H. Motomura and K. Matsuura (eds) Fishes of Yaku-shima Island
National Museum of Nature and Science, Tokyo, 10 March 2010

Distributional range extension of a scorpionfish,
Scorpaenodes quadrispinosus, in the Indo–Pacific,
and comments on synonymy of S. parvipinnis(Scorpaeniformes: Scorpaenidae)
Hiroyuki Motomura1*, Gota Ogihara1 and Kiyoshi Hagiwara2
1

Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan
Yokosuka City Museum, 95 Fukadadai, Yokosuka, Kanagawa 238-0016, Japan*
Corresponding author: e-mail: motomura@kaum.kagoshima-u.ac.jp
2

Abstract Scorpaenodes quadrispinosus, previously known only from type specimens from Fiji and
the Marshall Islands, is reported here on the basis of 27 specimens from the eastern Indian Ocean(Christmas Island, Australia) and the Pacific Ocean (East Asia, Micronesia, Melanesia, Timor Sea,and Polynesia). Validity of diagnostic characters of S. quadrispinosus given in the original description is assessed and newly recognized diagnoses are provided. Paronescodes asperrimus is regarded
here as a junior synonym of S. parvipinnis.
Key words: Scorpaenidae, Scorpaenodes quadrispinosus, S. parvipinnis, Paronescodes asperrimus.

Introduction
The shallow water scorpionfish, Scorpaenodes quadrispinosus, was originally described by
Greenfield and Matsuura (2002) on the basis offive specimens (47.5–76.2 mm standard length)from Fiji (type locality) and the Marshall Islands.
Subsequently, Randall (2005) based his accountfor this species on the original description without
reference to additional specimens. No specimensof S. quadrispinosus have been reported since its
original description.
Ongoing taxonomic studies of Scorpaenodes
by the first author and colleagues (e.g., Motomura
et al., 2009) revealed that S. quadrispinosus, previously known only from Fiji and the MarshallIslands, is in fact widely distributed in the eastern
Indian Ocean and Pacific Ocean. The species isredescribed here on the basis of 27 new specimens, collected from Japan, Taiwan, Pohnpei,
Marshall Islands, Solomon Islands, Papua New
Guinea, Australia, and French Polynesia. The
validity of diagnostic characters of S. quadrispinosus given by Greenfield and Matsuura (2002)

is also assessed and newly recognized diagnostic
characters are provided.
Scorpaenodes parvipinnis, originally described as Scorpaena parvipinnis by Garrett(1864) on the basis of a single specimen (depositedat CAS, but destroyed by fire in 1906; Eschmeyer,1998) from the Hawaiian Islands, is closely related to S. quadrispinosus (see Greenfield andMatsuura, 2002; Randall, 2005). This similarity
is supported by the fact that most of our 27 specimens of S. quadrispinosus, except for newly collected specimens from Yaku-shima Island, Japan,
were registered as S. parvipinnis in museum collections.
Paronescodes asperrimus was originallydescribed as a new genus and species by Smith(1958) on the basis of 19 specimens reported as S.
parvipinnis by Smith (1957) from the Seychelles,
Mozambique, and Tanzania. The only publication
dealing with P. asperrimus since its original description is that of Eschmeyer (1986) in which it
was regarded as a junior synonym of S. parvipinnis without giving reasons or examined specimens.
17

H. Motomura et al.

In their description of Scorpaenodes quadrispinosus, Greenfield and Matsuura (2002) did not discuss Eschmeyer’s decision that P. asperrimus wasa junior synonym of S. parvipinnis. To clarify thissituation, we examined the 19 type specimens of P.
asperrimus in this study and confirmed the validity of S. quadrispinosus and taxonomic status of P.
asperrimus as a synonym.
Materials and methods
Counts and measurements follow Motomura
et al. (2005a–c) and Greenfield and Matsuura(2002) respectively. The last two soft rays of thedorsal and anal fins are counted as single rays,
each pair being associated with a single pterygiophore. Standard length is expressed as SL.
Terminology of head spines follow Randall andEschmeyer (2002: fig. 1) and Motomura (2004:
fig. 1), with the following additions: the spine occurring at the base of the uppermost preopercularspine is referred to as the supplemental preopercular spine (Eschmeyer, 1965); the spine occurring at the lateral surface of the lacrimal bone isreferred to as the lateral lacrimal spine (Motomura
and Senou, 2008: fig. 2); the coronal and pretympanic (as an extra spine) spines are as figured in
Chen (1981: fig. 1) and Motomura et al. (2004:
fig. 14b) respectively; and the longitudinal ridgeon the anterior portion of the lacrimal is referred
to as the lacrimal ridge. The formula for configuration of the supraneural bones, anterior neuralspines and anterior dorsal-fin pterygiophores follows Ahlstrom et al. (1976). The presence of aswimbladder was confirmed by dissection of theabdomen on the right side of KAUM–I. 11475.
The osteological characters were confirmed from
X-ray images taken of two specimens (YCM–P
34209, 34291). ‘Description’ is based only on
specimens examined in this study, and ‘Diagnosis’is based on this study and Greenfield and Matsuura (2002).
Specimens examined in this study have been
deposited in the California Academy of Sciences, San Francisco (CAS); Fish Division, Field
Museum of Natural History, Chicago (FMNH);
Kagoshima University Museum, Kagoshima(KAUM); Kanagawa Prefectural Museum of
Natural History, Odawara (KPM); South Afri18

can Institute for Aquatic Biodiversity, National
Research Foundation, Grahamstown (SAIAB);
Museum Support Center, Smithsonian Institute,
National Museum of Natural History, Suitland(USNM); Fish Department, Western AustralianMuseum of Natural Science, Welshpool (WAM);
and Yokosuka City Museum, Yokosuka (YCM).
Scorpaenodes quadrispinosus
Greenfield and Matsuura, 2002
[New English name: Furry Coral Scorpionfish][New Japanese name: Atsuhimesangokasago](Figs. 1–2; Table 1)
Scorpaenodes quadrispinosus Greenfield andMatsuura, 2002: 973, figs. 1, 2A (type locality: off Suva, Viti Levu Island, Fiji); Randall,
2005: 121 (color figure of Fijian specimen).
Material examined. 27 specimens, 27.3–99.8mm SL. JAPAN: KAUM–I. 11469, 74.7 mm
SL, Isso, Yaku-shima Island, Kagoshima, Japan,
30°27′23″N, 130°29′59″E, 0.1–5 m, KAUM
fish team,13 Aug. 2008; KAUM–I. 11475, 79.6mm SL, same data as KAUM–I. 11469; YCM–P
34209, 93.7 mm SL, Setouchi, Amami-oshimaIsland, Ryukyu Islands, Japan, 28 Aug. 1994;
YCM–P 34291, 99.8 mm SL, Hamazaki, Setouchi, Kakeroma Island, Amami-oshima Island,
Ryukyu Islands, Japan, 29 Aug. 1994. TAIWAN:
USNM 361037, 2 specimens, 62.3–64.9 mm
SL, Ch’uan-fan-shih, 21˚56′08″N, 120˚49′20″E,
5–6 m, V. Springer et al., 24 Apr. 1968; USNM
361042, 2 specimens, 58.5–81.2 mm SL, south ofChin-chiao-wan, 21˚55′15″N, 120˚49′45″E, 0–6m, V. Springer and J. Choat, 8 May 1968. POHNPEI: USNM 224495, 3 specimens, 33.7–81.7 mm
SL, Nanmatol, Ponape, 5˚51′N, 158˚20′E, 0–2m, V. Springer et al., 3 Sept. 1980. MARSHALLISLANDS: USNM 140488, 74.2 mm SL, Bikini
Atoll, between Bikini and Amen Islands, V. Brocket al., 21 July 1947; USNM 140489, 2 specimens,
55.8–65.3 mm SL, Bikini Atoll, Eman Island, L.
Schultz et al., 19 July 1947; USNM 140490, 64.0
mm SL, Bikini Atoll, Eman Island, L. Schultz etal., 17 July 1947; USNM 360997, 3 specimens,
72.1–74.4 mm SL, Bikar, Taka or Kwajalein
Atolls, A. Amerson, 15 Oct.–11 Nov. 1964. SOL-

Scorpaenodes quadrispinosus in the Indo-PacificA

B

Fig. 1. Fresh specimens of Scorpaenodes quadrispinosus from Japan. A, KAUM–I. 11475, 79.6 mm SL, Yaku-shima Island,
Kagoshima Prefecture; B, YCM–P 34209, 93.7 mm SL, Amami-oshima Island, Kagoshima Prefecture, photo by M.
Hayashi.

OMON ISLANDS: USNM 391125, 76.3 mm
SL, north end of Taumako Island, Duff Islands,
Santa Cruz Islands, 9˚51′00″S, 167˚09′30″E,
0–8 m, J. Williams et al., 21 Sept. 1998. PAPUANEW GUINEA: WAM P. 30624-028, 27.3 mm
SL, Madang, 5˚09′00″S, 145˚50′00″E, 9–12 m,
G. Allen et al., 28 Jan. 1993. AUSTRALIA:WAM P. 29061-007, 31.5 mm SL, Cartier Island,
Northern Territory, 12˚31′S, 123˚33′E, 10–12 m,
G. Allen, 20 Sept. 1986; WAM P. 29331-005, 2
specimens, 46.6–48.6 mm SL, Christmas Island,
10˚27′S, 105˚42′E, 0.1–3 m, G. Allen, 25 Feb.
1987. FRENCH POLYNESIA: USNM 379391,
4 specimens, 84.5–94.1 mm SL, east side of Isle
Karapoo rahi, Rapa, 27˚38′53″S, 144˚19′49″W,
0–4 m, J. Williams et al., 1 Nov. 2002.

Diagnosis. A species of Scorpaenodes withthe following combination of characters: 16–18(usually 17) pectoral-fin rays; 39–47 (mode 45)
scale rows in longitudinal series; 4 or 5 (5) gillrakers on upper limb, 7–9 (7) on lower limb,
11–13 (12) in total; anterior and posterior lacrimal
spines simple, poorly developed; dorsal lacrimalbone with 1 (rarely 2) small, dorsally directed
spine; no spine at lacrimal ridge; lateral lacrimal
spine simple, well developed; 3 suborbital spines,
first spine extending dorsally above ventral margin of orbit; an additional spine below second
suborbital spine; interorbital and coronal spines
usually present; no distinct black blotches on
opercle or subopercle.

19

H. Motomura et al.

Description. Proportional measurements, as
percentages of SL, are given in Table 1. Dorsal
fin with 13 spines and 9 soft rays (8 rays in 1
specimen); anal fin with 3 spines and 5 soft rays
(4 and 6 rays in 2 and 1 specimens respectively);
pectoral fin usually with 17 rays on each side ofbody (18 rays on each side in 3 specimens; asymmetrically 16 and 17 in 2 specimens, 17 and 18
in 4 specimens); 42–47 (mode 45) scale rows inlongitudinal series; 20–24 (23) pored lateral-line
scales; 4–8 (6) scales above lateral line, 18–25 (21)below; 4–7 (5) scale rows between last dorsalfin base and lateral line; 4 or 5 (5) gill rakers on
upper limb, 7–9 (7) on lower limb, 0 on hypobranchial, 11–13 (12) in total; 4 dorsal series of
caudal-procurrent rays, 4 ventral series; 2 dorsal
series of segmented, unbranched caudal-fin rays, 3ventral series; 6 dorsal series of branched caudalfin rays, 5 ventral series. Formula for configuration of anterior neural spines and anterior dorsal
pterygiophores //2+1/1/1/1/1/1/1/1/1/1+1/.
Body not strongly compressed anteriorly, progressively compressed posteriorly. Nape and anterior body not strongly arched. Body depth rela-

tively deep, less than head length. Numerous tiny,
slender tentacles on head and body. Short, broad
tentacle on posterior edge of low membranous
tube associated with anterior nostril. Pectoralfin axil without skin flaps. Scales covering dorsal
and lateral surfaces of head and mid-lateral surface of maxilla. Well-exposed ctenoid scales onlateral surface of trunk, scales becoming cycloid
on abdomen. Exposed cycloid scales (sometimes
with cteni) on anteroventral surface of body. Body
scales extending onto basal fin rays or membranes, except for bases of pelvic fins. Lateralline nearly straight from upper end of gill opening
to caudal-fin base. Underside of dentary without
distinct sensory pores on each side; pores covered
with skin. Pair of small pores, usually covered
with skin, behind symphysial knob of lower jaw
in ventral view.
Posterior margin of maxilla reaching or extending slightly beyond a vertical through posterior margin of orbit. No distinct longitudinal ridgeon lateral surface of maxilla. Width of symphysial
gap separating premaxillary teeth bands subequal
to width of each band. Upper and lower jaws with

Table1. Measurements, expressed as percentages of SL, of type (Greenfield and Matsuura, 2002) and non-type specimens(this study) of Scorpaenodes quadrispinosus. Means in parentheses.
This study
Greenfield and Matsuura (2002)
Non-types
Holotype
Paratypes
Indo-PacificFiji
Fiji and Marshall Islandsn = 27n=4Standard length (SL; mm)
27.3–99.8
75.0
47.5–76.2
Head length
40.6–45.2 (42.7)
37.7
35.8–43.4 (39.3)
Snout length
10.2–12.7 (11.2)
9.9
8.2–10.1 (9.2)
Orbit diameter
10.8–13.9 (12.7)
12.8
12.2–13.4 (12.7)
Interorbital width
6.8–8.7 (7.7)
8.0
7.0–8.4 (7.6)
Upper-jaw length
21.7–26.3 (23.8)
20.9
18.5–20.9 (19.3)
Predorsal-fin length
36.5–42.2 (39.1)
36.5
31.5–38.7 (34.7)
Body depth
31.9–37.9 (35.3)
35.1
32.7–35.1 (33.7)
Pectoral-fin length
25.7–31.3 (27.5)
26.7
26.6–29.1 (27.4)
Pelvic-fin length
21.1–27.3 (24.3)
23.2
25.4–27.4 (25.6)
Caudal-fin length
21.4–26.2 (24.3)
23.6
22.6–23.9 (23.2)
1st dorsal-fin spine length
3.9–6.1 (5.1)
4.5
4.2–5.9 (4.9)
2nd dorsal-fin spine length
5.7–8.9 (7.1)
7.1
6.1–8.7 (7.2)
3rd dorsal-fin spine length
7.5–10.6 (9.0)
9.7
7.0–10.0 (8.7)
4th dorsal-fin spine length
8.4–11.3 (9.8)
10.3
7.2–11.1 (9.7)
5th dorsal-fin spine length
7.8–11.6 (10.5)
10.8
9.3–11.1 (10.3)
6th dorsal-fin spine length
8.3–11.9 (10.5)
11.2
8.8–11.0 (10.0)
7th dorsal-fin spine length
8.9–11.9 (10.6)
10.4
9.1–11.5 (10.2)
8th dorsal-fin spine length
7.4–11.6 (10.1)
9.7
8.8–10.5 (9.9)
9th dorsal-fin spine length
7.1–11.5 (9.5)
8.5
8.0–9.5 (8.6)
10th dorsal-fin spine length
6.5–10.4 (8.6)
5.7
5.8–8.4 (7.0)
11th dorsal-fin spine length
4.1–8.4 (6.8)
4.7
3.4–6.5 (4.5)
12th dorsal-fin spine length
4.5–7.1 (5.6)
3.3
2.4–4.8 (3.5)
13th dorsal-fin spine length
8.0–10.4 (9.3)
7.7
7.8–9.9 (8.3)
1st anal-fin spine length
7.1–11.9 (9.5)
9.6
10.2–11.3 (10.4)
2nd anal-fin spine length
14.5–22.2 (18.2)
17.7
17.2–21.6 (18.7)
3rd anal-fin spine length
10.9–16.3 (14.0)
14.9
13.8–15.9 (14.8)
Longest anal-fin soft ray (2nd)17.2–22.7 (20.8)
20.8
21.1–23.1 (21.8)
Longest dorsal-fin soft ray (5th)15.5–18.0 (16.9)
16.8
17.0–17.4 (17.1)

20

Scorpaenodes quadrispinosus in the Indo-PacificFig. 2. Distributional recordsof Scorpaenodes quadrispinosus. ★ based on specimens examined inthis study; ● based on
Greenfield and Matsuura(2002).

villiform teeth, lacking canine teeth. Villiform
teeth forming V-shaped patch on vomer. Palatine
teeth absent. Underside of lower jaw withoutridges. Gill rakers relatively short and spinous;
slit present behind fourth gill arch. Swimbladderpresent, well developed.
Dorsal profile of snout steep, forming angleof 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 (sometimes absent), separated
by relatively shallow channel; interorbital spines
usually present at level of midline of eye. Interorbital space shallow, with about one-tenth oforbit extending above dorsal profile of head. Preocular spine simple, slightly flattened anteriorly
and posteriorly; anterior surface of spine without distinct ridges. Supraocular, postocular, andtympanic spines simple. Coronal spines usuallypresent (sometimes absent on one side of head)
between origins of tympanic spines. Pretympanic
spines absent. Occipital region flat, with no distinct ridges in front or rear of occiput. Parietal
and nuchal spines joined at base. Sphenotic withseveral small spines. Postorbital usually with tiny
spines. Pterotic spine simple, directed posteroven-trally. Upper and lower posttemporal spines present; length of upper spine about one-third of thatof lower spine. Supracleithral and cleithral spines
simple.
Lateral lacrimal spine simple, well developed.
Anterior and posterior lacrimal spines simple,
triangular or rounded, not strongly developed.
Dorsal lacrimal with 0–2 sharp spines directeddorsally. Suborbital ridge with 3 well developed
spines in a row; first spine canted dorsally, its tip
below posterior margin of pupil; second and third
spines behind orbit. An additional spine below
second 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
broader than that between first and second spines.
Preopercle, between uppermost preopercularspine and upper end of preopercle, without serrae or spines, and its margin completely covered
with scales. Upper opercular spine simple without median ridge. Lower opercular spine simple
with low median ridge. Space between upper andlower opercular spines without ridges. Posterior
tips of upper and lower opercular spines reachingand extending slightly beyond opercular marginrespectively.
Origin of first dorsal-fin spine above supra21

H. Motomura et al.

A

B

Fig. 3. Fresh specimens of Scorpaenodes parvipinnis from Japan. A, KAUM–I. 6601, 72.6 mm SL, Okinawa-jima Island,
Ryukyu Islands; B, KPM–NI 4855, 81.2 mm SL, Ie-jima Island, Ryukyu Islands, photo by H. Senou.

cleithral spine. Posterior margin of opercularmembrane reaching a vertical through base ofthird dorsal-fin spine. Posterior tip of pectoral fin
not reaching 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 not reaching origin of
anal fin. Origin of first anal-fin spine anterior to avertical through last dorsal-fin spine.
Color when fresh. Based on color photographsof KAUM and YCM specimens (Fig. 1). Body
strongly variegated, mainly from pale pink todense red, with 4 indistinct, vertical, dark bands
on lateral surface of body; first band above oper22

cle, second between middle of spinous portion of
dorsal fin and pectoral fin, third below ninth totwelfth spines of dorsal fin, and fourth below softrayed portion of dorsal fin. All fins, except forspinous portion of dorsal fin, pinkish to reddish,with small reddish to blackish spots scatteredon rays. Caudal-fin base with a vertical, reddish
band, its width less than pupil diameter. No distinct black blotches on opercle or subopercle.
Color of preserved specimens. Head and trunk
yellowish-white, mottled with black blotches;few black spots on ventral surface of body. Blackspots scattered on all fins, except for lower portion of pectoral fin.

Scorpaenodes quadrispinosus in the Indo-Pacific

Distribution and habitats. Currently known
from the eastern Indian Ocean (Christmas Island,
Australia) and the Pacific Ocean (East Asia,
Micronesia, Melanesia, Timor Sea, and Polynesia) (Fig. 2). The northernmost and easternmost
recorded ranges of the species are considered to
be Yaku-shima Island, Japan, and Rapa, French
Polynesia (Fig. 2). Yoshigou (2004: fig. 2) reported a single specimen of Scorpaenodes sp. from
Minamidaito-jima Island (ca. 400 km east of Okinawa Island), Japan, and gave an illustration of
the head of the specimen. His specimen is identified here as S. quadrispinosus. Collection data forthe species indicated capture depths less than 12
m. The species occurs on rocky reefs with strong
surge. Most specimens of S. quadrispinosus werecollected with S. parvipinnis.
Remarks. Characters of the present 27 specimens agreed with those of type specimens of S.
quadrispinosus given by Greenfield and Matsuura(2002). As Greenfield and Matsuura (2002) mentioned, S. quadrispinosus can be easily distinguished from all other congeners by the first suborbital spine (= Greenfield and Matsuura’s second
suborbital spine, their first spine is equivalent to
the lateral lacrimal spine in this study; see Motomura and Senou, 2008: fig. 2) extending dorsally above the ventral margin of the orbit (vs. spine
tip in line with suborbital ridge and not projecting
dorsally in the latter). Although some spines in
S. parvipinnis (Fig. 3) point dorsally, they do notreach the ventral margin of the orbit (Greenfieldand Matsuura, 2002: fig. 2; this study). Greenfield and Matsuura (2002) considered that thedorsally directed suborbital spine could possibly
be a synapomorphy uniting S. quadrispinosus
and S. parvipinnis. Detailed comparisons of S.
quadrispinosus with other congeners were given
in Greenfield and Matsuura (2002).
Greenfield and Matsuura (2002) distinguished
S. quadrispinosus from S. parvipinnis by lacking the dorsally directed spine on the lacrimal
ridge (vs. having the spine in the latter). However, examination of specimens of S. parvipinnis
in this study showed that they often lacked thespine from the lacrimal ridge, indicating that thecharacter is not always reliable for species’ discrimination. In addition, Greenfield and Matsuura(2002) mentioned four color pattern differences
between the two species; viz., S. quadrispinosus
is characterized by having four prominent dark
spots along the base of the spinous portion of thedorsal fin, a dark spot running between the basesof the parietal and tympanic spines, three rows ofdark spots on the caudal fin, and the general background color of cream with some pink overtones.
These color characters, however, vary considerably between individuals and cannot be used as
diagnoses. The density and pattern of colorationof S. quadrispinosus may be influenced by environmental conditions for protective mimicry or
camouflage.
Examination of specimens of S. quadrispinosus and S. parvipinnis during this study revealed
that S. quadrispinosus also differs from the latterin having lower counts of pectoral-fin rays (usually
17 vs. 18 in S. parvipinnis), scale rows in the longitudinal series [39–47 (mode 45) vs. 49–62 (52)],
scales above the lateral line [4–8 (6) vs. 9–11(10)], and scale rows between last dorsal-fin baseand lateral line [4–7 (5) vs. 7–8 (7)], and higher
counts of scales below the lateral line [18–25 (21)vs. 14–18 (15)], in addition to characters given
by Greenfield and Matsuura (2002) and Randall(2005), viz., three suborbital spines in S. quadrispinosus (vs. 5–15 spines in S. parvipinnis), andthe first suborbital spine extending dorsally above
the ventral margin of the orbit (vs. not reaching
the orbital margin).
A nominal species, Paronescodes asperrimus
Smith, 1958, was not mentioned in the originaldescription of S. quadrispinosus, although Eschmeyer (1986) regarded it as a junior synonymof S. parvipinnis, a close relative to S. quadrispinosus. Examination of all type specimens [holotype (Fig. 4) and 18 paratypes; listed below]of P. asperrimus in this study showed them to be
identical with S. parvipinnis; thus P. asperrimus isregarded here as a junior synonym of S. parvipinnis. Selected meristics of the type specimens of P.
asperrimus are as follows (data for holotype arepresented first, followed by data for paratypes inparentheses if different): Dorsal-fin rays XIII, 9;
anal-fin rays III, 5; pectoral fin rays 18 on eachside of body (18 on each side in 14 paratypes,
and 19 in a paratype; asymmetrically 17 and 18 in
23

H. Motomura et al.

Fig. 4. Holotype of Paronescodes asperrimus, SAIAB 112, 90.6 mm SL, La Digue Island, Seychelles.

2 paratypes, and 18 and 19 in 1 paratype); scalerows in longitudinal series 52 (49–55); pored
lateral-line scales 23; scales above lateral line 9(9–11), below 14 (14–18); scale rows between lastdorsal-fin base and lateral line 7 (7–8); scale rows
between sixth dorsal-fin base and lateral line 8(8–10); predorsal scale rows 11 (11–14); gill rakers on upper limb 4 (4–5), on lower limb 8 (7–8),
on hypobranchial 0, total 12 (11–13).
Comparative material examined. Typespecimens of Paronescodes asperrimus (=Scorpaenodes parvipinnis): SAIAB 112, holotype, 90.6 mm SL, La Digue Island, Seychelles,
04°22′00″S, 55°49′59″E, J. Smith and M. Smith;
SAIAB 377, paratype, 23.0 mm SL, Ibo Island,
Querimba Archipelago, Mozambique, 12°19′59″S,
40°37′00″E, Aug. 1951; SAIAB 378, 3 paratypes, 52.9–67.7 mm SL, Mafia Island, Tanzania,
07°50′60″S, 39°46′59″E, J. Smith, Nov. 1954;
SAIAB 379, paratype, 66.1 mm SL, Pinda Island,
Mozambique, 14°2167’S, 40°7667’E, J. Smith
and M. Smith, 1956; SAIAB 380, 4 paratypes,
60.5–77.8 mm SL, Assumption Island, Cosmoledo Islands, Seychelles, 09°45′00″S, 46°30′00″E,
J. Smith, Nov. 1954; SAIAB 381, paratype, 71.8mm SL, Aldabra Island, Seychelles, 09°25′59″S,
46°19′59″E, J. Smith and M. Smith, Nov. 1954;
SAIAB 382, paratype, 42.8 mm SL, Bazaruto
Island, Mozambique, 21°40′00″S, 35°30′00″E,
Sept. 1953; SAIAB 383, 2 paratypes, 65.1–
76.2 mm SL, Bazaruto Island, Mozambique,24

21°40′00″S, 35°30′00″E, Sept. 1953; SAIAB 384,
paratype, 72.3 mm SL, Zanzibar Island, Tanzania,
06°10′00″S, 39°10′59″E, Sept. 1952; SAIAB 385,
paratype, 58.3 mm SL, Zanzibar Island, Tanzania,
06°10′00″S, 39°10′59″E, Dec. 1957; SAIAB 386,
2 paratypes, 34.8–40.9 mm SL, collection data
unknown; SAIAB 387, paratype, 82.4 mm SL,
collection data unknown.
Type specimens of Scorpaenodes quadrispinosus: CAS 214498, holotype, 74.2 mm SL,
off Suva, Viti Levu Island, Fiji, 4.6–6.4 m, D.
Greenfield et al., 28 May 1999; FMNH 110236,
paratype, 47.8 mm SL, same data as CAS 214498;
USNM 367135, paratype, 76.8 mm SL, same data
as CAS 214498.
Scorpaenodes parvipinnis: FMNH 112040,
64.7 mm SL, Ashmore Reef, 10°09′31″S,144°35′26″E, FNQ team, 25 Jan. 1993; KAUM–
I. 255, 76.9 mm SL, south side of Shin-nittetsu
oil factory in Ishado, Nakagusuku, Nakagami,
Okinawa-jima Island, Ryukyu Islands, Japan,
26°17′44″N, 127°48′38″E, Y. Sakurai, 9 July
2006, washed up by Typhoon No. 3; KAUM–I.
256, 74.3 mm SL, same data as KAUM–I. 255;
KAUM–I. 6596, 77.2 mm SL, Nakagusuku-hama
Port, Hama, Nakagusuku, Nakagami, Okinawajima Island, Ryukyu Islands, Japan, 26°15′29″N,
127°47′38″E, Y. Sakurai, 13 July 2007, washed
up by Typhoon No. 4; KAUM–I. 6597, 72.2mm SL, same data as KAUM–I. 6596; KAUM–
I. 6598, 79.7 mm SL, same data as KAUM–I.

Scorpaenodes quadrispinosus in the Indo-Pacific6596; KAUM–I. 6599, 69.2 mm SL, same data
as KAUM–I. 6596; KAUM–I. 6600, 80.8 mm
SL, same data as KAUM–I. 6596; KAUM–I.
6601, 72.6 mm SL, same data as KAUM–I.
6596; KAUM–I. 6602, 75.4 mm SL, same data
as KAUM–I. 6596; KAUM–I. 7089, 60.8 mm
SL, same data as KAUM–I. 6596; KAUM–I.
20324, 57.9 mm SL, west of Kamazeno-hana,
Kurio, Yaku-shima Island, Kagoshima, Japan,
30°16′03″N, 130°24′48″E, 0–11 m, KAUM fish
team, 30 Oct. 2008; KAUM–I. 12818, 72.6 mm
SL, Sesoko Island, Okinawa-jima Island, Ryukyu
Islands, Japan, 26°38′N, 127°51′E, Y. Sakurai, 1Jan. 2008; KPM–NI 4855, 81.2 mm SL, Kananzaki, Ie-jima Island, Okinawa, Ryukyu Islands, 15
m, 6 June 1998; USNM 140483, 70.0 mm SL, Bikini Atoll, Namu Island, L. Schultz et al., 6 Aug.
1947; WAM P. 33056-008, 36.4 mm SL, Sekartaji, Nusa Penida, Indonesia, 08°48′S, 115°36′E,
15–20 m, G. Allen and M. Erdmann, Nov. 2008;
WAM P. 31522-005, 25.1 mm SL, Pulau Weh, Sumatra, Indonesia, 05°54′N, 95°13′E, 20–23 m, G.
Allen, 23 Jan. 1999; YCM–P 34221, 2 specimens,
78.7–94.0 mm SL, Hamazaki, Kakeroma Island,
Amami-oshima Island, Ryukyu Islands, Japan,
28°09′31″N, 129°11′00″E, 6–15 m, Sagami Bay
Marine Biological Research Club, 29 Aug. 1994;
YCM–P 36425, 87.0 mm SL, Saneku, Kakeroma
Island, Amami-oshima Island, Ryukyu Islands, Japan, 28°11′01″N, 129°15′32″E, 10–23 m, Sagami
Bay Marine Biological Research Club, 24 Aug.
1995.
Acknowledgments
The first author is especially grateful to W. Eschmeyer, T. Iwamoto, D. Catania and M. Hoang(CAS), M. A. Rogers and K. Swagel (FMNH), H.
Senou (KPM), O. Gon, W. Holleman, P. Heemstra, and E. Heemstra (SAIAB), J. Williams, L.
Palmer, S. Raredon and K. Murphy (USNM), S.
Morrison (WAM) for their kind hospitality during his stay at their institutions. The first author’s
visits to CAS, SAIAB, 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); to KPM
was supported by a Grant-in-Aid for Scientific
Research (A) from the Japan Society for the Pro-

motion of Science, Tokyo (JSPS; 19208019); to
FMNH was supported by Dispatch Grant to Overseas for Curators from MEXT; and to WAM was
supported by Travel Grant for Academic Meetingsfrom JSPS (211026). We thank Y. Haraguchi andstudents (KAUM) for their curatorial assistance,
T. Hashimoto (KAUM) for taking X-rays, and M.
McGrouther (Australian Museum, Sydney) for
reviewing the manuscript.
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