Pterois antennata ( Bloch, 1787 )

Matsunuma, Mizuki & Motomura, Hiroyuki, 2018, Redescription and Geographic Variations of Pterois antennata and First Record of Pterois paucispinula from French Polynesia (Scorpaenidae: Pteroinae), Species Diversity 23, pp. 95-114 : 96-106

Pterois antennata ( Bloch, 1787)

[English name: Spotfin Lionfish; standard Japanese name: Nettai-minokasago] ( Figs 1–10 ; Tables 1, 2)

Scorpaena antennata Bloch 1787: 21 , pl. 185 (type locality: Ambon, Indonesia) .

Pterois antennata: Cuvier and Valenciennes 1829: 265 (Waigeo, Indonesia); Günther 1860: 124 (BMNH catalog); Seale 1906: 83 ( French Polynesia); Weber 1913: 497 ( Indonesia); Fowler and Bean 1927: 15 (Tahiti, French Polynesia); Herre 1936: 264 ( Solomon Islands); Matsubara 1943: 345 ( Japan); Smith 1957a: 397 (Aldabra Atoll, Seychelles); Beaufort and Briggs 1962: 45 (Indo-Pacific); Schultz et al. 1966: 23, pl. 125B, fig. 138j (Bikini Atoll and Guam); Masuda et al. 1975: 339, pl. 144-C (in part; Japan); Chen 1981: 48, fig. 12 (Taiwan); Gloerfelt-Tarp and Kailola 1984: 113, unnumbered fig. (southern Indonesia); Shen 1984b: 193, unnumbered fig. (Taiwan); Shimizu 1984: 302, pl. 282-C (in part; Japan); Eschmeyer 1986: 466, pl. 25, fig. 149.7 (western Indian Ocean); Allen and Swainston 1988: 48, unnumbered fig. (northwestern Australia); Winterbottom et al. 1989: 20, fig. 103 (Chagos Archipelago); Randall et al. 1990: 12 (Rapa, French Polynesia); Allen and Smith-Vaniz 1994: 8 (Cocos Keeling Islands); Masuda and Kobayashi 1994: 77, unnumbered figs. ( Japan); Shao and Chen 1994: 237, pl. 56-4 (Taiwan); Kuiter 1996: 90, unnumbered fig. ( Australia); Randall et al. 1997b: 16 (Ogasawara Islands, Japan); Kuiter 1998: 55, unnumbered figs. ( Maldives); Fricke 1999: 157 (Mascarene Islands); Meyers 1999: 101, pl. 28F ( Micronesia); Poss 1999: 2329, unnumbered fig. (Indo-Pacific); Randall 1999: 10 ( Pitcairn Islands); Randall and Earle 2000: 10 (Marquesas Islands, French Polynesia); Hutchins 2001: 27 (Western Australia); Terashima et al. 2001: 26, unnumbered fig. ( Mauritius); Nakabo 2002: 569, unnumbered fig. ( Japan); Allen and Adrim 2003: 29 ( Indonesia); Allen et al. 2003: 370, unnumbered fig. (Indo-Pacific); Hutchins 2003: 465 (Dampier Archipelago, Australia); Hutchins 2004: 364 (Dampier Archipelago, Australia); Kimura et al. 2003: 39, unnumbered fig. (Sulawesi, Indonesia); Randall et al. 2003: 10 ( Tonga); Rao 2003: 129, fig. 204 (Andaman and Nicobar islands); Heemstra and Heemstra 2004: 144, unnumbered fig. (southern Africa); Allen et al. 2006: 879 ( Australia); Senou et al. 2006: 71 (Ie-jima island, Okinawa, Japan); Satapoomin 2007: 46, unnumbered fig. (Andaman Sea); Shao et al. 2008: 246 (southern Taiwan); Allen and Erdmann 2009: 596 (West Papua, Indonesia); Fricke et al. 2009: 39 ( Réunion); Motomura 2009: 65, unnumbered fig. (Andaman Sea); Hirata 2010: 34, right unnumbered figs. (in part; Ehime, Japan); Motomura et al. 2010: 89 (Yaku-shima island, Kagoshima, Japan); Mundy et al. 2010: 25 (Howland, Palmya and Kingman islands); Motomura and Aizawa 2011: 466, fig. 2E (Yaku-shima island, Kagoshima, Japan); Allen and Erdmann 2012: 220, unnumbered fig. (East Indies); Larson et al. 2013: 81 (Northern Territory, Australia); Motomura 2013: 43, unnumbered fig. (Iou-jima and Take-shima islands, Kagoshima, Japan); Nakabo and Kai 2013: 688, unnumbered fig. ( Japan); Fricke et al. 2014: 51 (Madang, Papua New Guinea); Delrieu-Trottin et al. 2015: 5 (Marquesas Islands, French Polynesia); Bacchet et al. 2016: 179, unnumbered fig. ( French Polynesia); Koeda et al. 2016: 20, fig. 85 (Yonaguni-jima island, Okinawa, Japan); Motomura and Harazaki 2017: 23 (Yaku-shima island, Kagoshima, Japan); Siu et al. 2017: 259 ( French Polynesia).

Pseudomonopterus antennatus: Bleeker 1863: 234 ( Ternate, Indonesia).

Pseudomonopterus (Pterois) antennatus: Bleeker 1876: 48 ( Indonesia) ; Bleeker 1875 –1878: pl. 413, fig. 5 (atlas).

Pterois ellioti Meek 1897: 169 , pl. 40 (type locality: Berbera , Somalia) .

Pteropterus antennata: Smith 1957b: 79 , pl. 5, fig. F (Indo- West Pacific); Whitley 1964: 56 ( Australia); Smith and Smith 1965: 56, pls 49F, 97B ( Seychelles); Mandrytsa 2001: 16 (phylogenetic material).

Dendrochirus zebra (not of Cuvier and Valenciennes): Shen 1984a: 32, pl. 32, fig. 264-24c (in part; Taiwan).

Pterois cf. antennata: Motomura and Struthers 2015: 1095 , fig. 156.5 (Kermadec Islands, New Zealand).

Lectotype of S. antennata . ZMB 796, 117.6 mm SL, Ambon, Indonesia.

Other type specimen. FMNH 484 , holotype of Pterois ellioti , 48.8 mm SL, Berbera , Somalia, D . G. Elliot.

Non-type specimens examined. 241 specimens, 12.8– 153.9 mm SL. JAPAN: Boso Peninsula: KPM-NI 22989 , 23.5 mm SL, Tateyama Bay , Chiba Prefecture (Pref.), 7 m, 9 September 2007 . Izu Peninsula: KPM-NI 26918 , 36.1 mm

SL, Izu Oceanic Park, Shizuoka Pref., 16 December 1989. Izu Islands: NSMT-P 30809 , 58.5 mm SL, Miyake-jima island , 34°07′N, 129°31′E, 19 August 1977 . Ogasawara Islands: HUMZ 53491, 118.1 mm SL, Miyanohama , Chichijima island, 7–8 m, 18 June 1976 . Suruga Bay: MSM-02-144, 42.1 mm SL, Mera Port, east of Suruga Bay , Shizuoka Pref ., 28 October 2001. Kii Peninsula: FAKU 99045 , 77.0 mm SL, FAKU 99086 , 84.5 mm SL, Shirahama, Wakayama Pref ., 30 January to 20 February 2011. Tosa Bay: BSKU 79021, 102.2 mm SL, Tosa-shimizu, Kochi Pref ., 15 December 2005; BSKU 86405 , 86.1 mm SL, off Saga, Kochi Pref ., 31 March 1999; BSKU 100096 , 38.0 mm SL, Okino-shima island , Kochi Pref ., 24 September 2009; BSKU 103166, 111.1 mm SL, no further locality; NSMT-P 90778 , 70.0 mm SL, north of Mo-shima Port, Okino-shima island , Kochi Pref ., 3 m, 22 July 2008. Hyuga-nada Sea: MUFS 16455 , 66.0 mm SL, off Kushima, Miyazaki Pref ., 21 October 1998. Satsuma Peninsula: KAUM –I . 38746, 116.5 mm SL, off Kushi, Bonotsu, Minami-satsuma, Kagoshima Pref., 31°18′37″N, 130°13′19″E, 2–25 m, 14 June 2011. Osumi Islands: KAUM –I . 29634, 72.8 mm SL, KAUM –I. 29635, 73.5 mm SL, off west coast of Iou-jima island , 30°47′04″N, 130°15′42″E, 5–20 m, 28 May 2010; KAUM –I . 29745, 100.6 mm SL, KAUM –I. 29746, 153.9 mm SL, off south coast of Iou-jima island , 30°46′32″N, 130°16′43″E, 10–60 m, 26 May 2010; KAUM –I . 37938, 65.8 mm SL, Iou-jima Port , Iou-jima island, 30°46′43″N, 130°16′43″E, 2 m, 18 May 2011; KAUM –I . 62198, 18.8 mm SL, off Urata, Nishinoomote , Tanega-shima island, 30°49′36″N, 131°02′11″N, 10–15 m, 13 June 2014 . Tokara Islands: BSKU 51076 , 58.5 mm SL, Kuchino-shima island , May 1980; KAUM –I . 77955, 74.9 mm SL, west side of Nishinohama Fishing Port , Kuchino-shima island, 29°59′30″N, 129°54′42″E, 5–10 m, 28 August 2015 . Amami Islands: BSKU 6942, 109.8 mm SL, Ankyaba , Amami-oshima island, 2 November 1956; KAUM –I . 57749, 52.9 mm SL, off Setouchi, Amami-oshima island, 28°07′50″N, 129°21′07″E, 3–15 m, 13 December 2013; KPM-NI 21471 , 93.3 mm SL, off Udonosu Beach , Yoron-jima island , 1.5–4m, 22 March 2007; YCM 36181, 39.2 mm SL, off Sukomo , Setouchi , Amami-oshima island , 21 August 1995; YCM 36552, 62.4 mm SL, off Sakinome Beach, Setouchi, Amami-oshima island , 26 August 1995 . Okinawa Islands: KAUM –I . 6436, 133.6 mm SL, KAUM –I. 6438, 120.1 mm SL, KAUM –I. 6586, 132.9 mm SL, Nakagusuku , Okinawa-jima island, 26°15′29″N, 127°47′38″E, 13 July 2007; KAUM –I . 32021, 77.7 mm SL, * KAUM –I. 32022, 99.1 mm SL, off Odo Beach, Itoman , Okinawa-jima island, 26°05′22″N, 127°42′24″E, 23 August 2010; KAUM –I . 43003, 130.3 mm SL, KAUM –I. 43004, 144.2 mm SL, Araha Beach, Nakagami , Okinawa-jima island, 26°18′25″N, 127°45′50″E, 29 May 2011; KAUM –I . 48356, 116.8 mm SL, off Omna , Okinawa-jima island, 26°30′18″N, 127°52′58″E, 1–3 m, 14 August 2011; KAUM –I . 48416, 71.6 mm SL, KAUM –I. 48419, 74.2 mm SL, KAUM –I. 48420, 46.4 mm SL, off Yomitan, Okinawa-jima island , Okinawa Pref ., 26°26′08″N, 127°42′45″E, 1–3 m, 21 October 2011; KAUM –I . 52186, 69.4 mm SL, off Yomitan , Okinawa-jima island, 26°22′01″N, 127°44′15″E, 16 July 2012; KAUM –I . 78775, 102.3 mm SL, Hama Fishing Port, Nakagusuku, Okinawa-jima island , 26°09′06″N, 127°28′25″E, 10 July 2015; URM-P 1324, 104.8 mm SL, Okinawa-jima island ; URM-P 18654, 116.4 mm SL, off Zanpa to Yomitan, Okinawa-jima island , December 1986 . Yaeyama Islands: BPBM 8712 , 2 specimens, 65.5–82.5 mm SL, reef about 0.5 miles off Ishigaki, Ishigaki-jima island, 22 May 1968; * NSMT-P 80768 , 72.8 mm SL, Amitori Bay , Iriomote-jima island , 9 June 1982; URM-P 36099, 77.3 mm SL, off Uehara Beach, Iriomotejima island , 6 August 1996 . TAIWAN: ASIZP 57316 , 95.3 mm SL, Wanlitong, Pingtung, 3 April 1979; ASIZP 59074 , 66.0 mm SL, Liuchiu, Pingtung, 11 December 1979; ASIZP 59571, 108.3 mm SL, Dong-gang , Yilan, 10 June 1979; ASIZP 59601 , 94.0 mm SL, Liuchiu, Pingtung, 25 July 1986; ASIZP 72229, 136.4 mm SL, Magong, Penghu, 20 May 2008; KAUM –I . 43830, 115.5 mm SL, KAUM –I. 43831, 117.2 mm SL, KAUM –I. 43832, 124.0 mm SL, KAUM –I. 43833, 118.8 mm SL, Liuchiu, Pingtung; NMMB-P 6893 , 94.1 mm SL, Shao-liuchiu Island, Pingtung, 22 October 2003; NMMB-P 13172 , 91.2 mm SL, Dong-kang , Pingtung, 1 March 1979; NMMB-P 7033 , 19.2 mm SL, Shao-liuchiu Island , Pingtung, 12 December 2003; ZMB 16148, 31.3 mm SL, Taiwan (no further locality) . PHILLIPINES: AMS I. 40144-010, 5 specimens, 14.0– 73.2 mm SL, northeast side of Maestro de Campo Islands, 12°56′77″N, 121°43′53″E, 25–28 m, 29 May 2000; NSMT-P 34677, 112.3 mm SL, north east coast of Makesi Island, Honda Bay, Palawan , 10 m, 9 November 1988; NTOU 20080414- 101, 106.4 mm SL, NTOU 20080414-116 , 68.1 mm SL, Cebu , 14 April 2008; SMF 10025, 66.2 mm SL, Philippines (no further locality), 1966; USNM 99021, 112.8 mm SL, Lode Bay , Destacado Island , 5 m, 13 March 1909 . PALAU: BPBM 9941 , 36.7 mm SL, Angulpelu Reef, Palau Islands, 110–170 ft (34–52 m), 22 April 1970; FMNH 118622 , 88.8 mm SL, central east coast of Fana Island , Sonsoral, 05°21′17″N, 132°13′57″E, 12 September 2008; FMNH 118623 , 112.0 mm SL, southernmost tip of Fringing Reef of Tobi Island , Hatohobei, 02°59′50″N, 131°07′23″E, 15 September 2008; FMNH 118624 , 80.0 mm SL, north east tip of Helen Reef, Hatohobei, 02°58′01″N, 131°50′06″E, 21 September 2008 . MARIANA ISLANDS: AMS I. 39371-001, 87.5 mm SL, Merizo Reef, 26 August 1967; BPBM 258 , 79.5 mm SL, 12 July 1900; BPBM 4389, 133.1 mm SL, 4 July 1923; SAIAB 86225 , 85.0 mm SL, Lau Lau Beach at Standing Pipe , south east side of Saipan, 15°09′48″N, 145°45′46″E, 20–27 m, 26 May 2003 . CAROLINE ISLANDS: USNM 223578 , 77.6 mm SL, south west side of Pohnpei, Caroline Islands , 06°52′N, 158°06′E, 0–120 ft (0–37 m), 15 September 1980 . WAKE ISLAND: BPBM 15280 , 3 specimens, 79.0– 105.1mm SL, lagoon near Peale Island , 10 June 1953 . MARSHALL ISLANDS: BPBM 4395 , 60.2 mm SL, Ebon Atoll, 1877; BPBM 6429 , 8 specimens, 82.5–118.8 mm SL, BPBM 6432 , 79.7 mm SL, Piirai Island, Eniwetok Atoll, 4 August 1968; BPBM 6433, 114.1 mm SL, north end of Piirai Island , Eniwetok Atoll , 14 August 1968; BPBM 6434, 125.1 mm SL, south end of Japtan Island , Eniwetok Atoll , 1 August 1968; * BPBM 6442 , 96.0 mm SL, Japtan Island , Eniwetok Atoll , 29 July 1968; BPBM 8281 , 50.7 mm SL, Bogen Island , Eniwetok Atoll , 25–70 ft (8–21 m), 3 December 1967; * FMNH 44140 , 22.8 mm SL, Bikini Atoll, Eman Island , 19 July 1947 . INDONESIA: KPM-NI 14031 , 74.5 mm SL, north of Bali; NMV A74072 , 73.4 mm SL, Banda Islands, 1874; NSMT-P 62052 , 72.3 mm SL, Lembeh Strait, Sulawesi, 15 m, 01°27′N, 125°14′E; NSMT-P 63740 , 84.8 mm SL, Indonesia (no further locality); NSMT-P 71523 , 49.0 mm SL, north east coast of Kodek Bay , Lombok, 15 m; NSMT-P 71676 , 76.6 mm SL, Gerupuk, Lombok, 5 m; NSMT-P 102043 , 72.2 mm SL, Meno Island , Lombok; NSMT-P 102044 , 74.4 mm SL, Meno Island , Lombok; SMF 4204 , 89.9 mm SL, south east of Sulawesi, 1909–1910; SPMN-PI 41699 , 59.1 mm SL, Indonesia (no further locality); ZIN 809 , 2 specimens, 108.4– 110.1 mm SL, Ambon, 1858 . PAPUA NEW GUINEA: NMV A 13698, 121.7 mm SL, Hood Bay, New Guinea Island, August 1882; OMNH 13808, 104.9 mm SL, Rabaul , New Britain Island , September 1958; USNM 380294 , 34.6 mm SL, east side of Jalun Island , Hermit Islands, 0–34 m, 2 November 1978; ZMB 15123, 2 specimens, 56.6–84.0 mm SL, New Guinea Island . SOLOMON ISLANDS: AMS I. 22128-085, 3 specimens, 41.1–103.1 mm SL, 3 km from Munda, Roviana Lagoon, New Georgie Island , 08°18′S, 157°15′E, 1 December 1980; AMS I . 39040-045, 57.5 mm SL, south of Neo Village, west coast of Tomotu Island, Santa Cruz Islands , 10°40′S, 165°48′E, 10–30 m, 29 September 1998 . VANUATU: AMS I. 15847-004, 2 specimens, 56.2– 107.8 mm SL, Vanuatu (no further locality); MNHN 1987- 1822 , 89.1 mm SL, off Port Vila, 17°40′59″S, 167°49′59″E, October 1961; USNM 356588 , 53.4 mm SL, Judy Reef, off northwest tip of Tongoa Island , Shepherd Islands , 16°52′30″S, 168°31′30″E, 75–108 ft (23–33 m), 9 June 1996 . NEW CALEDONIA: USNM 324419 , 60.2 mm SL, Bagaat Islet, Ouvea Atoll , Loyalty Islands, 20°37′18″S, 166°16′08″E, 50–71 ft (15–22 m), 16 November 1991 . WALLIS AND FUTUNA: USNM 374803 , 81.5 mm SL, Ile Uvea, Wallis Islands , 13°22′05″S, 176°10′45″W, 7–10 m, 14 November 2000 . PHOENIX ISLANDS: BPBM 25487 , 3 specimens, 89.0– 129.8 mm SL, Kanton Island , 10 February 1950 . LINE ISLANDS: BPBM 6967 , 2 specimens, 60.4–83.3 mm SL, southeast of Sand Island, Palmyra Atoll, Line Islands, 3–12 ft (0.9–4 m), 12 November 1968; * BPBM 14258 , 80.0 mm SL, southwest end of Cook Islet at lagoon entrance, Kiritimati Atoll , Line Islands, 0–2 m, 24 March 1972; BPBM 15413 , 26.0 mm SL, Palmyra , Sand Island , Line Islands, 20 June 1962; BPBM 37599 , 17.9 mm SL, Kiritimati Atoll , Line Islands, 35–40 feet (11–12 m), September 1996; CAS 66059, 3 specimens, 78.5–90.1 mm SL, inside reef on west side of Paradise Island Jetty, Palmyra Atoll, Line Islands, 05°52′30″N, 162°04′42″E, 0.5–10 ft (0.2–3 m), 16 August 1951 . TONGA: USNM 337707 , 3 specimens, 78.1–106.9 mm SL, Luamoko Island, Vava’u Group, Tonga Islands , 18°40′55″S, 174°06′09″W, 70–101 ft (21–31 m), 15 November 1993 . COOK ISLANDS: BPBM 5281 , 67.3 mm SL, Raratonga Island, 15 June 1929; BPBM 11152 , 2 specimens, 36.0– 80.6 mm SL, Rarotonga Island, 75–100 ft (23–30 m), 11 March 1971; FMNH 16311 , 88.8 mm SL, FMNH 16312 , 87.7 mm SL, FMNH 16313 , 49.2 mm SL, Aitutaki Island , 1930 . SOCIETY ISLANDS: AMS I. 28946-012, 2 specimens, 81.4–82.4 mm SL, on barrier reef side on channel, 1.5 km west of Tareu Pass, Moorea, 17°19′S, 149°53′W, 19 February 1989; BPBM 5851 , 98.6 mm SL, outer edge of inshore reef at Papeere , Papetoai Bay , Tahiti , 40 ft (12 m), 14 September 1967; BPBM 6114 , 81.2 mm SL, Atimaouo , Papara , Tahiti , August 1967; BPBM 6927, 118.4 mm SL, outside reef about 1/ 4 mile east of Teauraa Pass , Papara, Tahiti, 90 ft (27 m), 26 February 1969; BPBM 7476 , 3 specimens, 52.6– 65.9 mm SL, Papara, Tahiti, 20 February 1969; BPBM 8922 , 78.4 mm SL, Popote Bay , Papara, Tahiti, 130 ft (43 m), 6 March 1969; FMNH 75884 , 76.3 mm SL, Tahiti, 27 February 1962; MNHN 1984-279 , 87.3 mm SL, off Temae, Moorea, 17°28′59″S, 149°48′00″W, 2 m, September 1983; MNHN 2008-718 , 44.8 mm SL, Moorea, 1 m, 15 March 2006; MNHN 2008-861, 116.9 mm SL, Moorea, 35 m, 18 March 2006; USNM 197734 , 58.5 mm SL, off Tahiti Hotel Arabiri Arie, 25 January 1963; USNM 392525 , 95.4 mm SL, Paroa, Moorea, 17°36′23″S, 149°50′00″W, 15 March 2006 . AUSTRAL ISLANS: AMNH 72813, 122.1 mm SL, north side of Tarohoi Island, Rapa Island, Tubuai Islands , 27°36′S, 144°18′W, 15 April 1970; AMNH 72814 , 4 specimens, 22.4– 93.1 mm SL, Teahateke Point in Anatauri Bay , Rapa Island , Tubuai Islands , 27°36′S, 144°18′W, 12 April 1970; BPBM 11191, 143.4 mm SL, west side off Avera, Rurutu Island , 140–175 ft (43–53 m), 28 February 1971; BPBM 11201 , 82.4 mm SL, west side off Avera, Rurutu Island , 150–190 ft (46–58m), 28 February 1971; BPBM 11246 , 63.1 mm SL, Haurei Bay , Rapa Island , Tubuai Islands, 40–50 ft (12–15 m), 11 February 1971; BPBM 12864 , 80.7 mm SL, Angairao Bay , Rapa Island , Tubuai Islands, 30 January 1976; BPBM 12996 , 2 specimens, 69.5–72.4 mm SL, south side of exposed reef at entrance to Haurei Bay , Rapa Island , Tubuai Islands, 14 February 1971; BPBM 13020, 138.5 mm SL, southwest side of Karapoo Island , Rapa Island , Tubuai Islands, 60 ft (18 m), 16 February 2017; USNM 379753 , 2 specimens, 45.3–100.1 mm SL, north coast off Pt . Kauira, Rapa Island, Tubuai Islands , 27°34′04″S, 144°20′46″W, 2–6 m, 15 November 2002 . TUAMOTU ARHCIPELAGO: CAS 66072, 67.1 mm SL, west side of Raroia, north of Ngaru Pass, Mataira Islet , west side of Raroia Atoll , 16°01′S, 142°26′W, 0–6 ft (0–2 m), 17 July 1952; CAS 66073, 47.4 mm SL, channel between Nengonengo and Kukina islets, west side of Raroia Atoll , 16°00′S, 142°26′W, 0–1.5 ft (0–0.5 m), 5 July 1952 . MARQUESAS ISLANDS: AMS I. 22015-018, 64.6 mm SL, Anaho Bay, Nuku Hiva, 8°55′S, 140°10′W, 1 August 1976; BPBM 4388 , 69.0 mm SL, Marquesas Islands; BPBM 10415 , 73.2 mm SL, Haka Puuae , Taipi Vai , Nuku Hiva , 9 September 1956; BPBM 11035 , 2 specimens, 21.4–41.6mm SL, south side of Vaeho Bay , off entrance to large cave, Ua Pou Island , 0–30 feet (0–9 m), 29 April 1971; BPBM 11114 , 2 specimens, 30.5–56.3 mm SL, point at north side of Hanauu Bay , Fatu Hiva, 40–60 feet (12–18 m), 21 April 1971; USNM 340483 , 78.7 mm SL, Taiohai, Nuku Hiva, 1 February 1957; USNM 399831 , 81.2 mm SL, west coast of Mohotane Island , 09°57′38″S, 138°50′17″W, 6–20 m, 14 October 2008 . PITCAIRN ISLANDS: BPBM 13261, 134.8 mm SL, off Bounty Bay, Pitcairn Island, 100–130 ft (30–40m), 26 December 1970; BPBM 16627, 109.7 mm SL, north side of atoll, Oeno Island , 70 ft (21m), 19 December 1970; BPBM 16636 , 90.7 mm SL, Pitcairn Island , 90–100 feet (27–30 m), 23 December 1970 . CORAL SEA: AMS I. 4119, 51.6 mm SL, Lord Howe Island , 1898; AMS I . 20756-044, 96.1 mm SL, Raine Island, Queensland , 1 February 1979; AMS I . 20965- 025, 88.8 mm SL, northeast of Dunk Islands, Queensland , 17°56′S, 146°43′E, 10–20 m, 27 February 1979; AMS I . 23847-027, 83.1 mm SL, south side of Yonge Reef, Lizard Island, Queensland , 14°35′S, 145°36′E, 8 m, 6 January 1975 . CHRISTMAS ISLAND: AMS I. 20441-005, 83.2 mm SL, Christmas Island, 30 May 1978; FAKU 113358 , 69.0 mm SL, Flyingfish Cave, Christmas Island , 13–26 m, 28 June 1986; * WAM-P 26092.016, 12.8 mm SL, Christmas Island , 10°29′S, 105°40′E, 14 m, 25 May 1978 . KERMADEC ISLANDS: NMNZ P. 009501, 145.0 mm SL, Raoul Island , 29°13′S, 178°10′W, 1956 . ANDAMAN SEA: PMBC 12055 , 89.0 mm SL, Phuket; ROM 69890, 2 specimens, 103.5– 111.6 mm SL, Ko Racha Yai Island , 07°28′15″N, 98°19′35″E, 18 November 1993 . ANDAMAN AND NICOBAR ISLANDS: SMF 28932, 122.9 mm SL, Nicobar Island, Andaman and Nicobar Islands , 11 August 1958 . SRI LANKA: SMF 4327 , 85.2 mm SL, Trincomalee , 21 May 1958 . MALDIVES: * BPBM 32927 , 70.7 mm SL, Maaniyafushi Island, South Male Atoll , 25–30 m, 17 March 1988; FMNH 75838 , 2 specimens, 85.5–101.5 mm SL, south of Male Atoll , Dunidu Island , 19 April 1964; SMF 5386 , 78.9 mm SL, Maldives (no further locality), 1 February 1958; SMF 9770 , 30.5 mm SL, Maldives (no further locality), 17 January 1958 . OMAN: CAS 73270, 107.7 mm SL, Christian Cemetery Cove , 2 m, 1 April 1988 . YEMEN: SMF 4625, 128.2 mm SL, SMF 4626, 105.1 mm SL, Abd al Kuri Island , 10 December 1957 . SOMALIA: CAS 208379, 102.3 mm SL, Gulf of Aden , 11°21′06″N, 49°11′54″E, 27 October 1985 . KENYA: LACM 31005-50, 101.8 mm SL, north side of Mombasa Island , 4°02′S, 39°41′E, 10 September 1970; SAIAB 14400 , 95.3 mm SL, Mombasa, 5°55′S, 39°40′E, August 1957; SMF 9100 , 67.5 mm SL, Mombasa, 24 January 1967; SMF 9932 , 74.3 mm SL, Mombasa, 27 January 1969 . TANZANIA: SAIAB 14398 , 96.2 mm SL, Zanzibar Island , 7°50′S, 39°11′E, 21 September 1952 . COMOROS: CAS 32512, 56.4 mm SL, reef in front of Hotel Coelacanthe, Grande Comoros, 25–30 m, 14 February 1975; SAIAB 34555 , 2 specimens, 61.0– 79.7 mm SL, Grande Comoros , 12°11′S, 43°01′E, 18 October 1986 . MADAGASCAR: MNHN 1992-669 , 82.7 mm SL, off Toliara, 23°19′59″S, 43°30′00″E, 18 August 1968; SAIAB 52777 , 76.8 mm SL, off Berofia Island, Radama Islands , 14°58′S, 47°42′E, 30–32 m, 30 August 1995 . SOUTH AFRICA: CAS 48671, 114.2 mm SL, Sodwana, Zululand, 1 April 1979; SAIAB 8835, 138.6 mm SL, Sodwana Bay , 28°29′S, 32°41′E, 3 April 1979; SAIAB 40493, 124.4 mm SL, Aliwal Shoal, Cathedral, off Umkomaas, Natal, 31°45′S, 30°49′E, 12 August 1992; SAIAB 46164, 114.3 mm SL, Anchors Reef, Aliwal Shoal, Natal, 31°45′S, 30°49′E, 17 June 1994; SAIAB 46397, 101.1 mm SL, Cathedral, Aliwal Shoal, Natal, 31°45′S, 30°49′E, 20 June 1994; SAIAB 62275 , 137.0 mm SL, Protea Bank, 30°50′S, 30°29′E, 10 March 2000 . SEYCHELLES: BPBM 20078, 100.2 mm SL, cast coast of Cerf Island , 6 m, 27 October 1973; USNM

360991, 2 specimens, 107.9–142.5 mm SL, south west of West Island , just south of La Passe du Bois, Aldabra Atoll, 9°23′54″S, 46°12′36″E, 0–3 m, 19 August 1967 . MAURITIUS: CAS 48693, 2 specimens, 35.9–37.4 mm SL, Mascarene Islands , 7 March 1971; MNHN 6556 , 64.6 mm SL, Mauritius (no further locality); SAIAB 2149 , 2 specimens, 44.2–97.2 mm SL, 1 mile east of Beauchamp , 21°44′S, 57°46′E, 7 March 1971; SAIAB 52107 , 2 specimens, 79.5– 99.6 mm SL, off rocks end of Victoria Avenue , Albion , 21°48′S, 57°24′E, 10 May 1995; USNM 349900 , 88.1 mm SL, west coast of Albion, Mascarene Islands, ca . 0–5 m, 10–15 May 1995; USNM 349899 , 46.4 mm SL, 30 m north of Pass , west coast of Flic En Flac, 20°16′S, 57°22′E, 4–10 m, 5 May 1995; ZMB 13713, 119.6 mm SL, Mauritius (no further locality) . RÉUNION: MNHN 6557 , 96.3 mm SL, Réunion (no further locality) . DOUBTFUL RECORD: SMF 12887, 77.1 mm SL, Red Sea (no further locality), 1967 . NO DATA: FAKU 102893, 123.9 mm SL; KPM-NI 6191 , 76.2 mm SL, aquarium trade .

Diagnosis. A species of Pterois distinguished from other species of the genus by the following combination of characters: dorsal-fin rays usually XII, 11 (rarely XII, 10, XII, 12 or XIII, 10); pectoral-fin rays usually 17 (rarely 15, 16 or 18); more than 10 narrow (but of irregular width) transverse lateral bands on body, including caudal peduncle; pectoral-fin membranes with 3–47 (0 or 1 in larvae <21.4 mm SL) blotches; posterior portion of upper pectoral-fin rays (free of membrane) without bands.

Description. Morphometrics and selected meristics shown in Tables 1 and 2. Meristics shown in Table 1 not repeated here. Pelvic-fin rays I, 5. Branchiostegal rays 7.

Body oblong, moderately compressed posteriorly; depth moderate, body depth at pelvic-fin origin 73% (55–93%) of longest dorsal-fin length. Caudal peduncle moderately short, depth subequal to length. Head moderately large. Several sensory pores on underside of each dentary; pore behind symphysial knob of lower jaw in ventral view; pore covered by lower-jaw lip on each side of symphysial knob in ventral view. Several small sensory pores on dorsal surface of each interorbital canal at mid-orbit level. Postorbital sensory canal short, slightly extending ventrally from sphenotic spine base. Isolated postorbital sensory canal (shown in Fig. 1 ) usually absent (including lectotype) (rarely present in other specimens).

Three (rarely four) short barbels on snout tip, their length subequal to anterior nasal tentacle length; numerous minute cirri on ca. 5 (or 4) thin skin ridges on frontal surface of snout bulge. Short tentacle on posterior edge of low, membranous tube associated with anterior nostril, its tip slightly extending beyond posterior margin of posterior nostril when depressed posteriorly. Supraocular possessing extremely long tentacle, relative length of tentacle [178.6% (75.5–342.8) of orbit diameter] decreasing with growth; thin skin ridges along lateral margins with 6 (or 0–5) short lateral branches. A short simple tentacle on anteroventral portion of lacrimal (just anterior to ventrally positioned sensory pore), its length subequal to that of snout tip barbel. A short elongate, leaf-like skin flap on ventralmost corner of

DS: dorsal-fin spine base; LL: lateral line; SR: scale rows. H and L indicate holotype of P.ellioti and lectotype of P. antennata , respectively. Larvae <21.4mm SL possessed II anal-fin spines; 2 specimens possessed 15/17 pectoral-fin rays.

posterior lacrimal plate, its length subequal to anterior lacrimal tentacle [59% (54.0–88.3%) of orbital diameter], its tip reaching posterior margin of maxilla when laid flat (small specimens with relatively long flap, its tip extending far beyond posterior margin of maxilla). Two small leaf-like skin flaps on posterior margin of preopercle, their lengths about half of orbit diameter with tips extending slightly beyond posterior margin of interopercle when laid flat; upper flap situated between 3rd and 4th preopercular spines, lower flap just below 4th preopercular spine base. A small skin flap anterodorsally on orbit surface, its length slightly greater than diameter of posterior nasal pore. No other skin flaps on head or body.

Body, including caudal peduncle, generally covered with well-developed ctenoid scales; thorax and abdomen covered with weakly ctenoid or cycloid scales. Dorsal- and anal-fin bases with 1 or 2 rows of weakly ctenoid or cycloid basal scales, of size about one-fifth to one-third of that of body scales. Pectoral-fin base largely covered with small cycloid scales. Caudal-fin base with ca. 3–5 rows of small cycloid basal scales. Head generally covered with ctenoid scales; snout, both jaws, maxilla, mandible and preocular bone without scales. Orbital surface without scales. Dorsal surfaces of supraocular and postocular bones covered with poorly developed ctenoid scales, most with a single spinule. Cheek region entirely covered with exposed ctenoid scales, suborbital scales not reaching a vertical through mid-orbit (rarely extending anteriorly beyond mid-orbit level in large non-type specimens); suborbital pit without scales (rarely a few ctenoid scales present).

Dorsal profile of snout relatively steep, forming angle of ca. 40° to horizontal axis of head and body. Mandible smooth without ridges. Mouth moderately large, slightly oblique, forming angle of ca. 30° to horizontal axis of head and body; upper edge of posterior maxilla swollen laterally, forming low ridge; posterior margin of maxilla just reaching a vertical through midorbit. Symphyseal gap separating premaxillary teeth bands slightly broader than width of each band; both jaws with band of small, slender conical teeth; ca. 8 (or 9, 10) and 8 (or 9–11) teeth rows at widest portions of upper and lower jaws, respectively; both teeth bands becoming narrow posteriorly; small conical teeth in ca. 8 (or 9, 10), rows forming blunt V-shaped patch on vomer; no palatine teeth.

Nasal spine with 1 (rarely 2 in large specimens) spinous point ( Fig. 1 ). Preocular spine with 1 (sometimes 2 in large specimens) spinous point directed upward. Supraocular with blunt spinous point covered by skin, at supraocular tentacle base. Postocular with 12 (8 on right side of lectotype) (1–14) small spines directed laterally along posterodorsal margin of orbit. Interorbital ridge weakly developed, diverging and reduced posteriorly, connected to coronal spine base. Coronal spine with 1 (or 2) spinous point directed posteriorly (dorsal view). Tympanic spine with 1 (or 2; 0 in small non-type specimens <30 mm SL) spinous point directed almost posteriorly; origin of tympanic spine slightly posterior to that of coronal spine. No spines between coronal and tympanic spines (1 or 2 small spines rarely present in non-type specimens). Occipital region flat; anterior margin of occipital region sloped transversely from between origin of coronal spines, slightly curved posteriorly (dorsal view); anterior border of occipital region rarely with spines in non-type specimens. Parietal spine with 2 (1–3) spinous points on relatively long ridge, diverging posteriorly. Nuchal spine with 2 (or 1) spinous points, its base completely conjoined with that of parietal spine. No spinous points on parietal-nuchal ridge in small non-type specimens <30 mm SL. Coronal and parietal spines well separated, small spine rarely present between them. Sphenotic with 5 (6 on right side of lectotype) (0–8) small spines, surrounding short sensory canal. Pterotic spine with 3 (0–6) spinous points on relatively short ridge. Lower posttemporal spine with 0 (2 in right side of lectotype) (or 1–3) spinous points on relatively short ridge. Cleithrum with upper short and from specimens of Pterois antennata , including primary types of Scorpaena antennata and Pterois ellioti , and French Polynesian specimen of

P. paucispinula .

lower long ridges, possessing 1 (0–2) and 1 (0–7) spinous points, respectively. Small specimens <30 mm SL lacking sphenotic, pterotic and cleithrum spines.

Lateral lacrimal ridge with 2 (3 in right side of lectotype) (0–6) spinous points; other ridges on lacrimal moderately spinous (spines absent in small specimens). Suborbital ridge strongly spinous with numerous lower supplemental spines, totaling 10 (0–37) spines (small specimens <30 mm SL lacking spine on suborbital ridge; becoming spinous with growth); suborbital pit without spines. Anterior lacrimal spine absent. Posterior lacrimal spine broad, plate-like, with 2 (1–9) spinous points on distal margin, all spinous points exposed, except for point at posterior lacrimal tentacle base. Preopercle with 5 spines, lowermost 2 spines minute, indis- tinct (covered with skin); first (uppermost) spine directed posterodorsally; second spine directed posteroventrally; third spine plate-like, with no (1 or 2) spinous points; no supplemental spine at each preopercular spine. Opercular spine absent.

Origin of 1st dorsal-fin spine above origin of lower posttemporal spine base; bases of 1st and 2nd dorsal-fin spines closer than those of subsequent adjacent spines; 8th (6th to 9th; usually 8th) spine longest; 11th (penultimate) [or 12th (last)] spine shortest; length of penultimate spine 36% (24–100%) and 91% (71–185%) of that of 10th (antepenultimate) and 12th spines, respectively; membrane of spinous portion of dorsal fin strongly incised. Dorsal-fin soft rays all branched; 6th (3rd to 7th; usually 5th) ray longest, but dis- tinctly shorter than longest dorsal-fin spine; posteriormost ray free from caudal peduncle. Anal-fin origin below last dorsal-fin spine base; 3rd spine longest; length of 1st spine 52% (39–65%) and 43% (30–58%) of that of 2nd and 3rd spines, respectively. Anal-fin soft rays all branched; 3rd (or 2nd; usually 3rd) ray longest, its length moderately greater than that of longest dorsal-fin soft ray; posteriormost ray free from caudal peduncle. Pectoral-fin rays extremely long, filamentous, all rays unbranched in all stages, posterior portion of upper rays free from membrane (about half of longest ray associated with membrane); lower 6 (7 or 8) rays weakly thickened; 2nd (2nd to 6th) ray longest, tip of longest ray extending beyond caudal-fin base when laid flat [relative length of rays (% SL) decreasing with growth]. Pelvic-fin spine base below 3rd dorsal-fin spine base; all soft rays branched; 3rd (2nd to 4th; usually 3rd) soft ray longest, its tip extending far beyond a vertical through 1st anal-fin spine base, but not reaching that of posteriormost anal-fin soft ray base; 5th soft ray with membranous connection to abdomen for ca. one-third of ray length. Soft rays of dorsal, anal and pelvic fins all unbranched in small non-type specimens <ca. 30 mm SL. Caudal fin rounded, with 3 procurrent rays, 2 segmented unbranched rays and 5 segmented branched rays in both dorsal and ventral series; procurrent rays well developed, exposed.

Color when fresh. Based on photographs of fresh nontype specimens ( Fig. 2 ). Ground color of head and body pink to pale red; whitish ventrally. Lips whitish. Supraocular tentacle same as ground color of head, with 3–7 dark red to black bands; flap on orbit surface dark red to black; other skin flaps or tentacles on head pale red. A broad reddish- brown band with narrow white margin from supraocular tentacle base, obliquely crossing eye and reaching posterior margin of preopercle. Two reddish-brown bands with narrow white margins saddling nape; anterior band short, just behind eye, at level between coronal spine and origin of parietal spine; posterior band just behind anterior band at level of parietal ridge, reaching central posteroventral margin of opercle at level of upper origin of pectoral-fin base. Numerous variable width, reddish-brown bands on body (extending onto dorsal and anal fins) and caudal peduncle (their number increasing with growth, up to ca. 30 in specimens> 100 mm SL), those below mid dorsal-fin spinous portion usually broadest; 1 or 2 reddish bands on thorax. Lateral bands on caudal peduncle narrow, slightly oblique. Ground color of dorsal-fin spines same as on body, with 1–5 (5 on longest spine) reddish-brown bands. Membranes of soft-rayed portions of dorsal and anal fins, and caudal fin semi-translucent; numerous small reddish-brown to black spots on those fin rays, additional small white spots between each dark spot. Pectoral-fin membrane pale red to dark brown, blackish marginally, with ca. 4 or 5 broad dark bands on basal portion; rays pale red to dark red, whitish posteriorly, posterior portions of upper rays (free of membrane) without bands; variable size dark blotches on membranes. Pelvic-fin membrane dark red, grading to nearly black prox- imally, with rays white or pale red.

Color of preserved specimens. Based on all examined specimens ( Figs 3 , 4 ). Head and body creamy-white; bands on head and body becoming brown to black (diffuse in long-term preserved or poor condition specimens) [lectotype with ca. 20 (mostly diffuse) (ca. 10–25 in non-type specimens) lateral bands on body]; supraocular tentacle with 6 (3–7) brown or black bands; ca. 63 (27–139) small spots on soft-rayed portion of dorsal fin, ca. 65 (22–89) on that of anal fin, ca. 52 (44–170) on caudal fin, number of spots increasing with growth; ca. 9 (3–47) variable size brown to black blotches on pectoral fin.

Distribution. Pterois antennata is found in the Indo-Pacific region from the east coast of Africa eastward to French Polynesia and from northern Australia north to Japan ( Fig. 5 ). One doubtful record has been reported from the Red Sea, without a precise locality (SMF 12887). Although the Red Sea was included in the distribution of P. antennata by Fricke et al. (2014), no additional records have been reported from the Red Sea ( Golani and Bogorodsky 2010; Matsunuma et al. 2016) and its presence there is doubtful. Collection data from 42 lots indicate that this species occurs at depths of 0–60 m, with most records from less than 20 m.

Figures

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 5mm.

Fig. 2. Fresh specimens of Pterois antennata from various localities at different growth stages. (A) KAUM–I. 62198, 18.8mm SL, Japan; (B) KAUM–I. 29635, 73.5mm SL, Japan; (C) KAUM–I. 32021, 77.7mm SL, Japan; (D) USNM 399831, 81.2mm SL, French Polynesia; (E) BPBM 5851, 98.6 mm SL, French Polynesia; (F) KAUM–I. 38746, 116.5mm SL, Japan; (G) BPBM 6927, 118.4mm SL, French Polynesia; (H) KAUM–I. 29746, 153.9mm SL, Japan. Photos: J. T. Williams (D) and J. E. Randall (E, G).

Fig. 3. Primary types identified as Pterois antennata. (A) lectotype of Scorpaena antennata, ZMB 796, 117.6mm SL, Ambon, Indonesia; (B) holotype of Pterois ellioti, FMNH 484, 48.8mm SL, Berbera, Somalia.

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

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 Islands; KE, Kermadec Islands; LI, Line Islands; PH, Phoenix Islands.

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, 12.8mm SL, Christmas Island, Australia.

Fig. 8. Relationship of number of blotches on pectoral fin (counted on both sides) to standard length (mm) in Pterois antennata from French Polynesia (FP) and Pitcairn Islands (PT) (triangles); Cook (CO), Kermadec (KE), Line (LI) and Phoenix (PH) islands (squares); and other localities (circles). See also Fig. 5.

Fig. 9. Pectoral fin of Pterois antennata from various localities. (A–C) inner surface in fresh specimens and (D–H) outer surface in preserved 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, 120.9mm SL, French Polynesia; (G) BPBM 11191, 143.4mm SL, French Polynesia; (H) NMNZ 9501, 145.0mm SL, New Zealand (Kermadec Islands).

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 locality. 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 branches indicate bootstrap probabilities in 1,000 bootstrap replications. Evolutionary distances computed using the uncorrected p-distance.