Description of two new species of the genus Priolepis from the Indo-Pacific with redescription of Priolepis profunda and Priolepis psygmophilia

Ichthyol Res (2010) 57:373 388 DOI 10.1007/s10228-010-0170-6 FULL PAPER Description of two new species of the genus Priolepis from the Indo-Pacific with redescription of Priolepis profunda and Priolepis psygmophilia Douglass F. Hoese Helen K. Larson Received: 12 December 2006 / Revised: 19 April 2010 / Accepted: 18 May 2010 Ó The Ichthyological Society of Japan 2010 Abstract Priolepis akihitoi is described as new from Australia, New Caledonia and Japan. The species is distinctive in having a transverse papilla pattern, second dorsal rays usually I, 11, predorsal fully scaled and dark bars dorsally on the caudal fin. Priolepis cyanocephala is described as new from eastern Australia and Lord Howe Island. It differs from other species in having a reduced transverse papilla pattern, second dorsal rays I, 10, predorsal largely naked and head with vertical bands, but no bands on the body. Priolepis profunda is redescribed based on material from northwestern Australia and Thailand, and photos of specimens from Indonesia, New Guinea and the Philippines. Priolepis psygmophilia is redescribed based on recently obtained material from the Kermadec Islands and other material from Lord Howe Island, Easter Island and Rapa Island. Keywords Introduction Gobiidae South Pacific Japan Priolepis The genus Priolepis Valenciennes in Cuvier and Valenciennes (1837) has been revised in a series of papers by Winterbottom and Burridge (1989, 1992, 1993a, b, c). D. F. Hoese (&) Australian Museum, 6 College St., Sydney, NSW 2010, Australia e-mail: Doug.Hoese@austmus.gov.au H. K. Larson Museum and Art Gallery of the Northern Territory, P.O. Box 4646, Darwin, NT 0801, Australia e-mail: Helen.Larson@nt.gov.au They treated 26 Indo-Pacific species, noted the three western Atlantic and one eastern Atlantic species, and divided the genus in three separate groups, based on papilla pattern and extent of development of predorsal scales. Subsequently, Goren and Baranes (1995) described an additional species from the Red Sea, and Nogawa and Endo (2007) described a species from Japan, bringing the total described species to 32. We describe two additional new species from the Indo-Pacific region. Brief examination of material from Australia and New Guinea suggests that additional undescribed species undoubtedly exist. The genus Priolepis is regarded here in a broad sense, but includes species typically with a relatively narrow gill opening. Generally, the head and body are banded. As noted by Whitley (1932), Priolepis has priority over Zonogobius Bleeker 1874. The name was based on Priolepis mica of an Ehrenberg manuscript, and a description is given by Valenciennes in Cuvier and Valenciennes (1837), suggesting it might be identical to his Gobius semidoliatus Valenciennes in Cuvier and Valenciennes (1837). Consequently, the generic name was introduced in synonymy, but made available by Whitley (1932). Under the code of Zoological Nomenclature a name introduced in synonymy becomes available if used prior to 1961. Quisquilius Jordan and Evermann 1903 had consistently been regarded as distinct from Priolepis, based on the absence of predorsal scales in Priolepis. This character, however, is not of significance in this group, since in the related genus Trimma Jordan and Seale 1906, the nape varies from naked to fully scaled. Furthermore, in some species typically referred to Quisquilius, such as one of the species redescribed here, the predorsal scales are incompletely developed and highly variable. Consequently, we agree with Winterbottom and Burridge (1989, 1992) in regarding Quisquilius as a synonym of Priolepis.

374 D. F. Hoese and H. K. Larson Relationships of Priolepis have not been well studied. Hoese and Brothers (1976) suggested that Priolepis (as Zonogobius and Quisquilius) was related to Trimma, Paratrimma Hoese and Brothers 1976, Lythrypnus Jordan and Evermann 1896 and Eviota Jenkins 1903. Winterbottom and Burridge (1993a) suggested a relationship with Egglestonichthys Miller and Wongrat 1979. They also discussed possible relationships and apparent paraphyly in relation to Trimma and Trimmatom Winterbottom and Emery 1981. Miller (1978) noted the strong similarity in coloration of the Atlantic species, Gorogobius nigricinctus (Delais 1951), to Priolepis (as Quisquilius and Zonogobius). However, he concluded that the similarity was convergent and that the genus was most likely related to other European genera, particularly Corcyrogobius Miller 1972, based on papillae and 27 vertebrae. It is suggested here that Lubricogobius Tanaka 1915 and Larsonella Randall and Senou 2001 are also probably related to the group based on their overall similarity to Priolepis, differing from Priolepis in having few or no scales on the body. Lubricogobius ornatus Fourmanoir 1966 and Larsonella have scales confined to the caudal peduncle, while other species lack all body scales. Larsonella was defined largely on the basis of the presence of caudal peduncle scales. Apparently the authors were not aware that Lubricogobius ornatus has scales on the caudal peduncle. Consequently, the presence of these scales in Lubricogobius ornatus suggests that Larsonella may not be separable from Lubricogobius. The single species of Larsonella does differ from Lubricogobius in having a depressed head, and we tentatively retain the genus Larsonella. Further studies are clearly needed to determine the validity of Larsonella. Relations of the various genera will be treated separately. Currently, over 250 Australian shore fishes are restricted to the southwest Pacific (Lord Howe Island, Norfolk Island, Kermadec Islands, New Caledonia or New Zealand). Randall and Meléndez (1987) noted that some species from Easter Island were restricted to the south Pacific. While the number of species found in the south Pacific is small, Randall et al. (1990) noted that 10% of the species found at Easter Island and Rapa Island were restricted to the south Pacific. There are also examples of species endemic to those or other south Pacific islands with their nearest relatives also in the south Pacific (Hoese and Larson 2005). We describe one species here that is restricted to the southwest Pacific and redescribe a species that appears to range from the Kermadec Islands to Easter Island, although there is some apparent differentiation between Rapa and Easter Islands. The nearest relative of that species appears to be in the Hawaiian Islands. Previously only a single gobiid fish, Kelloggella oligolepis (Jenkins 1903), has been recorded from Easter Island (de Buen 1963; Hoese 1975). Between 1958 and 1969 three expeditions collected four additional species: Gnatholepis Bleeker 1874, a species of Hetereleotris Bleeker 1874, the species of Priolepis treated herein and Trimma unisquamis (Gosline 1959). The Gnatholepis was subsequently identified as Gnatholepis cauerensis pascuensis by Randall and Greenfield (2001) and the Hetereleotris as Pascua caudilinea by Randall (2005). Hoese and Larson (2005) synonymized Pascua with Hetereleotris, although Randall (2006) argued for retention of Pascua. Although few gobiids occur at Easter Island, the presence of Priolepis is not surprising, because the genus is widespread in the Indo-west Pacific and Atlantic. In addition to the 250 species from Australia restricted to the south Pacific, 63 species are known from the northwest Pacific, many of those found in other parts of the southwest Pacific, but not known from more tropical regions of Indonesia, New Guinea or the Philippines. Consequently, about 7% of the Australian fauna is found only in the south Pacific and/or the northwest Pacific (Hoese et al. 2006). One species included here is known only from the southwest Pacific, one from the south Pacific and one from the southwest Pacific and the northwest Pacific. Extensive material has become available from depths of 30 120 m due to trawling efforts of the CSIRO Marine Laboratory and the Northern Territory Fisheries. From that material we have specimens that we believe belong to Priolepis profunda (Weber 1909), and we redescribe that species noting differences in our material from that mentioned in the original description and comparing our material to type material. Materials and methods Institution abbreviations for material examined follow Leviton et al. (1985), except that LICPP is now BLIH, Biological Laboratory of the Imperial Household Agency, Tokyo. Measurements were taken with dial calipers or with an ocular micrometer, and all are point-to-point measurements. Methods of measurement and character definitions follow those given by Hubbs and Lagler (1958), except as follows. All fish lengths are standard length (SL), taken from tip of snout to the end of the hypural plate. Head width: maximum width at cheeks; head depth: depth of head at vertical and posterior margin of preoperculum; suborbital margin to lower margin of upper lip: shortest distance from anteroventral margin of eye to lower edge of upper lip. The longitudinal scale count is taken from the upper pectoral base along the midline to the end of the hypural plate. The transverse scale count (TRB) is taken from the anal-fin origin upward and backward to base of the first dorsal fin. In material examined lists, the institution

New Priolepis 375 catalogue number is followed by the number of specimens examined and the size in SL in parentheses. Cephalic papilla line terminology follows Hoese (1983). Two letters are used for each type of line. The first letter indicates whether the line is more-or-less vertical (V) or longitudinal (L), and the second letter denotes the orientation of the papilla axis in relation to the axis of the papilla line, either along the line (L) or perpendicular to the axis of the line (T). Data are included for types and non-types. In many cases material was poorly preserved, and it was not possible to obtain counts for all characters for all specimens, without severely damaging the specimens. Values for the holotypes and syntypes are indicated by an asterisk. The pterygiophore formula follows Birdsong et al. (1988). Priolepis Valenciennes in Cuvier and Valenciennes (1837) Priolepis Valenciennes in Cuvier and Valenciennes (1837): 67 (type species Priolepis mica Valenciennes in Cuvier and Valenciennes (1837) = Gobius semidoliatus Valenciennes in Cuvier and Valenciennes (1837), by monotypy). Zonogobius Bleeker 1874: 323 (type species Gobius semifasciatus Kner 1868 = Priolepis semidoliatus Valenciennes in Cuvier and Valenciennes (1837), by original designation). Quisquilius Jordan and Evermann 1903: 203 (type species Quisquilius eugenius Jordan and Evermann 1903, by original designation). Pleurogobius Seale 1910: 536 [type species Pleurogobius boulengeri Seale 1910 = Priolepis cincta (Regan 1908), by monotypy, an apparent error for Pterogobius Gill 1863]. Cingulogobius Herre 1927: 201 [type species Pleurogobius boulengeri Seale 1910 = Priolepis cincta (Regan 1908), by original designation]. Fig. 1 Dorsal view (top), lateral (middle) and ventral (bottom) views of sensory papillae in Priolepis akihitoi based on WAM P.26662-018, 38 mm SL, from northwestern Australia. Dashed line indicates anteriormost extent of scales Diagnosis. Head pores absent; mouth very oblique, forming an angle of about 35 60 with body axis; pelvic interspinal membrane usually absent, present in some species; pelvic fins connected by interradial membrane to form single plate; 5 branched pelvic rays; fifth usually shorter than fourth; gill-opening broad, upper attachment of membrane above pectoral fin base, lower attachment on isthmus just anterior to posterior margin of preoperculum; body covered with ctenoid scales, head, nape, prepelvic area and base of pectoral fin naked or scaled; cheek naked or rarely with a few scales dorsally; operculum scaled or naked; tongue tip truncate, sometimes with slight notch at tip; pectoral rays branched, except one or two upper rays and one lower ray; teeth in outer row of upper jaw wideset; nostrils at end of short tube; posterior nostril immediately adjacent to anterior margin of orbit; head slightly broader than deep; inner face of gill rakers on first arch with small denticles; vertebrae 10? 16; pterygiophore formula 3(22110); head papilla pattern a developed or reduced transverse pattern (Figs. 1, 2, 3, 4). Remarks. As recognized here, the genus contains the 32 previously described species from the Indo-Pacific and Atlantic (Winterbottom and Burridge 1992, 1993a, b, c; Goren and Baranes 1995). Species are found in a variety of habitats, often associated with coral, rock or rubble, ranging in depths from the intertidal to depths of 400 m. Various species have been described under the generic names Priolepis, Zonogobius or Quisquilius, which we exclude from Priolepis. Examination of the holotype (ZMA 110.952) from Indonesia and material recently collected from the Timor Sea, Australia (NTM S.13427-028) of Quisquilius macrophthalmus Weber 1909 indicates that it has a depressed head, narrow gill opening and

376 D. F. Hoese and H. K. Larson Fig. 2 Dorsal view (top), lateral (middle) and ventral (bottom) views of sensory papillae of Priolepis cyanocephala AMS I.19717, 32 mm SL. Dashed line indicates anteriormost extent of scales; arrow indicates first dorsal origin Fig. 4 Dorsal view (top), lateral (middle) and ventral (bottom) views of sensory papillae of Priolepis psygmophilia, composite based on Easter Island material. Dashed line indicates maximum forward extent of predorsal scales; arrow indicates first dorsal origin; rectangular area before first dorsal fin naked Fig. 3 Dorsal view (top), lateral (middle) and ventral (bottom) views of sensory papillae of Priolepis profunda, composite, based on syntypes and Australian material. Dashed line indicates anteriormost extent of scales; arrow indicates first dorsal origin sensory papillae characteristic of the genus Cabillus Smith 1959. This species differs from C. lacertops Smith 1959 and C. tongarevae (Fowler 1927) in having predorsal scales. The holotype of Quisquilius malayanus Herre 1936 (CAS-SU 30963) has a well-developed pelvic frenum, well-developed head pores and a slightly oblique mouth. The papillae of the lateralis system below the eye are arranged in five or six vertical rows, interrupted by a single longitudinal row characteristic of Drombus Jordan and Seale 1905. Other nominal species of Zonogobius or Quisquilius are now placed in Trimma by Winterbottom (1984, 1995). Winterbottom and Burridge (1992, 1993a, b) divided the genus into three groups based on development of sensory papilla and predorsal scales. The first group includes species with a well-developed transverse sensory papilla pattern (Figs. 1, 3). Two species are included here in this group (Priolepis akihitoi, n.sp. and P. profunda). In the second group, the sensory papilla are reduced (see Figs. 2, 4), and predorsal scales cross the midline. In this group we include Priolepis psygmophilia Winterbottom and Burridge 1993c. The third group includes species with a reduced transverse papilla pattern

New Priolepis 377 and no scales on the predorsal midline. Winterbottom and Burridge (1993c) included Priolepis psygmophilia in this group. However, recently collected material indicates that the extent of predorsal scale coverage increases with size. Priolepis akihitoi sp. nov. (Figs. 1, 5a, b) Priolepis sp.: Akihito et al. 1984: 248, pl. 239D (Izu Peninsula, Japan). Priolepis RW sp. 8: Winterbottom and Burridge 1992: 1935 (key). Priolepis sp.: Akihito et al. 2002: 1173 (Izu Peninsula, Uwakai of Ehime Prefecture, Kashiwajima Island of Kochi Prefecture, Japan). Holotype. NSMT-P 69916 (ex LICPP 19820168), 53 mm SL female, Japan, Futo, Ito City, Shizuoka Prefecture, Japan, 25 m depth, 30 June 1982. Paratypes. AMS I.29544-003, 2 specimens (45 53 mm SL), Igaya, Miyake-jima, Japan, 21 August 1976; BMNH 2005.1.25.1, 1 (41), New Caledonia, 6 8 m, R. Lubbock; BPBM 19009, 1 (48), Abe-no-hama, Miyake-jima, Japan, 22 May 1975, J. E. Randall, J. Shepart, K. Meyer and L. Bell; BLIH 19820169, 1 (19), taken with holotype; NSMT- P34314, 1 (59), Jogasaki, Izu Oceanic Park, east coast, Izu Peninsula, Honshu, Japan, 19 December 1990, M. Aizawa; QM I.37607, 1 (31), Shag Rock, eastern side off North Stradbroke Island, Queensland, Australia, J. Johnson and M. Ekins, 15 December 2005, 7 10 m depth; ROM 64553, 2 (20.5 28.0), New Caledonia, west side of Isla Redika, New Caledonia, R. Winterbottom and G. Klassen, 1991; YCM 10016, 2 (52 57), Japan, Futo, Ito City, Shizuoka Prefecture, 25 m depth. Non-type material. BPBM 1 (17.5), Wistari Reef, Queensland, J. Randall, 22 January 1973; WAM P.26662-018, 1 (38), Shark Bay, Western Australia, B. Hutchins, 10 April 1979. Diagnosis. Eyes slightly elevated resulting in a shallow pit in interorbital area and a shallow groove along dorsoposterior margin of eye. Predorsal completely covered with ctenoid scales reaching to above mideye, anteriormost scales sometimes cycloid, no naked patch immediately before the first dorsal spine. Pectoral base covered with small cycloid scales. Prepelvic area covered with 11 13 rows of small cycloid scales. Cheek usually without scales, rarely with 1 or 2 scales near dorsoposterior margin of preoperculum. Operculum with a patch of small ctenoid scales dorsally. Second dorsal rays usually I, 10; anal rays usually I, 8; pectoral rays usually 20. Transverse pattern of sensory papilla rows on cheek (Fig. 1). Pelvic fins connected to form a plate, no interspinal membrane. Head and body with narrow light bands, with dark edges; a large elongate black spot anteriorly on first dorsal fin; dorsal part of caudal fin with 3 4 short vertical dark brown bars. Mouth oblique, forming an angle of about 40 45 with body axis. Description. Based on 14 specimens. First dorsal VI* (in 14); segmented caudal rays 17* (11); branched caudal rays 7/6 (2), 7/7* (8), 8/7 (1); procurrent caudal rays 5 6 above and 5 below; gill rakers on outer face of first arch 4? 1? 12 (1), 4? 1? 13 (2). Other counts are given in Tables 1, 2, 3, 4, 5 and 6. Head more or less rounded, head depth subequal to or slightly less than width at posterior preopercular margin, length 28.8 35.2% SL. Cheeks only slightly bulbous. Head width at posterior preopercular margin 22.6 24.8% SL. Depth at posterior preopercular margin 20.7 22.7% SL. Postorbital length subequal to distance from tip of snout to posterior end of eye. Snout slightly convex in side view, rounded in dorsal view, slightly less than eye length. Mouth oblique, with lower jaw protruding slightly beyond tip of upper jaw. Posterior end of jaws under mideye in young to anterior quarter of pupil in adult; anterior margin of jaws in line with lower half of pupil in young and lower margin of eye in adult. Interorbital narrow, fleshy interorbital 1 mm in largest specimens, width about one quarter eye length. Suborbital region between eye and upper lip broad, slightly less than half eye length. Mental frenum indistinct. Anterior nostril at end of short tube about 1 nostril diameter above upper lip. Posterior nostril at end of short tube just before and not in contact with anterior margin of eye, about 2 3 nostril diameters from anterior nostril. Gill opening extends forward to below a point just before posterior preopercular margin. Gill rakers on outer face of first arch slender, posterior rakers slightly shorter than filament length to slightly longer than gill filaments; rakers on inner face of first arch and rakers on other arches short and denticulate on distal tip. Tongue tip with shallow notch or emarginate. Body compressed, tapering posteriorly. Dentition. Teeth conical and slightly curved. Teeth in outer row of upper jaw wideset and enlarged; 5 6 inner rows of small teeth tapering laterally to 2 rows; an innermost row of slightly enlarged inwardly directed teeth. Teeth in outer row of lower jaw wideset, enlarged and confined to anterior half of jaw; 4 6 inner rows of small teeth; an innermost row of slightly enlarged erect teeth, teeth just behind angle of jaw distinctly larger than other teeth in row in males. Fins. Dorsal, anal and pelvic soft rays and most pectoral fin rays branched. First dorsal fin origin above a point behind pelvic insertion; first dorsal fin low and rounded, without filamentous spines; fourth and fifth dorsal spines extend beyond others when fin depressed. Pectoral fins long, reaching to beyond middle of anus in young and to

378 D. F. Hoese and H. K. Larson Fig. 5 a Holotype of Priolepis akihitoi sp. nov., NSMT-P 69916, 53 mm SL female; b freshly collected specimen of Priolepis akihitoi, from Western Australia, WAM P.26662-018, 38 mm SL, photo by J.B. Hutchins; c live holotype of Priolepis cyanocephala sp. nov., AMS I.19717, 32 mm SL, photo by R. Kuiter; d freshly preserved holotype of Priolepis cyanocephala, AMS I.19717, 32 mm SL, photo by D. Hoese Fig. 6 a Freshly collected specimen of Priolepis profunda from off Shark Bay, Western Australia, WAM P.26188-026, 28 mm SL, photo by J.B. Hutchins; b live individual of Priolepis profunda from New Guinea, photo by R. Steene; c freshly collected specimen of Priolepis psygmophilia, BPBM 6753, 27 mm SL, Easter Island, photo by J.E. Randall; d freshly collected specimen of Priolepis psygmophilia, BPBM 17306, 35 mm SL, Rapa Island, photo by J.E. Randall

New Priolepis 379 Table 1 Second dorsal ray counts in various species of Priolepis I,9 I,10 I,11 I,12 P. akihitoi 3 11* P. cyanocephala 1 1* P. profunda 4 18* P. psygmophilia (Kermadec Is.) 11* P. psygmophilia (Lord Howe Is.) 2 P. psygmophilia (Rapa Is.) 4 26 2 P. psygmophilia (Easter Is.) 2 21 An asterisk indicates count of holotype or syntypes Table 2 Anal ray counts in various species of Priolepis I, 7 I, 8 I, 9 P. akihitoi 1 13* P. cyanocephala 2* P. profunda 22* P. psygmophilia (Kermadec IlIs.) 1 10* P. psygmophilia (Lord Howe Is.) 2 P. psygmophilia (Rapa Is.) 31 P. psygmophilia (Easter Is.) 1 22 An asterisk indicates count of holotype or syntypes Table 3 Pectoral ray counts in various species of Priolepis 18 19 20 21 P. akihitoi (Japan) 5* 3 P. akihitoi (Australia) 2 P. akihitoi (New Caledonia) 1 1 1 P. cyanocephala 2* P. profunda 2 14* 4* 1 P. psygmophilia (Kermadec Is.) 1 8* 2 P. psygmophilia (Lord Howe Is.) 2 P. psygmophilia (Rapa Is.) 4 25 2 P. psygmophilia (Easter Is.) 6 14 2 An asterisk indicates count of holotype or syntypes Table 4 Predorsal scale counts in two species of Priolepis 14 15 16 17 18 19 20 21 22 23 24 25 P. akihitoi (Japan) 1* 1 3 3 P. akihitoi 2 (Australia) P. akihitoi (New Caledonia) 1 1 1 P. profunda 2* 3* 3 3 2 2 1 An asterisk indicates count of holotype or syntypes anal origin in adult. Pelvic fins long, fused into a plate, without interspinal membrane, fins reach to above middle of anus in juveniles and to anal origin in adult, origin below Table 5 Transverse scale count in various species of Priolepis 7 8 9 10 11 12 13 14 15 16 P. akihitoi (Japan) 3 2* 1 P. akihitoi (Australia) 2 P. akihitoi (New Caledonia) 1 1 1 P. cyanocephala 2* P. profunda 3 5* 3 1 P. psygmophilia (Kermadec 7* 4 Is.) P. psygmophilia (Lord Howe 1 Is.) P. psygmophilia (Rapa Is.) 3 14 17 P. psygmophilia (Easter Is.) 3 12 7 An asterisk indicates count of holotype or syntypes Table 6 Longitudinal scale count in various species of Priolepis 22 23 24 25 26 27 28 29 30 31 32 P. akihitoi (Japan) 2 2 1* 1 1 P. akihitoi 1 1 (Australia) P. akihitoi (New Caledonia) 1 2 P. cyanocephala 2* P. profunda 3* 4* 2 3 2 P. psygmophilia (Kermadec Is.) 1 2 3 5* P. psygmophilia 1 (Lord Howe Is.) P psygmophilia (Rapa Is.) 2 4 18 5 2 P. psygmophilia (Easter Is.) 1 11 7 1 An asterisk indicates count of holotype or syntypes a point just before lower pectoral insertion, 21.6 26.0% SL, fifth ray highly branched, length about 78 90% length of fourth ray. Caudal fin with rounded margin, about 1.4 1.7 in head. Squamation. Body scales ctenoid, except for cycloid scales on midline of belly. Predorsal scales small and irregularly arranged largely ctenoid, usually with scales between posterior margin of eye and postocular transverse papilla row. Belly completely scaled. Pectoral base and prepelvic area fully covered with small cycloid scales. Upper quarter of operculum with small scales, usually ctenoid. Base of caudal fin with small ctenoid scales. Head papilla pattern. Well-developed transverse pattern. Interorbital region with papillae following rim of orbit, posteriorly with 4 6 papilla arranged in a V with the apex pointing forward. Anteriorly a similarly arranged line of papillae with V pointing backward and normally with no

380 D. F. Hoese and H. K. Larson more than 4 papilla (2 on each side). Five VT rows below eye, an upper LT row meeting fifth row, fifth row with disjunct ventral segment below LT line. A sixth oblique line extending from eye posteroventrally to near middle of operculum. A lower LT line extending from middle of jaws posteriorly meeting and normally extending beyond fifth VT line. Two lines extending from anterior interorbital row to a point just medial to posterior nostril. Outer preopercular mandibular line interrupted at posterior end of jaws, with the forward section curved upward at the posterior end; line ending at angle of preoperculum. Inner preopercular mandibular line interrupted just behind posterior end of jaws. A single row of papilla on each side of chin converging posteriorly forming a V, but not meeting; each side with 18 22 papillae. Other papillae are shown in Fig. 1. The papilla pattern of the holotype is illustrated in Akihito et al. (1984: 239). The holotype is atypical in having the fifth vertical cheek line extending well below the upper longitudinal line on the cheek. Other material from Japan agrees with the specimen illustrated here, based on a specimen from Western Australia (WAM P.26662-018, 38 mm SL). Coloration of freshly collected material. Based on photo of holotype published in Akihito et al. (1984). Head and body brown. Each scale pocket with light brown center surrounded darker pigment, giving appearance of rows of faint, diamond-shaped marks on sides of body. Head with dark brown band, narrower than pupil, from anteroventral margin of eye to posterior end of jaws. Thin dark brown vertical band, narrower than pupil, extending from posteroventral margin of eye downward on cheek. Dark brown band connecting eyes just behind middle of eye; a curved brown bar connecting posterior margin of eyes. A broad brown, oblique band running along the preopercular margin connecting to a curved band crossing head above operculum. A brown blotch dorsally on operculum connecting to dark band crossing nape just before first dorsal fin. Thin faint yellow bar crosses base of pectoral fin. Body with 7 broad brown vertical bands, about twice width of lighter interspaces; first band below front half of first dorsal fin; second band below end of second dorsal fin; third band below anterior part of second dorsal fin; fourth band below posterior half of second dorsal fin; 3 bands on caudal peduncle. Pectoral fin yellowish-gray. Distal margins of first and second dorsal fins and anal fin, pelvic fins and caudal fin with bluish-white margin first dorsal fin with large black spot basally between first 3 dorsal spines fins, followed by smaller spots centered on lower third of fourth to sixth dorsal spines. Anal fin yellow with white mottling. Caudal fin with 4 dark brown wavy yellow bands on dorsal one-quarter of fin, rest of fin orange. A photo provided by Dr. B. Hutchins of a specimen from Shark Bay, Western Australia, is similar in coloration, except that the dark bands are only slightly broader than lighter interspaces (Fig. 5b). Also the darker edges to the scale pockets are only prominent on the dark bands and not in the lighter interspaces. Coloration of preserved material. Similar to fresh coloration, except as follows: contrast between dark bands and lighter interspaces less prominent, yellow and white becoming light brown. Melanophore edging less prominent. Distribution. Priolepis akihitoi is known from Japan, Shark Bay, Western Australia, Sydney, New South Wales, Australia, Moreton Bay and the southern Great Barrier Reef, Queensland, Australia and New Caledonia from depths of 3 25 m. Etymology. Named for the Emperor of Japan, Akihito, for his significant and innovative contributions to the systematics of gobioid fishes. Remarks. This species is known under the name Kokuten-benkeihaze in Japan and Emperor Reefgoby in Australia. This species falls into the group of Priolepis with a transverse papilla pattern reviewed by Winterbottom and Burridge (1993b), included in their key as Priolepis RW sp. 8. The group also includes the recently described Priolepis winterbottomi Nogawa and Endo 2007. The species differs from all species in that group, except for P. profunda, in usually having a second dorsal count of I, 10 (versus I, 9). It differs from P. profunda in having opercular scales (versus usually no scales) and in having the dark bands dorsally on the caudal fin. In addition, the interorbital is very narrow with the interorbital papilla arranged in a V, rather than transverse. The species has been observed by the first author from Sydney, but no specimens have been collected. The single specimen from Shark Bay differs in a some features including color differences noted above and in having cycloid scales on the nape and operculum. Counts agree with other specimens of the species. The specimens may represent a separate species and are not included in the type series. Priolepis cyanocephala sp. nov. (Figs. 2, 5c, d) Holotype. AMS I.19717-001, 32 mm SL female, Clovelly, New South Wales, Australia, P. Zorn, January 1977, 12 m. Paratype. AMS I.17417-001, 30 mm SL, male, Philip Rock, Lord Howe Island, G. Allen and J. Randall, 25 February 1973, 27 m. Diagnosis. No bony interorbital trench, but eyes elevated well above the interorbital region forming a nonbony trench. Head broader than deep, distinctly depressed. Predorsal largely naked with body scales extending forward to just above pectoral origin to just above posterior

New Priolepis 381 quarter of operculum. No opercular scales. Second dorsal rays I, 10; anal rays I, 9; pectoral rays 19. Pelvic fins connected to form plate, no interspinal membrane. Each scale pocket with concentrated xanthophores along margin forming a diamond-shaped mark in life, but fading in preserved material, but with a concentration of melanophores along margins of scale pockets visible. Head with four vertical bands in life, without vertical bars on body. Cheek sensory papillae a reduced transverse pattern (Fig. 2). Mouth oblique, forming an angle of about 35 40 with body axis. Description. Morphometrics based only on holotype due to poor condition of paratype. First dorsal VI* (in 2); segmented caudal rays 17 in holotype (caudal damaged in paratype); branched caudal rays 15 in holotype; procurrent caudal rays 6 above and 5 below; gill rakers on limb of outer arch 3? 1? 13 in paratype. Other counts are given in Tables 1, 2, 3, 4, 5 and 6. Head more or less rounded, head depth subequal to width at posterior preopercular margin, length, 31.3% SL in holotype. Cheeks bulbous, slightly protruding. Width at posterior preopercular margin 24.4% SL in holotype. Depth at posterior preopercular margin 19.4% SL. Postorbital length subequal to distance from tip of snout to posterior margin of eye. Snout slightly convex in side view, rounded in dorsal view, length much less than eye diameter. Mouth oblique, with lower jaw protruding slightly beyond tip of upper jaw. Posterior tip of mouth ends under anterior margin of pupil; anteriorly, mouth extends dorsal to an imaginary horizontal line from just above lower margin of pupil. Interorbital, a narrow depression (width 0.5 mm in holotype), about one-sixth as wide as eye. Suborbital very narrow, less than one-quarter eye diameter. Mental frenum indistinct. Nostrils at end of short tube; posterior nostril adjacent to anterior margin of orbit, but not in contact with eye and separated from anterior nostril by a \2 nostril diameters; anterior nostril almost in contact with upper lip. Gill rakers pointed, much longer than wide, posterior rakers subequal to or slightly longer than filament length; rakers on inner face of first arch and rakers on other arches short and denticulate on distal tip. Tongue tip emarginate. Body compressed, tapering posteriorly. Dentition. Upper jaw with outer row of 10 teeth on each side composed of enlarged, curved, fixed, widely separated teeth; inside this row, 4 or 5 rows of small conical teeth, converging laterally to 1 or 2 rows; inside these rows an inner row of 3 4 curved, enlarged teeth on each side; teeth of middle and inner rows depressible. At anterior tip of lower jaw a row of enlarged, widely separated teeth, numbering 6 on each side, with posterior most tooth in row largest; a smaller tooth often interposed between each pair of larger ones; inside this row, several irregularly placed teeth forming about four rows anteriorly, tapering to 1 or 2 rows, laterally and posteriorly; inside these rows, a row of enlarged curved teeth anteriorly, with largest tooth at mouth angle, size of teeth decreasing anteriorly and posteriorly. Fins. Dorsal, anal and pelvic soft rays and most pectoral fin rays branched. First dorsal fin origin above a point well behind pelvic insertion. Tip of pectoral fin reaches to above anal fin origin. Fifth pelvic soft ray highly branched and long (tips broken off); pelvic fin reaches just short of anus, length 25.9% SL. Posterior rays of dorsal and anal fins subequal in length in female, but fins damaged in male paratype; posterior most ray of each fin shorter than body depth at anal origin, about two thirds body depth at anal origin. Third dorsal spines longest, but only slightly longer than first and third spine, no filamentous spines. Caudal fin margin rounded, about 1.1 in head length. Squamation. Scales largely ctenoid; those on ventral surface of belly, on base of pectoral and on prepelvic area cycloid. Prepelvic area scaled posteriorly only, just before pelvic fin. No scales on cheek or operculum. No cycloid scales at base of caudal fin. Head papilla pattern. Reduced transverse pattern. Interorbital region with papillae following rim of orbit, anteriorly a single papilla on each side and posteriorly a single papilla just behind middle of eye. A short longitudinal (LL) line from middle of upper jaw extending posteriorly with 3 papillae, each presumably representing the rudiments of a VL line, meeting a short vertical (VT) line under middle of eye, with 3 or 4 papillae, a fifth papilla along posteroventral margin of eye. A short horizontal (LT) line on cheek from middle of eye to near end of preoperculum. A lower LT line extending from middle of jaws posteriorly almost to below end of eye. Two short converging lines expending from anterior interorbital row to a point just medial to posterior nostril. Outer preopercular mandibular line interrupted at posterior end of jaws, with no deflection in line, ending at angle of preoperculum. Inner preopercular mandibular line interrupted just behind posterior end of jaws. A single row of papillae on each side of chin converging posteriorly forming a V, but not meeting. Other papillae are shown in Fig. 2. Live coloration of holotype. Head bluish-white, body white. Each scale pocket with white center surrounded by yellow pigment, giving appearance of rows of diamondshaped marks on sides of body. Head with yellow band, narrower than pupil, from anteroventral margin of eye to posterior end of jaws. Faint yellow vertical band, narrower than pupil, extending from posteroventral margin of eye downward on cheek. Yellow band connecting eyes just behind middle of eye; a curved yellow bar connecting posterior margin of eyes. A thin yellow, oblique band running along preopercular margin connecting to a curved band crossing head above operculum. A faint yellow blotch

382 D. F. Hoese and H. K. Larson dorsally on operculum. Thin faint yellow bar crosses base of pectoral fin. No bands on body. Pectoral fin yellowishgray. Distal margins of first and second dorsal fins and anal fin, and caudal fin with bluish-white margin; dorsal fins with 4 rows of yellow spots (slightly smaller than pupil diameter); anal fin yellow; caudal fin with six or seven wavy yellow bands on dorsal two-thirds of fin. Coloration of freshly collected material. Similar to live coloration, except as follows: head dark bluish-gray. Yellow spots and first dorsal fin small and faint. Spots on second dorsal fin small, much less than pupil diameter, arranged on rays, uppermost yellow spots diffuse and faint. Anal fin with little yellow pigment, largely white. Pelvic fins white. Coloration of preserved material. Head and body brown, no trace of bands on head and pectoral base. Scale pockets lined with melanophores, giving a diamond-shaped appearance. Fins clear. Distribution. The species is known only from two specimens from Sydney, New South Wales and Lord Howe Island, in depths of 12 27 m. Etymology. The specific name, cyanocephala, referring the blue head in live individuals: cyano (Greek = blue) and cephalus (Greek = head). Remarks. Priolepis cyanocephala falls into a group of species with a reduced papilla pattern and no predorsal scales treated by Winterbottom and Burridge (1993a). It differs from other species treated in that work by having an elongate fifth pelvic ray, with numerous branches, scales faintly outlined with melanophores in preserved material, strongly depressed head, high pectoral ray count (19) and thin yellow bars below the eye, which disappear in preserved material. In those characters and overall coloration, the species is most similar to P. psygmophilia Winterbottom and Burridge 1993c, which is treated below. Winterbottom and Burridge (1993c) noted differences in this species (as Priolepis DFH sp. 3) from P. psygmophilia including fewer second dorsal rays, having predorsal scales starting more posteriorly, lower dorsal and anal rays and concave interorbital (versus U-shaped). The high degree of variation in predorsal scales in this species would suggest that that feature might not be significant. Priolepis profunda (Weber 1909) (Figs. 3, 6a, b) Quisquilius profundus Weber 1909: 155 (Sapeh-Strasse, Sumbawa and Dongala, Palos-Bai, Celebes); Weber 1913: 483, fig. 100; Koumans 1953: 129, fig. 30 (as Quisquilius eugenius Jordan and Evermann 1903). Priolepis sp. 1: Kuiter and Tonozuka 2001: 688 (South of Bintang, Indonesia). Material examined. Syntypes: ZMA 110.951, 3 specimens (23 27 mm SL), Dongola, Palo-baai, Sulawesi, Indonesia, 36 m; RMNH 17153, 1 (26) Sapeh Strait, Sumbawa Island, Lesser Sunda Islands, Indonesia. Other material examined. AMS I. (as 05/83/68), 1 (30), Western Australia, off Dampier Archipelago, 36 38 m, 20 07 0 S, 117 28 0 E R/V Soela; BMNH 1889.7.20.64, 1 (25), Northern Territory, Arafura Sea 12 September 1874; CAS 53232, 1 (21), Gulf of Thailand, 29 m, 12 07 0 N, 100 48 0 E, 30 January 1960, R. L. Bolin; CSIRO (ex B.8870), 1 (25), CSIRO B.3982, 4 (22 34), Western Australia, NW Shelf, 60 m, 19 47 0 S, 117 23 0 E, 1983; NMV A.3562, 1 (30), Western Australia, North of Red Point, 114 m, 18 42 0 E, 119 37 0 E, 12 March 1981, M. Gomon; NTM S.10559-003, 2 (28 29), Western Australia, Dampier Archipelago, 53 m, 19 40 0 S, 116 50 0 E, 12 May 1982, L. Bullard; NTM S.11090-003, 4 (25 33), Western Australia, Northwest Shelf, 19 52 0 S, 117 32 0 E, 29 May 1983, L. Bullard; NTM S.11897-022, 1 (25), Northern Territory, North of Goulburn Island, 60 m, 11 April 1986, N.T. Fisheries NTM S.11924-030, 1 (30), Western Australia, north of Dampier Archipelago, 60 80 m, 20 04 0 S, 116 37 0 E, 10 June 1980, N.T. Fisheries; NTM S.12616-001, 1 (30), Western Australia, off Dampier Archipelago, NW Shelf, 36 38 m, 18 October 1983, G. Leyland; NTM S.13339-001, 1 (27), Western Australia, Joseph Bonaparte Gulf, Timor Sea, 70 m, 13 07 0 S, 128 56 0 E, 23 November 1990, N.T. Fisheries; NTM S.13542-011, 1 (32), Northern Territory, Arafura Sea, 60 61 m, 10 32.9 0 S, 134 10.7 0 E, 27 September 1992, R. Williams; NTM S.13560-008, 1 (27), Northern Territory, Arafura Sea, 61 62 m, 10 26.4 0 S, 134 19.6 0 E, 12 October 1992, R. Williams; WAM P.26188-026, 1 (28), Western Australia, 20 km E of Trimouille Island, 45 m, 20 21 0 S, 115 53 0 E, 11 May 1978, B. Hutchins on R/V Courageous. Diagnosis. Interorbital broad, with a slight depression between eyes. Predorsal scales covered with ctenoid scales to above mideye, a small naked patch immediately before first dorsal fin; no scales immediately behind eye before transverse postocular papilla row. Prepelvic area partly scaled immediately before pelvic origin, in 2 8 rows. Usually no opercular scales, but rarely 1 5 small scales dorsally. Cheek without scales. Second dorsal rays usually I, 10; anal rays usually I, 8; pectoral rays usually 19. Transverse pattern of sensory papilla rows on cheek (Fig. 3). Pelvic fins connected to form a plate, but no interspinal membrane; fifth ray with multiple branches, length about 75 80% of length of fourth ray. Head and body brown, a distinct large black blotch at anterior base of first dorsal fin, head and body with very thin blue bands in life. No bands dorsally on caudal fin. Mouth very oblique, jaws form an angle for 55 60 with body axis.

New Priolepis 383 Description. Based on 22 specimens. First dorsal VI* (21); segmented caudal rays 17* (15), 18(1); branched caudal rays 7/7* (5), 8/7 (8); procurrent caudal rays 5 7 above and 5 7 below; gill rakers on outer face of first arch 2? 1? 10 (1), 3? 1? 11 (2), 3? 1? 12 (2), 3? 1? 13 (1); lower gill rakers on outer face of second arch 11 (3), 12 (2). Other counts are given in Tables 1, 2, 3, 4, 5 and 6. Head slightly compressed, depth slightly greater than width at preopercular margin, length 29.8 33.8% SL. Cheek slightly bulbous. Head width at posterior preopercular margin 21.0 23.7% SL. Depth at posterior preopercular margin 22.4 23.8% SL. Postorbital length about equal to distance from tip of snout to posterior end of eye. Snout short, rounded in dorsal view, slightly convex in side view, slightly shorter than eye length. Anterior margin of jaws in line with middle of eye in adult; posterior end of jaws under anterior quarter of eye. Interorbital moderate greater than pupil diameter. Suborbital region between eye and upper lip narrow, subequal to pupil diameter, about half eye diameter. Mental frenum indistinct. Anterior nostril at end of short tube about one nostril diameter above upper lip. Posterior nostril at tip of short flared tube at anterordorsal margin of eye about 1 2 nostril diameters above and behind anterior nostril. Gill opening extends forward to below a point just before posterior preopercular margin. Gill rakers on outer face of first arch slender, posterior rakers shorter than gill filaments; rakers on inner face of first arch and rakers on other arches denticulate at distal tip. Tongue tip truncate to slightly notched. Body compressed, tapering posteriorly. Dentition. Teeth conical and slightly curved. Teeth in outer row of upper jaw wideset and enlarged; 4 5 inner rows of small teeth tapering laterally to two rows; an innermost row of slightly enlarged inwardly directed teeth. Teeth in outer row of lower jaw wideset, enlarged and confined to anterior half of jaw; 4 5 inner rows of small teeth; an innermost row of slightly enlarged erect teeth, teeth just behind angle of jaw slightly larger than other teeth in row. Fins. Dorsal, anal and pelvic soft rays and most pectoral fin rays branched. First dorsal fin with rounded margin, origin well behind pelvic insertion; no filamentous dorsal spines; fifth spine extends beyond others when fin depressed in males, fourth extends farthest in females. Pectoral fin long, length 30.5 31.9% SL, reaching to above first 2 4 segmented anal rays. Pelvic fins fused to a plate, no interspinal membrane, origin below a point before lower pectoral insertion, 25.4 27.6% SL, reaching posteriorly to below anus to first 1 or 2 segmented anal rays; fifth pelvic ray highly branched, subequal in length to or slightly shorter than fourth ray. Caudal fin with rounded margin, about 1.0 1.2 in head. Squamation. Body scales ctenoid, except for cycloid scales on midline of belly. Predorsal fully scaled to above middle of eye, scales small and irregularly arranged. Belly completely scaled. Pectoral base covered with cycloid scales. Prepelvic areas partly covered with 4 9 rows of small cycloid scales, with a small triangular naked patch anteriorly. Head papilla pattern. Well-developed transverse pattern. Interorbital region with two transverse rows of papillae, with 6 8 papillae in each row. No papillae along rim of eye before anterior interorbital line. Few papillae along posterodorsal margin of eye, one behind and below posterior interorbital line followed ventrally by a pair of papillae; a single papilla just below posterior margin of eye. Five VT rows below eye, an LT row meeting fifth row, fifth row with disjunct ventral segment below LT line. A sixth oblique line extending from eye posteroventrally to near middle of operculum. A lower LT line extending from middle of jaws posteriorly meeting fifth VT line, sometimes extending slightly behind the fifth VT line. An oblique line of papillae below upper lip, from first VT line to end of jaws. Usually 2 short LT lines extending forward from first VT line. Two lines extending from anterior interorbital row to a point just medial to posterior nostril. Outer preopercular mandibular line interrupted at posterior end of jaws, ending at angle of preoperculum. Inner preopercular mandibular line continuous from chin to dorsoposterior edge of preoperculum. Chin papillae in 2 lines tapering posteriorly to form a V; each side with around 15 18 papillae. Other papillae are shown in Fig. 3. Live coloration of specimen from New Guinea (based on photo provided by R. Steene, see Fig. 6b; the specimen was not collected, and identification is based on comparison with freshly collected specimen listed below). Head and body greenish-brown, belly orange. Head with very thin white bands (bands less than a half pupil diameter), edges of bands blue; first band extending from below anteroventral margin of eye to behind middle of upper jaw; second extending vertically from mideye across cheek, third extending from below posterior margin of eye across cheek to angle of preoperculum, dorsally connecting to a curved band crossing top of head. Fourth vertical band on mid operculum connecting dorsally to a curved bar crossing top of head. Body with 8 very thin white vertical or slightly oblique bands sloping downward and backward; first band below first dorsal origin; second band below fourth dorsal spine, connecting to oblique bar extending forward on lower quarter of first dorsal fin; third band below second dorsal origin; fourth and fifth bands below second dorsal fin; sixth band below a point just behind end of second dorsal fin; seventh band on middle of caudal peduncle and eighth band at end of caudal peduncle extending over base of caudal fin. Pectoral base with thin

384 D. F. Hoese and H. K. Larson white oblique band extending full depth of pectoral-fin base. First dorsal fin with dusky area extending along membranes just above bases of first three dorsal spines; distinct white band on lower quarter of fin connecting to band below middle of fin and paler grey band above connecting to body band between dorsal fins. Second dorsal pale yellowish-orange, with small white spots forming oblique stripes; five yellowish-brown stripes on lower half of fin; distal quarter of fin grayish-orange. Pectoral fin a translucent orange. Caudal fin yellowish-orange, with distal white margin. Kuiter and Tonozuka (2001) have given a live picture of this species agreeing in general with the description above, except that the fins are all a bright yellow. Coloration of freshly collected material (based on photo of 28 mm SL male provided by B. Hutchins, WAM P.26188-026, Fig. 6a). Head and body a dusky orange, paler ventrally, belly whitish; head and body densely covered with minute brown spots, dark spotting denser along scale pockets on dorsal scales. Head with very thin bluish white bands (bands less than a quarter pupil diameter), edges of bands dark blue; first band extending from below anteroventral margin of eye to middle of upper jaw; second extending vertically from mideye across cheek, third extending from below posterior margin of eye across cheek, well in front of posterior preopercular margin, dorsally connecting to a curved band crossing top of head. Fourth vertical band on mid operculum connecting dorsally to a curved bar crossing top of head. Body with 8 very thin bluish-white vertical or slightly oblique bands sloping downward and backward; first below first dorsal origin; second below fourth dorsal spine; third below second dorsal origin; fourth and fifth below second dorsal fin; sixth below a point just behind end of second dorsal fin; seventh on middle of caudal peduncle and eighth at base of caudal fin. Pectoral base with a broken thin bluish-white oblique band. First dorsal fin with distinct irregularly shaped black spot extending along membranes just above bases of first three dorsal spines. Small white spot between second and third dorsal spines along margin of black spot, followed dorsally by small black spot; remainder of fin with scattered orange pigment, paler on upper half of fin. Second dorsal pale yellowish-orange, with dense small clear spots; three rows of yellowish-brown dark spots forming rows on lower half of fin. Anal fin similar to second dorsal fin, but without brown spots. Pelvic fin whitish-orange. Pectoral fin a translucent orange. Caudal fin yellowish-brown basally and dusky distally. Coloration of preserved material. Head and body dark brown. Scales uniformly colored, without dark pigment around scale pockets. Interorbital with 3 faint transverse brown bands. A thin brown band between anterior nostrils, obscure in some specimens. Head with thin, faint, dark band, much narrower than pupil, from just behind anterior margin of eye to posterior end of jaws. Faint dark band, narrower than pupil, extending downward on cheek from below middle of eye. A thin dark band extending posteroventrally from end of eye to angle of preoperculum, band continues dorsally across nape to meet band on other side of head. Sometimes with a short band from upper pectoral insertion extending toward, but not meeting posterior nape band operculum with a thin dark band extending across nape to join band on other side of head. Typically no bands on body, but sometime 1 or more bands, first from dorsal origin extending ventrally to pectoral fin; second from middle of first dorsal fin, extending ventrally to sides of belly; third from second dorsal origin to anal origin; fourth from middle of second dorsal fin extending ventrally to midside; fifth from near posterior end of second dorsal fin, extending ventrally onto midsides; sixth from just behind second dorsal origin onto midsides; seventh from just behind middle of caudal peduncle onto midsides; eighth at base of caudal fin, most distinctive dorsally. Faint dark bar extending from dorsoanterior part of pectoral base obliquely to middle of pectoral fin base. A large black blotch basally on first dorsal from first to third spine, base of fin grey behind spot, above band a clear area followed dorsally by a short dark brown to black horizontal band extending about half way along fin, blotch more intense anteriorly in male giving appearance of a dark spot when viewed from above. Other fins grey to white. Distribution. Priolepis profunda is known only from Thailand, northwestern Australia, Indonesia, New Guinea and Samar Sea, the Philippines (based on photo provided by E. Murdy), from depths of 45 114 m, but is probably more widespread. Remarks. The identification of this species is tentative. The black anteriorly on the first dorsal spine was not mentioned in the original description, and the spot is not apparent on the syntypes. Similarly, the original description mentions dark spots forming irregular bands on the caudal fin, which are not present in any material, including freshly collected material. Also the bands shown in a subsequent figure (Weber 1913) shows light bands that are much broader than those in material examined here. All of the syntypes are females, and in females from Australia, the black pigment is less intense and part of a band at the base of the fin. The dark spots on the second dorsal fin fade in preserved material, and it is possible that females might have those spots in life. The material examined here agrees in other features with the syntypes. Most specimens examined lack scales on the operculum, but one had a single scale and another specimen 5 small scales dorsally on the operculum.