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A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarification of Cryptocentrus leptocephalus and C. melanopus.

INTRODUCTION

Cryptocentrus contains 34 described and recognised species. A few species, which live around coral reefs, are generally well known, but most of the species live in silty environments adjacent to reefs, mangroves or on the continental shelf. Many are known only from a few trawled specimens. Species found in shallow waters are known to live with alpheid shrimps. Information is presented here to clarify the identity of some of the previously described species, which have been confused or believed incorrectly to be undescribed. Three of the species treated here live adjacent to coral reefs, but often in silty waters well away from coral. One of these species is well known and often photographed, and typically is referred to as the ventral barred goby. Workers have suspected that it represented an undescribed species, but evidence is presented here that it was described over 75 years ago.

Cryptocentrus is superficially similar to Amblyeleotris, and the two genera are often difficult to separate in the field. Hoese and Steene (1978) provided diagnoses of the two genera and noted that Cryptocentrus has the upper longitudinal cheek papilla row extending forward to the second or third transverse row, under the eye (vs. under fifth or sixth row); the mandibular papillae are arranged in two parallel rows extending posteriorly on the sides of the chin (vs. a single papillae set in a pit at each side of chin); the gill opening is more restricted, usually below or in front of the posterior preopercular margin (vs. behind the margin); scales are normally cycloid, except for 2 or 3 species (vs. always with some ctenoid scales); dorsal and anal rays vary from I,9 to 12 (vs. I,12-19). Randall (2004) incorrectly reported the papillae set in pits, first reported by Hoese & Steene (1978) on the chin, as being behind the upper jaw. He also reported it as unique, but the same pattern found in Amblyeleotris occurs in Ctenogobiops, as illustrated by Randall, Shao & Chen (2003) and Vanderhorstia, as illustrated by Shibukawa & Suzuki (2004). Hoese and Larson (2004) provided additional diagnostic features for Cryptocentrus. Cryptocentrus species generally have a larger mouth reaching well behind the eyes (versus usually ending below eye) and most Cryptocentrus species are deeper bodied and have a proportionally larger head, than those of Amblyeleotris. In Cryptocentrus the lower rakers on the inner face of the first arch and outer face of the second arch are not ossified, but in Amblyeleotris they are ossified. Unossified rakers are rare in gobioid fishes, but are also known in Callogobius.

METHODS

Methods of counts and measurements follow Hoese & Larson (2004). Institution abbreviations for material examined follows Leviton et al. (1985). After the institution catalogue number the number of specimens is given followed by the size range in mm SL (standard length). Counts for the holotype are marked with an asterisk. All measurements were taken with dial callipers and expressed as range of percent standard lengths.

Cryptocentrus sericus Herre, 1932

Figs 1-5

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

Cryptocentrus sericus Herre, 1932: 440 (type locality: Canton market, China).

Cryptocentrus sp. 1 - Akihito in Masuda et al. 1984: 259, pl 243L (Japan, hoshizorahaze).

Cryptocentrus sp. 3 - Okamura & Amaoka 1997: 608 (Japan).

Cryptocentrus sp. - Hayashi et al. 1990: 135, pl. 2, fig. 4 (Amami-oshima Island, Ryukyu c new Japanese name of Futahoshi-takanoha haze); Hayashi & Shiratori 2003: 133 (Japan, ventral barred goby); Kuiter & Debelius 2006: 653 (Indonesia, Ventral-Barred Goby).

Cryptocentrus sp. B. - Myers 1999: 240, pl.152G, H (Yaeyamas and Palau); Suzuki et al. 2004: 322 (Japan).

Cryptocentrus n.sp. 1 - Larson, Jafaar & Lim 2008: 148 (Singapore).

Material Examined: HOLOTYPE: CAS-SU 25725, 75.5 mm SL female from fish market at Canton, China. OTHER MATERIAL. Hong Kong, China NTM (uncatalogued) 2(66-70): Australia: AMS I.19474-003, 1(39), Linnett Reef, Queensland, 25 m; WAM P.31650-006, 3(23-67), Cassini Island, Western Australia, 20-30 m. Philippines: WAM P.32882-007, 1(55), Palawan, Pinasil Island, 15-17 m; WAM P.33000-003, 1(58), Palawan, Imorigue Channel, 12-15 m. Indonesia: WAM P.31303-004, 1(43), Bintan Island, Carston's reef, 12 m; WAM P.31525-007, 1 (68), Sumatra, 15-40 m; WAM P.32814-002, 1(38), Papua, Pulau Batuputih, Fak Fak coast, 3-5 m. Brunei: WAM P.33030-003, 2(35-42), Brunei patches east, 12-15 m. Thailand: AMS I.42904-003, 1(71), Ka Island, east coast; AMS I.42911-001, 3(54-79), Pu Island, northwest coast; AMS I.42911-002, 2(39-40).

In addition, 41 distribution records of this species can be found under the name Cryptocentrus sp. 2 at FishPix of the Kanagawa Prefectural Museum of Natural History, Odawara and the Museum of Science and Nature, Tokyo (http://fishpix.kahaku.go.jp/fishimage-e/search.html), as follows: Amami-oshima Island, Ryukyu Islands, Japan: KPM-NR 36538; KPM-NR 40704. Ishigaki-jima Island, Yaeyama Group, Ryukyu Islands, Japan: KPM-NR 81680 (yellow phase). Iriomote-jima Island, Yaeyama Group, Ryukyu Islands, Japan: KPM-NR 81680. Okinawa Island, Ryukyu Islands, Japan: KPM-NR 7112, 25 m; KPM-NR 11688, 20m (yellow phase); KPM-NR 15880, 20m (yellow phase); KPM-NR 28237, 25m; KPM-NR 52478; KPM-NR 61359 (yellow phase); KPM-NR 69230, 3 m; KPM-NR 80827, 7 m. Zamami-jima Island (Kerama Group), Ryukyu Islands, Japan: KPM-NR 42170; KPM-NR 87476, 17 m. Hong Kong: KPM-NR 34758; KPM-NR 34759; KPM-NR 34760 (yellow phase); KPM-NR 34761 (yellow phase); KPM-NR 34762; KPM-NR 34763; KPM-NR 64667; KPM-NR 64669; KPM-NR 64712, (yellow phase); KPM-NR 69167, 3m; KPM-NR 69178, 2m; KPM-NR 69187, 4m; KPM-NR 69191, 4m; KPM-NR 69192, 4m, (yellow phase); bKPM-NR 69196, 4m. Negros Island, Philippines: KPM-NR 35855 (yellow phase); KPM-NR 35856 (yellow phase); KPM-NR 35857; Boracay Island, Philippines: KPM-NR 61019 (yellow phase); KPM-NR 80440, 15m (yellow phase). Sulawesi, Indonesia: KPM-NR 11476, 15-20m; KPM-NR 85968, 6-17m. Sabah, Borneo: KPM-NR 27873, 1m; KPM-NR 31690, 22 m.

Diagnosis. Mental frenum absent. Mouth large; ending well beyond end of eye (by about one pupil diameter or slightly less); jaws forming angle of 22-28[degrees] with body axis; upper margin of upper jaw in line with point about one-quarter to one third pupil diameter below eye. Cheeks slightly bulbous. Fleshy interorbital very narrow, much less than half pupil diameter. Gill opening reaching to below a point just under posterior preopercular margin. Head papillae minute and not on distinct ridges. Scales entirely cycloid. Predorsal area partly scaled, with scales reaching forward to just before posterior preopercular margin to midway between eye and posterior preopercular margin. Cheek naked or with one or two minute embedded cycloid scales; operculum naked or with 1-5 small embedded cycloid scales. Pectoral fin base usually naked, sometimes with small patch of scales centrally. Prepelvic area partly to fully scaled with 6-16 rows before pelvic fin origin. Belly fully covered with cycloid scales, except for small naked area just behind pelvic insertion. First dorsal fin moderately high, with truncate margin, first spine longest but not distinctly prolonged into filament. Cheek often with two short black stripes, one just above posterior end of jaws and second just above lower stripe. First dorsal fin with two black, oblique stripes. Pelvic fin with 4-6 curved black bars in females, fin usually uniformly dark in males or with one or two darks bands posteriorly. Pelvic fin large; reaching to just short of anus. Second dorsal-fin rays usually I,10; anal fin rays usually I,9; pectoral fin rays 16-17; longitudinal scale count 68-74; predorsal scale count 20-27; transverse scale count (TRB) 21-27.

Description: Based on 16 specimens, 28-79 mm SL. Counts of holotype indicated by asterisk. Numbers in parentheses after counts indicate the number of specimens with that count.

First dorsal spines VI(16)*; second dorsal rays I,9(1), I,10(14)*, I,11(1); anal rays I,9(16)*; pectoral rays 16(3), 17(13)*; longitudinal scale count 64(2), 65(2), 67(1), 68(1), 70(1)*, 71(1), 74(1), 78(1); predorsal scale count 18(1), 19(1)*,20(3), 22 (4), 23(1), 24(1); transverse scale count (TRB) 21,(1), 24(2), 25(2), 26(4)*, 27(1); gill rakers on outer face of first arch 2+10(1), 3+1+10(1), 2+1+11(1), 2+1+12(1), 4+1+12(1)*; gill rakers on outer face of second arch 3+1+12(1), 3+1+13(1), 4+1+13(1), 3+1+15(1)*; segmented caudal rays 9/7(1) (2), 9/8(13)*; branched caudal rays 7/6(12)*, 7/7(2); vertebrae 10+16 (holotype).

Head strongly compressed. Snout rounded in dorsal view; steeply oblique (slightly convex) in side view. Eye large and elevated, with shallow groove behind, about twice snout length. Anterior nostril at end of short tube, with broad flat rim, just above upper margin of upper lip. Posterior nostril a large pore about one nostril diameter behind anterior nostril, and one nostril diameter from eye. Preoperculum long, distance from end of eye to upper posterior preopercular subequal to distance from snout to an eye length behind eye. Postorbital long, slightly shorter than distance from tip of snout to posterior preopercular margin. Body slender, depth at anal origin 16.7-18.1% SL. Upper jaw long, reaching about one pupil diameter behind eye (13.3-16.1% SL). Teeth conical and curved. Teeth in upper jaw: outer row of teeth curved, enlarged and wide-set, teeth larger near angle of jaw; three to four rows of smaller, depressible teeth anteriorly, tapering to two to three rows posteriorly, teeth in rows pointing inward into mouth; an innermost row of one to two enlarged backwardly directed teeth anteriorly on each side of jaw. Teeth in lower jaw: teeth in outer row curved, conical, wide-set, covering anterior end of dentary only, three to four inner rows of smaller conical teeth anteriorly and two to three rows posteriorly, innermost row of teeth enlarged and larger than teeth in outer row, with largest teeth just behind bend in dentary. Tongue tip rounded. Gill rakers on outer face of first arch slender, denticulate on posterior margin, rakers much shorter than filament length. Rakers on inner face of first arch and other arches short and denticulate at distal tip. Body covered with cycloid scales. Head largely naked, midline of nape with scales extending to above preoperculum, operculum and preoperculum naked or with a few small embedded scales. Adpressed length of first dorsal fin 22.2-28.9% SL, origin just above pelvic fin insertion. Second dorsal fin base 29.3-32.8% SL. Anal fin base 20.0-23.8% SL. Pectoral fin with rounded margin, reaching to above or just before pelvic fin tip, to or just before anus in adults, length, 22.2-28.0% SL. Pelvic disc large, reaching to anus or slightly beyond, but not to anal fin origin; pelvic length 23.3-29.0% SL in adult. Caudal fin with rounded posterior margin, central rays longest; length 26.7-32.7% SL.

Head pores. Posterior nasal pore slightly median to posterior nostril; median anterior interorbital pore just before eyes; median posterior interorbital above posterior margin of pupil; postorbital pore behind upper quarter of eye; infraorbital pore below and behind postorbital pore, behind and in line with upper margin of pupil; lateral canal pore above middle of preoperculum; terminal lateral canal pore above posterior preopercular margin; a long tube with pores at each end above operculum; three preopercular pores, upper in line with point just below middle of eye; middle pore midway between upper and lower pores.

Papillae, based largely on holotype. Head papilla pattern transverse (Fig. 4). Cheek with five VT lines; first from below middle of eye to middle of jaws; second incomplete and extending upward from just behind first row, not meeting eye, ending just below or before upper LT line; third interrupted by upper LT line; fourth slightly oblique cut into two parts by upper LT line with upper part nor meeting upper LT line; fifth a short oblique line before infraorbital pore and a vertical section ventrally below upper LT line. Upper LT reaching to near posterior preopercular margin, lower LT line ending under or just behind fifth VT line. An oblique line extending above upper jaw toward anterior nostril. Three short oblique lines from anterior part of upper jaw to first VT line. Preopercular-mandibular series with outer LT line interrupted just behind upper jaw; inner line composed of single row of papillae, not interrupted behind jaws. A transverse (TT) line behind each eye. Chin papillae arranged in two posteriorly converging LT lines.

[FIGURE 4 OMITTED]

Coloration in alcohol. Head and body light brown. Head with two dark brown to black, short, horizontal stripes above the posterior two-thirds of the upper lip, faint or absent in specimens above 55 mm SL; a dark brown oval spot about half pupil diameter at posterior end of preoperculum, followed by similar sized spot anteriorly on operculum, spots often fused to form larger oval spot or short stripe, often faint or absent in specimens over 55 mm SL; dark brown oval spot, oriented obliquely on upper operculum; dark brown to black stripe above; sometimes with oval dark spot, height about half pupil diameter behind eye; short horizontal thin black stripe above posterior preoperculum, sometimes absent; similar short horizontal stripe above operculum, contiguous with lighter brown band crossing nape; cheek with numerous scattered small light brown oval spots. Body with two vertical brown bands below first dorsal fin, often with irregular margins and sometimes partly interconnected and forming mottling; from second dorsal origin posteriorly 5 vertical oval brown spots present along midside, first just behind first dorsal fin origin, second below middle of second dorsal fin, third below posterior end of second dorsal fin, fourth anteriorly on caudal peduncle and fifth just before base of caudal fin, spots elongating into bands extending from near base of second dorsal fin to base of anal fin in specimens above 55 mm SL; usually with very thin brown vertical lines between broader body bands; fins dusky, pelvic fin with 3-5 dark bands in females and 1-3 in males, often very faint in preserved material, particularly in males.

Live Coloration. Basic color pattern as shown in Figures 1-3. Some underwater photos (Fig. 5, Fishpix and photo from H. Debelius) show an overall yellow or yellowish-orange coloration of the head, body and fins. In these the dark spots on the midside are faint, but visible. Coloration of freshly collected specimens differs little from the live coloration. Yellow and banded color phases have been observed and collected on the Great Barrier Reef of Cryptocentrus cinctus Herre and these have been observed to change color (R. Kuiter, pers. comm.).

[FIGURE 5 OMITTED]

Variation. Besides the live coloration variation and sexual dimorphism mentioned above, the largest specimens from Thailand and Hong Kong differ slightly in having the first two dorsal spines more elongate and some have the midside spots larger, appearing as bands in specimens above 55 mm SL. In addition the dark cheek spots and bars are faint or absent in the largest specimens from Thailand and in one specimen from Hong Kong. These specimens, however, are the largest specimens examined and the differences are probably related to size. Photos of 54 mm SL male from Thailand show a very similar coloration to the specimen from the Great Barrier Reef shown here. Similarly, a 67 mm SL specimen from Western Australia has bands on the side narrower than in specimens from Thailand, but with the dark cheek bars and spots distinct. A 68 mm specimen from Sumatra has dark body bands as broad as in larger specimens from Thailand, but with distinct dark cheek stripes and spots. Photos of freshly collected specimens from Thailand indicates that specimens with broad bands on the body and faint or no cheek dark marks are the xanthic form. Helmut Debelius provided information indicating he believed the two forms to represent a single species from observations in the field.

Distribution. The species is widely distributed in the Western Pacific (Western Australia and the Great Barrier Reef in Australia, New Guinea, Brunei, Thailand, Malaysia, the Philippines, Indonesia, Palau, China and Ryukyu Islands, Japan) and is known to be associated with alpheid shrimps (Hayashi & Shiratori 2003). It occurs over a depth range of 1 to at least 30 m.

Relationships. Hoese & Larson (2004) recognised five distinct species complexes of Cryptocentrus. The groups were based on various morphological and color characteristics. The Cryptocentrus cryptocentrus complex is distinctive in having a bony ventral projection from the operculum; the C. strigilliceps complex is distinctive in having transverse rows of mandibular papillae and ctenoid scales on the body; the C. bulbiceps complex is distinctive in having a wedge-shaped patch of predorsal scales and the head with several oblique thin lines sloping backward and upward; the C. pavoninoides complex is distinctive in having 8-10 short rows of papillae radiating from the eye along the ventral and posteroventral margins of the eye onto the cheek; and the C. leucostictus complex is distinctive in having a very slender body, with a white stripe on the midline of the head, often extending onto the body. The various complexes provide convenient grouping, but these may not necessarily represent monophyletic groups. The sixth group possessed no unifying characterists other than lacking features of the other groups. The species in this group include: Cryptocentrus caeruleopunctatus (Ruppell, 1830), Cryptocentrus cinctus (Herre, 1936), Cryptocentrus fasciatus (Playfair, 1866), Cryptocentrus geniornatus Herre, 1935, Cryptocentrus leptocephalus Bleeker, 1876, Cryptocentrus lutheri Klausewitz, 1960, Cryptocentrus melanopus (Bleeker, 1860), Cryptocentrus shigensis Kuroda, 1956, Cryptocentrus sericus Herre, 1932 and Cryptocentrus yatsui Tomiyama, 1936. All of the species in this group have similar meristic features of 16-18 pectoral rays and longitudinal scale count of 60-80 and the head usually with small pearly white spots. The species described here falls into this last group. Cryptocentrus sericus differs from all other species in the group, except C. fasciatus, in having predorsal scales on the midline. Cryptocentrus sericus differs from C. fasciatus (see Fig. 6) in having three preopercular pores (vs. 2), narrow dark bars or oval spots along the midside (versus 5 dark vertical bands extending from back to ventral surface of body, often with irregular margins), blue cheek spots rounded (vs. obliquely elongated) and anterior margin of first dorsal fin elevated, with first two dorsal spines the longest (vs. truncate to rounded first dorsal margin with third or fourth spine the longest).

[FIGURE 6 OMITTED]

IDENTITY OF CRYPTOCENTRUS LEPTOCEPHALUS.

It should be noted that there two color phases typically referred to as Cryptocentrus leptocephalus. Russell, Fraser & Larson (2010) noted that Cryptocentrus melanopus, described from a Castelnau painting of a specimen from Singapore belongs to the group and suggested that it was a senior synonym of Cryptocentrus leptocephalus. While further work is needed on the group, Cryptocentrus melanopus matches one of the color forms with large red spots on the head; an elongate red stripe on upper pectoral fin base; a broad red bar from the ventral margin of the eye to the middle of the upper jaw, rarely broken into two parts; and the first dark body band below the second dorsal fin sloping anteriorly ending entirely before anal fin, followed by 4 thin oblique bands. The Castelnau painting shows only 3 oblique bands behind the first band below the second dorsal fin, but agrees in other features with the species referred to here as Cryptocentrus melanopus. In some material of Cryptocentrus melanopus from Australia some of the bands may fuse under the second dorsal fin, resulting in three bands on the body behind the second dorsal fin origin. Cryptocentrus melanopus has also been described as Cryptocentrus cheni Herre, 1933 and Smilogobius obliquus Herre, 1934. Cryptocentrus melanopus has sometimes been identified as Cryptocentrus singapurensis (Herre, 1936). The figure of the holotype given by Herre (1936) generally matches C. melanopus in having the first band under the second dorsal fin ending before the anal fin and in having a very dark area posteriorly with two or three bands. However, the drawing shows small spots as in C. leptocephalus. The specimen

now labelled holotype (CAS SU 29807) is faded, but appears to have 4 bars beginning from the second dorsal origin to the caudal peduncle, the body pale posteriorly and the first bar in contact with the anal spine, as in Cryptocentrus leptocephalus. The current number of apparent paratypes (CAS SU 16963, 35 specimens; BMNH 1937.9.22, 1 specimen and ZMH H.2864, 2 specimens) is greater than the 33 listed by Herre and it is possible that there has been a mixup in the specimens. However, it is more likely that Herre's drawing was actually a composite based on the holotype and paratypes. Both species are included in the paratype series. Because of this uncertainty and the confusion of separating the two species, we use the older name Cryptocentrus melanopus here (Figs 7-9: note Fig. 9 is on the back cover). Cryptocentrus leptocephalus (including the holotype RMNH 4665) has more numerous and smaller spots on the head; small round spots on upper pectoral fin base; no broad bar below the eye to the jaws; and the first band below the second dorsal fin partly overlapping the anterior base of the anal fin, followed by three sloping bands (Figs 10-12). Males of both species have spots on the dorsal fins, while females have red bands on the membranes between the fin rays and spines.

[FIGURE 7 OMITTED]

[FIGURE 8 OMITTED]

[FIGURE 10 OMITTED]

[FIGURE 11 OMITTED]

[FIGURE 12 OMITTED]

In Australia Cryptocentrus leptocephalus has been collected from mangroves on coral reefs, while Cryptocentrus melanopus has been recorded from sandy reef flats, with algae. Johnson & Gill (2006) recorded and illustrated C. melanopus from the Gulf of Carpentaria (as C. leptocephalus). Both differ from Cryptocentrus sericus in having more anal rays (I,10), no predorsal scales and oblique dark bands on the side of the body.

ACKNOWLEDGEMENTS

Barry Russell kindly provided information and photos of Castelnau's painting of Cryptocentrus melanopus. Bill Eschmeyer (CAS), Sue Morrison (WAM), Anthony Gill and James Maclaine (BMNH), Marinus Boeseman (RMNH), Madam Bauchot (MNHN) and Katsusuke Meguro all provided access to type and other material of Cryptocentrus. Rick Winterbottom kindly provided three photos used here and photos of freshly collected specimens of Cryptocentrus sericus from Thailand and Palau. Mark Mohlmann provided material of Cryptocentrus sericus from Thailand taken by Ukkrit Satapoomin. Rudie Kuiter and Gerald Allen collected Australian material. Roger Steene and Jack Randall provided underwater photos of Cryptocentrus sericus from Australia, New Guinea and Palau and Roger Steene provided underwater photos of Cryptocentrus melanopus. Helmut Debelius provided photos and information about Cryptocentrus sericus. Hiroshi Senou kindly provided us with access to and use of information from Fishpix.

Received: 12 March 2011 - Accepted: 13 May 2011

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Doug Hoese (1), Koichi Shibukawa (2) and Jiro Sakaue (3)

(1.) Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. Email: doug.hoese@austmus.gov.au

(2.) Nagao Natural Environment Foundation, 3-10-10 Shitaya, Taito-ku, Tokyo 110-0004, Japan. E-mail: kshibukawa@nagaofoundation.or.jp

(3.) Southern Marine Laboratory, P.O. Box 1598 Koror, Republic of Palau 96940. E-mail: sakaue01@sml-palau.co
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Author:Hoese, Doug; Shibukawa, Koichi; Sakaue, Jiro
Publication:aqua: International Journal of Ichthyology
Article Type:Report
Geographic Code:8AUST
Date:Jul 10, 2011
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