Species Diversity 19: 133–139
25 November 2014
DOI: 10.12782/sd.19.2.133

Sebastapistes perplexa, a New Species of Scorpionfish(Teleostei: Scorpaenidae) from Japan
Hiroyuki Motomura1,4, Masahiro Aizawa2 and Hiromitsu Endo3
1 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp

2 Imperial Household Agency, 1-1 Chiyoda, Chiyoda-ku, Tokyo 100-8111, Japan
3 Laboratory of Marine Biology, Faculty of Science, Kochi University,

2-5-1 Akebono, Kochi 780-8520, Japan
4 Corresponding author(Received 28 July 2014; Accepted 24 September 2014)
A new scorpionfish, Sebastapistes perplexa, is described on the basis of 43 specimens from the Pacific coast of Japan,
where it ranges from the Boso Peninsula to Shikoku, including the Izu Islands, in depths of 2–48 m. The new species ischaracterized by having 12–15 (mode 14) pectoral-fin rays; 33–42 (37) scale rows in longitudinal series; 4 or 5 (5) scalerows above the lateral line, 9–13 (10) scale rows below the lateral line; 3 or 4 (4) scale rows between the sixth dorsal-fin
spine base and the lateral line; 3–5 (4) scale rows between the last dorsal-fin spine base and the lateral line; 2–4 (3) predorsal scale rows; 11–17 (15) gill rakers; palatine teeth present; ctenoid body scales; simple anterior and posterior lacrimal
spines; no lateral lacrimal spine or ridge; the first and second suborbital ridges fused to form a single ridge with one suborbital spine at its end; a median ridge on lower opercular spine; the opercular spines not covered by scales; and a dark blotch
usually visible on the subopercle and pectoral-fin base (distinct in preserved specimens). Sebastapistes perplexa appears to
be restricted to temperate waters whereas its congeners are primarily tropical and subtropical species. The new species iscompared in detail to two congeneric species that share some diagnostic features with it, and to the sympatric S. strongia(Cuvier in Cuvier and Valenciennes, 1829).
Key Words: Actinopterygii, morphology, Sebastapistes strongia, Sebastapistes ballieui, Sebastapistes mauritiana.

Introduction
The Indo-Pacific scorpionfish genus Sebastapistes Gill in
Streets, 1877 is characterized by having 12 dorsal-fin spines,
teeth on the palatines, the posterior lacrimal spine directed
posteroventrally, and pored lateral-line scales continuing
onto the caudal-fin base, and by the lack of a deep occipitalpit (Poss 1999; Motomura 2009). Some species of the deepwater scorpionfish genus Neomerinthe Fowler, 1935 sharethese characters with Sebastapistes, but can be distinguishedfrom the latter by the head squamation and spines, somemeristics, and habitat depths. The species of Sebastapistes
are usually distributed in shallow coastal waters of the tropics (Motomura et al. 2006b), but one species, Sebastapistes
strongia (Cuvier in Cuvier and Valenciennes, 1829), extends
to inshore waters of temperate regions (Nakabo 2002).
During revisionary studies of the genus Sebastapistes,
unidentified specimens of this genus from shallow temperate waters off the Pacific coast of Japan were found in fish
collections of Japanese museums, and additional specimens
were collected from the Boso Peninsula, which forms theeastern edge of Tokyo Bay. These specimens are describedherein in detail as a new species endemic to Japanese temperate waters. Sebastapistes strongia, a primarily tropical andsubtropical species, co-occurs with the new species at similar habitats and depths in temperate Japanese waters, butS. strongia is clearly distinguished from the new species inseveral aspects, including coloration, head spine structure,and meristics. In this paper, the new species is comparedwith the other nominal species of Sebastapistes, including S.
strongia.

Materials and Methods
Measurements generally follow Motomura (2004a, b),
except for head width (Motomura et al. 2005b, 2006a),
maxillary depth (Motomura et al. 2006b), and body depth(Motomura et al. 2012). Counts follow Motomura et al.
(2005a–c) and Motomura and Johnson (2006), with predorsal scale row counts following Motomura et al. (2006b). The
last two soft rays of the dorsal and anal fins are counted as
single rays, each pair being associated with a single pterygiophore. Counts of preopercular spines begin with the uppermost spine. Standard length is expressed as SL. Terminology of head spines follows Randall and Eschmeyer (2002:
fig. 1) and Motomura (2004b: fig. 1) with the following
additions: the spine at the base of the uppermost preopercular spine is referred to as the supplemental preopercularspine (Eschmeyer 1965); the spine on the lateral surface ofthe lacrimal bone is referred to as the lateral lacrimal spine(Motomura and Senou 2008: fig. 2; Motomura et al. 2011b:
fig. 1); and the coronal and pretympanic spines (the latter© 2014 The Japanese Society of Systematic Zoology

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Fig. 1. Color photographs of Sebastapistes perplexa, showing variations of fresh coloration and supraocular tentacle lengths, and sexual
dichromatism. A, CMNH-ZF 15719, female holotype, 48.8 mm SL; B, KAUM–I. 65652, male paratype, 38.2 mm SL; C, KAUM–I. 20724, maleparatype, 35.4 mm SL; D, KAUM–I. 20716, female paratype, 33.1 mm SL; E, CMNH-ZF 15718, female paratype, 46.6 mm SL.

as an extra spine) are as figured in Chen (1981: fig. 1) andMotomura et al. (2004: fig. 14b) respectively. The specimens
examined in this study are deposited in the Laboratory ofMarine Biology, Faculty of Science, Kochi University, Kochi(BSKU); Coastal Branch of Natural History Museum andInstitute, Chiba, Katsuura (CMNH); Kyoto University Museum, Kyoto (FAKU); Kagoshima University Museum, Kagoshima (KAUM); Kanagawa Prefectural Museum of Natural History, Odawara (KPM); Muséum National d’Histoire
Naturelle, Paris (MNHN); National Museum of Nature andScience, Tsukuba (NSMT); Museum Support Center, Smithsonian Institution National Museum of Natural History,

Suitland (USNM); and Yokosuka City Museum, Yokosuka(YCM). Underwater photographs of the new species referred to in this study are registered at the Image Databaseof Fishes in KPM (KPM-NR) and the color description oflive individuals is based on the KPM-NR photographs.
Comparative material of Sebastapistes was listed in Motomura (2009), Motomura and Senou (2009), and Motomura
et al. (2011a). Type specimens of nominal species mentioned in this paper are as follows: MNHN 6883, 2 syntypesof Scorpaena ballieui Sauvage in Vaillant and Sauvage, 1875,
31.9–85.7 mm SL, Hawaii; MNHN 8993, syntype of S. ballieui, 79.2 mm SL, Hawaii; MNHN 9557, 2 syntypes of S.

New species of Sebastapistes from Japan

135ballieui, 63.1–74.2 mm SL, Hawaii; USNM 50691, holotype
of Sebastapistes corallicola Jenkins, 1903, 85.9 mm SL, Honolulu, Oahu Island, Hawaii; MNHN 5711, holotype of Scorpaena mauritiana Cuvier in Cuvier and Valenciennes, 1829,
61.8 mm SL, Mauritius; MNHN 4069, holotype of Scorpaena
megastoma Sauvage, 1878, 65.9 mm SL, Réunion; FAKU
6083, holotype of Scorpaena hatizyoensis Matsubara, 1943,
62.9 mm SL, Hachijo Island, Izu Islands, Japan.
Sebastapistes perplexa n. sp.
[New standard Japanese name: Akamadara-fusakasago][New English name: Temperate Scorpionfish](Figs 1–2)
Holotype. CMNH-ZF 15719, female, 48.8 mm SL,
Hirane off Hasama, Tateyama, Chiba Prefecture, Japan,
34°58′42″N, 139°47′33″E, 15 m, M. Aizawa, hand net, 28
July 2006.
Paratypes. 42 specimens, 19.6–50.7 mm SL, all from
Pacific coast of Japan. BOSO PENINSULA: CMNH-ZF
7049, female, 40.2 mm SL, Hatto-ne off Isomura, Kamogawa, 35°05′20″N, 140°07′15″E, 22 m, H. Tachikawa, handnet, 1 October, 2003; CMNH-ZF 7050, female, 30.4 mm
SL, Isomuradashi off Ubara, Katsuura, 35°07′00″N,
140°16′58″E, 20 m, M. Aizawa, hand net, 3 October 2003;
CMNH-ZF 7620, female, 30.4 mm SL, CMNH-ZF 7621,
female, 50.7 mm SL, Mankurou off Ishomura, Kamogawa, 35°05′22″N, 140°07′17″E, 15 m, M. Aizawa, hand net,
5 March 2004; CMNH-ZF 12096, female, 38.2 mm SL,
CMNH-ZF 12097, female, 42.8 mm SL, Igaijima off Ishomura, Kamogawa, 35°05′18″N, 140°07′13″E, 15 m, M. Aizawa,
hand net, 4 December 2004; CMNH-ZF 15717, female,
49.4 mm SL, CMNH-ZF 15718, female, 46.6 mm SL, collected with holotype; KAUM–I. 17390, female, 30.1 mm SL,
KAUM–I. 17595, male, 30.8 mm SL, Takane off Hasama,
Tateyama, 34°58′38″N, 139°47′19″E, 20 m, M. Aizawa, handnet, 10 December 2008; KAUM–I. 20716, female, 33.1 mm
SL, Hirane off Hasama, Tateyama, 34°58′42″N, 139°47′33″E,
15 m, M. Aizawa, hand net, 2 December 2008; KAUM–I.
20724, male, 35.4 mm SL, Takane off Hasama, Tateyama,
34°58′38″N, 139°47′19″E, 20 m, M. Aizawa, hand net, 2December 2008; KAUM–I. 24764, female, 48.3 mm SL, off
Field Science Center of Tateyama Station (Tokyo University of Marine Science and Technology), Sakata, Tateyama,
34°58′37″N, 139°46′10″E, 7 m, M. Watai, hand net, 17 June
2009; KAUM–I. 24765, female, 43.1 mm SL, KAUM–I.
24766, female, 42.2 mm SL, off Field Science Center of Tate­yama Station (Tokyo University of Marine Science andTechnology), Sakata, Tateyama, 34°58′37″N, 139°46′10″E,
7 m, M. Watai, hand net, 18 June 2009; KAUM–I. 25864,
female, 34.7 mm SL, off Field Science Center of Tateyama
Station (Tokyo University of Marine Science and Technology), Sakata, Tateyama, 34°58′37″N, 139°46′10″E, 7 m, M.
Watai, hand net, 10 November 2009; KAUM–I. 65652, male,38.2 mm SL, Takane off Hasama, Tateyama, 34°58′38″N,
139°47′19″E, 20 m, M. Aizawa, hand net, 25 September
2007; KPM-NI 22993, female, 35.8 mm SL, Banda, Tate­yama Bay, 7 m, A. Murase, hand net, 9 September 2007;

Fig. 2. Lateral view of head of Sebastapistes perplexa. KAUM–I.
17595, male paratype, 30.8 mm SL. Numerous small papillae andsensory pores on head not illustrated. Scale bar: 2 mm.

NSMT-P 18962, male, 23.8 mm SL, southwest off Kamogawa, 34°58′N, 140°01′E, 48 m, 13 November 1973; NSMT-P
57904, 3 females, 42.1–49.2 mm SL, Kominato, Awa, 3 July
1973; NSMT-P 72660, female, 48.6 mm SL, off Kominato,
Kamogawa, 25 June 1981; YCM-P 42102, female, 34.2 mm
SL, Hirane off Hasama, Tateyama, 34°58′42″N, 139°47′33″E,
Sagami Bay Marine Biological Research Club, hand net,
14 December 2002. IZU PENINSULA (SAGAMI BAY):FAKU 59112, female, 47.0 mm SL, Izu Oceanic Park, Futo,
Ito, 20–25 m, T. Murai, hand net, 22 June 1992; FAKU
59211, female, 20.3 mm SL, Izu Oceanic Park, Futo, Ito,
20 m, T. Murai, hand net, 14 July 1992; FAKU 59221, male,23.2 mm SL, FAKU 59222, male, 19.6 mm SL, Izu Oceanic Park, Futo, Ito, 26 m, T. Murai, hand net, 15 July 1992;
KPM-NI 21838, female, 47.1 mm SL, Izu Oceanic Park,
Futo, Ito, K. Suzuki, hand net, 13 June 1994; KPM-NI 34456,
male, 28.0 mm SL, Izu Oceanic Park, Futo, Ito, 5 m, K. Okamoto, hand net, 2 July 1993; KPM-NI 34457, male, 23.4 mm
SL, Izu Oceanic Park, Futo, Ito, 5 m, K. Suzuki, hand net,
July 1993; KPM-NI 34462, female, 28.5 mm SL, Izu Oceanic Park, Futo, Ito, 5 m, 29 July 1993; KPM-NI 34465, male,
31.5 mm SL, off Futo Port, Ito, 6 m, K. Suzuki, hand net, 10
August 1993. IZU ISLANDS: KPM-NI 24975, 24.0 mm SL,
KPM-NI 24976, 25.3 mm SL, Nazumado, Hachijo Island, 18
m, S. Kato, hand net, 17 July 2007; NSMT-P 30818, female,
35.3 mm SL, Chotaro-ike tide-pool, southern Miyake Island,
34°03′N, 139°32′E, 2 m, J. Shepard, 6 May 1976. SHIKOKU(KOCHI PREFECTURE): BSKU 91169, female, 24.9 mm
SL, BSKU 91170, male, 27.2 mm SL, Kuboura, Okinoshimaisland, Sukumo, 32°44′N, 132°33′E, 5–15 m, T. Hirata, handnet, 24 July 2007; BSKU 95359, female, 25.8 mm SL, Himeshima island, Sukumo, 32°44′N, 132°29′E, 5–15 m, H. Endoet al., hand net, 23 July 2003; NSMT-P 77472, 25.0 mm

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SL, NSMT-P 77473, 27.2 mm SL, Akazaki, Moshima Port,
Okinoshima island, Sukumo, 3 m, E. Katayama, hand net,
23 July 2007; NSMT-P 90812, male, 27.0 mm SL, north of
Moshima Port, Okinoshima island, Sukumo, BSKU members, hand net, 23 July 2008.
Underwater photographs. 48 photographs, all fromJapan. IZU PENINSULA (SURUGA BAY): KPM-NR
1446, Ose, 16 m, KPM-NR 4898, Ose, 13 m; KPM-NR 7881,
7884, 7885, Ose, 5 m; KPM-NR 11677, Toda, 3 m; KPM-NR
16022, Ose, 18 m; KPM-NR 22886, Ose; KPM-NR 27444–
27445, Matsuzaki, 10 m; KPM-NR 34360, Ose; KPM-NR
40649, Ose; KPM-NR 63834, Koganezaki; KPM-NR 84849,
Ose, 12 m; KPM-NR 94114, Ose, 10 m; KPM-NR 97355,
Ose, 20 m. IZU PENINSULA (SAGAMI BAY): KPM-NR
6875, 6876, Futo, 24 m; KPM-NR 6877–6882, Futo, 5 m;
KPM-NR 11775, Atami, 10 m; KPM-NR 13628, Futo, 5 m;
KPM-NR 14837, Atami, 6 m; KPM-NR 14854, Atami, 13 m;
KPM-NR 15084, Atami; KPM-NR 15092, Atami, 12 m;
KPM-NR 15107, Atami, 15 m; KPM-NR 15108, Atami,
12 m; KPM-NR 15180, Futo, 5 m; KPM-NR 22887, Futo;
KPM-NR 29333, Futo; KPM-NR 36060, Izu Peninsula, detailed locality unknown; KPM-NR 80038, Futo, 9 m; KPMNR 80329, Futo, 5 m; KPM-NR 88092, Kawana, 12 m;
KPM-NR 88788, Kawana, 16 m; KPM-NR 89187, Kawana,
10 m; KPM-NR 95616, Futo, 15 m; KPM-NR 95617, Futo,
10 m. IZU ISLANDS: KPM-NR 7149, Hachijo Island, 32 m;
KPM-NR 39023, Izu-oshima island. KII PENINSULA(WAKAYAMA PREFECTURE): KPM-NR 33259, Kushimoto, 16 m; KPM-NI 84635, Kumano, 15 m; KPM-NR
96045, Shirahama, 20 m.
Diagnosis. A species of Sebastapistes with the following
combination of characters: 12–15 (mode 14) pectoral-finrays; 33–42 (37) scale rows in longitudinal series; 4 or 5 (5)scale rows above lateral line, 9–13 (10) scale rows below lateral line; 3 or 4 (4) scale rows between sixth dorsal-fin spinebase and lateral line; 3–5 (4) scale rows between last dorsalfin spine base and lateral line; 2–4 (3) predorsal scale rows;
11–17 (15) gill rakers; palatine teeth present; ctenoid bodyscales; anterior and posterior lacrimal spines simple; laterallacrimal spine and ridge absent; first and second suborbital
ridges fused to form single ridge with one suborbital spine
at its end; lower opercular spine with median ridge; opercular spines not covered by scales; interorbital ridges poorly
developed, with shallow channel between ridges; coronalspines usually absent, but poorly developed spines rarely
present; dark blotch usually on subopercle and pectoralfin base (distinct in preserved specimens); indistinct dark
blotch on spinous portion of dorsal fin in males; largest recorded specimen 50.7 mm SL.
Description. In the description below (including thecolor description of preserved specimens and specimens
when fresh), the data and description of the holotype arepresented first, followed by data for paratypes in parentheses (range and mode for counts, range and mean for proportional measurements) when different.
Dorsal-fin rays XII, 9 (XI, 10 in one paratype; XII, 10 in
one paratype); anal-fin rays III, 5 (III, 4 in 2 paratypes; III,
6 in 2 paratypes); pectoral-fin rays 14 on each side (13 oneach side in 2 paratypes; 15 on each side in 5 paratypes;
asymmetrically 14 and 15 in 3 paratypes; asymmetrically 12
and 14 in one paratype); scale rows in longitudinal series 38(33–42, mode 37); pored lateral-line scales 23 (20–23, 22);
scale rows above lateral line 4 (4 or 5, 5); scale rows below
lateral line 11 (9–13, 10); scale rows between sixth dorsalfin spine base and lateral line 4 (3 or 4, 4); scale rows between last dorsal-fin spine base and lateral line 4 (3–5, 4);predorsal scale rows 3 (2–4, 3); gill rakers on upper limb
5 (4–6, 4); gill rakers on lower limb 10 (7–11, 10), including 2 (0–3, 2) rakers on hypobranchial; total gill rakers 15(11–17, 15). Following morphometrics expressed as percentage of SL: body depth 40.2 (33.6–41.1, mean 36.8); bodywidth 24.0 (15.9–24.8, 20.4); head length 42.0 (38.5–44.2,
40.9); snout length 11.3 (10.3–12.7, 11.3); orbit diameter
11.9 (10.9–14.5, 12.9); interorbital width at middle of eye
7.2 (5.6–7.7, 6.8); interorbital width between preocular spine
bases 6.4 (4.7–6.8, 5.7); head width 15.6 (12.6–16.4, 14.7);
upper-jaw length 20.9 (19.5–22.5, 20.9); maxillary depth
5.5 (5.5–7.0, 6.1); suborbital space 1.8 (1.1–2.6, 1.8); postorbital length 20.1 (15.9–20.3, 18.8); distance between tips of
opercular spines 5.7 (4.5–7.4, 6.0); pre-dorsal-fin length 38.1
(32.7–38.5, 35.8); pre-anal-fin length 65.0 (61.3–67.9, 65.5);
pre-pelvic-fin length 39.1 (36.8–42.9, 39.7); first dorsalfin spine length 6.4 (5.6–7.9, 6.6); second dorsal-fin spine
length 12.3 (10.2–14.7, 12.8); third dorsal-fin spine length
17.8 (16.4–20.4, 18.2); fourth dorsal-fin spine length 17.4
(14.8–21.0, 18.2); fifth dorsal-fin spine length 17.0 (14.9–
19.4, 17.2); eleventh dorsal-fin spine length 11.5 (9.5–12.6,
10.9); twelfth dorsal-fin spine length 15.8 (12.0–16.3, 14.7);
longest dorsal-fin soft ray length 24.0 (18.1–23.0, 21.2); firstanal-fin spine length 8.0 (6.4–9.6, 8.1); second anal-fin spine
length 16.6 (15.5–19.4, 17.5); third anal-fin spine length 15.6
(13.4–17.5, 15.5); longest anal-fin soft ray length 22.5 (18.7–
23.0, 21.2); pectoral-fin length 37.9 (31.0–37.9, 35.0); pelvicfin spine length 16.2 (14.2–18.8, 16.1); longest pelvic-fin
soft ray length 25.6 (20.6–27.2, 24.3); caudal-fin length 27.9
(21.8–32.3, 28.3); caudal-peduncle length 20.9 (17.5–22.3,
19.6); caudal-peduncle depth 10.5 (9.6–11.8, 10.6).
Body moderately compressed anteriorly, progressively
more compressed posteriorly. Nape and anterior part ofbody moderately arched (less arched in young and juveniles). Body moderately deep, but body depth less thanhead length. All pectoral-fin rays unbranched (1–3 middle
rays branched in some adults); seventh (sixth to eighth) ray
longest. Second (or third) soft ray longest among dorsal-fin
rays. Second soft ray longest among pelvic- and anal-fin rays.
On head (Fig. 2), supraocular tentacle slender and short, its
length less than orbit diameter (length variable, from absent
to three times greater than orbit diameter; see Fig. 1). Posterior lacrimal spine associated with short, fleshy tentacle,
length of latter less than (or subequal to) that of supraocular tentacle; posterior lacrimal spine tentacle linked posteriorly to head by skin. Distinct tentacle on posterior edge oflow membranous tube associated with anterior nostril; thistentacle reaching to anterior margin of orbit when laid back.
Pectoral-fin axil without skin flaps. Swimbladder absent.
Well-exposed, weakly ctenoid scales covering surface ofNew species of Sebastapistes from Japanbody; exposed cycloid scales (some scales with weak ctenii) on base of pectoral fin; embedded cycloid scales (somescales exposed) on anteroventral surface of body; exposed
cycloid scales on opercle between tips of opercular spines.
Body scales not extending onto rays or membranes of fins,except on base of caudal fin. Embedded cycloid scales oncheek. Lateral line sloping downward from second pored
scale. Underside of dentary with three well-developed sensory pores on each side, first pore below anterior margin oforbit, second pore below anterior end of posterior lacrimal
spine base, third pore located on posterior margin of dentary. Pair of small pores behind symphysial knob of lower
jaw in ventral view.
Mouth large, oblique, forming angle of about 15 (15–30)
degrees to horizontal axis of head and body (Fig. 2). Posterior margin of maxilla just short of vertical drawn throughposterior margin of orbit. No distinct longitudinal ridgeon lateral surface of maxilla. Lower jaw with symphysialknob. Width of symphysial gap separating premaxillaryteeth bands less than width of each band. Villiform teeth
on upper and lower jaws, but no canine teeth; tooth band
narrowing posteriorly. Tooth bands of upper and lower jaws
subequal in width. Four (4–5) rows of small teeth at frontof vomer, becoming 3 (1–3) rows posteriorly, forming Vshaped patch on vomer. About 2–4 rows of small teeth oneach palatine. Underside of lower jaw without ridges.
Dorsal profile of snout steep, forming angle of about 60(50–60) degrees to horizontal axis of head and body. Nasal
spine simple, directed dorsally, its tip extending well beyond
top of ascending process of premaxilla in lateral view. Ascending process of premaxilla not quite intruding into interorbital space, its posterior margin not extending beyond
vertical drawn from anterior margin of posterior nostril in
dorsal view when mouth completely closed. Median interorbital ridge absent. Interorbital ridges poorly developed anteriorly, separated by shallow channel, beginning posterior
to nasal spines and then conjoined at level of origins of tympanic spines, forming indistinct low ridge (forming lump inyoung and juveniles) at front of occiput; diverging anteriorly
and posteriorly in dorsal view, distance between ridges narrowest at level of vertical drawn through middle of pupil. Interorbital space shallow, about one-tenth of orbit extending
above dorsal profile of head. Preocular spine simple, directed dorsally; tip of spine extending well above level of upper
margin of pupil in lateral view; flattened anteriorly andposteriorly; anterior surface of spine without distinct ridge.
Supraocular spine simple, not canted laterally; its length
less than that of postocular spines. Postocular spine simple,
slightly canted laterally; its basal width subequal to widthof tympanic spine base. Tympanic spine simple, slightly
canted laterally, located posterior to postocular spine; posterior end of tympanic spine base not reaching (extending
beyond in some juveniles) level of origin of parietal spinebase in dorsal view. Interorbital and pretympanic spines absent. Coronal spines absent (poorly developed spines present in three paratypes). Occipital region slightly concave(nearly flat or slightly convex in young and juveniles), without distinct transverse ridges at rear of occiput. Occiput sur-137

rounded laterally by bases of tympanic and parietal spines.
Parietal spine simple. Nuchal spine simple; nuchal and parietal spines joined at base. Sphenotic with 2 minute spines.
Postorbital without spines. Pterotic spine simple; length ofspine base less than (or subequal to) that of parietal spine
base. No spine (or single small spine) in area surroundedby parietal, pterotic, and posttemporal spines. Upper posttemporal spine simple. Lower posttemporal spine simple, its
basal length subequal to that of pterotic spine. Supracleithralspine simple. Cleithral spine flattened, pointed.
Lateral lacrimal spine and ridge absent. Anterior tip ofanteriorly directed lacrimal ridge embedded within skin.
Anterior lacrimal spine simple, directed ventroanteriorly,
its tip reaching dorsal margin of upper lip. Posterior lacrimal spine simple, directed ventroposteriorly; posterior lacrimal spines larger than anterior lacrimal spine. Suborbital
ridge single, with spine at end of ridge. Space between ventral margin of eye and suborbital ridge remarkably narrow.
Suborbital pit absent. Preopercle with five spines; uppermost
spine largest, with supplemental preopercular spine on its
base. Preopercle, between its upper end and uppermost preopercular spine, without serrae or spines. Upper opercularspine simple with indistinct low ridge along it upper edge.
Lower opercular spine simple with distinct median ridge.
Space between upper and lower opercular spines withoutridges. Posterior tips of upper and lower opercular spines
not reaching opercular margin (tip of lower spine sometimes reaching to opercular margin).
Origin of first dorsal-fin spine above supracleithral spine.
Posterior tip of pectoral fin extending beyond verticaldrawn through base of third anal-fin spine. Origin of pelvicfin spine slightly posterior to vertical drawn through originof first pectoral-fin ray. Posterior tip of depressed pelvic finjust reaching to anal-fin origin (extending beyond anus, but
not reaching to anal-fin origin, in most specimens). Originof first anal-fin spine anterior to vertical drawn through lastdorsal-fin spine base.
Color of preserved specimens—Body white, mottled withbrown blotches. Subopercle with single indistinct (distinct
in most specimens) dark brown blotch, its size subequal topupil diameter. No distinct marking on opercle. Unpaired
fins semi-transparent, with poorly defined pale brown
blotches. No distinct dark brown blotch on spinous portionof dorsal fin (distinct blotch present between sixth and tenthspines in males). Pectoral fin semi-transparent, with distinct
dark brown blotch, its size greater than pupil diameter, basally between seventh and twelfth rays.
Color when fresh (Fig. 1)—Body reddish, mottled withdark red blotches; anterior part of caudal peduncle chalky
red. Underside of head pale red (or chalky red). Subopercle
with black blotch, its size subequal to pupil diameter. Spinous portion of dorsal fin reddish, with white membranes
distally; no black blotch (black blotch on membranes between sixth and tenth spines in males). Soft-rayed portionof dorsal fin and anal fin reddish, with broad white band
obliquely across middle of each fin. Pelvic fin red, withwhite margin. Pectoral fin semi-transparent red, with poorly
defined black blotch, its size greater than pupil diameter, ba-

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sally between seventh and twelfth rays. Caudal fin reddish,with irregular vertical broad white band.
Color of live individuals—Based on underwater photographs listed above. Body and fin coloration variegated from
whitish, yellowish, and reddish to brownish and blackish.
Many individuals with broad white saddle, its anterior edge
posterior to eye and posterior edge at opercular margin.
Other coloration very similar to that of fresh specimens described above.
Distribution and habitats. This species is known only
from the Pacific coast of Japan, where it ranges from theBoso Peninsula (Chiba Prefecture) to Shikoku (Kochi Prefecture), including the Izu Islands. The type series was collected from the Boso Peninsula, Izu Peninsula, Hachijo
Island, Miyake Island, and Shikoku in depths of 2–48 m.
Underwater photographs of S. perplexa were taken off the
Izu Peninsula, at Izu-oshima and Hachijo islands in the Izu
Islands, and off the Kii Peninsula in depths of 3–32 m. This
species mainly inhabits rocky reefs and also occurs on sandy
bottoms scattered among rocky reefs.
Although the ichthyofauna of the Ryukyu Islands andKyushu, southern Japan, has been well surveyed (e.g.,Motomura and Matsuura 2010, 2014; Motomura et al.
2013), no examples of S. perplexa have been collected there.
In addition, the first author has examined numerous specimens of Sebastapistes from the Indo-Pacific Ocean but
found no specimens of S. perplexa from tropical waters.
These facts indicate that S. perplexa is restricted to temperate waters of Japan.
Etymology. Derived from the Latin perplexa meaning“confused”, in reference to the confused taxonomic status of
the species. The species is well known by local scuba divers
and frequently photographed by them; it is often seen motionless on rocky surfaces in shallow waters and therefore
presents a fine subject for photography. However, this species has been regarded to date as juveniles of Scorpaena or
Parascorpaena, or as Sebastapistes strongia.
Remarks. One female specimen (BSKU 91169, 24.9 mmSL) from Kochi possessed expanded gonads with welldeveloped ova, and some eggs were released when theindividual was captured, indicating that S. perplexa is asmall species among its congeners. Although the smallest
mature female was 24.9 mm SL, the species reaches at least
50.7 mm SL (CMNH-ZF 7621, female, Chiba).
The males of some species of Scorpaenidae, e.g., Parascorpaena moultoni (Whitley, 1961) and several species of Scorpaenopsis, are known to possess a large black blotch on themembrane of the spinous portion of the dorsal fin (e.g., Poss
1999; Randall and Eschmeyer 2002; Motomura et al. 2005c).
Some species, e.g., Scorpaena bergi Evermann and Marsh,
1900 and Scorpaenodes varipinnis Smith, 1957, have a black
blotch in both sexes whereas others, e.g., Parascorpaena
mossambica (Peters, 1855) and Sebastapistes strongia, lack a
blotch in either sex (Eschmeyer 1965; Poss 1999; Motomura2009). In Sebastapistes perplexa, only males have a dark
blotch on the membranes between the sixth and tenthspines of the dorsal fin. Although some scorpaenids develop a black blotch on the spinous portion of the dorsal fin of“mature” males only (e.g., Randall and Eschmeyer 2002), a
blotch was present in all sizes of male S. perplexa available inthis study (23.2 to 38.2 mm SL), indicating that its presence
may not be associated with the onset of sexual maturity.
Sebastapistes perplexa differs from all except five of its
available nominal congeners in having the first and second suborbital ridges fused, forming a single ridge witha single suborbital spine at its end, and a single posterior
lacrimal spine (Fig. 2). The other five nominal species are
S. mauritiana (type locality: Mauritius), S. ballieui (Hawaiian Islands), S. megastoma (Réunion Island), S. corallicola (Hawaiian Islands), and S. hatizyoensis (Izu Islands).
Motomura et al. (2011a) regarded S. mauritiana as a seniorsynonym of S. megastoma. Sebastapistes corallicola and S.
hatizyoensis were regarded as junior synonyms of Scorpaena
ballieui by Eschmeyer and Randall (1975) and Randall(2007), and of S. mauritiana by Nakabo (2002), respectively.
The validity of these synonymies was confirmed in the present study on the basis of examination of type specimens(listed in Material and methods).
Sebastapistes perplexa differs from the two valid species, S. ballieui and S. mauritiana, in having a simple posterior lacrimal spine (vs a spine with two spinous points
in the latter two species), modally 14 pectoral-fin rays (vs16 rays), poorly developed interorbital ridges with a shallow channel between the ridge (vs well-developed ridges
with a deep channel between them), and in lacking a laterallacrimal ridge or spine (vs spine usually present; if absent,a well-developed ridge present) and coronal spines (onlythree of 38 type specimens with poorly developed coronalspines vs well-developed spines always present). The poorly
defined blackish blotch on the subopercle and pectoral-finbase found in most specimens of S. perplexa (distinct in preserved specimens) is lacking in S. ballieui and S. mauritiana.
The Indo-Pacific species S. strongia co-occurs with S. perplexa in the latter’s entire distributional range, and is similar
to S. perplexa in overall body appearance. However, in addition to the above-mentioned suborbital ridge feature, S. perplexa can be easily distinguished from S. strongia by its having one suborbital spine (vs usually two, rarely three spines
in S. strongia) and modally 14 pectoral-fin rays (vs 15 rays;
see Motomura 2009: table 2). While the nape between theanterior four dorsal-fin spine bases and the upper opercular margin of S. strongia usually has a poorly defined blackish blotch (Motomura 2009), S. perplexa lacks such a blotch.
The blackish blotch on the subopercle and pectoral-fin baseof S. perplexa is not found in S. strongia. Scale numbers on
various parts of the body in S. perplexa tend to be lower
than those in S. strongia: e.g., 33–42 (mode 37) scale rows
in longitudinal series in S. perplexa vs 40–44 (43) in S. strongia; 4 or 5 (5) scale rows above the lateral line vs 5 or 6 (6);
9–13 (10) scale rows below the lateral line vs 11–13 (12); 3
or 4 (4) scale rows between the sixth dorsal-fin spine baseand the lateral line vs 5–7 (6); 3–5 (4) scale rows between
the last dorsal-fin spine base and the lateral line vs 5 or 6(5); and 2–4 (3) predorsal scale rows vs 4–6 (5) (Motomura
2009: table 2).

New species of Sebastapistes from Japan

Acknowledgments
The first author is especially grateful to G. Shinohara, E.
Katayama, and M. Nakae (NSMT), H. Senou (KPM), Y. Kai
and T. Nakabo (FAKU), R. Causse, P. Béarez, P. Pruvost, C.
Ferrara, and Z. Gabsi (MNHN), and J. Williams, S. Raredon,and all other staff (USNM) for their assistance during his
stays at their museums. We thank K. Hagiwara (YCM) for
loans of specimens, M. Watai (Fisheries Research Agency,
Yokohama) for donating specimens of the new species to
KAUM, M. Meguro (Foundation of Kagoshima Environmental Research & Service, Kagoshima) for his assistancein taking some measurements, students and volunteers of
KAUM for their curatorial assistance, and G. Yearsley (Ellipsis Editing) for checking an early version of this manuscript. This study was supported in part by Grants-in-Aid
for Scientific Research (A: 26241027, B: 24370041, and C:
23580259 and 26450265) from the Japan Society for the
Promotion of Science, Tokyo, Japan (JSPS), JSPS AsianCore Program “Establishment of Research and Education
Network on Coastal Marine Science in Southeast Asia”, theCoastal Area Capability Enhancement in Southeast Asia
Project of the Research Institute for Humanity and Nature,
Kyoto, Japan, and the “Biological Properties of Biodiversity
Hotspots in Japan” project of the National Museum of Nature and Science, Tsukuba, Japan.

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