Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019)

DESCRIPTION OF A NEW SPECIES OF (: BEMBRIDAE) FROM THE ANDAMAN SEA, THAILAND

Saki Kishimoto1*, Toshio Kawai2, Fumihito Tashiro3 and Charatsee Aungtonya4

1Graduate School of Fisheries Sciences, Hokkaido University, Hakodate, Hokkaido 041-8611, Japan 2Faculty of Fisheries Sciences, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan 3Fisheries Science Center, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan 4Phuket Marine Biological Center, Muang District, Phuket 83000, Thailand *Corresponding author: [email protected]

ABSTRACT: A new species of Bembradium (Scorpaeniformes: Bembridae) is described on the basis of a single specimen collected in 284–300 m in the Andaman Sea off Phuket, Thailand. The new species, Bembradium magnoculum, differs from all congeners by the following combination of characters: short and wide snout (length 79.9% eye diameter; width at anterior margin of anterior nostril 13.4% SL), large eye (13.2% SL), long upper jaw (20.1% SL), long preanus length (64.8% SL), long predorsal-fin length (41.9% SL), short second dorsal-fin base (57.1% HL), long anal-fin base (23.9% SL), short lower caudal peduncle length (25.8% HL), short pectoral fin (45.9% HL), short pelvic-fin spine (16.0% HL), small spines of similar size on nasal bone and broad dermal flap of anterior nostril.

Keywords: Pisces, Bembradium magnoculum sp. nov., Indian Ocean, BIOSHELF

INTRODUCTION the specimen was found to be a new species. The study provides a formal description of the new The genus Bembradium Gilbert, 1905, placed species. in the family Bembridae, was established for Bembradium roseum Gilbert, 1905, based on a MATERIALS AND METHODS specimen captured off Hawaii. A second species, Bembradium furici Fourmanoir and Rivaton, 1979, Specimens examined in this study are in the was described from a New Caledonian specimen. Kyoto University, Japan (FAKU), Muséum national Since then, this genus has included only the two d’Histoire naturelle, France (MNHN), Phuket species, B. roseum recorded from Hawaii, Emperor Marine Biological Center, Thailand (PMBC), and Seamount Chain, Japan, Taiwan, and New Caledonia, National Museum of Natural History, Smithsonian in depths of 138–800 m (Gilbert, 1905; Chave and Institution, USA (USNM). Mundy, 1994; Rivaton and Bourret, 1999; Yamada Counts and proportional measurements gen- et al., 2007; Ho et al., 2009; Fricke et al., 2011; erally follow Hubbs and Lagler (1958), and scale Nakabo and Kai, 2013), and B. furici still known counts follow Nakabo (2002). Snout width was only from New Caledonia in 360–400 m (Fourmanoir measured at the anterior margin of the orbit, and and Rivaton, 1979; Poss, 1999; Fricke et al., 2011). at the anterior margin of the anterior nostril. Standard In 1999, a single specimen identified as and head lengths are abbreviated as SL and HL, Bembradium was caught in the Andaman Sea off respectively. Measurements were made to the Phuket, Thailand, during the Biodiversity of the nearest 0.1 mm with a digital caliper and a divider. Andaman Sea Shelf (BIOSHELF) project operated Terminology of head spines generally follows by the Phuket Marine Biological Center of Thailand. Knapp et al. (2000) except for the nasal ridge. Nasal As a result of a comparison of the specimen with ridge is on the nasal bone. types and other specimens of the two congeners, 10

Phuket mar. biol. Cent. Res. Bull.

TAXONOMY Opercle with two smooth ridges (Fig. 2). Operclular Family Bembridae flap well developed. Gill rakers on upper limb Genus Bembradium Gilbert, 1905 three plus 12 (left) or 11 (right) on lower limb, including single raker at angle. Lateral line running Bembradium Gilbert, 1905: 637 (type species nearly parallel to dorsal contour of body. First dorsal Bembradium roseum Gilbert, 1905); neuter. fin separated from second fin. First dorsal fin originating above pectoral-fin base. Anal fin origi- Generic diagnosis. Anterior part of head strongly nating slightly posterior to vertical through origin depressed, becoming less so posteriorly; lower jaw of second dorsal fin. Pectoral fin rounded; its tip short and not protruding; many small spines on reaching to vertical through second anal ray base. suborbital ridge; origin of pectoral fin posterior to Posterior tip of depressed pelvic fin reaching vertical vertical through pelvic fin origin; lateral line running through base of seventh dorsal fin spine. Posterior near middle of body; body scales large (Gilbert, margin of caudal fin slightly rounded. 1905; Nelson et al., 2016). Some small spines of similar size on nasal bone. Lachrymal with (left side) or without (right side) Bembradium magnoculum sp. nov. single small spine. Two ridges with small spines Figs. 1A, 1B, 2, 3A; Tables 1, 2 on suborbital region; length of spines on anterior ridge [12 (left side) or 11 (right side)] much shorter Holotype. PMBC 30062, 115.9 mm SL, 7°30'N than on posterior ridge [18 (left side) or 16 (right 97°50'E to 7°31'N 97°51'E, 284–300 m depth, side)]. Single preocular spine in front of eye; tip 9 Nov. 1999, Agassiz trawl, R/V Chakratong of spine forked (left side) or simple (right side). Tongyai, St. I6. Interorbital region with 17 (left side) or 18 (right side) supraorbital spines. Fourth and fifth infraor- Diagnosis. Snout length 79.9% eye diameter; snout bital bones serrated. Postocular region with 18 (left width at anterior margin of anterior nostril 13.4% SL; side) or 14 (right side) small spines. Pterotic with eye diameter 13.2% SL; upper-jaw length 20.1% ridge possessing three spines. Parietal with single SL; preanus length 64.8% SL; predorsal-fin length prominent spine. Two nuchal spines. Single robust 41.9% SL; length of second dorsal-fin base 57.1% spine on preopercle with small supplemental spine HL; length of anal-fin base 23.9% SL; length of just before robust spine; robust spine not reaching lower caudal peduncle 25.8% HL; pectoral-fin to posterior margin of opercle (Fig. 2). length 45.9% HL; pelvic-fin spine length 16.0% Teeth on jaws, vomer, palatine and first basi- HL; size of spines on nasal bone small and of branchial conical, uniformly very small; upper-jaw similar size; dermal flap of anterior nostril broad. teeth forming separate tooth-bands on each side. Scales ctenoid on body, cheek, occiput, isthmus, Description. Counts and measurements are provided jugular, pectoral-fin base and basal 1/6 of pectoral in Tables 1 and 2. Body elongate and gradually fin; scales on body largest, those on pectoral fin compressed posteriorly, its profile slightly arched smallest. dorsally and nearly straight ventrally, deepest at the Coloration when fresh unknown. When pre- first spine of first dorsal fin. Head large, depressed, served in alcohol, head, body and all fins uniformly especially anteriorly. Snout short and wide, strongly yellowish brown. depressed, with broad dermal flap on anterior nostril (Fig. 3A). Mouth subterminal; maxilla Distribution. Known from only the sea floor at extending beyond anterior margin of eye, but not 284–300 m depth off Phuket, Thailand in the reaching past center of eye. Orbit and eye very southern Andaman Sea. large; interorbital region very narrow and grooved. 11

Description of a new species of Bembradium

Etymology. The specific name magnoculum from Key to the species of Bembradium the Latin magnus, large, and oculus, eye, refers to 1a. Dermal flap of anterior nostril narrow; eye small the large eyes of this species. (9.8–11.8% SL)..…...... B. roseum 1b. Dermal flap of anterior nostril broad; eye large Remarks. In this study, a new species is added to genus Bembradium, bringing the total number of (12.4–13.4% SL)…………...... 2 species in this genus to three. They can be 2a. Posteriormost spine on nasal bone much stronger distinguished from each other by differences in than others; snout long (87.8–98.6% eye diameter) snout length (79.9% eye diameter in B. magnoculum …...………………...... … B. furici vs. 87.8–98.6 in B. furici and 86.4–115.7 in B. roseum), 2b. Spines on nasal bone small and of similar size; snout width at anterior margin of anterior nostril snout short (79.9% eye diameter)….…..…… (13.4% SL in B. magnoculum vs. 10.5–12.9 in B. furici and 9.9–11.6 in B. roseum), eye diameter ………...... …...... … B. magnoculum sp. nov. (13.2% SL in B. magnoculum and 12.4–13.4 in B. furici vs. 9.8–11.8 in B. roseum), upper-jaw length Comparative materials (20.1% SL in B. magnoculum vs. 17.9–19.5 in B. Bembradium furici: MNHN 1978-0470, holotype furici and 17.7–19.4 in B. roseum), preanus length of B. furici, 120.2 mm SL, west of the Pines Island, (64.8% SL in B. magnoculum vs. 58.0–64.3 in B. New Caledonia, 360 m depth, trawl; MNHN 2004- roseum), predorsal-fin length (41.9% SL in B. 2577, 3 specimens, 104.2–117.1 mm SL, 380–389 m magnoculum vs. 38.5–40.5 in B. furici and 35.7– depth, 23 Oct. 2003; MNHN 2004-2691, 1 specimen, 41.0 in B. roseum), length of second dorsal-fin base 99.6 mm SL, 395–400 m depth, 5 Nov. 2003. (57.1% HL in B. magnoculum and 48.9–58.0 in B. furici vs. 58.8–73.6 in B. roseum), length of anal-fin Bembradium roseum: USNM 51617, holotype of base (23.9% SL in B. magnoculum and 23.3–27.6 B. roseum, 71.7 mm SL, Pailolo Channel, Hawaii, in B. roseum vs. 21.7–22.0 in B. furici), length of 252 m depth; FAKU 1818, 1820, 1821, 3 specimens, lower caudal peduncle (25.8% HL in B. magnoculum 85.3–97.8 mm SL, off Owase, Mie, Japan, 13 May vs. 30.9–35.0 in B. furici vs. 27.4–38.6 in B. roseum), 1936; FAKU 1890, 1 specimen, 70.7 mm SL, off pectoral-fin length (45.9% HL in B. magnoculum Owase, Mie, Japan, 27 May 1936; FAKU 3103, 1 vs. 47.9–53.5 in B. furici and 47.6–60.5 in B. roseum), specimen, 75.3 mm SL, Miya Market, Aichi, pelvic-fin spine length (16.0% HL inB. magnoculum Japan, 7 Nov. 1936; FAKU 72512–72514, 3 specimens, vs. 19.7–21.5 in B. furici and 16.6–22.9 in B. 97.4–116.0 mm SL, Emperor Seamount Chain, 23 roseum), size of spines on nasal bone (small and Nov. 1994; FAKU 72541, 72549, 2 specimens, of similar size in B. magnoculum and B. roseum 104.3–104.9 mm SL, Emperor Seamount Chain, vs. posteriormost spine much stronger than others 1994; FAKU 72577, 1 specimen, 110.4 mm SL, in B. furici) and shape of dermal flap of anterior Emperor Seamount Chain, 14 Nov. 1994. nostril (broad in B. magnoculum and B. furici vs. narrow in B. roseum; the flap broken in holotype of B. roseum) (Fig. 3; Tables 1, 2). 12

Phuket mar. biol. Cent. Res. Bull.

Figure 1. Three species of Bembradium. (A) B. magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL; (B) dorsal view of head of B. magnoculum sp. nov., PMBC 30062, holotype; (C) B. furici, MNHN 1978-0470, holotype, 120.2 mm SL; (D) B. roseum, USNM 51617, holotype, 71.7 mm SL. Scale bars 10 mm.

Figure 2. Lateral view of head of Bembradium magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL. Spines: LAC, lachrymal; NC, nuchal; NR, nasal ridge; PA, parietal; PO, preopercular; POC, preocular; PSOC, postocular; PT, pterotic; SBO, suborbital; SPO, supraorbital; SS, supplemental preopercular. Scale bar 10 mm. 13

Description of a new species of Bembradium

Figure 3. Dermal flap on anterior nostril: (A)B. magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL; (B) B. roseum, FAKU 72541, 104.3 mm SL. Scale bars 1 mm.

ACKNOWLEDGEMENTS Development Institute, Thailand), and staff of the Reference Collection of PMBC for their kind help We sincerely thank Yoshiaki Kai (FAKU) for during research in PMBC; J. Williams, J. Clayton, S. loaning specimens and providing valuable com- Raredon and K. Murphy (USNM) and P. Pruvost, G. ments; William J. Richards (National Marine Fisheries Zora, L. M. Duque-Vélez and A. Laurent (MNHN) Service, Southeast Fisheries Science Center / NOAA) for their organization and assistance during FT’s for his critical reading of an early manuscript and visit. We also thank Hisashi Imamura (Hokkaido English correction; Somkiat Khokiattiwong (former University, Japan) and Naohide Nakayama (formerly PMBC Director), Somchai Bussarawit (former FAKU) who supported our examination. Specialist, Marine and Coastal Resources and 14

Phuket mar. biol. Cent. Res. Bull. . et al * 2 26 I, 5 3–5 5–8 2–3.5 10–11 23–27 27–28 others 70.7–116.0 VIII–XI-11–12 6–7+6–7=12–14 2–4+7–12=10–15 B. roseum 11 26 28 I, 5 71.7 IX-12 23–25 3+6=9 No data No data No data No data holotype * 1 10 I, 5 23–30 28–29 No data No data No data No data No data others IX-11–12 99.6–117.1 2–3+7=9–10 B. furici 10 29 I, 5 120.2 IX-11 24–25 2+8=10 No data No data No data No data No data holotype . sp. nov. 4 2 5 11 25 26 28 I, 5 115.9 IX-12 7+6=13 holotype Bembradium 3+11–12=14–15 B. magnoculum partly from Formanoir and Rivaton (1979), Kanayama (1982), Yatou (1985), Ochiai (1988), Imamura (1997), Yamada Yamada (1985), Ochiai (1988), Imamura (1997), Yatou partly from Formanoir and Rivaton (1979), Kanayama (1982), partly from Poss (1999). . (2009), and Nakabo Kai (2013). et al B. roseum B. furici Meristic counts of three species Dorsal fin Anal fin Caudal fin Pectoral fin Pelvic fin Gill rakers Vertebrae Lateral-line scales Predorsal scales Scales above lateral line Scales below lateral line *Data for *Data for Standard length (mm) Counts Table 1. Table 1 2 (2007), Ho 15

Description of a new species of Bembradium others* 2.9–4.5 1.6–1.9 9.9–11.6 10.0–11.4 10.0–11.8 14.7–21.9 12.1–18.1 41.4–47.6 13.5–18.8 16.2–20.5 12.0–16.3 14.9–16.2 16.6–19.4 17.9–19.4 58.7–64.3 37.2–41.0 62.7–68.1 15.3–23.0 25.3–29.1 23.3–27.2 70.7–116.0 B. roseum 9.8 2.4 11.3 71.7 15.9 15.5 39.6 15.6 17.9 12.6 10.5 12.6 15.9 17.7 58.0 35.7 61.2 20.4 29.1 27.6 No data holotype others 1.5–1.8 No data No data 11.7–12.3 11.8–14.5 18.3–20.6 15.9–17.5 44.4–46.8 18.3–21.5 17.6–19.5 10.5–12.9 12.4–13.4 14.4–15.2 18.3–19.2 17.9–19.5 38.5–40.5 63.1–66.9 18.1–21.1 22.9–26.1 21.7–22.0 99.6–117.1 B. furici 20.5 18.3 43.8 13.4 13.1 19.3 120.2 No data No data No data No data No data No data No data No data No data No data No data No data No data No data holotype . sp. nov. 3.2 2.0 20.8 16.5 46.8 18.1 20.3 10.5 15.4 13.4 13.2 16.6 19.4 20.1 64.8 41.9 67.7 19.7 26.7 23.9 115.9 Bembradium holotype B. magnoculum (mm) Measurements relative to SL and HL for three species of and HL Measurements relative to SL Body depth at first dorsal fin Body depth at second dorsal fin Body width Head length Head depth Head width Snout length of orbit Snout width at anterior margin of anterior nostril Snout width at anterior margin Eye diameter Orbital diameter Post orbital length Interorbital width length Upper-jaw Preanus length Predorsal-fin length Preanal-fin length Length of first dorsal-fin base Length of second dorsal-fin base Length of anal-fin base Standard length Measurements (% SL) Table 2. Table 16

Phuket mar. biol. Cent. Res. Bull. others* 5.8–7.4 7.1–9.7 11.9–16.0 12.8–17.7 19.2–25.8 14.3–18.0 58.8–69.5 27.4–38.6 47.6–60.5 16.6–21.9 B. roseum 7.5 9.1 16.3 14.1 73.6 35.6 22.9 No data No data No data holotype others 7.2–7.5 8.9–10.0 15.0–17.5 14.0–15.8 21.6–22.8 18.7–19.4 48.9–58.0 30.9–35.0 47.9–51.4 19.7–21.5 B. furici 7.6 17.8 14.0 23.5 18.1 31.9 53.5 No data No data No data holotype . (2009). et al sp. nov. 7.5 7.5 13.8 12.1 21.5 16.7 57.1 25.8 45.9 16.0 holotype B. magnoculum partly from Kanayama (1982), Yatou (1985) and Ho Yatou partly from Kanayama (1982), B. roseum (con't) Caudal peduncle depth Length of upper caudal peduncle Length of lower caudal peduncle Pectoral-fin length Pelvic-fin length Pelvic-fin spine length Length of second dorsal-fin base Length of lower caudal peduncle Pectoral-fin length Pelvic-fin spine length Measurements (% HL) Table 2. Table *Data for 17

Description of a new species of Bembradium

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Manuscript received: 26 June 2018 Accepted: 11 October 2018