Tus (Actinopterygii: Teleostei: Platycephalidae), with Comments on the Composition of the Type Series

Home , Platycephalidae, Platycephalus, Platycephalus speculator

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Species Diversity, 2006, 11. 295 306

Rediagnosis of the Marbled Flathead, Platycephalus marmora-

tus (Actinopterygii: Teleostei: Platycephalidae), with Comments on the Composition of the Type Series

Hisashi Imamura

7:he Hbkkaido Uhiversity Museum, thculty ofFisheries, Hbkkaido University, 3-1-1 Minato-cho, Htikodate, HOhkaido, 041-8611 Japan E-mail: [email protected]

(Received 8 May 2ee6; Accepted 30 August 2006)

The taxonomic characters of the marbled flathead, Platycephalus marmoratus Stead, 1908, are described and evaluated. In addition to the marbled

body and dark brown to black caudal fin, the fo11owing combination of char-

acters separates it from the other 14 species of the genus Platycephalus: 13 second dor$al and anal fin rays; 63-70 scales in the Iateral line, with all pored

scales lacking spines; snout and interorbit naked; no strong canine teeth on the jaws, palatines, and vomer; interopercular fiap absent; and skinny sensory tubes on the suborbitals and preopercle partially covering the cheek region. It has been confirmed that the type series of the species was composed of the holotype and one paratype, although the paratype is presumably no

lenger extant,

Key Words: Actinepterygii, Platycephalidae, Pla(ycqphaius marmoratus, taxonomic characters, diagnosis, redescription, paratype.

Introduction

Stead (1908) described a new species of flathead, Platycephalus marmoratus "marbled Stead, 1908, and proposed flathead" as its English name, in reference to the marbled markings on the body and head. No other species of Platycephalus Bloch, 1795 [sensu Imamura (1996), who synonymized IVeqplatycqphalus Castelnau, 1872 under the genus] has a marbled patter'n on the body and head. In alcohol, specimens of Platycqphalus marmoratus also have a more extensive area of dark brown to black in the caudal fin than is present in other species of Platycephalus. Al- though P. marmorattts is easily distinguished from its congeners when coloration

is reasonably well retained, these characters may not be available in long-pre- served specimens (for example, the holotype of this species is considerably faded) (Fig, 1), Although severa] authors have briefly described some of the morphologi- cal characters of P. marmoratus, e.g., Knapp (1991) and Imamura (1996), who pro- vided selected meristic values, and osteological and mycological characters, re-

spectively, no study has compared P. marmoratus with its congeners in detail, The purpose of this study is to redescribe Platycephatus marmoratus based on the holotype and other material, compare its taxonomic characters (other than color) with other species of Platycephalus, and present an emended diagnosis. Also, the status of the paratype ofP. marmoratus is clarified and a previous inter-

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296 Hisashi Imamura

pretation of composition of the type series is corrected.

Materials and Methods

Counts and measurements were made fbllowing Hubbs and Lagler (1958), Gill rakers were counted on the right side. Measurements were made with calipers to

the nearest O,1mm. Length of the first dorsal fin base excludes the isolated spine between the dorsal fins. Terminology of head spines fbllows Knapp et al, (2000). Institutional acronyms are from Leviton et al. (1985) except for SALAB (South African Institute fbr Aquatic Biodiversity, formerly RUSI), Standard and head

lengths are abbreviated as SL and HL, respectively.

Taxonomic Account

Platycophatus marmoratus Stead, 1908 (Figs 1-6)

Platycephalus marmoratus Stead, 1908: 9, pls 3-5 (type locality: Port Stephens, Aus- tralia); McCulloch 1929: 402; Coleman 1980: 109, an unnumbered color figure; Hutchins and Thompson 1983: 78, fig, 112; Hutchins and Swainston 1986: 127, fig. 204; Paxton et al. 1989: 469; Knapp 1991: 29; Kuiter 1993: 104, two unnumbered color figures; Imamura 1996: 125; Hutchins 2001i 28; Grant 2004: 196, pl. 89, an unnumbered co]or plate.

Material examined. Holotype: AMS I.15260, 310.4mm SL, Port Stephens, June 1904. Other material. Seven specimens, all collected from New South Wales, Aus- tralia: AMS I.15279, 471.6mm SL, data unknown, cell. D. Stead; AMS I,20721-O02,

270.9mm SL, Coffs Harbour, 30018'S, 153008'E, 80m depth, 13 August 1978, colL J. Hooper; AMS I.20870-OOI, 151.8mm SL (dissected by Imamura 1996), Bondi Beach, Sydney, 33053'S, 151015'E, 30 October 1977, co}1. R. Kuiter; AMS I.22129-O04, 309.5mm SL, North Coast, 1978, colL J. Hooper; AMS I,25663-O13, 169.5mm SL, southeast of Yamba, 29034'S, 153e25'E, 49-53m depth, 24 March 1985, coll. FRV Kti- pala; AMS I.25665-026, 184.6mm SL, northeast of Yamba, 29021'S, 153029'E, 49-51m depth, 21 May 1985, coll, FRV Ktipala; AMS I.27322-O06, 173.0mm SL, off Tuncurry, 32eo8'S-32o06'S, 152e31'E-152033'E, 26ul7m depth, 1 October 1985, co!L FRV KI]pata. Diagnosis (emended). A species of Platycqphalus with 13 second dorsal and anal fin rays; 63-70 pored scales in lateral line, each with one exterior opening posteriorly and lacking spine; 2+8-11 (7-11 in Knapp 1991) gill rakers; snout and interorbit naked; lower opercular spine slightly longer than upper one; no strong canine teeth on jaws, palatine, or vomer; interopercular flap absent; margin of interopercle not scalloped; skinny sensory tubes from suborbitals and preopercle partially covering cheek region. Description. Data are presented first fbr all specimens, then for the holotype (in parentheses). Counts: dorsal fin spines I-VI-VII-I or I-VII-VIII (I-VII); secend dorsal fin rays 13

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Fig, 1 Pfalyeophahas inatrno;atus Stead, l90g, AMS holetypo,310A,mm 5E,,Pori

$lephelts NSW, Autstr,]lia, entire emmal in dorgal vie"IIA260

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ing 2Yambevip"sPlan,eeptzakis masmfiratus Steact, 1908, AMS i25663026, 1846mm Si, northeast ot

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298 Hisashi Imamura

3mm

Fig. 3. Iris lappet (left eye) of PtaCycephalus marmoratus Stead, 1908, AMS I.25665-026, 184.6mm SL, northeast of Yamba, NSW, Australia.

(13); anal fin rays 13 (13); pectoral fin rays 1-2 (upper, unbranched)+11-14 (middle, branched)+6-8 (lower, unbranched)-2or22 (2+12+7=21); pelvic fin rays I, 5 (I, 5); branched caudal fin rays 5-6 (upper)+5 6 (lower)-11-12 (6+6=:12); pored scales in lateral line 63-70 (66), all scales witheut spines; oblique body scale rows slanting downward and backward above lateral line 107-119 (108); gill rakers 2+8-11=10-13 (2+8-10), `V･h Proportions as SL: HL 28.1-29.7 (28.9); predorsal length 31,5-33.6 (32.6); length of first dorsal fin base 15.8-20.4 (17,7); length of second dorsal fin base 34.9- 36,5 (36.1); length of anal fin base 38.3-40.8 (38.8); caudal peduncle length 6.9 10.6 (7,9); caudal peduncle depth 3.8-4.4 (3.8); snout length 8.8-9.4 (9,2); orbital diameter 4.5-5.9 (4.9); upper jaw length 11.1-12.3 (11.5); lower jaw length 15,4-16.2 (15.4); interorbital width 3.8-6,3 (5.1); postorbital length 14.1-15.8 (15.2); suborbital width 2.5-3.0 (2.8); pectoral fin length 13.5-15.9 (14.2); pelvic fin length 21.9-24.1 (22,1); caudal fin length 15,5-19.8 (17,5); length of first spine of first dorsal fin O.6-3.2 (O.6); length of second spine of first dorsal fin 13.4-14.8 (14.8); length of first ray of second dorsal fin 7,2-13,1 (10,7); length of first anal fin ray 6.8-10.1 (6.8). Proportions as 9･,b HL: snout length 29,8-32.3 (31,9); orbital diameter 15.2-20.1 (16.8); upper jaw length 38.1-42,1 (39,8); lower jaw length 53,3-55.5 (53.3); interorbital width 13.6-21.4 (17.6); postorbital length 50.1-53.4 (52.5); suborbital width 8.3- 10.2 (9.7), Body greatly depressed, mostly covered with ctenoid scales, but some cycloid scales on undersurface. Head greatly flattened, length 3,4-3,5 (3,5) in SL; snout and interorbit naked, occipital region with or without several scales embedded under skin (without), nape mostly scaled. Snout robust, longer than eye diameter, length 3,1-3.4 (3.1) in HL. Upper sudece of eye without papillae. Iris lappet usually broad, well expanded and simple dorsally, and broad, simple, and weakly cenvex ventrally (Fig. 3), weakly concave dorsally only in AMS I.22129-O04. Interorbital width 4.7-7,4 (5.7) in HI., varying with growth, smaller than orbital diameter in smaller specimens, becoming larger than orbital diameter by 270 mm SL. Spines and ridges weakly developed on top and side ef head (Fig. 4). Nasal lacking spines, Lachrymal with single antrorse spine. Single preocular spine present. Suborbital and supraor- bital ridges entirely smooth, Single postorbital spine with broad base present in

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Taxonomic characters of a platycephalid 299

LAC PRO

po NN pso

× N

10 mm

Fig, 4. Dorsal view of head of Ptatycephalus marmoratus Stead, 1908, AMS I,25665-026, 184,6

mm SL, northeast of Yamba, NSW, Australia. Abbreviations: LAC, lachrymal; LO, lower oper-

cular; NC, nuchal; PO, preopercular; PRO, preocular; PSO, postoeular; UO, upper epereular.

smaller specimens, blunt, obscure, or absent in largest specimens (blunt), Pterotic, frontal, and supraoccipital with entirely smooth ridges. Parietal usually with ridge lacking spines, rarely having one spine embedded under skin posteriorly (lacking spine). Supratemporal, posttemporal, and supracleithrum with or without spine (spine present only on supracleithrum). Preopercle usually with two spines, with an additional spine ventrally only in AMS I. 20710-O02; lower spine slightly longer than upper one, not reaching posterior margin of opercle; upper spine lacking sup- plementary spine. Opercle with two spines, lacking prominent ridge, Interopercu- lar flap absent; margin of interopercle not sca]leped (Fig. 5). Maxilla reaching slightly beyond anterior margin of orbit, length 2.4-2.6 (2.5) in HL, Teeth in band on jaws and palatine, and in single V-shaped patch on vomer; tooth band on upper jaw lacking distinct notch medially (Fig, 6). Upper jaw with several small canine teeth anteriorly; remainder of jaw with small- to moderate-size conical teeth, Lower jaw with two or three tooth rows anteriorly and two posteriorly; inner row

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300 Hisashi Imamura

10 mm

Fig. 5. Ventrolateral view of head of Platycephalus marmoratus Stead, 1908, AMS I.25665-026,

184.6mm SL, northeast of Yamba, NSW, Australia, showing the smooth interopercle and the

skinny sensory tubes arising from the suborbitals and preopercle.

with longer conical teeth; outer row with small- to moderate-size conical teeth.

Palatine with two tooth rows: inner row with longer and stouter conical teeth, outer row with small conical teeth. Vemer with small- to moderate-size canine teeth anteriorly and a few canine teeth posteriorly. Lip margins without papillae. Skinny sensory tubes on suborbitals and preopercle weakly developed, partially covering cheek region (Fig. 5). Pored scales in lateral line each with one exterior opening posteriorly; opening in most pored scales directed posteroventrally, in several scales posterodorsally. First dorsal fin originating posterior to opercular margin. First and second dorsal fin narrowly separated. Pectoral fin rounded posteriorly, length 1.9-2.2 (2.0) in HL. Posterior tip of pelvic fin reaching to between origin of anal fin and base of second anal fin ray (between bases of first and second fin rays), length 1.2-1,4 (1,3) in HL. Caudal fin slightly rounded posteriorly, length 1.5-1.9 (1.7) in HL, Cblor in alcohoL Body and head marbled with dark brown, brown, and pale ir- regular bands and spots above, pale below. Head with single indistinct, dark brown band crossing occipital and anterior opercular regions, Body with five dark

brown bands or sometimes with three dark brown spots replacing the three ante-

rior bands; bands continuous with blackish spots on side of body, First and second dorsal fins with small, dark-brownish spots along rays. Pectoral fin pale brown with white lower margin; small brown spots present or absent on upper portien. Pelvic fin dark brown with pale brown basal portion and white outer margin. Anal fin with brownish pigment along rays; membranes of posterior portion of anal fin dusky. Caudal fin dark brown, gradually becoming black posteriorly; posterior margin white; anterior portion of upper margin with or without several brownish oblique bands. Distribution. Known from southern Western Australia and eastern coast of Australia, from southern Queensland to Bass Strait, Tasmania (Stead 1908; Paxton

et al. 1989; Kuiter 1993; Hutchins 2001; this study).

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Taxonomic characters of a platycephalid 301

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Comparison. In addition to the marble-like markings on the body and head, and the dark brown to blaek caudal fin, Platycephalus marmoratus can easily be separated from the other 14 species in the genus Platycqphal"s by the fo11owing characters: number of second dorsal and anal fin rays, number of pored scales in the lateral line, presence ofa spine on the anteriormost pored scales in the lateral line, gill raker counts, scaled area of head, teeth, interopercular flap, and skinny sensory tubes on cheek region. Of these, the number of spine-bearing anteriormost pored scales in the lateral line and the extent of scalation on the head are sufficient in combination to distinguish R marmoratus from any other species of Platy- caphalus. The counts discussed here are summarized in Table 1. IVumbers of second dorsal and anal fin rays, Platycqphalus marmoratus has 13 rays in both the second dorsal and anal fins. Nthough intraspecific variation in these numbers occurs in several species, they are usefu1 in distinguishing P. mar-

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302 Hisashi lmamura

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Taxonomic characters of a platyeephalid 303

Ptatycephatus is more varied (59-93) than in other platycephalid genera, which usually have ca, 50-55 (except fbr Elates Jordan and Seale, 1907 with 83-107 and Onigocia Jerdan and Thempson, 1913 with fewer than 45; e.g., Knapp 1999). Despite intraspecific variation in the number of lateral line scales, the counts for P. marmoratus (63-70) do not overlap with those of the fo11owing species: P. aurimaculatus Knapp, 1987 (81-85), P. bassensis (71-82), P. caeruleQpunctatus (83-92), P. .fttscus Cuvier, 1829 (71-79), P. Iaevigatus Cuvier, 1829 (81-92), and P. speculator Klun- zinger, 1872 (80-93) (Knapp 1991; this study). The number of anteriomiost pored scales in the Iateral line that have a spine is also a valuable character. Ptatycephalus marmoratus differs from all members of the genus, except fbr P. arenarius Ramsay and Ogilby, 1886, P. chautiodous Knapp, 1991, and P. Iaevigatus, in lacking any pored lateral line scales with a spine (vs, IB such scales in the others) (Knapp 1991, 1999; this study). AJUmber of gilt rakers. The gill raker count is 2+7-11 in P. nzarmoratus, whereas there is either one, or else three or more rakers on the upper gill arch in P. arenarius (3 or 4), P. aurimaculatus (1), P. bassensis (4 or 5), P. caeruleopunctatus (4-6), P. Iongispinis Cuvier, 1829 (5-7), and P. speculator (3 or 4). In addition, there are 12 or more rakers on the lower gill arch of P. bassensis (13-15), P. caeruleopunctatus (14-17), and P, longispinis (16-19) (Knapp 1991; this study). Scated area of head, In P, marmoratus, the snout and interorbit are naked. In contrast, the interorbit and posterior portion of the snout are covered by scales in all other species of Piatycephalus except fbr P. chautiodous and P. taevigat"s, in which the snout, interorbit, and occipital region are naked (this study). In the examined specimen of P. aurimaculatus it is unclear whether the posterior portions of the snout and interorbit are naked, or the scales there have been detached. Knapp (1991) stated that the top of the head in this latter species has scales. 71eeth. Although P. marmoratus has several small canine teeth anteriorly in the upper jaw, there are no canine teeth in other regions. In centrast, large and strong canine teeth are present anteriorly on the upper jaw, lewer jaw, and palatine of P. aurimaculatus, P. chauliodous, P. conatus Waite and McCulloch, 1915, and P. richardsoni Castelnau, 1872 (Knapp 1991; this study). interQpercular flap. Presence or absence of the interopercular fiap has been recognized as an important taxonemic character for Platycephalidae. Platy- cephalus marmoratus lacks the flap (see also Imamura 1996), whereas it is present in P. arenarius, P. caeruleQpunctatus, P. cultellatus Richardson, 1846, P. endrachtensis Quoy and Gaimard, 1825, P. .fttsc"s, P, indicus Linnaeus, l785, P. Iaevigatus, and P. speculator (e.g,, Knapp 1999), Although Imamura (1996) stated that P, bassensis lacks an interopercular flap, the present study has revealed that it is present or absent in different individuals of the species, Skinay sensoilv tubes on cheek ragion. Imamura et al, (1995) demonstrated that the skinny sensory tubes on the cheek region, which extend from the suborbitals and preopercle, are useful characters for platycephalid taxonomy. In additien to P, marmoratus, only P. chauliodous, R taevigatus, and P. Iongispints have sensory tubes partially covering the cheek region. In P, longispinis only the tubes from the preopercle partially cover the posterior cheek region, Others. In addition to the above-mentioned characters, P. marmoratus is sepa- rable from several members of Platycephalus in respect to the fo11owing characters: the lower preopercular spine being slightly longer than the upper one (vs,

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304 Hisashi Imamura

upper longer than lower in P. Iaevigatus and lower much longer than upper in P. tongispinis; Kuiter 1993; this study); the margin of the interopercle not being scalloped (vs. scalloped in P. chautiodous; Knapp 1991; this study), and the pored scales in the lateral line having one exterior opening posteriorly (vs, two in P. richardsoni; Imamura 1996). Remarks. Eschmeyer et al. (1998) indicated that Platycephalus marmoratus might have been described from the holotype and two paratypes. They flagged the possibility that AMS I.15279 is one of the paratypes by inserting a question mark before the catalogue number. Stead (1908) clearly stated, however, that he described the species based on two specimens, one collected from Port Stephens and

one from Coffs Harbour, and he designated the specimen from Port Stephens as the

holotype. It is therefore obvious that the type series of P. marmoratus comprises the holotype and a single paratype. Although AMS I.15279 was collected by Stead, the specimen has no attached collection data. Stead (1908) stated that the total length of the paratype was 490 mm, whereas the present author's measurement of the length of AMS I.15279 is 550.3mm. The total length of the holotype, 367mm, as measured by Stead (1908) and 368.8mm as measured in the present study, are very similar. Therefore, AMS I,15279 is, most likely, a non-type specimen of P, marmoratus. No specimen of P, marmorattts collected from CoffS Harbour befbre 1908 is deposited at AMS (McGrouther pers, comm., 23 March 2006). Accordingly, the whereabouts of the single paratype are unknewn and it has probably been lost.

Comparative material examined Platycqphalus arenariLts (9 specimens from Western Australia, Queensland, and New South Wales, Australia): AMS B.7136, holotype, 238.9 mm SL; AMS I.22831- 032, 306.2mm SL; AMS I.34399-O16, 123.0mm SL; AMS I,34361-O02, 2 of 4 specimens,

135,6, 206.5mm SL; AMS I.41262-023, 126.8mm SL; AMS I.41674-O13, 97.2mm SL;

AMS I.41877-038, 145.9 mm SL; AMS L43521-OOI, 130.9 mm SL. Platycephalus aurinzacutatus (1 specimen from Tasmania, Australia): AMS I,26328-OOI, paratype, 322.4 mm SL, Platycqphalus bassensis (7 specimens from New South Wales and Tasmania): AMS I.12794, 197.4 mm SL; AMS L12796, 1 of 2 specimens, 175.7 mm SL; AMS I.14150,

I.14151, 2 specimens, 114.2, 200.8 mm SL; AMS I.17561-OOI, 155.0mm SL; AMS I.19832- O04, 212,4mm SLI AMS I.34823-OOI, 302,8 mm SL. Ptatycephalus caeruteQpunctatus (9 specimens from New South Wales): AMS IA,5700, 166.3mm SL; AMS IB.6296, 141.8mm SL; AMS I.3163, lectotype, 290.6mm SL; AMS I.3947, 266.3mm SL; AMS I.14928, 242,Omm SL; AMS I,9567, 205,8mm SL; AMS I,15202, 216.8mm SL; AMS I.19832-O02, 338.2 mm SL; AMS I.39988-O03, 132.6 mm SL, Platycephalus chauliodous (1 specimen from Western Australia): AMS I,20231- O07, holotype, 312.6mm SL. Platycaphalus conatus (8 specimens from South Australia and Western Aus- tralia): AMS E.750, 232.5mm SL; AMS L13649, paratype, 242,7mm SL; AMS I,13650, paratype, 269.1mm SL; AMS I.12393, I,12394, 2 specimens, 234.2, 236,6mm SL; AMS I.18710-O07, 2 specimens, 158.0, 173.9mm SL; AMS I.25551-OOI, 303.3mm SL, Platycephatus cuttellat"s (9 specimens from the Western Pacific): AMS IA,3201- 3202, 2 specimens, 224.2, 242.3mm SL; HUMZ 190447, 233.4mm SL; NSMT-P 55619, 125.8mm SL; NSMT-P 56060, 235.7mm SL; NSMT-P 67375, 202.6mm SL; NSMT-P

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Taxonomic characters ofaplatycephalid 305

68928, 230,6mm SL; NSMT-P 69319, 2 specimens, 244.2, 282.8mm SL. Platycophalus endrachtensis (8 specimens from Western Australia and Queens- land): AMS I.13140, 262,5mm SL; AMS I.20978-084, 291.3mm SL; AMS I,19468-O18, 1 of 4 specimens, 138,9mm SL; AMS I.21244-OOI, 150.5mm SL; AMS I.25508-O03, 98,2mm SL; AMS I.34314-O03, 260.9mm SL; AMS I,34397-025, 187.4mm SL; AMS I.34398-OO3, 208.1 mm SL, Platycephalus .fitscus (10 specimens from New South Wales): AMS I.16741-OOI, 241.4mm SL; AMS I.17178-OIO, 2 specimens, 251,5, 316.9mm SL; AMS I.21149-032,

212.2mm SL; AMS I.36105-OOI, 1 of 2 specimens, 183,7mm SL; AMS I.41262-024,

218.3 mm SL; AMS I.41264-020, 2 of 5 specimens, 163.5, 188.6 mm SL; AMS I.41287-020,

1 of 3 specimens, 186.0m SL; AMS I.41874-Oll, 203,6 mm SL. Platycephalus indicus (10 specimens from the Indian Ocean and Western Pa-

cific): CAS 29657, 240,6mm SL, Pakistan; CAS 58591, 280.7mm SL, India; FRLM

30658, 114.3mm SL, Andaman Sea; HUMZ 193328, 191.lmm SL, Indonesia; NSMT-P 71056, 2 specimens, 242.5, 256.4mm SL, Phuket; SAIAB (RUSI) 35752, 229,8mm SL, Taiwan; SU 13165, 313.2mm SL, Philippines; SU 35770, 203.5mm SL, India; SU 39878, 182.6 mm SL, Myanmar, Platycqphatus laevigatus (3 specimens from Victoria, Australia): AMS I,7543, I.7544, 2 specimens, both 271,1 mm SL; AMS I.7592, 287,3 mm SL. Platycophalus longispinis (12 specimens from New South Wales and Tasma- nia): AMS I,14178, 216.3mm SL; AMS I.16355-OOI, holotype, 233.0mm SL; AMS

I.16887-OOI, 5 of 7 specimens, 126.4-190.0mm SL; AMS I.17268-OOI, 1 of 4 specimens, 204.0mm SL; AMS I.20720-OOI, 208.0mm SL; AMS I.41259-OOI, 3 specimens, 180.3- 204.4 mm SL. Pla(),cephalus richardsoni (4 specimens from New South Wales): AMS B.6541, holotype of Ptatycephal"s nzacrodon Ogilby, 1885, 280.6mm SL; AMS I.23871-O02, 193.8 mm SL; AMS I.26026-OOI, 266.2mm SL; AMS I.40292-OOI, 230.6mm SL. Platycephalus speculator (2 specimens from South Australia and Western Aus- tralia): AMS I,16354-OOI, holotype of Platycephalus castelnaui Macleay, 1881, 281,1

mm SL; AMS I.17615-OIO, 144.3 mm SL.

Acknowledgments

I am deeply indebted to M. McGrouther (AMS) for reading the draft manu- script and for providing support when I visited AMS. I also express my thanks to all other staff of the Fish Section of AMS for their support during my werk at the museum. My special thanks also go to M. E, Anderson (SAIAB), D. Catania (CAS), S, Kimura (FRLM), and K. Matsuura and G. Shinohara (NSMT) fbr making materi-

als available.

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