O 一角．，．．．．．．．．．．．．eg ，，，．〔 ∩ 〔【寓臼 O ミ馬「蟻蟻馬蟻 q ヘ気 q 儀気 q 気「「つののの賄の［コ［d q ρ 眺 O H 【面 O 臼山

一 NII-ElectronicN 工工 Eleotronio Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid svstematics 5

Fig, 1. Lateral aspect of head ef Heminodus phitmpinus, HUMZ 193268, 174.5 mm SL. Abbrevia- tions: ECP, ectopterygoid; ETH, ethmoid; FRO, frontal; HYO, hyomandibular; IO, infiraorbital; LET, lateral ethmoid; NA, nasal; OP, opercle; PA, parietal; POP, preopercle; PT, posttemporal; PTO, pterotic; SCL, supracleithrum; SOP, suboperc]e; SI]O, sphenotic; ST,

supratemporal, Scale bari 5mm.

A B c

Fig. 2, Diagrammatic illustration showing relationships among fOurth and fitth infraorbitals and hyomandibular, A, TS 1-O; B, TS 1-1; C, coded as ?. Abbreviations: HYO, hyomandibular;

IO, infraorbitaL

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

6 Toshio Kawai

' ttt .t. t '',]-,"//'1.ts.i.:i'I/'i.,i.il}･'･,1',',･;'S!'･.,t.t .. ttttttttt. t. .ttt zrl-/ ttt..' pv t t tttttttt.... x B t t.ttttt.t t. 't='/': "' ±=. t' -/ 'xdr' gS･ -x

Fig. 3. Lateral aspect of neurocranium. A, Garlgariscus prionocephatus, BSKU 17658, 148.0mm SL; B, lleminodus philippinus, HUMZ 193268, 174.5mm SL; C, Paraheminodus murrop,i, BSKU

4599, 139.3mm SL; D, Peristedion orientale, HUMZ 36744, 125.4mm SL; E, Satyrichtlrys anziscus, HUMZ 51851, 147.5mm SL; E Satyrichthys riqEfett, HUMZ 90522, 145,8mm SL, Abbre- viations: BO, basioccipital; EO, exoccipital; ETH, ethmoid; FRO, frontal; IC, intercalar; IO, in- fr-aorbital; LET, ]ateral ethmoid; NA, nasal; PA, parietal; PAS, parasphenoid; PRO, prootic; PT, posttemporal: PTO, pterotic; PTS, pterosphenoid; PV, prevomer; SPO, sphenotic; ST,

supratemporal, Scale bars: 5mm.

7LS 4. Upper jaw teeth (O: present; 1: absent) (Fig. 4). The upper jaw teeth are prcsent in Gargarisc"s, Hkiminodus, Paraheminodus, Triglidae, and Hoplichthyi- dae (TS 4-O), but absent in Peristedion and SaCyrichttrys (TS 4-1). 71S 5. Interopercle and epihyal (O: directly attached posteromedially; 1: connected via long Iigament posteriorly) (Fig. 5). The interopercle is directly attached to the epihyal posteromedially in Peristedion and Triglidae (TS 5-O), but these bones are connected via a long ligament posteriorly in Gargariscus, Heminodus, Paraheminodus, Satyrichtbys, and Hoplichthyidae (TS 5-1).

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristeditd systematics 7 A RC B MAX/ 7 PRM

D - / gyt.vktey× ;)o

-'rfo"tL ` " <:otta!L. bl.. }

"/J/J--ev4 Fe - C ("7----x. xx----- u N2> xt.. t y 7' " /V-'x...=.!..3-7k " ee,JVA ct,< ,tfx e.sfNtt

Fig. 4. Lateral aspect of jaws, A, Gargariscus prionocephalus, BSKU 17658, 148.0mm SL; B, Heminodus phitippinus, HUMZ 193268, 174.5mm SL; C, Paraheminodus murrayi, BSKU 4599, 139.3mm SL; J), Peristedion orientale, HUMZ 36744, 125.4mm SL; E, Satyrichtltys amiscus, HUMZ 51851, 147.5mrn SL; F, SaCyrichthys riqtfeli, HUMZ 90522, 145.8mm SL, Abbreviationsi ANG, anguloarticular; DEN, dentary; MAX, maxilla; PRM, premaxilla; RC, rostral cartilage;

RET, ratroarticular, Scale bars: 5mm.

TS 6 Ventral region of cartilaginous band between ceratohyal and epihyal (Oi not extending to ventral margin of ceratohyal; 1: extending to ventrai margin of ceratohyal)

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology

8 Toshio Kawai B

x------Lv <

li iiiiiiilii E F " "m u ' ----!- --tcrf

XV N \rvtgii}iti :=:r

5, aspect of hyoid arch, Fig. Lateral A, Gargariscus prionocophalus, BSKU 17658, 148,Omm SLi B, Htirninodus IIUMZ phitippinus, 193268, 174.5mm SL; C, Paraheminodus murrayi, BSKU 4599, 139.3mm SI,; D, Peristedion orientale, HUMZ 36744, 125,4mm SL: E, Sa4yrichtnj's amiscus, HUMZ 51851, 147.5mm SL: F, SaCyrichtttys riqtfizti, HUMZ 90522, 145.8mm SL. Abbre- viations: BR, branchiostegal;CH, ceratohyal; DHH, dorsal hypohyal; EH, epihyal; IH, inter- hyal; IOP, tnteropercle; VHH, ventral hypohyal. Scale bars: 5mm.

serrulatus, and Hoplichthyidae (TS 6-O), whereas this region extend to the ventral margin of the ceratohyal in GaTErariscus, the other examined species of Peristedion and Satyrichtiays, and Triglidae (TS 6-1). TS Z Shape of third basibranchial (O: slender; 1: robust) (Fig. 6). The third basibranchial is slender in Peristedion, Triglidae, and Hoplichthyidae (TS 7-O), but robust in Garzgariscus, Heminodus, Pczraheminodus, and Satyrichthys (TS 7-1). 7S 8. Second pharyngobranchial and first epibranchial (O: directly connected; 1: connected via interarcual cartilage; 2: connected via long, thin ligament> (Fig. 7). The second pharyngobranchial is directly connected with the first epibranchial in the Triglidae and Hoplichthyidae (TS 8-O); both bones are connected via an interarcual eartilage in Gangariscus, Peristedion, Satyrichtdys adeni, and S, riE:tXizli (TS 8-1); and both bones are connected via a long, thin ligament in Ileminodus, Parahemin- odus, SatyrichtJtys amtscus, S. gilberti, S. orientalis, and S. serrulatus (TS 8-2), rs 9. Cartilaginous band between scapula and coracoid (O: moderately broad;

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematies 9 iBg4

Fig. 6. Dorsal aspect of lower branchial arch. A, Peristedion orientale, HUMZ 177798, 119.5mm SL; B, Heminodus phittppinus, HUMZ 193268, 174.5mm SL; C, SaCyrichthys amiscus, HUMZ

51838, 139,8mm SL. Abbreviations: BB, basibranehial; BH, basihyal; CB, ceratobranchial; HB,

hypobranchial. Scale bars: 5mm,

1: extremely broad) (Fig. 8). The cartilaginous band between the scapula and the coracoid is moderately broad in Peristedion, Triglidae, and Hoplichthyidae (TS 9- O), but extremely broad in the other peristediids (TS 9-1), 71S la Number of postcleithra (O: one; 1: two) (Fig, 8). Gar:gariscus, Ueminodus, Paraheminodus, Peristedion antillarum, P. gracile, P. thompsoni Fowler, 1952, Satyrichtdys, Triglidae, and Hoplichthyidae have a single postcleithrum (TS 10-O), whereas the other examined species of Peristedion have two posteleithra (TS 10-1). 7LS 11. First epineural (O: present; 1: absent) (Fig. 9). An epineural is present on the first vertebra in Peristedion, Triglidae, and Hoplichthyidae (TS 11-O), but absent in Garlgariscus, Heminodus, Parahenzinodus, and Sa(yrichtlrys (TS 11-1). 71S 12, Se¢ ond epineural (O: present; 1: absent) (Fig. 9). An epineural is present on the second vertebra in Gargariscus, Pertstedion, Satyrichtbys adeni, S. riqtfeli, Triglidae, and Heplichthyidae (TS 12-O), whereas it is absent in Heminodus, Para- heminodus, Satyrichtnjs amiscus, S. gilberti, S, ertentalis, and S, serrulatus (TS 12- 1). TS 13, Dorsal stay (0: autogenous; 1: fused) (Fig. 10). The dorsal stay is autoge-

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

10 Toshio Kawai

EBfiv-e m PB4IAC c D

Qlj ,

E F .-..-vi{i{) f) gtrts x

Fig. 7. Dorsal aspect of upper branchial arch. A, (]angariscus prionocephalus, BSKU 1765B, 148.0mm SL; B, Hetninodus philippinus, HUMZ 193268, 174.5mm SL; C, Paraheminodus mur-

rayi, BSKU 4599, 139.3mm SL; D, Peristedton orientale, HUMZ 36744, 125.4mm SL; E,

Satyrichthys amiscus, HUMZ 51851, 147.5mm SL; F, SaCyrichtPrys riqffl?li, HUMZ 90522,

145.8mm SL, Abbreviations: EB, epibranchial; IAC, interarcual cartilage; PB, pharyngob-

ranchial. Scale bars] 5mm.

nous in Gangariscus, Triglidae, and Hoplichthyidae (TS 13-O), whereas it is fused with the posteriormost dorsal proximal pterygiophore in Heminodus, Parahemin- odus, Peristedion, and SaCyrichthys (TS 13-1), 71S 14, Second preural centrum and its hemal spine (O: autogenous; 1: fused) (Fig. 11), The second preural centrum and its hemal spine arc autogenous in Gar- gariscus, Peristedion antillarum, P, barbi.uer

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systernatics 11

COR D

Fig, 8. Lateral aspect et' peetoral girdle. A, Gailg'ariscus prionocephalus, BSKU 17658, 148.0mm

SL; B, Heminodus philippinus, HUMZ 193268, 174.5mm SL; C, Paraheminodus murrqyi, BSKU

4599, 139.3mm SL; D, Figristedion ortentale, HUMZ 36744, 125.4mm SL; E, Satyrichthys amiscus, HUMZ 51838, 139,8 mm SL; F, Satyrichtlzys riqtfeli, HUMZ 90522, 145.8mm SI.. Abbre-

viations: ACT, acttnost; CLE, cleithrum; COR, coracoid; PC, postcleithrum; SCA, scapula;

SCL, supracleithrum. Scale bars: 5 mm.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

12 Toshio Kawai

Fig. 9. Lateral aspect of anterior and middle vertebrae and associated bones. A, Gangariscus

prionocephalus, BSKU 17658, 148.0mm SL; B, Peristedion orientate, HUMZ 36744, 125.4 mm SL; C, Satyrichthys amiscus, HUMZ 51851, 147.5mm SL, Abbreviations: DP, distal pterygiophore; EN, epineural rib; PP, proximal pterygiophore; PR, pleural rib; R, soft ray; S, spine. Seale bars: 5mm.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediidi systematics 13 B

Fig, 10, Lateral aspect of posterior vertebra and assoefiated bones, A, Gargariscus priono-

cephalus, BSKU 17658, 148.0mm SL; B, Peristedion tiorhynchus, HUMZ 178284, 144.3mm SL.

Abbreviationsi DP, distal pterygiophore; PP, proximal pterygiophore; R, soft ray; STA, stay.

Scale bars: 5mm. B P

1/t

Fig. 11. Lateral aspect of caudal skeleton. A, Gangariscus prionocephalus, BSKU 17658, 148.0 mm SL; B, Heminodus philippinus, HUMZ 193268, 174.5mm SI.; C, Satyrichthys amtsc"s, HUMZ 51851, 147.5mm SL; D, Satyrichtlays riqfil?li, HUMZ 90522, 145.llmm SL. Abbreviations:

EU, epural; HS, hemal spine; LCI', lower caudal plate; PU, preural centrum; UCP, upper cau-

dal plate. Scale bars: 5 mm.

S. serrulatus, and Hoplichthyidae (TS 14-1). ZS 15 Third preural centrum and its hemal spine (O: autogenous; 1: fused) (Fig. 11). The third preural centrum and its hemal spine are autogenous in Gar- gariscus, Peristedion antillarum, P. brevirostre, P. gracile, P. thompsoni,

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

14 Toshio Kawai

LAP

Flg. 12, Lateral aspect of cheek and cephalic muscles. A, Heminodus phitmpinus, HumZ 193268, 174,5mm SL; B, Peristedion ecuadorense, HUMZ 31502, 133.8mm Sl. Abbreviations: A,

adduetor mandibu]ae; AAP, adductor arcus palatini; DO, dilator epercu]i: LAP, levator arcus palatini; LO, levator operculi. Scale bars: 5mm.

Satyrichthys adeni, S, riqfifeii, and Triglidae, except fbr Lepidotrigla kanagashira Kamohqra, 1936 and Ptenygotrigla henzisticta (Temminck and Schlegel, 1843) (TS 15- O), whereas both bones are fused in Heminodus, Paraheminodus, the other examined species ot' Peristedion and Satyrichtdys, Lepidotrigla kana.crashira, Ptery- gotrigla hemisticta, and Hoplichthyidae (TS 15-1). Myology. TS 16 Shape of section A2 of adductor mandibulae (O: wide; Ii narrow) (Fig. 12). The adductor mandibulae section A2 is wide in Garlgarisc"s, H2mi- nodus, Paraheminodus, Satyrichthys, and Triglidae (TS 16-O), but narrow in Periste- dion (TS 16-1). The condition of this muscle can not be categorized as such in the

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid svstematics 15 B

Fig. 13. Medial aspect of cheek muscle and lower jaw. A, Peristedion orientale, HUMZ 177798, 119.5mm SL; B, Satyrichthys amiscus. HUMZ 51851, 147.5mm SL. Abbreviations: A, adductor

mandibulae; AW, adductor mandibulae section w; BB, barbelius; IM, intermandibularis.

Scale bars: 5mm.

Hoplichthyidae (thus eoded ?), because sections Al and A2 are fused. TS IZ Barbelius (O: absent; 1: present) (Fig, 13). The barbelius, supporting the chin barbels, is absent in Gargariscus, IIeminodus, Paraheminodus, Sa4);richthys, Triglidae, and Hoplichthyidae (TS 17-O), This muscle is only present in Peristedion (TS 17-1). TS 18 Connection of obliquus dorsalis (O: directly connected to the third and fourth epibranchials; 1: connected to the third and fourth epibranchials by a long tendon) (Fig. 14). The obliquus dorsa]is is directly connected to the third and fourth epibranchials in Gargariscus, Heminodus, Paraheminodus, Peristedion, SaC)srichtbys adeni, S. riqffeti, Triglidae, and Hoplichthyidae (TS 18-O), whereas this muscle is connected to these bones by a long tendon in Satyrichtltys amiscus, S. gitberti, S. orientalis, and S. serrulatus (TS l8-1). TS 19. Extensor proprius (O: attached to pectoral girdle; 1: separated from pectoral girdle) (Fig. 15). The extensor proprius is attaehed to the pecteral girdle in Gargariscus, Paraheminodus, Peristedion, Satyrichthys, Triglidae, Hbplichthys giZberti Jordan and Richardson, 1908, and Il Zangsdoittii Cuvier, 1829 (TS 19-O), but not in Hdeminodus and Hbplichthys haswelli McCulloch, 1907 (TS 19-1). 71S 20. Baudelot's ligament (O: originating frem basioccipital and first vertebra; 1: originating only from first vertebra; 2: originating only from basioccipital), The

Baudelot's ligament originates from the basioecipital and first vertebra in Bellator, Chelidonichthys, Lepidotrigla, and Prionotas (TS 20-O), but only from the first vertebra in Gaugariscus, Paraheminodus, Peristedion, Satyrichtdys, Pteilygotrigla, and Hoplichthyidae (TS 20-1), and only from the basioccipital in Heminodus (TS 20-2). External morphology. TS 2J. Barbels on lower jaw (O: absent; 1: present, not branched except posteriormost barbels; 2: present, all barbels branched) (Fig, 16). Barbels are absent on the lower jaw in the Triglidae and Hoplichthyidae (TS 21-O), but in all peristediid taxa the lower jaw is provided with barbels. Furthermore, in Gar:gariscus, HOminodus, Paraheminodus, Peristedion antillarunz, and Sat),richthys, only the posteriormost barbels are branched (TS 21-1), whereas in Peristedion species besides P. antillarum, all barbels are branched (TS 21-2). 7iS 22. Antrorse spines of posterior part of upper lateral row of bony plates (O: absent; 1: present) (Fig. 17). Antrorse spines are absent on the more posterior bony plates of the upper lateral row in Gangariscus, Heminodus, Paraheminodus,

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

l6 Toshio Kawai A B

TDP OBD

Fig. 14, Dersal aspect of upper branchial muscles after removal of TDA in rtght side. A, Hemtnodus philtpm)inus, HUMZ 193268, 174.5mm SL; B, Satyrichthys amiscus, HUMZ 51838, 139.8mm SL. Abbreviations: LE, levator externus; LEP, levator posterior; LI, levator internus; OBD, obliquus dorsalis; OBP, obliquus posterior; RD, radial dorsalis; SO, sphincter oesophagi; TDA, transversus dorsalis anterior; TDP, transversus dorsalis posterior. Scale bars: 5mm,

Satyrichtbys amiscus, S. orientalis, and Hoplichthyidae (TS 22-O), whereas these spines are present in Peristedion and the other examined species of Satyrichtltys (TS 22-1). In the outgroup Triglidae there are no bony plates on the body; therefbre, this taxon is coded as? for this character. 71S 23. Posterior pairs of bony plates in lower Iateral rows (Oi separated by ventral plate rows; 1: sutured contralaterally along midline) (Fig. 18). The more posterior pairs of bony plates in the lower lateral rows are separated in Garlgariscus, Heminodus, Paraheminodus, and Satyrichtdys (TS 23-O), but sutured along the midline in Peristedion. The Triglidae and Hoplichthyidae are coded as? fbr this character, because there are no bony plates on the body in the Triglidae, and no lower

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 17

B SP

Fig. 15. Dorsal aspect of pelvic fin muscles after removal of ADSP and EXP ln right side. A, Heminodus philippinus, HUMZ 193268, 174,5mm SL; B, Satyrichtltys amiscus, HUMZ 51838, 139.8mm SL. Abbreviationsi ABSP, abductor superficialis pelvicus; ADPP, adductor profun- dus pelvicus; ADSP, adduetor superficialis pelvicus; ARDP, arrector dorsalis pelvicus; EXP, extensor proprius. Arrow indicates connection of EXP and pectoral gird!e. Seale bars: 5 mm.

A Lip B

Chin

Fig. 16. Diagrammatic illustration of barbels on lower jaw, showing TS 21-1 (A) and 21-2 (B).

lateral rows of bony plates in the Hoplichthyidae.

Interrelationships within Peristediidae The characters derived from the above 23 transformation series were used in an analysis to infer the interrelationships of and within the family Peristediidae. Two equally parsimonious trees were obtained, The statistics of the trees are: consistency index O,625, rescaled consistency index O.559, and tree length 40. The characters supporting each clade in the strict consensus trees, including reversals, are shown in Fig. 19, The clade including all peristediids is supported by the fbur synapomor[phies TS 4-1, 8-1, 13-1, and 21-1, Each elade recognized within Peristediidae is described

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology

18 Toshio Kawai A

Fig. 17, Ventral aspect of posterior region of beny plates on lateral Iine. A, Paraheminodus mttrrayi, HUMZ 191555, 132,6 mm SL; B, Peristedion orientale, HUMZ 80336, 117.3 mm SI].

Ventral row

anterior B -

Fig. I8. Diagrammatic illustration ot' ventral surtace of caudal peduncle, showing TS 23-O (A) and 23-1(B).

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 19

Gargariscus prionocephaius

Satyrichthys rieheti

S. adeni Heminodusphifippinus Paraheminodusmunayt

Satyrdchthys gilberti

S. senulatus

S. orientafis

S. amiscus

Peristedion antiIlarum

P. gracile

P. thompsoni

P. bnevirostre

P. miniatum

R truncatum

R greyae

P. Iongispatha

P. catapht:actum

P. barbiger

P. ecuadorense

P. weberi

P. Iiorhynchus

P. niet:straszi

P. orientale 60r

Fig. 19. Stricl eonsensus tree ot' two equally parsimonious cladograms of Peristediidae. Num- ' bering of characters and states corresponds to that in the text. indicates autapomorphic `[r" character, Abbreviation indicates reversal.

below. Clade al. Gargariscus, Henzinodus, Paraheminodus, and Sa4yrichthys. This clade is supported by the three autapomorphies TS 7-1, 9-1, and 11-1. Clade a2. Peristedion, This clade is supperted by the three synapomorphies TS 16-1, 17-1, and 22-1, the first two of which are autapomorphic. Clade bl. Gargariscus, Satyrichthys adeni, and S. riqffbii. The single synapomorphy TS 6-1 supports this clade.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology

20 Toshio Kawai

Clade b2. Heminodus, Paraheminodus, Satyrichthys amiscus, S, gilberti, S. orientalis, and S, serrulatus. This clade is supported by fbur synapomorphies, viz., TS 2-1, 8-2, 12-1, and 14-1, Of these features, TS 2-1, 8-2, and 12-1 are autapomorphic.

Two different topologies are equally well supported in this clade: the first tree shows sister relationships between clade d3 and the clade including Heminoalus philippinus Smith, 1917 (species dl) and Paraheminodus murropi (GUnther, 1880) (species d2); and the second has a trichotomy among dl, d2, and d3. The clade including lfeminodus and Paraheminodus is supported by the single synapomorphy

TS 4-0r in the fbrmer topology. Species cl. Garzgariscus prionocqphalus (Dumeril, 1869). This taxon is supported by the two reversal character states TS 4-Or and 13-Or. Clade c2. Satyrichtbys adeni and S. rieXfeti. This clade is supported by the single synapomorphy TS 22-1, Species dl. Heminodus philippinus. This taxon is supported by the two synapomorphies TS 19-1 and 20-2. 0f these features, TS 20-2 is autapomorphic. Species d2. Paraheminodus murrqyt, This taxon has no derived charaeter states of its own, Clade d3. Satyrichtbys amiscus, S. gilberti, S. orientalis, and S. serrulatus, This c]ade is supported by the single autapomorphy TS 18-1, and it is further divided into two clades, the first including Satyrichthys gilberti and S. serruZatus, and the second including S. amiscus and S. orientalis. The former and latter clades are

each supported by a single character state, TS 22-1 and 6-1 respectively, Species el. Peristedion antillar"nz. No apomorphic character state has been identified for this species. Clade e2. Peristedion harbi.crer, P. brevirostre, P, catqphractum, R ecuadorense, P. gracile, P. greyae, P. Iiordynchus, P. Ion.uispatha, P. miniatum, P. nierstraszi, P. orientale, P. thompsoni, P. truncatum, and P. weberi. This elade is supported by the

single autapomorphy TS 21-2. Species fl. Peristedion gracite, There is no unique apomorphy for this species among the scored characters. Clade M. Peristedion barbiger, P. brevirostre, P. catophractum, P. ecuadorense, P. grqyae, P. Iiorhynchas, P. Iongispatha, P. miniatum, P. nierstraszi, P, orientale, P, thompsoni, P. truncat"m, and P. weberi. This clade is supported by the single

synapomorphy TS 6-1. Species gl, Peristedion thompsoni. There is no unique apomoxphy for this species among the scored characters. Clade g2. Peristedion barbiger, P. brevirostre, P. catqphractum, P. ecuadorense, P. grqyae, P. Iiorltynchus, P. Iongispatha, P. miniatum, P. nierstraszi, P. orientale, P, truncatum, and P, weberi, The single autapomerphy TS 10-1 supports this clade. Species hl. Peristedion brevirostre. There is no unique apomorphy for this species among the scered characters, Clade h2. Peristedion barbiger, P, catophractum, P. ecuadorense, P. grqJ;ae, P. Iiorbynchus, R longispatha, R mintatum, P, nierstraszi, P. orientale, R truncatum, and P, tveberi. This clade is supported by the single synapomorphy TS 15-1. Species il to i7. No unique apomorphic characters have been identified fbr any of these terminal taxa, Clade i8. Peristedion liorhynchus, P. nierstraszi, P. orientale, and P. weberi. This clade is supported by the single synapomorphy TS 14-1. Interrelationships

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systernatics 21 g･ opxG`eg8 g8-"o9tseg .8 eoeeq･"oq E E･ g E g it o ts e X Q cr>

-Genu$

Fig. 20. Phylogenetic relationships within Peristediidae and ranking at generic leve!,Num- bering of clades correspond to that in Fig. 19.

among the four species in this c]ade have not been clarified. Only P, ortentale is characterized by the single apomorphic character TS 6-0r,

Classification

Based on the inferred phylogenetic pattern, I give generic rank to the clades or species a2, cl, c2, dl, d2, and d3; this action minimizes the taxonomic changes at the genus level (Fig. 20). As a result, the following six genera are recognized within the fami]y: Gangariscus, Heminodus, Paraheminodus, Peristedion, Satyrichthys, and Scalicus.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology

22 Toshio Kawai

Genus Gargariscus Smith, 1917 [Japanese namei Oni-kihoubou-zoku] (Fig, 21A)

Gargariscus Smith,1917: 145. Gender: masculine.

Type species. Gaizgariscus semidentatus Smith, 1917 aunior synonym of Peris- tethidion prionocephatum Dumeril, 1869). Composition. Gat:gariscus prionocephalus (Dumeril, 1869), Diagnosis. Upper jaw teeth present; lateral margin of head extremely indented; posterior parts of lower lateral rows of bony plates separated from each other; alld barbels on lower jaw not branched exeept for posteriormost lip and chin barbels. Remarks. Previous authors (e.g,, Kamohara 1952a; Richards 1999; Shinohara et al. 2001) recognized Gargariscus as menotypic, including in it only G. priono- cqphal"s, the senior synonym of the type species of the genus, Gargariscus semidentatus.

Genus Heminodus Smith, 1917 [Japanese name: Ito-kihoubou-zoku] (Fig. 21B)

Heminodus Smith,1917:146. Gender: masculine.

Type species. Hentinodus philippinus Smith, 1917. Composition. Heminodus philmpintts Smith, 1917. Diagnosis. Upper jaw teeth present; lateral margin of head smooth; posterior parts of lower lateral rows of bony plates separated from each other; and one or two very short barbels present. Remarks. This genus is menotypic, including only the type species Hemi- nodus philippinus, Kawai and Nakaya (2007) showed that Heminodus jqponicus Kamohara, 1952 is ajunior synonym ofH philippinus.

Genus Paraheminodus Kamehara, 1957 rJapanese name: Koutou-kihoubou-zoku] (Fig, 21C)

Paraheminodus Kamohara,1957: 4, Gender: masculine.

Type species. Satyrichthys laticephalus Kamohara, 1952, Compesition. Paraheminodus kamoharai Kawai, Imamura, and Nakaya, 2004, P. Iaticephalus (Kamohara, 1952), and P. murrayi (GUnther, 1880), Diagnosis. Upper jaw teeth present; lateral margin of head smooth; posterior parts of lower lateral rows of bony plates separated from each other; barbels on lower jaw not branched except for posteriormost barbels. Remarks. The genus Paraheminodus, established by Kamohara (1957) for Satyrichthys laticophaZus (type species), is characterized by having a tooth band on

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristedijd systematics 23

¢ :- wr･'-s

ec'.l ,-."=

Fig. 21. Lateral aspect of peristediid species. A, Gangariscus prionocqphalus (=G. semidentatus; type species of Gargariscus) (after Richards 1999); B, Heminodus philippinus, type species of Heminodus (from Richards 1999); C, Paraheminodus laticephalus, type species of Para- heminodus (from Kamohara 1952b).

the upper jaw and well-developed barbels on the lower jaw. Ochiai and Yatou (1984) treated Paraheminodus as a junior synonym of Satyrichth",s, without comment. Since then, the status ef the genus has been the subject of disagreement. Shinohara et al. (2001) and Yamada (2002) fo11owed Ochiai and Yateu (1984), whereas other authors (e,g,, Miller 1974; del Cerro and Lloris 1997; Kawai et al. 2004a, b) recognized Paraheminodus as valid. The present analysis shows that Paraheminodus murrayi and Satyrichthys riqtfeli (the type speeies of SaCyrichtbys) belong to diflerent clades. Because no anatomical information is available for S. Iaticephalus

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

24 Toshio Kawai

relationship of S, laticephalus and P. kamoharai to P, murrayi, These include the presence of a tooth band on the upper jaw, contralaterally separated posterior parts of the lower lateral rows of bony plates, no branched barbels except fbr the posteriormost ones, well-developed posteriormost lip barbels, and a smooth lateral margin of the head. Consequently, S. Iaticephatus and P, kamoharai are linked to P. marrayi. The appropriate generic name for this group is thus Paraheminodus.

Genus Peristedion Lacapede, 1801 [Japanese name: Kihoubou-zoku] (Fig. 22A)

Peristedion Lacepede, 1801: 368, Gender: ncuter. Octonus Rafinesque, 1810: 29, 54 [type species: Octonus olosteon Rafinesque, 1810 Ounior synonym of Trigla cataphracta I,innaeus, 1758)]. Gender: masculine. Peristethidium Agassiz, 1846: 280 [unjustified emendation of Peristedion Lacepede, 1801; also incorrectly spelled as Peristethidion by several authors]. Gender: neuter. Peristethus Kaup, 1858: 336 [type species: Trigla catophracta Linnaeus, l758]. Gen- der: masculine. Polyacantichtbys Kaup, 1873: 82 [type species: Pertstedion orientale Temminck and Schlegel, 1843]. Gender: masculine. Peristedtum Jordan and Gilbert, 1883: 732 [unjustified emendation of Peristedion Lacenede, 1801]. Gender: neuter. VZitusiculus Jordan and Evermann, 1896: 489 [type species: lleristedion imberbe Poey, 1861]. Gender: masculine. Panichtdys Whitley, 1933: 97 [type species: Peristedion pict"ratum McCulloch, 1926 aunior synonym of Peristethus liorh"?nchus GUnther, ]872)], Gender: masculine.

Type species. Peristedion malarmat Lac6pede, 1801 Gunior synonym of TrigZa catophracta Linnaeus, 1758). Composition. Peristedion altipinnis Regan, 1903, P. amblygeays Fowler, 1938, P. antillarum Teague, 1961, P, barbtger (Garman, 1899), P. brevirostre (GUnther, 1860), P, catqphractunz (Linnaeus, 1758), P. crustosum (Garman, 1899), P, ecuadorense Teague, 1961, R gracile Goode and Bean, 1896, P. grayae Miller, 1967, P. imberbe Poey, 1961, P. Iiorbynchus (GUnther, 1872), P, longispatha (Goode and Bean, 1886), P. miniatum (GQode, 1880), P. nierstraszi Weber, 1913, P. orientate Tem- minck and Schlegel, l843, P. paucibarbiger Castro-Aguirre and Garcia-Dominguez, 1984, P. riversandersoni (Alcoek, 1894), R thompsoni Fowler, 1952, P, truncatum (GUnther, 1880), P. unicuspis Miller, 1967, and R weberi Smith, 1934, Diagnosis. Upper jaw teeth absent; lateral margin of head smooth; and posterior pairs of bony plates in lower lateral rows eontralaterally sutured along midline. Remarks. The genus Peristedion is the largest in the family, and 15 of its species were examined anatomica]]y during this study, AIthough materia] was not available for anatomical examination, I refor P. altipinnis, P. amblygempJs, P, crustosum, P, imberbe, P. paucibarhiger, R riversandeTsoni, and P. unicuspis to Periste-

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 25

f7.

Fig. 22. Lateral aspect ef peristediid species. A, Peristedion catqphractum (==P. malarnzat; type species of Peristedion) (firom Miller and Richards 1981); B, Satyrichthys riqfiizli, type species of Satyrichthys (from Richards 1999); C, Scalicus amiscus, type species of Scaticus (from Kamohara 1952a).

dion because all of them possess the autapomorphy of clade e2, i,e., all barbels are branched (TS 21-2), Peristedion has several junior synonyms. The type species of the genus, P, malarmat, as well as Octonus olosteon, type species of Octonus, are considerect to be junior synonyms of Trigla catqphracta (type species of Peristethus). Peristethidium and Peristedium are unjustified emendations of Peristedton (see Eschmeyer 1998), Possession of a rudimentary postrorse spine on the preopereule has been regarded as one of the diagnostic characters of Peristedion (e.g., Kamohara 1952a; Ya- mada 2002). This spine is fu11y developed, not rudimentary, in Peristedion brevirostre and P. mintatum, however, and such variability makes this character is un- reliable fbr recogriition of the genus.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

26 Toshio Kawai

Genus SatyrichthJ,s Kaup, 1873 [Japanese name: Iso-kihoubou-zoku] (Fig. 22B>

Salyrichthys Kaup, 1873: 82. Gender: masculine, Acanthostedion Fowler, 1943: 75 [type species: Acanthostedion rugosum Fowler, 1943]. Gender: neuter.

Type species. Peristethus rtqtX/eli Kaup, 1859, Composition. The following 10 species are assigned to Satyrichthys: S. adeni (Lloyd, 1907), S. clavilqpis Fowler, 1938, S. isokawae Yatou and Okamura, 1985, S. Iingi (Whitley, 1933), S, longiceps (Fowler, 1943), S, magnus Yatou, 1985, S. moluccensis (Bleeker, 1851), S. riqffeli (Kaup, 1859), S. rugostts (Fowler, 1943), and S. wetchi (Herre, 1925). Diagnosis. Upper jaw teeth absent; lateral margin of head smooth; posterior parts of lower ]ateral rows of bony plates separated from each other; barbels on lower jaw not branched except for posteriormost lip and chin barbels (some species without ehin barbels); number of dorsal fin soft rays fewer than 20; number of anal fin soft rays fewer than 19, Remarks. This study demonstrates that the previous concept of Satyrichthys is not monophyletic. Species that have been assigned to the genus comprise two clades, viz,, clades c2 and d3 (Fig. 19). Miller (1974) and Yamada (2002) pointed out that Satyrichtdys could be divided into two groups of species by the numbers of dorsal and anal fin soft rays, and these groups correspond to the two clades recognized in this study, In fact, all members of clade c2 are characterized by low numbers of dorsal and anal fin soft rays (at most 19 dorsal and 18 anal), whereas all members of clade d3 have high numbers (at least 20 in each fin) (Kamohara 1952a; Miller 1974; Yamada 2002), The polarity of this character was not assessed in the present study. No specimens of the fbllowing taxa currently assigned to Satyrichthys were available ibr anatomical examination: S. ctavilmpis, S. isokawae, S. Iingi, S. Iongiceps, S. magnus, S. moluccensis, S rugosus (the type species of Acanthostedion), and S. welchi. Thus, I could not definitely assign these species to clade c2 based on synapomorphic characters. However, these eight species are

linked to clade c2 by certain external features, including the absence of teeth on the upper jaw, the separated posterior parts of the lower lateral rows of bony plates on both sides, ne branched barbels except for the posteriormost ones, a smooth lateral margin of the head, and low numbers of dorsal and anal fin seft rays. In light of this, I also include these eight species in this clade.

Genus Scalicus Jordan, 1923 [Japanese name: Hige-kihoubou-zoku] (Fig. 22C)

Scaticus Jordan. 1923: 216. Gender: masculine. Aremoperistedion Fowler, 1938: 126 [type species: Nemoperistedion orientale Fowler, 1938]. Gender: neuter.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 27

Type species. Pertstedion amiscus Jordan and Starks, 1904. Composition. The fbllowing eight species are assigned to Scalicusi S, amiscus (Jordan and Starks, 1904), S. engyceros (GUnther, 1872), comb. nov., S. gilberti (Jor- dan, 1921), comb. nov., S. hians (Gilbert and Cramer, 1897), comb. nov., S. investigatoris (Alcock, 1898), comb. nov., S. orientalis (Fowler, 1938), comb. nov., S. q"adratorostratus (Fourmanoir and Rivaton, 1979), eomb. nov., and S. serrulatus (AIcock, 1898), comb. nov, Diagnosis. Upper jaw teeth absent; lateral margin of head smooth; posterior pairs of lower lateral rows of bony plates separated from each other; barbels en Iower jaw not branched except for posteriormost lip and chin barbels; and number of dorsal and anal fin soft rays greater than 19. Remarks. Jordan (1923) established the new genus Scaticus for its type speeies Peristedion amiscus. Subsequent authors (e.g,, Miller 1974; Yatou 1985; Yamada 2002) have referred this species to Satyrichthys, with the result that Scalicus has been placed under the synonymy of Satyrichthys, This study shows that R amiscus and three other species heretofore assigned to Satyrichtdys (e.g., Kamohara 1952a; Yamada 2002), i.e., S. gilberti, S. orientalis, and S. serrulatus, foum a clade not closely related to Satyrichthys (Fig. 19). This clade can be ranked as a full genus, and by priority Scalicus is the correct available name for it. As was mentioned before, Scalicus differs from Satyrichthys in having more dorsal fin soft rays (more than 19 vs. Iess than 20 in Satyrichthys) and anal fin seft rays (more than 19 vs. Iess than 19 in Satyrichtdys) (Kamohara 1952a; Miller 1974; Yamada 2002). Peristethus engyceros, Peristedion hians, Peristedion investigatoris, and Peristedion quadTatorostratum are also referable to Scalicus based on features of the external morphology, although ne specimens for anatomical examination

were available for these species. These features include the absence of a tooth band on the upper jaw, contralateral separation of the posterior parts of the lower lateral rows of bony plates, no branched barbels except fbr the posteriormost ones, a smooth lateral margin of the head, and high numbers of dorsal and anal fin rays. In total, the eight species mentioned above are recognized in the genus.

Key to the genera of the family Peristediidae 1. Upperjaw teeth present ...... ,...... 2 - Upperjawteethabsent...... ,...... 4 2. Lateral margin of head extremely indented...... Gangariscus - Lateral margin of head smooth (except injuveniles) ...... ,.....3 3. Longest barbel very short...... ,...... H2minodus - Longest barbel moderately long ...... ,....,.,..,.,Parahenzinodus 4. Posterior pairs of bony plates in lower lateral rows contralaterally sutured alongmidline...... Peristedion - Posterior parts of lower lateral rows of bony plates separated by ventral rows ofplates...... ,...... ,....,...... 5 5. Dorsal fin soft rays fewer than 20 ...... Satyrichthys

- Dorsal fin soft rays more than 19...... ,....Scalicus

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

28 Toshio Kawai

Acknowledgements

I am very gratefu1 to Prof. Kazuhiro Nakaya (HUMZ) for his guidance during this study and his critical reading of the manuscript. I also thank Prof. Mamoru Yabe (HUMZ) and Assoc, Prof, Hisashi Imamura (the Hokkaido University Mu- seum) for their critical reading of the manuscript and helpfuI advice. I sincerely thank Prof. Emeritus Kunio Amaoka (Hokkaido University) for his valuable advice on phylogeny and anatomieal morphology. My thanks also go to the follewing persons for loans or gifts of specimensi M. McGrouther (AMS); J. Maclaine (BMNH); Y. Machida, K. Sasaki, and H. Endo (BSKU); T. Mitsuhashi (Cisto Co., Ltd.); M. Sato (Hakodate Museum); M. Yoshi,da (Japan Deep Sea Trawlers Association); H. Motomura (Kagoshima University Mu- seum); G, Duhamel and P. Pruvost (MNHN); Y. Iwatsuki (MUFS); T. Kanda (Nobeoka Marine Science Station, Miyazaki University); K, Matsuura and G. Shi- nohara (NSMT); K. Yamamoto (Seikai National Fisheries Research Institute); C. R. Robins (UF); T, Yoshino (URM); J. T. Williams and S. J. Raredon (USNM); and E. Okuto and M. Yamane (Nagato, Yamaguchi, Japan). I express my thanks to M. Shi- mazaki (Earth Environmental Service Co., Ltd,), T, Takahashi (FAKU), R. Fujii (Gifu Prefectural Research Institute fbr Freshwater Fish and Aquatic Environ- ments), M. Takahashi (Hokkaido Federation of Fisheries Cooperative Associa- tions), Y. Choi (Kunsan National University), H. Miyahara (Kyowa Concrete Indus- try Co,, Ltd,), B,-J, Kim (National Institute of Biological Resources, Korea), F. Muto (National Research Institute of Far Sea Fisheries), K. Sato (Okinawa Churaumi Aquarium), K. Hoshino and N. Suzuki (Seikai Fisheries National Research Insti- tute), M, Aibara (Tokyo Institute of Technology), H. Matsubara (Tokyo University of Agriculture), and HUMZ colleagues for helpfu1 suggestion, and K. Suda and F. Tashiro (HUMZ), A. Williston (MCZ), Y. Takata (NSMT), and M. J. P. van Oijen (RMNH) for offering information of specimens. I also express my appreciation to Kochi University, the Food and Agriculture Organization of the United Nations (FAO), and the Ichthyological Society of Japan fbr permission to reproduce illus- trations. I sineerely thank T. Komai (Natural History Museum and Institute, Chiba) for his patient review of the manuscript and for offering many usefu1 com- ments. Finally, my special thanks go to my parents, Syun Kawai and Noriko Kawai, for their continued encouragement, This manuscript is a part of doetoral thesis submitted to the Graduate School of Fisheries Sciences, Hokkaido Univer- sity.

Appendix: List of Material Examined

Material used for dissection Peristediidae. Gargariscus prionocephatus: BSKU 17658, 1 specimen, 148.0mm SL. South China Sea (7u26.3'N, 109"14,9'E to 7e30.2'N, 109e13,2'E), 265-286m, 11 July 1972. IIeminodus philippinus: HUMZ 193268, 1 specimen, 174,5mm SL, ofi' Sumatra (sa47.0'S, 102e36.6'E te 5c48,1'S, 102'35.1'E), Indonesia, Indian Ocean, 386-387m, 22 July 2005. Paraheminodusnzurrayi.' BSKU 4599, 1 specimen, 139.3mm SIi,Mimase FishMarket, Kochi, Japan, 30 April 1955. Peristedion antillarum: HUMZ 187919, 1 specimen, 140.0mm SL, off St, Kitts and Nevis (17'15'N, 62022'W), Leeward Islands,

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 29

Caribbean Sea, 580m, 8 December 1969; HUMZ 187920, 1 specimen, 155.3mm SL, eff Honduras (16035'N, 80"10'W), Caribbean Sea, 576m, 18 May 1962. Peristedion barbiger: HUMZ 168029, 1 specimen, 170.8rnm SL, off Peru (3o38'S, 81o12'W to 3o4o'S, 81014'W), 235--256m, 10 August 1999; HUMZ 173577, 1 specimen, 122.2mm SL, off Peru (3e40'S, 81oOl'W to 3041'S, 810e2'W), 326-332m, 6 May 2000. Peristedion brevirostre: HUMZ 187909, 1 specimen, 101.5mm SL, off Cuba (23005'N, 78049'W), 411-457m, 7 November 1961; HUMZ 187910, 1 specimen, 107.8mm SL, off Cuba (23o04'N, 78e46rW), 362m, 15 December 1969. Peristedion catqphractum: HUMZ 187905, 1 specimen, 135.8 mm SL, off Gabon (1030.00'S, 8027.30'E), 201 m, 3 September 1963; MNHN 2006-1319, 1 specimen, 110,2 mm SL, Gulf of Guinea (OoOO', 5aOO'E), Peri- stedion ecuadorense: HUMZ 31502, 31513, 2 specimens, 133.8, 136.2mm SL, off Suri- name C6n57'N, 56035'W to 7044'N, 53041'W), 725m, 6 December 1973. Peristedion gracile: HUMZ 187914, 1 specimen, 95.5mm SL, otf FLorida (24023.21'N, 82o14,49'W), 84-124m, 24 May 1989; HUMZ 187924, 1 specimen, 134.0mm SL, off Tobago (11030'N, 60o46'W), Trinidad and Tobago, 366-439m, 22 September 1964. Peristedion greyae: HUMZ 187907, 1 specimen, 112.3mm SL, off Florida (24016.276'N, 82044.486'W), 393-400m, 10 May 2000; HUMZ 187908, 1 specimen, 100.4mm SL, off FIorida (24017.408'N, 82052,48'W), 360-366m, 12 May 1999. Peristedion liorh:vnchus: HUMZ 178284, 1 specimen, 144,3mm SL, collection data unknown. Peristeclion longispatha: HUMZ 187921, 1 specimen, 165.3mm SL, off Colombia (11010'N, 74027'W), Caribbean Sea, 457-512m, 17 May 1964; HUMZ 187922, l specimen, 160.9mm SL, off Panama (9o15'N, 81o32'W), Caribbean Sea, 457m, 25 May 1962. Peristedionminiatum: HUMZ 187912, 1 specimen, 110.5mm SL, off ELorida (24e19.318'N, 82059.066'W), 270-274m, 14 May 1999. Peristedion nierstraszi: HUMZ 35722, 1 specimen, 97.0mm SL, collection data unknown. Peristedion orientale: HUMZ 36744, 189794, 2 specimens, 125.4, 109.7mm SL, collection data unknown; HUMZ 177798, 1 specimen, 119.5mm SL,Mi- mase FishMarket, Kochi, Japan, 22 January 2001, Peristedion thompsoni: HUMZ 187917, 1 specimen, 122.4mm SL, off Florida (24018.58'N, 82e16,39'W), 21 April 1988; HUMZ 187918, 1 specimen, 127,Omm SL, off Florida (24019.32'N, 82o33.04'W), 181-183m, 8 May 1998, Peristedton truncatum: HUMZ 187915, 1 speeimen, 121,6mm SL, off Florida (24014,32'N, 82034.46'W), 545-549m, 8 May 1988; HUMZ 187916, 1 specimen, 126.5mm SL, off Florida (24014.43'N, 81028.07'W), 552-558m, 6 May 1998. Peri- stedion weberi: MNHN 2006-1320, 1 specimen, 125.5mm SL, Mozambique Channel (23a20'S, 43031'E), January 1986. Satyrichtdys adeni: HUMZ 81520, 1 specimen, 296.9mm SL, Saya de Malha Bank (10046'S, 61005'E), Indian Ocean, 125m, 10 Decem- ber 1978; HUMZ 90034, 1 specimen, 269.4rnm SL, Andaman Sea, 16 January 1971. Satyrtchtdys amiscus: HUMZ 51838, 51851, 2 specimens, 139.8, 147.5mm SL, off Owase, Mie, Japan, 350m, 13 March 1976. Satyrichtbys gilberti: HUMZ 99830, 99832, 2 specimens, l56.5, 161.0mm SL, Central Pacific (19057'N, 155"OO'W), 379m, 7 Febru- ary 1983, Satyrichthys orientalis: HUMZ 39019, 1 specimen, 81,2mm SL, Mimase Fish Market, Kechi, Japan, 2 March 1974; HUMZ 177601, 1 specimen, ca. 88mm SL, Mimase Fish Market, Kochi, Japan, 22 January 2001. Satyrichthys riqffeli: HUMZ 90522, 1 specimen, 145.8mm SL, East China Sea (31000'N, 127028'E), 8 April 1981; HUMZ 90539, 1 speeimen, 130.4mm SL, East China Sea (31COI'N, 126"53'E), 8 April 1981. Satyrichthys serrulatus: HUMZ 51714, 1 specimen, 135.2mm SL, off Omaezaki, Shizuoka, Japan, 200-350m, 10 Mareh 1976; HUMZ 80137, 1 specimen, 136.0mm SL, off Cape Ashizuri, Kochi, Japan, 240m, 30 November 1978. Triglidae. Bellator egretta (Goode and Bean, 1896): HUMZ 69393, 1 specimen, 81 mm SL (dissected by H.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

r]'oshio :IO Kawai

Imamura), off South Carolina (32"06'N, 79013'W), 8 November 1973. Chelidonichthvs spinosus (McClelland, 1844)i HUMZ 108541, 1 specimen, 200 mm SL (dissected by H. Imamura), East China Sea (30045.08'N, 127014.05'E), 106m, 21 November 1985; HUMZ 178722, 1 specimen, 226.2mm SL, shopping center at Hakodate, Hokkaido, Japan, 2001. Lapidotrigla alata (Houttuyn, 1782): HUMZ 178493, 1 specimen,

126.6mm SL, Irino Fishing Port, Kochi, Japan, 2 December 2001. Lepidotri.ola .auen-

theri Hilgendorf, 1879: HUMZ 188807, 1 specimen, 155.2nmi SL, East China Sea (32o26.6'N, 127e48.5'E to 32e25.8'N, 127"49.9'E), 152-157m, 11 February 2004. Lopi- dotrigla hime Matsubara and Hiyama, 1932: HUMZ 188810, 1 specimen, 130.1 mm SL, East China Sea (28'58,6'N, 126"27.0'E to 28056.9'N, 126026.4'E), 124-110m, 1 March 2004, Lepidotrigla kanagashira: HUMZ 188676, 1 specimen, 127.4mm SL, East China Sea (30e26,2'N, 127049.4'E to 30"27.8'N, 127D49.8'E), 271 m, 5 March 2004. Lepidotrigta kishinoayei Snyder, 1911: HUMZ 188867, 1 specimen, 101.2mm SL, East China Sea (28057.7'N, 124057,O'E to 28"59.3'N, 124057.0'E), 90-84m, 17 February 2004. Lqpidotrigla microptera Ganther, 1873: HUMZ 178031, 152.3mm SL, 1 specimen, shopping center at Hakodate, Hokkaido, Japan, 28 June 2001, Prionotus carolintts (Linnaeus, 1771): HUMZ 69398, 1 specimen, 145.6mm SL, off South Carolina (32o50'N, 78o58'W), 24 October 1973. Prionotus stearnsii Jordan and Swain, 1884: HUMZ 32456, 1 specimen, 98mm SL (dissected by H. Imamura), off Suriname (7011.07'N, 54058,03'W), 81m, 8 December 1973. Pter:ygotrigla hemisticta: HUMZ 184937, 1 specimen, 125.6mm SL, East China Sea (28044,9'N, 126e44.6'E to 28"46.3'N, 126045.9'E), 13 March 2003. Ptenygotri.uta macrordynchus Kamehara, 1936: HUMZ 190324, 1 specimen, 95mm SL (dissected by H, Imamura), collected data unknown. Pter:vgotrigla muitioceZlata (Matsubara, 1937): HUMZ 188880, 1 specimen, 231.4mm SL, East China Sea (27054,4'N, 126025.7'E to 27053.4'N, 126024.5'E), 265-266m, 27 Feb- ruary 20e4. Hoplichthyidae. Hbptichthys gilberti: HUMZ 51736, 1 specimen, 152mm SL (dissected by H. Imamura), otT Omaezaki, Shizuoka, Japan, 200-350m, 10 March 1976; HUMZ 178506, 1 specimen, 146.lmm SL, TQsa Bay, Japan, 150m, 5 December 2001. HOplichthys haswelli: HUMZ 50268, 1 specimen, 341mm SL (dissected by H. Imamura), off New Zealand (43003'S, 176027'W), 558m, 13 June 1975; HUMZ 91386, 1 specimen, 328.8 mm SL, off New Zealand (47026'S, 169015'E), 30 Janu- ary 1981. HOplichtdys langsdor:fii: HUMZ 75337, 1 specimen, 156 mm SL (dissected by H. Imamura), Mimase Fish Market, Kochi, Japan, 27 February 1978; HUMZ 177869, 1 specimen, 166.4 mm SL, Mimase Fish Market, Kochi, Japan, 15 January 2001.

Material of Peristediidae used for examination of external morphology Gargariscus prionocephalus: BSKU 17657, 1 specimen, 149.0mm SL, South China Sea (7026.3'N, 109014,9'E to 7a30.2'N, le9e13.2'E), 265-286m, 11 July 1972; BSKU 87460, 1 specimen, 20g,5mm SL, Mimase Fish Market, Koehi, Japan, 7 April

2000; MUFS 11756, 1 specimen, 248,8mm SL, collection data unknown; NSMT-P

54649, 2 specimens, 132.3, 166.9mm SL, Sanya Fish Harbor, south coast of Hainan, China, 4 March 1997. Ueminodtts philippinus: USNM 78250, holotype, 140,5mm SI., off Tawi-Tawi, Mindanao Sea, Philippines; BSKU 948, holotype of Heminodus joponicus, 74.8 mm SL, Mimase Fish Market, Kochi, Japan, 3 February 1951; HUMZ 193267, 1 speeimen, 163.8mm SL, off Sumatra, Indonesia, 22 July 2005; HUMZ 194104, 1 specimen, 132.5mm SL, off Sumatra (5045,7'S, 102032.1'E to 5e46.6'S, 102"33,3'E), Indonesia, 418-405m, 31 May 2005. Paraheminodus kamoharai: BSKU 15243, holotype, 106.9mm SL, Sulu Sea (8Ull,8'N, 117058.0'E), Philippines, 285m, 26

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 31

May 1972; BSKU 15242, 15244, paratypes, 2 specimens, 107.3, 114.8mm SL, same data as holotype. Paraheminodus laticqphalus: BSKU 1612, holotype, 111.6mm SL, Mi- mase Fish Market, Kochi, Japan, 24 January 1952. Paraheminodus murrayi: BMNH 1879.5.14.265, holotype, 154.3mm SL, Banda Sea, Indonesia; BSKU 1566, 1 paratype of Paraheminodus kochiensis Kamohara, 1957, 109.7mm SL, Mimase Fish Market, Kochi, Japan, 20 January 1952; BSKU 1610, 1611, 2 paratypes of P. kochiensis, 117.4, 124.9mm SL, Mimase Fish Market, Kochi, Japan, 24 January 1952; BSKU 4598,

4599, 2 paratypes of P. kochiensis, 101.0, 139,3mm SL, Mimase Fish Market, Kochi, Japan, 30 April 1955; BSKU 7265, holotype of P. kochiensis, 138,3mm SL, Mimase Fish Market, Kochi, Japan, 13 May 1957; BSKU 7398, 7399, 2 paratypes of P, kochiensis, 75,3, 83.9mm SL, Mimase Fish Market, Kochi, Japan, 16 January 1951; BSKU

12986, 1 specimen, 154.6mm SL, collection data unknown; BSKU 40ee8, 1 specimen,

90.lmm SL, Mimase Fish Market, Koehi, Japan, 16 April 1984; HUMZ 191555, 1 specimen, 132.6mm SL, off Sumatra (5046.52'S, 102e41.25'E to 5046.02'S, 102e39.53'E), Indonesia, 639-546m, 26 September 2004; MUFS 4578, 1 specimen, 117.8mm SL, To- toro, Miyazaki, Japan, 24 February 1976. Peristedion amblygeays: USNM 9g870, holotype, 132.9mm SL, off San Fernando Point Light (16"30.36'N, 120:11,06'E), west coast of Luzon, Philippines, 82m, 10 May 1909. Peristedion barbiger: HUMZ 167926, 167927, 2 specimens, 145.4, 138.5mm SL, off Peru (4o04'S, 81o]O'W to 3e3s'S, soosgrW), 300-316m, 18 August 1999; HUMZ 180062, 180063, 180066, 3 specimens, 168.8-175.4mm SL off Peru (5G18'S, 81018'W), September 2001; HUMZ 185666, 1 spcc- imen, 155,6mm SL, off Peru (3058'S, 81011'W to 3058'S, 81010rW>, 456-421m, 20 Sep- tember 2002; HUMZ 185991, 1 specimen, 78.2mm SL, off Peru, 2002; HUMZ 189054, 189055, 2 specimens, 161.6, 186.2 mm SL, off Peru (3o49'S, 81aOO'W to 3o48'S, 80o59'W), 157-155m, 18 October 2003; HUMZ 189168, 1 specimen, 143.2mm SL, off Peru (5C18'S, 81e20'W to 5e16'S, 81e20'W), 134-138m, 16 October 2003. Peristedion liordynchus: BMNH 1871.7.20.53, holotype, 188.3mm SL, Celebes Sea, Manado, Indonesia; AMS E.4628, holotype of Peristedion picturatum McCulloch, 1926, 127.5mm SL, east of Flinders (39057'S, 148020'E), Australia, 182m, 4 December l913; AMS L19205-OO2, 3 specimens, 138.3-l61.3mm SL, east of Broken Bay (33033'S, 151"59'E), Australia, 384m, 26 May 1976; AMS I.20435-O05, 2 specimens, 131.0, 135.0mm SL, off North Soli- tary Island (29047'S, 153e44'E), Australia, 438m, 2 August 1978; AMS i.22807-028, 3 specimens, 173.8-183.lmm SL, 175km north of Port Hedland (18G32'S, 118o17'E), Australia, 204m, 2 April 1982; AMS I.39994-OOI, 6 specimens, 130.6-150.3mm SL, off Bermagui (36044'S, 150C20'E), Australia, 375m, 29 February 2000; BSKU 27393, 1 specimen, 222.6mm SL, Okinawa Trough (27000,2'N, 125046.0'E), East China Sea, 302-309m, 3 March 1978; BSKU 32664, 1 specimen, 208.0mm SL, Okinawa Trough (2scso.O'N, 124020,O'E), East China Sea, 250-290 m, 20 September 1979; HUMZ 71452, 1

specimen, 158.8mm SL, Port Onahama, Iwaki, Fukushima, Japan, 13 November

1977; HUMZ 79424, 1 specimen, 199.3mm SL, off Cape Ashizuri, Kechi, Japan, 240m, 30 November 1978. Peristedion orientale: HUMZ 51710, 51711, 51713, 3 specimens, 121.8-150.0mm SL, off Omaezaki, Shizuoka, Japan, 200-350m, 10 March 1976;

HUMZ 80336, 1 specimen, 117.3mm SL, Mimase Fish Market, Kochi, Japan, 15-17

December 1978; HUMZ 171263, 1 specimen, 136.0mm SL, off Daikei, Taiwan, 2 May 2000; HUMZ 177144, 1 specimen, 115.5mm SL, off Iwaki (37"OO'N, 141u18'E to 37o02'N, 141018'E), Fukushima, Japan, 148m, 25 October 2000; HUMZ 181718-181724, 7 specimens, 60,7-87.3mm SL, off Hitachi (36032'N, 140C59'E to 36030'N, 140S57'E), Ibaraki, Japan, 144-161m, 23 October 2001; HUMZ 182809, 1 specimen, 94.7mm SL, off Hi-

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

32 Toshio Kawai

tachi (36023'N, 140a53'E to 36022'N, 140u52'E), Ibaraki, Japan; HUMZ 185189, 185191, 185193-185197, 7 specimens, 118.2-135,8mm SL, Donggang Fishing Port, Pingtung County, Taiwan, 28 August 2002; MUFS 8568, 20929, 20931, 20932, 4 specimens, 119.5-131.2mm SL, off Miyazaki, Japan, 5 October 1987. Satyrichtdys amiscus:

USNM 51428, holotype, 79,7mm SL, off Manazuru Poim, Sagami Bay, Japan, 279m, 5 May 1900; HUMZ 178436, 1 specimen, 72.5mm SL, off Kuji (40n09'N, 142o12'E), Iwate, Japan, 243m, 28 October 2001; HUMZ 178678, 1 specimen, 68.7mm SL, off Iwaki (37eOO'N, 141o27'E to 37002'N, 141029'E), Fukushima, Japan, 237-252m, 20 Oc- tober 2001; HUMZ 181703, 1 specimen, 73,Omm SL, off Hitachi (36028'N, 140a58'E to 36031'N, 141eOO'E), Ibaraki, Japan, 232-257rn, 23 October 2001. Satyrichtdys clavilqpis: USNM 98868, holotype, 200,9mm SL, eff Sombrero Island (13048,45'N, 120e41.51'E), Balayan Bay and Verde Island Passage, Philippines, 290m, 21 January 1908. Satyrichthys gilberti: USNM 84102, holotype, 111.1mm SL, off lava flow from Mauna Loa, Alika, Kau Dist,, Hawaii, 386m, 27 March 1902; HUMZ 99829, 99831, 99833, 3 specimens, 172,1-185.0mm SL, Central Pacific (19=57'N, 155uOO'W), 379m, 7 February 1983. Satyrichthys hians: USNM 47730, lectotype, 154.7mm SL, Hawaii (21012'N, 157049'W), 539m, 4 December 1891. Satyrichtdys isokawae: BSKU 29614, holotype, ca. 290mm SL, Okinawa Trough (25045'N, 124004'E), 220m, 21 September 1979. Satyrichtnj,s tingi: AMS IA 5690, holotype, 248.5mm SL, South Head of Sydney Harbor (33051'S, 151"21'E), Australia, 5 April 1933. Satyrichtbys tongicqps: USNM 99516, holotype, 155.0mm SL, off Tayabas Light (13C46.45'N, 121o35.08'E), Marinduque Island and vicinity, Philippines, 347m, 2 March 1909, Satyrichthys magnus: BSKU 26638, holotype, 486.7mm SL, Okinawa Trough (26o32'N, 125e04'E), 207-215m, 25 January 1978, Satyrichtdys orientalis: USNM 98876, holotype, 146.lmm SL, otT- Makyan (OO16.30'N, 127030.00'E), Indonesia, 497m, 29 November 1909, Satyrichtdys riqffeli: HUMZ 94895, 1 specimen, 151.0mm SL, East China Sea (31"OO'N, l27028'E), 122m, 8 April 1982; HUMZ 109576, 1 specimen, 231.1mm SL, off Sarawak, Borneo, South China Sea. Sa4yrichtdys rugos"s: USNM 99502, paratype, 1 specimen (one of 7 paratypes), 44,3mm SL, off Malabrigo Light (13"27.20'N, 121017.45'E), east coast of Mindoro, Philippines, 197m, 2 February 1908. Satyrichtin]s welchi: NSMT-P 54651, 1 specimen, 165.lmm SL, Sanya Fish Harbor,

south coast of Hainan, China, 4 March 1997,

References

Agassiz, L, 1846. IVbmenclatorts Zbologict index [J)ziversatis. Sumtibus et typis Jent et

Gassmann, Soloduri, 393 pp. de! Cerro, L. and I.loris, D. 1997. Gurnard fishes (Scorpaenlibrmes, Triglidae) from off New Caledonia, with description of five new species. in: Seret, B, (Ed.) Resuttats des Cam- pagnes MusORS7'OM. Vol, 17. Memoires du Museum National d'Histoire Naturelle 174: 91-124, Eschmeyer, W. N. 1998. C)titalQg ofFishes. Vols 1-3. California Academy of Science, San Fran- clsco, 2905 pp.

Fowler, H. W. 1938. Descriptions of new fishes obtained by the United States Bureau of Fish-

"Albatross", eries steamer ehiefly in Philippine seas and adjacent waters. Proceedings of

the United States National Museum 85: 31-135.

Fowler, H. W. 1943. Contributions to the biology of the Philippjne Archipelago and adjacent

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Peristediid systematics 33

regions. Descriptions and figures of new fishes obtained in Philippine seas and adjacent

"Albatross", waters by the United States Bureau of Fisheries steamer United States Na-

tional Museum Bulletin 100: i-iii+ 53-91.

Gill, T, l888, On the classification of the mail-cheeked fishes. Proceedings of the United States

National Museum 11: 567-592.

Gree,nwood, P, H,, Rosen, D. E., Weitzman, S. H. and Myers, G. S. 1966. Phyletic studies of

teleostean fishes, with a provisional classification of living forms. Bulletin of the Ameri- can Museum of Natural History 131: 339ul56.

Hubbs, C, I]. and Lag]er, K. F. 1958. Fishes of the Great Lakes region. Bulletin of the Cran- -213, brook Institute of Science 26: 1 pls 1-44. Imamura, H, 1996. Phylogeny of the family Platycephalidae and related taxa (Piscesi Scor- paenifbrmes). Speeies Diversity 1: 123-233.

Imamura, H. 2004. Phylogenetic relatienships and new classification of the superfamily Scor- paenoidea (Actinopterygii: Perciformes). Species Diversity 9: 1-36. Jordan, D. S. 1923. A classification of fishes inc]uding families and genera as far as known. Stanford University Publications, University Series, Biological Sciences 3: 77-243+i-x (index), Jordan, D. S. and Eveumann, B. W. 1896. A check-list of the fishes and fish-like vertebrates of

North and Middle Amerjca. Report of Commissioner of Fish and Fisheries 2]: 207-584.

Jordan, D. S. and Gilbert, C. H. 1883. Synopsis of the fishes of North America. Bulletin of the

United States National Museum 16i i liv+1-･101S.

Kamohara, T. 1952a. Studies on the family Peristediidae found in Japan. Japanese Journal of

Ichthyology 2: 1-13. [In Japanese with English summary] Kamohara, T, 1952b, Additions to the offshore bottom fishes of Prov. Tosa, Japan, with de-

scriptions of two new species, Research Reports of the Kochi University 1: 1-3.

Kamohara, T. 1957, Notes on twenty additions to marine fish fauna of Prov. Tosa, Japan, including one new genus (family Peristediidae). Researeh Reports of the Kochi University

6; 1-6,

Kaup, J. J. 1858, Einiges Uber die Acanthopterygiens a joue cuirassee Cuv. Archiv fUr Naturgeschichte 24: 329-343,

Kaup, J. J, IS73. Ueber die Familie Triglidae nebst einigen Worten Uber die CIassification.

Archiv fUr Naturgeschichte 39: 71-93. Kawai, T,, Imamura, H. and Nakaya, K. 2004a. Paraheminodu.s kochiensis Kamohara, 1957 (Teleostei: Peristediidae), a juntor synonym of Paraheminodus murrayi (Ganther, 1880), with a comparison of Iikeraheminodus murrqJ,i and Paraheminodus laticephalus (Kamo- hara, 1952). Ichthyological Research 51: 73--76,

Kawai, T., Imamura, H. and Nakaya, K, 2004b, A new species of armored sea robin, Para- heminodus kamoharai (Teleostei: Peristediidae), from the Philippines. Ichthyological Re-

search 51: 12e130.

Kawai, T. and Nakaya, K. 2007. Redescription ef a r'are armored gurnard, Hlaminod"s philippinus (Actinopterygii: Teleostei: Peristediidae). Species Diversity 12: 167-173. Lacepede, B. G. E. 1801. Histoire IVictturelle des Poissons, Vol. 3. Prasson, Paris, lxvi+558 pp., 34 pls.

Leviton, A. E., Gibbs, R. H. Jr., Heal, E. and Dawson, C. E. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic eodes for institutional resource collections in herpetology and ichthyology, Copeia 1985: 802B32. Matsubara, K. 1943. Studies on the seorpaenoid fishes of Japan. Anatomy, phylogeny and tax- onomy (I). Transactions of the Sigenkagaku Kenkyusyo 1: 1-170.

NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

34 Toshio Kawai

Miller, G. C. 1974. Fische des Indischen Ozeans. Ergebnisse der ichthy･ologischen Unter- "Meteor]' suchungen wtihrend der Expedition des Forschungsschiffes in den Indischen Ozean, Oktober 1964 bis Mai 1965, A. Systematischer Teil, XIV, Scorpaenifbrmes (2) Fam- ily Peristediidae. Meteor Forschungsergebnisse Reihe D 18: 61-72,

Miller, G. C. and Richards, W. J. 1981. Peristediidae. in: Fischer, W., Bianchi, G. and Scott, W, B. (Eds) IF:AO Species Identijication Sheets for Fishet:], Purposes, Eastem Central Atlantic; Fishitzg Area 34 and Part of4Z Vol. 3, FAO, Reme, Nelson, J. S. 1976. Fishes of the Wbrld, John Wiley & Sons, New York, xiii+416 pp, Nelson, J, S, 1984, Ftshes of the Idortd. Second Edition, John Wiley & Sons, New York, xv+523 pp.Nelson, J. S. 2006. Fishes of the PVbrld, Fburth Edition. John Wiley & Sons, New York, xix+601 pp.Oehiai, A. and Yatou, T. 1984. Peristediidae, Pp. 334-336, pls 301-302, 364. in: Masuda, H,,

Arnaoka, K., Araga, C,, Uyeno, T. and Yoshino, T. (Eds) 7Vie Fishes of the tlapanese Archi- pegago. Tokai University Press, Tokyo.

Rafinesque, C. S. 1810, frzdice d7ttiotogia Siciliana. Ossia, catatogo Metodico dei Nbmi Lattni,

itatiani, e Siciliani dei Pesci, che si Rinvengono in Sicilia. Presso Giovanni del Nobolo, Messina, 70 pp., 2 pls. Richards, W. J. 1999. Triglidae. Pp. 2359-2382, In: Carpenter, K. E. and Niem, V. H, (Eds) E40 Eipecies ldentijication Guide for Fisheny Purposes. The Living A4arine Resources of the VVestern Central Pacijic. Vbl. 4. Boay Fishes Part 2 (7L4ugilidae to C]txrangidae). FAO, Rome.

Shinohara, G., Endo, H,, Matsuura, K., Machida, Y. and Honda, H. 2001. Annotated checklist

of the deepwater fishes frorn Tosa Bay, Japan. National Science Museum Monographs 20: 283-343.

Smith, H. M. 1917. New genera of deep-water gurnards (Peristediidae) from the Philippine Is- lands. Proceedings of the Biological Society of Washington 30: 145-146.

Smith W, L. and Wheeler, W. C, 2e04. Polyphyly of the mail-cheeked fishes (Teleostei: Scor- paenifbrmes): evidence t'rom mitochondrial and nuclear sequence data. Molecular Phylo- genetics and Evolution 32: 627-646, Swoffbrd, D, L. 2002, PAUP*. Phylogenetic Analysis [lsing Parsimony ('and Other Methodt9. Version 4.0. Sinauer Associates, Sunderland, Massachusetts.

Teague, G. W. 1961. The armored sea-robins of America, a revision of the American species of the t'amily Peristediidae, Anales del Museo de Historia Natural, Montevideo 7: 1-27.

Washington, B. B., Eschmeyer, W, N. and Howe, K, M, 1984. Scoi[paenifbrTnes: relationships.

Pp. 438-447. I}i: Moser, H. G., Richards, W. J., Cohen, D. M., Fahay, M, P,, Kendall, A. W. Jr, and Richardson, S. I,. (Eds) Ontogeay and Svstematics ofFishes. American Society ot' Ichthyologists and Herpetologists, I]awrence, Kansas,

Whitley, G, P. 1933, Studies in ichthyology, No. 7. Records of the Australian Museum 19:

60-112, pls 11-15.

Winterbottom, R. 1974, A deseriptive synonymy of the striated muscles of the Teleostei. Pro-

ceedings of the Academy of Natural Sciences of Philadelphia I25: 225 317.

Yamada, U. 2002. Peristediidae, Pp, 610-613, 1523. in: Nakabe, T, (Ed,) Fishes of .lapan with Pictorial Ifeys to the Species, English Edition, Tokai University Press, Tokyo,

Yatou, T. 1985. Satyrichtbys amtscus (Jordan et Starks). Pp. 5S4, 585, 726. In: Okamura, O, (Ed,) Fishes ofthe Okinawa Trough and the A(Vacent ivaters. Vol. 2. Japan Fisheries Resources Conservation Association, Tokyo,

NII-Electronic Library Service