Species Diversity 23: 95–114 25 May 2018 DOI: 10.12782/specdiv.23.95

Redescription and Geographic Variations of antennata and First Record of Pterois paucispinula from French Polynesia (: )

Mizuki Matsunuma1,3 and Hiroyuki Motomura2 1 Department of Environmental Management, Faculty of Agriculture, Kindai University, 3327-204 Nakamachi, Nara 631-8505, Japan E-mail: [email protected] 2 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan 3 Corresponding author (Received 13 November 2017; Accepted 8 March 2018)

Pterois antennata (Bloch, 1787) (Scorpaenidae: Pteroinae) was redescribed on the basis of 243 specimens from throughout the Indo-Pacific region, including the lectotype ofScorpaena antennata and holotype of Pterois ellioti Meek, 1897, a junior synonym of the former. The species is diagnosed by the following combination of characters: dorsal-fin rays usually XII, 11; pectoral-fin rays usually 17; more than ten narrow (but of irregular width) transverse lateral bands on body, including caudal peduncle; pectoral-fin membranes with 3–47 blotches; posterior portion of upper pectoral-fin rays (free of membrane) without bands. Geographic variation in the number of pectoral-fin blotches and ontogenetic morphological changes within P. antennata are also discussed in detail. The first record ofPterois paucispinula Matsunuma and Motomura, 2014 from French Polynesia reported. Key Words: Pteropterus, synonymy, distribution, diagnosis, morphology.

Introduction Materials and Methods

Pterois antennata, the second oldest nominal species in Measurements generally followed Motomura (2004a, b), the genus Pterois Oken, 1817 (Scorpaenidae: Pteroinae), head width and maxillary depth following Motomura et al. was originally described by Bloch (1787) as Scorpaena an- (2005b, 2006a) and Motomura et al. (2006b), respectively. tennata on the basis of two specimens from Ambon, Indo- The following measurements were also made: body depth nesia. The species has since been recorded from through- at anal-fin origin—taken at level of first anal-fin spine base; out the Indo-Pacific region, from the east coast of Africa to 1st–9th pectoral-fin ray length—distance from uppermost French Polynesia, and from northern Australia to southern pectoral-fin ray origin to each ray tip. Supraocular tentacle Japan (Smith 1957b; Poss 1999; Matsunuma and Motomura length was measured between the tentacle base and ex- 2011). Pterois ellioti, described by Meek (1897) on the basis treme tentacle tip. Counts generally followed Motomura et of a single specimen from Berbera, Somalia, western Indian al. (2005a–c) and Motomura and Johnson (2006), counts Ocean, has been recently treated as a junior synonym of P. of pre-dorsal-fin scale rows and scale rows above the later- antennata [e.g., Eschmeyer (1998), Matsunuma and Moto- al line following Motomura et al. (2006b) and Matsunuma mura (2014) and Eschmeyer et al. (2017)], but without de- and Motomura (2013b), respectively. Oblique, horizon- tailed discussion; nor has the validity of the former been de- tal and vertical cheek scale rows followed Matsunuma and termined following direct comparison of the primary types Motomura (2013a). The last two soft rays of the dorsal and of the two nominal species. anal fins were counted as single rays, each pair being associ- During a revisionary study of Pterois, specimens of P. an - ated with a single pterygiophore. Counts and measurements tennata from the central-south Pacific Ocean were found to were made on the left side whenever possible, except for have generally more numerous dark pectoral-fin blotches, pectoral-fin rays (counted on both sides). Meristic data only compared with those from other localities, such being re- were taken from specimens marked by an asterisk in the garded as intraspecific geographic variation. In addition, a lists of specimens examined. In the latter, sampling method direct comparison of the primary types of S. antennata and and collector data for the specimens were omitted. Standard P. el li oti confirmed their synonymy, as proposed in previous length is abbreviated as SL. studies. A redescription of P. antennata, including a discus- Head spine terminology shown in Fig. 1 followed Ran- sion of geographic variation is provided. A single specimen dall and Eschmeyer (2001: fig. 1), Motomura (2004b: fig. 1), of Pterois paucispinula Matsunuma and Motomura, 2014 and Matsunuma and Motomura (2013b: fig. 1). Supplemen- from French Polynesia represents the easternmost record of tal preopercular spine and lateral lacrimal spine terminol- that species in the Pacific Ocean. ogy followed Eschmeyer (1965) and Motomura and Senou

© 2018 The Japanese Society of Systematic Zoology 96 M. Matsunuma and H. Motomura

(2008: fig. 2), respectively. In the redescription of P. anten- Poss 1999: 2329, unnumbered fig. (Indo-Pacific); Ran- nata, data for the lectotype are given first, followed by other dall 1999: 10 (Pitcairn Islands); Randall and Earle 2000: specimen data, including the holotype of P. el li oti (if differ- 10 (Marquesas Islands, French Polynesia); Hutchins 2001: ent) in parentheses. Data on lengths of dorsal-fin spines for 27 (Western Australia); Terashima et al. 2001: 26, unnum- specimens abnormally possessing 13 dorsal-fin spines are bered fig. (Mauritius); Nakabo 2002: 569, unnumbered excluded from the description and Table 2. Institutional fig. (Japan); Allen and Adrim 2003: 29 (Indonesia); Allen codes follow Sabaj (2016), with the following addition: Na- et al. 2003: 370, unnumbered fig. (Indo-Pacific); Hutchins tional Museum of Nature and Science (NSMT), Tsukuba, 2003: 465 (Dampier Archipelago, Australia); Hutchins Japan. A distributional map was prepared using Quantum 2004: 364 (Dampier Archipelago, Australia); Kimura et al. GIS 2.2 (Quantum GIS Development Team 2014), with data 2003: 39, unnumbered fig. (Sulawesi, Indonesia); Randall from Natural Earth. et al. 2003: 10 (Tonga); Rao 2003: 129, fig. 204 (Andaman For molecular analyses, partial cytochrome c oxidase and Nicobar islands); Heemstra and Heemstra 2004: 144, subunit 1 (COI) sequence data for Pterois spp. and Scor- unnumbered fig. (southern Africa); Allen et al. 2006: 879 paenodes parvipinnis (Garrett, 1864) (as an out group) were (Australia); Senou et al. 2006: 71 (Ie-jima island, Oki- taken from DDBJ/EMBL/GenBank which includes sequenc- nawa, Japan); Satapoomin 2007: 46, unnumbered fig. es analyzed by Hubert et al. (2012). Identifications of the (Andaman Sea); Shao et al. 2008: 246 (southern Taiwan); following sequences were determined by our examination Allen and Erdmann 2009: 596 (West Papua, Indonesia); of voucher specimens: P. antennata: DDBJ/EMBL/GenBank Fricke et al. 2009: 39 (Réunion); Motomura 2009: 65, un- accession number: JQ432077 (voucher specimen: USNM numbered fig. (Andaman Sea); Hirata 2010: 34, right un- 392525) and KU943359 (ASIZP 72229); P. paucispinula: numbered figs. (in part; Ehime, Japan); Motomura et al. JQ432074 (USNM 392530). Identification of the sequence, 2010: 89 (Yaku-shima island, Kagoshima, Japan); Mundy JQ432080 (P. radi ata ), was determined from a photograph et al. 2010: 25 (Howland, Palmya and Kingman islands); of the voucher specimen (USNM 392519). Sequence diver- Motomura and Aizawa 2011: 466, fig. 2E (Yaku-shima gences (uncorrected p-distance) among samples were cal- island, Kagoshima, Japan); Allen and Erdmann 2012: culated by MEGA 7 (Kumar et al. 2016), and a dendrogram 220, unnumbered fig. (East Indies); Larson et al. 2013: was constructed by the neighbor-joining algorithm. 81 (Northern Territory, Australia); Motomura 2013: 43, unnumbered fig. (Iou-jima and Take-shima islands, Pterois antennata (Bloch, 1787) Kago­shima, Japan); Nakabo and Kai 2013: 688, unnum- [English name: Spotfin Lionfish; standard Japanese name: bered fig. (Japan); Fricke et al. 2014: 51 (Madang, Papua Nettai-minokasago] New Guinea); Delrieu-Trottin et al. 2015: 5 (Marquesas (Figs 1–10; Tables 1, 2) Islands, French Polynesia); Bacchet et al. 2016: 179, un- numbered fig. (French Polynesia); Koeda et al. 2016: 20, Scorpaena antennata Bloch 1787: 21, pl. 185 (type locality: fig. 85 (Yonaguni-jima island, Okinawa, Japan); Motomu- Ambon, Indonesia). ra and Harazaki 2017: 23 (Yaku-shima island, Kagoshima, Pterois antennata: Cuvier and Valenciennes 1829: 265 Japan); Siu et al. 2017: 259 (French Polynesia). (Waigeo, Indonesia); Günther 1860: 124 (BMNH cata- Pseudomonopterus antennatus: Bleeker 1863: 234 (Ternate, log); Seale 1906: 83 (French Polynesia); Weber 1913: 497 Indonesia). (Indonesia); Fowler and Bean 1927: 15 (Tahiti, French Pseudomonopterus (Pterois) antennatus: Bleeker 1876: 48 Polynesia); Herre 1936: 264 (Solomon Islands); Matsub- (Indonesia); Bleeker 1875–1878: pl. 413, fig. 5 (atlas). ara 1943: 345 (Japan); Smith 1957a: 397 (Aldabra Atoll, Pterois ellioti Meek 1897: 169, pl. 40 (type locality: Berbera, Seychelles); Beaufort and Briggs 1962: 45 (Indo-Pacific); Somalia). Schultz et al. 1966: 23, pl. 125B, fig. 138j (Bikini Atoll Pteropterus antennata: Smith 1957b: 79, pl. 5, fig. F (Indo- and Guam); Masuda et al. 1975: 339, pl. 144-C (in part; West Pacific); Whitley 1964: 56 (Australia); Smith and Japan); Chen 1981: 48, fig. 12 (Taiwan); Gloerfelt-Tarp Smith 1965: 56, pls 49F, 97B (Seychelles); Mandrytsa and Kailola 1984: 113, unnumbered fig. (southern Indo- 2001: 16 (phylogenetic material). nesia); Shen 1984b: 193, unnumbered fig. (Taiwan); Shi- (not of Cuvier and Valenciennes): Shen mizu 1984: 302, pl. 282-C (in part; Japan); Eschmeyer 1984a: 32, pl. 32, fig. 264-24c (in part; Taiwan). 1986: 466, pl. 25, fig. 149.7 (western Indian Ocean); Allen Pterois cf. antennata: Motomura and Struthers 2015: 1095, and Swainston 1988: 48, unnumbered fig. (northwestern fig. 156.5 (Kermadec Islands, New Zealand). Australia); Winterbottom et al. 1989: 20, fig. 103 (Chagos Archipelago); Randall et al. 1990: 12 (Rapa, French Poly- Lectotype of S. antennata. ZMB 796, 117.6 mm SL, nesia); Allen and Smith-Vaniz 1994: 8 (Cocos Keeling Ambon, Indonesia. Islands); Masuda and Kobayashi 1994: 77, unnumbered Other type specimen. FMNH 484, holotype of Pterois figs. (Japan); Shao and Chen 1994: 237, pl. 56-4 (Taiwan); ellioti, 48.8 mm SL, Berbera, Somalia, D. G. Elliot. Kuiter 1996: 90, unnumbered fig. (Australia); Randall et Non-type specimens examined. 241 specimens, 12.8– al. 1997b: 16 (Ogasawara Islands, Japan); Kuiter 1998: 153.9 mm SL. JAPAN: Boso Peninsula: KPM-NI 22989, 55, unnumbered figs. (Maldives); Fricke 1999: 157 (Mas- 23.5 mm SL, Tateyama Bay, Chiba Prefecture (Pref.), 7 m, 9 carene Islands); Meyers 1999: 101, pl. 28F (Micronesia); September 2007. Izu Peninsula: KPM-NI 26918, 36.1 mm Redescription of Pterois antennata 97

SL, Izu Oceanic Park, Shizuoka Pref., 16 December 1989. Hama Fishing Port, Nakagusuku, Okinawa-jima island, Izu Islands: NSMT-P 30809, 58.5 mm SL, Miyake-jima is- 26°09′06″N, 127°28′25″E, 10 July 2015; URM-P 1324, land, 34°07′N, 129°31′E, 19 August 1977. Ogasawara Is- 104.8 mm SL, Okinawa-jima island; URM-P 18654, lands: HUMZ 53491, 118.1 mm SL, Miyanohama, Chichi- 116.4 mm SL, off Zanpa to Yomitan, Okinawa-jima island, jima island, 7–8 m, 18 June 1976. Suruga Bay: MSM-02-144, December 1986. Yaeyama Islands: BPBM 8712, 2 speci- 42.1 mm SL, Mera Port, east of Suruga Bay, Shizuoka Pref., mens, 65.5–82.5 mm SL, reef about 0.5 miles off Ishigaki, 28 October 2001. Kii Peninsula: FAKU 99045, 77.0 mm SL, Ishigaki-jima island, 22 May 1968; *NSMT-P 80768, FAKU 99086, 84.5 mm SL, Shirahama, Waka­yama Pref., 30 72.8 mm SL, Amitori Bay, Iriomote-jima island, 9 June 1982; January to 20 February 2011. Tosa Bay: BSKU 79021, URM-P 36099, 77.3 mm SL, off Uehara Beach, Iriomote- 102.2 mm SL, Tosa-shimizu, Kochi Pref., 15 December 2005; jima island, 6 August 1996. TAIWAN: ASIZP 57316, BSKU 86405, 86.1 mm SL, off Saga, Kochi Pref., 31 March 95.3 mm SL, Wanlitong, Pingtung, 3 April 1979; ASIZP 1999; BSKU 100096, 38.0 mm SL, Okino-shima island, 59074, 66.0 mm SL, Liuchiu, Pingtung, 11 December 1979; Kochi Pref., 24 September 2009; BSKU 103166, 111.1 mm ASIZP 59571, 108.3 mm SL, Dong-gang, Yilan, 10 June SL, no further locality; NSMT-P 90778, 70.0 mm SL, north 1979; ASIZP 59601, 94.0 mm SL, Liuchiu, Pingtung, 25 July of Mo-shima Port, Okino-shima island, Kochi Pref., 3 m, 22 1986; ASIZP 72229, 136.4 mm SL, Magong, Penghu, 20 May July 2008. Hyuga-nada Sea: MUFS 16455, 66.0 mm SL, off 2008; KAUM–I. 43830, 115.5 mm SL, KAUM–I. 43831, Kushima, Miyazaki Pref., 21 October 1998. Satsuma Penin- 117.2 mm SL, KAUM–I. 43832, 124.0 mm SL, KAUM–I. sula: KAUM–I. 38746, 116.5 mm SL, off Kushi, Bonotsu, 43833, 118.8 mm SL, Liuchiu, Pingtung; NMMB-P 6893, Minami-satsuma, Kagoshima Pref., 31°18′37″N, 94.1 mm SL, Shao-liuchiu Island, Pingtung, 22 October 130°13′19″E, 2–25 m, 14 June 2011. Osumi Islands: 2003; NMMB-P 13172, 91.2 mm SL, Dong-kang, Pingtung, KAUM–I. 29634, 72.8 mm SL, KAUM–I. 29635, 73.5 mm 1 March 1979; NMMB-P 7033, 19.2 mm SL, Shao-liuchiu Is- SL, off west coast of Iou-jima island, 30°47′04″N, land, Pingtung, 12 December 2003; ZMB 16148, 31.3 mm 130°15′42″E, 5–20 m, 28 May 2010; KAUM–I. 29745, SL, Taiwan (no further locality). PHILLIPINES: AMS I. 100.6 mm SL, KAUM–I. 29746, 153.9 mm SL, off south coast 40144-010, 5 specimens, 14.0–73.2 mm SL, northeast side of of Iou-jima island, 30°46′32″N, 130°16′43″E, 10–60 m, 26 Maestro de Campo Islands, 12°56′77″N, 121°43′53″E, May 2010; KAUM–I. 37938, 65.8 mm SL, Iou-jima Port, 25–28 m, 29 May 2000; NSMT-P 34677, 112.3 mm SL, north Iou-jima island, 30°46′43″N, 130°16′43″E, 2 m, 18 May east coast of Makesi Island, Honda Bay, Palawan, 10 m, 9 2011; KAUM–I. 62198, 18.8 mm SL, off Urata, Nishi- November 1988; NTOU 20080414-101, 106.4 mm SL, noomote, Tanega-shima island, 30°49′36″N, 131°02′11″N, NTOU 20080414-116, 68.1 mm SL, Cebu, 14 April 2008; 10–15 m, 13 June 2014. Tokara Islands: BSKU 51076, SMF 10025, 66.2 mm SL, Philippines (no further locality), 58.5 mm SL, Kuchino-shima island, May 1980; KAUM–I. 1966; USNM 99021, 112.8 mm SL, Lode Bay, Destacado Is- 77955, 74.9 mm SL, west side of Nishinohama Fishing Port, land, 5 m, 13 March 1909. PAL AU: BPBM 9941, 36.7 mm Kuchino-shima island, 29°59′30″N, 129°54′42″E, 5–10 m, 28 SL, Angulpelu Reef, Palau Islands, 110–170 ft (34–52 m), 22 August 2015. Amami Islands: BSKU 6942, 109.8 mm SL, April 1970; FMNH 118622, 88.8 mm SL, central east coast of Ankyaba, Amami-oshima island, 2 November 1956; Fana Island, Sonsoral, 05°21′17″N, 132°13′57″E, 12 Septem- KAUM–I. 57749, 52.9 mm SL, off Setouchi, Amami-oshima ber 2008; FMNH 118623, 112.0 mm SL, southernmost tip of island, 28°07′50″N, 129°21′07″E, 3–15 m, 13 December Fringing Reef of Tobi Island, Hatohobei, 02°59′50″N, 2013; KPM-NI 21471, 93.3 mm SL, off Udonosu Beach, 131°07′23″E, 15 September 2008; FMNH 118624, 80.0 mm Yoron-jima island, 1.5–4 m, 22 March 2007; YCM 36181, SL, north east tip of Helen Reef, Hatohobei, 02°58′01″N, 39.2 mm SL, off Sukomo, Setouchi, Amami-oshima island, 131°50′06″E, 21 September 2008. MARIANA ISLANDS: 21 August 1995; YCM 36552, 62.4 mm SL, off Sakinome AMS I.39371-001, 87.5 mm SL, Merizo Reef, 26 August Beach, Setouchi, Amami-oshima island, 26 August 1995. 1967; BPBM 258, 79.5 mm SL, 12 July 1900; BPBM 4389, Okinawa Islands: KAUM–I. 6436, 133.6 mm SL, KAUM–I. 133.1 mm SL, 4 July 1923; SAIAB 86225, 85.0 mm SL, Lau 6438, 120.1 mm SL, KAUM–I. 6586, 132.9 mm SL, Nakagu- Lau Beach at Standing Pipe, south east side of Saipan, suku, Okinawa-jima island, 26°15′29″N, 127°47′38″E, 13 15°09′48″N, 145°45′46″E, 20–27 m, 26 May 2003. July 2007; KAUM–I. 32021, 77.7 mm SL, *KAUM–I. 32022, CAROLINE ISLANDS: USNM 223578, 77.6 mm SL, south 99.1 mm SL, off Odo Beach, Itoman, Okinawa-jima island, west side of Pohnpei, Caroline Islands, 06°52′N, 158°06′E, 26°05′22″N, 127°42′24″E, 23 August 2010; KAUM–I. 43003, 0–120 ft (0–37 m), 15 September 1980. WAKE ISLAND: 130.3 mm SL, KAUM–I. 43004, 144.2 mm SL, Araha Beach, BPBM 15280, 3 specimens, 79.0–105.1 mm SL, lagoon near Nakagami, Okinawa-jima island, 26°18′25″N, 127°45′50″E, Peale Island, 10 June 1953. MARSHALL ISLANDS: BPBM 29 May 2011; KAUM–I. 48356, 116.8 mm SL, off Omna, 4395, 60.2 mm SL, Ebon Atoll, 1877; BPBM 6429, 8 speci- Okinawa-jima island, 26°30′18″N, 127°52′58″E, 1–3 m, 14 mens, 82.5–118.8 mm SL, BPBM 6432, 79.7 mm SL, Piirai August 2011; KAUM–I. 48416, 71.6 mm SL, KAUM–I. Island, Eniwetok Atoll, 4 August 1968; BPBM 6433, 48419, 74.2 mm SL, KAUM–I. 48420, 46.4 mm SL, off Yomi- 114.1 mm SL, north end of Piirai Island, Eniwetok Atoll, 14 tan, Okinawa-jima island, Okinawa Pref., 26°26′08″N, August 1968; BPBM 6434, 125.1 mm SL, south end of Japtan 127°42′45″E, 1–3 m, 21 October 2011; KAUM–I. 52186, Island, Eniwetok Atoll, 1 August 1968; *BPBM 6442, 69.4 mm SL, off Yomitan, Okinawa-jima island, 26°22′01″N, 96.0 mm SL, Japtan Island, Eniwetok Atoll, 29 July 1968; 127°44′15″E, 16 July 2012; KAUM–I. 78775, 102.3 mm SL, BPBM 8281, 50.7 mm SL, Bogen Island, Eniwetok Atoll, 98 M. Matsunuma and H. Motomura

25–70 ft (8–21 m), 3 December 1967; *FMNH 44140, mens, 81.4–82.4 mm SL, on barrier reef side on channel, 22.8 mm SL, Bikini Atoll, Eman Island, 19 July 1947. 1.5 km west of Tareu Pass, Moorea, 17°19′S, 149°53′W, 19 INDONESIA: KPM-NI 14031, 74.5 mm SL, north of Bali; February 1989; BPBM 5851, 98.6 mm SL, outer edge of in- NMV A74072, 73.4 mm SL, Banda Islands, 1874; NSMT-P shore reef at Papeere, Papetoai Bay, Tahiti, 40 ft (12 m), 14 62052, 72.3 mm SL, Lembeh Strait, Sulawesi, 15 m, 01°27′N, September 1967; BPBM 6114, 81.2 mm SL, Atimaouo, Papa- 125°14′E; NSMT-P 63740, 84.8 mm SL, Indonesia (no fur- ra, Tahiti, August 1967; BPBM 6927, 118.4 mm SL, outside ther locality); NSMT-P 71523, 49.0 mm SL, north east coast reef about 1/4 mile east of Teauraa Pass, Papara, Tahiti, 90 ft of Kodek Bay, Lombok, 15 m; NSMT-P 71676, 76.6 mm SL, (27 m), 26 February 1969; BPBM 7476, 3 specimens, 52.6– Gerupuk, Lombok, 5 m; NSMT-P 102043, 72.2 mm SL, 65.9 mm SL, Papara, Tahiti, 20 February 1969; BPBM 8922, Meno Island, Lombok; NSMT-P 102044, 74.4 mm SL, Meno 78.4 mm SL, Popote Bay, Papara, Tahiti, 130 ft (43 m), 6 Island, Lombok; SMF 4204, 89.9 mm SL, south east of Su- March 1969; FMNH 75884, 76.3 mm SL, Tahiti, 27 February lawesi, 1909–1910; SPMN-PI 41699, 59.1 mm SL, Indonesia 1962; MNHN 1984-279, 87.3 mm SL, off Temae, Moorea, (no further locality); ZIN 809, 2 specimens, 108.4– 17°28′59″S, 149°48′00″W, 2 m, September 1983; MNHN 110.1 mm SL, Ambon, 1858. PAPUA NEW GUINEA: NMV 2008-718, 44.8 mm SL, Moorea, 1 m, 15 March 2006; A 13698, 121.7 mm SL, Hood Bay, New Guinea Island, Au- MNHN 2008-861, 116.9 mm SL, Moorea, 35 m, 18 March gust 1882; OMNH 13808, 104.9 mm SL, Rabaul, New Brit- 2006; USNM 197734, 58.5 mm SL, off Tahiti Hotel Arabiri ain Island, September 1958; USNM 380294, 34.6 mm SL, Arie, 25 January 1963; USNM 392525, 95.4 mm SL, Paroa, east side of Jalun Island, Hermit Islands, 0–34 m, 2 Novem- Moorea, 17°36′23″S, 149°50′00″W, 15 March 2006. ber 1978; ZMB 15123, 2 specimens, 56.6–84.0 mm SL, New AUSTRAL ISLANS: AMNH 72813, 122.1 mm SL, north Guinea Island. SOLOMON ISLANDS: AMS I. 22128-085, 3 side of Tarohoi Island, Rapa Island, Tubuai Islands, 27°36′S, specimens, 41.1–103.1 mm SL, 3 km from Munda, Roviana 144°18′W, 15 April 1970; AMNH 72814, 4 specimens, 22.4– Lagoon, New Georgie Island, 08°18′S, 157°15′E, 1 Decem- 93.1 mm SL, Teahateke Point in Anatauri Bay, Rapa Island, ber 1980; AMS I. 39040-045, 57.5 mm SL, south of Neo Vil- Tubuai Islands, 27°36′S, 144°18′W, 12 April 1970; BPBM lage, west coast of Tomotu Island, Santa Cruz Islands, 11191, 143.4 mm SL, west side off Avera, Rurutu Island, 10°40′S, 165°48′E, 10–30 m, 29 September 1998. 140–175 ft (43–53 m), 28 February 1971; BPBM 11201, VANUATU: AMS I. 15847-004, 2 specimens, 56.2– 82.4 mm SL, west side off Avera, Rurutu Island, 150–190 ft 107.8 mm SL, Vanuatu (no further locality); MNHN 1987- (46–58 m), 28 February 1971; BPBM 11246, 63.1 mm SL, 1822, 89.1 mm SL, off Port Vila, 17°40′59″S, 167°49′59″E, Haurei Bay, Rapa Island, Tubuai Islands, 40–50 ft (12–15 m), October 1961; USNM 356588, 53.4 mm SL, Judy Reef, off 11 February 1971; BPBM 12864, 80.7 mm SL, Angairao Bay, northwest tip of Tongoa Island, Shepherd Islands, Rapa Island, Tubuai Islands, 30 January 1976; BPBM 12996, 16°52′30″S, 168°31′30″E, 75–108 ft (23–33 m), 9 June 1996. 2 specimens, 69.5–72.4 mm SL, south side of exposed reef at NEW CALEDONIA: USNM 324419, 60.2 mm SL, Bagaat entrance to Haurei Bay, Rapa Island, Tubuai Islands, 14 Feb- Islet, Ouvea Atoll, Loyalty Islands, 20°37′18″S, 166°16′08″E, ruary 1971; BPBM 13020, 138.5 mm SL, southwest side of 50–71 ft (15–22 m), 16 November 1991. WALLIS AND Karapoo Island, Rapa Island, Tubuai Islands, 60 ft (18 m), 16 FUTUNA: USNM 374803, 81.5 mm SL, Ile Uvea, Wallis Is- February 2017; USNM 379753, 2 specimens, 45.3–100.1 mm lands, 13°22′05″S, 176°10′45″W, 7–10 m, 14 November SL, north coast off Pt. Kauira, Rapa Island, Tubuai Islands, 2000. PHOENIX ISLANDS: BPBM 25487, 3 specimens, 27°34′04″S, 144°20′46″W, 2–6 m, 15 November 2002. 89.0–129.8 mm SL, Kanton Island, 10 February 1950. LINE TUAMOTU ARHCIPELAGO: CAS 66072, 67.1 mm SL, ISLANDS: BPBM 6967, 2 specimens, 60.4–83.3 mm SL, west side of Raroia, north of Ngaru Pass, Mataira Islet, west southeast of Sand Island, Palmyra Atoll, Line Islands, 3–12 ft side of Raroia Atoll, 16°01′S, 142°26′W, 0–6 ft (0–2 m), 17 (0.9–4 m), 12 November 1968; *BPBM 14258, 80.0 mm SL, July 1952; CAS 66073, 47.4 mm SL, channel between Nen- southwest end of Cook Islet at lagoon entrance, Kiritimati gonengo and Kukina islets, west side of Raroia Atoll, Atoll, Line Islands, 0–2 m, 24 March 1972; BPBM 15413, 16°00′S, 142°26′W, 0–1.5 ft (0–0.5 m), 5 July 1952. 26.0 mm SL, Palmyra, Sand Island, Line Islands, 20 June MARQUESAS ISLANDS: AMS I. 22015-018, 64.6 mm SL, 1962; BPBM 37599, 17.9 mm SL, Kiritimati Atoll, Line Is- Anaho Bay, Nuku Hiva, 8°55′S, 140°10′W, 1 August 1976; lands, 35–40 feet (11–12 m), September 1996; CAS 66059, 3 BPBM 4388, 69.0 mm SL, Marquesas Islands; BPBM 10415, specimens, 78.5–90.1 mm SL, inside reef on west side of Par- 73.2 mm SL, Haka Puuae, Taipi Vai, Nuku Hiva, 9 Septem- adise Island Jetty, Palmyra Atoll, Line Islands, 05°52′30″N, ber 1956; BPBM 11035, 2 specimens, 21.4–41.6 mm SL, 162°04′42″E, 0.5–10 ft (0.2–3 m), 16 August 1951. TONGA: south side of Vaeho Bay, off entrance to large cave, Ua Pou USNM 337707, 3 specimens, 78.1–106.9 mm SL, Luamoko Island, 0–30 feet (0–9 m), 29 April 1971; BPBM 11114, 2 Island, Vava’u Group, Tonga Islands, 18°40′55″S, specimens, 30.5–56.3 mm SL, point at north side of Hanauu 174°06′09″W, 70–101 ft (21–31 m), 15 November 1993. Bay, Fatu Hiva, 40–60 feet (12–18 m), 21 April 1971; USNM COOK ISLANDS: BPBM 5281, 67.3 mm SL, Raratonga Is- 340483, 78.7 mm SL, Taiohai, Nuku Hiva, 1 February 1957; land, 15 June 1929; BPBM 11152, 2 specimens, 36.0– USNM 399831, 81.2 mm SL, west coast of Mohotane Island, 80.6 mm SL, Rarotonga Island, 75–100 ft (23–30 m), 11 09°57′38″S, 138°50′17″W, 6–20 m, 14 October 2008. March 1971; FMNH 16311, 88.8 mm SL, FMNH 16312, PITCAIRN ISLANDS: BPBM 13261, 134.8 mm SL, off 87.7 mm SL, FMNH 16313, 49.2 mm SL, Aitutaki Island, Bounty Bay, Pitcairn Island, 100–130 ft (30–40 m), 26 De- 1930. SOCIETY ISLANDS: AMS I. 28946-012, 2 speci- cember 1970; BPBM 16627, 109.7 mm SL, north side of Redescription of Pterois antennata 99 atoll, Oeno Island, 70 ft (21 m), 19 December 1970; BPBM 360991, 2 specimens, 107.9–142.5 mm SL, south west of 16636, 90.7 mm SL, Pitcairn Island, 90–100 feet (27–30 m), West Island, just south of La Passe du Bois, Aldabra Atoll, 23 December 1970. CORAL SEA: AMS I. 4119, 51.6 mm SL, 9°23′54″S, 46°12′36″E, 0–3 m, 19 August 1967. Lord Howe Island, 1898; AMS I. 20756-044, 96.1 mm SL, MAURITIUS: CAS 48693, 2 specimens, 35.9–37.4 mm SL, Raine Island, Queensland, 1 February 1979; AMS I. 20965- Mascarene Islands, 7 March 1971; MNHN 6556, 64.6 mm 025, 88.8 mm SL, northeast of Dunk Islands, Queensland, SL, Mauritius (no further locality); SAIAB 2149, 2 speci- 17°56′S, 146°43′E, 10–20 m, 27 February 1979; AMS mens, 44.2–97.2 mm SL, 1 mile east of Beauchamp, 21°44′S, I.23847-027, 83.1 mm SL, south side of Yonge Reef, Lizard 57°46′E, 7 March 1971; SAIAB 52107, 2 specimens, 79.5– Island, Queensland, 14°35′S, 145°36′E, 8 m, 6 January 1975. 99.6 mm SL, off rocks end of Victoria Avenue, Albion, CHRISTMAS ISLAND: AMS I. 20441-005, 83.2 mm SL, 21°48′S, 57°24′E, 10 May 1995; USNM 349900, 88.1 mm SL, Christmas Island, 30 May 1978; FAKU 113358, 69.0 mm SL, west coast of Albion, Mascarene Islands, ca. 0–5 m, 10–15 Flyingfish Cave, Christmas Island, 13–26 m, 28 June 1986; May 1995; USNM 349899, 46.4 mm SL, 30 m north of Pass, *WAM-P 26092.016, 12.8 mm SL, Christmas Island, west coast of Flic En Flac, 20°16′S, 57°22′E, 4–10 m, 5 May 10°29′S, 105°40′E, 14 m, 25 May 1978. KERMADEC 1995; ZMB 13713, 119.6 mm SL, Mauritius (no further lo- ISLANDS: NMNZ P. 009501, 145.0 mm SL, Raoul Island, cality). RÉUNION: MNHN 6557, 96.3 mm SL, Réunion (no 29°13′S, 178°10′W, 1956. ANDAMAN SEA: PMBC 12055, further locality). DOUBTFUL RECORD: SMF 12887, 89.0 mm SL, Phuket; ROM 69890, 2 specimens, 103.5– 77.1 mm SL, Red Sea (no further locality), 1967. NO DATA: 111.6 mm SL, Ko Racha Yai Island, 07°28′15″N, 98°19′35″E, FAKU 102893, 123.9 mm SL; KPM-NI 6191, 76.2 mm SL, 18 November 1993. ANDAMAN AND NICOBAR aquarium trade. ISLANDS: SMF 28932, 122.9 mm SL, Nicobar Island, Anda- Diagnosis. A species of Pterois distinguished from man and Nicobar Islands, 11 August 1958. SRI LANKA: other species of the genus by the following combination of SMF 4327, 85.2 mm SL, Trincomalee, 21 May 1958. characters: dorsal-fin rays usually XII, 11 (rarely XII, 10, MALDIVES: *BPBM 32927, 70.7 mm SL, Maaniyafushi Is- XII, 12 or XIII, 10); pectoral-fin rays usually 17 (rarely 15, land, South Male Atoll, 25–30 m, 17 March 1988; FMNH 16 or 18); more than 10 narrow (but of irregular width) 75838, 2 specimens, 85.5–101.5 mm SL, south of Male Atoll, transverse lateral bands on body, including caudal pe- Dunidu Island, 19 April 1964; SMF 5386, 78.9 mm SL, Mal- duncle; pectoral-fin membranes with 3–47 (0 or 1 in larvae dives (no further locality), 1 February 1958; SMF 9770, <21.4 mm SL) blotches; posterior portion of upper pecto- 30.5 mm SL, Maldives (no further locality), 17 January 1958. ral-fin rays (free of membrane) without bands. OMAN: CAS 73270, 107.7 mm SL, Christian Cemetery Description. Morphometrics and selected meristics Cove, 2 m, 1 April 1988. YEMEN: SMF 4625, 128.2 mm SL, shown in Tables 1 and 2. Meristics shown in Table 1 not re- SMF 4626, 105.1 mm SL, Abd al Kuri Island, 10 December peated here. Pelvic-fin rays I, 5. Branchiostegal rays 7. 1957. SOMALIA: CAS 208379, 102.3 mm SL, Gulf of Aden, Body oblong, moderately compressed posteriorly; depth 11°21′06″N, 49°11′54″E, 27 October 1985. KENYA: LACM moderate, body depth at pelvic-fin origin 73% (55–93%) 31005-50, 101.8 mm SL, north side of Mombasa Island, of longest dorsal-fin length. Caudal peduncle moderately 4°02′S, 39°41′E, 10 September 1970; SAIAB 14400, 95.3 mm short, depth subequal to length. Head moderately large. SL, Mombasa, 5°55′S, 39°40′E, August 1957; SMF 9100, Several sensory pores on underside of each dentary; pore 67.5 mm SL, Mombasa, 24 January 1967; SMF 9932, behind symphysial knob of lower jaw in ventral view; pore 74.3 mm SL, Mombasa, 27 January 1969. TANZANIA: covered by lower-jaw lip on each side of symphysial knob in SAIAB 14398, 96.2 mm SL, Zanzibar Island, 7°50′S, 39°11′E, ventral view. Several small sensory pores on dorsal surface 21 September 1952. COMOROS: CAS 32512, 56.4 mm SL, of each interorbital canal at mid-orbit level. Postorbital sen- reef in front of Hotel Coelacanthe, Grande Comoros, sory canal short, slightly extending ventrally from sphenotic 25–30 m, 14 February 1975; SAIAB 34555, 2 specimens, spine base. Isolated postorbital sensory canal (shown in Fig. 61.0–79.7 mm SL, Grande Comoros, 12°11′S, 43°01′E, 18 1) usually absent (including lectotype) (rarely present in October 1986. MADAGASCAR: MNHN 1992-669, other specimens). 82.7 mm SL, off Toliara, 23°19′59″S, 43°30′00″E, 18 August Three (rarely four) short barbels on snout tip, their 1968; SAIAB 52777, 76.8 mm SL, off Berofia Island, Radama length subequal to anterior nasal tentacle length; numer- Islands, 14°58′S, 47°42′E, 30–32 m, 30 August 1995. ous minute cirri on ca. 5 (or 4) thin skin ridges on frontal SOUTH AFRICA: CAS 48671, 114.2 mm SL, Sodwana, Zu- surface of snout bulge. Short tentacle on posterior edge of luland, 1 April 1979; SAIAB 8835, 138.6 mm SL, Sodwana low, membranous tube associated with anterior nostril, its Bay, 28°29′S, 32°41′E, 3 April 1979; SAIAB 40493, tip slightly extending beyond posterior margin of posterior 124.4 mm SL, Aliwal Shoal, Cathedral, off Umkomaas, nostril when depressed posteriorly. Supraocular possessing Natal, 31°45′S, 30°49′E, 12 August 1992; SAIAB 46164, extremely long tentacle, relative length of tentacle [178.6% 114.3 mm SL, Anchors Reef, Aliwal Shoal, Natal, 31°45′S, (75.5–342.8) of orbit diameter] decreasing with growth; thin 30°49′E, 17 June 1994; SAIAB 46397, 101.1 mm SL, Cathe- skin ridges along lateral margins with 6 (or 0–5) short lat- dral, Aliwal Shoal, Natal, 31°45′S, 30°49′E, 20 June 1994; eral branches. A short simple tentacle on anteroventral por- SAIAB 62275, 137.0 mm SL, Protea Bank, 30°50′S, 30°29′E, tion of lacrimal (just anterior to ventrally positioned sen- 10 March 2000. SEYCHELLES: BPBM 20078, 100.2 mm SL, sory pore), its length subequal to that of snout tip barbel. A cast coast of Cerf Island, 6 m, 27 October 1973; USNM short elongate, leaf-like skin flap on ventralmost corner of 100 M. Matsunuma and H. Motomura

Table 1. Frequency distribution of selected meristic values from specimens of Pterois antennata (including primary types of Scorpaena antennata and Pterois ellioti).

DS: dorsal-fin spine base; LL: lateral line; SR: scale rows. H and L indicate holotype of P. el li oti and lectotype of P. antennata, respectively. Larvae <21.4 mm SL possessed II anal-fin spines; 2 specimens possessed 15/17 pectoral-fin rays. posterior lacrimal plate, its length subequal to anterior lac- premaxillary teeth bands slightly broader than width of each rimal tentacle [59% (54.0–88.3%) of orbital diameter], its tip band; both jaws with band of small, slender conical teeth; reaching posterior margin of maxilla when laid flat (small ca. 8 (or 9, 10) and 8 (or 9–11) teeth rows at widest portions specimens with relatively long flap, its tip extending far be- of upper and lower jaws, respectively; both teeth bands be- yond posterior margin of maxilla). Two small leaf-like skin coming narrow posteriorly; small conical teeth in ca. 8 (or 9, flaps on posterior margin of preopercle, their lengths about 10), rows forming blunt V-shaped patch on vomer; no pala- half of orbit diameter with tips extending slightly beyond tine teeth. posterior margin of interopercle when laid flat; upper flap Nasal spine with 1 (rarely 2 in large specimens) spi- situated between 3rd and 4th preopercular spines, lower flap nous point (Fig. 1). Preocular spine with 1 (sometimes 2 just below 4th preopercular spine base. A small skin flap an- in large specimens) spinous point directed upward. Supra- terodorsally on orbit surface, its length slightly greater than ocular with blunt spinous point covered by skin, at supra- diameter of posterior nasal pore. No other skin flaps on ocular tentacle base. Postocular with 12 (8 on right side of head or body. lectotype) (1–14) small spines directed laterally along pos- Body, including caudal peduncle, generally covered with terodorsal margin of orbit. Interorbital ridge weakly devel- well-developed ctenoid scales; thorax and abdomen cov- oped, diverging and reduced posteriorly, connected to cor- ered with weakly ctenoid or cycloid scales. Dorsal- and onal spine base. Coronal spine with 1 (or 2) spinous point anal-fin bases with 1 or 2 rows of weakly ctenoid or cycloid directed posteriorly (dorsal view). Tympanic spine with basal scales, of size about one-fifth to one-third of that of 1 (or 2; 0 in small non-type specimens <30 mm SL) spi- body scales. Pectoral-fin base largely covered with small nous point directed almost posteriorly; origin of tympanic cycloid scales. Caudal-fin base withca . 3–5 rows of small spine slightly posterior to that of coronal spine. No spines cycloid basal scales. Head generally covered with ctenoid between coronal and tympanic spines (1 or 2 small spines scales; snout, both jaws, maxilla, mandible and preocular rarely present in non-type specimens). Occipital region flat; bone without scales. Orbital surface without scales. Dorsal anterior margin of occipital region sloped transversely from surfaces of supraocular and postocular bones covered with between origin of coronal spines, slightly curved posteri- poorly developed ctenoid scales, most with a single spinule. orly (dorsal view); anterior border of occipital region rarely Cheek region entirely covered with exposed ctenoid scales, with spines in non-type specimens. Parietal spine with 2 suborbital scales not reaching a vertical through mid-orbit (1–3) spinous points on relatively long ridge, diverging pos- (rarely extending anteriorly beyond mid-orbit level in large teriorly. Nuchal spine with 2 (or 1) spinous points, its base non-type specimens); suborbital pit without scales (rarely a completely conjoined with that of parietal spine. No spinous few ctenoid scales present). points on parietal-nuchal ridge in small non-type specimens Dorsal profile of snout relatively steep, forming angle <30 mm SL. Coronal and parietal spines well separated, of ca. 40° to horizontal axis of head and body. Mandible small spine rarely present between them. Sphenotic with 5 smooth without ridges. Mouth moderately large, slightly (6 on right side of lectotype) (0–8) small spines, surround- oblique, forming angle of ca. 30° to horizontal axis of head ing short sensory canal. Pterotic spine with 3 (0–6) spinous and body; upper edge of posterior maxilla swollen laterally, points on relatively short ridge. Lower posttemporal spine forming low ridge; posterior margin of maxilla just reach- with 0 (2 in right side of lectotype) (or 1–3) spinous points ing a vertical through midorbit. Symphyseal gap separating on relatively short ridge. Cleithrum with upper short and Redescription of Pterois antennata 101

Table 2. Morphometric values, expressed as percentages of standard length (orbit diameter for supraocular tentacle length), recorded from specimens of Pterois antennata, including primary types of Scorpaena antennata and Pterois ellioti, and French Polynesian specimen of P. paucispinula.

P. antennata P. paucispinula Lectotype of S. antennata Holotype of P. el li oti Non-type specimens USNM 392530 ZMB 796 FMNH 484 (n=241) (n=1) SL (mm) 117.6 48.4 12.8–153.9 95.9 Body depth (% SL) 37.6 35.5 31.3–43.6 (38.5) 36.0 Body depth at anal-fin origin 30.6 30.5 23.6–34.4 (30.2) 29.5 Body width 24.7 24.0 16.8–29.4 (24.6) 24.3 Head length 37.9 39.5 38.2–44.8 (41.4) 40.3 Head width 13.4 13.3 12.3–16.1 (13.9) 12.9 Snout length 12.2 11.7 10.8–14.4 (12.4) 11.4 Orbit diameter (OD) 10.7 12.3 10.4–15.9 (12.8) 13.2 Interorbital width at mid-orbit 7.9 7.6 6.8–9.6 (8.2) 7.4 Interorbital width at preocular spine base 6.1 8.4 5.2–8.9 (7.0) 6.8 Upper-jaw length 17.6 17.2 17.0–20.6 (18.6) 19.9 Maxilla depth 6.9 6.6 6.1–8.6 (7.2) 7.9 Postorbital length 18.4 16.8 16.2–20.8 (18.5) 16.6 Suborbital depth 1.8 0.8 0.3–2.2 (1.0) 1.3 Pre-dorsal-fin length 34.3 33.4 31.4–39.2 (35.6) 36.3 Pre-anal-fin length 71.7 71.9 68.9–77.0 (72.6) 75.1 Pre-pelvic-fin length 37.1 40.4 35.9–43.9 (39.6) 36.7 1st dorsal-fin spine length 19.0 — 13.2–25.1 (18.8) 14.3 2nd dorsal-fin spine length 31.0 — 16.3–40.3 (29.8) 22.7 3rd dorsal-fin spine length 34.5 — 24.4–48.4 (34.5) 28.6 4th dorsal-fin spine length 38.5 — 28.2–50.3 (39.3) 39.0 5th dorsal-fin spine length 47.7 38.1 30.6–61.5 (46.6) 41.2 6th dorsal-fin spine length 51.0 — 34.0–65.4 (51.1) 45.4 7th dorsal-fin spine length 51.7 40.8 33.8–67.4 (54.0) 46.8 8th dorsal-fin spine length 51.7 — 34.9–68.7 (54.1) 47.8 9th dorsal-fin spine length 49.7 — 30.2–68.1 (51.7) 47.3 10th dorsal-fin spine length 36.1 31.4 12.6–57.6 (35.2) 43.6 11th dorsal-fin spine length 12.1 — 10.0–25.5 (14.2) 29.2 12th dorsal-fin spine length 13.3 — 11.2–19.3 (14.3) 14.0 13th dorsal-fin spine length — — — 15.0 Longest dorsal-fin spine length 51.7 — 35.4–68.7 (54.5) 47.8 1st dorsal-fin soft ray length 19.1 22.1 16.5–28.4 (21.9) 20.4 Longest dorsal-fin soft ray length 29.4 (6th) 31.6 (5th) 26.1–35.9 (31.2) 27.3 1st anal-fin spine length 7.6 8.6 6.0–11.0 (8.5) 6.9 2nd anal-fin spine length 14.5 17.8 13.0–20.4 (16.0) 13.9 3rd anal-fin spine length 17.8 18.9 15.4–23.0 (19.2) 17.7 1st anal-fin soft ray length 27.3 30.7 24.1–37.3 (30.4) 27.4 Longest anal-fin soft ray length 29.8 (3rd) — 27.0–39.5 (33.4) 28.4 1st pectoral-fin ray length 91.8 — 63.1–129.3 (93.7) 84.8 2nd pectoral-fin ray length 103.1 — 71.4–140.2 (101.0) 90.8 3rd pectoral-fin ray length 100.3 — 72.2–138.9 (99.0) 89.6 4th pectoral-fin ray length 94.6 80.5 66.4–135.3 (99.3) 88.7 5th pectoral-fin ray length 89.5 94.9 67.9–129.8 (97.7) 83.7 6th pectoral-fin ray length 95.7 93.2 66.8–117.9 (95.2) 81.9 7th pectoral-fin ray length 92.2 88.5 61.1–113.2 (89.8) 79.4 8th pectoral-fin ray length 82.7 84.2 53.9–101.8 (82.2) 74.7 9th pectoral-fin ray length 72.8 68.4 51.9–93.7 (72.8) 70.7 Longest pectoral-fin length 103.1 (2nd) 94.9 (5th) 72.3–140.2 (102.3) 90.8 Pelvic-fin spine length 18.9 19.3 16.2–24.7 (20.5) 19.5 Longest pelvic-fin soft ray length 43.0 (3rd) 49.6 (3rd) 38.9–57.9 (48.0) 43.0 Caudal-fin length 32.7 40.8 31.3–51.3 (38.8) 34.9 Caudal-peduncle length 15.3 16.0 14.2–18.4 (16.5) 15.2 Caudal-peduncle depth 10.9 11.9 9.2–12.8 (11.3) 10.8 Supraocular tentacle length (% OD) 178.6 225.0 75.5–342.8 (200.2) 166.9 Means of all specimens given in parentheses for P. antennata; values of dorsal-fin spine length for P. antennata specimens with 13 dorsal-fin spines excluded. 102 M. Matsunuma and H. Motomura

Fig. 1. Lateral (A) and dorsal (B) views of head and supraocular tentacle (C) of Pterois antennata (USNM 340483, 78.7 mm SL). Shaded areas indicate skin flaps. 1, nasal spine; 2, preocular spine; 3, supraocular spine; 4, postocular spine; 5, coronal spine; 6, tympanic spine; 7, parietal spine; 8, nuchal spine; 9, sphenotic spine; 10, pterotic spine; 11, lower posttemporal spine; 12, supracleithral spine; 13, preopercular spine; 14, suborbital ridge/spine; 15, lateral lacrimal ridge/spine; 16, posterior lacrimal spine; 17, postorbital spine/exposed sensory canal. Scale bars 5 mm. lower long ridges, possessing 1 (0–2) and 1 (0–7) spinous tinct (covered with skin); first (uppermost) spine directed points, respectively. Small specimens <30 mm SL lacking posterodorsally; second spine directed posteroventrally; sphenotic, pterotic and cleithrum spines. third spine plate-like, with no (1 or 2) spinous points; no Lateral lacrimal ridge with 2 (3 in right side of lectotype) supplemental spine at each preopercular spine. Opercular (0–6) spinous points; other ridges on lacrimal moderately spine absent. spinous (spines absent in small specimens). Suborbital ridge Origin of 1st dorsal-fin spine above origin of lower post- strongly spinous with numerous lower supplemental spines, temporal spine base; bases of 1st and 2nd dorsal-fin spines totaling 10 (0–37) spines (small specimens <30 mm SL closer than those of subsequent adjacent spines; 8th (6th lacking spine on suborbital ridge; becoming spinous with to 9th; usually 8th) spine longest; 11th (penultimate) [or growth); suborbital pit without spines. Anterior lacrimal 12th (last)] spine shortest; length of penultimate spine 36% spine absent. Posterior lacrimal spine broad, plate-like, with (24–100%) and 91% (71–185%) of that of 10th (antepenul- 2 (1–9) spinous points on distal margin, all spinous points timate) and 12th spines, respectively; membrane of spinous exposed, except for point at posterior lacrimal tentacle base. portion of dorsal fin strongly incised. Dorsal-fin soft rays all Preopercle with 5 spines, lowermost 2 spines minute, indis- branched; 6th (3rd to 7th; usually 5th) ray longest, but dis- Redescription of Pterois antennata 103 tinctly shorter than longest dorsal-fin spine; posteriormost 52% (39–65%) and 43% (30–58%) of that of 2nd and 3rd ray free from caudal peduncle. Anal-fin origin below last spines, respectively. Anal-fin soft rays all branched; 3rd (or dorsal-fin spine base; 3rd spine longest; length of 1st spine 2nd; usually 3rd) ray longest, its length moderately greater

Fig. 2. Fresh specimens of Pterois antennata from various localities at different growth stages. (A) KAUM–I. 62198, 18.8 mm SL, Japan; (B) KAUM–I. 29635, 73.5 mm SL, Japan; (C) KAUM–I. 32021, 77.7 mm SL, Japan; (D) USNM 399831, 81.2 mm SL, French Polynesia; (E) BPBM 5851, 98.6 mm SL, French Polynesia; (F) KAUM–I. 38746, 116.5 mm SL, Japan; (G) BPBM 6927, 118.4 mm SL, French Polynesia; (H) KAUM–I. 29746, 153.9 mm SL, Japan. Photos: J. T. Williams (D) and J. E. Randall (E, G). 104 M. Matsunuma and H. Motomura than that of longest dorsal-fin soft ray; posteriormost ray brown band with narrow white margin from supraocular free from caudal peduncle. Pectoral-fin rays extremely tentacle base, obliquely crossing eye and reaching posterior long, filamentous, all rays unbranched in all stages, poste- margin of preopercle. Two reddish-brown bands with nar- rior portion of upper rays free from membrane (about half row white margins saddling nape; anterior band short, just of longest ray associated with membrane); lower 6 (7 or 8) behind eye, at level between coronal spine and origin of rays weakly thickened; 2nd (2nd to 6th) ray longest, tip of parietal spine; posterior band just behind anterior band at longest ray extending beyond caudal-fin base when laid level of parietal ridge, reaching central posteroventral mar- flat [relative length of rays (% SL) decreasing with growth]. gin of opercle at level of upper origin of pectoral-fin base. Pelvic-fin spine base below 3rd dorsal-fin spine base; all soft Numerous variable width, reddish-brown bands on body rays branched; 3rd (2nd to 4th; usually 3rd) soft ray longest, (extending onto dorsal and anal fins) and caudal peduncle its tip extending far beyond a vertical through 1st anal-fin (their number increasing with growth, up to ca. 30 in speci- spine base, but not reaching that of posteriormost anal-fin mens >100 mm SL), those below mid dorsal-fin spinous soft ray base; 5th soft ray with membranous connection to portion usually broadest; 1 or 2 reddish bands on thorax. abdomen for ca. one-third of ray length. Soft rays of dorsal, Lateral bands on caudal peduncle narrow, slightly oblique. anal and pelvic fins all unbranched in small non-type speci- Ground color of dorsal-fin spines same as on body, with mens < ca. 30 mm SL. Caudal fin rounded, with 3 procur- 1–5 (5 on longest spine) reddish-brown bands. Membranes rent rays, 2 segmented unbranched rays and 5 segmented of soft-rayed portions of dorsal and anal fins, and caudal fin branched rays in both dorsal and ventral series; procurrent semi-translucent; numerous small reddish-brown to black rays well developed, exposed. spots on those fin rays, additional small white spots between Color when fresh. Based on photographs of fresh non- each dark spot. Pectoral-fin membrane pale red to dark type specimens (Fig. 2). Ground color of head and body brown, blackish marginally, with ca. 4 or 5 broad dark bands pink to pale red; whitish ventrally. Lips whitish. Supraocu- on basal portion; rays pale red to dark red, whitish poste- lar tentacle same as ground color of head, with 3–7 dark red riorly, posterior portions of upper rays (free of membrane) to black bands; flap on orbit surface dark red to black; other without bands; variable size dark blotches on membranes. skin flaps or tentacles on head pale red. A broad reddish- Pelvic-fin membrane dark red, grading to nearly black prox-

Fig. 3. Primary types identified as Pterois antennata. (A) lectotype of Scorpaena antennata, ZMB 796, 117.6 mm SL, Ambon, Indonesia; (B) holotype of Pterois ellioti, FMNH 484, 48.8 mm SL, Berbera, Somalia. Redescription of Pterois antennata 105 imally, with rays white or pale red. on head and body becoming brown to black (diffuse in Color of preserved specimens. Based on all examined long-term preserved or poor condition specimens) [lec- specimens (Figs 3, 4). Head and body creamy-white; bands totype with ca. 20 (mostly diffuse) (ca. 10–25 in non-type

Fig. 4. Preserved specimens of Pterois antennata from various localities at different growth stages. (A) USNM 356588, 53.4 mm SL, Vanu- atu; (B) USNM 337707 (1 of 3 specimens), 78.1 mm SL, Tonga (right side, reversed); (C) KPM-NI 21471, 93.3 mm SL, Japan; (D) USNM 99021, 112.8 mm SL, Philippines; (E) USNM 360991 (1 of 2 specimens), 142.5 mm SL, Seychelles; (F) BPBM 11114 (1 of 2), 56.3 mm SL, French Polynesia; (G) USNM 399831, 81.2 mm SL, French Polynesia (right side, reversed); (H) AMNH 72814 (1 of 4), 87.5 mm SL, French Polynesia; (I) BPBM 6927, 120.9 mm SL, French Polynesia; (J) NMNZ 9501, 145.0 mm SL, New Zealand (Kermadec Islands). Note: speci- mens from central-south Pacific Ocean (F–J) possess relatively higher numbers of pectoral-fin blotches compared with those from other localities (A–E). 106 M. Matsunuma and H. Motomura

Fig. 5. Distributional map of Pterois antennata based on examined specimens. Star, triangle and circles indicate localities of lectotype of Scorpaena antennata, holotype of Pterois ellioti, and other specimens, respectively. FP, French Polynesia; PT, Pitcairn Islands; CO, Cook Is- lands; KE, Kermadec Islands; LI, Line Islands; PH, Phoenix Islands. specimens) lateral bands on body]; supraocular tentacle pl. 97-1–2) from Wakayama Prefecture, Japan was identified with 6 (3–7) brown or black bands; ca. 63 (27–139) small here as P. paucispinula, having 13 relatively longer dorsal- spots on soft-rayed portion of dorsal fin, ca. 65 (22–89) fin spines. Pterois antennata figured by Masuda et al. (1975) on that of anal fin, ca. 52 (44–170) on caudal fin, number and Shimizu (1984) from Japan, Randall et al. (1997a) and of spots increasing with growth; ca. 9 (3–47) variable size Randall (2005) from Bali, and Hirata (2010: left unnum- brown to black blotches on pectoral fin. bered fig.) from Ehime Prefecture, Japan were also identi- Distribution. Pterois antennata is found in the In- cal with P. paucispinula, having XIII, 10 dorsal-fin rays, 18 do-Pacific region from the east coast of Africa eastward pectoral-fin rays and reddish bands on the latter. Jones and to French Polynesia and from northern Australia north Kumaran (1980) recorded P. antennata from the Laccadive to Japan (Fig. 5). One doubtful record has been reported Archipelago in the Indian Ocean, but their description and from the Red Sea, without a precise locality (SMF 12887). figure were indicative of P. mombasae. Although Meyers Although the Red Sea was included in the distribution of (1999) and Randall (2005) stated the number of dorsal-fin P. antennata by Fricke et al. (2014), no additional records spines as 13 for P. antennata, such was either a simple error have been reported from the Red Sea (Golani and Bogoro- or their description may have included P. paucispinula or dsky 2010; Matsunuma et al. 2016) and its presence there is P. mombasae. Dendrochirus zebra from Taiwan, figured by doubtful. Collection data from 42 lots indicate that this spe- Shen (1984a: pl. 32, fig. 264-24c), was identified here as P. cies occurs at depths of 0–60 m, with most records from less antennata. than 20 m. Comparisons. Smith (1957b) regarded Pteropterus as a valid genus, distinguishing it from Pterois on the basis of several external characters, including: body covered dorso- Discussion laterally with ctenoid scales (vs. all scales cycloid in Pterois); fewer than 65 scale rows in the longitudinal series (vs. more Although Smith (1957b) and Mandrytsa (2001) included than 65); suborbital ridge close to orbit (vs. widely sepa- the species in Pteropterus Swainson, 1839, that genus has rated); and pectoral-fin rays more than 15 (vs. less than 15). been generally regarded as a junior synonym of Pterois in According to Smith (1957b), the following seven nominal recent studies, including Eschmeyer (1986), Poss (1999) and species of Pterois, currently regarded as valid, were better Matsunuma and Motomura (2014, 2015). However, recent assigned to Pteropterus: P. antennata, Pterois brevipectora- molecular studies have indicated two distinct lineages with- lis (Mandrytsa, 2002), Pterois cincta Rüppell, 1838, Pterois in Pterois (Kochzius et al. 2003; Freshwater et al. 2009) and a mombasae (Smith, 1957), Pterois paucispinula Matsunuma phylogenetic revision of the genus is necessary to determine and Motomura, 2014, Pterois radiata (Cuvier in Cuvier and the validity of Pteropterus. Valenciennes, 1829) and Pterois sphex Jordan and Ever- Literature citations of P. antennata have sometimes con- mann, 1903. fused that species with a related congener, either P. mom - Among the above group of species, P. antennata is char- basae or P. paucispinula (see Matsunuma and Motomura acterized by having usually XII, 11 (rarely XII, 10, XII, 12 or 2014). However, P. antennata differs from the latter two spe- XIII, 10) dorsal-fin rays, compared with usually XIII, 10 in cies in having usually XII, 11 dorsal-fin rays (usually XIII, the others (except P. cincta and P. radi ata with XII, 11 dor- 10 in the latter) and the posterior portion of the pectoral- sal-fin rays) (Matsunuma and Motomura 2013b, 2014, 2015; fin rays (free of membrane) without bands (with bands). this study). Pterois antennata can be also distinguished from Pterois antennata figured by Ikeda and Nakabo (2015: 98, the others by having 15–18 (modally 17) pectoral-fin rays, Redescription of Pterois antennata 107 compared with 15 or 16 (16) in P. brevipectoralis; 15–17 P. el li oti given by Meek (1897), and the configuration of the (16) in P. cincta; 17–20 (18) in P. mombasae; 17–19 (18) in dorsal-, anal-, pectoral- and pelvic-fin rays, condition of the P. paucispinula; 15–17 (16) in P. radi ata ; and 16 in P. sphex head spines, body size, skin flaps and coloration, including (Eschmeyer and Randall 1975; Matsunuma and Motomura the number of bands on the supraocular tentacle, also in 2013b, 2014, 2015; this study). Although P. cincta and P. ra - agreement with the original description of P. el li oti . The cat- diata are similar to P. antennata in having usually XII, 11 alog of fishes collected during the Field Museum East Afri- dorsal-fin rays, they can be distinguished in coloration, P. can Expedition noted the locality for FMNH 484 as Berbera, antennata having usually more than ten narrow transverse Somalia, on the east coast of Africa. Clearly, “Berbera, West lateral bands on the body, including the caudal peduncle, Coast of Africa”, as stated by Meek (1897: 169) was in error. compared with five broad transverse bands on the body and Meek (1897) compared P. el li oti with P. volitans, but not a single broad horizontal band on the caudal peduncle in with P. antennata or the other congeners. Subsequently, P. cincta and P. radi ata (Figs 2, 4; see also Matsunuma and Herre (1952) and Beaufort and Briggs (1962) provided com- Motomura 2015). Moreover, P. antennata is characterized prehensive synonymies for P. antennata but did not include by 3–47 blotches on the pectoral-fin membranes (none in P. el li oti . Similarly, Smith (1957b) and Eschmeyer (1986), P. cincta and P. radi ata , and numerous narrow transverse who reviewed pteroini fishes in the western Indian Ocean, bands in P. sphex) and the posterior portions of the upper overlooked P. el li oti , although the species had been listed pectoral-fin rays without bands (with bands in P. brevipec- and regarded as a junior synonym of P. antennata (author toralis, P. mombasae and P. paucispinula) (Eschmeyer and given as Rüppell) by Boulenger (1898) in The Zoological Randall 1975; Matsunuma and Motomura 2013b, 2014, Record. More recently, Eschmeyer (1998), Matsunuma and 2015; this study). Motomura (2014, 2015) and Eschmeyer et al. (2017) recog- Synonymy. Pterois antennata was originally described nized P. el li oti as a junior synonym of P. antennata, all with- by Bloch (1787) as Scorpaena antennata, based on material out detailed discussion and erroneously giving the publica- from Ambon, Indonesia. Although Bloch (1787) did not tion year of Meek (1897) as 1899. clearly indicate the number of type specimens of S. anten- The holotype of P. el li oti possesses characters consistent nata, he stated (Bloch 1787: 22) that he had dissected two with the examined specimens of P. antennata, including specimens each of S. antennata and Scorpaena volitans (= P. the lectotype of the latter: dorsal-fin rays XII, 11; pectoral- volitans). A French translation of this account (Bloch 1788) fin rays 17 (both sides); scale rows in longitudinal series 51; also indicated the existence of more than one syntype of S. body and caudal peduncle with numerous narrow trans- antennata. Paepke and Fricke (1992) subsequently con- verse dark bands; pectoral fin with ca. 2 or 3 blotches (ap- sidered Bloch’s (1787) S. antennata to have been based on parently obscured) and posterior portions of soft rays (free two syntypes, supported by Bloch’s own handwritten cata- of membrane) without bands. Because no clear differences log. One of the two syntypes was sold at the beginning of are apparent between the holotype of Pterois ellioti Meek, the 19th century and is now apparently missing (Lichthen- 1897 and the lectotype of Scorpaena antennata Bloch, 1787, stein 1823; Paepke 1999). The other syntype, without the the former is regarded as a junior synonym of the latter. original label, was located and registered by H.-J. Paepke as Morphological changes with growth. Analysis of 49 ZMB 796 (Fig. 3A) (Paepke and Fricke 1992). Paepke (1999) proportional measurements of P. antennata showed growth- subsequently designated ZMB 796 as the lectotype of S. an- related changes in the proportions of some body parts (Fig. tennata. Examination of the specimen during this study re- 6). Body depth and width, postorbital length and suborbital vealed its dorsal-, anal-, pectoral- and pelvic-fin ray counts depth tended to increase relative to SL with growth (Fig. to be consistent with the original description and drawing 6A), whereas orbit diameter, interorbital width at the supra- of S. antennata given by Bloch (1787), as were the relatively ocular spine base, and length of the first and longest dorsal- short supraocular tentacles with four dark bands and lateral and anal-fin soft rays, pectoral-fin rays and pelvic-fin spine branches. Moreover, Bloch (1787) stated that his specimens and longest soft ray, and caudal fin become proportionally of S. antennata had been dissected to observe the gonads, shorter with growth (Fig. 6B). No distinctive ontogenetic ZMB 796 having been similarly dissected. Therefore, we fol- changes were apparent for other body proportions or mer- low Paepke (1999) and regard ZMB 796 as the lectotype of istic characters. However, the relative length of supraocular S. antennata. tentacle (% of orbit diameter) shortens with growth in sev- In his account of S. antennata, Bloch (1787) stated that the eral members of Pterois (see Matsunuma and Motomura specimens had been collected together with P. volitans (stat- 2013b, 2014, 2015), including P. antennata (Fig. 6C). Small ed by Bloch 1787: 22), the examples of the latter having been specimens of BPBM 110035 (1 of 2 specimens, 21.4 mm SL) collected from Ambon, Indonesia (Bloch 1787: 20). Thus, and WAM 26092-016 (12.8 mm SL; Fig. 7) possessed only the type locality of S. antennata is also regarded as Ambon. two anal-fin spines, the incomplete head spination and body Paepke (1999) stated the type locality of S. antennata to be squamation of those specimens confirming their larval sta- “East India” [the East Indies], based on Bloch’s catalog. tus. Meek’s (1897) description of P. el li oti did not identify the Geographic variations in the number of pectoral-fin catalog number of the holotype, Ibarra and Stewart (1987) blotches. Geographic variations in the number of blotches later identified it as FMNH 484 (Fig. 3B). The overall ap- on the pectoral fin were apparent in P. antennata (Figs 5, 8), pearance of FMNH 484 is consistent with the drawing of specimens from central-south Pacific Ocean island groups, 108 M. Matsunuma and H. Motomura

Fig. 6. Relationships of body width (A), interorbital width at preocular spine base (B) (both as % of SL) and supraocular tentacle length (C) [as % of orbit diameter (OD)] to standard length (mm) in Pterois antennata, showing ontogenetic changes.

Fig. 7. Preserved larval specimen of Pterois antennata, WAM-P 26092.016, 12.8 mm SL, Christmas Island, Australia. including the Phoenix, Kermadec, Line, Cook and Pitcairn groups, plus French Polynesia, tending to have a greater number compared with those from other localities (Fig. 9). Pterois antennata from French Polynesia, figured by Bacchet et al. (2016), had a highly blotched pectoral fin. However, there was no clear geographic separation of groups. Three specimens from Tonga (USNM 337707, 78.6–106.9 mm SL) possessed relatively fewer pectoral-fin blotches (6–8). In contrast, notwithstanding the body size Fig. 8. Relationship of number of blotches on pectoral fin difference, a specimen (NMNZ P.009501, 145.0 mm SL) (counted on both sides) to standard length (mm) in Pterois anten- nata from French Polynesia (FP) and Pitcairn Islands (PT) (tri- from Raoul Island, Kermadec Islands (south of Tonga) pos- angles); Cook (CO), Kermadec (KE), Line (LI) and Phoenix (PH) sessed 36 or 37 blotches, being included in the central-south islands (squares); and other localities (circles). See also Fig. 5. Pacific population. However, a second specimen (NMNZ P.050178, 110 mm SL) from Raoul Island possessed ca. 11 from selected species of Pterois is shown in Fig. 10. In the blotches [counted from Motomura and Struthers (2015: fig. present analysis, identification of P. antennata, P. radi ata 156.5 as Pterois cf. antennata)]. Moreover, no distinctive dif- and P. paucispinula clades were supported by examinations ferences were recognized in other morphological characters of voucher specimens representing one or more sequences. or COI sequences among geographic populations (Fig. 10). Identification of P. cf. mombasae was based on the DDBJ/ Therefore, differences in pectoral-fin coloration are regarded EMBL/GenBank database, a voucher specimen not being here as intraspecific geographic variation of P. antennata. available for this study. Similar geographic variations in the number of pectoral-fin Among sequences included in Hubert et al.’s (2012) rays were reported for P. radi ata in the Indo-Pacific region analysis as P. antennata (DDBJ/EMBL/GenBank accession by Matsunuma and Motomura (2015). numbers: JQ432074–432077, JQ350293 and JQ350294), Genetic lineages within P. antennata. Although Hu- JQ432076 and JQ432077 were included in “one lineage bert et al. (2012) recognized three genetic lineages within found only in French Polynesia”. The voucher specimen of P. antennata, based on COI sequences, their sequences of P. JQ432077 (= USNM 392525) was also identified here as P. antennata in fact included P. paucispinula. A neighbor-join- antennata. In the present analysis, both French Polynesian ing tree based on 548 basepairs of the COI gene sequence sequences were included in a clade with sequences from the Redescription of Pterois antennata 109

Fig. 9. Pectoral fin of Pterois antennata from various localities. (A–C) inner surface in fresh specimens and (D–H) outer surface in pre- served specimens. (A) KAUM–I. 32022, 99.1 mm SL, Japan; (B) KAUM–I. 29745, 100.6 mm SL, Japan; (C) KAUM–I. 29746, 153.9 mm SL, Japan; (D) USNM 360991 (1 of 2 specimens), 142.5 mm SL, Seychelles; (E) BPBM 25487 (1 of 3), 112.9 mm SL, Phoenix Islands; (F) BPBM 6927, 120.9 mm SL, French Polynesia; (G) BPBM 11191, 143.4 mm SL, French Polynesia; (H) NMNZ 9501, 145.0 mm SL, New Zealand (Ker- madec Islands).

Pacific Ocean, including Taiwan, Vietnam and Northern The voucher specimen of JQ350294 (from Réunion) was Mariana Islands (Fig. 10). also identified here as P. antennata (from a high-resolution Hubert et al. (2012) also recognized a “second lineage photograph of the specimen provided by N. Hubert). In the only found in the Indian Ocean”, represented by JQ350294. present analysis, the Réunion sequence represented a sister 110 M. Matsunuma and H. Motomura clade of the Pacific Ocean clade, as in Hubert et al. (2012). of JQ350293 (from Madagascar), JQ432074 and JQ432075 Although numerous examples of P. antennata from the In- (both from French Polynesia). However, the voucher speci- do-Pacific region were examined during this study, no dis- men of JQ432074 (= USNM 392530) was re-identified here tinctive morphological differences were recognized between as P. paucispinula (see below). Therefore, the Pacific Ocean specimens from the Indian and Pacific oceans. In contrast, (French Polynesia) subclade of “the third lineage” of Hubert as noted above, geographical variations in the number of et al. (2012) could represent P. paucispinula. In contrast, al- pectoral-fin blotches were apparent between specimens though the voucher specimen of JQ350293 from Madagas- from the central-south Pacific Ocean, including French car was also identical with P. antennata, judging from a pho- Polynesia, and other localities, but no distinctive divergence tograph of the specimen, the sequence formed a clade with in COI sequences occurred among the Pacific Ocean clade, sequences of P. cf. mombasae from Sri Lanka and South Af- including sequences from Taiwan, Vietnam, Northern Mari- rica in the present analysis (Fig. 10). Further investigations ana Islands, and French Polynesia (Fig. 10). of both mitochondrial and nuclear loci are needed to deter- In addition, Hubert et al. (2012) recognized a “third lin- mine the genomic differentiation among P. antennata and eage found in both Indian and Pacific oceans” on the basis related taxa.

First Record of Pterois paucispinula from French Polynesia

An example of P. paucispinula from Moorea, Society Is- lands, French Polynesia (USNM 392530, 95.9 mm SL; Fig. 11) represents the first record of the species from the south- central Pacific Ocean, previous records having been re- stricted to the southeastern Indian Ocean and western Pa- cific (from northern Australia east to the Wallis and Futuna Islands) (Matsunuma and Motomura 2014). The specimen conformed to P. paucispinula in both meristic and mor- phometric values, as follows: XIII, 11 dorsal-fin rays; III, 6 anal-fin rays; 19 pectoral-fin rays (both sides); 56 longitudi- nal series scale rows; 25 pored lateral-line scales; 8/14 scale rows above/below lateral line; 9 scale rows between last dorsal-fin spine and lateral line; 8 scale rows between sixth dorsal-fin spine and lateral line; 8 pre-dorsal-fin scale rows; 4 (upper)+10 (lower)=14 (total) gill rakers; body depth at anal-fin origin 29.5% SL; head width 12.9% SL; postorbital length 16.6% SL; longest dorsal-fin spine length 47.8% SL; caudal-peduncle depth 10.8% SL (Table 2). Color character- istics included; pectoral fin with several bands on posterior portion of rays (free of membrane); several narrow to wide vertical bands laterally on body and caudal peduncle. In ad- dition, the area of ctenoid scales on the body was restricted

Fig. 10. Neighbor-joining tree based on 548 basepairs of COI gene sequence. All sequence data taken from DDBJ/EMBL/ GenBank and identified by accession number and collection local- ity. Sequences shown in bold type were included in Hubert et al.’s (2012) analysis (see text). Identifications of asterisked sequences based on examination of voucher specimens. Numbers at branch- es indicate bootstrap probabilities in 1,000 bootstrap replications. Fig. 11. Fresh specimen of Pterois paucispinula from French Evolutionary distances computed using the uncorrected p-distance. Polynesia, USNM 392530, 95.9 mm SL. Photo: J. T. Williams. Redescription of Pterois antennata 111

(usually dorsolateral region only at ca. 90 mm SL, some- iwa (NSMT), T. Satoh, S. Chiba, E. Katayama and F. Tanaka times extending below lateral line onto ventrolateral region, (formerly NSMT); I.-S. Chen (NTOU); K. Hatooka (for- and on pectoral-fin base and laterally on caudal peduncle in merly OMNH); C. Aungtonya (PMBC) and U. Satapoomin adults). These characters are consistent with the diagnostic (formerly PMBC); R. Bills and O. Gon (SAIAB); T. Alper- characters of the species given by Matsunuma and Motomu- mann, H. Zetzsche (SMF) and F. Krupp (formerly SMF); K. ra (2014). The French Polynesian specimen differed slightly Shibukawa (SPMN); T. Yoshino and Y. Sakurai (formerly from western Pacific specimens of P. paucispinula, having a URM); J. Williams (USNM); S. Morrison (formerly WAM); relatively greater number (37) of blotches on the pectoral- K. Hagiwara (YCM) and M. Hayashi (formerly YCM); M. fin membranes, compared with 14–28 blotches in the latter, V. Nazarkin and A. V. Balushkin (ZIN); and P. Bartsch and similar to the geographic variation in number of pectoral- C. Lamour (ZMB) for their kind hospitality during the first fin blotches in P. antennata. author’s visits to their institutions. We are indebted to J. Although Naranji et al. (2016) recently recorded P. paucis- Sparks and B. Brown (AMNH); D. Catania (CAS); R. Feeney pinula from India (based on four specimens), their basis for (LACM); C. Struthers (NMNZ); and R. Winterbottom and identification was not given. Although the Indian specimens H. López-Fernández (ROM) for providing opportunities to were not available for this study, a photographed specimen examine specimens. We are grateful also to S. Chiba (for- (Naranji et al. 2016: fig. 1) is most like P. mombasae in over- merly NSMT) for instructing the first author in molecular all body appearance, including relatively shorter dorsal-fin analysis; N. Hubert (Université de La Réunion, France), J. spines. The latter species is currently known from northern Randall and J. Williams for providing photographs of speci- Australia north to southern Japan and east to French Poly- mens; H.-C. Ho and S. Koizumi (formerly URM) for donat- nesia (Matsunuma and Motomura 2014; this study). ing specimens; M. Itou, Y. Haraguchi and M. Takayama, and other volunteers and students of KAUM for curatorial as- sistance and collection of specimens; R. Fricke for help with Comparative material examined German translation; G. Hardy (Ngunguru, New Zealand) for reading the manuscript and providing help with English; Pterois brevipectoralis: 10 specimens (49.2–117.9 mm SL) and S. Poss (CAS) and anonymous reviewers provided help- listed in Matsunuma and Motomura (2013b). Pterois cinc- ful comments on the manuscript. This study was supported ta: 31 specimens (47.1–159.4 mm SL) listed in Matsunuma in part by JSPS KAKENHI Grant Numbers JP19770067, and Motomura (2015). Pterois mombasae: 37 specimens JP26241027, JP24370041, JP23580259, and JP26450265; (26.1–160.1 mm SL) (including holotype) listed in Matsunu- JSPS Fellow (PD: 16J00047); the JSPS Core-to-Core Pro- ma and Motomura (2014, 2015). Pterois paucispinula: 41 gram: B Asia-Africa Science Platforms; the “Biological specimens (22.7–143.5 mm SL) (including holotype) listed Properties of Biodiversity Hotspots in Japan” project of the in Matsunuma and Motomura (2014, 2015), with the fol- National Museum of Nature and Science, Tsukuba, Japan; lowing additions. BPBM 22234, 56.3 mm SL, Caban Island, “Establishment of Research and Education Network on Luzon, Philippines, 30 m, 2 September 1977; USNM 392530, Biodiversity and Its Conservation in the Satsunan Islands” 95.9 mm SL, Moorea, Society Islands, French Polynesia, project of Kagoshima University adopted by the Ministry of 17°29′41″S, 149°51′43″W, 14 March 2006. Pterois radiata: Education, Culture, Sports, Science and Technology, Japan; 187 specimens (21.9–152.3 mm SL) (including neotype of P. and the “Island Research” project by Kagoshima University. radiata and holotype of Pterois vittata) listed in Matsunuma and Motomura (2015), with the following addition. BPBM 4393, 51.8 mm SL, Apiang, Gilbert Islands. Pterois sphex: six References specimens (40.6–111.7 mm SL) (including holotype) listed in Matsunuma and Motomura (2013b). Allen, G. R. and Adrim, M. 2003. Coral reef fishes of Indonesia. Zoo- logical Studies 42: 1–72. Allen, G. R., Cross, N. J., and Hoese, D. F. 2006. Scorpaenidae. Lionfish- Acknowledgments es, rockfishes, scorpionfishes, stingfishes, stonefishes, waspfishes. Pp. 876–892. In: Hoese, D. F., Bray, D. J., Paxton, J. R., and Allen, G. R. (Eds) Zoological Catalogue of Australia. Volume 35, Parts We are especially grateful to M. McGrouther, A. Hay 1–3: Fishes. CSIRO Publishing, Collingwood. and S. Reader (AMS); S.-P. 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