Phylogenetic Systematics (Teleostei

Japanese Society of Systematic Zoology

NII-Electronic Library Service

JapaneseSociety ofSystematic Zoology

SpeciesDiversity,2008, 13, 1･ 34

Phylogenetic Systematics of the Family

(Teleostei: Actinopterygii)

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 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 (Miller1974; 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




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 themonophyly 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 thePeristediidae are a monophyletie group, but that the sister group is the familyTriglidae, 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 hoplichthyidspecies (see Appendix), and used primarily these 40 species fbr the phylogeneticanalysis.

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-




Peristediid systematics 3

nized in this study based on the available information, although the assignmentshould 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, the13 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 themonophyly 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

was treated as "unordered".

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 characteroptimization, 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 isundeterminable (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). Thefifth inhraorbital and hyomandibular are clearly sutured in Gailgariscus, Periste-

dion, Satyrichth{vs adeni, S. riofeli, Triglidae, and Hoplichthyidae (TS 2-O), whereasthey 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).



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Peristediid svstematics 5

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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, infiraor-

bital; LET, lateral ethmoid; NA, nasal; OP, opercle; PA, parietal; POP, preopercle; PT, post-

temporal; 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




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Fig. 3. Lateral aspect of neurocranium. A, Garlgariscus prionocephatus, BSKU 17658, 148.0mmSL; 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 areprcsent 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: con-

nected via long Iigament posteriorly) (Fig. 5). The interopercle is directly attachedto the epihyal posteromedially in Peristedion and Triglidae (TS 5-O), but thesebones are connected via a long ligament posteriorly in Gargariscus, Heminodus,Paraheminodus, Satyrichtbys, and Hoplichthyidae (TS 5-1).




Peristeditd systematics 7

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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 (Oinot extending to ventral margin of ceratohyal; 1: extending to ventrai margin of

ceratohyal) <Fig. 5), The ventral region of the cartilaginous band between the cera-

tohyal and the epihyal does not extend to the ventral margin of the ceratohyal in

Heminodus, Parahenzinodus, Peristedion antillarum Teague, 1961, P. gracile Goode

and Bean, 1896, P. orientale Temminck and Schlegel, 1843, Satyrichtdys gilberti, S.



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4599, 139.3mm SI,; D, Peristedion orientale, HUMZ 36744, 125,4mm SL:

amiscus, HUMZ 51851, 147.5mm SL: F, SaCyrichtttys ri

viations: BR, branchiostegal;hyal; IOP, tnteropercle; VHH, ventral hypohyal. Scale bars: 5mm.

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qtfizti, HUMZ 90522, 145.8mm SL. Abbre-CH, ceratohyal; DHH, dorsal hypohyal; EH, epihyal; IH, inter-

serrulatus, and Hoplichthyidae (TS 6-O), whereas this region extend to the ventralmargin of the ceratohyal in GaTErariscus, the other examined species of Peristedionand Satyrichtiays, and Triglidae (TS 6-1). TS Z Shape of third basibranchial (O: slender; 1: robust) (Fig. 6). The third basi-branchial is slender in Peristedion, Triglidae, and Hoplichthyidae (TS 7-O), but ro-

bust 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 inthe Triglidae and Hoplichthyidae (TS 8-O); both bones are connected via an interar-cual 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;




Peristediid systematies 9

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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, HUMZ51838, 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 thecoracoid 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 onthe 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 presenton 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-




10 Toshio Kawai

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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 fusedwith 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 <Garman, 1899), R brevirostre

(GUnther, 1860), P. catqphractum (Linnaeus, 1758), P, ecuadorense Teague, 1961, P.gracile, P. greyae Miller, 1967, P, longispatha (Goode and Bean, 1886), P. miniatum

(Goode, 1880), P. thonzpsoni, P, truncatunz (GUnther, 1880), Satyrichthys adeni, S. ri-

ql[feli, and Triglidae (TS 14-O), whereas both bones are fused in Henzinodus, Para-heminodus, Peristedion liorbynchus (GUnther, 1872), P, nierstraszi Weber, 1913, P.orientale, P. weberi Smith, 1934, Satyrichtbys amiscus, S, githerti, S. orientalis,




Peristediid systernatics 11

COR

D

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




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. Sealebars: 5mm.




Peristediidi systematics 13

A

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.

P

B

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

(Fig. 11). The third preural centrum and its hemal spine

gariscus, Peristedion antillarum, P. brevirostre, P.

(O: autogenous; 1: fused)

are autogenous in Gar-gracile, P. thompsoni,




14 Toshio Kawai

LAP

Flg. 12, Lateral aspect of cheek and cephalic muscles. A, Heminodus phitmpinus, HumZ193268, 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 kanagashiraKamohqra, 1936 and Ptenygotrigla henzisticta (Temminck and Schlegel, 1843) (TS 15-O), whereas both bones are fused in Heminodus, Paraheminodus, the other exam-

ined 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 nar-

row) (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




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, adductormandibulae; 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 thechin 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 andfourth epibranchials; 1: connected to the third and fourth epibranchials by a longtendon) (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 thismuscle 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 pec-toral girdle) (Fig. 15). The extensor proprius is attaehed to the pecteral girdle inGargariscus, Paraheminodus, Peristedion, Satyrichthys, Triglidae, Hbplichthys

giZberti Jordan and Richardson, 1908, and Il Zangsdoittii Cuvier, 1829 (TS 19-O), butnot 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 verte-bra 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, notbranched 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 inPeristedion 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,




l6 Toshio Kawai

A

TDP

OBD

so

BP

EP

B

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 ven-

tral plate rows; 1: sutured contralaterally along midline) (Fig. 18). The more poste-rior pairs of bony plates in the lower lateral rows are separated in Garlgariscus,Heminodus, Paraheminodus, and Satyrichtdys (TS 23-O), but sutured along the mid-

line in Peristedion. The Triglidae and Hoplichthyidae are coded as? fbr this char-acter, because there are no bony plates on the body in the Triglidae, and no lower





AD

SP

B

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: con-

sistency index O,625, rescaled consistency index O.559, and tree length 40. The char-

acters 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




18

A

Toshio Kawai

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

Ventral row

B anterior-

N

!

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




Peristediid systematics

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

19

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 autapomorphiccharacter, Abbreviation

`[r" indicates reversal.

below.

Clade al. Gargariscus, Henzinodus, Paraheminodus, and Sa4yrichthys. Thisclade 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 synapo-

morphy TS 6-1 supports this clade.




20 Toshio Kawai

Clade b2. Heminodus, Paraheminodus, Satyrichthys amiscus, S, gilberti, S. ori-

entalis, 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 treeshows 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 in-cluding 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 sup-

ported 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 sin-

gle 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 charaeterstates 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 thesecond 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 beenidentified 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 thesingle 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 singlesynapomorphy TS 6-1.

Species gl, Peristedion thompsoni. There is no unique apomoxphy for thisspecies 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




g8-"o9tse

cl

opxG`eg8

Peristediid systernatics

g･g .8

E E･ gE g ito ts eX Q cr>

eoeeq･"oq

Fig. 20. Phylogenetic relationships within

bering of clades correspond to that in Fig.

-Genu$

Peristediidae and ranking at generic leve!,

19.

21

Num-

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.




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; alldbarbels on lower jaw not branched exeept for posteriormost lip and chin barbels.

Remarks. Previous authors (e.g,, Kamohara 1952a; Richards 1999; Shinoharaet 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 semi-dentatus.

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; posteriorparts of lower lateral rows of bony plates separated from each other; and one ortwo very short barbels present. Remarks. This genus is menotypic, including only the type species Hemi-nodus philippinus, Kawai and Nakaya (2007) showed that Heminodus jqponicusKamohara, 1952 is ajunior synonym ofH philippinus.

Genus Paraheminodus Kamehara, 1957rJapanese 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; posteriorparts 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) forSatyrichthys laticophaZus (type species), is characterized by having a tooth band on




Peristedijd systematics 23

¢ :- wr･'-s

ec'.l ,-."=

.

Fig. 21. Lateral aspect of peristediid species. A, Gangariscus prionocqphalus (=G. semidenta-tus; 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 au-

thors (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 <the type species of Para-heminod"s) and P, kamoharai, definite determinatien of the phylogenetic positionof these species is not easy. Nevertheless, some external characters suggest a close




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 posteriorparts of the lower lateral rows of bony plates, no branched barbels except fbr theposteriormost ones, well-developed posteriormost lip barbels, and a smooth lateralmargin of the head. Consequently, S. Iaticephatus and P, kamoharai are linked toP. 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: mascu-

line.

Type species. Peristedion malarmat Lac6pede, 1801 Gunior synonym of TrigZacatophracta 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 poste-rior pairs of bony plates in lower lateral rows eontralaterally sutured along mid-

line.

Remarks. The genus Peristedion is the largest in the family, and 15 of itsspecies were examined anatomica]]y during this study, AIthough materia] was not

available for anatomical examination, I refor P. altipinnis, P. amblygempJs, P, crus-tosum, P, imberbe, P. paucibarhiger, R riversandeTsoni, and P. unicuspis to Periste-





A

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 arebranched (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 re-

garded as one of the diagnostic characters of Peristedion (e.g., Kamohara 1952a; Ya-

mada 2002). This spine is fu11y developed, not rudimentary, in Peristedion brevi-rostre and P. mintatum, however, and such variability makes this character is un-

reliable fbr recogriition of the genus.




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. moluc-censis (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; posteriorparts of lower ]ateral rows of bony plates separated from each other; barbels on

lower jaw not branched except for posteriormost lip and chin barbels (somespecies without ehin barbels); number of dorsal fin soft rays fewer than 20; numberof anal fin soft rays fewer than 19,

Remarks. This study demonstrates that the previous concept of Satyrichthysis not monophyletic. Species that have been assigned to the genus comprise twoclades, 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 recog-

nized in this study, In fact, all members of clade c2 are characterized by low num-

bers 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 toSatyrichthys 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 toclade c2 based on synapomorphic characters. However, these eight species are

linked to clade c2 by certain external features, including the absence of teeth onthe upper jaw, the separated posterior parts of the lower lateral rows of bony plateson both sides, ne branched barbels except for the posteriormost ones, a smooth lat-eral margin of the head, and low numbers of dorsal and anal fin seft rays. In lightof 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.





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. investi-gatoris (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; posteriorpairs 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; Yamada2002) have referred this species to Satyrichthys, with the result that Scalicus hasbeen 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 seftrays (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 featuresof 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 lat-

eral 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




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 ad-

vice 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 ChuraumiAquarium), K. Hoshino and N. Suzuki (Seikai Fisheries National Research Insti-

tute), M, Aibara (Tokyo Institute of Technology), H. Matsubara (Tokyo Universityof 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 toKochi 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,





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 barbi-

ger: 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 brevi-rostre: 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), CaribbeanSea, 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, collectiondata 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 spec-

imen, 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, MimaseFish 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.




:IOr]'oshio

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, EastChina 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: HUMZ190324, 1 specimen, 95mm SL (dissected by H, Imamura), collected data unknown.Pter:vgotrigla muitioceZlata (Matsubara, 1937): HUMZ 188880, 1 specimen, 231.4mmSL, 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 (dis-sected 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 byH. 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, SouthChina 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 April2000; MUFS 11756, 1 specimen, 248,8mm SL, collection data unknown; NSMT-P54649, 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; HUMZ194104, 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: BSKU15243, holotype, 106.9mm SL, Sulu Sea (8Ull,8'N, 117058.0'E), Philippines, 285m, 26





May 1972; BSKU 15242, 15244, paratypes, 2 specimens, 107.3, 114.8mm SL, same dataas holotype. Paraheminodus laticqphalus: BSKU 1612, holotype, 111.6mm SL, Mi-mase Fish Market, Kochi, Japan, 24 January 1952. Paraheminodus murrayi: BMNH1879.5.14.265, holotype, 154.3mm SL, Banda Sea, Indonesia; BSKU 1566, 1 paratypeof 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, MimaseFish Market, Kochi, Japan, 13 May 1957; BSKU 7398, 7399, 2 paratypes of P, kochien-sis, 75,3, 83.9mm SL, Mimase Fish Market, Kochi, Japan, 16 January 1951; BSKU12986, 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), westcoast 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; AMSE.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, offBermagui (36044'S, 150C20'E), Australia, 375m, 29 February 2000; BSKU 27393, 1specimen, 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, 1specimen, 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, 15-17December 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 speci-

mens, 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-




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, PingtungCounty, 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 clavi-

lqpis: 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 fromMauna 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: USNM99516, 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 November1909, 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, offSarawak, Borneo, South China Sea. Sa4yrichtdys rugos"s: USNM 99502, paratype, 1specimen (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,

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