The Beagle, Records o f the Museums and Art Galleries of the Northern Territory, 1999 15: 23-42

A REVIEW OF THE MANGROVE GOBY HEMIGOBIUS (GOBIOIDEI, , )

HELEN K. LARSON Museum and Art Gallery of the Northern Territory PO Box 4646, Darwin N T 0801, AUSTRALIA ' (e-mail: [email protected])

ABSTRACT The gobiid genus Hemigobius is reviewed and found to comprise two , which are redescribed. The genus is considered to belong to the subfamily Gobionellinae, is closely related to the genera and , and can be distinguished from them by a combination of characters. Hemigobius is restricted to mangrove habitats in northern Australia and South-east Asia.

KEYWORDS: Gobioidei, Gobiidae, Gobionellinae, Hemigobius, mangroves, northern Australia, South-east Asia.

INTRODUCTION were “... too badly preserved to decide it with certainty” (1931: 102). The holotype of The gobiid fish genus Hemigobius Gobius hoevenii (RMNH 4457) is a valid Bleeker, 1874, consists of six nominal species of Hemigobius, redescribed below. species, which have been variously placed in Three probable syntypes of Gobius Hemigobius, Mugilogobius or Pseudogobius. tambujon (RMNH 4458) are The generic name apparently has not been species (Larson, in prep.). used other than in Koumans (1931, 1953), Fowler (1940) erected the genus Miller (1987), Kottelat et al. (1993) and Sphenentogobius, for his new species Larson (1995; in press). vanderbilti. He remarked upon the elevated Bleeker (1874) placed his new genus rear ramus of the mandible, and the Hemigobius in the group Subphalanx etymology of his genus Sphenentogobius Eugobii, in which he put most of the gobiid reflects this: “o<|>r)v wedge + evto^ within + genera comprising his Phalanx Gobiini. He Gobius”. He considered Sphenentogobius to characterised Hemigobius by the truncate be “apparently related to Redigobius". teeth of its type species, Gobius melanurus During revision of the gobiid fish genus Bleeker. Mugilogobius Smiti and its nominal relatives Koumans (1931: 101) erected the genus in the subfamily Gobionellinae (Larson Microgobius (non Poey), based on Bleeker’s 1995; Larson in press), Hemigobius Bleeker “museum name” written on the bottles of was found to be most closely related to two species at Leiden museum (RMNH), Pseudogobius. and listed it as a junior synonym of Hemigobius and Pseudogobius group Bleeker (along with together due to their derived fifth Pseudogobius, given as another Bleeker ceratobranchial, lip and gut morphology, museum name). Koumans included under mouth position and abbreviate headpore Microgobius non Poey: Gobius hoevenii patterns (they both lack preopercular pores Bleeker, 1851, and Gobius tambujon and the rear part of oculoscapular canal, but Bleeker, 1854, based on Bleeker’s jar labels. headpores are always present) (Larson in Koumans indicated that the latter two prep.). Each genus has an autapomorphic species were probably Stigmatogobius, but long gut coiling pattern, differing from most

23 Helen K. Larson

of the Mugilogobius group of genera. on this group of gobionellines by Aurich Hemigobius has a long coiled gut (1938) and Miller (1987, 1989). reminiscent of (Larson Pterygiophore formula follows Birdsong et 1995; in press). The gut morphology of al. (1988). Transverse scale counts are taken Pseudogobius resembles a short version of by counting the number of scale rows from that in (Geevarghese 1983; Larson the anal fin origin diagonally upward and in press). Other related genera have short, back toward the second dorsal fin base. Head typically “carnivorous goby” gut formé. length is taken to the upper attachment of the Hemigobius has 17 segmented caudal rays, opercular membrane. Interorbital width is while Pseudogobius has 16. Pseudogobius least fleshy width (not least bony width). In has two s papillae rows on the snout, and the descriptions, an asterisk indicates counts Hemigobius has three, as in Mugilogobius. of the holotype. Numbers in parentheses Hemigobius has the fine villi on the head after counts indicate the number of characteristic of Mugilogobius, while specimens with that count, or the range of Pseudogobius does not possess them. counts. Vertebral counts and other Miller (1987) was in error when he stated osteological information was obtained by that Hemigobius lacked head canals. The radiography and clearing and double- author assumes that Miller made this staining. statement on the basis of his examination of Synonymies are not complete, as it was the type of Gobius melanurus Bleeker, 1849 not always possible to determine from (which is also the type of Hemigobius descriptions or illustrations what species was bleekeri Koumans, 1953), the only material referred to in various publications (for of this genus referred to in his paper (Miller example, Stigmatogobius hoevenii could be 1987). This specimen (RMNH 4501) has Hemigobius hoevenii or Mugilogobius badly abraded skin covering the very short chulae). Synonymies are given where the canals, so that the headpores appear to be identity was verified by examination of absent. However, the cheek scales, specimens or the description was characteristically shaped dentary and unequivocal. flattened teeth are present. Examination of Abbreviations for institutions referred to fresh specimens of this species revealed the are: AMS - The Australian Museum, canal and headpore arrangement Sydney; ANSP - Academy of Natural characteristic of this genus. Miller Sciences, Philadelphia; BMNH - The apparently was not aware of the identity of Natural History Museum, London; CAS - Gobius hoevenii Bleeker, 1851, as he refers California Academy of Sciences, San to Hemigobius as being monotypic (based on Francisco; CMK - Collection Maurice Koumans 1953). In the same paper, Miller Kottelat, Comol, Switzerland; KUMF - erroneously included Sphenentogobius Kasetsart University Museum of Fisheries, Fowler as a synonym of Redigobius (no Bangkok; NIFI - National Inland Fisheries reasons are given, although it is likely that he Institute, Bangkok; RMNH - Nationaal did so in agreement with Fowler’s statement, Naturhistorisches Museum, Leiden; NTM - in the original description of the genus, that Museum and Art Gallery of the Northern it was related to Redigobius). Territory, Darwin; URM - University of the Ryukyus, Naha; USNM - National Museum of Natural History, Washington; WAM - METHODS Western Australian Museum, Perth; ZMH - Measurements were taken using Zoologische Museum, Hamburg; ZRC - electronic callipers and dissecting Zoological Reference Collection, University microscope. Counts and methods generally of Singapore; ZSM - Zoologische Staat­ follow Hubbs and Lagler (1970), except as sammlung, München. indicated below. Papillae pattern Other abbreviations used: HL - head terminology is based on that of Sanzo length; SL - standard length; TRB - (1911), due to its use in previous literature transverse scale rows backward.

24 Review of the mangrove goby Hemigobius

SYSTEMATICS extends upward and forward, well above quadrate. Mandibular ramus elevated and Hemigobius Bleeker, 1874 curved anteriorly in H. mingi, elevated but angled backward in H. hoevenii. Fifth Hemigobius Bleeker, 1874 (Gobius ceratobranchia^ triangular, very open and melanurus Bleeker, 1849: 31, Java; = lattice-like in structure. Hemigobius bleekeri Koumans, 1953: 191, Found in mangroves in Indo-west replacement name for Gobius melanurus Pacific. Bleeker, by original designation and monotypy). Key to species ofHemigobius Microgobius Koumans, 1931 (listed as synonym of Stigmatogobius). Preoccupied 1. Pectoral rays 13-17, usually 15-16; by Microgobius Poey. body relatively slender, depth at anus about Sphenentogobius Fowler, 1940 (S. 4-5.5 times in SL; about six distinct diagonal vanderbilti Fowler, 1940: 396, figs 8-11, bars along sides; first dorsal fin with two Sumatra, by original designation and broad dark bands, not forming monotypy). spot...... H. hoevenii (Bleeker, 1851) Diagnosis. Distinguished by following (Thailand, Hong Kong, Malaysia, combination of characters. Second dorsal Singapore, Philippines, Borneo, New rays 1,6-8, modally 1,7; anal rays 1,6-9, Guinea, northern Australia) modally 1,7; pectoral rays 13-17; 17 1A. Pectoral rays 13-15, usually 14; segmented caudal rays in 9/8 pattern; some body relatively deep, depth at anus about headpores present with no lateral canal over 3.5-4.5 in SL; scale margins outlined, preoperculum, no preopercular pores, no forming reticulated pattern, with four or five nasal pores, and pairs of interorbital pores indistinct dorsal saddles sometimes present; not connecting across interorbital space; 25- very distinct black spot posteriorly on first 34 lateral scales; circumpeduncular scales dorsal...... H. mingi (Herre, 1936) strongly modally 12; predorsal scales 7-12, (Thailand, Singapore, Sumatra, Java) extending close up to behind eyes, margins of anteriormost scales scalloped; preopercle at least partly scaled; interorbital and top of Hemigobius hoevenii (Bleeker, 1851) snout with fine villi; gili rakers without (Figs 1-6, Tables 1-4) spines; papillose flaps or pads present at insertion of first gili arch onto roof of mouth; Gobius hoevenii Bleeker, 1851: 426-427 jaws small, lower jaw symphysis usually (Sambas, in river, Borneo). raised; thin, folded lower lip; teeth small and Vaimosa crassa Herre, 1945: 403 (brook flattened in females (at least), usually near Un Long, Hong Kong). conical in males; anterior nostril in short Stigmatogobius hoevenii - Koumans tube oriented down and forward over upper 1953: 125 (in part). lip, preorbital usually curved outward Microgobius hoevenii - Bleeker 1983: pi. slightly around base of nostril; genital 438, fig. 17. papilla slender, flattened and pointed in Mugilogobius obliquifasciata Wu and Ni, males, conical and blunt-tipped in females; 1985: 93-95 (Haikou, Hainan Island, China). intestine very long and tightly coiled. - Anon. 1986: 272-273. Dorsal pterygiophore formula 3-12210; Mugilogobius obliquifasciatus - Zhu two epurals, rarely one; one to three anal 1988: fig. 162. pterygiophores before haemal spine of first Hemigobius crassa - Davis 1988: 164. caudal vertebra; neural spine on first Pseudogobius hoevenii - Murphy 1990: vertebra usually short and broad. Palatine 155. and pterygoid short, with broad, T-shaped Mugilogobius latifrons - Nguyen 1991: heads; palatine larger and more robust than 334-335, fig. 143. pterygoid. Metapterygoid deep, well Hemigobius hoevenii - Kottelat et al. separated from quadrate, anterior process 1993: 146.

25 Helen K. Larson

Material examined. 80 specimens (8.5- 91, 8, Sungei Seletar; ZRC 21872-906, 35, 37). THAILAND: URM P. 12662, 2(28-28), Mandai Kecil; ZRC 20476-78, 3, Mandai mangrove swamp at Phuket, H. Senou and V. mangroves; NTM S.13957-009, 35, Sungei Chavalit, 2 November 1983; ex URM Pandan; CMK 8223, 6, Kranji mangrove; P.6677, 3(22.5-25.5), Ranong, 9 March CAS 40136, 8, Serangoon; ZRC 29185, 1, 1982. MALAYSIA: ZSM 27559, 4(23- Sungei Buloh; NTM S. 13968-008, 13, 26.5), mangrove at Bamgangan, SW of Sungei Pandan; NTM S.13959-012, 10, Sandakan, Sabah, Kettner, Krumenacher and Sungei Buloh; NTM S. 13961-007, 27, Witte, 13 March 1988. SINGAPORE: Mandai Kecil. BRUNEI: NTM S. 13052- Paratype of Vaimosa fusca, CAS 32987,25.5 002, 1, Pulau Berambang. NTM S. 13053- mm SL male, mangrove swamp, Kranji 004, 2, Pulau Berambang. PHILIPPINES: River, A. Herre, March 1937. NTM USNM 99613, 1, Port Dupon; CAS 38637, S. 14235-004, 11(8.5-27), Sungei Buloh 46, Panay, Capiz; CAS 26381,1, Lake Buhi. mangroves, K. Lim, 30 January 1991. INDONESIA: CMK 7265, 6, Tanjung BRUNEI: NTM S.12812-002, 2(18-24), Mayong, Padang, Sumatra. PAPUA NEW Kedalayan River, from Nypa leaf axils, R. GUINEA: USNM 316170, 1, Dani. Hanley and S. Choy, 7 April 1989. AUSTRALIA, NORTHERN TERRITORY: INDONESIA: Holotype of Gobius hoevenii, NTM S. 10419-006, 2, Elizabeth River, RMNH 4457, 32 mm SL female, in river, Darwin Harbour; NTM S. 10420-002, 1, Sambas, Borneo. BMNH 1935.5.27.28, Elizabeth River, Darwin Harbour; NTM 1(36), (possibly from Sulawesi), Arnold. 5. 11845-004, 1, Leanyer Swamp; NTM HONG KONG: Holotype of Vaimosa S. 14289-001, 12, Leader’s Creek, Gunn crassa, 35 mm SL female, CAS/SU 39848, Point; NTM S. 14290-002, 20, Leader’s Un Long, New Territories, A.W. Herre, 23 Creek, Gunn Point. NO DATA: RMNH February 1941. PHILIPPINES: CAS 38636, 14049,1. 30(19-33), Coron, Busuanga, 22-30 June Diagnosis. Hemigobius with body 1940, A.W. Herre. PAPUA NEW GUINEA: robust, compressed, with rounded, WAM P.26751-006, 2(21.5-37), Tureture somewhat depressed head, snout rounded village, Binaturi, G. Allen, 29 September and overhanging upper lip; second dorsal 1979. AUSTRALIA: QUEENSLAND: rays 1,6-8 (modally 1,7); anal rays 1,6-7 AMS 1.23262-001,8(18-32), The Esplanade, (modally 1,7); pectoral rays 13-16; Cairns, D. Hoese and D. Rennis, 2 October longitudinal scales 27-32; TRB 7-12; 1982. AUSTRALIA, NORTHERN anteriormost predorsal scale largest, 8-10 TERRITORY: NTM S.11065-002, 13(13- scales, extending close up behind eyes; 21.5), Leanyer Swamp, Darwin, T. Davis, 5 mouth enlarged in males; scales on body March 1980. WESTERN AUSTRALIA: ctenoid; first dorsal fin low, rounded, with AMS 1.25521-009, 1(27.5), Crab Creek, no filamentous spines; body greyish to Broome, D. Hoese, D. Rennis, 20 September brownish with six diagonal blackish bars 1985. across sides, marbled to ocellate black spot Other material examined (but not used in on caudal base, roughly vertical black line description). 275 specimens from the on opercle extending to underneath head; following localities. HONG KONG: known from mangrove areas of the Indo- Paratype of Vaimosa crassa, CAS/SU Malayan Archipelago, Papua New Guinea 39849, 1, 35 mm SL. THAILAND: URM and northern Australia. P.13336, 1, Khung Kraben Bay; NTM Description. Based on 40 specimens, 15- S.13953-014, 6, Klong Bang Sai, Phuket; 37 mm SL. Counts of holotype of Gobius URM P. 13344, 2, Khung Kraben Bay; NTM hoevenii indicated by asterisk. S. 14288-002, 6, Ta-Chalab. MALAYSIA: First dorsal VI*; second dorsal 1,7-8 CAS 33168, 3, Kabili River, North Borneo. (mean 1,7*); anal 1,7-9 (mean 1,7*), pectoral SINGAPORE: ZRC 20635-40, 6, Sungei rays 13-16* (mean 16), segmented caudal Punggol; ZRC 20238-45, 8, Sungei Punggol; rays always 17*; caudal ray pattern 6/6 to ZRC 27450, 1, Siglap Canal; ZRC 21084- 9/8 (modally 8/7*); branched caudal rays 12-

26 Review of the mangrove goby Hemigobius

Table 1. Frequency distribution of fin ray counts in HL 3.2-4.1 (mean 3.6) in SL. Depth at Hemigobius species. posterior preopercular margin 1.5-1.9 (mean 1.7) in HL. Width at posterior preopercular Second margin 1.2-1.5 (mean 1.3) in HL. Mouth dorsal rays Anal rays Pectoral raysterminal to subterminal, almost horizontal, Species 6 7 8 6 7 8 9 13 14 15 16 17with rounded snout overhanging upper lip; jaws forming angle of about 7-15° with body hoevenii -25 15 - 27 12 1 1 2 17 19 1 axis; jaws generally reaching at least to mingi 1 34 1 1 35 - - 2 29 5 • - below anterior half of eye, and to rear edge of eye in large (sexually mature) males. Upper jaw 1.7-3.6 (mean 2.8) in HL; in Table 2. Frequency distribution of longitudinal counts in males, 1.7-3.5 (mean 2.6); in females, 2.1- Hemigobius species. 3.6 (mean 2.9) in SL. Upper lip narrow, smooth, without fleshy fimbriae; lower lip Species 25 26 27 28 29 30 31 32 33 thin, 34 reduced to narrow (mugilid-like) fold, free along posterior half of jaw, anterior half hoevenii 2 3 10 426561 1 of lip fused to underside of head, lower lip mingi - - 4 4 17 9 1 1 - fold often thicker and extending further forward in mature males; lower jaw slightly curved upward, in large specimens low ridge Table 3. Frequency distribution of transverse backward at symphysis sometimes present. Eyes large, scale counts in Hemigobius species. dorsolateral, high on head, sometimes forming part of dorsal profile, 3.1-4.2 (mean Species 7 8 9 10 11 12 3.5) in HL. Snout rounded, inflated over top lip, 2.8-4.1 (mean 3.3) in HL. Interorbital hoevenii 1 11 10 10 7 1 broad, flat, 2.3-5.3 (mean 3.0) in HL. Top of mingi - - 11 19 5 1 head, from just behind eyes up to snout tip, often with fine villi, these sometimes 17 (modally 15, broken in holotype); relatively sparse on anteriormost nape scales unsegmented (procurrent) caudal rays 6/7 to (villi may only be visible in specimens with 8/8 (modally 7/7); longitudinal scale count well-preserved mucous coat). Body often 25-32 (mean 29; 27 in holotype); TRB 7*-12 quite round in abdominal region in females, (mean 10); predorsal scale count 8-10 (mean depth at anal origin 4.2-5.6 (mean 4.9) in SL. 9*); circumpeduncular scales 12 (13 in one). Caudal peduncle compressed, length 3.3-4.1 Gili rakers on outer face of first arch 2+8 to (mean 3.7) in SL. Caudal peduncle depth 4+9 (modally 2+8). Dorsal pterygiophore 5.5-7.9 (mean 6.9) in SL. formula 3-12210 (in 12). Vertebrae 10+16 First dorsal fin low, rounded, tips of (in nine), 10+17 (in one), 11+15 (in one), second to fourth spines free, second or third 11+16 (in one). Neural spine of first vertebra spines longest or subequal; spines always very short and broad, sharply bent falling short of second dorsal fin origin when posteriorly at halfway point (in 11). Two depressed. First dorsal spine always shorter epurals (in 10) or one very broad epural (in than next three. Third dorsal spine length two). Two (in two) or three (in 10) anal 5.1-13.2 (mean 7.6) in SL. Fourth dorsal pterygiophores before haemal spine of first spine length 6.3-10.0 (mean 7.8) in SL. caudal vertebra. Lachrymal not enlarged, Second dorsal and anal fins low, relatively slender. posteriormost rays usually longer than Body approximately rounded anteriorly; anteriormost, rays reaching more than compressed posteriorly. Head depressed, halfway to caudal fin base when depressed. especially in mature males, width always Pectoral fin short and rounded, central rays greater than depth, cheeks may be inflated in longest, 4.4-5.5 (mean 4.9) in SL; rays males; profile blunt to rounded; nape often usually all branched but for uppermost. convex behind flattened broad interorbital, Pelvic fins short, rounded, reaching half (or

27 Helen K. Larson less) distance to anus, 4.9-6.8 (mean 5.7) in males and females. In males, teeth in outer SL. Caudal fin round, 3.5-4.9 (mean 3.9) in row of upper jaw enlarged, conical and SL. curved (Fig. 1), row present across front of No mental fraenum, chin smooth. jaw only; inner row teeth small, conical, Anterior nostril in short tube, placed just stout, blunt or pointed, extending whole behind upper lip, tube oriented forward and length of jaw, inner teeth more widely down; preorbital straight, not curved to spaced along sides of jaw than those accommodate nostril. Posterior nostril oval, crowded together across front. Lower jaw with or without low rim, placed about teeth in males stout, conical, slightly curved halfway between anterior margin of eye and and pointed; teeth in both rows about equal edge of preorbital (usually slightly closer to in size but anteriormost two to four teeth in eye). Gili opening narrow, usually extending inner row usually enlarged and pointing from just beyond lower pectoral base backward; usually no teeth present on rear forward to just under opercle. Inner edge of third to half of jaw. In females, both rows of shoulder girdle smooth with no ridge or upper jaw teeth very small, flattened, with flange (in 16) or with low bony ridge or blunt tips, teeth with tips bent to one side flange formed by partly exposed cleithrum along side of jaw; both rows of teeth (in 10), which may be bent laterally. Gili extending along entire length of jaw. Lower rakers on outer face of first arch very short, jaw teeth in females very small, curved and pointed and without spines, longest raker at pointed; teeth slightly larger near median angle of arch; several thin lobes or flaps on symphysis, teeth at side of jaw slightly fleshy pads on roof of mouth just above first increasing in size posteriorly, with arch; rakers on inner face of first arch long posteriormost tooth considerably larger, and slender; outer rakers on second arch stout and strongly curved. similar to, but smaller than, those on first Predorsal sedes medium, largest scales arch; outer rakers on third and fourth arches anteriormost, scales reaching forward to very tiny or absent; inner rakers on second close behind eyes; edge of first few and third arches progressively longer and predorsal scales often crenulate or scalloped finer than first arch inner rakers; inner rakers (most conspicuous in larger specimens). of fourth arch small, fine and closely spaced. Operculum covered with cycloid scales. Tongue tip usually rounded, somewhat Cheek below eye naked, often one to three bilobed in northern Australian and Papua scales on upper preopercle close behind eye. New Guinea specimens. Teeth in two rows Pectoral base covered with cycloid scales. in each jaw; tooth form differing between Prepelvic area covered with small cycloid

Table 4. Measurements (mm) of Hemigobius hoevenii (Bleeker, 1851).

Character Holotype Males Males Males Mean Females Females Females Mean Minimum Maximum n = 18 Minimum Maximum n = 22

Head Length 7.9 4.1 10.0 7.3 4.2 9.5 6.5 Head Depth 5.3 2.3 6.3 4.4 2.3 5.4 3.8 Head Width 6.3 3.1 8.0 5.7 3.1 7.2 5.0 Body Depth 7.0 2.7 8.6 5.4 3.1 7.4 4.8 Body Width - 1.5 5.3 3.2 1.8 5.7 2.9 Caud. Ped. Length 8.6 4.5 10.6 7.1 3.8 9.3 6.3 Caud. Ped. Depth 5.4 1.9 6.5 3.9 2.0 5.4 3.4 Snout 1.1 3.3 2.3 1.2 3.1 1.9 Eye 2.5 1.2 3.0 2.1 1.3 2.9 1.9 Jaw - 1.2 5.5 3.2 1.2 3.7 2.3 Interorbit 1.5 1.2 4.1 2.6 1.2 3.7 2.3 Pectoral 2.8 7.6 5.3 3.0 6.7 4.7 Pelvic 4.7 2.5 6.9 4.6 2.6 5.7 4.1 Caudal 3.8 10.2 6.7 4.1 7.7 5.8 Longest D1 spine - 1.7 5.1 4.1 1.8 4.4 3.2

28 Review of the mangrove goby Hemigobius

hyomandibular 7

maxilla metapterygoid / \ V'C \ palatine \ J I M premaxilla symplectic [yj pterygoid

quadrate

preoperde

dentary articular

Fig. 1. Jaws and Suspensorium of Hemigobius hoevenii, male, ex NTM S. 11065-002, Leanyer Swamp, Northern Territory. Scale bar = 1 mm. Black areas are cartilage. scales. Belly scales mostly cycloid, anterior row i extending up behind symphysis, so that half to third ctenoid. Body scales ctenoid up four papillae appear to be present). to pectoral base, ctenoid scales sometimes Coloration of fresh material. An extending above posterior part of opercle. illustratibn is given in Bleeker (1983: pi. Genital papilla in female short, rounded, 438, fig. 17; as Microgobius hoevenii) of this slightly flattened toward blunt tip; papilla in species. It shows the bands on the first dorsal male slender, elongate and flattened, fin, and the oblique blackish bars over narrowing toward pointed tip; small finger­ lighter background (rather pinkish, with like protrusion at tip may be present. lower half of head yellow). Head pores present, in reduced pattern From colour photograph in Zhu (1988: (Fig. 2). Anterior and posterior interorbital fig. 162), identified as Mugilogobius pores paired; latter pair never joined by obliquifasciatus Wu and Ni, of two H. canal, always separate. Postorbital and hoevenii in an aquarium. Head and body infraorbital pores present. No preopercular greyish yellow to greyish pink, with dark pores. No lateral canal over preoperculum. brown oblique bars and other markings, Sensory papillae pattem longitudinal, as in edges of oblique bars quite black, darker Figure 2. Papilla row p consisting of widely than colour in bars’ centres. Greyish pink spaced papillae; row largely replaced by streak, with brown streak on either side, interorbital canal. Cheek papillae rows short; extending from front of eye to upper jaw. Iris papillae rows rather broken-up, scattered, very dark brown. Ocellate black caudal spot rows a, cp and c composed of few large surrounded by almost pinkish colour. Both papillae; rows b and d short, composed of dorsals translucent yellowish grey with sniall, close-set papillae. Preopercular margin blackish brown markings; second dorsal papilla row e often broken up into short with broad transparent margin. Caudal fin sections or pairs of papillae following margin. translucent, with few brownish streaks along Three s rows present on snout, of one papilla fin ray bases. Pectoral base with distinct each. Single ƒ row papilla on each side of black line near upper edge, and indistinct mandibular symphysis, behind lip (sometimes pinkish band along bases of fin rays.

29 Helen K. Larson

»o'*

cp

Fig. 2. Hemigobius hoevenii papillae pattem. CAS/SU 38636, Coron, Busuanga, Philippines. Scale bar = 1 mm.

Live specimens from Singapore of opercle down and across branchiostegal mangroves noted (by author) as being membranes, becoming diffuse at isthmus; mostly pinkish with dark brown bars and black line may be almost indistinguishable other markings. Lower half of body pale from brown mottling on head in heavily- pinkish to pinkish brown, upper half light pigmented specimens. brown to pinkish. Pectoral fins yellowish Six broad blackish to brown oblique bars with gold mark on ventral half of fin base. crossing back and side, bars oriented Chrome yellow area just above and anterior anteriorly; first bar crossing nape in front of to black caudal base spot. first dorsal fin and extending (diffusely) onto Live juveniles from mangroves near opercle, last bar beginning below uppermost Darwin, Northern Territory, with dull few rays of caudal fin. Posteriormost bars whitish yellow body with brownish black often more distinct, due to contrasting lighter markings. Peritoneum silvery blue, dorsally background. Bars sometimes pointed blotched with dark brown. Light iridescent ventrally or partly broken up and patch present on each side of chest, and interconnecting with interspersed mottled similar blue pigment on lower pectoral fin dark blotches or spots. At upper base of rays and scattered across branchiostegal caudal fin, distinct round black spot present, rays. First dorsal fin mainly black, surrounded by light brown or whitish; spot translucent anteroventrally, with bright partly ocellate in many specimens. Below orange stripe through centre. this spot, variably shaped blackish oblique Coloration of preserved material. Head blotch present; blotch forming spot, streak or and body grey to light brown, lighter ocellate curved black line. Belly and ventral ventrally and sometimes posteriorly (Figs 3- part of body whitish or mottled with brown. 4). Top and side of head indistinctly mottled Pectoral base with small black spot or short with brown to greyish brown; usually most horizontal stripe near upper edge. distinct markings being two brown streaks Peritoneum dark brown, fading at lower from front edge of eye to upper lip, areas sides toward belly. between and on either side of streaks usually First dorsal fin divided into three nearly paler than surroundings. Black to dark equal bands: lowermost band dark grey to brown line running along anteriormost edge brown, central band narrowest, clear to

30 Review of the mangrove gob> Heimgohii o

Fig. 3 . Hemigobius hoevenii f e m a le . 29 mm SL. ZRC 21S72-2I906. Singapore

Fig. 4. Hemigobius hoevenii, incile. 31 ram SL, NTM S. 1396S-008. Singapore

whitish, outermost band usually dusky Comparisons. This species can be brownish, with ups of fin spines darkest. distinguished from the other known species I.owermosi band usually with black spot in the genus. H. m in g i, by having more between fourth and sixth spines; sometimes pectoral rays (usually 15-16 versus 14 in II. second, poorly developed, black spot present m ingi), the cheek below the eye being naked near base of third spine. Second dorsal tin (versus two or three rows of scales below the dusky, with broad white to translucent margin, eye), two dark bands, not forming a black and about three indistinct rows of shon dark spot, on the first dorsal fin (versus distinct vertical blotches (blotches oriented along fin black spot posteriorly), body shape (rather rays); uppermost row of blotches usually flat-headed and slender bodied versus forming continuous dark edge just below square-headed and deep-bodied) and in live white marginal band; blotches in lowermost colour. row sometimes coalescing and forming three This species superficially resembles a dark patches evenly spaced along fin base. Mugilogobius (in ils si/e. having a Anal fin plain dark grey to brown, with broad somewhat depressed head with rounded white to translucent margin. Pectoral fin snout, similar body form and preference for translucent to dusky, with blackish pigment shallow mangrove habitat), bul can be along fin rays. Pel vies pale to dusky grey with distinguished by possessing headpores. 17 broad whitish margin, fraenum whitish. segmented caudal rays, a long coiled gui and Caudal tin mostly plain greyish, with two to a relatively small mouth with reduced lips. three \cnical curving dark hands behind Distribution. Specimens are known ocellate black spot, irregular dark spots and from Mong Kong. Borneo. Brunei. Sabah. streaks sometimes present; posteliormost edge Thailand. Singapore, the Philippines. Papua of fin often whitish. New Guinea and Nonhem Australia (Fig. 5).

31 Helen K. Larson

•V species.

■ O co Hemigobius

o o o CO o <£> CO Fig. Fig. 5. Distribution of

32 Review oí the mangrove goby Hemigobius

A i

tig . 6. Microgobius hoevenii ( - Hemigobius hoevenii), from B lee 1er 1983: pi. 438. fig. 17. Courtesy of the Smithsonian Institution Press.

Zhu (1988; English Appendix, un* interorbital width given is closer to the least paginated) stated that this species "...is found bony interort>ild! width, not least fleshy in brackish water in the estuary of the Nandu width. Both Doug Hoese (AMS) and the River (Chinai and in freshwater ol the author independently examined the holotype Hainan Island". A colour slide sent by 1- and obtained similar measurements. Shiung Chen (Bristol University) ol a RMNH 4457 is a little small <32 mm SL.) specimen he collected from Taiwan, for agreement with the type specimen in confirms the species' occurrence at that Bleekcr’s description. The type was given as island. 45 mm TL by Blecket. The greatest caudal Ecology Hemigobius hoevenii can be fin length obtained for a female was 7 7 mm. .{»lile abundant in mangrove estuaries and which would only bring the type to about 40 >trcams. It is most easily observed in the mm TL. The longest caudal fin was recorded shallows and in small isolated pools, where for a 36 mm SL male (10.2 intii CL». they lie concealed under leaf litter and Allowance should be made, however, for the detritus. Two specimens have been collected missing jaws and damaged head of the type from Nypa palm leaf axils; the palms were and the specimen is here accepted as being fringing a flowing estuarina river. the holotype. Murphy (1990) reports that this species Bleeker’s (1983) ligure of Microgobius i Pseudogobius hoevenii) feeds very close hoevenii (Fig. 6) show s the oblique dark bars ‘he water’s edge at low tide in Singapore of the species. Bleeker (1851) considered mangroves and that lie has observed a that Gobius hoevenii was related to G. M ve i men captured by a large wolt'-spidor of poicilosoma <= Pseudogobius poicilosomus). tile genus Thalassius (the goby was actually Koumans (1953. 125) confused this twice the body length of the spider I. species with , placing Remarks. Blocker’s female holotype of species belonging to V/. chulae in synonymy G. hoevenii (RMNH 4457» is in very poor with Stigmatogobius hoevenii ( = condition: the tail is broken, the jaws are Hemigobius hoevenii). In his 1953 work, he missing and the skin front the top of the head describes the imerorbital pores of fi'J upper cheeks is missing. The bony Hemigobius but the colour pattern of M irooves of the imerorbit.il canals can he chulae (Koumans 195.' 125-126). ■■served, and the counts and proportions Type specimens of Mugilogobius ;rce with those of more recently-collected obliquifasciatus Wu and Ni. 1985, were specimens. The small imerorbital width unavailable for study. However, the senior given her« for the holotype probably reflects author of this species. Dr Wu of Shanghai the condition of the specimen, therefore the 1 niversity. confirmed via correspondence

3 3 Helen K. Larson

Table 5. Measurements (mm) of Hemigobius mingi (Herre, 1936).

Character Holotype Males Males Males Mean Females Females Females Mean Minimum Maximum n = 18 Minimum Maximum n = 18

Head Length 10.6 4.3 11.5 8.2 4.6 12.0 9.7 Head Depth 6.9 2.7 8.9 5.6 3.0 8.6 6.5 Head Width 7.3 3.3 8.3 5.9 3.4 8.9 7.0 Body Depth 10.8 3.3 13.1 7.7 3.7 12.1 9.3 Body Width -; 1.7 6.8 4.1 3.6 7.8 5.6 Caud. Ped. Length 12.8 3.9 12.8 8.7 4.5 15.1 10.1 Caud. Ped. Depth 7.1 1.9 8.3 - 4.9 2.1 9.0 5.7 Snout 3.3 1.2 3.7 2.6 1.3 4.1 3.1 Eye 3.5 1.4 3.6 2.7 1.6 3.7 3.0 Jaw 3.0 1.3 3.7 2.6 1.6 3.8 3.0 Interorbit 4.9 1.9 5.2 3.8 2.3 6.1 4.4 Pectoral 7.8 2.8 8.0 5.7 4.5 9.7 6.8 Pelvic 6.7 2.5 8.1 5.2 2.8 7.8 5.9 Caudal 11.0 3.9 12.0 8.3 6.5 11.9 9.8 Longest D1 spine 6.2 2.2 6.7 5.5 3.5 6.9 5.9 and photographs that the species was the Hemigobius mingi (Herre, 1936) same as H. hoevenii. (Figs 5, 7-12; Tables 1-3, 5) Herre’s (1939) record of Vaimosa hoeveni (sic) from Middle Andaman Island probably Gobius melanurus Bleeker, 1849: 31 refers to M. chulae. Hemigobius hoevenii has (Java) [not Gobius melanurus Gmelin]. - no more than 10 predorsal scales (Herre Günther 1961: 33. gives 13 predorsal scales for his Andaman Hemigobius melanurus Koumans - 1931: specimen), whileM. chulae has 11-15 scales. 78; - Bleeker 1983: pi. 433, fig. 9. Munro (1967), in a key, refers to a record mingi Herre, 1936: 8-9, pi. of Stigmatogobius hoeveni from West New IV (Pulau Ubin, Singapore). - Fowler 1938: Guinea, which could be of Hemigobius 266; - Koumans 1940; 151. hoevenii, Mugilogobius chulae or a currently Sphenentogobius vanderbilti Fowler, undescribed Mugilogobius (Larson in press) 1940: 396-397, figs 8-11 (Medan, Sumatra). (location of Munro’s specimens unknown). - Böhlke 1984: 111. Chatterjee’s (1980) record of Stig­ Stigmatogobius mingi - Koumans 1953: matogobius hoevenii from West Bengal is 118-119. probably not a Hemigobius hoevenii or a Hemigobius bleekeri Koumans, 1953: Mugilogobius species, but possibly a 191-192, fig. 47 (replacement name for Drombus, as he illustrates the fish as having Gobius melanurus Bleeker, 1849, not two transverse papillae rows on the cheek. Gobius melanurus Bloch and Schneider, Also, his predorsal scale count of 10-11 is 1801). - Kottelat et al. 1993: 146, pi. 67. little high for H. hoevenii. Unfortunately, Material examined. 68 specimens (11.3- Chatteijee did not give sufficient information 55). INDONESIA: Lectotype of Gobius to allow the reader to confidently identify the melanurus and Hemigobius bleekeri, RMNH species from the text and drawings. It is 4501, 55 mm SL female, Java, in sea (Sunda unclear what he meant by the “nasal sensory Archipelago on jar label). Paralectotypes of canal-pores” illustrated in Figure IB; they Gobius melanurus and Hemigobius bleekeri, may be the large sensory papillae belonging ex RMNH 4501, 2 (40.5-55), same data as to the nasal c series. lectotype. Holotype of Sphenentogobius Nguyen’s (1991) record of Mugilogobius vanderbilti, ANSP 68714, 40.5 mm female, latifrons from Ha Nam Ninh in Vietnam is Medan, Sumatra, Vanderbilt Expedition, 23 accompanied by a drawing that is clearly of May 1939. SINGAPORE: Holotype of H. hoevenii. Gnatholepis mingi, GAS 30960, 43 mm SL

34 Review of the mangrove goby Hemigobius male, Pulau Ubin, A.W. Herre, 1934. ZRC rays 13-15 (mean 14; holotype with 15 on 20263-72, 10(28-47), Sungai Punggol, 22 right, 14 on left); segmented caudal rays March 1966; ZRC 20192-37, 46(11.3-41.2), always 17*; caudal ray pattern 6/6 to 9/8 Sungei Punggol, C.K. Quek and M. Dali, 19 (modally 8/7*); branched caudal rays 12-17 October 1965; CMK 8322, 2(39-41), Kranji (mean 15*); unsegmented (procurrent) mangroves near Sungei Buloh, M. Kottelat caudal rays 6/7 to 8/8; longitudinal scale and D. Murphy, 8 April 1992; NTM count 27-32* (mean 29); TRB 9*-12 (mean S.14235-003, 4(14.5-22), Sungei Buloh 10); predorsal scale count 8-10* (mean 9); mangroves, K. Lim, 30 January 1992; ZMH circumpeduncular scales 12* or 13 (in one). 19308, 2(26.5-35), aquarium import, Gili rakers on outer face of first arch 5+12 to Reichelt, 1 December 1090. THAILAND: 6+12 (modally 5+12). Dorsal pterygiophore URM P.6677, 4(33-40), Ranong, 9 March formula 3-12210 (in 12). Vertebrae 10+16 1982; NTM S.14288-001, 1(46), in ponds (in nine), 10+17 (in one), 11+15 (in one), within research station, Ta-Chaluab, 11+16 (in one). Neural spine of first vertebra Chantaburi Province, Mahidolia Project, very short and broad, sharply bent NIFI, Chulalongkorn University and posteriorly at halfway point (in 11). Two Mahidol University parties, 2 June 1990; epurals (in 10) or one very broad epural (in KUMF uncatalogued, 1(40.5), Tak Bai two). Two (in two) or three (in 10) anal canal, Narathiwat Province, D. Tanwilai, 25 pterygiophores before haemal spine of first September 1984; CMK 5419, 1(32.5), Ban caudal vertebra. Lachrymal enlarged, almost Pliu, near Chantaburi, M. Kottelat, 21 March square. 1980. NO LOCALITY: RMNH 12580,2(28- Body rounded anteriorly (especially in 35.5). females); belly rounded and body Other material examined (but not used in compressed posteriorly in both sexes. Head description). 28 specimens, from the square in cross-section, depth about following localities. SINGAPORE: ZRC equalling width; profile blunt to rounded; 30165-70, 6, Sungei Buloh East mangroves; nape profile often curving behind flattened NTM S.13961-006, 12, Mandai Kecil; NTM broad interorbital, HL 3.1-4.7 (mean 3.7) in S.13957-008, 9, Sungei Pandan. SL. Depth at posterior preopercular margin THAILAND: USNM 316180, 1, Cheh 1.3-1.6 (mean 1.5) in HL. Width at posterior Bilang, Satul, Satul Province. preopercular margin 1.3-1.6 (mean 1.4) in Diagnosis. Deep bodied Hemigobius, HL. Mouth small, subterminal, slightly with blunt head and compressed body; oblique, with rounded snout overhanging second dorsal rays 1,6-8 (modally 1,7); anal upper lip; jaws forming angle of about 15- rays 1,6-7 (modally 1,7); pectoral rays 13-15; 20° with body axis; jaws generally reaching longitudinal scales 27-32; TRB 9-12; to below anterior margin of eye in both predorsal scales large, 8-10, edges often sexes. Upper lip narrow, smooth, without scalloped, extending close up behind eyes; fleshy fimbriae; lower lip reduced, narrow body scales ctenoid; side of lower jaw lip fold free at lower comer of jaw, rest of lip strongly curved upward, forming ridge at smoothly fused to underside of head; lower symphysis; first dorsal fin low, rounded, jaw curving upward and forming ridge at with no filamentous spines; body greyish to symphysis (resembling jaw of mugilid). brownish, scales often with blackish centres Upper jaw 2.8-3.5 (mean 3.2) in HL. Eyes and five to six white diagonal bars (very large, lateral, high on head, sometimes prominent in life), marbled or somewhat forming part of dorsal profile, 2.7-3.6 (mean ocellate blackish spot on caudal base, eyes 3.2) in HL. Snout bluntly rounded, and blue when live; known only from mangrove inflated over top lip, 2.8-3.6 (mean 3.2) in areas in Thailand, Singapore and Indonesia. HL. Interorbital broad, flat, 1.8-4.1 (mean Description. Based on 36 specimens, 2.3) in HL. Top of head above preopercular 13.5-34 mm SL. Counts of holotype of margin up to close behind posterior nostril Gobius melanurus indicated by asterisk. often with fine villi, these relatively sparse First dorsal VI*; second dorsal 1,6-1,8 on scales when compared with naked skin (mean 1,7*); anal 1,6-7 (mean 1,7*); pectoral (villi often only visible in specimens with

35 Helen K. Larson well-preserved mucous coat; visible in small pointed teeth, tooth band widest toward holotype). Body usually quite round in rear of jaw; teeth covering raised mandibular abdominal region, especially in females, symphysis. No difference in teeth between depth at anal origin 3.5-4.Ó (mean 4.0) in SL. males and females. Caudal peduncle long, compressed, length Predorsal scales medium to large, with 3.3-4.5 (mean 3.6) in SL. Caudal peduncle largest scales anteriormost, scales reaching depth 5.5-7.3 (mean 6.4) in SL. forward to close behind eyes; edge of First dorsal fin low, rounded, tips of predorsal scales usually crenulate or second to fourth spines free, second or third scalloped, most conspicuous in larger spines longest or subequal; spines always specimens. Operculum covered with cycloid falling short of second dorsal origin when scales. Cheek with two or three rows of depressed. First dorsal spine usually shorter cycloid scales; single row of scales below than next three. Second dorsal spine length eye always present in adults, may be absent 3.5-6.7 (mean 4.5) in SL. Third dorsal spine in specimens about 14 mm SL or less; often length 2.2-6.4 (mean 4.9) in SL. Fourth one or two scales on upper preopercle close dorsal spine length 4.5-6.8 (mean 5.5) in SL. behind eye. Pectoral base covered with Second dorsal and anal fins low, cycloid scales. Prepelvic area covered with posteriormost rays usually longer than small cycloid scales. Belly scales ctenoid; anteriormost, rays only extending about half few small cycloid scales around anus length of caudal peduncle when depressed. sometimes present. Body scales ctenoid up Pectoral fin small, short and rounded, central to pectoral base, ctenoid scales sometimes rays longest, 4.7-6.1 (mean 5.4) in SL; in extending above posterior part of opercle. adults, rays all branched but for lower and Genital papilla in female short, conical uppermost rays (uppermost ray or two and rather pointed, slightly flattened toward usually unbranched). Pelvic fins short, tip; papilla in male slender, flattened and rounded, reaching half (or less) distance to pointed to slightly rounded at tip; may be anus, 5.2-8.0 (mean 6.0) in SL. Caudal fin short or elongate (up to half length of anal rounded, 3.4-5.0 (mean 3.8) in SL. fin spine). No mental fraenum, chin smooth. Anterior nostril in very short tube, placed just behind Head pores present, in reduced pattern upper lip, tube oriented forward. Posterior (Fig. 8). Anterior and posterior interorbital nostril oval, with low rim, placed halfway pores paired; latter pair of pores never joined between anterior margin of eye and edge of by canal, always separate. Postorbital and preorbital. Gili opening usually extending infraorbital pores present. No preopercular forward to just under opercle. Inner edge of pores. No oculoscapular canal over shoulder girdle smooth with no ridge or preoperculum. In small (juvenile) flange (in 21) or with low bony ridge or specimens, anterior interorbital pores may flange (in 10). Gili rakers on outer face of first be absent. arch very short, pointed and without spines, Sensory papillae pattern longitudinal, as longest two rakers on either side of angle of in Figure 8. Papilla row p consisting of arch; tiny papillose flaps on pads above first widely spaced papillae; row largely replaced arch; rakers on inner face of first arch more by interorbital canal. Cheek papillae rows slender; outer rakers on other arches similar to short, interrupted by scale rows on cheek; those on first arch; inner rakers on other papillae rows rather broken-up, scattered, arches twice length of first arch inner rakers. rows a, cp and c composed of few large Tongue tip usually blunt, or with tip concave papillae; rows b and d short, composed of in centre (almost bilobed in few specimens). small, close-set papillae. Three s rows Outer teeth in upper jaw very small, flattened, present on snout, consisting of one or two slightly curved, with pointed or somewhat papillae each. Single ƒ row papilla on each spatulate tips (Fig. 7); behind this row, one or side of mandibular symphysis, behind lip. two rows of very small sharp teeth often Coloration of fresh material. A dead partly concealed by flesh of mouth. Lower specimen is shown in Kottelat et al. (1993: jaw with band of three to five rows of very 146, pi. 67), and a coloured illustration in

36 Review of the mangrove goby Hemigobius

premaxilia hyomandibular

} ( ' ( palatine metapterygoid

maxilla \ffv pterygoid

symplectic

quadrate dentary articular preopercle

Fig. 7. Jaws and Suspensorium of Hemigobius mingi, female, ex URM R6677, Ranong, Thailand. Scale bar = 1 mm. Black areas are cartilage.

Bleeker (1983: pi. 433, fig. 9). The living markings. Caudal fin greyish to pinkish with fish are considerably more conspicuous. grey vertical banding and spotting. Juveniles Notes taken by the author, from living with most intense fin colours. Singapore specimens, follow. Coloration of preserved material. Head Head and body yellowish brown with and body dark grey to light brown, usually pale yellowish to whitish yellow bars paler on lower abdomen and belly. Most alternating with dark brownish bars on side scales with darker spot near centre; often of body. Body bars most clearly defined in giving appearance of indistinct thin lines young fish; pale bars appear wider in young. midlaterally. Six variably oblique white bars Area around dark spot at base of caudal present, crossing dorsal midline (most slightly more intensely yellow than yellowish distinct in well-preserved fresh specimens): (pale) body bars. Juveniles and specimens up first bar crossing above opercle, second bar to about 25 mm with three bright blue-white below first dorsal fin, third at gap between to whitish yellow or white bars across top of dorsals, fourth at midpoint of second dorsal, nape, beginning not far behind eyes, bars can fifth across caudal peduncle and sixth be irregular or asymmetric in shape; (vertical) crossing caudal base (Figs 9-11). anteriormost bar often rounded, forming Bars breaking up laterally, with anteriormost conspicuous blotch or spot. Underside of bar often absent in larger specimens. At head silver, silvery blue or deep blue; blue caudal base behind white bar, black spot on sometimes extending up onto cheek, opercle upper half of base or vertically paired black and pectoral base. All fish with distinctive spots usually visible; spot diffuse in large pale blue eyes; blue visible when viewed adults, intense black in juveniles. Top and from above. Inside of lips and mouth golden sides of head dark grey, sometimes mottled, yellow to brownish yellow. but without any distinct pattern. Juvenile First dorsal fin with dense black spot specimens often with one or more white occupying rear half of fin; anterior half of fin spots on anteriormost predorsal scale. bright orange. Second dorsal and anal fins Peritoneum dull brownish, darkest dorsally, pinkish to yellow with darker brownish fading toward pale belly.

37 Helen K. Larson

Flg. 8. Hemigobius mingi, headpores and papillae pattem. ZRC 20263-72, Sungei Punggol, Singapore. Scale bar = 1 mm.

First dorsal fin with distal half whitish to (Fig. 5). The species has recently been translucent, with narrow black to brown collected by the author from Brunei (Bandar margin; proximal half dusky to brown with Seri Begawan, Tutong, Kuala Belait), but the large black spot posteriorly, occupying space material was not available to include in this between third or fourth spines and rear of study. fin. Second dorsal fin translucent whitish, Ecology. In Singapore, this species is with three indistinct dusky stripes; stripes common in muddy mangrove pools back from often broken up into series of vertically the main tidal flow, especially among aligned oval brown to blackish spots, Rhizophora thickets. The behaviour of although uppermost stripe usually present as Hemigobius mingi is interesting in that the fish continuous stripe; fin margin translucent hover above the substrate in an almost vertical whitish. Anal fin dusky, translucent posture, in small groups, with the bright proximally and at anterior base of first few silvery-white bars across the dorsal surface fin rays. Caudal fin dark grey to brown, with making the fish quite conspicuous against the about six or seven vertical rows of small dark background. This behaviour is unusual in spots and short streaks. Pectorals clear to that many mangrove gobies conceal whitish, fin rays with very narrow blackish themselves under leaf litter or in holes, with edges. Pelvics plain light brown to whitish. the conspicuously marked and Comparisons. Characters distinguishing being exceptions. The anteriormost this species from the only other known species spot (or bar) in this species is reminiscent of in the genus, H. hoevenii, are given under that visible in the ricefishes (Oryzias ) which Comparisons for that species. Hemigobius are syntopic with Hemigobius in the Indo- mingi has an autapomorphy in the shape of the Malayan Archipelago. lachrymal, which is quite large for this group It is apparently fairly tolerant of poor of , and almost square. conditions, as some specimens collected in Distribution. Specimens are known Singapore (Mandai Kecil) came from pools from Singapore, Thailand, Sumatra and Java in which the substrate consisted mostly of

38 Review ol the m.ui|:rx»ve goby Hemigobius

Fig. 9. Hemigobius mingi. Holotype of Gnatholepis mingi Herre. 44 mm SL. CAS 30960. Pulau Ubin. Singapore.

Fig. IO. Hemigobius mingi. Holotype Of Sphenentogobius vanderbilti Fowler. 42.5 mm SL. ANSP 68714. Vledun. Sumatra.

Fig. 11. Hemigobius mingi, male. 4(1 mm SL. one of ZRC 20263-20272. Sungai Punggol. Singapore.

ud and rotting garbage (although ii must be Remarks. Koumans »1953» created the nlmitted that the Hemigobius and other name bleekeri as a replacement name for gobie?» present appealed temporarily stunned Gobius melanurus Bleeker "not Gobius by the gases released from beneath the melanurus Broussonet not Gobius substrate as the author moved about the melanurus Gnielin in Linnaeus ... not Gobius pools dipnetling fish). melanurus B!. Schn. This \sas an

t»i H e le n K 1.ÜMIII

Fig. 12. Hemigobius minui, from Bleeker I''S.* (pi. 433. fig. M|. Courtesy ol the Smithsonian Institution Press unnecessary replacement, as is outlined name for the taxon is Gnatholepis mingi below. Herre. 1936. Broussonet (1782). at the end of his It is uncertain as to what the species description of Gobius ocellaris, listed four melanurus of Gmclin and Lacepèdc actually sets of gobioid groups and a character is (described as "pinna dorsal i unica, cauda distinguishing each group from Gahi us nigra" by Gmclin). Richardson (1846) ocellaris. Here he gate the name Gobiis referred to specimens from Canton. China, melanura and a character "pinnis dorsalibus as melanurus (after Broussonet iluabus". The character of "two dorsal manuscript and figure). No specimens are fins’Vas provided to distinguish G ocellaris known, and Richardson’s description is from Gobiis anguillaris and Gobiis based on a drawing and notes, which melanura by its possessing two dorsal fins, indicate that the fish had a pointed tail, a implying that the latter two species have one black spot on the base of the fin and one fin. Gobiis anguillaris and G. melanurus are dorsal fin (Richardson 1846). It is possible not distinguished front each other by any that Gobioides melanurus is an amblyopinc. statement. Therefore the name melanura or even a channid. Broussonet is not available as no description Miller (1987) referred to dic holotype of exists. Eschmeyer (1998) stated the name Hemigobius bleekeri Koumans as being Gobiis melanura Broussonet. 1782. as not RMNH 4501. a 40 nun SL ("40+8") female. available (no distinguishing features». Koumans (1953) stated "Types seen”, but Gmelin's ( 1789) use of the name Gobius did noi designate a ty pe of H. bleekeri, nor is mclanuros constitutes a description, as he there any indication in the specimen jar as to lists two characters (single dorsal fin. black which specimens he considered to he types caudal fin), therefore that name is available In 1988. RMNH 4501 consisted of three Lacepede (1800) correctly latinized the possible syntype specimens. 40.5-55.0 mm Greek -os to -im when he used the emended SL. of which the largest and smallest are name melanurus. and he placed the species female. In the original description. Bleeker in the genus Gobioides (along with three (1849) wrote "33 millimetr". that is. 33 mm other species). Bloch and Schneider (1801) IL. which would make these types of also referred to Gmelin’s name and corrected melanurus (and bleekeri) rather larger than the spelling toGobius melanurus. he described (he did not indicate how many Bleeker (1849) created the name Gobius specimens he had). The description may melanurus for his species from Java, without have been based on a male, as Bleeker reference to Broussonet or Gmclin. w rote: ". . appendice anali conica acula. ."; Blecker’s name is thus a primary homonym however, (his description could equally of Gmelin’s emended Gobius melanurus, the apply to some female specimens. The figure first available name. The next available given in Bleeker of Hemigobius melanurus

441 Review of the mangrove goby Hemigobius

(1988: pi. 433, fig. 9) (Fig. 12), drawn life- der gesamten Hydrobiologie und Hydrographie sized, is of a 55 mm SL fish, probably the 38(1/2): 125-183. large female in RMNH 4501. In the interests Birdsong, R.S., Murdy, E.O. and Pezold, F.L. 1988. of stability, this 55 mm SL female specimen A study of the vertebral column and median (RMNH 4501) is here designated as fin osteology in gobioid fishes with comments lectotype of Gobius melanurus Bleeker and on gobioid relationships. Bulletin of Marine Hemigobius bleekeri Koumans. Science 42(2): 174-214. Bleeker, P. 1849. Bijdrage tot de kennis der Blennioïden en Gobioïden van den Soenda- ACKNOWLEDGMENTS Molukschen Archipel, met beschrijving van 42 nieuwe soorten. Verhandelingen van het My thanks to curators and staff at the Bataviaasch Genootschap van kunsten en institutions who loaned specimens, wetenschappen 22(6): 1-40. especially Jerry Allen, WAM; Genie Böhlke,4 Bleeker, P. 1851. Vijfde bijdrage tot de kennis der ANSP; Dave Catania, CAS; Bill Eschmeyer, ichthyologische fauna van Borneo met CAS; Doug Hoese, AMS; Susan Jewett, beschrijving van eenige nieuwe soorten van USNM; Maurice Kottelat, CMK; zoetwatervisschen. Natuurkundig Tijdschrift (previously at ZSM); Kelvin Lim, ZRC; voor Nederlandsch-lndie 2: 415-442. Mark McGrouther, AMS; Nigel Merrett, Bleeker, P. 1854. Ichthyologische waaremingen BMNH; Peter Ng, ZRC; Martien van Oijen, gedaan op verschillende reizen in de residentie RMNH; Chavalit Vidthayanon, NIFI; H. Bantam. Natuurkundig Tijdschrift voor Wilkens, ZMH; Prachit Wongrat, KUMF; Nederlandsch-lndie 7: 309-326. Thosaporn Wongratana, Chulalongkorn Bleeker, P. 1874. Esquisse d’un système naturel des University, Bangkok; Tetsuo Yoshino, URM. Gobioides. Archives Néerlandaises des Many thanks to those people who helped Sciences exactes et naturelles 9: 289-331. provide fresh specimens or information: I- Bleeker, P. 1983. Atlas Ichthyologique des Indes Shiung Chen, Bristol University; Satish Orientales Néêrlandaises, par M.- P. Bleeker. Choy, formerly of University of Brunei; (Plates originally planned for planned tomes Russell Hanley, formerly of NTM; Ukkrit XI-XIV published here for the first tíme). Satapoomin, Biological Research Centre, Smithsonian Institution Press: Washington. Phuket; Dave Wilson, Territory Wildlife Bloch, M.E. and Schneider, J.G. 1801. Systema Park, Darwin; Dr Wu Han-ling, Shanghai Ichthyologiae. Iconibus ex Illustratum. Post University. Obitum Auctoris Opus Inchoatum Absolvit. My thanks to Tony Gili, BMNH, for his Correxit, Interpolavit Jo. Gottlob Schneider. useful comments on versions on this Berlin. Facsimile, Cramer and Swann: manuscript. And many thanks also to Rex Codicote. Williams, NTM technical officer, for his Broussonet, P.M.A. 1782. Ichthyologia sistens black and white photography and map piscium descriptions et icones. Emsley: preparation. London. Chatterjee, T.K. 1980. Record of Stigmatogobius hoevenii (Bleeker) from the Gangetic Delta, REFERENCES west Bengal, with a key to the Indian species of Stigmatogobius. Bulletin of the Zoological Anon. (Pearl River Fisheries Research Institute, Survey of India 2(2&3): 229-231. Shanghai Fisheries University, East China Sea Davis, T.L.O. 1988. Temporal changes in the fish Fisheries Research Institute, Fisheries School fauna entering a tidal swamp system in of Guangdong Province). 1986. The tropical Australia. Environmental Biology of freshwater and estuaries fishes of Hainan Fishes 21(3): 161-172. Island. Guangdong Science and Technology Eschmeyer, W.N. (ed.). 1998. Catalog o f fishes. Press: Guangzhou (in Chinese). California Academy of Sciences: San Francisco. Aurich, H.J. 1938. Mitteilung XXVin der Wallacea- Fowler, H.W. 1940. Zoological results of the George Expedition Woltereck, Die Gobiiden. Vanderbilt Sumatran Expedition, 1936-1939. (Ordnung: Gobioidea). Internationale Revue Part II - The fishes. Proceedings o f the

41 Helen K. Larson

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