Gnatholepis anjerensis ( Bleeker, 1851 )

Larson, Helen K. & Buckle, Duncan J., 2012, A revision of the goby genus Gnatholepis Bleeker (Teleostei, Gobiidae, Gobionellinae), with description of a new species, Zootaxa 3529 (1), pp. 1-69 : 15-21

Gnatholepis anjerensis ( Bleeker, 1851)

( Figs 1 , 4–7 , 11B ; Tables 5 –9)

Gobius anjerensis Bleeker, 1851: 251 , pl. 1, fig. 11 (Anjer, Java Boenakeng, off Ujung Padang, Sulawesi, Indonesia).

Gobius capistratus Peters, 1855a: 251 and 1855b: 443 (Ibo, Mozambique).— Klunzinger 1871: 476–477 (Red Sea); Jatzow and Lenz 1898: 507 ( Zanzibar).

Gobius deltoides Seale, 1901: 125 (Agaña, Guam, Mariana Islands).— Koumans 1940: 156–157; Böhlke 1984: 105.

Gnatholepis deltoides —Yoshino in Masuda et al. 1984: 252 (Yaeyama Islands); Larson and Murdy 2001: 3601; Kuiter and Tonozuka 2001: 672, figs A–C (Bali); Nakabo 2002: 1212.

Gnatholepis anjerensis — Yoshino in Masuda et al. 1984: 251 (Ryukyu Islands); Wass 1984: 28 ( Samoa); Randall and Randall 1987: 309 (Enewatak Atoll); Randall and Goren 1993: 13 ( Maldives); Goren and Dor 1994: 64 (Red Sea); Allen and Smith–Vaniz 1994: 16 (Cocos (Keeling)); Randall 1995: 336 ( Oman); Myers 1999: 255, plate 161D ( Micronesia); Fricke 1999: 512 (Mascarene Islands); Larson in Randall and Lim 2000: 637 (South China Sea region); Hutchins 2001: 43 (Western Australia); Larson and Murdy 2001: 3601; Randall and Greenfield 2001: 3; Kuiter and Tonozuka 2001: 671, figs C, D (Bali, Indonesia); Nakabo 2002: 1212; Hayashi and Shiratori 2003: 96, fig. 176; Allen and Adrim 2003: 58 (Papua to Sumatra); Manilo and Bogorodsky 2003: S118; Myers and Donaldson 2003: 638 (Marianas); Senou et al. 2004: 244–245; Randall et al. 2004: 27 ( Tonga); Heemstra et al. 2004: 3329 (Rodrigues); Hoese and Larson 2006: 1654; Williams et al. 2006: 259 (Wallis Islands); Kimura et al. 2009: 276 (west coast of Thailand); Fricke et al. 2009: 101.

Acentrogobius cauerensis —Dor 1984: 237 (Red Sea).

Gnatholepis cauerensis —(in part) Maugé 1986: 369 (Transkei northwards); Winterbottom and Emery, 1986: 35 (Chagos Archipelago); Myers 1991: 238, plate 124A ( Guam); Kulbicki and Williams 1997: 22 (Ouvéa Atoll, New Caledonia).

Gnatholepis sp. —Randall et al. 1993: 387 ( Midway Atoll).

Gnatholepis sp.1 — Kuiter and Tonozuka 2001: 671, fig. (Mabul, Malaysia).

Diagnosis. A large stocky Gnatholepis (up to 84 mm SL) with ctenoid scales on head, predorsal midline and pectoral fin base, distinct flap present at end of lower lip; at least three dark spots along first spine of first and second dorsal fins, followed by at least three rows of dark spots or short streaks, pectoral fin with rows of fine dark and white speckles; second dorsal and anal fin rays nearly always I,11; pectoral rays 14–17, modally 16; lateral scales 25–29, modally 27; predorsal scales 7–13, usually 9–10, mostly ctenoid, may be cycloid anteriorly or toward midline.

Material examined. SOUTH AFRICA: SAIAB 2576 , 1 (84), pool at Thompson’s Bay , Kwa-Zulu Natal , R. van Elst , 30 May 1972 . MOZAMBIQUE: SAIAB 19636 , 3 (41–52), Inhaca Island , 5 August 1948 ; SAIAB 16933 , Naca , 27 August 1956 ; SAM 28944, 5 (26–50.6), Ilha Magaruque , S of Bazaruto, B. Kensley and party, 24 March 1973 . KENYA: NTM S.13965-005, 1(45.5), Gazi Bay , H. Coene, July–August 1993 . MADAGASCAR: USNM 260667 , 12 (31–51), Tulear [Toliara] Harbour, 11 August 1964 ; AMS I.28108-038, 4(18.5–44), Andilans Beach, Nosy Be, 20 September 2005 . RODRIGUES: SAIAB 70332 , 1 (51.5), Yacht Casse, P. Heemstra and party, 21 October 2001 . MAURITIUS: SAIAB 2112 , 5 (26–51), La Morne Brabant, THF-SA-28,7 March 1971 ; SAIAB 2036 , 3 (39–44), La Morne Brabant, 6 March 1971 ; SAIAB 2317 , 4 (37–45.5), 2 miles N of Trou d’Eau Douce , T.H. Fraser , 27 March 1971 ; SAIAB 57877 , 3 , S of Pointe Petite Riviere , 22 April 1995 ; SAIAB 57889 , 2 (42–46), tidepools at end Avenue Victory, Albion, 25 April 1995 . EGYPT: USNM 313779 , 1 (43), bay on NW end of Giftun Al Kebir Island, Hurghada, Strait of Jubal , Egypt, H.A. Fehlmann and party, 3 January 1965 . ISRAEL: AMS I.21874-001, 2(35.5–39), Shurat el Mankta, Gulf of Aqaba , L. Fishelson, 15 September 1967 ; MNHN 1977-778 , 1 (45), Gulf of Aqaba , M.L. Bauchot and J.E. Randall . YEMEN: USNM 327464 , 6 (31–45.5), Zubair Island , Yemen, E. Clark, 30 September 1967 . INDONESIA: Neotype of Gobius anjerensis, BPBM 26651, 31 mm SL male, Bunaken Island, off Manado , Sulawesi, J.E. Randall and party, 28 August 1978 ; NSMT-P.64544, 3(26–27.5), Baguala Bay , Ambon, K. Shibukawa, 1 December 1998 ; NSMT-P.63406, 1(20.5), Lembeh Island , Bitung, Sulawesi, 13 July 2000 ; USNM 211025 , 19 (19–34), bay off Tuhaka , Ihamahu, Pulau Saparua, Moluccas , V.G. Springer and party, 18 January 1973 . GUAM: AMS I.40826-001, 1(44), Tumon Bay , H. Larson, 4 May 1969 ; Syntypes of Gobius deltoides, ANSP 84134, 10(28–43), Agana, A. Seale, June 1900; BPBM 267 , 4 (26–44.5), Agana, A. Seale, 2 June 1900 ; USNM 109399 , 1 (39), A. Seale, 2 June 1900 . REPUBLIC OF MICRONESIA: USNM 224767 , 4 (28–35), N coast Ponape, Caroline Islands , 4 September 1980 . AUSTRALIA: AMS I.28999-010, 2(31.5–36.5), E end of Pulao Kambing, Cocos-Keeling Islands, G . R. Allen , 12 February 1989 ; NTM S.11973-037, 2(32–37.5), lagoon entrance by West Islet , Ashmore Reef, Timor Sea, H. Larson, 26 July 1986 ; NTM S.11983-037, 1(40.5), W end Raine Island , Queensland , AIMS-AMS survey, 13 February 1979 ; AMS I.35852-003, 3(45–60), first lagoon, north channel, One Tree Island , J. Caley, 7 April 1990 ; ex-USNM 308212, 2(54–56), reef flat on SE side One Tree Island , Queensland , V.G. Springer and party, 27 November 1966 . SOLOMON ISLANDS: USNM 358895 , 1 (37), Santa Cruz Island , 22 September 1998 . FIJI: USNM 243060 , 4 (37–49), main harbour on W side Matuku Island , Lau Group, 23 April 1982 . FRENCH POLYNESIA: AMS I.21874-001, 2(35–38), mouth of creek at head of Opunohu Bay , Moorea , R. Galzin , 15 October 1990 ; AMS I.30940-006, 8(16–37), mouth of creek at head of Opunohu Bay , Moorea , R. Galzin , October 1990 ; CAS 51530, 6 (40.5–51), almost-enclosed lagoon, Taravao , Tahiti, J.E. Randall, 21 April 1956 .

Other material; no data taken. SOUTH AFRICA: SAM 25625, 1, Inhaca, B. Kensley and party. MOZAMBIQUE: SAIAB 16943, 2, Ibo Island, Querimba Archipelago; SAIAB 57255, 4, Querimba Archipelago; SAIAB 13674, 1, Querimba Archipelago; SAIAB 16928, 2, Pinda Island; SAIAB 16934, 1, Pinda Island. EGYPT: BPBM 18334, 4, Ras Mohammed, S end Sinai Peninsula; BPBM 19840, 10, beach N of Nuweiba el-Muzeini, Gulf of Aqaba. SUDAN: BPBM 19760, 1, N of Port Sudan, Red Sea; BPBM 19750, 4, Suakin Harbour, Red Sea. SAUDI ARABIA: BPBM 30405, 1, about 5 miles N of Marifa, Persian Gulf. MADAGASCAR: USNM 327542, 3. OMAN: BPBM 36143, 1, Masirah Island; BPBM 34406, 1, Hoone’s Bay, near Mirbat; BPBM 36032, 10, Sawda Island. ALDABRA: SAIAB 16930, 3; SAIAB 16940, 2; USNM 327760, 60, Aldabra Atoll. SEYCHELLES: USNM 308213, 46, Farquhar Atoll. PHILIPPINES: AMS I.21931-005, 6, Lapu Lapu market, Cebu, Mactan Island. INDONESIA: NTM S.15420-009, 1, Pahepa, Sangir Island; AMS I.23679-002, 2, Denpasar, Bali. THAILAND: NSMT-P.61750, 1, Ko Lon, Phuket. MARIANA ISLANDS: NTM S.14712-001, 2, Sanvitores Beach, Guam. MICRONESIA: USNM 224763, 59, Ponape, Caroline Islands. MARSHALL ISLANDS: USNM 368807, 5. VANUATU: AMS IB.3607, 1, Vila, New Hebrides. SOCIETY ISLANDS: MNHN 1984-168, 5, Tiahura, Moorea; CAS 53895, 2, Moorea; CAS 58267, 2, Moorea; CAS 58268, 2, Moorea; CAS 53896, 2, Papeete Harbour, Tahiti.

Description. Based on 66 specimens, 26–84 mm SL. An asterisk indicates the counts of the neotype of Gobius anjerensis .

First dorsal VI*; second dorsal I,10–I,11* (mean I,10.9); anal I,10–12 (mean I,11.0*), pectoral rays 14–17 (mean 16.0*), segmented caudal rays nearly always 17*; caudal ray pattern nearly always 9/8*; branched caudal rays 6/6 to 8/7 (mean 7/6*); lateral scale count 25–29 (mean 26.6, 26 in neotype); TRB 9*–12 (mean 10.4); predorsal scales 7*–13 (mean 9.3); circumpeduncular scales 11–13 (mean 12.1*). Gill rakers on outer face of first arch 1–2 + 3–4 (in 6, modally 1+4).

Body compressed, width at anus 11.9–28.1% (mean 15.2%) of SL. Body rather stocky in appearance, body depth at anus 19.6–26.6% (mean 23.3%) of SL, body depth at first dorsal fin origin 14.3–27.7% (mean 23.3%) of SL. Head compressed, broader ventrally, slightly deeper than wide, HL 26.6–33.3% (mean 29.3%) of SL; head depth at posterior preopercular margin 62.0–83.7% (mean 71.2%) of HL; head width at posterior preopercular margin 53.7–82.2% (mean 68.5%) of HL; head profile bluntly pointed; nape slightly convex behind eyes. Mouth subterminal to nearly terminal, slightly oblique; jaws generally reaching to below anterior margin of eye; upper jaw length 32.0–41.5% (mean 35.9%) of HL. Upper lip smooth, narrower than lower, lower lip papillose close to teeth, with twist or fold posteriorly, forming triangular flap, lip interrupted at chin. Eye moderate to relatively small in large specimens, dorsolateral, 22.7–30.5% (mean 27.1%) of HL; preorbital width 18.2–27.8% (mean 22.3%) of HL. Snout bluntly pointed, 22.4–44.4% (mean 34.6%) of HL; posterior naris round to almost triangular, close to anterior margin of eye; anterior naris in short tube, higher on posterior margin of eye, about level with middle of eye or somewhat ventral to it. Interorbital narrow, 4.7–11.3% (mean 7.9%) of HL. Caudal peduncle compressed, length 12.9–18.6% (mean 16.1%) of SL; caudal peduncle depth 10.7–13.8% (mean 11.8%) of SL.

First dorsal fin rounded to square, with no spines greatly elongate; third or fourth spine usually longest; when adpressed, spine tips reaching to first to third element of second dorsal fin. Third dorsal spine modally longest, 15.4–19.6% (mean 17.6%) of SL; fourth dorsal spine length 15.0–19.8% (mean 17.7%) of SL. Second dorsal fin as tall as first dorsal fin, rays longer anteriorly than posteriorly, fin pointed to slightly rounded posteriorly. Anal fin lower than second dorsal fin, anteriormost rays shorter than posterior few rays; fin pointed to slightly rounded posteriorly. Second dorsal and anal fin rays, when adpressed, usually reaching caudal fin in adults. Pectoral fin somewhat pointed, central rays longest, 23.5–28.7% (mean 26.0%) of SL; fin reaching back to above first few anal fin rays. Pelvic fins fused, frenum with distinctive finely fimbriate margin, fins round to somewhat oval, reaching

Gill opening restricted, extending anteriorly to lower edge of pectoral base or to just under opercle. One or two slender gill rakers on outer face of first arch, closest to angle of arch, remainder of rakers very short, pointed; pointed papillae present along inner face of first arch; outer rakers on second gill arch consisting of two series of papillae, one low and one of slightly pointed papillae; outer rakers on remaining arches low, broad-based. Inner face of upper limb of first gill arch, and to lesser extent, upper limbs of other arches, covered with low dense fleshy papillae which may form clumps or groups; dorsal portion of arch may have short fleshy protuberances ending in one or several papillae. About one-third of first gill arch bound by membrane to inner face of opercle. Tongue short, tip bilobed to concave.

Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest, curved and pointed, largest teeth at front of jaw on either side of symphysis; innermost row teeth very small, sharp and evenly sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw but outer row teeth may be smaller (in females); posteriormost one to three outer row teeth enlarged and somewhat recurved in males.

Ctenoid predorsal scales on nape of variable extent, from just over opercle to close up behind eyes; nape midline scales may be partly cycloid from behind eyes to rear margin of preopercle, nape midline never entirely with cycloid scales ( Fig. 4 ). Opercle covered with scales, cycloid or ctenoid or mix of both. Preopercular scales usually cycloid, upper or rear part of preopercle may include ctenoid scales; cycloid scales may extend anterior to vertical dark cheek bar below eye. Breast with cycloid scales reaching up to below mid-opercle or as far forward as rear edge of preopercle. Pectoral fin base covered with cycloid scales, some may be ctenoid. Belly scales along midline usually cycloid; ctenoid scales present posteriorly on midline in many specimens.

Africa and E-F, G. thompsoni from Tobago and Brazil respectively. Hatching indicates ctenoid scales.

Head pores with anterior nasal pore just anterior to anterior naris, posterior nasal pore beside each posterior naris, a pair of anterior interorbital pores, a single posterior interorbital pore, a postorbital pore, a terminal pore over opercle and an anterior and posterior temporal pore in short separate posterior portion of the oculoscapular canal over opercle; three preopercular pores present ( Fig. 1 ).

Sensory papillae arranged in transverse pattern ( Fig. 1 ); rows on opercle may be partly concealed by scales; papillae on chin in two short longitudinal rows on either side of midline (see Larson & Buckle 2005: fig. 2B).

Coloration of fresh material. Fresh-dead specimens are illustrated in Randall and Greenfield (2001: pl. IA–F), while good illustrations of living fish are in Senou et al. (2004: 244–245), Kuiter and Tonozuka (2001: 672, as G. deltoides and G. sp.1) and Myers (1991: plate 124A, 1999: plate 161D). There is a little variation in colour pattern ( Figs 5–6 ).

Living fish show characteristic small bright blue-white to pearly white spots and blotches on the side of the head (may form oblique to horizontal rough lines on cheek and opercle), pectoral fin base and scattered along the middle of the side of the body, mostly immediately behind the pectoral fin or less often, extending down the anterior half of the body ( Fig. 5 ). A characteristic pale to deep gold or orange spot is present above the pectoral fin base, that may be surrounded by a grey to purplish grey to brownish blotch or a short brownish horizontal line, with one or several short blue-white to pearly white short horizontal lines beneath ( Fig. 6 ). Body colour pattern similar to preserved specimens but blotches and short lines on body are purplish or reddish brown to orange brown in colour, with the large lateral blotches or bars being browner than the overlying short lines and spots. The first dorsal fin always has three to four wavy blackish to reddish-brown lines beginning on the first spine; with a

Coloration of preserved material. The illustrations in Randall and Greenfield (2001) show some of the variations in colour pattern. The basic pattern is as follows.

Head and body yellowish white to pale brown (depending upon preservation) with five (sometimes indistinct) brown saddles crossing dorsum, posterior part of each saddle merging into one of six vertical broad diffuse brown bars or elongate to rounded blotches along mid-side of body, posteriormost bar (on caudal peduncle just before mid-base of caudal fin) often indiscernible. Two to four irregular rows of small dark brown spots along dorsal quarter of body, mid-side of body may be marked by series of short brown horizontal lines and spots, which may be enlarged and merging with underlying vertical diffuse brown bars. Specimens may be heavily spotted, with most scales on upper two-thirds of body having a brown spot.

Predorsal, snout and side of head finely speckled and spotted with brown, opercle with brown line extending from centre of preopercular margin obliquely back to rear opercular corner. Blackish to dark brown cheek-bar from ventral edge of eye running vertically to slightly obliquely down to end on lower preopercular edge, but not extending onto branchiostegal membranes. Blackish to dark brown line from (usually) rear half of iris running over top of eye, rarely meeting its counterpart in interorbital space. Upper lip crossed by dusky bar (at level with nares) and similar dusky bar crossing centre of lip; bars on lip may be marbled, vermiculate or broken into series of irregular spots and short streaks. Cheek with diffuse brownish horizontal bar crossing vertical cheek-bar; brownish bar variable in shape and intensity, its pigment often surrounding the round pale spots that were white when fish alive. Above pectoral fin base, brownish to dark brown blotch or short horizontal bar (may be diffuse to nearly indiscernible) surrounding small pale round spot (not always discernible in preserved material); brown blotch or bar may be partly connected to brown line or series of brown spots running from behind eye along top of preopercle. Pectoral fin base with horizontal brown line across dorsal half, line extending onto lower part of fin and ending in diffuse brown blotch.

First dorsal fin transparent, with three to four wavy brown lines (may break up into series of spots)

beginning at first spine; beginning of first three wavy lines marked by characteristic darker brown to blackish spot on front of spine. Second dorsal fin similar but spots on face of fin spine much less distinct; wavy lines may transform into many series of spots or coalesce into plain brown pigment, which may form vertical streaks on membrane, especially on posterior half of fin. Anal fin plain dusky to brownish, may be darker distally. Caudal fin transparent to translucent brownish with pale brown streaks or rows of fine brown spots along fin membranes, especially on dorsal half of fin. Pectoral fins transparent to translucent dusky, distinctively finely speckled with darker brown, in irregular rows. Pelvic fins and frenum plain dusky to brownish, darker brown toward midline of fused fins.

Distribution. Gnatholepis anjerensis is known from South Africa, Mozambique, Zanzibar, Kenya, Madagascar, Aldabra, Seychelles, Rodrigues, Mauritius, Maldives, Chagos, Egypt, Israel, Oman, Yemen, Thailand, Malaysia, Indonesia, Philippines, Japan, Guam, Micronesia, Marshall Islands, Vanuatu, Australia (Cocos-Keeling Islands to One Tree Island), Solomon Islands, New Caledonia, Tonga, Fiji, Wallis Island and the Society Islands.

Ecology. Very little is reported of the natural history of this species other than Randall and Greenfield (2001), who provided information on preferred habitat (though this data is mixed with that for other species). Gnatholepis anjerensis is usually found in lagoonal situations over sand near coral or coral rock, rubble, Halimeda or seagrasses at a depth range of 0.25– 12 m. Coene and Ollevier (2000) found G. anjerensis in Kenya mostly at sites with seagrass, feeding on a general range of benthic invertebrates.

Cole (1990), in her discussion of hermaphroditism in some coral reef gobies, confirmed that Gnatholepis anjerensis was a gonochore and did not change sex (a feature of the subfamily Gobionellinae ). Her collection of 69 females and 54 males showed a sex ratio of 1.3 females per male.

Thacker’s (2004a) work on the population structure of South Pacific G. anjerensis and G. cauerensis (which she refers to as G. scapulostigma ) showed that the populations of G. anjerensis from Fiji to the Tuamotus were all much younger (about 88,000 years) than the ages of the island groups in which they were found. Population

Comparisons. When alive, this species is most easily distinguished from its congeners by having the first dorsal fin rather square to rounded in shape (the fifth D1 spine is nearly as long as the third to fourth spines), in having the first spine banded by three to four black spots or short streaks; the pectoral fin is covered with fine white and dark speckles, a characteristic small round gold spot is above the pectoral fin base (often with a short dark line or elongate blotch under it) and the nape and body is covered with irregular rows of small blackish to reddish spots, with five to six diffuse dusky blotches along the lower mid-side of the body. When preserved, G. anjerensis can be distinguished by its ctenoid predorsal scales, slightly stocky body and square first dorsal fin shape; most of the colour pattern remains visible but for the small round gold spot (which may be hard to discern) above the pectoral fin base.

Remarks. Gobius anjerensis was based on a Kuhl and van Hasselt figure of a 48 mm specimen, and is the type species of the genus Gnatholepis . A neotype (BPBM 26651) was designated by Randall and Greenfield (2001: 3, fig. 2). Randall and Greenfield (2001) picked a fairly small (31 mm SL; the species reaches 84 mm SL) and slightly fragile specimen as neotype of Gobius anjerensis (from northern Sulawesi, some distance from the type locality of Anjer in Java, but forgivable in that Anjer was destroyed in the 1883 eruption of Krakatau). Some of Randall and Greenfield’s counts do not agree with ours. For example the lateral scale count for the neotype is given as 30, but we counted 26 scale rows on this specimen and their Figure 2 of the neotype also shows 26 scales. Although Randall and Greenfield (2001) say that they counted lateral scales “from the upper edge of the gill opening”, they also say that scales are crowded behind the gill opening and that they counted all the small scales here. It appears that they were counting lateral rows beginning right behind the gill opening and thus their counts would be several rows greater than ours.

Gobius capistratus Peters, 1855 , was described from Ibo, Mozambique, with two syntypes (ZMB 2103). Randall and Greenfield (2001) synonymised this with G. anjerensis but gave no reason for doing so and apparently did not examine type material. Peters’ specimens were described and figured in Sauvage (1891: 356, pl. 38, fig. 5) and a partial translation of Peters’ description was provided in Günther (1861: 36). Klunzinger (1871) considered that G. ophthalmotaenia was a synonym of G. capistratus . Jeff Leis photographed the syntypes at ZMB and the fine light and dark speckling on the pectoral fins is visible, as are the blue-white spots on the lower part of the body and side of the head (in both specimens) ( Fig. 7 ).

Gobius deltoides Seale, 1901 , was described from 18 syntypes from Guam (ANSP 84134, BPBM 267, USNM 109399). ANSP 84134 (formerly BPBM 267 ) includes a 14 mm characoid.

Some specimens identified by Randall and Greenfield as G. anjerensis (AMS I.20757-072, Raine Island; AMS I.27156-035, Middleton Reef; AMS IB.4004, Heron Island), were subsequently identified by us as G. cauerensis (n.b. one specimen in AMS I.20757-072 was G. anjerensis , the other 10 were G. cauerensis ). These may be some of the anomalous specimens that they mentioned as having distinct longitudinal lines on the body, a colour pattern not usually found in G. anjerensis ( Randall & Greenfield 2001: 4–5) . Their definition of the species G. anjerensis should be taken from the description of the neotype only, as their view of this species is confused.

Thacker’s (2004b) definitions of G. anjerensis and G. knighti do not agree with ours. From her discussion ( Thacker 2004b: 578–580, fig. 3), it seems that she based her decision as to what name with which to label each taxon, on whether or not the fish had a dark blotch or spot, with or without a pale centre, above the pectoral fin base (these she called G. scapulostigma ), a dark line or dash above the pectoral fin base (these she called G. knighti ), or had no distinct spot at all above the pectoral fin base (these she called G. anjerensis ). Specimens identified as G. anjerensis from AMS by Thacker, were re-examined by AMS fish section staff on our behalf and were found to include Gnatholepis gymnocara (specimens from Yeppoon, Sabina Point and Townshend Island, all in Queensland). Two lots from the Northern Territory (East Arm; Gunn Point) were found to be G. argus , not G. anjerensis , as pointed out by Larson and Buckle (2005).

Kuiter and Tonozuka (2001: 671, figs C, D) show specimens from Gilimanuk and Lipah, Bali, Indonesia, which appear to be oddly-marked specimens of G. anjerensis . They show the black spots along the leading edge of the first dorsal spine but the fin is patterned with rows of oval spots and the body is very darkly mottled. Otherwise they closely resemble G. yoshinoi .

Hayashi and Shiratori’s Figure 176 is of G. anjerensis (2003: 96) , while Figure 175 is of G. yoshinoi (see

Figures

FIGURE 1. Sensory pores and papillae in Gnatholepis anjerensis, SAIAB 2112, Mauritius, 46 mm SL male. Scales omitted.

FIGURE 2. Canonical analysis of principal co-ordinates plot for the seven Gnatholepis species with vectors overlaid. CAP axes 1 and 2 explain 62% of the variation from 13 canonical axes.

FIGURE 4. Diagram showing variation in ctenoid and cycloid scale arrangement on the predorsal (nape) region in specimens of A-B, Gnatholepis anjerensis from Mauritius and Ambon respectively, C-D, G. cauerensis, both from Aliwal Shoal, South

FIGURE 5. Gnatholepis anjerensis, 2 m depth, Shams Alam, south of Marsa Alam, Egypt, Red Sea. Photo by Sergey Bogorodsky.

FIGURE 6. Gnatholepis anjerensis, Kosi River mouth, Kwa Zulu Natal, South Africa. Photo by Dennis King.

FIGURE 7. Gnatholepis anjerensis, syntypes of Gobius capistratus, ZMB 2103. Photo by Jeff Leis.

FIGURE 11. Gnatholepis caudimaculata n. sp., Sharm, Sinai, Egypt. Photo by Ole Brett.