Species Diversity, 2010, 15, 71–81

Thysanichthys evides, a Senior Synonym of Sebastellalittoralis, and a Valid Species of Scorpaenodes(Actinopterygii: Scorpaenidae)
Hiroyuki Motomura1, Sakda Arbsuwan2 and Prachya Musikasinthorn2
1

The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima, 890-0065 Japan
E-mail: motomura@kaum.kagoshima-u.ac.jp
2
Department of Fishery Biology, Faculty of Fisheries, Kasetsart University,
Chatuchak, Bangkok 10900, Thailand(Received 5 March 2010; Accepted 1 June 2010)
The nominal species, Thysanichthys evides Jordan and Thompson, 1914
(Scorpaenidae), originally described from Misaki, Japan, has not been reported since its original description. Examination of the type specimens ofT. evides showed them to be identical with the holotype and non-type specimens of a species previously widely-regarded as Scorpaenodes littoralis(Tanaka, 1917), originally described as Sebastella littoralis, also from Misaki.
Thysanichthys evides is herein regarded as a senior synonym of Sebastellalittoralis, and is a valid species of Scorpaenodes.
Key Words: Teleostei, Scorpaenidae, Scorpaenodes, synonymy, Thysanichthys
evides, Sebastella littoralis.

Introduction
The scorpionfish Scorpaenodes littoralis (Tanaka, 1917) (q.v.), originally described as a new genus and species Sebastella littoralis on the basis of a single
specimen from Misaki, Kanagawa Prefecture, Japan, is characterized by having
coronal and interorbital spines, and a dark blotch on the subopercle (Poss 1999;
Nakabo 2002). This species is widely distributed in the Indo-Pacific, including East
Asia (Ishida 1997; Randall et al. 1997; Xinbo 2006), Hawaii (Mundy 2005), French
Polynesia (Randall 2005), Australasia (Hutchins 2001; Paulin et al. 2001), the Arabian Sea (Manilo and Bogorodsky 2003), and South Africa (Eschmeyer 1986). However, it has never been recorded from equatorial regions, and thus has an antitropical distribution (Poss 1999; this study). It is common on shallow rocky reefs intemperate and subtropical regions, especially along the Pacific coast of southern
Japan (Motomura and Iwatsuki 1997; Shinohara et al. 2000; Senou et al. 2002, 2006).
A detailed study on its biology was published by Yoneda et al. (2000).
Scorpaenodes littoralis has been widely recognized as a valid species (e.g., Poss
1999; Greenfield and Matsuura 2002); however, our examination of the holotypes ofThysanichthys evides Jordan and Thompson, 1914 (q.v.) and Sebastella littoralis(both from Misaki, Japan) revealed that they represent the same species. Thus, T.
evides is a senior synonym of Scorpaenodes littoralis, and is regarded here as a

72

Hiroyuki Motomura et al.

valid species of Scorpaenodes. In this paper we document this synonymy through amorphological comparison of non-type material of S. littoralis with the type specimens of T. evides.

Material and Methods
Measurements follow Motomura (2004a, b), with head width following Motomura et al. (2005b, 2006a) and maxillary depth following Motomura et al. (2006b).
Counts follow Motomura et al. (2005a–c) and Motomura and Johnson (2006). The
last two soft rays of the dorsal and anal fins are counted as single rays, each pairbeing associated with a single pterygiophore. 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 preopercular spine (Eschmeyer 1965); the spine on the lateral surface of the lacrimal bone is referred to as
the lateral lacrimal spine (Motomura and Senou 2008, fig. 2); and the coronal andpretympanic (as an extra spine) spines are those figured by Chen (1981, fig. 1) andMotomura et al. (2004, fig. 14b) respectively. Institutional codes are as follows: Biodiversity Research Center, Academia Sinica, Taipei (ASIZP); Laboratory of Marine
Biology, Faculty of Science, Kochi University, Kochi (BSKU); California Academyof Sciences, San Francisco [CAS (SU)]; Carnegie Museum of Natural History, Pittsburgh (CM); Field Museum of Natural History, Chicago (FMNH); Kagoshima University Museum, Kagoshima (KAUM); Museum of Comparative Zoology, Harvard
University, Cambridge (MCZ); Museum of New Zealand Te Papa Tongarewa(NMNZ); National Museum of Nature and Science, Tokyo (NSMT); Museum Support Center of the Smithsonian Institution National Museum of Natural History,
Suitland (USNM); Western Australian Museum of Natural Science, Welshpool(WAM); and University Museum, University of Tokyo, Tokyo (ZUMT).
Type specimens examined in this study — Thysanichthys evides: FMNH 57082(formerly CM 6019a), holotype, 73.8 mm SL, Misaki, Kanagawa Prefecture, Pacificcoast of central Japan (Fig. 1A); SU 22611, 5 paratypes, 39.7–68.4 mm SL, Misaki. Sebastella littoralis: ZUMT 7439, holotype, 75.5 mm SL, Misaki (Fig. 1B).
Non-type specimens previously identified as Scorpaenodes littoralis examinedin this study (51 specimens, 27.9–82.7 mm SL) — JAPAN: BSKU 13974, 80.0 mm SL,
13976, 73.6 mm SL, 13977, 67.8 mm SL, Usa, Tosa, Kochi, Sep. 1968; BSKU 39342, 68.6mm SL, Inojiri, Usa, Tosa, Kochi, 11 Jun. 1983; BSKU 40079, 66.5 mm SL, 40080, 65.3mm SL, 40081, 53.7 mm SL, Uchinoura Bay, Susaki, Kochi, 16 May, 1984; BSKU
45650, 58.7 mm SL, Kashiwa-jima Island, Kochi, 10 m, 23 Apr. 1989; BSKU 53919, 71.4mm SL, Irino Fishing Port, Kuroshio, Kochi, 12 Apr. 2002; BSKU 60855, 81.8 mm SL,
off Susaki, Kochi, 8 Nov. 2002; BSKU 65503, 78.3 mm SL, 70331, 56.9 mm SL, 70332,
49.4 mm SL, 70333, 55.5 mm SL, 70334, 62.5 mm SL, 70335, 71.5 mm SL, west side of
Futanarabi-jima, off Okino-shima Island, Sukumo, Kochi, 22 Jul. 2003; BSKU 67339,
77.3 mm SL, Ashizuri Port, Tosashimizu, Kochi, 6 Aug. 1998; BSKU 87688, 69.2 mm
SL, Iburi, Tosashimizu, Kochi, 13 Jun. 2000; BSKU 91538, 67.2 mm SL, Saga Fishing
Port, Kuroshio, Kochi, 17 Jun. 2007; KAUM-I. 1510, 56.4 mm SL, Tagami, Kumano,
Nishinoomote, Tanega-shima Island, Kagoshima, 30°3847N, 131°0234E, M.
Takayama, 4 Jan. 2007; KAUM-I. 3209, 40.2 mm SL, east of Sakinoyama, Kataura,

Validity of Scorpaenodes evides

73

Fig. 1. Scorpaenodes evides. A, Holotype of Thysanichthys evides, FMNH 57082, 73.8 mm SL,
Misaki, Kanagawa, Japan; B, holotype of Sebastella littoralis, ZUMT 7439, 75.5 mm SL, Misaki, Kanagawa, Japan; C, non-type of Scorpaenodes evides, KAUM-I. 4371, 46.6 mm SL, Minamisatsuma, Kagoshima, Japan.

74

Hiroyuki Motomura et al.

Kasasa, Minamisatsuma, Kagoshima, 31°2544N, 130°1149E, 27 m, M. Ito, 18 Oct.
2006; KAUM-I. 3899, 61.0 mm SL, off Chiringa-shima Island, Ibusuki, Kagoshima
Bay, 31°1638N, 130°4018E, 25 m, Orita Fishery, 20 May 2007; KAUM-I. 4371, 46.6mm SL (Fig. 1C), 4775, 70.0 mm SL, Hirasaki, Bonotsu, Minamisatsuma,
Kagoshima, 31°2033N, 130°1226E, 3–5 m, M. Ito, 20 Jun. 2007; KAUM-I. 5142, 45.8mm SL, 5143, 68.8 mm SL, 5144, 50.0 mm SL, 5145, 57.3 mm SL, 5146, 50.3 mm SL, 5147,
48.0 mm SL, 5148, 39.7 mm SL, 13350, 44.3 mm SL, 13537, 44.6 mm SL, 13538, 49.4 mm
SL, 13539, 41.1 mm SL, 13540, 31.3 mm SL, 13541, 43.2 mm SL, coast of Hanase, Wakiura, Kaimon, Ibusuki, Kagoshima, 31°1136N, 130°3027E, Jun. 1962; KAUM-I.
10765, 69.1 mm SL, Ohto Fishing Port, Kasasa, Minamisatsuma, Kagoshima,
31°2519N, 130°1017E, 4–5 m, M. Ito, 6 Jul. 2008; KAUM-I. 13262, 82.1 mm SL,
Kataura Fishing Port, Kasasa, Minamisatsuma, Kagoshima, 31°25N, 130°10E, 2 m,
M. Ito, 14 Sep. 2008; KAUM-I. 18249, 50.9 mm SL, 18250, 45.9 mm SL, 18255, 58.8 mm
SL, 18311, 44.0 mm SL, 18313, 58.0 mm SL, 18314, 82.7 mm SL, 18317, 73.2 mm SL,
18348, 56.2 mm SL, Ryukyu Islands, 3 Jun. 1966. TAIWAN: ASIZP 58888, 2, 56.7 and58.0 mm SL, Longdong, Taipei, 25°11N, 121°54E, L.-T. He, 29 Jul. 1993; ASIZP 63590,
2, 27.9 and 41.7 mm SL, Dapengwan, Pingtung, 22°44N, 120°47E, P.-L. Lin, 15 May
2000.
Comparative material of Scorpaenodes cf. evides examined in this study (50specimens, 19.6–76.3 mm SL) — SOUTH AFRICA: CAS 48681, 2, 69.6 and 73.2 mm
SL, wreck off Brighton, Natal, P. Heemstra, 29 Apr. 1980. SRI LANKA: CAS 207730,
4, 44.3–56.9 mm SL, Trincomalee, 7.6 m, C. Koenig, 4 Apr. 1970. AUSTRALIA: CAS
13855, 3, 66.4–70.7 mm SL, One Tree Island, Great Barrier Reef, Queensland, 6.1 m,
B. Collette et al., 23 Nov. 1969; MCZ 90620, 4, 45.6–62.9 mm SL, Elizabeth and Middleton Reefs, Great Barrier Reef, Queensland, 0–10 m, D. Williams, 1984; NMNZ P.
20605, 35.0 mm SL, Maroubra, south Sydney, New South Wales, 34°00N, 151°15E, J.
Moreland and J. Yaldwyn, 23 Mar. 1967; WAM P. 27099-007, 2, 65.0 and 65.7 mm SL,
Port Hacking, New South Wales, 34°04S, 151°06E, 6–7 m, J. Hutchins, 28 Jan. 1981;
WAM P. 27590-011, 3, 37.5–67.2 mm SL, north end of Dicks Island, Abrolhos Islands,
28°30S, 113°46E, 30–32 m, G. Allen and N. Sinclair, 17 Apr. 1982. NEW ZEALAND:NMNZ P. 17768, 3, 19.6–25.7 mm SL, NMNZ P. 17774, 2, 30.8 and 36.6 mm SL, Meyer
Island, off Raoul Island, Kermadecs, 29°10S, 177°50W, K. Bailey and M. Francis, 19Aug. 1985; NMNZ P. 41051, 4, 20.0–33.7 mm SL, west side of Meyer Island, off Raoul
Island, Kermadecs, 29°14S, 177°53W, 7 Nov. 2004; NMNZ P. 41067, 59.7 mm SL,
south end of Denham Bay, Raoul Island, Kermadecs, 29°16S, 177°57W, 5 Nov. 2004;
NMNZ P. 41084, 8, 23.0–54.1 mm SL, off Smith’s Bluff, Denham Bay, Raoul Island,
Kermadecs, 29°16S, 178°57W, A. Stewart et al., 10 Nov. 2004; NMNZ P. 41097, Denham Bay, Raoul Island, Kermadecs, A. Stewart, 9 Nov. 2004. FRENCH POLYNESIA: USNM 379396, 2, 36.2 and 58.0 mm SL, off south side of Tarakoi Islet, Rapa,
27°3533S, 144°1812W, 20 m, J. Williams, 23 Sep. 2004. HAWAIIAN ISLANDS:
CAS 13476, 2, 60.9 and 64.3 mm SL, Hauula Park, Honolulu, Oahu, W. Gosline et al.,28 Jun. 1949; CAS 13477, 74.0 mm SL, Kaena Point, Oahu, W. Gosline and class, 4Mar. 1950; CAS 33915, 76.3 mm SL, Hauula Park, Honolulu, Oahu, W. Gosline et al.,
11 May 1952.

Validity of Scorpaenodes evides

75

Results and Discussion
The specimens examined here were recognized as Scorpaenodes Bleeker, 1857
(q.v.) because they had 13 dorsal-fin spines, three well-developed anal-fin spines,
and exposed, spine-shaped caudal-fin procurrent rays, and lacked palatine teeth,characters diagnostic of Scorpaenodes (e.g., Greenfield and Matsuura 2002; Motomura et al. 2010; Motomura unpubl. data). Sebastella has previously been regarded
as a junior synonym of Scorpaenodes [type species: Scorpaena polylepis Bleeker,
1851 (q.v.)] by Eschmeyer (1969) and Mandrytsa (2001).
Characters of the holotype (FMNH 57082, formerly registered as CM 6019a) andparatypes of Thysanichthys evides agreed with those of the holotype of Sebastellalittoralis (ZUMT 7439) and a number of non-type specimens previously identified as
Scorpaenodes littoralis. All specimens were characterized by having coronal andinterorbital spines, and a dark blotch on the subopercle, a character combination
unique to Scorpaenodes littoralis among the Indo-Pacific Scorpaenodes. Although
the holotype of T. evides (Fig. 1A) had an unusually high body depth (40.1% of SL)
compared with those of the holotype of Sebastella littoralis and non-type specimensof Scorpaenodes littoralis (30.8–38.2% of SL, mean 33.0%), X-ray examination of theformer showed a strong curvature of the spine caused by deformed vertebrae.
Although T. evides has not been reported since its original description and
Scorpaenodes littoralis has been widely used as a valid name by numerous researchers (see Introduction) [thereby meeting the condition of the Article 23.9.1.2of the International Code of Zoological Nomenclature (International Commission
on Zoological Nomenclature 1999)], the species-group name of the former publishedin 1914 is of insufficient antiquity (post-1899) to meet the Article 23.9.1.1 of theCode. Thus, the provisions of the Article 23.9 for “reversal of precedence” cannot
be applied so as to validate Sebastella littoralis. Thysanichthys evides is regarded
herein as a valid species, Scorpaenodes evides, being a senior synonym of Sebastella littoralis. Although it is not directly relevant to the nomenclatural considerations here, the status of Thysanichthys Jordan and Starks, 1904 (q.v.) as a valid
monotypic genus with palatine teeth (type species: Thysanichthys crossotus Jordan
and Starks, 1904) (Poss 1999; Nakabo 2002) is also confirmed.
Meristics and morphometrics of Scorpaenodes evides from East Asia, including
the type specimens of Sebastella littoralis and T. evides, are given in Table 1. Fifty
specimens (19.6–76.3 mm SL) collected from outside the East Asian region werealso examined for comparative purposes in this study (see Material and Methods)
but were not included in Table 1 because they possibly represented more than one
species or geographic subspecies or, at least, geographically distinct populations.
We have recognized at least seven geographic populations in the species currently
identified as Scorpaenodes evides, i.e. those from the southeastern coast of Africa,
the northwestern and northeastern Indian Ocean, Australasia, East Asia, theHawaiian Islands, and the southeastern Pacific, some of which represent disjunctdistributions that make inter-populational gene flow unlikely. Molecular analysis
and more detailed morphological comparisons will be required in the future to assess the taxonomic status of these populations. Incidentally, Scorpaenodes littoralis
figured by Kim et al. (2005: 218, fig. 392) from Korea was re-identified here as Scorpaena miostoma Günther, 1877, and Kim et al.’s (2005: 217, fig. 389) Scorpaena miostoma was re-identified as S. neglecta Temminck and Schlegel, 1843.

76

Hiroyuki Motomura et al.

Table 1. Meristic and morphometric characters of Scorpaenodes evides, including types ofThysanichthys evides and Sebastella littoralis.

Standard length (SL, mm)
Meristics
Dorsal-fin rays

Thysanichthys
evidesSebastellalittoralis

Scorpaenodes
evidesHolotype
Paratypes
FMNH 57082
n5
Japan
Japan

Holotype
ZUMT 7439
Japan

Non-typesn51Japan andTaiwan73.8

39.7–68.4

75.5

27.9–82.7

XIII, 9

XIII, 9

XIII, 9

18/18

18/18

III, 545
25
5
5

17–19/
17–18
III, 5
43–47
24–26
4–6
4–5

III, 545
24
5
6

XIII-XIV(XIII),
6–9 (9)16–19 (18)/
17–19 (18)III, 542–49 (48)
22–26 (25)
4–6 (5)
3–6 (4)

Pectoral-fin rays
(one side/other side)Anal-fin rays
Scale rows in longitudinal series
Pored lateral-line scales
Scales above lateral line
Scale rows between last dorsal-fin
spine base and lateral line
Scale rows between 6th dorsal-fin
spine base and lateral line
Gill rakers (upper+ceratohyal+hypohyal)

5

4–5

5

3–5 (4)583
16

5–67–92–4
14–18693
18

5–6 (5)6–10 (9)1–4 (3)14–20 (17)Morphometrics (% of SL)
Body depth
Body width
Head length
Snout length
Orbit diameter
Interorbital width1
Interorbital width2
Head width
Upper-jaw length
Maxillary depth
Postorbital length
Between tips of opercular spines
Pre-dorsal-fin lengthPre-anal-fin length
Pre-pelvic-fin length
1st dorsal-fin spine length
2nd dorsal-fin spine length
3rd dorsal-fin spine length
4th dorsal-fin spine length
5th dorsal-fin spine length
6th dorsal-fin spine length
12th dorsal-fin spine length
13th dorsal-fin spine length

40.1
25.1
42.6
10.6
12.9
7.1
5.8
14.8
23.2
6.9
20.2
4.7
40.5
72.1
40.0
6.6
12.1
14.6
16.9
16.3
16.0
7.2
11.4

32.8–41.9
19.2–22.6
38.8–42.3
9.5–11.6
13.7–14.5
4.8–6.1
4.8–5.7
16.1–17.3
21.1–22.7
6.4–7.1
15.8–19.6
4.6–6.1
35.9–42.7
69.9–73.6
38.1–43.9
4.5–6.5
7.1–11.4
12.6–15.2
13.5–15.1
14.2–16.7
14.2–15.5
6.4–7.6
10.8–14.3

34.6
19.1
40.9
10.6
11.4
6.4
5.7
14.8
22.1
8.5
20.7
5.0
37.1
72.9
41.7
6.8
–
13.8
14.0
14.2
–
7.3
10.5

30.8–38.2 (33.0)
17.0–25.8 (21.6)
36.9–45.4 (41.6)
9.1–12.1 (10.4)
11.0–15.6 (13.2)
4.9–7.3 (5.9)
4.8–6.3 (5.4)
13.2–20.5 (16.9)
19.9–24.3 (21.8)
6.0–8.2 (6.9)
17.2–21.4 (19.3)
4.0–6.8 (5.1)
36.7–43.6 (39.1)
68.9–77.2 (71.6)
35.3–45.4 (39.0)
5.1–8.8 (6.8)
8.8–15.1 (11.1)
11.7–17.2 (14.1)
12.3–19.1 (14.8)
11.7–17.6 (14.8)
12.7–18.7 (15.0)
4.9–9.7 (7.1)
8.7–13.1 (11.0)

Validity of Scorpaenodes evides
Table 1.

77

Continued.

Morphometrics (% of SL)
Longest dorsal-fin ray length
1st anal-fin spine length
2nd anal-fin spine length
3rd anal-fin spine length
Longest anal-fin ray length
Pectoral-fin ray length
Pelvic-fin spine length
Longest pelvic-fin ray length
Caudal-fin length
Caudal-peduncle length
Caudal-peduncle depth

Thysanichthys
evidesSebastellalittoralis

Scorpaenodes
evidesHolotype
Paratypes
FMNH 57082
n5
Japan
Japan

Holotype
ZUMT 7439
Japan

Non-typesn51Japan andTaiwan

18.2
–
20.4
17.5
22.5
33.9
16.4
13.3
27.2
16.3
10.2

15.6–19.9 (17.2)
7.2–10.7 (8.7)
18.6–26.4 (22.0)
13.2–21.8 (16.8)
17.8–25.6 (21.5)
28.9–38.8 (33.5)
14.8–20.3 (17.8)
19.5–26.5 (23.3)
22.9–30.4 (27.0)
14.1–27.7 (18.8)
9.0–11.9 (10.8)

16.5
9.2
21.5
–
–
32.3
19.2
26.7
27.0
17.1
10.5

–
7.3–9.7
20.7–22.6
16.5–18.0
19.3–23.9
33.3–35.9
16.8–19.7
22.3–27.6
25.4–27.4
17.8–19.4
10.3–11.8

Morphometrics are expressed as percentages of standard length. Means in parentheses include type data.
1
At vertical midline of eye; 2 at posterior end of preocular spine base; – broken.

A morphological description of Scorpaenodes evides from East Asia follows.
Body not strongly compressed anteriorly, progressively compressed posteriorly.
Nape and anterior body not strongly arched. Body depth moderate, less than headlength. Short, broad tentacle on posterior edge of low membranous tube associatedwith anterior nostril. Pectoral-fin axil without skin flaps. Numerous scales covering occiput, area behind orbit, and cheek; few scales in interorbital space; noscales on lateral surface of maxilla and anteroventral surface of head. Well-exposed ctenoid scales on lateral surface of trunk, scales becoming cycloid on abdomen. Exposed cycloid scales on anteroventral surface of body and pectoral-finbase. Body scales extending onto basal parts of fin rays or fin membranes, except
for bases of pelvic fins. Lateral line nearly straight from upper end of gill opening
to caudal-fin base. Underside of dentary with three distinct sensory pores on eachside. Pair of small pores behind symphysial knob of lower jaw in ventral view. Posterior margin of maxilla usually not reaching (or rarely extending slightly beyond)a vertical through posterior margin of orbit. No distinct longitudinal ridge on lateral surface of maxilla. Width of symphysial gap separating premaxillary teeth
bands subequal to width of each band. Upper and lower jaws with villiform teeth,
lacking canine teeth. Villiform teeth forming Y-shaped patch on vomer. Palatine
teeth absent. Underside of lower jaw without ridges. Gill rakers relatively shortand spinous; slit present behind fourth gill arch. Swimbladder present. Dorsal profile of snout steep, forming angle of about 40 degrees to horizontal axis of head andbody. Ascending process of premaxilla slightly intruding into interorbital space,its posterior margin extending well beyond level of posterior margin of posterior
nostril in dorsal view. Median interorbital ridge absent. Interorbital ridges weakly
developed, separated by relatively shallow channel; interorbital spines usually

78

Hiroyuki Motomura et al.

present at level of midline of eye. Interorbital space shallow, with about one-tenthof orbit extending above dorsal profile of head. Preocular spine simple, slightly
flattened anteriorly and posteriorly; anterior surface of spine without distinct
ridges. Supraocular, postocular, and tympanic spines simple. Coronal spines usually present (sometimes absent on one side of head) between origins of tympanic
spines. Pretympanic spines absent. Occipital region flat, with no distinct ridges oneither front or rear part of occiput. Parietal and nuchal spines joined at base. Sphenotic with several small spines. Tube-shaped postorbital bone usually without
spines. Pterotic spine simple, directed posteroventrally. Upper and lower posttemporal spines present; length of upper spine about one-third of that of lower spine.
Supracleithral and cleithral spines simple. Lateral lacrimal spine simple, well developed. Anterior and posterior lacrimal spines simple, triangular or rounded, notstrongly developed. Dorsal margin of lacrimal without spines. Suborbital ridgewith 2 well developed spines; first spine tip below posterior margin of pupil; second
spine at end of suborbital ridge. Additional weak spine sometimes present below
tip of first suborbital spine. No space between ventral margin of eye and suborbital
ridge. Suborbital pit small. Preopercle with 3 spines; uppermost spine largest, with
supplemental preopercular spine on its base; space between second and third
spines slightly broader than that between first and second spines. Preopercle, between uppermost preopercular spine and upper end of preopercle, usually without
serrae or spines (rarely with small spines). Upper and lower opercular spines simple without distinct median ridge. Space between upper and lower opercularspines without ridges. Posterior tip of upper opercular spine not reaching opercular margin; posterior tip of lower opercular spine reaching or extending slightly
beyond opercular margin. Origin of first dorsal-fin spine above supracleithralspine. Posterior margin of opercular membrane reaching a vertical through baseof third dorsal-fin spine. Posterior tip of pectoral fin extending slightly beyond a
vertical through origin of anal fin. Origin of pelvic-fin spine slightly posterior to avertical through origin of pectoral fin. Posterior tip of depressed pelvic fin notreaching origin of anal fin. Origin of first anal-fin spine anterior to a verticalthrough last dorsal-fin spine.
Fresh coloration of East Asian S. evides as follows: head reddish-black dorsallyand laterally, reddish ventrally, with distinct blackish blotch on subopercle(blotch subequal in size to pupil diameter); trunk variegated, mottled with brown,
red, and/or orange blotches and spots. Spinous portion of dorsal fin blackish anterobasally, remaining parts reddish or orange with white spots scattered on membranes; no distinct black blotch on posterior spinous portion of dorsal fin. Softrayed portion of dorsal fin semitransparent with reddish areas basally and distally. Pectoral, pelvic, and anal fins with reddish spots (anal fin sometimes withbroad red band basally). Caudal fin semitransparent with reddish submarginal
band; reddish spots on caudal-fin rays. In underwater photographs of S. evides inJapan, many individuals with broad, white saddle, its anterior edge behind eye,and posterior edge level with middle of spinous portion of dorsal fin (saddle width
subequal to head length).

Validity of Scorpaenodes evides

79

Acknowledgements
We are especially grateful to K.-T. Shao, H.-C. Ho, and Y.-C. Liao (ASIZP); W.
Eschmeyer, T. Iwamoto, D. Catania, and M. Hoang (CAS); H. Endo (BSKU); M. A.
Rogers and K. Swagel (FMNH); K. Hartel and A. Williston (MCZ); C. Roberts, A.
Stewart, and C. Struthers (NMNZ); G. Shinohara (NSMT); J. Williams, L. Palmer,
S. Raredon, and K. Murphy (USNM); S. Morrison (WAM); and K. Sakamoto (ZUMT)
for specimen loans and their kind hospitality during the first author’s stays attheir institutions. The first author’s visits to ASIZP, CAS, and USNM were supported by a Grant-in-Aid for Young Scientists (B) from the Ministry of Education,
Science, Sports and Culture, Tokyo (MEXT; 19770067); those to BSKU and NSMT by
a Grant-in-Aid for Scientific Research (A) from the Japan Society for the Promotion of Science, Tokyo (JSPS; 19208019); those to FMNH and MCZ by a Dispatch
Grant to Overseas for Curators from MEXT; that to NMNZ by the Biosystematics
of New Zealand Exclusive Economic Zone Fishes Program (New Zealand Foundation for Research, Science, and Technology contract MNZX0203); and that to WAM
by a Travel Grant for Academic Meetings from JSPS (211026). The second and third
authors’ visits to KAUM were supported by the JSPS International Training Program and a Grant-in-Aid for Scientific Research (B) from JSPS (20405012), respectively. We thank Y. Haraguchi and students (KAUM) for their curatorial assistance, K. Swagel and P. Willink (FMNH) for taking a photograph and X-rays of theholotype of T. evides, K. Kuriiwa (NSMT) for taking a photograph of the holotype of
S. littoralis, and G. Hardy (Ngunguru, New Zealand) for reviewing the manuscript.

References
Bleeker, P. 1851. Nieuwe bijdrage tot de kennis der Percoïdei, Scleroparei, Sciaenoïdei,Maenoïdei, Chaetodontoïdei en Scomberoïdei van den Soenda-Molukschen Archipel.
Natuurkundig Tijdschrift voor Nederlandsch Indië 2: 163–179.
Bleeker, P. 1857. Bijdrage tot de kennis der ichthyologische fauna van de Sangi-eilanden.
Natuurkundig Tijdschrift voor Nederlandsch Indië 13: 369–380.
Chen, L.-C. 1981. Scorpaenid fishes of Taiwan. Quarterly Journal of the Taiwan Museum 34:
1–60.
Eschmeyer, W. N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including
four new species. Bulletin of Marine Science 15: 84–164.
Eschmeyer, W. N. 1969. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from the Indian Ocean, with comments on the limits of the genus. Occasional Papers of the California Academy of Sciences 76: 1–11.
Eschmeyer, W. N. 1986. Family No. 149: Scorpaenidae. Pp. 463–478. In: Smith, M. M. andHeemstra, P. C. (Eds) Smith’s Sea Fishes. J. L. B. Smith Institute of Ichthyology, Grahamstown.
Greenfield, D. W. and Matsuura, K. 2002. Scorpaenodes quadrispinosus: a new Indo-Pacificscorpionfish (Teleostei: Scorpaenidae). Copeia 2002: 973–978.
Hutchins, J. B. 2001. Checklist of the fishes of Western Australia. Records of the Western Australian Museum. Supplement 63: 9–50.
International Commission on Zoological Nomenclature 1999. International Code of Zoological
Nomenclature, Fourth Edition. International Trust for Zoological Nomenclature, London,

80

Hiroyuki Motomura et al.

xxix306 pp.
Ishida, M. 1997. Scorpaenidae. Pp. 189–209. In: Okamura, O. and Amaoka, K. (Eds) Nihon no
Kaisuigyo [Sea Fishes of Japan]. Yama-kei Publishers, Tokyo. [In Japanese]
Jordan, D. S. and Starks, E. C. 1904. A review of the scorpaenoid fishes of Japan. Proceedingsof the United States National Museum 27: 91–175.
Jordan, D. S. and Thompson, W. F. 1914. Record of the fishes obtained in Japan in 1911. Memoirs of the Carnegie Museum 6: 205–313.
Kim, I.-S., Choi, Y., Lee, C.-L., Lee, Y.-J., Kim, B.-J. and Kim, J.-H. 2005. Illustrated Book of Korean Fishes. Kyo-Hak Publishing, Seoul, 615 pp.
Mandrytsa, S. A. 2001. Seismosensornaia Sistema; Klassifikatsiia Skorpenovadnykh Ryb (Scorpaeniformes: Scorpaenoidei) [Lateral Line System and Classification of Scorpaenoid Fishes(Scorpaeniformes: Scorpaenoidei)]. Perm University Press, Perm, 393 pp. [In Russian]
Manilo, L. G. and Bogorodsky, S. V. 2003. Taxonomic composition, diversity and distributionof coastal fishes of the Arabian Sea. Journal of Ichthyology 43, Supplement 1: S75–S149.
Motomura, H. 2004a. New species of scorpionfish, Scorpaena cocosensis (Scorpaeniformes:
Scorpaenidae) from the Cocos Islands, Costa Rica, eastern Pacific Ocean. Copeia 2004:
818–824.
Motomura, H. 2004b. Revision of the scorpionfish genus Neosebastes (Scorpaeniformes:
Neosebastidae) with descriptions of five new species. Indo-Pacific Fishes 37: 1–76.
Motomura, H., Fricke, R. and Eschmeyer, W. N. 2005a. Redescription of a poorly known scorpionfish, Scorpaena canariensis (Sauvage), and a first record of Pontinus leda Eschmeyerfrom the Northern Hemisphere (Scorpaeniformes: Scorpaenidae). Stuttgarter Beiträge
zur Naturkunde. Serie A, Biologie 674: 1–15.
Motomura, H. and Iwatsuki, Y. 1997. A preliminary report of scorpaenid, synanceiid,
tetrarogid and aploactinid fishes in Miyazaki waters, southern Japan. Bulletin of theFaculty of Agriculture, Miyazaki University 44: 127–138.
Motomura, H. and Johnson, J. W. 2006. Validity of the poorly known scorpionfish, Rhinopias
eschmeyeri, with redescriptions of R. frondosa and R. aphanes (Scorpaeniformes: Scorpaenidae). Copeia 2006: 500–515.
Motomura, H., Last, P. R. and Gomon, M. F. 2006a. A new species of the scorpionfish genus
Maxillicosta from the southeast coast of Australia, with a redescription of M. whitleyi(Scorpaeniformes: Neosebastidae). Copeia 2006: 445–459.
Motomura, H., Last, P. R. and Yearsley, G. K. 2005b. Scorpaena bulacephala, a new species ofscorpionfish (Scorpaeniformes: Scorpaenidae) from the northern Tasman Sea. Zootaxa
1043: 17–32.
Motomura, H., Last, P. R. and Yearsley, G. K. 2006b. New species of shallow water scorpionfish (Scorpaenidae: Scorpaena) from the central coast of Western Australia. Copeia
2006: 360–369.
Motomura, H., Ogihara, G. and Hagiwara, K. 2010. Distributional range extension of a scorpionfish (Scorpaeniformes: Scorpaenidae), Scorpaenodes quadrispinosus, in the Indo-Pacific, and comments on synonymy of S. parvipinnis. Pp. 17–26. In: Motomura, H. and Matsuura, K. (Eds) Fishes of Yaku-shima Island — a World Heritage Island in the Osumi
Group, Kagoshima Prefecture, Southern Japan. National Museum of Nature and Science,
Tokyo.
Motomura, H., Paulin, C. D. and Stewart, A. L. 2005c. First records of Scorpaena onaria (Scorpaeniformes: Scorpaenidae) from the southwestern Pacific Ocean, and comparisons with
the Northern Hemisphere population. New Zealand Journal of Marine and Freshwater
Research 39: 865–880.

Validity of Scorpaenodes evides

81

Motomura, H. and Senou, H. 2008. A new species of the scorpionfish genus Scorpaena (Scorpaenidae) from Izu Peninsula, Pacific coast of Japan. Journal of Fish Biology 72:
1761–1772.
Motomura, H., Yoshino, T. and Takamura, N. 2004. Review of the scorpionfish genus Scorpaenopsis (Scorpaeniformes: Scorpaenidae) in Japanese waters with three new records
and an assessment of standard Japanese names. Japanese Journal of Ichthyology 51:
89–115.
Mundy, B. C. 2005. Checklist of the fishes of the Hawaiian Archipelago. Bishop Museum Bulletins in Zoology 6: 1–704.
Nakabo, T. 2002. Scorpaenidae. Scorpionfishes. Pp. 565–595, 1519–1522. In: Nakabo, T. (Ed.)
Fishes of Japan with Pictorial Keys to the Species, English Edition. Tokai University
Press, Tokyo.
Paulin, C., Stewart, A., Roberts, C. and McMillan, P. 2001. New Zealand Fishes. A Complete
Guide. Te Papa Press, Wellington, xiv279 pp.
Poss, S. G. 1999. Scorpaenidae. Pp. 2659–2756. In: Carpenter, K. E. and Niem, V. H. (Eds) FAO
Species Identification Guide for Fishery Purposes. The Living Marine Resources of theWestern Central Pacific. Vol. 4. Bony Fishes Part 2 (Mugilidae to Carangidae). FAO, Rome.
Randall, J. E. 2005. Reef and Shore Fishes of the South Pacific. New Caledonia to Tahiti and the
Pitcairn Islands. University of Hawai’i Press, Honolulu, xii707 pp.
Randall, J. E., Ida, H., Kato, K., Pyle, R. L. and Earle, J. L. 1997. Annotated checklist of the inshore fishes of the Ogasawara Islands. National Science Museum Monographs 11: 1–74.
Randall, J. E. and Eschmeyer, W. N. 2002 (dated 2001). Revision of the Indo-Pacific scorpionfish genus Scorpaenopsis, with descriptions of eight new species. Indo-Pacific Fishes
34: 1–79.
Senou, H., Matsuura, K. and Shinohara, G. 2006. Checklist of fishes in the Sagami Sea withzoogeographical comments on shallow water fishes occurring along the coastlines under
the influence of the Kuroshio Current. Memoirs of the National Science Museum, Tokyo
41: 389–542.
Senou, H., Shinohara, G., Matsuura, K., Furuse, K., Kato, S. and Kikuchi, T. 2002. Fishes ofHachijo-jima Island, Izu Islands group, Tokyo, Japan. Memoirs of the National Science
Museum, Tokyo 38: 195–237.
Shinohara, G., Sato, Y. and Matsuura, K. 2000. Coastal fishes of Ishima Island, Tokushima,
Japan. Memoirs of the National Science Museum, Tokyo 33: 175–186.
Tanaka, S. 1917. [Eleven new species of fish from Japan]. Zoological Magazine 29: 7–12. [InJapanese]
Xinbo, J. 2006. Fauna Sinica. Ostichthyes, Scorpaeniformes. Science Press, Beijing, xv739 pp.
Yoneda, M., Miura, H., Mitsuhashi, M., Matsuyama, M. and Matsuura, S. 2000. Sexual maturation, annual reproductive cycle, and spawning periodicity of the shore scorpionfish,
Scorpaenodes littoralis. Environmental Biology of Fishes 58: 307–319.