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Home , Acanthistius, Adelosebastes, Apistidae, Apistus carinatus, Aploactis aspera, Bellator (fish)

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Species Diversity. 2004, 9, 1-36

Phylogenetic Relationships and New Classification of the

Superfamily Scorpaenoidea (: )

Hisashi Imamura

lhe HOickaido University Museum, hacudy ofFisheries, HOkkaido Ultiversity,

3-1-1 Minato-cho. Htikodate, Hbkkaido, 041-8611 .Jtipan E-maii;imamura/(ti/museum.hokudai.ac.jp

(Received 10 October 2e02; Accepted 10 August 2003)

The phylogenetic relationships of the supeirlami!y Scorpaenoidea are re- constructed cladistically. based on speeimens belonging to 18 families, 59

genera, and 86species, by using osteological and myological characters rec- ognized in 111 transformation series. The tbllowing relationships are in- ferred; <1) the former Scerpaenoidei is paraphyletic; (2) the family Sebasti- dae is not monophyletic, most of the genera included having initially branched ofi' other ingroup taxa: (3) the family and Trachyscor- pia have a sister relationship, being nested within the paraphylatic Scor- paenidae; and <4) the family Neosebastidae is the sister group of the former Platycephaloidei, In conelusion, the Scorpaenoidea is reclassified into 20

families, accepting several redefined taxa, such as the , Sebastolo-

bidae, and Seorpaenidae. Key Words: Scorpaenoidea, monophyly, phylogony, classification.

Introduction

The percifbrm superfamily Scorpaenoidea, including the former scorpaeni- fbrm suborders Scorpaenoidei and Platycephaloidei (sensu Imamura 1996; Esch- meyer 1998), was proposed by Imamura and Yabe (2002), who separated the poly- phyletic Scorpaenifbrmes into the perciform Scorpaenoidea and Cottoidei. Al- though Imamura (1996) had earlier recognized the monophyly of the super:lamily and inferred its phylogenetic relationshtps, his cladogram included many unre- solved relationships. Furthermore, although many ichthyologists have recognized the former Scorpaenoidei as a valid taxonomic unit (e.g., Greenwood et al, 1966; Nelson 1976, 1984, 1994; Ishida 1994; Shinohara 1994; Imamura 1996; Imamura and Shinohara 1998; Eschmeyer 1998; Mandrytsa 2001), neither Shinohara (1994) nor Imamura (1996) could find any synapomorphies supporting the monophyly of that taxon. Imamura and Yabe (2002) did not provide taxonomic names corresponding to the former suborders Scorpaenoidei and Platycephaloidei. Recently, although Mandrytsa (2001) initially reconstructed the phylogenetic relationships of Scorpaenoidei (excluding Platycephaloidei) using PAUP ver. 3.1.1, he did not accept the findings and reanalyzed by hand (but it is unclear what methodology he fo11owed) both the fami!ial relationships within his Scorpaenoidei and the intergeneric relationships within each family, He recognized the mono- phyly of his Scorpaenoidei. excluding the fetmilies (sensu Nelson

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2 Hisashi Imamura

1994) and Pataecidae, on the basis of two derived characters (presence of three neu- romasts on the first infraorbital and insertion of the first two dorsal proximal pterygiophores into the space between the second and third neural spines), using

Percoidei as an outgroup. However, these characters are found not acceptable in the present study, as is explained below. In addition, Mandrytsa (2001) recognized several paraphyletic taxa (e.g., subfamilies Scerpaeninae and Neocentropogoni- nae). The purposes of the present study are to reconstruct the phylogenetic relation- ships of the Scorpaenoidea and to propose a new classification based on the rela- tionships.

Methods and Materials

The terminology fbllows Imamura (1996) for osteolog}i, except fbr infraorbitals and extrascapulars, which fo11ow Imamura (2000) and Imamura and Yabe (2002), respectively, and Winterbottom (1974) for myology, exeept for the swimbladder muscles, which fbllow Hallacher (1974). Institutional abbreviations fbllow Leviton et al. (1985). Although (sensu Baldwin and Johnson 1993) was inferred to be closely related to Scorpaenoidea by Imamura and Yabe (2002), the most closely related serranid taxon to Scorpaenoidea has not been specMed; in this study, therefore, three subfamilies (, Epinephelinae, and Serraninae) were used as separate outgroups (Fig, 1). The data were analyzed using PAUP' 4.0blO (Swoflbrd 2002), including the heuristic search option. Character evolution "ordered" "unordered" was assumed as (Wagner parsimony) or (Fitch parsimony) "loss" when the transformation series (TS) contained a derived or character order was equivocal owing to many modifications. The initial familial classification of the scorpaenoids fbllowed Ishida (1994), except for the plectrogeniid PlectrQgenium, and Imamura (1996).

Material examined Scorpaenoidea. : (Bloch, 1801), HUMZ 37372 and one uncatalegued specimen (2, 93-102mm SL). Aploactinidae: Aploactis aspera (Richardson, 1845), HUMZ 37483 (55mm); Erisphex potti (Steindachner, 1896), HUMZ 64275, 108774 (2, 74-87mm). : jqponica Cuvier, 1829, HUMZ 49409, 49411, 49412 (3, 160-182mm), Congiopodidae: Alertichthys btacki (Moreland, 1960), HUMZ 66621 (161 mm); coriaceus Paulin and More- land, 1979, HUMZ 91192 (142mm), Gnathanacanthidae: Gnathanacanthus goetzeei Bleeker, 1855, AMS I. 20188001 (47mm) and AMS IB. 668 (158mm, dissected by Ishida 1994). Hoplichthyidae: Hbplichthys gilberti Jordan and Richardson, 1908, HUMZ 51736 (152mm); U haswelli McCulloch, 1907, HUMZ 50268 (341mm); Il tangsdoi:fii Cuvier, 1829, HUMZ 75337 (156mm). : Maxillicosta raoulensis Eschmeyer and Poss, 1976, HUMZ 110722 (88mm); AJeosebastes thetidis (Waite, 1899), HUMZ 21124 (196mm). Parabembridae: Parabembras curta (Tem- minck and Schlegel, 1843), HUMZ 108335, 108460, 108778 (3, 125-165mm). Pataecidae: Pataecus fronto Richardson, 1844, AMS I. 38479002 (110mm), : Periste- dion orientale Temminck and Schlegel, 1843, HUMZ 106621 (152mm); Satyrichtdys serrutatus (Alcock, 1898), HUMZ 108541 (200mm). Plectrogeniidae: ro-

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Phylogeny of Scorpaenoidea 3

seum Gilbert, 1905, HUMZ 75105, 79190 (2, 85-133mm); Ptectrogenium nanum Gilbert, 1905, HUMZ 37337 (52mm). : (Quoy and Gaimard, 1824), HUMZ 63987 (132mm); Ponttnus macrocephalus (Sauvage, 1882), HUMZ 99503 (121mm); volitans (Linnaeus, 1758), NSMT-P 54350 (93mm); Scorpaena izensis Jordan and Starks, 1904, HUMZ 79974 (134mm); Scorlpaenodes lit- toralis (Tanaka, 1917), HUMZ 101394 (74mm); ScorpaenQpsis neglecta Heckel, 1837, HUMZ (uncatalogued, 125mm); Taenianotus triacanthus Lacepede, 18e2, USNM 106917 (60 mm). Sebastidae: AdeZosebastes latens Eschmeyer, Abe and Nakano, 1979, HUMZ 72035, 75569 (2, 206-2e7mm); hiigendoi:t7 (Steindachner and DOderlein, 1884), HUMZ 39743 (136mm); H percoides (Richardson and Solander, 1842), HUMZ 91501 (164mm); Hbzukius gayotensis Barsukov and Fedrov, 1975, HUMZ 71936 (232mm); matsubarae Hilgendorf 1880, HUMZ 90303 (114 mm); S. owstoni (Jordan and Thompson, 1914), HUMZ 42641 (183 mm); S. thompsoni (Jordan and Hubbs, 1925), HUMZ 92291 (156mm); marmoratus (Cuvier, 1829), HUMZ 69895, 117714 (2, 125-157mm); macrochir(GUnther, 1880), HUMZ 59268, 68329 (2, 112-169 mm); Tracbyscorpia eschnzeyeri Whitley. 1970, NSMT- P 41332 (235 mm). Setarchidae: imus Garman, 1899, HUMZ 751]2 and one uncatalogued specimen (2, 85-113mm); longimanus (Alcock, 1894), HUMZ 79505 (125 mm). : ChoridactyIus multibarbus Richardsen, 1848, CAS 15067 (71mm); erosa (Cuvier, 1829), HUMZ 64250 (63mm); japonicus (Cuvier, 1829), HUMZ 79103, 140587 (2, 104-143 mm); monodactylus (Bloch and Schneider, 1801), HUMZ 101864 (95mm SL); S,nanceia horrida (Lin- naeus, 1766), CAS 15073 (106mm). Tetrarogidae: TetrarQge niger (Cuvier, 1829), URM-P 31819 (18mm, data from G. Shinohara). : Bellotor egretta (Goode and Bean, 1896), HUMZ 69393 (87 mm); Chelidonichthys spinosus (McClelland, 1844), HUMZ 108541 (200mm); micrQptera GUnther, 1873, HUMZ 48942 (159 mm); stearnsii Jordan and Swain, l884, HUMZ 32456 (98mm); Pter.v- gotrigla macrortrynchus Kamehara, 1936, HUMZ (uncatalogued, 95mm). In addition, platycephalid specimens, including l6 genera and 38 , were also examined, such being listed in Imamura (1996).

Comparative materials Serranidae. Anthiinae: Ptectranthias keltoggi azumonus (Jordan and Richard- son, 1910), HUMZ 79389 (104 mm). Epinephelinae: Diploprion bdesciatum Kuhl and van Hasselt, 1828, HUMZ 87031 (122mm); EZpinqphelus awoara (Temminck and Schlegel, 1843), HUMZ 97035 (125mm); Niphon sptnosus Cuvier, 1829, HUMZ (uncat- alogued, 126mm). Serraninae: ChelidQperca hirundinacea (Valenciennes, 1831), HUMZ 36481 (102 mm). A further 31 percoid species that were previously examined were listed in Ima- mura (2000),

Phylogenetic Analysis of Scorpaenoidea

Monophyly of Scorpaenoidea Imamura and Yabe (2002) recognized the monophyly of the Scorpaenoidea on the basis of three synapomorphies: presence of a suborbital stay, the parietal sen-

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sory canal bony structure with spines, and an extrinsic swimbladder muscle de- rived from the obliquus superioris (Fig. 1). They also suggested that the single pos- tocular spine in the larval stage may be a synapomorphy of Scorpaenoidea and

Serranidae, or of the supe family together with the Anthiinae and Epinephelinae,

Although most or many members of the scorpaenoid superfamily have a back- wardly-directed opercular spine that crosses the subopercle and also possess a cau- dal fin muscular element, the adductor dorsalis, Imamura and Yabe (2002) inferred that these characters are the synapomorphies of Scorpaenoidea plus Serranidae (= their Scorpaenoidei) (Fig. 1). The family Congiopodidae (sensu Nelson 1994=Congiopodidae+Zanclorhyn- chidae sensu Mandrytsa 2001) was included in the suborder Congiopodoidei by Mandrytsa (2001), However, the family possesses a suborbital stay, a parietal sen- sory canal with spines (spines absent in CongiQpodus), an extrinsic swimbladder muscle derived from the obliquus superioris, and a backwardly-directed opercular spine that crosses the subopercle, although it does not have the adductor dorsalis. Because the adductor dorsalis is also absent in several scorpaenoids (e.g., Synan- ceiidae, Aploactinidae, Triglidae, and ), a condition regarded as a reversal (see Imamura and Yabe 2002), the family is included in the fo11owing phy- logenetic analysis of the superfamily Scorpaenoidea. Mandrytsa (2001) excluded the family Pataecidae from his Scorpaenoidei, placing it in the perciform suborder Pataecoidei. Although the family does not possess a suborbital stay, spines associ- ated with the parietal sensory canal, and adductor dorsalis, it does have an extrin- sic swimbladder muscle originating from the posteroventral border of the neuro- cranium and inserted onto the vertebrae, This muscle agrees well with that recog- Serranidae Scorpaenoidea (OUTGROUP) (INGROUP)

l with

larval stage

Fig. 1. Phylogenetic relationships of the Scorpaenoidea (ingroup) and Serranidae (outgroup) inferred by Imamura and Yabe (2002).

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Phylogeny of Scorpaenoidea 5

nized in many scorpaenoids (and also Cegiopodidae). In addition, the Patecidae has a backwardly-directed opercular spine that crosses the subopercle (see also Ishida 1994, fig. 3D); therefore, this family is also included in the phylogenetic analysis below. In the Scorpaenoidea, the configuration of the suborbital stay can be divided into three morphotypes (exeept in Pataecidae, where it is absent-see TS 8), these characters being included in the phylogenetic analysis. There are no spines associ- ated with the parietal sensory canal in Tetraroge, CongiQpodus, Gnathanacanthus, and Pataecus (Imamura and Yabe 2002), their presence or absence also being in- cluded in the analysis (see TS 24). Other variations considered inc!ude the bony el- ements associated with the extrtnsic muscle (see TS 101).

Characters used for phylogenetic analysis Usefu1 characters for the analysis of scorpaenoid phylogenetic relationships were found in 111 transformation series, The character matrix is shown in Table 1,

Autapomorphic characters, recognized only in the terminal taxa, are not included in the fo11owing character list and phylogenetic analysis. Many centradictions were found between the character scoring in this study and that in Ishida (1994). Some were due to different identifications of bony, carti- laginous, and muscular elements (e.g., sixth inhaorbital, second pharyngo- branchial, interarcual cartilage, and pelvic fin muscles), but it was often unclear whether Ishida's (1994) character recognition was simply incorrect or differed owing to intraspecific variation. The derived characters recognized in this study

as acceptable for analysis exclude those known to be subject to intraspecific varia- tien. Except where otherwise noted, all characters were appraised by direct exami-

nation of specimens. In this study, I do not discuss several previously recognized transformation se- ries [i.e,, TS 21 (entopterygoid), 31 (configuration of actinosts), 32 (cartilaginous band between scapula and coracoid), and 59 (muscle bundles serving free pectoral fin rays) of Imamura (1996)], because the morphological gaps between the charac- ters in these transformation series were fi11ed by the newly examined specimens

and the characters ceuld not be clearly divided into distinct morphotypes. In addi- tion, Imamura's (1996) TS 46 (first and second neural spines) is also not discussed, because the homology of the vertebrae supporting these spines is unclear (see dis- "Classification"). cussion under TS 1. First to third inhaorbitals: O, closely associated; 1, separated. The first to third infraorbitals are closely associated with each other in Serranidae (e.g., Bald- win and Johnson 1993). They are cempletely separated from each other in Gnathanacanthidae and Pataecidae. TS 2. First and third infraorbitals: O, separated by second infraorbital; 1, at- tached; 2, separated by second infraorbital and ectopterygoid (unordered). The first and third infraorbitals are separated by the second inhaorbital in Serranidae (e.g., Katayama 1959). Characters 2-1 and 2-2 are restricted to Triglidae and Peristedi- idae, respectively (Fig, 2). TS 3. Middle portion of second infraorbital sensory canal: O, bridge absent; 1, bridge present with no sensory openings; 2, bridge present with a single sensory opening; 3, bridge present with more than two sensory openings (unordered). In most scorpaenoids examined, the sensory canal of the second infraorbital has

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Table I.Charactermatrix of superfamily Scorpaenoidea.

TransformatienSeries Taxon

156 !O11-15leP20 21-252613031-3536-4041-45 4650 'Outgroup

Serraninae oueoeoooooOOAOOooooooooooeooooooooooooooOOAOOooooooooooAOAOOooooooeooooloeooooooooeooooeoooooooOOOAOAOOIOooooooooooooooooooooooooeoooooooooooooooooooo Anthiinae Epinephelinae lngroup Sebastis(t"s OOIOOOOIOOOOIOOOOIOOOOIOOeoleoeoleoeolooooleoO0300oo3e?O030?OOIOOoelooC)OIO?oolo'g(]OIC)[}OOIOOOOIOOuoleoOOIOOOOIOOOOIOOO[)100OOIOOOOIO?OOIO?1010010100O0200O0200O0200O0200O0200O0200O1211OllllOllllO1211O12110211002210OODOOOOIOOoeleoooleDoe2ooO0200O0200O13001131e1131111311103iO1032010320ll:llOll3101132111321O1311Ol311Ol:l12O1311O1311O1311O1311{]1:l12Ol;'SllO1320O1321O131AOOO12Ol310O131eO1311O1310O1300O1300O1300O1300O1300O1301O1301O1300e13ooe1301O1300oooooooooooooooooooooooooooooo10000leooo11OOO110001100011000llOOO1100011eooneooOOIOOoooooooooooooeooeoeooooooOOIOOoooooOIOOOOOIOO11100ooooooooooooeoooooooooooooooooeoooooooooOOIIOeolloUOIIOOOIIOOOIIOOOIItOOI11ooloeOOAOOoooooooooooooooooooooooooooooooeoooooooe2(}ooeoooooooooonoooooooooOOOIOOOOBOOOOBOOO020oooooOO020oooooooooo20C)OO10020OO021ooeol2000120101200AOOO121ooooooooooooooo10AOOoooouoooooooooo11UOI2100010000elooo2002120021OO021AOAOOOOOIOOOOIOoooleOOOIOOOOIOOOOIOOOOIOOOOIOC)OOIIOOOIOOOO12OOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOIOOOOO1210e12ooe12OC]Ol210012OOO12OOI121010310113OOO02OOO02oooleOIOIOOOIIO00110OOOIOeooloelOllOIOIOOIO12OIOIOOIOIO1101211012OOIIDOIOIOooooooooooooooooooooooooooooooooonoOOOOIOOOOIOOOOIOOOOIOOOOIooooooooooooeoenooooOIOOOoooo{)OOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOIIIOOOIIOOOOIOOOOIooooooooooeooooeoooo10000ooooooloelOIOOIoloeloloeoOIOOOOIIOIOIIOI1000010000OOOIOOOOIOOOOIOOOOIO1001010010100101001011110oooleOOOIIOOOIIOOOIO{}OOIOOOOIOoootoOIOIOOIILOOIIIOOIIIOOI!IOOIIIOOIIIOOIOIOOIOIOOI020OI020olo2eelo2oOOOIOODOIOOOOIOOOOIOOOOIOoooleOOOIOOOOIOOOOIOOOOIOOOOI{}OOOIOOOOIOOIIIOOOOIOoooooooooooooooooooeeooooooeoooooooOOIOOooooooooooooooouoooooeoooooooooooooooooooooooOOOOIOOOIOOOOOIOOIOI20101OOIOIOIIelOllOl22001Ollel2000121001ooeooooooooooooOOIOOoooooooooo201102011020110OOIIOOOIIOlooel101elOOIIOOOAIOoooooOOOAOOOOIOOOOIOoeoAoOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOIOIOeooloOOOIOOOO]AooeloOIOIBOII12Olll?OIO12OlllOOII12O?212ell12eoe1220012QllllellllOOOIOOIOIOOOOIO10010OOOIOOOOIOOOOIOOOOI{]OOOIOOOOIOOOOIOOOIIOO?210llOIOOOOIOOIOOOOIOOOOIOOOOIOOOOIOUOoooooOIOOOOIOOOooeoloooooOOOOIDeoooOOOOIoooooooooooooooOIOIOOIOOOOIOOOOIOOOelo2o21030ilOlleloolOIOOIOOOOIOIOOIOIOOIO120lOIOOOOIOOOooooooooooooooooooooOO130OOI:SOOO130OO13DOO130OO020[}OILOOOI:30OOEUO Sebastes Hoi･ttkius Htalicotenus Sebastoiobus Sco!paena Scorpaenopsis Taertianotus Scmpaenodes PteroisDendrochirus

Pontlnus Tr'achyscorpia Setarches Ectreposebastes ApistusTeti'aroge

S.vnanceia EJ'osaInimictts

Chortdact.ytus

,'LVtlnt')usAploactis

ErisphexCongi.opodus

Aterti('hthi,s Grv.(zthanacatithus Pataecus Neoset)astes .,T･laxiilic/osta Piecrrogenium Bemhradiurn Parabernhras Bernbras Chelidon.i('hthvs Lepidott'igla Ptet),buotri.uta i'rlonotus BellatorPeristedion

Satyrichth)'s Elopiichthys P]atycephalidae

Nurribers us$igned to tyanstbmnation series and to inC:2andthe3. Polytno]'phic characters are coded as follows/ O andcharacters1, A;1 andcerrespond2, B: O anc]to2,those D:text.O, 1,and 2. E: O, 1T

2, and 3. F.

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Phylogenyof Scorpaenoidea 7

Table 1,Continued.

TransformationSeries

51-5556 60 61 65 65 7e71-7576- 8081 85 86-9U91-95 96 10e 101-105106-110111

oooooooooooooooeoooo oeooo OOAAOOOAAOOOAOOooeooooooooooooooooooooooooooooeooo ooooo ooeoo ooooo oeeoe oooooooooooooooooo

OOOOO OOOOA ooooo ooooe ooeoo oDooo ooooo

AOAOA OOOOF ooooo ooooe ooeoo ooooA ooooe

oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooOOOIOooooooooooOOOOIeoe21ooe21oooooooeoloooo]OD030.ooooooooooOIOOOOIOOOOIOOO0200012eoo0200002000120001200012100120001210002100oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooeooooooo{}o{}()OOOOOIOOOOI10001OOIOIOOIOI101{]110100oo(}ooOO120eooooooooooooooOO121oeoeoOOIII101111011110111OOIIIOOIII111il111111111110121OOO02ooeo2OOO02ooeo2OOO02OOO02OOO03OOO03OOO03oeoo3OOO03OOO03OOO02OOO02OOO02OOO03OOO03Oll03OOOO13OOO03OIO03OIO03110031000311103oooo:s10003110D3110e3OOO03OOO03oooa3oooo:sOOOOIOooDDOO023OO023OOO13oooo:iOOO03OIO03oloonOOO03OOO03OOI1OOOIIOOOIIOOOIIOOOIIOOOIIOOOIIOOOIIO30110OOIOOeolooOOIOOOOIOOoooooOOO{K)oeoooOOIIO3311133111OOIII3311033110:S311033110C2111OOIIIOOIIIllllA33111oooooooooo2200022000ooeoooooooOOOIIOOIIOOOOIOooo?oOOOIOO131113:llllOOIIOOOIAOooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooonoooooooooooooooo1OOOO1?OOO10000110001101011000]lelo11eoo11eoo10{)10100101110111111oooooooooonoooovooooooeoooDeooODOIOooeloooelooooooooooo111101111011110OAOOOooooooooooooooooooooooooooooooooooooooooooooooooooooeoooooooooooooooooooeooooeoooooooooeooOIOOIOIOOIIAOOOoooooOOIOOOOIOOOOIOOllOOOlllOOAOIII121LOooe?oOOOOIoooooooooooooeoOOODOloo2eooe2oOO020OO020OO0201002010020ooo?oooooooooooeooloeooloOOOIOOOOIOOOOIOOOOIOOOOIOOOIIOoeooooolleoolleoooooooooo(}o{}(]{)eoeoo101012011012110121101011011111l21111111111111l12001110011110llOIOooooooeoolOOOIIOOOIIoooooOOOIOOIOIIOIOIIOIOIIOIOIIOIOIIOOOIIOOOIIO?OAIOOOAIoooeooeoeooooooooooooeooooooooooooooooooooooooo{}(]()ooooooooeoooooooeDooOIOOOOIOOOOOIOOOOOIOOOOIOOOOIOOOOIOooooooooooOO?10OOIIO20001200012000010110OOOIOooooooooooooeoloooooeoooooooooooooeooooooooooooeooooooooooooOOOIIOOOAOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOOIOOOI1OOOo?ooOllOOOOIOOOOIUOOOIOOOOIOOOOIOOooooooooooooooooooooooeolooeoleoooloooooooooo11oeo11100OllOOOIOOOooeooooooooooooOOIOOoo{)(]ooooooOllOlOllOlOllOlOllOlellOlOllOlOllOlOOOAOOOOIOoooooooeoooooooooeoooooooooeooooeooeoeooooooooooooooooooooooooooooooooooooooooooooeooeoeooOOOIIOOOIIOOOOI10001oeool101el10101OOOII10111OOOOAO?111ooooooeooooooeooooooooo{}ooooooOIOIIOIOIIOIOIIOIOIIOIOIIOIOOIOIoo1oAoelOOOOIloeoo200002000020000200C}O20000200002001030010100101001010010200002oeoo20010200101101020011201?O201?O2001020010200102001120011201?O2el?o200102eOll30000200002000C}200002000020000OIOIOOIOIO201?OOIOIO3101e201?O201?O20AIO20000ooooooooeoooooooooooooouoooouoooooooooooooooooooooOOIOOOOIOOoooooooooooooooooooo10000IOOIO11010llOIO1101011010?10101001110Ull?2010?2011OOOIO10010ooooo10000oooooooooo10000eoooo?OIO(]?OIOO?oooooooooeoooo?1021?1021?1021?OAOAooo0oo(}oooooooooc}222222oo2o22oooooooeoeo33B0

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A

5mm

Fig. 2. Lateral view of infiraorbitals. A, Beltator agretta, HUMZ 69393, 87mm; B,

orientate, HUMZ le6621, 152 mm. ECP, ectopterygoid; IO, infraorbitals. Open and solid arrows

indicate, respectively, the opening ln the mid-ventral region of the second infraorbital and

that in the third infraorbital directed upwards.

openings at the junctiens with the first and third infraorbitals (character 3-1) (Figs 2B, 3A-C). On the other hand, Neosebastidae, Plectrogeniidae, Parabembridae, Be- mbridae, Chelidonichthys, Prionotus, , , and HOplichtdys

haswetli have an additional opening in the mid-ventral region of the second infra- orbital (character 3-2) (Figs 2A, 3D), whereas Scot:paenodes, Pterois, Dendrochirus, HOplichthysgilberti, and Il tangsdotzfii have more than two openings in that region (character 3-3), Bridges and hence openings from the second infraorbital sensory canal are absent in the Anthiinae and Serraninae, and also in some members of Epinephelinae (e,g., AporQps and Lioprqpoma; see Baldwin and Johnson 1993) (character 3-O). In other epinephelines examined (IViphon and llpinephelus), the canal has two openings at the anterior and posterior junctions, but lacks an addi- tional opening(s) in the middle portion (character 3-1), TS 4. Third infraorbital and lateral ethmoid: O, separated; 1, attached, The lat- eral ethmoid is attached only to the first infraorbital in Serranidae. Character 4-1 occurs only in Triglidae and Peristediidae. TS 5. Third and fifth infraorbitals: O, separated by fourth infiraorbital; 1, at- tached. The third and fifth infraorbitals are separated by the fburth infraorbital in Serranidae (e.g., Katayama 1959; Baldwin and Johnson 1993), as well as in most scorpaenoids, Attachment of these bones occurs only in Triglidae (Fig. 2A). In Pterois, Dendrochirus, Setarchidae, and Congiopodidae, the fifth infraorbital is ab- "?" sent and this character is coded as (see also TS 9). TS 6. Direction of posterior opening of third infraorbital, continuous with sen- sory canal of fourth to sixth infraorbitals: O, upward; 1, backward. Serranidae and most ingroup taxa exhibit character 6-O (e.g., Baldwin and Johnson 1993) (Figs 2, 3A-B, D), whereas eharacter 6-1 occurs in Scorpaenidae, Trachyscorzpia, and Setarchidae (Fig, 3C). TS 7. Position of upward (or backward) epening of third infraorbital: O, on dor- sal (or posterior) margin of element; 1, below (or anterior to) margin. In Scotlpae-

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Phylogeny of Scorpaenoidea 9

A B K 10 lill Nfr

5mm c e D

･"

}ii CI:I.

× 5mm 5mm

Fig. 3. Lateral view of infraorbitals. A, , HUMZ 69895, 125mm; B, Hbzukius guyotensis, HUMZ 71936, 232mm; C, ScorpaenQpsis neglecta, HUMZ uncat., 125mm;

D, Nbosebastes thetidis, HUMZ 21124, 196mm. IO, infraorbitals; SS, suborbital stay. Open and

solid arrows indicate, respectively, the opening in the mid-ventral region of the second infra-

orbital and that in the third infiraorbita1 directed upwards or backwards.

nodes, Pterois, Dendrochirus, Pataecidae, and Sebastidae (except Sebastolobus and Trachyscorlpia), the upward (or backward) opening of the third infraorbital is situ- ated along the dorsal (or posterior) margin of the element (Figs 2, 3A-B), as in Ser- ranidae (e.g., Baldwin and Johnson 1993). This epening is positioned below (or an- terior to) the margin in the other ingroup taxa (Fig. 3C-D). TS 8. Suborbital stay on third inhaorbital: O, absent; 1, present, its posterior end pointed or bluntly pointed and not strongly connected with preopercle; 2, pres- ent, its posterior end narrowly truncated and strongly connected with preopercle; 3, present, its pesterior end broad and strongly connected with preopercle (ordered as O-1-2-3), A suborbital stay is absent in Serranidae (Katayama 1959; Baldwin and Johnson 1993) and Pataecidae, whereas character 8-1 occurs in Sebastiscus (Fig. 3A) and Sebastes, character 8-2 in Adelosebastes, Hbzukius (Fig. 3B), and Heli- cotenus, and character 8-3 in the remaining taxa (Figs 2, 3C-D), TS 9. Fourth and fifth infiraorbitals: O, both present; 1, fburth absent; 2, both absent (ordered as O-1-2). In Serranidae, the fourth and fifth infraorbitals are both present (e.g., Katayama 1959; Baldwin and Johnson 1993). Trachyscot:pia and Scor- paenidae (except Pterois and Dendrochirus) have a single infraorbital element pos- terior to the eye (Fig. 3C). In these taxa, a prominent gap exists between this ele- ment and the third infraorbital. Thus, the element could be identified as the fifth infraorbital, the gap representing the loss of the fourth infraorbital. Apistidae, Tetrarogidae, Synanceiidae, Aploactinidae, Gnathanacanthidae, Pataecidae,

Neosebastidae, and Plectrogeniidae atso have a single element, but lack an exten- sive gap (Fig. 3D). A larval synanceiid, Inimicus J'aponicus (10.1mm), with such a gap has also been reported (Imamura and Yabe 1998). In addition, among the scor-

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10 Hisashi Imamura

paenoids having both the fourth and fifth infraorbitals (e.g., Sebastiscus, H02ukius, and Chelidonichthys), either both bones have a single bridge in the middle portion, the fifth has such a bridge but the fourth does not, or both the fourth and fifth lack a bridge (Figs 2, 3A-B). If a fusion of these elements has occurred, the fused ele- ment should have either a single bridge in the upper portion, two bridges in the upper and lower pertions, or no bridges, However, the above-listed taxa have a sin- gle bridge in the middle portion of the element (Fig. 3C-D); thus the fusion of the fourth and fifth infraorbitals cannot be assumed. I assumed that the fourth infraor- bital is also absent in the above-listed taxa, the gap between the third and fifth being considered to decrease with growth, On the other hand, both the fourth and fifth infraorbitals are absent in Pterois, Dendrochirus, Setarchidae, and Congiopo- didae. TS 10. Sixth inflraorbital: O, attached to sphenotic; 1, fused to sphenotic; 2, ab- sent (unordered). A sixth infraorbital (=dermosphenotic) is present and attached to the dorsal aspect of the sphenotic in the Serranidae examined, Character 10-1 is

recognized in Scorpaenopsis, Ttienianotus, Setarchidae, Apistidae, Tetrarogidae, Synanceiidae (except ), Erisphex, Alertichtbys, a specimen of Gnathanacanthus goetzeei (AMS IB. 668), Ptectrogenium, Prionotus, Betlator, and Hoplichthyidae, and character 10-2 in Synanceia, Aploactis, and Pataecidae, When a suture line on the dorsal surface of the sphenotic was absent, but a bony tubular structure or canal was present, it was assumed that the sixth infraorbital and sphenotic were fused (character 10-1). The sixth infraorbital was considered absent (character 10-2) when both a suture line and tubular structure or canal were ab- sent. TS 11. Sensory canal of third infiraorbital: O, continuous with that of infraor- bitals posterior to eye (fburth and fifth); 1, not continuous with that of inhaorbital posterior to eye (fifth). The sensory canal from the third infraorbital to the neuro- cranium is continuous through the fourth and fifth infraorbitals, situated poste- rior to the eye, in the Serranidae examined, Character 11-1 is recognized in Scor- paenidae, Trachysco]lpia, Setarchidae, and Atertichthys, in which the fourth infra- orbital is absent (see TS 9). TS 12. Sensory canal of head: O, continuous with that of infraorbital; 1, not continuous with that of infraorbital. Charaeter 12-1 occurs in Scorpaenidae (except Scor:paena and ScorpaenQpsis), , Setarchidae, Erisphex, and Aler-

tichthys, See comment under TS 11 fbr the serranid condition. TS 13. Tubercles on neurocranium: O, absent; 1, present. Tubercles on the dor- sal surface of the neurocranium are absent in most scorpaenoids and Serranidae, except tn Diploprion, in which stout tubercles are present (the neurocranium is "rugose" in the tribe Diploprionini; Baldwin and Johnson 1993). Tubercles are also present in Apistidae, Minous, Congiopodidae, Triglidae, Peristediidae, Ho- plichthyidae, and some platycephalids (e,g,, and ). TS 14. Nasal and neurocranium: O, loosely attached; 1, sutured. The nasal is loosely attached to the neurocranium in most scorpaenoids and Serranidae exam-

ined. Character 14-1 is restricted to Triglidae and Peristediidae. TS 15. Nasals on both sides: O, separated; 1, sutured medially, The nasals are separated in the Serranidae and most ingroup taxa examined, but are sutured me-

dially in Peristediidae. TS 16. Number of tooth plates on prevemer: O, one: 1, two; 2, none (unordered).

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Phylogeny of Scorpaenoidea 11

There is a single tooth plate on the prevomer in the Serranidae examined and most scorpaenoids, although Baldwin and Smith (1998) reported the epinepheline Dipro- prion having two tooth plates, Character 16-1 is found in Minous, Bembradium, Lqpidotrigla, Prionotus, and Platycephalidae (Elates and Onigociinae), and charac- ter 16-2 in IZienianotus, Choridacts,lus, Congiopodidae, Gnathanacanthidae, Pterly- gotrigia, and Peristediidae. TS 17. Lateral ethmoids: O, separated; 1, meeting in midline. The lateral eth- moids are separated by the ethmoid in Serranidae (e.g., Katayama 1959) and most ingroup taxa, but meet in the midline in Triglidae (Lopidotrigla, PteioJgotrigla, and Beltator) (see Imamura 1996, fig. 9). TS 18. Parasphenoid and pterosphenoid: O, separated; 1, connected. In the Ser- ranidae examined, except Eipinephelus, the parasphenoid and pterosphenoid are separated by the prootic, A parasphenoid-pterosphenoid connection is fbund only in Atertichthys, Pataeeidae, a specimen of Bembradium roseum (HUMZ 75105), and several platycephalids (e,g., Elates and Poptltocuticeps) in the ingiroup, as well as in Eipinqphelus. TS 19. Basisphenoid: O, present, posterior margin connected directly with neu- rocranium; 1, present, posterior margin free from neurocranium; 2, absent (un- ordered). The posterior margin of the basisphenoid is cennected directly with the neurocranium in the Serranidae examined. Eschmeyer and Collette (1966) and Ishida (1994) found that all of the genera in the Setarchidae had character 19-1, whereas character 19-2 was recognized for this family in the present study, as also in Matsubara (1943), It is assumed, therefore, that this osteological character "1" '`2" varies intraspecifically in the family, Accordingly, it is coded and for Setarchidae, Character 19-1 is also apparent in Trachyscor:pia and a specimen of Gnathanacanthus goetzeei (AMS I, 20188001), whereas character 19-2 occurs in Apistidae, th,nanceia, Minous, tttploactis, Pataeeidae, Peristediidae, and Ho- plichthyidae. TS 20. Prootic and intercalar: O, in contact; 1, separated. The prootic and inter- calar are in contact in Serranidae (Katayama 1959), but separated in Aploac- tinidae, Congiopodidae, Pataecidae, Lqpidotrigta, Peristediidae, and Hoplichthy- idae. TS 21. Intercalar and posttemporal: O, with ligamentous articulation; 1, su- tured, The intercalar and ventral limb of the posttemporal are articulated via a lig- ament (therefore separable) in the Serranidae examined. Character 21-1 is found in Erosa, Minous, Congiepodidae, and Peristediidae. TS 22. Lateral pore on pterotic: O, absent; 1, present, A lateral pore on the pterotic, continueus with the preoperculo-mandibular sensory canal, is absent in many ingroup taxa, but present in Maxillicosta, Triglidae, Peristediidae, and Platy- cephalidae (see Imamura 1996, fig. 20). Among the Serranidae examined, character 22-O is fbund in the Anthiinae and Serraninae, and character 22-1 in Epinephelinae. TS 23. Skinny sensory canal between pterotic and preopercle: O, present; 1, ab- sent. A skinny sensory canal is present between the pterotic and preopercle in the Serranidae examined. In the ingroup, this canal is absent in Tetrarogidae, Erisphex, Congiopodidae, Plectrogeniidae, and Hoplichthyidae. TS 24. Spines associated with sensory canal on parietal: O, absent; 1, present. All Serranidae lack spines and a sensory canal on the parietal, except for the epi- nepheline Niphon (which has a sensory canal but no spines). Most scorpaenoids

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12 Hisashi Imamura

A

rt.

5mm 5mm

Fig. 4. Lateral view ofpremaxilla. A, Helocotenus hiigendotzf7, HUMZ 39743, 136mm; B, Cheti-

donichthys spinosus, HUMZ 108541, 2oomm. Arrow indicates notch between ascending process and cranial condyle.

possess spines assoctated with the sensory canal on the parietal, although spines are absent in 7letraroge, CongtQpodus, Gnathanacanthidae, and Pataecidae (see also Imamura and Yabe 2002). TS 25. Baudelot's ligament: O, originating from basioccipital; 1, originating from basioccipital and first vertebra; 2, originating from first vertebra; 3, absent (unordered). Character 25-1 is recognized in 7keenianotus and Chelidonichthys, character 25-2 in Pterois, Synanceiidae, Aploactinidae, Gnathanaeanthidae, Patae- cidae, Pter:ygotrigla, Peristediidae, and Hbptichthys haswelli and H gitberti, and character 25-3 in Congiopodidae and Il langsdoi:fii. Other ingreup members and Serranidae show character 25-O (e.g., Gosline 1966), TS 26. Ascending process of premaxilla: O, continuous with remaining part of premaxilla; 1, separated, The ascending process of the premaxilla is continuous with the remaining part of the bene in most ingroup taxa (Fig. 4) and Serranidae examined, whereas the process is separate in Parabembridae, Hoplichthyidae, and Platycephalidae (see Imamura 1996, fig. 21). TS 27. Notch between ascending process and cranial condyle: O, present; 1, ab- sent. In the Serranidae examined and most ingroup taxa, a notch is present be- tween the ascending process and the cranial condyle (Fig. 4A), whereas it is absent in Apistidae, Triglidae (Fig. 4B), and Peristediidae. TS 28. Teeth on jaws: O, present; 1, absent. Jaw teeth are present in the Ser- ranidae and most scorpaenoids exainined (Fig. 4), but absent in Congiopodus and Peristediidae. TS 29. Palatine and ectopterygoid: O, connected; 1, separated, The palatine and ectopterygoid are connected in Serranidae (e.g., Gosline 1966; Baldwin and John- son 1993) and most scorpaenoids (Fig. 5A-C), whereas they are widely separated in Congiopodidae (Fig. 5D), TS 30. Teeth on palatine: O, present; 1, absent. Palatine teeth are present in the Serranidae examined. Character 30-1 is fbund in Scorpaenopsis (Fig. 5B), Taeniano- tus, Scor:paenodes, PteTois, Dendrochirus, Synanceiidae (Fig. 5C), Congiopodidae (Fig. 5D), Aploactinidae, Gnathanacanthidae, Pataecidae, Chelidonichthys, Lepi- dotrigta, Pter:ygotrigta, and Peristediidae. TS 31. Ectopterygoid and metapterygoid: O, separated; 1, meeting medially,

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Phylogeny of Scorpaenoidea 13

The ectopterygoid and metapterygoid are separated in the examined specimens of

Anthiinae, Serraninae, and IViphon, whereas they meet medially in DipIQprion and Epinephelus (Baldwin and Johnson 1993). Character 31-1 is found in Heticolenus, Se- bastolobus, Scoilpaena, Scoilpaenopsis (Fig. 5B), and Ttzenianotus, TS 32. Metapterygoid lamina: O, present; 1, absent. The metapterygoid lamina is typically present in Serranidae (e.g., Katayama 1959). Character 32-1 is fbund in 71aenianotus, Apistidae, Tetrarogidae, Synanceiidae (Fig. 5C), Aploactinidae, Con- giopodidae (Fig. 5D), Gnathanacanthidae, Pataecidae, and Hoplichthyidae. TS 33. Space between metapterygoid and hyomandibula: O, prominent; 1, rudi- mentary or absent. A prominent space for the insertion of the levator arcus pala- tini is present between the metapterygoid and hyomandibula in the Serranidae ex- amined. Character 33-1 is present in Ttzenianotus, Tetraroge, Synaneeiidae (Fig. 5C), and Hoplichthyidae. TS 34. Preepercular margin: O, with serrations; 1, with prominent spine(s); 2, without serrations or prominent spines (unordered). In most scorpaenoids, the preopercle margin has five or fewer prominent spines posteriorly (Fig. 5A-C), whereas it lacks prominent spines in Congiopodidae (Fig. 5D), Gnathanacanthidae, and Pataecidae. On the other hand, a serrated preopercular margin is present in Serraninae and some anthiines (e.g., Plectranthias), and a margin with spine(s) in other anthiines (e.g., Acanthistius) and Epinephelinae (Baldwin and Johnson 1993;

B x t

5mm D

(tJ-l--=z

5mm 5mm

Fig. 5. Lateral view of suspensorium. A, Sehastiscus marmoratus, HUMZ 69895, 125rnm; B,

Scorpaenopsis naglecta, HUMZ uneat., 125mm; C, , CAS 15073, 106mm; D,

Alertichtbys blacki, HUMZ 66621, 161 mm. ECP, ectopterygoid; ENP, endopterygoid; HM, hye-

mandibula; IOP, interopercle; ML, metapterygoid lamina; MP, metapterygoid; OP, opercle; PAL, palatine; POP, preopercle; QUA, quadrate; SOP, subopercle; SYM, symplectic.

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14 Hisashi Imamura

present study). TS 35. Backwardly-directed opercular spine: O, present; 1, absent. A back- wardly-directed opercular spine is present in Serranidae (e.g., Gosline 1966; John- son 1983; Imamura and Yabe 2002) and most scorpaenoids (Fig. 5), whereas it is ab- sent in Pterots and Dendrochirus, TS 36. Basihyal: O, essified; 1, cartilaginous; 2, absent (unordered). The basi- hyal is ossified in the Serranidae examined, but cartilaginous in Peristediidae (see Imamura 1996, fig. 11C) and absent in Choridac(ytus, Congiopodus, Gnathanacan-

thidae, Pataecidae, Chetidonichthys, Lepidotrtgta, and PterlJ;gotrigta, TS 37. Branchiostegal rays: O, seven; 1, six; 2, five (ordered as O-1-2). Seven branchiostegal rays are present in Serranidae (e.g., Katayama 1959) and most scor- paenoids. Six branchiostegal rays are found in Aploactinidae, Alertichtdys, and Pataecidae, and five rays in Congiopodus. TS 38. Interarcual cartilage: O, present; 1, absent. The interarcual cartilage is typically present in Serranidae (e.g., Baldwin 1990), but not in Scorzpaenopsis, Synanceiidae (except Synanceia and Erosa), Aploactinidae, Alertichthys, Bembra- diutn, Triglidae, Satyrichtbys, Hoplichthyidae, and Platycephalidae (except Platy- cephalus, Etates, Ratabulus, and T)'iysanQphiois cirronasa). Identification of this cartilage fo11owed Imamura (1996) and Imamura and Yabe (1998). TS 39. Tooth plate on second epibranchial: O, absent; 1, present. An autoge- nous tooth plate is present at the articulation between the second pharyngob- ranchial and second epibranchial in the serranines and epinephelines, whereas it is absent in the anthiines (e.g., Baldwin 1990, see also her figs 20 and 23). On the other hand, a tooth plate is sutured to the middle portion of the second epi- branchial in Triglidae, Hoplichthyidae, and P!atycephalidae (see Imamura 1996, fig, 30), It is assumed that the autogenous tooth plate that occurs in several ser- ranids and the sutured tooth plate characteristic of scorpaenoid families are not homologous. TS 40. Tooth plate on third epibranchial: O, present; 1, absent. A tooth plate is present on the third epibranchial in the Serranidae examined, but absent in Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodidae, Gnathanacanthidae (Fig. 6C), Pataecidae, and Peristediidae. TS 41. First pharyngobranchial: O, ossified; 1, cartilaginous; 2, absent (un- ordered). The first pharyngobranchial is ossified in the Serranidae examined, car- tilaginous in Bembradiuni and Hoplichthyidae, and absent in Atertichthys. TS 42, Tooth plate on second pharyngobranchial: O, present; 1, absent. In Ser- ranidae, the second pharyngobranchial has a medial tooth plate (Baldwin 1990), such also being present in most scorpaenoids (Fig. 6A). A tooth plate is absent in Pontinus, Tetrarogidae, Synanceiidae, lirisphex, Gnathanacanthidae (Fig. 6C), Pataecidae, Maxitticosta (Fig. 6B), and Hoplichthyidae. In Aploactis and Satyrichthys, the second to fourth pharyngobranchials are continuous, with only a single large tooth plate present, Because the relationship of this tooth plate to the "?", second pharyngobranchial is unclear, it is coded as TS 43. Second to fourth pharyngobranchials: O, separated; 1, third and fburth continuous; 2, second to fburth continuous (ordered as O-1-2). The second to fburth pharyngobranchials are separated in the Serranidae examined, Character 43-1 is reeognized in Synanceiidae (except inimicus), Erisphex, Gnathanacanthidae (Fig, 6C), Pataecidae, and Peristedion, and character 43-2 in Aploactis and Satyrichtdys,

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Phylogeny of Scorpaenoidea 15

PB c

5mm 3mm 1mm

Fig. 6. Medial view of upper branchial arch. A, Heticotenus hiig. endotlfi, HUMZ 39743, 136mm; B, Maxitlicosta raouiensis, HUMZ 110722, 88mm; C, Gnathanacanthus goetzeei, AMS I. 20188001, 47mm. EB, epibranchials; IAC, interarcual cartilage; PB, pharyngobranchial; TP, tooth plate on second epibranchial.

In the taxa with isolated second to fourth pharyngobranchials (e,g., Helicolenus, see Fig, 6A), the first and second epibranchials are connected with the second pharyngobranchial (directly or via interarcual cartilage), and the third and fourth epibranchials with the third and fourth pharyngobranchials, respectively. Each epibranchial in the taxa having character 43-1 or 43-2 is connected with the respec- tive pharyngobranchial element (Fig, 6C). Therefbre, the disappearance of the pharyngobranchial elements was not considered in this analysis. If a pharyngob- ranchial element(s) were to disappear, the corresponding epibranchial(s) is as- sumed to lose its connection with the former, the epibranchial becoming free from the pharyngobranchial, TS 44. Medial extrascapular: O, present; 1, absent, In the Serranidae examined, the medial extrascapular is typically present. Although this element is also pres- ent tn Sebastiscus (Fig. 7A), Sebastes otvstont, and Helieotenus percoides, it is absent in most scorpaenoids (Fig. 7B-C). TS 45. Lateral extrascapular: O, single element with three sensory openings; 1, two elements (horizontal and longitudinal tubes) with two sensory openings; 2, sin- gle element (longitudinal tube) with two sensory openings (unordered), The lateral extrascapular has the form of a single element with three sensory openings in the Serranidae examined and many scorpaenoids (Fig, 7A). Character 45-1 occurs in ECtreposebastes, except on the right side of a specimen of E imus (HUMZ 75112) showing eharacter 45-O, Gnathanacanthidae, Pataecidae (Fig. 7C), and the right side of a specimen of 7Ietraroge niger, and character 45-2 in Synanceiidae (except Erosa and Choridactl,lus) (Fig, 7B), Aploactinidae, Congiopodidae, and the right side of a specimen of Tetrarage niger. The condition of Erosa is unclear, the bone being firmly sutured and the number ef openings net confirmed (thus coded as "?))).

TS 46. Cleithrum and coracoid: O, without ventromedial connection; 1, with

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16 Hisashi lmamura

A

-V, 1' zz,,- ttt - /-

3mm 3mm 3mm

Fig. 7. Lateral view of medial and lateral extrascapulars, and posttemporal. A, Sebastiscus

marmoratus, HUMZ 69895, 125mm; B, Inimicusjcrponicus, HUMZ 140587, 143mm; C, Pataec"s

fronto, AMS I. 38479002, 110mm. LE, lateral extrascapular; ME, medial extrascapular; PT, posttempora].

partial ventromedial connection; 2, with complete ventromedial connection (or- dered as O-1-2). The cleithrum is not connected ventromedially with the ceracoid in the Serranidae examined. Character 46-1 is present in Minous, and character 46-2 in . TS 47. Uppermost actinost and scapula: O, separated; 1, fused. The uppermost actinost is separated firom the scapula in Serranidae (e.g., Katayama 1959; Baldwin and Johnson 1993). These elements are fused in Sebastidae (except Sebastolobus), Scorpaena, ScorpaenQpsis, Apistidae, Tetrarogidae, Apistidae, Synanceiidae,

Aploactinidae, Alertichthys, Gnathanacanthidae, Pataecidae, and Neosebastidae. TS 48. Number of postcleithra: O, two; 1, one; 2, zero (ordered as O-1-2). Two postcleithra are present in Serranidae (e.g., Katayama 1959), Character 48-1 is fbund in Triglidae, Satyrichtbys, and Hoplichthyidae, and character 48-2 in Patae- cidae. In Platycephalidae, all three conditions are found (e.g., two postcleithra in Ratabulus, one in , and none in Elates). TS 49. Number of firee lower pectoral fin rays: O, zero; 1, one; 2, two; 3, three or more (ordered as O-1-2-3). There are no free lower pectoral fin rays in the Ser- ranidae examined, Character 49-1 is found in Apistidae and Minous, character 49-2 in inimicus and Peristediidae, and character 49-3 in Choridactylus, Triglidae, and Hoplichthyidae. TS 50. Branched pectoral fin rays: O, present; 1, absent. In the Serranidae ex- amined, at least some of the pectoral fin rays are branched. Pectoral fins with only unbranched rays are found in Taenianotus, Pontinus, Pterois, Minous, Aploac-

tinidae, Congiopodidae, Gnathanacanthidae, and Pataeeidae. TS 51. Cartilaginous caps on anterior portion of pelvis: O, separated; 1, fused. The cartilaginous caps on the anterior portion of the pelvis are separated in the Serranidae examined, as well as in most scorpaenoids. Fusion is evident in Trigli- dae (except Lqpidotrigla and PteT:ygotrigta), Peristediidae, and Hoplichthyidae. TS 52. Posterior pelvic fbssa: O, absent; 1, present, opposing fbssae meeting; 2, present, opposing fossae separated (unordered). Posterior pelvic t'ossae (sensu Ima- mura 1996) are absent in the Serranidae examined. Opposing fossae meet in Plec-

trogeniidae and Parabembridae, but are separate in Bembridae, Triglidae, Periste- diidae, Hopliehthyidae, and Platycephalidae (see Imamura 1996, fig, 5). Fossae are

absent in the other scorpaenoids.

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Phylogeny of Scorpaenoidea 17

TS 53. 0pposing pesteromedial parts of pelvis: O, sutured; 1, separated. The op- posing posteromedial parts of the pelvis are sutured in the Serranidae (e.g,, Katayama 1959) and most ingroup taxa, but separated in Peristedion, Hoplichthy- idae, and Platycephalidae (see Imamura 1996, fig. 5). TS 54. Number of pelvic fin rays: O, six; 1, five; 2, four; 3, none (ordered as O-1-2- 3). Pelvic fin rays number six (one spine and five soft rays) in Serranidae (e,g,, Katayama 1984) and most scorpaenoids, five in Erosa, four in Aploactinidae, and zero in Pataecidae. TS 55. Branched pelvic fin rays: O, present; 1, absent. The pelvic fin rays are branched in Serranidae, but unbranched in Minous, Aploactinidae, Alertichthys, and Gnathanacanthidae. Pelvic fin rays are absent in Pataecidae and thus are "?". coded as TS 56. Number of spines on first proximal pterygiephore: O, two; 1, one. The first dorsal fin proximal pterygiophore supports two spines in Serranidae, except for several epinephelines (e.g., ]EZpinephelus and Liopropoma) (e.g., Kendall 1976), in addition to most scorpaenoids. Character 56-1 is apparent in Minous, Con- giopodidae, Triglidae (except Prionotus and Bellator), Peristediidae, Hoplichthy- idae, and Platycephalidae (see Imamura 1996, fig. 34), TS 57. Dorsal spines: O, stout; 1, slender. Weak, slender spines are restricted to Peristediidae and Hoplichthyidae, whereas they are strong and stout in the other scorpaenoids and Serranidae examined, TS 58. First spine on first anal fin proxlmal pterygiophore: O, present; 1, ab- sent. Three anal spines (two on the ventral edge of the first anal fin proximal ptery- giophore and one on the edge of the second pterygiophore) usually occur in Ser- ranidae, although the epinephelines Betonoperca and DiptQprion have just two spines (the first spine is considered to be absent in the former, and the third spine reduced to a soft ray in the latter) (e,g,, Katayama 1984; Nelson 1994; Baldwin and Smith 1998; present study). The first spine on the first pterygiophore is absent in Aploactinidae, Congiopodidae, Pataecidae, Bembradium, Bembridae, Triglidae, Peristediidae, Hoplichthyidae, and Platycephalidae. TS 59. Second element on first anal fin proximal pterygiophore: O, spine; 1, soft ray; 2, absent (unordered). Character 59-1 is tbund in Bembridae, Triglidae, Peri- stediidae, and Hoplichthyidae, and character 59-2 in Pataecidae, Bembradium, and Platycephalidae. See TS 58 fbr the serranid conditien, TS 60. Ray on second anal fin proximal pterygiophore: O, spine; 1, soft ray. Character 60-1 is fbund in Synanceiidae (except Synanceia and Erosa), Aploac- tinidae, CongiQpodus, Bembradium, Bembridae, Triglidae, Peristediidae, Ho- plichthyidae, and Platycephalidae. See TS 58 for the serranid condition, TS 61. Branched dorsal and anal fin rays: O, present; 1, absent. The dorsal and anal fins have branched soft rays in the Serranidae examined and most scor- paeneids. Character 61-1 is present in Mtnous, Aploactinidae, Alertichthys, Gnathanacanthidae, and Pataecidae. TS 62. Number of rays supported by posteriormost proximal pterygiophores: O, two; 1, one, The posteriormost proximal pterygiophore of both the dorsal and anal fins supports two rays in the Serranidae examined. Character 62-1 is evident in Tetrarogidae, inimicus, Choridactytus, Minous, Erisphex, Gnathanacanthidae,

Pataecidae, and Peristediidae, TS 63. Anterior dorsal proximal pterygiophores: O, separated from neurocra-

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18 Hisashi Imamura

nium; 1, sutured to neurocranium. The anterior dorsal proximal pterygiophores are separated from the neurocranium in most scorpaenoids, as well as in Ser- ranidae (Katayama 1959; Gosline 1966; Baldwin and Johnson 1993). Character 63-1 is restricted to Tetrarogidae and Erisphex, TS 64. Dorsal fin proximal pterygiophores: O, not exposed; 1, laterally exposed along first dorsal fin; 2, laterally exposed along both dorsal fins (ordered as O-1-2), The dorsal fin proximal pterygiophores are not exposed laterally in the Serranidae examined and most scorpaenoids. Character 64-1 is apparent in , and

character 64-2 in Chelidontchtltys and Lopidotrigta. TS 65. Number of supraneurals: O, three; 1, two; 2, one; 3, zere (ordered as O-1-2- 3). Within Serranidae, Anthiinae has two or three supraneurals, Serraninae three, and Epinephelinae zero to three (Kendall 1976), Character 65-1 is found in Parabe- mbridae, character 65-2 in Sebastidae, Pontinus, Setarches, and a single specimen of Bembrasjoponica (HUMZ 49409), and character 65-3 in the remaining taxa exam- ined. TS 66. Dorsal fin stay: e, present (separate and ossified); 1, fused with proximal pterygiophore; 2, present (cartilage); 3, absent (unerdered), The dorsal and anal fin stays are apparent as separate ossified elements in the Serranidae examined as well as in many ingroup taxa. In typical scorpaenoids, the suture lines between the proximal pterygiophore and dorsal and anal fin stays are below or level with the base of the last fin rays. Such is not the case in Gnathanacanthidae, although a bony element extends posteriorly to the base; this is interpreted as a fusien of the proximal pterygiophore and stay (character 66-1). Character 66-2 is apparent in a specimen of Erisphex potti (HUMZ 108774) and Plectrogeniidae, and character 66-3 in 7keenianotus, Tetrarogidae, Synanceiidae (except Erosa), Aploactis, Pataecidae, and Satyrichtlrys. TS 67. Anal fin stay: O, present (separate and ossified); 1, fused with proximal pterygiophore; 2, present (cartilage); 3, absent (unordered). See above (TS 66) for the serranid condition and the identification of the anal fin stay, Character 67-1 is apparent in Gnathanacanthidae, character 67-2 in Ertsphex and Plectrogeniidae, and character 67-3 in Tetrarogidae, Synanceiidae (except Erosa), Aptoactis, Patae- cidae, and Peristediidae. TS 68. First and second hypurals: O, separated; 1, continuous. In the serranid caudal skeleton, the fo11owing autogenous elements are recognized: three to five hypurals [first and second fused in such as the anthiine Pseudoanthias and serra- nine ChelidQperca hirundinacea (but autogenous in the examined specimen of C. hirundinacea), and third and fourth fused in such as C hirundinacea (but not in the specimen examined), Pseudoanthias, and the epinepheline Eipinephetus areota- tus (but anterior portions not fused)], parhypural, hemal spines on second and third preura] centra, three epurals, urostyle, and single uroneural (e,g., Fujita 1990). Character 68-1 is seen in Sebastidae (except Tracdyscorpia), Scorpaenidae, Apistidae, Tetrarogidae, Synanceiidae, Aploactinidae, Cengiopodidae, Gnatha- nacanthidae, Pataecidae, Lepidotrigla, Peristediidae, Hoplichthyidae, and Platy- cephalidae. TS 69. Third and fburth hypurals: O, separated; 1, continuous. Character 69-1 is apparent in Sebastidae (except Trachyscor:pia), Scorpaena, Scoilpaenopsis, Ttienia- notus, Apistidae, Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodidae, Gnathanacanthidae, Pataecidae, Triglidae (except Prionotus), Peristediidae, Ho-

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Phylogeny of Scorpaenoidea 19

plichthyidae, and most platycephalids. In Prionotus, the elements are continuous ``?" anteriorly and separated posteriorly, being coded as in this study. In the Platy-

cephalidae, only Ratabulus has separated third and fourth hypurals. See TS 68 for the serranid condition, TS 70, Fifth hypural: O, present; 1, absent. The fifth hypural is absent in Tetrarogidae, Synanceia, Erosa, Erisphex, Congiopodidae, one specimen of Gnathanacanthus goetzeei (AMS I. 20188001), Pataecidae, Chelidonichtbys, and Peri- stediidae. Although Ishida (1994) reported Gnathanacanthus goetxeei (AMS IB, 668) as not having the fifth hypural, that element was recognized during reexamination

of the specimen, See TS 68 for the serranid ¢ ondition. TS 71. Lower hypural plate and parhypural: O, separated; 1, fused. The lower hypural plate and parhypural are fused in Apistidae, Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodidae, Gnathanacanthidae, Pataecidae, Peristediidae, and

Hoplichthyidae. See TS 68 fbr the serranid condition. TS 72. Hemal spine and third preural centrum: O, separated; 1, fused. The hemal spine and third preural centrum are fused in Synanceiidae (except Synan- ceia), Aploactinidae, Gnathanacanthidae, Pataecidae, Peristediidae, Hoplichthy- idae, and Platycephalidae (Elates). See TS 68 fbr the serranid condition. TS 73. Hemal spine and second preural centrum: O, separated; 1, fused. Charac- ter 73-1 is fbund in Gnathanacanthidae, Pataecidae, Peristediidae, and Ho- plichthyidae. See TS 68 for the serranid condition, TS 74. Urostyle and upper hypural plate: O, separated; 1, fused. Character 74-1 is found in Inimicus, Minous, Cengiopodidae, Pataecidae, Triglidae (except Priono- tus and Bellator), Peristediidae, and Hoplichthyidae. See TS 68 for the serranid condition. TS 75. Urostyle and lower hypural plate: O, separated; 1, fused. Character 75-1 is restricted to Gnathanacanthidae and Pataecidae. See TS 68 for the serranid con- dition. TS 76. Uroneural: O, present; 1, absent. The uroneural is absent in Jltimicus, Congiopodidae, one specimen of Gnathanacanthus goetzeei (AMS IB. 668), Pataeci- dae, Chelidonichtbys, and Peristediidae. See TS 68 for the serranid condition. TS 77. Number of epurals: O, three; 1, two; 2, one (ordered as O-1-2). Two epu- rals are found in Synanceia, Erosa, one specimen of Inimicus japonicus (HUMZ 79103), and Congiopodidae, and a single epural in Pataecidae only. See TS 68 for the

serranid condition. TS 78. Branched caudal fin rays: O, present; 1, absent. The caudal fin rays are branched in most ingroup taxa, as well as in the serranids (e.g, Johnson 1984), Character 78-1 is fbund in Minous, Aploactinidae, Alertichthys, Gnathanacanthi-

dae, and Pataecidae, TS 79. Posterier end of adductor mandibulae section 1: O, connected to preop- ercle (and hyomandibula); 1, connected only to hyomandibula; 2, free from poste- rior bony element(s) (unordered). The adductor mandibulae section 1 (Al) of the Serranidae examined is connected to the preopercle (and hyemandibula), as in most ingroup taxa, Character 79-1 is present in Gnathanacanthidae and Pataeci- dae, and character 79-2 in Triglidae and Peristediidae, TS 80. A16: O, absent; 1, present. In the Serranidae examined, Al is not divided into subsections, but Alfi is recognized in Synanceia, Erosa (Fig. 8B, D), Gnathanacanthidae, and Maxilltcosta. In Gnathanacanthidae, Alfi is further di-

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20 Hisashi [mamura

vided into Alfi' and Alfi", which condition can be regarded as an autapomorphy of

this taxon. TS 81. 0rigin of A2-3: O, lateral to levator arcus palatini; 1, partially medial to levator arcus palatini; 2, completely medial to levator arcus palatini; 3, not closely associated with levator arcus palatini (unordered). A2-3 is located lateral to the lev- ator arcus palatini in the Serranidae examined and many scorpaenoids. Character 81-1 is found in Apistidae, Synanceiidae (Fig. 8B, D), Aploactinidae, Congiopodi- dae, Gnathanacanthidae, and Pataecidae, character 81-2 in Tetrarogidae (Fig. 8A, C), and character 81-3 in Taenianotus, Although Ishida (1994) stated that A2-3 of Aploactis is located medial to the lev- ator arcus palatini (i.e., character 81-2 herein), character 81-1 was found in the A. aspera examined in this study. Ishida (1994) also recognized Pataecidae as having

A2-3 originating from the neurocranium, but did not comment Qn the relationship of A2-3 to the levator arcus palatini. It was confirmed in the present study that the posterodorsal portion of A2-3 originates from the neurocranium [as diseussed by Ishida (1994)], the muscle also being partially medial to the levator areus palatini (see also Ishida 1994, fig. 3D). A2-3 originating from the neurocranium is an autapo- morphic condition in the Pataecidae.

/ B

A '-

x 3mm × 5mm

biig. 8. Lateral view of cheek and other cephalic muscles: A and B, superficial aspect; C and

D, lateral aspect after removal of superficial muscles. A and C, Tetraroge niger, URM-P 31819,

18mm; B and D, Erosa erosa, HUMZ 64250, a3mm. Al, AIP, and A2-3, sections of adductor

mandibulae; AAP, adductor arcus palatini; DO, dilatator operculi; LAP, Ievator arcus pala-

tini; I,O, levator operculi.

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Phylogeny of' Scorpaenoidea 21

TS 82. Position of adductor arcus pa]atini: O, dorsal surface of entopterygoid; 1, medial margin of entopterygoid; 2, ventral surface of entopterygoid (ordered as O-1- 2). The adductor arcus palatini is situated on the ventral surface of the entoptery- goid in the Serranidae examined, as well as in most scorpaenoids, Character 82-1 is apparent in C7zoridactylus, Aploactinidae, Congiopodidae, Gnathanacanthidae, Pataecidae, and Triglidae, and character 82-2 in Synanceia, Erosa, and Minous. In "?", Hoplichthyidae, the entopterygoid is absent, and this character is coded as TS 83. 0rigin of levator operculi: O, pterotic; 1, pterotic and posttemperal; 2, posttemporal (ordered as O-1-2). The levator operculi originates only from the pterotic in the Serranidae examined and most ingroup taxa. Character 83-1 is ap- parent in Taenianotus, Pterois, Dendrochirus, Apistidae, Tetrarogidae (Fig, 8A, C), Synanceiidae (Fig, 8C, D), Aploactinidae, Atertichtttys, and Gnathanacanthidae, and character 83-2 in Congiqpodus, TS 84. Hyohyoides inferioris: O, absent; 1, present, The hyohyoides inferioris is absent in the Serranidae examined and also in Sebastiscus, Scorpaenodes, Ponti-

nus, Trachyscorlpia, Setarchidae, Apistidae, Congiopodidae, Neosebastidae, Parabembridae, HOptichtdys gilberti, H langsdoizfii, and some p]atycephalids (e.g., ; see Imamura 1996, fig. 41). Character 84-1 is apparent in the other scor- paenoids. TS 85. Anterior portion of transversus dorsalis anterior: O, branched; 1, un- branched. The anterior portion of the transversus dorsalis anterior is branched in the Serranidae examined. An unbranched transversus dorsalis anterior is fbund in Apistidae, Choridactylus, Minous, Aploactinidae, Maxitlicosta, Plectrogeniidae, Triglidae, Peristediidae, Hoplichthyidae, and Platycephalidae. TS 86. Posterior portion of transversus dorsalis anterior: O, unbranched; 1, branched, not continuous with anterior branch of same muscle; 2, branched, con- tinuous with anterior branch of same muscle, comprising circular muscle ele- ments (ordered as O-1-2). Character 86-1 is seen in Pataecidae (Fig. 9B), and charac-

BTDA c

O

,

LE

LEN 3mm 3mm 3mm

Fig. 9. Medial view of upper branchial museles. A, Sebastes owstoni, HUMZ 42641, 183 mm; B,

Pataecusf}"onto, AMS I. 38479002, 110mm; C, Alertichtdys blacki, HUMZ 66621, 161 mm. AD, ad-

ductor; LE, levator externus; LEP, levator posterior; LI, levator internus; OD, obliquus dor-

salis; RD, retractor dorsalis; SO, sphincter oesophagi; TDA, transversus dorsalis anterior;

TDP, transversus dorsalis posterior.

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22 Hisashi Imamura

ter 86-2 in Congiopodidae (Fig, 9C) and Gnathanacanthidae, whereas character 86-O is fbund in the other scorpaenoids (Fig. 9A) as well as the Serranidae examined. TS 87. 0bliquus dorsalis II: O, absent; 1, present. The obliquus dorsalis II is ab- sent in the Serranidae examined. Character 87-1 is restricted to the Setarchidae. TS 88. Posterior levator internus: O, lateral to obliquus dorsalis; 1, sandwiched by obliquus dorsalis. The posterior levator internus is located lateral to the obliquus dorsalis in the Serranidae examined. Character 88-1 is fbund in Apisti- dae, Erisphex, and Pataecidae (Fig. 9B). Ishida (1994) recognized Aploactis as also having character 88-1, but this could not be confirmed, owing to the damaged upper gill arch of the specimen of Aploactts examined in this study (therefore coded as Llt.J77).

TS 89. Levator externus III: O, present; 1, absent. The levator externus III is ab- sent in Tetrarogidae, Synanceia, Erosa, inimicus, Aploactinidae, Pataecidae (Fig. 9B), IVeosebastes, Hoplichthyidae, and some Platycephalidae (e.g., Suggrundus meerdervoortii and inagocia japonica). This muscle is present in the Serranidae ex- amined and other scorpaenoids. TS 90. Levator pesterior: O, present; 1, absent. Character 90-1 is observed in the Cengiopodidae (Fig. 9C), Bembradium, and Hoplichthyidae, whereas the other in- group taxa and the Serranidae examined show character 90-O. TS 91. Adductores I-III: O, absent; 1, present. Character 91-O is found in the Ser- ranidae examined and most scorpaenoids, whereas character 91-1 is restricted to Congiopodidae (Fig. 9C), as pointed out by Ishida (1994), TS 92. Transversus ventralis anterior and posterior: O, overlapping; 1, sepa- rated, The transversus ventralis anterior and posterior overlap in the Serranidae examined, as well as in most scorpaenoids (Fig. 10A, B), whereas they are sepa- rated in 7benianotus, Pterois, Congiopodidae (Fig. 10C), Gnathanacanthidae, Pataecidae, Triglidae, and Peristediidae. TS 93. Transversus ventralis posterior: O, without a tendon anteriorly; 1, with a tendon anteriorly. The transversus ventralis does not have an anterior tendon in Apistidae (Fig, 10B), Tetrarogidae, Synanceiidae, Aploactinidae, CongiQpodus, Pataecidae, Neosebastidae, Plectrogenium, Parabembridae, Bembridae, Hopli-

A B c

n

PHE PHI ! 1/ X

5mm 3mm 3mm

Fig. 10. Ventral view of lower branchial muscles, A, Setarches longimanus, HUMZ 79505, 125

mm; B, Apistus carinatus, HUMZ 37372, 93mm; C, Atertichthys blacki, HUMZ 66621. 161 mm.

PHE, pharyngoclavicularis externus; PHL pharyngociavicularis internus; RC, rectus com-

munis; RV, rectus ventralis; TVA, transversus ventralis anterior; TVP, transversus ventralis posterlor.

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Phylogeny of Scorpaenoidea 23

chthyidae, and Platycephalidae, nor in the Serranidae examined, whereas it has the tendon anteriorly in the other scorpaenoids (Fig. 10A, C). TS 94. Protractor pectoralis: O, sheet-like muscle only; 1, composed of sheet- like anterior and robust posterior elements. The pretractor pectoralis comprises sheet-]ike anterior and robust posterior elements in Iloptichtlays gitberti, H langs- dor:fii, and Platycephalidae. In the other scorpaenoids and the Serranidae exam-

ined, character 94-O exist.

TS 95. Division of bundles of adductor superficialis serving firee pectoral fin rays: O, absent; 1, present. The bundles of the adductor superficialis serving each fin ray are not subdivided in the Serranidae examined and most scorpaenoids. Character 95-1 is apparent in inimicus, Minous, Choridactytus, Triglidae, and Peri- stediidae. TS 96. 0rigin of coracoradialis: O, posteromedial face of posterier process of coracoid; 1, posteromedial face of posterior process of coracoid and posterolateral face of posteroventral precess of cleithrum. In the Serranidae examined and most scorpaenoids, the coracoradialis connects the lowermost actinost and the postero- medial face of the posterior process of the coracoid. Character 96-1 is seen in Chori- dacCylus, Aploactinidae, and Atertichtbys. Although Ishida (1994) stated that the coracoradialis is absent in Apistus and Gnathanacanthus, a tiny ceracoradialis was found during the present study [including the right side of a specimen of Gnathanacanthus goetzeei examined by Ishida (1994) (AMS IB. 668, element on left side not confirmed owing to previous dissection)]. TS 97. Dorsal elements of pelvic fin muscles: O, not attached to pectoral girdle; 1, attached to pectoral girdle. In the Serranidae examined, the dorsal elements of the pelvic fin muscle (adductor superficialis pelvicus, adductor profundus pelvi- cus, and extensor proprius) are not attached to the pectoral girdle. Character 97-1 is found in Triglidae, Peristediidae, and Hoplichthyidae (except HOplichthys haswelli). TS 98. Extensor proprius: O, present; 1, absent. The extensor proprius is found in the Serranidae examined, and also in most scorpaenoids. This muscle is absent in Aploactinidae, Alertichthys, and Pataecidae. Ishida (1994) stated that the exten- sor proprius is present and that the adductor superficialis pelvicus and adductor profundus pelvicus are united in Alertichthys and Zanclorhynchus; however, pres- ent observations confirmed that the latter two muscles are separated, and the ex- tensor proprius absent, in Alertichthys, In this instance, muscle identification was based on their fin ray insertions (inserted onto several inner rays-extensor pro- prius vs. all rays including spine-adductor superficialis pelvicus) (Winterbottom 1974; present study), It is suspected that Ishida (1994) identMed the adductor super- ficialis pelvicus as the extensor proprius. TS 99. Flexor ventralis externus: O, present; 1, absent. Character 99-1 is found tn Synanceia, Erosa, Congiopedidae, Pataecidae, and Triglidae, with character gg-o occurring in the other scorpaenoids and the Serranidae examined. TS 100. Adductor dorsalis: e, present; 1, absent, In the Serranidae examined, the adductor dorsalis is absent in the epinephelines DipIQprion and Niphon, whereas it is present in the others. Character 1oo-O is found in Sebastidae, Seor- paenidae, Setrachidae, Apistidae, Tetrarogidae, a specimen of Gnathanacanthus goetzeei (AMS IB. 668), Neosebastidae, Plectrogeniidae, Parabembridae, and Bem- bridae, and character leO-1 in the remaining taxa.

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24 Hisashi Imamura

TS 101. Extrinsic muscle: O, absent; 1, present, connected to neurocranium an- terierly and swimbladder pesteriorly; 2, present, connected to neurocranium ante- riorly and vertebrae posteriorly; 3, present, free from neuroeranium anteriorly and connected to vertebrae posterierly (unordered). There is no extrinsic muscle derived from the obliquus superioris in the Serranidae examined nor in Cheli-

donichthys, Lepidotrigla, or I'rionotus. Character 101-1 is found in Sebastiscus,

Scorpaenodes, Pterois, Dendrochirus, and Apistidae, character 101-3 in Ttienia-

notus, IVeosebastes, and Bellator, and character 101-2 in the remaining taxa. TS 102. Intrinsic muscle: O, absent; 1, present. The intrinsic muscle is present in Apistidae and Triglidae (except Ptenygotrigla), whereas it is absent in other in- group taxa, as well as in Serranidae, TS 103. 0bliquus superioris: O, extending to neurocranium; 1, not extending to neurocranium, In the Serranidae, the obliquus superioris extends to the neurocra- nium. Character 103-1 is recognized in Svnanceia, Erosa, Congiopodidae, Ptery- gotrigla, Peristediidae, Hbpiichtdys haswetli, and IL langsdot:tii. TS 104. 0bliquus superioris and Baudelot's ligament: O, ebliquus superioris penetrated by Baudelet's ligament; 1, obliquus superioris bypassing and ]ying ven- trally to Baudelot's ligament. The obliquus superioris is penetrated by Baudelot's ljgament in Serranidae, Character 104-1 is found in Scoilpaenopsis, Taenianotus,

Scoizpaenodes, Pterois, Dendrochirus, Setarchidae, Apistidae, Tetrarogidae, Synan- ceiidae, Aploactinidae, Gnathanacanthidae, Pataecidae, Triglidae, and Hbplichthys gilberti. Iii Synanceia, Erosa, Congiopodidae, Pteilygotrigla, Peristediidae, Hb- plichtirys haswelli, and H. Iangsdor:fii, the obliquus superioris is absent (also Baudelot's ligament in Congiopodidae and Il langsdot:fii), and the association of

"?"). the muscle and ligament is uncertain (thus coded as TS 105. Supracarinalis anterior: O, present; 1, absent. The supracarinalis ante- rior is absent in Tetrarogidae, Aploactinidae, and Pataecidae, but present in the other scorpaenoids and the Serranidae examined. TS 106. Inclinator dorsalis associated with first dorsal spine: O, present; 1, ab- sent. An inclinator dorsalis associated with the first dorsal spine is present in the Serranidae examined. On the other hand, it is absent in Apistidae, Tetrarogidae,

Synanceiidae, Aploactinidae, Pataecidae, Maxitlicosta, and Parabembridae. In taxa without a first dorsal spine (see TS 56), the condition of the muscle was not deter- "`?"). mined (thus coded as TS 107. Gill membranes: O, free from isthmus; 1, narrowly fused with isthmus, gil1 opening wide; 2, broadly fused with isthmus, gil1 opening narrow (ordered as O- 1-2). In the Serranidae examined, the gil1 membranes are free from the isthmus, Character 107-1 is fbund in Synanceiidae, Peristediidae, and Hoplichthyidae, and

character 107-2 in Congiopodidae.

TS 108. Sensory ducts in : O, simple; 1, with two or more branches. The sensory ducts in the lateral line scales are simple in the Serranidae examined. Character 108-1 occurs in Pterois, Dendrochirus, Chelidonichtdys, Lopidotrigla, and some platycephalids (e.g,, Thysanophrlys, Onigocia, and Rogadius). TS 109. Lateral line: O, with scales; 1, with tube-like bones; 2, with bony plates (unordered), Character 109-1 oceurs in Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodidae, Gnathanacanthidae, and Pataecidae, and character 109-2 in Peri-

stediidae and Hoplichthyidae. In the other scorpaenoids and Serranidae examined the lateral line has scales.

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Phylogeny of Scorpaenoidea 25

TS 110. Spines on lateral Iine s ¢ ales: O, absent; 1, present, The lateral line scales lack spines in the Serranidae examined. Character 110-1 is apparent in Ap}oactinidae, Alertichthys, Peristediidae, Hoplichthyidae, and some platy- cephalids (e. g., Grammoptites and Solitas). TS 111. Body scales: O, present; 1, absent except in dorsal region; 2, entirely ab- sent; 3, present as bony plates (unordered), Character 111-1 is observed in HO- plichtdys hasivelli, character 111-2 in Tetrarogidae, Synanceiidae, Congtopodus, Gnathanacanthidae, Pataecidae, HOplichth.vs gtlberti, and lt langsdoi:fii, and char- acter 111-3 in Peristediidae. The body is covered with scales in the other ingroup taxa, as well as in the Serranidae examined, Although Nakabo (2002a) considered body scales to be absent in Taenianotus, embedded scales were fbund in this during the present study. This charaeter is an autapomorphy of [ltzenianotus.

Phylogenetic relationships Analysis of the data matrix fbr the Scorpaenoidea (Table 1) using PAUP* 4.0blO resulted in three equally most parsimonious trees (TL=419, CI=O.37), The different topologies resulted from the placement of the three serranid subfamilies included as outgroups, whereas the relationships of the ingroup taxa were repre- sented by a single tree (Fig. 11), Two characters, the suborbital stay with its end pointed (character 8-1) and the presence of spines associated with the parietal sen- sory canal (character 24-1), occurred at the ingroup node. In addition to four synapomorphies of the Scorpaenoidea, the monophyly of this superfamily is addi-

tionally supported by two further synapomorphies, characterized by successive conditions (reversal or change to another derived character): characters 47-1 (up- permost actinost and scapula fused) and 93-1 (transversus ventralis posterior with a tendon anteriorly). Only the presence, but not the condition of TS 101 (extrinsic muscle) at the ingroup node was confirmed. At this node, the muscle was either connected posteriorly with the swimbladder (character 101-1) or with vertebrae (101-2) fbllowing analysis by ACCTRAN, whereas characters 101-1 and 101-2 charac- terized clactes IA and IB, respectively, when DELTRAN was employed. Characters "r" labeled indicate reversals. Both unambiguous characters, which do not change

the evolutionary interpretation, and the ambiguous characters, in which the inter- pretation is changed when different character optimization (ACCTRAN or DEL- TRAN) is employed, are listed below. Clade IA. Sebastiscus-although this clade has no derived characters accord- ing to ACCTRAN, it is supported by character 101-1 according to DELTRAN. Clade IB. All ingroups minus Sebastiscus-this clade is supported by charae- ter 84-1 (also 44-1 according to ACCTRAN and 101-2 according to DELTRAN), Clade 2A. Sebastes-this clade has no derived characters. Clade 2B. All scorpaenoids minus Sebastiscus and Sebastes-clade 2B is sup- ported by character 8-2 (also 44-1 a ¢ cording to DELTRAN). Clade 3A. Adelosebastes-this clade has no apomorphic characters. Clade 3B. -altheugh this clade had no derived characters arnong those included in the analysis, Hozukius has an autapomorphic character, honey- comb-like concavities on the posterodorsal surface of the neurocranium (Matsu- bara 1934, 1943; present study). Clade 3C. All scorpaenoids minus Sebastiscus, Sebastes, Adelosebastes, and Hbzukius-this clade is supported unambiguously by character 31-1.

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26 HisashiImamura

tA Sebesttseus Sabasas Adelomatas Harukins Hollootenus setretstolotus Scotpaetta Sco,paenopsis Teenienotus Seotpsenodes Pterots Dend,vchbus

Ponthas

Ttl" tyscorPla Seterches Ectreposebastes Aptstus retrervge Synanceia Erose inimicus Choridectytus MVnous Aptaactls Etisphex Conglopadus Alartichthys Gnathensaenthus Pateecus tuax"deosta Ploctrogenium Bemt"edium Perabembras Bembras

Chelldonk hthys LepldOtriig1ta PtetygebtDla Ptlonotus

Betiator Peristection Setyrk"ithys Hopdehtirys Platyceptalldee 26B

Fig. 1]. Phylogenetic reLationships of the supert'amily Scorpaenoidea. Nunibers assigned to clades correspond to those in text.

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Phylogeny of Scorpaenoidea 27

Clade 4A. HOIicotenus-this clade has no derived characters. Clade 4B. All scorpaenoids minus Sebastidae, except for Sebastiscus and Tra- chyscorpia-this clade shares unambiguous characters 7-1 and 8-3. Clade 5A. Sebastolobus--clade 5A is supported unambiguously by character 47-Or.

Clade 5B. All scorpaenoids minus Sebastidae, except fbr Trachyscoilpia-this clade is supported unambiguously by characters 9-1 and 65-3, Clade 6A. Scorpaenidae, T}'achyscorzpia, and Setarchidae-this clade is sup- ported unambiguously by characters 6-1 and 11-1. Clade 6B. All scorpaenoids aside from Sebastidae, Scorpaenidae, and Setar- chidae--this clade is supported by characters 31-Or and 93-Or. Clade 7A. Sconpaena-this clade has no apomorphic characters, Clade 7B. Scorpaenidae minus Scoilpaena, and Trachysconpia and Setarchi- dae-this clade is supported by characters 30-1 and 104-1 (also 10-1 according to AC- CTRAN), Clade 8A. Scorpaenqpsis-this clade is supported by character 38-1 (also 10-1 according to DELTRAN), Clade 8B. Scorpaenidae minus Scoilpaena and Scor:paenopsis, and 7'rachyscor- pia and Setarchidae-this clade is supported unambiguously by characters 12-1 and 47-Or, Clade 9A. Taenianotus-this clade is supported by eharacters 16-2, 25-1, 32-1, 33-1, 50-1, 66-3, 81-3, 83-1, 92-1, and 101-3 (also 10-1 according to DELTRAN). This clade also has embedded bedy scales as an autapomorphy. Clade 9B. Scorpaenidae minus Sconpaena, Scoizpaenopsis, and Ttienianotus,

and Traclayscorpia and Setarchidae-this clade is supported by characters 31-Or and 69-O (also 10-Or and 84-Or according to ACCTRAN). Clade 10A. Scorpaenodes, Pterois, and Dendrochirus-this clade is supported

unambiguously by characters 3-3, 7-Or, and 101-1. Clade 10B. Pontinus, Trachyscorpia, and Setarchidae--clade 10B is supported by characters 30-Or and 65-2r (also 104-Or according to ACCTRAN and 84-Or accord-

ing to DELTRAN). Clade 11A. Scotlpaenodes-this clade has no derived characters according to

ACCTRAN but is supported by character 84-er according to DELTRAN. Clade 11B. Pterois and Dendrochirus-this clade is supperted by characters 9- 2, 35-1, 83-1, and 108-1 (also 84-lr according to ACCTRAN), Pterois is supported un- ambiguously by characters 25-2, 50-1, and 92-1, whereas Dendrochirus has no de- rived characters. Clade 12A. Pontinus-this clade is supported by characters 42-1 and 50-1 (also 104-Or according to DELTRAN). Clade 12B. Trachysconpia and Setarchidae-clade 12B is supported unambigu- ously by characters 19-1 and 68-O. Clade 13A. Trachyscorlpia-this clade has no characters according to ACC- TRAN but is supported by character 104-Or according to DELTRAN. Clade 13B. Setarchidae (Setarches+EctreposebastesFthis clade is supported by characters 9-2, 10-1 (reversed according to ACCTRAN), and 87-1 (also 104-lr ac- cording to ACCTRAN). Within this clade, Setarches has no derived characters,

whereas Ectreposebastes has unambiguous character 65-3r. According to Ishida (1994), Ectreposebastes has an autapomorphy, a bifid pharyngoclavicularis inter-

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28 Hisashi Imamura

nus, although this condition could not be confirmed in the specimens examined. Clade 14A. Apistidae, Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodi- dae, Gnathanacanthidae, and Pataecidae-this clade is supported by characters 10- 1, 32-1, 71-1, 81-1, 83-1, 104-1, and 106-1. Clade 14B. Neosebastidae, Plectrogeniidae, Parabembridae, Bembridae, Trig-

lidae, Peristediidae, Hoplichthyidae, and Platycephalidae----this clade is supported

unambiguously by characters 3-2, 68-O, and 69-O. Clade 15A. Apistidae (1tpistus)-clade 15A is supported unambiguously by characters 13-1, 19-2, 27-1, 49-1, 84-Or, 85-1, 88-1, 101-1, and 102-1. Clade 15B. Tetrarogidae, Synanceiidae, Aploactinidae, Congiopodidae, Gnathanacanthidae, and Pataecidae--this clade is supported by characters 40-1, 42- 1, 45-2, 66-3, 67-3, 70-1, 89-1, 109-1, and 111-2 (also 33-1, 62-1, and 72-1 according to ACC- TRAN). Clade 16A. Tetrarogidae (7letraroge)-this clade is supported by characters 23- 1, 24-Or, 63-1, 81-2, and 105-1 (also 33-1 and 62-1 according to DELTRAN). Clade 16B. Synanceiidae, Aploactinidae, Congiopodidae, Gnathanaeanthidae, and Pataecidaorthis clade is supported by characters 25-2, 30-1, 43-1, 82-1, and 100-1 (also 72-1 according te DELTRAN). Clade 17A. Synanceiidaerthis clade bears apemorphic character 107-1 (also 33-1 according to DELTRAN), Clade 17B. Aploactinidae, Congiopodidae, Gnathanacanthidae, and Pataeci- daorthis clade is supported by characters 20-1, 37-1, 50-1, 55-1, 58-1, 61-1, and 78-1 (also 33-Or and 98-1 according to ACCTRAN). Clade 18A. Synanceia and Erosa-clade 18A is supported by characters 77-1, 80-1, 82-2, 99-1, and 103-1 (also 62-Or according to ACCTRAN), Synanceia has unam- biguous characters 10-2, 19-2, and 72-Or, and Erosa, 21-1, 54-1, 66-Or, and 67-Or. Clade 18B. Inimicus, Choridactylus, and Minous-this clade is supported by characters 38-1, 60-1, 70-Or, and 95-1 (also 49-2 and 74-1 according to ACCTRAN and 49-1 and 62-1 according to DELTRAN). Clade 19A. inimicus this clade is supported by characters 43-Or, 76-1, and 82- Or (also 49-2 and 74-1 according to DELTRAN), Clade 19B. Ohoridactylus and Minoustlade 19B is supported unambiguously by characters 46-1, 85-1, and 89-Or (also 16-1 or 16-2 according to ACCTRAN). Chori- dactylus is supported by characters 36-2, 45-Or, 46-2, 49-3, and 96-1 (also 74-Or accord- ing to ACCTRAN and 16-2 according to DELTRAN), and Minous by 13-1, 19-2, 21-1, 50-1, 55-1, 56-1, 61-1, 78-1, and 82-2 (also 49-lr according to ACCTRAN and 16-1 and 74- 1 according to DELTRAN). Clade 20A. Aploactinidae (Aptoactis+ErisphexFthis clade is supported un- ambiguously by characters 38-1, 54-2, 60-1, 85-1, 96-1, 105-1, 110-1, and 111-Or (also 88-1 according to ACCTRAN and 98-1 according to DELTRAN). Aploactis is character- ized by characters 10-2, 19-2, 43-2, and 70-Or

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Phylogeny of Scorpaenoidea 29

84-Or, 90-1, 91-1, 103-1, and 107-2 (also 62-Or according to ACCTRAN and 16-2, 74-1, 77- 1, 86-2, 89-Or, and 99-1 according to DELTRAN), Congiopodus has characters 10-Or, 28-1, 37-2, 47-Or, 55-Or, 60-1, 61-Or, 78-Or, and 83-2 (also 98-Or according to ACCTRAN

and 24-Or and 36-2 according to DELTRAN), and Alertichthys, 11-1, 12-1, 18-1, 38-1, 41- 2, 93-lr, 96-1, 110-1, and 111-Or (also 24-1 and 36-Or according to ACCTRAN and 98-1 according to DELTRAN), Clade 21B. Gnathanacanthidae (Gnathanacanthus) and Pataecidae (Patae- cus>･---this clade js supported by characters 1-1, 45-1, 73-1, 75-l, and 79-1 (also 24-Or, 36-2, and 62-1 according to DELTRAN), Gnathanacanthus has characters 20-Or, 37- Or, 58-Or, ua-1, 67-1, 80-1, 93-lr, and 106-Or (also 74-Or, 77-Or, 98-Or, and 99-Or according te ACCTRAN and 16-2, 86-2, and 89-Or according to DELTRAN), and Pataecus, 7-Or, 8-Or, 10-2, 18-1, 19-2, 48-2, 54-3, 59-2, 77-2, 83-Or, 88-1, and 105-1 (also 16-Or, 86-lr, and 89- lr according to ACCTRAN and 74-1, 98-1, and 99-1 according to DELTRAN). Gnathanacanthus also has an autapomorphic character, the presence of Alfi' and P". According to Ishida (1994), all genera of the Pataecidae (Pataecus, Aetqpcus, and IVeopataecus) have, as autapomorphic characters, A2-3 originating from the neuro- cranium and the absence of all pelvic fin muscle elements [Ishida's (1994) charac- ters 5-3, 23-1, 24-1, 25-1, 26-1, 28-1, and 29-1]. This family has five to 11 anal spines (Washington et al. 1984b), the increased number of spines being an autapomorphy of the family, Mandrytsa (2001) pointed out that Pataecidae has an autapomorphic extracleithrum (sensu Mandrytsa 2001). Clade 22A. Neosebastidae (IVeosebastestMaxiUicostaFthis clade is supported by character 84-Or. AJigosebastes is characterized unambiguously by characters 89-1 and 101-3, and Maxillicosta, by 22-1, 42-1, 80-1, 85-1, and 106-1, Clade 22B. Plectrogeniidae, Parabembridae, Bembridae, Triglidae, Peristedi- idae, Hoplichthyidae, and Platycephalidaorthis clade is supported by characters 47-Or and 52-1. Clade 23A. Plectrogeniidae (PZectrogenium+BembradiumFthis clade is sup- ported unambigueusly by characters 23-1, 66-2, 67-2, and 85-1, Plectrogenium has an unambiguous character 10-1, and Bembradtum, 16-1, 38-1, 41-1, 58-1, 59-2, 60-1, 90-1, and 93-lr. Clade 23B. Parabembridae, Bembridae, Triglidae, Peristediidae, Hoplichthy- idae, and Platycephalidaorthis clade is supported unambiguously by eharacter 9- or. Clade 24A. Parabembridae (ParabembrasFthis clade is supported unam- biguously by characters 26-1, 65-]r, 84-Or, and 106-1. Clade 24B, Bembridae, Triglidae, Peristediidae, Hoplichthyidae, and Platy- cephalidaexlade 24B is supported unambiguously by characters 52-2, 58-1, 59-1, and 60-1. Clade 25A. Bembridae (Bembras)-this clade has no apomorphic characters. Clade 25B. Triglidae, Peristediidae, Hoplichthyidae, and Platycephalidae- this clade is supported by characters 22-1, 39-1, 56-1, 85-1, and 100-1 (also 13-1, 38-1, 48-1, 53-1, 68-lr, and 69-lr according to ACCTRAN). Clade 26A. Triglidae, Peristediidae, and Hoplichthyidae-clade 26A is sup- ported by characters 51-1, 79-2, 97-1, and 104-1 (also 4-1, 14-1, 27-1, 49-3, 74-1, 92-1, 93- lr, and 95-1 according to ACCTRAN and 13-1, 38-1, 48-1, 49-2, and 69-lr aecording to DELTRAN), Clade 26B. Platycephalidae-this clade is supported by characters 3-lr, 26-1,

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30 Hisashi Imamura

59-2, and 94-1 (also 53-1 and 68-lr according to DELTRAN), This family also has three autapomorphic conditions: the urohyal and first basibranchial sutured, the

sternohyoideus inserted onto both the lateral and ventral aspects of the urohyal, and the adductor superficialis pelvicus connected to the pelvic fin rays via a long, strong tendon (Imamura 1996). Clade 27A. Triglidaerthis clade is supperted by characters 2-1, 5-1, 82-1, 99-1, 101-Or, and 102-1 (also 53-Or and 68-Or according to ACCTRAN and 4-1, 14-1, 27-1, 49-3, 92-1, 93-lr, and 95-1 according to DELTRAN). Clade 27B. Peristediidae and Hoplichthyidae-this clade is supported by char- acters 19-2, 20-1, 25-2, 57-1, 71-1, 72-1, 73-1, 107-1, 109-2, and 110-1 (also 103-1 according

to ACCTRAN or 70-lr and 74-1 according to DELTRAN), Following ACCTRAN analysis, the character of TS 111 at the node of this clade was not confirmed (111-1, -2, and -3 thereby being equivocal). Clade 28A. Chelidonichtbys, Lepidotrigla, and Ptetl)?gotrigla-this clade is sup- ported by characters 30-1 and 36-2 (also 64-2 and 108-1 according to ACCTRAN and 64-1 and 74-1 according to DELTRAN). Clade 28B. Bellator and Prionotus-this clade is supported by characters 10-1 and 56-Or (also 74-Or according to ACCTRAN). Prionotus has unambiguous charac- ter 16-1, and Betlator, 17-1 and 101-3. Clade 29A. Chetidonichthys-this clade is supported by characters 25-1, 70-1, and 76-1 (also 64-2 and 108-1 according to DELTRAN). Clade 29B. Lepidotrigla and Ptei:ygotrigla-this clade is supported unambigu-

ously by the characters 3-lr, 17-1, and 51-Or. Following ACCTRAN analysis, it is un-

clear whether this clade is also supperted by 16-1 or 16-2, Lepidotrigla is supported by 20-1 and 68-lr (also 16-1, 66-2, and 108-1 accerding to DELTRAN), and Pter:y- gotrigla by 25-2, 101-2r, 102-Or, and 103-1 (also 66-lr and 108-Or according to ACC- TRAN and 16-2 according to DELTRAN). Clade 30A. Peristediidae (Peristedion+SatyrichtdysFthis clade is supported unambiguously by characters 2-2, 15-1, 16-2, 21-1, 28-1, 30-1, 36-1, 39-er, 40-1, 43-1, 62-1, 67-3, 70-1, and 76-1 (also 49-2r according to ACCTRAN and 4-1, 14-1, 27-1, 92-1, 93-lr, 95-1, 103-1, and 111-3 according to DELTRAN). Peristedion has characters 3-lr, 38-Or, and 48-Or (also 53-1 according to DELTRAN), whereas Sa(yrichthys has 43-2 and 66-3 (also 53-Or according to ACCTRAN), Clade 30B. Hoplichthyidae (HOplichthysFthis clade is supported by charac- ters 10-1, 220r, 23-1, 26-1, 32-1, 33-1, 41-1, 42-1, 89-1, and 90-1 (also 4-Or, 14-Or, 27-Or, 92- Or, 93-Or, and 95-Or according to ACCTRAN and 49-3 and 53-1 according to DEL-

TRAN). Hoplichthyidae also has two autapomorphic characters: absence of the en- topterygeid, and articulated hypohyals and basihyal (Imamura 1996).

Classification

The phylogenetic analysis of the Scorpaenoidea demonstrates that the former Scorpaenoidei was a paraphyletic group. Although Ishida (1994) regarded the for- mer Scorpaenoidei as monophyletic, supported by two synapomorphies [presence ofa suborbital stay (divided into characters 8-1, -2, and -3 in this study) and num- ber of supraneurals reduced to one (character 65-2 in this study)], his synapomor- phies have now been recognized in all or most former platycephaloids, the analysis

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Phylogeny of Scorpaenoidea 31

placing characters 8-1 (suborbital stay with the end pointed) and 65-2 at the node of Scorpaenoidea. Mandrytsa (2001) also supported the monophyly of his Scor- paenoidei, excluding Congiopodidae (sensu Nelson 1994) and Pataecidae, on the basis of two derived characters (presence of three neuromasts on the first infraor- bital and first two dorsal proximal pterygiophores being inserted into the space be- tween the second and third neural spines). Three neuromasts on the first infraer- bital are also recognized in several outgroups (e.g., DiptQprion, Grammistes, IViphon) (Baldwin and Johnson 1993; present study), as well as in many percoids (present study), and Mandrytsa's (2001) eharacter polarity of neuromast number is not acceptable. Dorsal fin pterygiophore insertion was not included in the present phylogenetic analysis, because it is unclear whether the first vertebra was absent or the dorsal fin pterygiophores had shilted anteriorly in the Congiopodidae (first "first" "second" pterygiophore inserted into the space between the and neural

spines in the congiopodids vs. inserted into the space between the second and third

neural spines in other scorpaenoids and most outgroup taxa) and the homology of each anterior dorsal fin pterygiophore in the super:lamily is uncertain, Although the dorsal pterygiophore insertion pattern in the former Platycephaloidei (first and second dorsal pterygiophores inserted into the spaces between the second and third, and third and fburth neural spines, respectively, see Imamura 1996, fig. 57A) is also recognized in most serranids (Johnson 1984; present study), this pattern can be considered a reversal based on the reconstructed relationships. Therefore, the former Scorpaenoidei cannot be recognized cladistically as a valid taxon, and the supedemily Scorpaenoidea should not be divided into two major taxa correspon- ding to the former Scorpaenoidei and Platycephaloidei. Although Mandrytsa (2001) did not include Congiopodidae and Pataecidae in his Scorpaenoidei, it was inferred in this study that Congiopodidae (clade 21A) and Pataecidae (lower branch of clade 21B) are deeply nested within the superlamily Scorpaenoidea; the absence of a suborbital stay, absence of spines associated with the parietal sensory canal, and absence of the adductor dorsalis in the Pataecidae represent reversals, These families are included in the Scorpaenoidea as redefined here. Although Mandrytsa (2001) split the Congiopodidae into two families, Con- giopodidae (including only Congiopodus) and Zanclorhynchidae (containing Aler- tichthys and Zanclorltynchus), the latter is, for now, not recognized here, because

Zanclorhynchus, the genus of the family, was not examined and Congiopodi- dae (sensu Nelson 1994) was well defined by 17 unambiguous derived characters in this study. The family Sebastidae (sensu Eschmeyer 1998) includes the following seven valid genera: Sebastiscus, Sebastes, Adelosebastes, HOzukius, Helicolenus, Sebas- totebus, and Trachysconpia. However, none of these genera could be shown to have a sister relationship with any other, The genus Tracdyscorpia (clade 13A) being nested within the scorpaenids (clade 6A), it should be included in Scorpaenidae; this interpretation agrees with Mandrytsa (2001). The other sebastid genera ini- tially branched off successively from the other scorpaenoids. Although Eschmeyer (1998) recognized two subfamilies (Sebastinae including Sebastiscus, Sebastes, Hbzukius, and Heticolenus, and Sebastolobinae containing Adetosebastes, Sebas- tolobus, and Tracdyscot:pia) in the Sebastidae, neither is monophyletic. It is cladis- tically reasonable to treat each genus, except for [llrachyscoilpia, as a separate fam- ily, in which case new family-group names would need to be established for Se-

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bastiscus, Adelosebastes, HOzukius, and Llalicolenus among the above-mentioned genera. However, of them, Sebastes, Sebastiscus, and Helicolenus are known to be ovoviviparous (e,g., Washington et al, 1984a), which can be regarded as an apomor- phic condition because outgroups (Serranidae) are known to spawn the eggs (e.g., Kendal1 1984). I could not include characters related to reproduction in the present analysis, because such information is still unknown for many species of Scor- paenoidea, including Adelosebastes and HOz"kius. In this study, I provisionally rec- ognize a paraphyletic Sebastidae, including the examined Sebastiscus (clade IA), Sebastes (clade 2A), Adetosebastes (clade 3A), HOzukius (clade 3B), and Helicolenus (clade 4A), until further reproductive infbrmation is aecumulated, rather than es- tablish several new family groups without an analysis including characters re- lated to reproduction. Mandrytsa (2001) included Thysanichthys as well in his fam- ily Sebastidae. Although this genus was not examined in the present study, I also provisionally include it in the newly-defined Sebastidae, fo11owing Mandrytsa (2001). I recognize the tamily Sebastolobidae including only Sebastotobus (clade 5A), because this genus has different mode of reproduction than several members of the Sebastidae; i.e., the eggs of Sebastolobus are extruded in bilobed gelatinous egg masses, which fioat at the sea surface (Washington et al. 1984a). On the basis of the common absence of the hyohyoides inferioris, Ishida (1994) inferred that the families Setarchidae (inc}uding Setarches, Lioscorlpius, and Ectre- posebastes) and Neosebastidae (containing NLiosebastes and Maxilticosta) have a sister relationship. Although none of the examined specimens belonging to these families had that muscle (character 84-O), this character was determined to be ple- siomorphic for the Scorpaenoidea in the present analysis, so Ishida's (1994) conclu- sion is not supported. Rather, it appears that Setarchidae (clade 13B) is nested within Scerpaenidae (clade 6A) and has a sister relationship with Tracdyscoi:pia (clade 13 A), whereas Neosebastidae (clade 22A) is the sister group of the fbrmer Platycephaloidei (clade 14B), The former relationship agrees with the cladegram in Mandrytsa (2001, fig, 225) analyzed by PAUP, although that author did not accept such a relationship. Mandrytsa (2001> classified the family Scorpaenidae into two subfamilies (Scorpaeninae, including three tribes-Scorpaenini, Pteroini, and Setarchini; and

Taenianotinae, containing two tribes Taenianotini and Pteroidichthyini). How-

ever, such a classification is not acceptable cladistically, because his Scorpaeninae is a paraphyletic group derived from the non-rigorous cladogram (Mandrytsa 2001, fig. 235) used for his classification. In addition, many disagreements exist between the relationships presented by Mandrytsa (2001) and the present study. Accord- ingly, Scorpaenidae has not been subdivided into subfamilies or tribes in this

study, although this may be done in the future. Scorpaenidae, containing Tra-

chyscor:pia and the former Setarchidae, is included in clade 6A. The family Tetrarogidae was subdivided into fbur subfamilies (Tetraroginae, Centropogoniae, Cocotropsinae, and Neocentropogoninae) by Mandrytsa (2001); however, judging from his cladogram of the family (Mandrytsa 2001, fig. 232), Neo- centropogoninae is a paraphyletic group. Although only a single genus (Tetraroge: clade 16A) was examined in the present study, Mandrytsa's (2001) classification of Tetrarogidae cannot be fbllowed cladistically; therefbre, I have refrained from sub-

dividing the family into subfamilies. Eschmeyer (1998) recognized the family Synanceiidae as comprising three sub-

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Phylogeny of Scorpaenoidea 33

families (Synanceiinae, Choridactylinae, and Minoinae), although Ishida (1994) had not recognized any subfamilies in that family. Eschmeyer et al. (1979b) had earlier included two genera (Choridactylus and inimicus> in the Choridactylinae; however, this concept of Choridactylinae is paraphyletic, because Choridactylus (upper branch of clade 19B) and Minous (lower branch of clade 19B), the latter being included in Minoinae, have a sister relationship. Mandrytsa (2001) hypothe- sized Choridactylus and inimicus as having a sister relationship supported by two characters (opposing parietal sensory canals not continuous, and absence of the sixth infraorbital). Although the former character was not used in the present phy- logenetic analysis, it is considered here that Choridactytus and inimicus both have the sixth infraorbital, but fused with the sphenotic (character 10-1). Because char- acter 10-1 supports clade 14A, it is not a synapemorphy of Choridactylus and Iltimi-

cus. The sister relationship between Choridactylus and Minous is supported herein by three unambiguous characters (46-1, 86-1, and 89-Or). Of these, 46-1 (cleithrum and coracoid partially connected ventromedially) is an autapomorphy among the Scorpaeneidea, It is proposed here that the former subfamily Choridactylinae be divided into two subfamilies: the redefined Choridactylinae (including Choridacty- tus: upper branch of clade 19B) and Inimicinae (containing inimicus: clade 19A), which had been treated as a junior synonym of Choridactylinae (see Eschmeyer et al. 1979b). Synanceiinae (sensu Eschmeyer and Rama Rao 1973) (clade 18A) is re- tained in this study, although some genera of this subfamily (Dampierosa, Leptosy- nanceia, , and ) were not examined. Minoinae

(lower branch of clade 19B) is a monotypic subfamily, as recognized by Eschmeyer et al, (1979a). To summarize, the family Synanceiidae (clade 17A) is subdivided into four subfamilies, a classification that matches that of Mandrytsa (2001). Imamura (1996) recognized the following seven families in the former Platy- cephalioidei (clade 22B), based on phylogenetic relationships: Plectrogeniidae (clade 23A), Parabembridae (clade 24A), Bembridae (clade 25A), Triglidae (clade 27A), Peristediidae (clade 30A), Hoplichthyidae (clade 30B), and Platycephalidae (clade 26A). Their relationships, as suggested by Imamura (1996), were confirmed in this study and these families are retained, although Nakabo (2002b) did net rec- ognize Plectrogeniidae, including Plectrogenium and Bembradiunz. Richards and Jones (2002) recently presented a detailed study of triglid interrelationships. In conclusion, a new classification of Scerpaenoidea is herein proposed, the su- perfamily including the fo11owing 20 families: Sebastidae (paraphyletic, including clades IA, 2A, 3A, 3B, and 4A), Sebastolobidae (clade 5A), Scorpaenidae (clade 6A), Apistidae (clade 15A), Tetrarogidae (clade 16A), Synanceiidae [including Synancei- inae (clade 18A), Inimicinae (clade 19A), Choridactylinae (upper branch of clade 19B), and Minoinae (lower branch of clade 19B)] (clade 17A), Aploactinidae (includ- ing the Aploactininae, Bathyaploactininae, and Matsubarichthyinae: Mandrytsa 2001) (clade 20A), Congiopodidae (clade 21A), Gnathanacanthidae (upper branch of clade 21B), Pataecidae (lower branch of elade 21B), Caracanthidae (not examined in this study), Eschmeyeridae (not examined in this study; see Mandrytsa 2001), Neosebastidae (clade 22A), Plectrogeniidae (clade 23A), Parabembridae (clade 24A), Bembridae (clade 25A), Triglidae (clade 27A), Peristediidae (clade 30A), Ho- plichthyidae (clade 30B), and Platycephalidae (including the Platycephalinae and Onigociinae: Imamura 1996) (clade 26B).

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34 Hisashi Imamura

Acknowledgments

I express my sincere thanks to J. S. Nelson and K. Jackson (UAMZ) fbr reading the draft manuscript and providing important comments. I alse sincerely thank G. Shinohara (NSMT), who provided anatomical data for Tetraroge niger, I am grate- fu1 to T. Iwamoto (CAS), S, L, Jewett and L. Palmer (USNM), K. Matsuura (NSMT), and M. McGrouther (AMS) for providing materials, and also thank G. S. Hardy (Ngunguru, New Zealand) for English corrections.

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