Ichthyol Res
DOI 10.1007/s10228-014-0451-6

FULL PAPER

Pterois paucispinula, a new species of lionfish (Scorpaenidae:
Pteroinae) from the western Pacific Ocean
Mizuki Matsunuma • Hiroyuki Motomura

Received: 5 September 2014 / Revised: 12 November 2014 / Accepted: 13 November 2014
Ó The Ichthyological Society of Japan 2014

Abstract A new species of lionfish (Scorpaenidae:
Pteroinae), Pterois paucispinula sp. nov., is described on
the basis of 37 specimens from the western Pacific Ocean.
The new species is closely related to and has been previously confused with Pterois mombasae (Smith 1957), both
species sharing usually XIII, 10 dorsal-fin rays, usuallymore than 18 pectoral-fin rays, numerous black blotches on
the pectoral-fin membrane, and several bands on the posterior portion of the pectoral-fin rays (free from membrane). However, P. paucispinula differs from similarly
sized P. mombasae in having a lesser body depth at theanal-fin origin, head width, postorbital length, and caudalpeduncle depth, in addition to a slightly higher number ofscale rows below the lateral line. Pterois paucispinula also
differs from the typical form of P. mombasae, occurring off
the east coast of Africa and in the central Indian Ocean andAndaman Sea, by having usually 18 pectoral-fin rays (vs.

This article was registered in the Official Register of Zoological
Nomenclature (ZooBank) as AB446EA3-8698-4527-8FC8F8B4437BC095.
This article was published as an Online First article on the online
publication date shown on this page. The article should be cited byusing the doi number.

Electronic supplementary material The online version of thisarticle (doi:10.1007/s10228-014-0451-6) contains supplementarymaterial, which is available to authorized users.
M. Matsunuma (&)
Seikai National Fisheries Research Institute, 1551-8 Taira,Nagasaki 851-2213, Japane-mail: k1139853@kadai.jp
H. Motomura
The Kagoshima University Museum, 1-21-30 Korimoto,Kagoshima 890-0065, Japane-mail: motomura@kaum.kagoshima-u.ac.jp

usually 19 in the latter) and relatively long dorsal-fin
spines, with the longest dorsal-fin spine length
42.9–51.7 % of standard length (SL) (vs. 35.1–44.8 % ofSL). Although a Sri Lankan population of P. mombasae issimilar to P. paucispinula in the above characters (usually
18 pectoral-fin rays and longest dorsal-fin spine length
37.5–51.1 % of SL), such differences among P. mombasae
are regarded as intra-specific geographical variations.
Additionally, young and adult P. mombasae have ctenoid
scales on the pectoral-fin base, ventrolateral portion of the
body (below the lateral line) and laterally on the caudal
peduncle. These regions in all examined P. paucispinula
usually have only cycloid or at most a few ctenoid scales,
thereby providing a consistent basis for identification ofboth species, including the Sri Lankan population of
P. mombasae. Pterois mombasae is distributed in the
Indian Ocean from the east coast of Africa to the Andaman
Sea, whereas P. paucispinula is recorded from the western
Pacific Ocean, from northern Australia to southern Japanand eastward to the Wallis and Futuna Islands.
Keywords Pterois mombasae  Pteropterus  Geographic
variation  Distribution

Introduction
The Indo-Pacific genus Pterois Oken 1817 (Scorpaenidae:
Pteroinae) has been characterized as having a combinationof three anal-fin spines, all pectoral-fin rays unbranched,
the parietal spine of males not elongated, and the mandiblelacking a spine or ridge (Eschmeyer and Randall 1975;
Poss 1999). Although Smith (1957) and Mandrytsa (2001,
2002) regarded Pteropterus Swainson 1839 as valid, being
distinguished from Pterois by several characters, including123M. Matsunuma, H. Motomura

pectoral-fin ray and body scale number, scale morphology
and eye position, a comprehensive phylogenetic study of
Pteroinae is yet to be made, and the relationships among
Pterois, Pteropterus and related genera remain unclear.
The treatment of Pteropterus as a junior synonym of
Pterois by most recent studies (e.g., Herre 1952; Eschmeyer and Randall 1975; Poss 1999) is tentatively followed here (see also Matsunuma and Motomura, 2011,2013b).
Pterois mombasae, originally described by Smith (1957)(as Pteropterus mombasae) from a single specimen from
Mombasa, Kenya, has since been widely recorded in the
Indo-West Pacific, ranging from the east coast of Africa to
Wallis and Futuna Islands, and from northern Australia tosouthern Japan (Eschmeyer 1986; Poss 1999; Matsunuma
and Motomura 2011; Motomura et al. 2011a; Allen andErdmann 2012). However, careful examination has recently
shown that western Pacific and Indian Ocean specimens canbe separated on the basis of meristics, morphometrics andscale morphology. The western Pacific specimens are
therefore described herein as a new species of Pterois.

Materials and methods
Measurements generally followed Motomura (2004b, c),with head width and maxillary depth following Motomura
et al. (2005b, 2006a) and Motomura et al. (2006b),
respectively. Body depth at the anal-fin origin and first to
ninth pectoral-fin ray length measurements followed
Matsunuma and Motomura (2013b). The length of the supraocular tentacle was taken as the distance from the flap
base to the extreme distal point on the flap. Counts generally followed Motomura et al. (2005a–c) and Motomura
and Johnson (2006), with counts of pre-dorsal-fin scalerows and scales above the lateral line following Motomura
et al. (2006b) and Matsunuma and Motomura (2013b),
respectively. The last two soft rays of the dorsal and anal
fins were counted as single rays, each pair being associatedwith a single pterygiophore. Counts and measurements
were made on the left side whenever possible, except forpectoral-fin rays (counted on both sides). The numbers ofscale spinules in each of the following regions were represented as the mean numbers of spinules counted on 3–10
scales (Fig. 1): dorsolateral scales (Fig. 1a), 4–6 scales
along the scale row between the first pored lateral line scale
and the dorsal-fin base, not including the small uppermost
scale; cheek scales (Fig. 1b), 3–6 scales along a more orless horizontal row, beginning from the second scale abovethat located at the posteroventral corner of the cheek(junction of suborbital and postorbital ridges), not including small anterior scales (much smaller than posteriorscales); preopercular scales (Fig. 1c), 4–10 scales along a123

horizontal scale row, beginning from the posteriormost
scale just below the upper posterior-most scale, notincluding small anterior scales; ventrolateral scales(Fig. 1d), 5–6 scales along a row below the pored lateralline scale below the last dorsal-fin spine base; caudalpeduncle scales (Fig. 1e), 4–6 scales surrounding the pored
lateral-line scale located at the approximate midportion of
the caudal peduncle; pectoral-fin base scales (Fig. 1f), 4–6
scales along a row on the pectoral girdle, beginning fromthe scale located at the most laterally expanded point on thegirdle and running posteroventrally.
The mitochondrial DNA (mtDNA) cytochrome b gene
was examined, and DNA extraction from a single specimenof P. paucispinula (KAUM–I. 48423, Okinawa Prefecture,
Japan) was performed with the Gentra Puregene Tissue Kit(QIAGEN) following the manufacturer’s protocols. Using
total DNA and the primer pair AJG15 (Akihito et al. 2000)
and Simt12-Fb (Kuriiwa et al. 2007), approximately 2.3 kbp
fragments, including the cytochrome b gene and control
region of mtDNA, were amplified under the following PCR
conditions. Thirty amplification cycles were performed at
94 °C for 45 sec, 56 °C for 30 sec and 72 °C for 2 min.
Sequencing reactions were carried out using the BigDye
Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems) with the primer L15369-CYB (Miya and Nishida2000), following the manufacturer’s protocols and utilizing
the automated genetic analyzer model 3500 (Applied Biosystems). The obtained DNA sequences were subjected tomultiple alignment using Clustal X (Thompson et al. 1997).
A sequence is available on DDBJ/EMBL/GenBank accession number AB986281. Comparative sequences of
P. mombasae from Kenya were also available (DDBJ/
EMBL/GenBank accession numbers AJ429427–429428)(Kochzius et al. 2003). The sequence difference (p-distance) was calculated using MEGA 6 (Tamura et al. 2013).
Head spine terminology generally followed Randall andEschmeyer (2002: fig. 1), Motomura (2004c: fig. 1), andMatsunuma and Motomura (2013b). Supplemental preopercular spine and lateral lacrimal spine terminology followed Eschmeyer (1965) and Motomura and Senou (2008:
fig. 2), respectively.
A principal component analysis (PCA) was conducted
for the following nine measurements considered to influence interspecific and intraspecific differences: body width,
head width, maxillary depth, postorbital length, sixth–
eighth and twelfth dorsal-fin spine lengths, and caudalpeduncle depth. All measurements were standardized bylog transformation and taken from the following 12 specimens of P. paucispinula, including the holotype, and 13
specimens of P. mombasae. P. paucispinula: AMS I.
37179-002 (Australia); BSKU 61061, BSKU 61062, BSKU
72569, BSKU 86402, BSKU 86404, CMNH–ZF 15450,
NSMT-P 110789 (two specimens), URM-P 4265, URM-PNew Pterois from western Pacific Ocean
Fig. 1 Positions of head andbody scales (shown as shaded)utilized for spinule counts inthis study (see Materials andmethods). a Dorsolateral scales;b cheek scales; c preopercularscales; d ventrolateralscales;e caudal-peduncle scales;
f pectoral-fin base scales.
Arrowheads in a, d ande indicate first pored lateral-line
scale, pored lateral-line scale
below last dorsal-fin spine baseand pored lateral-line scale at
midpoint of caudal peduncle,
respectively. Schematic
drawings of scales based on
Pterois mombasae, MNHN
1992-494, 112.7 mm SL

4266, URM-P 4268 (Japan); and P. mombasae: PMBC
(uncataloged; one of two specimens) (Andaman Sea coastof Thailand); CAS 75355 (India); AMS I. 33775-001 (three
specimens), SMF 4209, SMF 4322 (Sri Lanka); BPBM
36050 (Oman); SMF 10692, 12797, 12818 (Tanzania);
SMF 4210, MNHN 1992-494 (Madagascar). Damaged
specimens, including the holotype of P. mombasae, were
eliminated from the analysis, because they lacked one ormore aforementioned measurements, such as dorsal-fin
spine length. The first two components (PC1 and PC2)
were obtained using the software package R 3.1.0 (R CoreTeam 2014) based on the function prcomp.
A distributional map was prepared using Quantum GIS
2.2 (Quantum GIS Development Team 2014), with datafrom Natural Earth. Standard length is abbreviated as SL. In
the description, features of the holotype are presented first,
followed by paratype data (if different) in parentheses.
Institutional codes followed Sabaj Pérez (2014), with thefollowing addition: NSMT, National Museum of Natureand Science, Tsukuba.

Pterois paucispinula sp. nov.
(Japanese name: Mizuhiki-minokasago) (Figs. 2, 3a–d, 4,5, 6–10, 11a; Tables 1–3)

Pterois antennata (not of Bloch): Masuda et al. 1975:
339, pl. 144, fig. C (Japan; in part; short description);
Shimizu 1984: 302, pl. 282, fig. C [Japan; in part; figure
from Masuda et al. (1975); short description]; Randall et al.
1997: 79, unnumbered fig. (Coral Sea; short description);
Randall 2005: 116, unnumbered fig. [South Pacific; in part;
figure from Randall et al. (1997); short description]; Hirata
2010: 34, left unnumbered fig. (Ehime, Japan; in part).
Pterois mombasae (not of Smith): Gloerfelt-Tarp andKailola 1984: 112, unnumbered fig. (northwestern Australia; short description); Eschmeyer 1986: 467, fig. 149.9;
pl. 25, fig. 149.9 (Indo-West Pacific; in part; short
description); Allen and Cross 1989: 446 (northwest coast ofWestern Australia; list); Randall 1995: 109, unnumberedfig. (Indo-West Pacific; in part; short description); Kuiter
1996: 90, unnumbered fig. (Australia; short description);
Allen 1997: 76, pl. 19, fig. 4 (Indo-West Pacific; in part;short description); Poss 1999: 2330, unnumbered fig.
(Indo-West Pacific; in part; keys); Hutchins 2001: 27(Western Australia; list); Allen and Adrim 2003: 29 (Flores
to Sumatra, Indonesia; list); Allen et al. 2003: 370,
unnumbered fig. (Indo-West Pacific; in part; shot description); Allen et al. 2006: 879 (northwest coast of Western
Australia); Kuiter and Debelius 2006: 193, unnumberedfigs. (Indo-Pacific; in part); Fricke et al. 2011: 378 (NewCaledonia; list); Matsunuma and Motomura 2011: 27,

123

M. Matsunuma, H. Motomura
Fig. 2 Fresh specimens of
Pterois paucispinula sp. nov. atdifferent stages. a KAUM–I.
6588, 103.1 mm SL, paratype,
Okinawa-jima island, Ryukyu
Islands, Japan; bKAUM–I.
39268,98.3 mm SL, paratype,southwestern Taiwan (Photo:
H.-C. Ho); c BSKU 61062,
90.2 mm SL, holotype,Kashiwa-jima island, Kochi
Prefecture, Japan (Photo:
BSKU); d KAUM–I. 48423,
39.3 mm SL, paratype,
Okinawa-jima island, Ryukyu
Islands, Japan; e CMNH–ZF
15450, 30.9 mm SL, paratype,Yaku-shima island, Kagoshima
Prefecture, Japan (Photo: M.
Aizawa)figs. 1–2, 3A, 4 (Japan; description); Matsunuma et al.
2011: 3, figs. 1, 2A (Japan; description); Motomura et al.
2011b: 98, fig. 1 (Taiwan; description); Kubota et al.,
2011: 14 (Japan; list); Allen and Erdmann 2012: 221,
unnumbered fig. (Indonesia, Philippines, Papua New Guinea, and Solomon Islands; short description, ecologicalnote).
Holotype. BSKU 61062, 90.2 mm SL, Kashiwa-jimaIsland, Kochi Prefecture, Japan, S. Manabe, hand net, 21
November 2002.

123

Paratypes. Thirty-six specimens of 22.7–143.5 mm
SL. JAPAN: BSKU 61061, 82.3 mm SL, collected withholotype; BSKU 72569, 79.3 mm SL, Irino, Kochi Prefecture, bottom trawl, 16 January 2004; BSKU 86402,
59.4 mm SL, Saga, Kochi Prefecture, bottom trawl, 22
March 1999; BSKU 86404, 56.3 mm SL, Saga, Kochi
Prefecture, bottom trawl, 31 March 1999; BSKU 100149,
22.7 mm SL, Okino-shima Island, Kochi Prefecture, handnet, 25 September 2009; CMNH–ZF 15450, 30.9 mm SL,
Isso, Yaku-shima Island, Kagoshima Prefecture

New Pterois from western Pacific Ocean

Fig. 3 Preserved specimens of Pterois paucispinula sp. nov. (a–d) and Sri Lankan (e–h) and other (i–l) specimens of P. mombasae
showing differences in overall appearance between the two species
and within P. mombasae. a USNM 265918, 53.7 mm SL, paratype,
Macclesfield Bank, South China Sea; b AMS I. 39038-030, 59.5 mm
SL, paratype, Nemya Island, Solomon Islands; c URM-P 4268,
84.1 mm SL, paratype, Okinawa-jima islands, Ryukyu Islands, Japan;d AMS I. 37179-002, 102.1 mm SL, paratype, Timor Sea, Australia;
e SMF 10728, 47.5 mm SL, Sri Lanka; f AMS I. 33775-001 (one ofthree), 60.6 mm SL, Kalpitya, Sri Lanka; g SMF 10055, 84.5 mm SL,
Sri Lanka; h SMF 4322, 99.7 mm SL, Trincomalee, Sri Lanka;
i SAIAB 46315, 38.4 mm SL, Cathedral Reef, South Africa; j SMF
12793, 78.1 mm SL, Seychelles; k SMF 12797, 82.4 mm SL, Dar es
Salaam, Tanzania; l MNHN 1992-494, 112.7 mm SL, Madagascar(30°270 2800 N, 130°290 2100 E), hand net, 13 July 2006;
FAKU 26425, 143.5 mm SL, Makurazaki, Kagoshima
Prefecture, 2 September 1956; FAKU 98973, 40.0 mm
SL, FAKU 98976, 46.3 mm SL, FAKU 99044, 124.0 mm
SL, Shirahama, Wakayama Prefecture, washed ashore;
FAKU 99087, 89.1 mm SL, Shirahama, Wakayama Prefecture, washed ashore, 20 February 2011; KAUM–I.
6582, 82.2 mm SL, KAUM–I. 6588, 103.1 mm SL, Nakagusuku, Okinawa-jima Island, Ryukyu Islands, Y. Sakurai, washed ashore, 13 July 2007; KAUM–I. 22824,
52.3 mm SL, Okinawa-jima Island, Ryukyu Islands,
before October 2008; KAUM–I. 48423, 39.3 mm SL,
Zanpa, Yomitan, Okinawa-jima Island, Ryukyu Islands(26°260 0800 N, 127°420 45E), 1–3 m, K. Matsuzaki, handnet, 28 October 2011; KPM-NI 26919 (formerly IOP1103), 26.2 mm SL, Futo, Ito, Shizuoka Prefecture, H.

Masuda, 7 January 1990; KPM-NI 26930 (formerly IOP3210), 49.5 mm SL, Futo, Ito, Shizuoka Prefecture, H.
Masuda; NSMT-P 54352, 38.3 mm SL, Susaki, Chiba
Prefecture, 25 m, K. Utsuki; NSMT-P 110789 (formerly
KSHS 22471 and KSHS 22473), two specimens,
83.6–112.1 mm SL, Kashiwa-jima Island, Kochi Prefecture, 22 April 1995; URM-P 4264, 134.7 mm SL, URM-P
4265, 101.1 mm SL, URM-P 4266, 90.3 mm SL, URM-P
4267, 92.7 mm SL, URM-P 4268, 84.1 mm SL, URM-P
4269, 63.3 mm SL, west of Sesoko-jima Island, Okinawajima Islands, Ryukyu Islands, 14 April 1977; URM-P
41467, 73.6 mm SL, Okinawa-jima Island, Ryukyu
Islands, washed ashore, 17 October 2001.
TAIWAN: KAUM–I. 39268,98.3 mm SL, off southwestern Taiwan (obtained at Hsin-Da Port, Kaohsiung), H.C. Ho., 3 April 2011.

123

M. Matsunuma, H. Motomura

Fig. 4 Distributional map of Pterois paucispinula sp. nov. (stars) and P. mombasae (triangles) based on specimens examined in this study.
H indicates holotype

Fig. 5 Holotype of Pteropterus mombasae, SAIAB 117, 128.8 mm
SL, reef near Mombasa, Kenya (Photo: T. Yoshida)

PHILIPPINES: SMF 12808, 70.6 mm SL, 1972; USNM
99920, 58.4 mm SL, Observation Island, Tawi Tawi
Group, Sulu Archipelago (4°590 1000 N, 119°510 0000 E), 51 m,
R/V Albatross, 24 February 1908.
SOUTH CHINA SEA: USNM 265918, 53.7 mm SL,
MacclesfieldBank(16°030 4200 N,
114°400 0000 E),
82.3–84.1 m, W. Chan on board of R/V Cape St. Mary, 21
June 1964.
AUSTRALIA: AMS I. 37179-002, 102.1 mm SL, Timor Sea (10°070 4900 S, 129°190 5500 E), 85 m, J. Lloyd,
trammel net, 2–3 October 1995.
SOLOMON ISLANDS: AMS I. 39038-030, 59.5 mm
SL, mouth of Nemya Bay, just north of Nemya Island,
Sanra Cruz Islands, AMS Team; USNM 382904, 66.5 mm
SL, deep channel just north of Nemya Island at mouth of
Nemya Bay, Ndendo Island, Santa Cruz Islands

123(10°490 3000 S, 165°490 4800 E), 5–35 m, J. Williams et al., 28
September 1998.
WALLIS AND FUTUNA ISLANDS: MNHN
1995-737, 96.3 mm SL, Futuna Island (14°190 0500 S,
178°040 0500 E), 245–440 m, R/V Alis, trawl, 11 May 1992.
NO DATA: ZUMT 46873, 138.4 mm SL.
Diagnosis. A species of Pterois distinguished from otherspecies of the genus by the following combination ofcharacters: dorsal-fin rays XIII, 10 (rarely XIII, 11); pectoral-fin rays 17–19 (modally 18); scales below lateral line
11–15 (13); body depth at anal-fin origin 26.0–32.8 (mean30.2) % SL; head width 12.6–15.6 (14.4) % SL; postorbital
length 15.4–20.0 (18.1) % SL; longest dorsal-fin spine
length 42.9–51.7 (47.8) % SL; caudal-peduncle depth
9.9–11.9 (11.0) % SL; pectoral fin with 14–28 relatively
large (subequal to pupil diameter) black blotches onmembrane, several bands on posterior portion of rays (notattached to membrane); several narrow to wide verticalbands laterally on body and caudal peduncle; area of ctenoid scales on body relatively small (usually dorsolateralregion only at ca. 90 mm SL, sometimes extending below
lateral line onto ventrolateral region, and on pectoral-finbase and laterally on caudal peduncle in adults [80 mm
SL).
Description. Morphometrics and selected meristics areshown in Tables 1–3. Meristics given in Diagnosis are notrepeated here. Anal-fin rays III, 6; pelvic-fin rays with I, 5.
Branchiostegal rays 7.
Body oblong, moderately compressed posteriorly; depth
moderate, subequal to (or slightly less than) length of
longest dorsal-fin spine. Three barbels on snout tip, their

New Pterois from western Pacific Ocean
Table 1 Morphometrics, expressed as percentages of standard length, of Pterois paucispinula sp. nov. and P. mombasae
Pterois paucispinula sp. nov.

Locality

Holotype

Paratypes

Japan (BSKU
61062)

western Pacific(n = 36)

Pterois mombasae
Mean

Holotype

Other specimens

Kenya (SAIAB
117)

India and Sri Lanka(n = 13)

Meanother Indian Ocean(n = 19)

SL (mm)90.2

22.7–143.5

128.8

47.5–136.5

26.1–128.8

Body D (%
SL)

43.0

33.0–44.6

39.9

40.1

36.8–46.6

36.1–43.6

40.1

Body Da

32.8

26.0–32.5

30.2

33.5

31.7–34.8

27.4–35.2

32.2

Body W

28.0

18.2–29.3

25.2

26.6

22.8–30.1

20.1–32.3

27.0

Head L

45.1

39.9–46.5

43.1

45.5

40.7–44.9

41.6–47.1

43.6

Head W

14.0

12.6–15.6

14.4

15.2

14.2–16.1

14.4–16.4

15.2

Snout L

14.2

11.0–14.3

12.3

14.1

10.9–12.8

11.4–14.2

12.2

Orbit diameter

14.0

11.7–17.1

14.3

12.9

13.0–15.6

12.4–16.2

14.3

Interorbital Wb

7.8

7.2–10.1

8.4

9.9

7.0–8.9

7.0–9.9

8.1

Interorbital Wc
Upper-jaw L

7.5
20.6

6.6–9.4
18.0–22.3

8.0
20.2

9.4
21.1

6.7–9.4
18.5–21.7

5.3–9.6
19.1–22.4

8.0
20.4

Maxillary D

8.1

7.1–8.9

8.0

7.8

6.4–7.5

6.5–8.9

7.5

Postorbital L

18.5

15.4–20.0

18.1

20.0

18.7–20.9

17.7–21.3

19.6

Suborbital D

1.9

0.4–1.9

1.2

2.0

0.5–2.1

0.4–2.4

1.3

Pre-dorsal-fin
L

39.7

33.7–42.0

37.8

36.5

33.7–37.9

35.0–39.9

36.1

Pre-anal-fin L

75.5

70.5–79.7

75.1

76.2

70.4–81.1

71.3–76.2

74.0

Pre-pelvic-finL
1st DS L

42.6

35.2–43.2

39.7

47.4

36.3–41.2

37.7–47.4

39.6

16.6

14.4–19.7

17.0

14.8

18.4–23.6

14.8–21.2

18.9

2nd DS L

25.6

21.2–30.0

26.0

21.0

24.2–34.1

21.0–28.7

26.6

3rd DS L

30.7

27.6–36.8

31.7

broken27.3–37.8

23.3–32.7

30.4

4th DS L

35.3

31.4–43.0

36.7

broken31.8–41.6

26.2–35.1

33.8

5th DS L

40.0

34.4–48.2

42.2

broken38.6–46.0

30.7–38.3

37.8

6th DS L

42.4

37.9–54.1

45.5

32.0

35.1–49.0

32.0–42.1

39.9

7th DS L

42.5

39.2–53.8

46.5

33.9

37.1–51.1

33.8–45.0

41.8

8th DS L

43.5

41.6–51.5

46.8

broken37.5–50.6

35.1–44.2

42.0

9th DS L

42.7

39.0–51.7

45.7

33.5

41.9–50.0

33.5–44.0

41.2

10th DS L

38.1

33.2–48.9

41.8

30.9

34.5–46.2

29.2–42.2

38.3

11th DS L

28.9

16.8–45.4

29.5

26.7

24.4–38.7

17.9–35.5

29.9

12th DS L

12.2

10.2–18.5

13.4

12.4

12.9–18.2

12.4–22.9

16.0

13th DS L

13.9

13.1–17.8

15.1

12.3

12.4–18.6

12.3–19.2

15.4

Longest DS L

43.7

42.9–51.7

47.8

unknown

37.5–51.1

35.1–44.8

42.1

1st DSR L
Longest DSR
L

20.5
27.8

18.8–26.2
26.5–33.0

22.8
29.5

brokenbroken19.6–25.9
24.6–31.8

17.3–26.4
24.3–32.9

22.4
27.4

1st AS L

6.9

6.3–11.0

8.3

6.3

7.0–10.8

6.3–9.8

8.9

2nd AS L

14.0

13.0–19.3

15.5

10.5

10.5–19.1

10.5–19.0

15.4

3rd AS L

18.1

14.5–22.6

18.4

14.4

15.7–22.5

14.4–22.3

18.5

1st ASR L

26.4

25.0–31.7

28.4

broken24.9–33.3

22.4–31.1

27.6

Longest ASR
L

28.4

28.7–34.3

31.0

broken27.2–35.7

26.1–32.6

30.1

1st P1 L
2nd P1 L

66.3
71.1

49.0–87.6
59.6–85.4

67.9
74.8

brokenbroken43.5–82.0
68.6–88.9

57.2–76.6
65.7–82.0

66.7
75.9

3rd P1 L

72.0

63.1–89.6

75.6

broken58.6–90.1

68.5–79.5

77.3

123

M. Matsunuma, H. Motomura
Table 1 continued
Pterois paucispinula sp. nov.
Holotype

Paratypes

Locality

Japan (BSKU
61062)

western Pacific(n = 36)

4th P1 L

76.6

65.3–89.3

5th P1 L

76.3

58.2–87.1

6th P1 L

73.2

7th P1 L

Pterois mombasae
Mean

Holotype

Other specimens

Mean

Kenya (SAIAB
117)

India and Sri Lanka(n = 13)other Indian Ocean(n = 19)76.2

broken64.8–90.7

70.5–82.5

77.5

74.9

broken70.6–88.0

69.1–83.2

76.4

60.7–85.2

73.4

broken70.1–87.8

66.6–81.9

76.0

69.8

56.1–84.8

71.8

52.7

62.1–86.7

52.7–79.7

71.6

8th P1 L

68.6

56.3–82.5

69.1

53.7

69.3–83.2

53.7–76.6

70.0

9th P1 L
Longest P1 L

62.2
76.6

55.1–81.6
65.7–89.3

65.5
77.2

brokenbroken61.1–73.0
76.2–90.7

57.2–73.1
71.4–83.2

66.3
74.3

P2S L

19.5

17.6–23.9

21.1

17.5

17.0–23.4

17.5–23.1

20.3

Longest P2SR
L

46.0

36.4–53.9

45.0

broken40.1–52.5

38.2–52.4

45.9

Caudal-fin L

34.8

32.6–42.4

37.3

broken32.0–41.4

32.5–43.0

36.9

Caudalpeduncle L

16.1

13.8–17.8

15.9

16.5

14.1–17.2

13.4–16.9

15.7

Caudalpeduncle D

11.1

10.2–11.9

11.0

11.6

11.1–13.0

11.0–12.8

11.9

AS anal-fin spine; ASR anal-fin soft ray; D depth; DS dorsal-fin spine; DSR dorsal-fin soft ray; L length; P1 pectoral-fin ray; P2S pelvic-fin spine;
P2SR pelvic-fin soft ray, W widthaAt the anal-fin originb

At the vertical midline of the eyecAt the posterior end of the preocular spine baselength subequal to anterior nasal tentacle length. Shortsimple tentacle on posterior edge of low, membranous tube
associated with anterior nostril, its tip extending beyond
posterior margin of posterior nostril when depressed posteriorly. Supraocular with a low skin flap (long tentaclewith 2–4 lateral branches in juveniles; low skin flap in large
adults). Anteroventral portion of lacrimal with a shorttentacle, its length subequal to that of barbels on snout tip.
A relatively large fan-like skin flap on tip of posterior
lacrimal spine, its tip extending slightly beyond posterior
margin of maxilla when laid flat. Two small fan-like flaps
on posterior margin of preopercle, their tips just reachingor extending slightly beyond posterior margin of interopercle when laid flat. Small fan-like flap usually present onventral portion of third preopercular spine in large
adults [ ca. 130 mm SL (absent in juveniles). A small skinflap anterodorsally on orbit surface, its length slightly
greater than diameter of posterior nasal pore. No other skinflaps on head or body.
Ctenoid scales covering preopercle, suborbital area
(exposed or embedded cycloid scales in juveniles \ ca.
30 mm SL), cheek, suprapostorbital region between
tympanic and sphenotic spine bases, occipital region(exposed or embedded cycloid scales in juveniles) andanterodorsal portion of opercle. Dorsal surface of123

postocular and frontal portion of preocular with mostly
cycloid scales, sometimes with weakly ctenoid scalespossessing 1–4 spines (cycloid scales only or scales
absent in juveniles). Scales generally absent from middorsal interorbital area, interorbital canal (sometimes arow of small cycloid scales in adults [ ca. 100 mm SL),
snout, lacrimal (several small scales, mostly cycloid but afew poorly developed ctenoid scales possessing 1–4
spines, on posterior lacrimal spine plate in largest
adults [ ca. 130 mm SL), maxilla [ZUMT 46873,
138.4 mm SL (largest examined specimen) with a patchof small cycloid scales on posterodorsal corner], lips,
mandibular and interopercle. Other regions of head covered with cycloid scales [all head scales cycloid in BSKU
100149, 22.7 mm SL (smallest examined specimen)].
Anterodorsal portion of body mostly above lateral line
and anterior to level of last dorsal-fin spine base, with
ctenoid scales; remainder of body with cycloid scales (allscales cycloid in BSKU 100149, area of body with ctenoid scales increasing with increasing SL). Soft-rayed
portions of dorsal and anal fins, pectoral-fin and caudal-finbases with small cycloid basal scales.
Mouth moderately large, slightly oblique, forming angleof ca. 30° (30–40°) to horizontal axis of head and body;
upper edge of posterior maxilla swollen laterally, formingNew Pterois from western Pacific Ocean
Table 3 Frequency distribution of selected meristics in Pterois paucispinula sp. nov. andP. mombasae, showing interspecific differencesand geographical variations in P. mombasae
Pectoral-fin rays

P. paucispinula

17/17

17/18

18/18

18/19

19/19

1

2

28H

2

4

1

9

2

19/20

20/20

P. mombasae
Sri Lanka and northern India
Andaman Sea (Thailand)1

Maldives and Seychelles

2

Oman

2

east Africa and Madagascar

4

P. paucispinula

1

9H a

1

1

Scales between last DS and LL

Scales between 6th DS and LL

Pre-dorsal-fin scale rows

6

7

8

6

7

8

4

5

6

7

8

4

18

8H

3

5

11H

4

8

12H

6

1

4

7

2

3

6

2

9

9

P. mombasae
Sri Lanka and northern India
Andaman Sea (Thailand)

1

2

8

1

1

Maldives and Seychelles

1

Oman

1

1

east Africa and Madagascar

5

5

2

2

2

1

2

1
2

7

1

3

2

1

1

4

2

2

DS and LL indicate dorsal-fin base and lateral line, respectivelyH

includes holotypeafrom original description of Pteropterus mombasae; right pectoral fin of holotype removed

low ridge; posterior margin of maxilla just reaching midorbit level. Symphyseal gap separating premaxillary teeth
bands distinctly broader than width of each band; upper
jaw with band of small, slender conical teeth; ca. 8 (8–10)
tooth rows at front of upper jaw; ca. 3–5 tooth rows at frontof lower jaw; small teeth in ca. 6 (6–15) rows forming
blunt V-shaped patch on vomer; no palatine teeth. Three
sensory pores on underside of each dentary, one small poreon each side of symphysial knob.
Dorsal profile of snout relatively steep, forming angle ofca. 40° (30–40°) to horizontal axis of head and body. Nasal
spine with 1 (1 or 2) spinous point. Preocular spine with 2(1–4) spinous points directed upward. Supraocular with 1
spinous point covered by skin. Postocular with 7 (1–13)
small spines directed laterally. Interorbital ridge notdeveloped, reduced posteriorly, connected to coronal spine
base. Coronal spine with a single point. Tympanic spinewith a single point (absent in BSKU 100149). Anterior
margin of occipital region sloped transversely frombetween bases of coronal spines, slightly curved posteriorly
in dorsal view. Parietal spine with 2 (0–4) spinous points;
its base relatively long, diverging posteriorly and notconnected to coronal spine base. Nuchal spine with 1 (1–3)
spinous point, its base completely conjoined with that ofparietal spine. Postorbital spine and exposed sensory canalabsent. Sphenotic with 5 (0–10) small spines. Pterotic with
3 (0–7) small spines. Lower post-temporal spine with asingle (1 or 2) small spine. Cleithrum with short upper andlong lower ridges; upper ridge with a single (0–2) somewhat flattened spine posteriorly; lower ridge with 2 (1–4)
spinous points posteriorly (single ridge without spine in
BSKU 100149).
Lateral lacrimal spine with 3 (0–3) spinous points (upperand lower ridges with 2–3 and 1–4 spines in large
adults [ ca. 130 mm SL); other ridges on lacrimal mostly
smooth (strongly spinous in large adults). Suborbital withupper main and lower supplemental spines; upper ridgewith 11 (2–18) small spines; lower spines 8 (0–14) (ridge
only without spines in BSKU 100149). Anterior lacrimal
spine absent. Posterior lacrimal spine broad, plate-like,with a single (1–5) spinous point on distal margin, tipcovered by skin (if more than two spines present, all spinous points exposed, except for a point situated at posterior
lacrimal skin flap base). Preopercle with 3 distinct similarlength spines; uppermost spine directed posterodorsally,
middle spine directed posteroventrally, lowermost spine
directed posteriorly; spine bases lacking supplemental
spines (1 or 2 supplemental spines in ZUMT 46873,

123

M. Matsunuma, H. Motomura

138.4 mm SL); lowermost spine broad, plate-like. Exposed
opercular spine absent.
Origin of first dorsal-fin spine above lower post-temporal spine; bases of first and second dorsal-fin spines
closer than those of subsequent adjacent spines; eighth(sometimes seventh or ninth) spine longest; sixth to ninth
spines almost same length; twelfth [rarely thirteenth (last)]
spine shortest, its length 42 % (36–73 %) and 88 %(64–107 %) that of antepenultimate and posteriormost
spines, respectively; membrane of spinous portion of dorsal
fin strongly incised. Dorsal-fin soft rays all branched; fifth(usually fourth or fifth, sometimes sixth, rarely third) ray
longest, but distinctly shorter than longest dorsal-fin spine;posteriormost ray joined by extremely low membrane tocaudal peduncle. Origin of first anal-fin spine below posteriormost dorsal-fin spine base (sometimes below firstsoft-ray base); third spine longest; length of first spine
49 % (45–63 %) and 38 % (38–60 %) of that of second
and third spines, respectively. Anal-fin soft rays all branched; third (sometimes second) ray longest, its length
subequal to that of longest dorsal-fin soft ray; posteriormost
ray free from caudal peduncle. Pectoral fin relatively long,
all rays unbranched, lower 7 (7–8) rays weakly thickened;
fourth (variably second to seventh) ray longest, tip of
longest ray extending beyond caudal-fin base, not reaching
posterior margin of caudal fin. Pelvic-fin spine base below
third (or fourth) dorsal-fin spine base; all pelvic-fin softrays branched; third soft ray longest, its tip extending
slightly beyond third (second to fifth) anal-fin soft ray basewhen depressed, not reaching posteriormost anal-fin softray base (tip of longest ray just reaching first anal-fin spinebase in URM-P 4265); posteriormost soft ray withmembranous connection to abdomen for ca. one-fourth of
ray length. Caudal fin rounded, with 3 procurrent rays, 2segmented unbranched rays and 5 segmented branched rays
in dorsal series, and 3 procurrent rays, 3 segmented
unbranched rays and 4 segmented branched rays in ventral
series (usually 3 procurrent, 2 segmented unbranched and 5
segmented branched rays in both series; rarely with 4
procurrent rays in ventral series). Caudal peduncle moderately short, deep, its depth 69 % (63–88 %) of caudalpeduncle length.
Color of fresh specimens (based on Fig. 2). Groundcolor of head and body pinkish creamy-white (or white).
A poorly defined dark red (or light brown) band on snout
reaching from anteroventral portion of eye to anteroventral margin of lacrimal. A poorly defined dark red (or lightbrown) band with narrow white margin extending frombelow eye, across posterodorsal corner of maxilla toposteroventral margin of preopercle. A relatively broad(broadest on head) reddish-black (or black) band withnarrow white margin from supraocular tentacle base,obliquely crossing eye and reaching posterior margin of
preopercle (uppermost and middle preopercular spine
bases); indistinct connection with a large black blotch(slightly smaller than eye) on subopercle. Three bands
saddling nape; first band reddish-black (or black), justbehind eye, at level between coronal spine and origin ofparietal spine; second band at level of parietal spine,reaching central posterior margin of opercle at level ofupper origin of pectoral-fin base; third band at level
between nuchal spine and dorsal-fin origin, reaching upper
origin of gill opening. Numerous (ca. 18) reddish-brown(or pale red to light brown) bands on side of body, narrow

Fig. 6 Comparisons of dorsal-fin spine length (as % of standard
length) between Pterois paucispinula sp. nov. and Sri Lankan andother specimens of P. mombasae. Symbols indicate means of eachdorsal-fin spine length (stars, P. paucispinula; circles, Sri Lankan
population of P. mombasae; triangles, other P. mombasae). Verticalbars indicate ranges of each dorsal-fin spine length (top and bottom
indicate maximum and minimum values, respectively). Polynomial
trendlines for means of each dorsal-fin spine length (solid line,
P. paucispinula; dashed line, Sri Lankan population of P. mombasae;
dotted line, other P. mombasae)

123

New Pterois from western Pacific Ocean

bands inserted between each relatively broad band runningtransverse and obliquely in anterior and posterior portionsof trunk region; a band below fourth to sixth dorsal-fin
spine bases broadest; caudal peduncle with ca. 6 (2–6)
poorly defined narrow bands running obliquely. Body
bands extending slightly onto dorsal-fin rays and membrane; more so onto anal-fin, a posteriormost trunk band
reaching from penultimate dorsal-fin soft ray base to
middle of third anal-fin spine. A moderately large black
blotch (slightly smaller than iris) situated just above
pectoral-fin base, partly (partly or mostly) behind pectoralfin, with a small white inner spot (ca. one-fourth to onethird of iris diameter). Two (or one) white margined
reddish bands on chest. Dorsal-fin membrane and soft rays
translucent, spines creamy-white; 2–6 reddish-brown (or,pale red to light brown) (lower portion) to black (upper
portion) bands on each dorsal-fin spine; ca. 70 (37–74)
small black spots on soft rays [juveniles (CMNH-ZF
15450, 30.9 mm SL and KAUM–I. 48423, 39.3 mm SL)with numerous small white spots on soft rays between
each black spot]. Anal-fin membrane and rays translucent;ca. 44 (26–50) small black spots on soft rays (juveniles
with numerous small white spots on soft rays between
each black spot). Pectoral-fin membrane translucent (orpale red), a poorly defined narrow black line marginally,
and a poorly defined broad black band basally on innersurface; ca. 21 (14–28) relatively large reddish-black (toblack) blotches (subequal to pupil diameter) on membraneassociated with non-thickened rays (first to tenth oreleventh rays); 2–3 (1–4) relatively small black blotches(slightly smaller than half pupil diameter) on each membrane between thickened rays (below eleventh ray); posterior portion of rays (free from membrane) creamy-white(or somewhat yellowish to reddish) with numerous brown
to reddish-black bands [ca. 10 (5–13) bands on first ray];pectoral-fin base with 2 white marginal dark red bands.
Pelvic-fin membrane dark reddish, becoming gradually
blackish toward base; rays creamy-white. Caudal-fin
membrane translucent with ca. 86 (48–98) small blackspots on ray (juveniles with numerous small white spots
on soft rays marginally between each black spot). Imagesof live specimens (KPM-NR 1425, 25926, 35565,
62458–62460, 63160, and 82854) registered in the Image
Database of Fishes in the Kanagawa Prefectural Museumof Natural History (KPM-NR; http://fishpix.kahaku.go.jp/
fishimage-e/index.html).
Color of preserved specimens (Fig. 3a–d). Head andbody creamy-white with brown to black markings; ca. 70(26–71) small spots on soft-rayed portion of dorsal fin, 44(26–47) on that of anal fin, 86 (41–96) on caudal fin.
Distribution. Pterois paucispinula is distributed in thewestern Pacific Ocean from northern Australia (Timor Sea)
to southern Japan; eastwards to Wallis and Futuna Islands(Fig. 4). Collection depth data from seven specimens
indicated a depth range of P. paucispinula between 1 and440 m. However, the deepest collected specimen (MNHN
1995-737, 96.3 mm SL, collected by trawl from off Wallis
and Futuna Islands in depths of 245–440 m) was most
likely taken in a shallow water layer. The species has been
photographed underwater several times (KPM-NR) at
Pacific coast locations in central Japan (Matsunuma andMotomura 2011).
Etymology. The specific name paucispinula is derived
from Latin for ‘‘few spines’’ in reference to the small
number of spinules on the head and body ctenoid scales, incontrast to its closely related congener, P. mombasae.

Pterois mombasae (Smith 1957)
Pteropterus mombasae Smith 1957: 80, fig. 7, pl. 6, fig. D(type locality: Mombasa, Kenya); Mandrytsa 2001: 16(phylogenetic material).
Pterois antennata (not of Bloch): Jones and Kumaran
1980: 637, fig. 544 (Kavarathi, Laccadive Islands, India;
short description).
Pterois mombasae: Kotthaus 1979: 21, fig. 473 (Gulf ofAden and off Somalia; short description); Poss and RamaRao 1983: 24, unnumbered fig. (Sri Lanka, India, east coastof Africa; sort description, ecological note); Eschmeyer
1986: 467, fig. 149.9; pl. 25, fig. 149.9 (Indo-West Pacific;
in part; short description); Randall 1995: 109, unnumberedfig. (Indo-West Pacific; in part; short description); Allen
1997: 76, pl. 19, fig. 4 (Indo-West Pacific; in part; short
description); Poss 1999: 2330, unnumbered fig. (Indo-West
Pacific; in part; keys); Allen et al. 2003: 370, unnumberedfig. (Indo-West Pacific; in part; short description); Kochzius et al. 2003: 396 (Kenya; molecular analysis material);
Kuiter and Debelius 2006: 193, unnumbered figs. (IndoPacific; in part; figures taken in Indonesia); Motomura
2009: 65, unnumbered fig. (Andaman Sea; short description); Satapoomin 2011: 53 (Andaman Sea; list).
Holotype. SAIAB117, 128.8 mm SL, reef near Mombasa, Kenya.
Other specimens examined. Thirty-two specimens of
26.1–136.5 mm SL. THAILAND (ANDAMAN SEA):PMBC (uncatalogued), two specimens, 70.3–75.0 mm SL,
Phuket.
SRI LANKA: AMS I. 33775-001, three specimens,
53.5–60.6 mm SL, Kalpitya (8.22N, 79.82E), P. Pethiyagodo; CAS 75358, 102.0 mm SL, Trincomalee(8°330 2400 N, 81°140 4200 E), 5–15 feet (ca. 1.5–4.6 m), T.
Iwamoto, 17 May 1970; FMNH 75819, 83.4 mm SL, L.
Woods et al., 4 March 1964; SMF 4209, 60.1 mm SL, 10
May 1957; SMF 4322, 99.7 mm SL, Trincomalee, R.
Jonklaas, 21 May 1958; SMF 10055, three specimens,123

M. Matsunuma, H. Motomura
Table 2 Frequency distribution of selected meristics in Pterois paucispinula sp. nov. and P. mombasae

P. paucispinula
P. mombasae

Dorsal-fin rays

Scale rows in longitudinal series

XIII, 10

XIII, 11

43

35H

2

H

30

3

44

45

1

1

1

2

Pored lateral-line scales

P. paucispinula
P. mombasae

P. paucispinula
P. mombasae
H

24

25

2

5H

2

17

46

48

49

50

51

3

2

2

14H

2

3

6

1

H

3

3

Scales above lateral line

26

27

3

2

6

7
8H

H

H

2

6

Upper gill rakers

Lower gill rakers

4

5

9

10

11

21

14H

1

20

14H

18

H

21

H

15

47

52

53

54

2

2

3

Scales below lateral line

8

9

11

12

16

5

1

5

17

4

H

4

17

13

14

15

12

6

2H

3

3

Total gill rakers
12

11

1

13

14

15

16

1

13

15

6H

10

16

4H

includes holotype

84.5–108.2 mm SL, 1958; SMF 10728, 47.5 mm SL, 1968;
SMF 11887, 75.8 mm SL, Trincomalee, October 1972.
INDIA: CAS 75355, 136.5 mm SL, Wadge Bank
(8°010 6000 N, 77°120 0000 E), 27–33 fathoms (ca. 49–60 m), C.
Koenig, 23 December 1969.
MALDIVES: BMNH 1901.12.31.26, 48.7 mm SL.
SEYCHELLES: SMF 12793, 78.1 mm SL, August
1969.
OMAN: BPBM 34447, 115.3 mm SL, Eagles’ Retreat,near Mirbat (16°580 0000 N, 54°420 5000 E), 6–8 m, J. Randall
and J. Mee, rotenone, 25 April 1990; BPBM 36050,
134.3 mm SL, off southwest side of Sawda Island, 21 m, J.
Randall et al., rotenone, 7 November 1993.
KENYA: SMF 10056, 87.0 mm SL, Mombasa, 21
February 1966.
TANZANIA: SMF 10692, 128.0 mm SL, Dar es Salaam, 7 August 1969; SMF 12797, 82.4 mm SL, Dar es
Salaam, 7 August 1969; SMF 12818, 111.1 mm SL, Dar es
Salaam, 1973.
MOZAMBIQUE: CAS 65996, 53.7 mm SL, southeastof Beira, 3 October 1964; SAIAB 82480, 81.6 mm SL, off
Mozambique (16°23.40 S, 40°020 E), P. Heemstra and E.
Heemstra, bottom trawl, 4 November 2007.
MADAGASCAR: MNHN 1992-494, 112.7 mm SL, 16
August 1972; SMF 4210, 79.2 mm SL, Nosy Be (13°150 S,
48°150 E), February 1922.
SOUTH AFRICA: SAIAB 10741, 102.4 mm SL, off
Durban, Oceanographic Research Institute; SAIAB 40285,
two specimens, 26.1–27.6 mm SL, 2 km south of Kosi Bay
mouth (26°530 S, 32°520 E), C. Buxton and P. Heemstra, 9August 1992; SAIAB 46315, 38.4 mm SL, Aliwal Shoal,
near Cathedral Reef (30°250 S, 30°490 E), C. Buxton and P.
Heemstra, 19 June 1994; SAIAB 58553, 37.2 mm SL,

123

Aliwal Shoal, near Hospital Road (30°180 S, 30°490 E), A.
Bentley and P. Heemstra, 14 June 1998.
Diagnosis. A species of Pterois distinguished from otherspecies of the genus by the following combination ofcharacters: dorsal-fin rays XIII, 10 (rarely XIII, 11); pectoral-fin rays 17–20 [modally 18 in Sri Lankan population;
19 elsewhere (typical form; see under Geographic variations)]; scales below lateral line 11–14 (modally 12); body
depth at anal-fin origin 27.4–35.2 (mean 32.2) % SL; headwidth 14.2–16.4 (15.2) % SL; postorbital length 17.7–21.3(19.6) % SL; longest dorsal-fin spine length 35.1–51.1(42.1) % SL; caudal-peduncle depth 11.0–13.0 (11.9) %SL; pectoral-fin with 6–48 relatively large (subequal topupil diameter) black blotches on membrane, and several
bands on posterior portion of rays (not attached to membrane); several narrow to wide vertical bands laterally on
body, including caudal peduncle; ctenoid scale-covered
area of body relatively extensive (entire body except for
abdomen at ca. 90 mm SL; pectoral-fin base, ventrolateral,and caudal-peduncle portions almost always covered with
ctenoid scales in young and adults greater than 50 mm SL).
Distribution. Although Matsunuma and Motomura(2011) and Matsunuma et al. (2011) recently recorded
P. mombasae from Japan, the former stating that the species is widely distributed in the Indo-West Pacific Ocean,
P. mombasae is now considered to be restricted to theIndian Ocean, from the east coast of Africa to the Andaman
Sea (Fig. 4). Depth collection data for two specimens
indicated a depth range 1.5–60 m. Kotthaus (1979) recorded the species from the Gulf of Aden and off Somalia in
depths of 55–65 m, and Jones and Kumaran (1980)
recorded it (as P. antennata) from the Laccadive Islands(off the west coast of India).

New Pterois from western Pacific Ocean
Fig. 7 Relationships of: a body
depth at anal-fin origin; b headwidth; c postorbital length;d sixth dorsal-fin spine length;e longest dorsal-fin spine length;and f caudal-peduncle depth (allas % of standard length) to
standard length (mm) in Pterois
paucispinula sp. nov. (stars), Sri
Lankan population of
P. mombasae (circles) and otherP. mombasae (triangles).
H indicates holotype

Remarks. A description and illustration of the so-called
P. antennata from the Laccadive Islands in Jones andKumaran (1980) clearly shows the typical form of
P. mombasae [19 pectoral-fin rays and relatively shortdorsal-fin spine length (length of the longest dorsal-fin
spine ca. 35.7 % SL; estimated from illustration)].
Although the specimens recorded by Kotthaus (1979) fromthe Gulf of Aden and off Somalia were not examined in thepresent study, his photograph of the Gulf of Aden specimen(fig. 374) is clearly identifiable as a typical example of
P. mombasae.

Geographical variations. Morphological variations
evident between specimens of P. mombasae from Sri
Lanka and the remaining localities justified recognition of a
distinct geographical (Sri Lankan) population [including
CAS 75355 (Wadge Bank, off the northern tip of the Indian
subcontinent) due to the close distributional proximity andmorphological similarity to the latter]. The Sri Lankan
specimens (n = 13, 47.5–136.5 mm SL; Fig. 3e–h) differed from other conspecifics (n = 20, 26.1–134.3 mm SL;
Fig. 3i–l) in having relatively longer dorsal-fin spines longest dorsal-fin spine length 37.5–51.1 (mean 45.1) %123

M. Matsunuma, H. Motomura

Fig. 8 Plots of principal component (PC) scores based on ninemeasurements for Pterois paucispinula sp. nov. (stars) and P. mombasae from Sri Lanka and India (circles), Andaman Sea (cross),
Oman (diamond) and other Indian Ocean localities (triangles)

SL [vs. 35.1–44.8 (39.8) % SL] (Figs. 6, 7d–e); usually one
less pectoral-fin ray 17–19 (modally 18) [vs. 18–20 (19)]; aslightly higher number of scales between the last dorsal-fin
spine base and the lateral line 6–9 (8) (vs. 7 or 8); scales
between the sixth dorsal-fin spine base and lateral line 7–9(8) [vs. 7–8 (7)]; and pre-dorsal-fin scale rows 4–7 (6) [4–7
(usually 4 or 5)] (Table 3). Although the relative lengths ofthe dorsal-fin spines of the Sri Lankan specimens werevariable, the range overlapping those of other material,
several examples of the former had clearly longer dorsalfin spines.
The results of PCA conducted for 9 measurements (seeabove) indicated subtle differences between the Sri Lankan and remaining specimens of P. mombasae (Fig. 8).
The first two principal components (PC1 and PC2)
accounted for 68.9 % of the variance of the original data,
PC1 being loaded at similarly low levels on all measurements, and PC2 relatively heavily loaded on headwidth and postorbital length [factor loadings are shown inElectronic Supplemental Material (ESM) Table S1]. Plotsfor the Sri Lankan and remaining P. mombasae slightly
overlapped. Although two specimens of P. mombasae
from the Andaman Sea appeared typical of the species inhaving relatively short dorsal-fin spines (longest dorsal-fin
spine length 40.7 % SL; n = 1) and 19–20 pectoral-finrays, PCA located them between the Sri Lankan andremaining specimens. Two Omani specimens, similar to
the Sri Lankan specimens in having 18 pectoral-fin rays
and relatively long dorsal-fin spines (longest dorsal-fin

123spine length 44.8 % SL; n = 1), were equivocal, one
falling into the grouping of Sri Lankan specimens(Fig. 8).
The Sri Lankan specimens of P. mombasae weresimilar to P. paucispinula in having usually 18 pectoralfin rays (Table 3) and relatively long dorsal-fin spines(Fig. 7; Table 1), the number of pectoral-fin rays usually
being considered as a useful diagnostic character forseveral Pteroinae species (Matsunuma et al. 2013b;
Matsunuma and Motomura 2013a–b), whereas dorsal-fin
spine lengths were variable within species and populations. In any case, with the exception of pectoral-fin ray
numbers and dorsal-fin spine lengths, the diagnostic
characters separating Sri Lankan P. mombasae fromP. paucispinula were consistent with those of P. mombasae elsewhere. Therefore, the Sri Lankan specimens
were identified herein as P. mombasae and the aforementioned differences between the two groups regarded
as intraspecific geographic variations. The non-Sri Lankan specimens were regarded as the typical form of
P. mombasae because of their close agreement with the
holotype of the species. Additional specimens fromeastern distributional borders, characterized by having aform intermediate between P. mombasae and P. paucispinula, should be examined in greater detail, including
genetic appraisal, with the prospect of further clarification of the phylogeography and relationships of the two
species.

Fig. 9 Relationship of supraocular tentacle length (as % of orbit
diameter) to standard length (mm) in Pterois paucispinula sp. nov.
(stars) and P. mombasae (triangles), showing relative decreases insupraocular tentacle length with growth

New Pterois from western Pacific Ocean
Fig. 10 Relationships ofnumber of spinules (NS) on:a dorsolateral scales; b cheekscales; c preopercular scales;
d ventrolateral scales; e caudalpeduncle scales; and f pectoralfin base scales to standard
length (mm) in Pterois
paucispinula sp. nov. (stars), Sri
Lankan population of
P. mombasae (circles) and otherP. mombasae (triangles).
H indicates holotype

Discussion
Morphological changes with growth. Almost all headspine numbers increase with growth in both Pterois paucispinula and P. mombasae, the number of nasal spines in
P. paucispinula, for example, being one in juvenile andyoung specimens, but sometimes two in specimens morethan ca. 100 mm SL, as in Parapterois (see Motomura
2004a; Matsunuma et al. 2013a) and Pterois brevipectoralis (Mandrytsa 2002) (see Matsunuma and Motomura
2013b). In contrast, the numbers of supraocular, coronal,

tympanic and preopercular spines remained unchanged
with growth, following complete development of the headspines.
Growth-related changes in the proportions of some bodyparts were also recognized in P. paucispinula andP. mombasae, as in several other members of the subfamilies Pteroinae (Matsunuma et al. 2013b; Matsunuma
and Motomura 2013b) and Scorpaeninae (e.g., Motomura
et al. 2005b, c, 2011b). Analysis of 54 measurements of
P. paucispinula indicated that relative lengths of the orbit
diameter and almost all fin rays, except for the first dorsal-

123

M. Matsunuma, H. Motomura

fin spine, first anal-fin spine, first pectoral-fin ray andlongest pelvic-fin soft ray, tended to decrease as a percentage of SL with growth (only the sixth dorsal-fin spine
shown in Fig. 7d). In contrast, body depth at the anal-fin
origin, postorbital length and suborbital depth became
relatively greater with growth (body depth and postorbital
length shown in Fig. 7a, c). Other growth-related changes
in P. mombasae included positive correlations in body
depth at the anal-fin origin and body width, and negative
correlations in orbit diameter, first dorsal-fin spine length,
thirteenth dorsal-fin spine length, dorsal-fin soft ray length,
all anal-fin ray lengths, pelvic-fin soft ray length and caudal-fin length.
The supraocular tentacle tends to become shorter(reduced in size) with growth in P. paucispinula andP. mombasae (Fig. 9), as in P. antennata (see Matsunuma
and Motomura 2011) and P. brevipectoralis (see Matsunuma and Motomura 2013b). The smallest examined
specimen of P. paucispinula (BSKU 100149, 22.7 mm SL)
had a long supraocular tentacle (length more than twice
orbit diameter), possessing six pairs of lateral branches andfive black bands, whereas a large specimen (URM-P 4264,
134.7 mm SL) had a low skin flap (ca. one-fifth orbit
diameter) on the supraocular without branches or bands.
The number of spinules on the head and body ctenoid
scales increases with growth in both P. paucispinula andP. mombasae (Fig. 10), as do the numbers of spots on the
median fins and blotches on the pectoral fin.
Species comparisons. Smith (1957) regarded Pteropterus as a valid genus, distinguishing it from Pterois on
the basis of several external characters, including: dorsolateral body covered with ctenoid scales (vs. all scales
cycloid in Pterois); fewer than 65 longitudinal series scalerows (vs. more than 65); suborbital ridge close to orbit (vs.
widely separated); and pectoral-fin rays more than 15 (vs.
less than 15). According to Smith (1957) and Mandrytsa(2001, 2002), the following five nominal species of Pterois,
currently regarded as valid, can be assigned to Pteropterus,
being distinguished from other congeners by the abovementioned characters: P. antennata, P. brevipectoralis,
P. mombasae, Pterois radiata (Cuvier in Cuvier andValenciennes 1829) and Pterois sphex Jordan and Evermann 1903. The new species clearly also belongs to
Pteropterus sensu Smith(1957). Examination of the available type specimens or original descriptions of all nominal
species belonging to Pteropterus sensu Smith(1957) indicates that P. paucispinula is readily distinguishable fromthe others, except for P. mombasae, by the combination of
the following characters: dorsal-fin rays usually XIII, 10;pectoral-fin rays usually more than 18; numerous blotcheson pectoral-fin membrane; and several bands posteriorly on
the pectoral-fin rays (not associated with fin membrane).
However, P. paucispinula differed from P. mombasae in

123

Fig. 11 Areas of ctenoid scales (shaded) in a Pterois paucispinula
sp. nov. and b P. mombasae. Based on a BSKU 61062, 90.2 mm SL,
holotype, Japan and b SMF 12797, 82.4 mm SL, Tanzania

having usually one fewer pectoral-fin rays, 17–19 (modally18) [vs. 18–20 (19) in P. mombasae, except for the Sri
Lankan population, 18 or 19 (18) rays]; usually one morescale below the lateral line 11–15 (13) [vs. 11–14 (12)];slightly shallower body depth at the anal-fin origin
26.0–32.8 (mean 30.2) % SL [vs. 27.4–35.2 (32.2) % SL],
head width 12.6–15.6 (14.4) % SL [vs. 14.2–16.4 (15.2) %SL] and caudal-peduncle depth 9.9–11.9 (11.0) % SL [vs.
11.0–13.0 (11.9) % SL]; and slightly shorter postorbital
length 15.4–20.0 (18.1) % SL [vs. 17.7–21.3 (19.6) % SL](Fig. 7a–c, f). Pterois paucispinula tended to have relatively longer dorsal-fin spines compared with P. mombasae, except for the Sri Lankan population, e.g., fifth dorsalfin spine 34.4–48.2 (mean 42.2) % SL in P. paucispinula[vs. 38.6–46.0 (42.2) % SL in the Sri Lankan population;
30.7–38.3 (34.3) % SL in remaining P. mombasae]; sixth
dorsal-fin spine 37.9–54.1 (45.5) % SL [vs. 35.1–49.0(43.1) % SL; 32.0–42.1 (37.4) % SL]; seventh dorsal-fin
spine 39.2–53.8 (46.5) % SL [vs. 37.1–51.1 (44.6) % SL;
33.8–45.0 (39.4) % SL]; eighth dorsal-fin spine 41.6–51.5(46.8) % SL [vs. 37.5–50.6 (45.1) % SL; 35.1–44.2(39.5) % SL]; ninth dorsal-fin spine 39.0–51.7 (45.7) % SL

New Pterois from western Pacific Ocean[vs. 41.9–50.0 (45.9) % SL; 33.5–44.0 (33.5) % SL]; andlongest dorsal-fin spine 42.9–51.7 (47.8) % SL [vs.
37.5–51.1 (45.1) % SL; 35.1–44.8 (39.8) % SL] (Figs. 6,7d–e). The result of PCA conducted for 10 measurementsof the two species showed only a slight overlap of theP. paucispinula and P. mombasae groups (Fig. 8), despite
the limited number of specimens available for the analysis.
Furthermore, P. paucispinula tended to have small areasof the head and body covered with ctenoid scales, incontrast with P. mombasae including the Sri Lankan population. Counts of ctenoid scale spinules on the head andbody showed significant differences between the two species (Fig. 10), notwithstanding a 108.2 mm SL specimenof P. mombasae (SMF 10055) from Sri Lanka having
undeveloped ctenoid scales entirely covering the head andbody, whereas an 84.5 mm SL specimen from the same lot
had an irregularly low number of spinules on the dorsolateral scales (Fig. 10). These two examples are regarded
herein as abnormal and are therefore excluded from thefollowing discussion. Pterois paucispinula is distinguishedfrom P. mombasae by having relatively fewer spinules on
the pectoral-fin base scales 0–2.0 (mean 0.3) [vs. 0–14.8(6.1) in P. mombasae], ventrolateral scales 0–11.7 (0.8)[vs. 2.8–42.0 (23.8)] and caudal-peduncle scales 0–4.5(1.1) [vs. 11.8–47.7 (25.1)] (Fig. 11d–f). Furthermore,
P. paucispinula lacks ctenoid scales on those areas until ca.
80 mm SL, with a small number of ctenoid scales in somelarger adults. Both species have ctenoid scales on the cheekand preopercle area, although P. paucispinula tends to havefewer spinules on the scales compared with P. mombasae,
viz., number of cheek scale spinules 0–8.8 (mean 2.8) in
P. paucispinula vs. 2.3–15.3 (8.2) in P. mombasae; preopercle scale spinules 0–6.7 (1.7) vs. 1.1–12.9 (5.6)(Fig. 10b–c). In contrast, no significant difference between
the species was apparent in spinule numbers on the dorsolateral scales (Fig. 10a). Areas on the head and bodycovered by ctenoid scales at ca. 90 mm SL in the two
species are shown in Fig. 11. At this stage (Fig. 11a),
P. paucispinula has large ctenoid scale areas on the cheekand preopercle, with smaller patches overlaying the suprapostorbital and anterodorsal part of the opercle. Additionally, ctenoid scales occur dorsolaterally on the body(bordered by the nuchal and lower post-temporal spine
bases, the lateral line and base of the spinous portion of thedorsal fin). At the same stage (Fig. 11b), P. mombasae has
similarly large ctenoid scale areas on the cheek and preopercle, but with the opercle also entirely covered with
ctenoid scales. Furthermore, the ctenoid scale area on the
body extends anteriorly to the suprapostorbital region. In
addition, P. mombasae has ctenoid scales over almost all of
the body, including the pectoral-fin base, chest and caudal
peduncle, but excluding the abdomen. Ctenoid scale areas
become more extensive with growth in both species, withlarge adult P. paucispinula having ctenoid scales on theventrolateral portion of the body, caudal peduncle, pectoral-fin base, a large area over the opercle and the lacrimal. However, the development of ctenoid scales is clearly
more rapid in P. mombasae.
Moreover, genetic analysis indicated a substantial
degree of genetic divergence between P. mombasae andP. paucispinula. Comparisons on the basis of 333 bp of the
mtDNA cytochrome b gene showed differences ranging
from 2.5 % to 2.7 % between the two species; in contrast,
the intraspecific sequence difference was 0.7 % in
P. mombasae. The sequence data for P. mombasae(AJ429427–429428) was taken from Kochzius et al.
(2003), who used material from Kenya.
Pterois paucispinula sp. nov. is also similar to Pterois
antennata in having relatively narrow bands on the bodyand the pectoral fins with more than 17 rays. However
P. paucispinula could be easily distinguished from P. antennata as follows: dorsal-fin rays usually XIII, 10 (rarely
XIII, 11) [vs. usually XII, 11 (rarely XII, 10 or XII, 12) in
P. antennata]; pectoral-fin rays usually more than 18(rarely 17) [vs. usually less than 17 (rarely 18)]; and several
bands on the posterior portion of the upper pectoral-fin rays
(free from membrane) (vs. bands absent) (Matsunuma andMotomura 2011; this study). Pterois sphex and P. brevipectoralis, known from the Hawaiian Islands and southwestern Indian Ocean (Mascarene Islands and Saya de
Malha Bank), respectively, are also similar to P. paucispinula in having dorsal-fin rays XIII, 10 and several bands
posteriorly on the upper pectoral-fin rays. However,P. paucispinula can be clearly distinguished from P. sphex
by having usually more than 18 pectoral-fin rays (vs. 15–16
in P. sphex), the body with approximately 18 relatively
narrow vertical bands (vs. 4–5 broad vertical bands withnarrow bands between) and several blotches, but without
distinct vertical bands on the pectoral-fin membrane (vs.
3–13 vertical narrow bands without blotches) (Eschmeyer
and Randall 1975; Matsunuma and Motomura 2013b; thisstudy). It can be distinguished from P. brevipectoralis byhaving more than 17 pectoral-fin rays (vs. less than 16 in
P. brevipectoralis) and skin flaps on the lacrimal andposterior margin of the preopercle not becoming larger
with growth (vs. becoming extremely large, length greaterthan orbit diameter in large adults) (Mandrytsa 2002;
Matsunuma and Motomura 2013b; this study). Pterois
paucispinula is also readily distinguished from P. radiataby pectoral-fin ray number (usually more than 18 vs. 16–17
in P. radiata) and the nature of the bands on the caudal
peduncle (2–6 narrow transverse bands vs. a single broad
horizontal band in P. radiata) (Smith 1957; this study).
Comparative materials examined. Pterois antennata:
80 specimens (14.0–153.9 mm SL): JAPAN: BSKU 6942,
109.8 mm SL; BSKU 51076, 58.5 mm SL; BSKU 79021,

123M. Matsunuma, H. Motomura

102.2 mm SL; BSKU 86405, 86.1 mm SL; BSKU 100096,
38.0 mm SL; BSKU 103166, 111.1 mm SL; FAKU 99045,
77.0 mm SL; FAKU 99086, 84.5 mm SL; HUMZ 53491,
118.1 mm SL; KAUM–I. 6436, 133.6 mm SL; KAUM–I.
6438, 120.1 mm SL; KAUM–I. 6586, 132.9 mm SL;
KAUM–I. 29634, 72.8 mm SL; KAUM–I. 29635,
73.5 mm SL; KAUM–I. 29745, 100.6 mm SL; KAUM–I.
29746, 153.9 mm SL; KPM-NR 21471, 93.3 mm SL;
KPM-NR 22989, 23.5 mm SL; KPM-NR 26918, 36.1 mm
SL; NSMT-P 30809, 58.5 mm SL; NSMT-P 80768,
72.8 mm SL; NSMT-P 90778, 70.0 mm SL; URM-P 1324,
104.8 mm SL; URM-P 18654, 116.4 mm SL; URM-P
36009, 77.3 mm SL. TAIWAN: ASIZP 57316, 95.3 mm
SL; ASIZP 59074, 66.0 mm SL; ZMB 16148, 31.3 mm
SL. PHILIPPINES: AMS I. 40144-010, five specimens,
14.0–73.2 mm SL; NSMT-P 34677, 112.3 mm SL; NTOUP 20080414-101, 106.4 mm SL; NTOU-P 20080414-116,
68.1 mm SL; SMF 10025, 66.2 mm SL; USNM 99021,
112.8 mm SL. INDONESIA: KPM-NR 14031, 74.5 mm
SL; NSMT-P 62052, 72.3 mm SL; NSMT-P 63740,
84.8 mm SL; NSMT-P 71523, 49.0 mm SL;NSMT-P
71676, 76.6 mm SL; NSMT-P 102043, 72.2 mm SL;
NSMT-P 102044, 49.0 mm SL; SMF 4204, 89.9 mm SL;
ZMB 796, lectotype of Scorpaena antennata Bloch 1787,
117.6 mm SL. MICRONESIA: USNM 223578, 77.6 mm
SL. GUAM: AMS I. 39371-001, 87.5 mm SL. PAPUANEW GUINEA: NMV A13698, 121.7 mm SL; OMNH
13808, 109.6 mm SL. SOLOMON ISLANDS: AMS I.
39040-045, 57.5 mm SL. VANUATU: USNM 356588,
53.4 mm SL. WALLIS AND FUTUNA: USNM 374803,
81.5 mm SL. TONGA: USNM 337077, three specimens,
78.1–106.9 mm SL. COOK ISLANDS: FMNH 16311,
three specimens, 49.2–88.8 mm SL; FMNH 44140,
22.8 mm SL. FRENCH POLYNESIA: MNHN 1984-279,
87.3 mm SL. AUSTRALIA: AMS I. 4119, 51.6 mm SL;
AMS I. 20441-005, 83.2 mm SL; AMS I. 20756-044,
96.1 mm SL; AMS I. 20965-025, 88.8 mm SL; AMS I.
23847-027, 83.1 mm SL; FAKU 113358, 69.0 mm SL.
INDIA: SMF 28932, 122.9 mm SL. SRI LANKA: SMF
4327, 85.2 mm SL. RED SEA: SMF 12887, 77.1 mm SL.
YEMEN: SMF 4625, 128.2 mm SL; SMF 4626, 105.1 mm
SL. SOMALIA: FMNH 484, holotype of Pterois ellioti
Meek 1899, 48.8 mm SL. KENYA: SMF 9100, 67.5 mm
SL; SMF 9932, 74.3 mm SL. MALDIVES: SMF 5386,
78.9 mm SL; SMF 9770, 30.5 mm SL. MAURITIUS:
USNM 349900, 88.1 mm SL.
Pterois brevipectoralis: 10 specimens (49.2–117.9 mmSL) listed in Matsunuma and Motomura (2013b).
Pterois radiata: 45 specimens (23.3–151.4 mm SL):JAPAN: BSKU 2712, 66.8 mm SL; BSKU 3239, 57.7 mm
SL; BSKU 3317, 46.4 mm SL; BSKU 63779, 61.6 mm SL;
BSKU 74108, 83.3 mm SL; BSKU 106150, 24.5 mm SL;
FAKU 6113, 98.7 mm SL; FAKU 98972, 56.9 mm SL;

123

KAUM–I. 41254, 93.9 mm SL; KPM-NR 16856,
103.0 mm SL; KPM-NR 22714, 56.3 mm SL; NSMT-P
31593, 23.3 mm SL; NSMT-P 55493, 109.0 mm SL;
NSMT-P 104342, 52.3 mm SL. TAIWAN: ASIZP 55046,
80.5 mm SL; ASIZP 56910, 123.8 mm SL; NMMB 10536,
129.3 mm SL;NTOU-P 2007-04-07-002, 80.2 mm SL;
NTOU-P 20070625-008, 67.2 mm SL; NTOU-P
20070710-201, 88.9 mm SL; NTOU-P 20071111-401,
105.5 mm SL; NTOU-P 20071111-701, 112.7 mm SL.
INDONESIA: KPM-NR 14056, 57.8 mm SL; KPM-NR
14057, 59.1 mm SL; NCIP 2024, 91.5 mm SL. KIRIBATI:
NSMT-P 23485, 49.2 mm SL; NSMT-P 23266, two specimens, 52.2–55.8 mm SL. SAMOA: NMV A5290,
126.9 mm SL. FRENCH POLYNESIA: FMNH 23176,
151.4 mm SL; LACM 54162.001, 142.0 mm SL; MNHN
1984-280, two specimens, 103.6–108.3 mm SL; MNHN
6625, holotype of Pterois (Pseudomonopterus) vittata
Sauvage 1878, 58.4 mm SL. MALDIVES: FMNH 75832,
three specimens, 65.4–120.2 mm SL. SEYCHELLES:
SMF 12794, two specimens, 76.5–94.9 mm SL; SMF
12882, 64.7 mm SL. SAUDI ARABIA: SMF 349, holotype
of Pterois cincta Rüppell 1838, 141.5 mm SL. TANZANIA: SMF 12882, 64.7 mm SL. COMOROS: SAIAB
30752, 88.5 mm SL. MADAGASCAR: AMS 28107-013,
90.8 mm SL. MAURITIUS: USNM 349900, 88.1 mm SL.
Pterois sphex: six specimens (40.6–111.7 mm SL),
including holotype of Pterois sphex Jordan and Evermann
1903, listed in Matsunuma and Motomura (2013b).
Acknowledgments We are deeply grateful to M. McGrouther, A.
Hay and S. Reader (AMS); K.-T. Shao, Y.-C. Liao and M.-Y. Lee(ASIZP); J. Maclaine and O. Crimmen (BMNH); H. Endo, T.
Yamakawa, N. Nakayama (BSKU) and E. Katayama (formerly
BSKU); T. Nakabo, Y. Kai, A. Tokairin (FAKU) and N. Muto (formerly FAKU); K. Swagel (FMNH);M. Yabe, H. Imamura, T. Kawai(HUMZ); H. Senou (KPM); R. Causse (MNHN); Z. Arifin, R. Pratiwi,
M. Adrim, I. Alhakim and K. Wibowo (NCIP); H.-C. Ho (NMMB);
M. Gomon and D. Bray (NMV); K. Matsuura and G. Shinohara(NSMT); I.-S. Chen (NTOU); K. Hatooka (OMNH); U. Satapoomin(PMBC); T. Alpermann, H. Zetzsche (SMF) and F. Krupp (formerlySMF); T. Yoshino (formerly URM); J. Williams (USNM); P. Bartschand C. Lamour (ZMB); and K. Sakamoto (ZUMT) for their kind
hospitality during the first author’s visits to their institutions. We areindebted to D. Catania (CAS), A. Suzumoto (BPBM), T. Noichi(CMNH), R. Feeney (LACM) and R. Bills (SAIAB) for providing
opportunities to examine specimens. We especially thank S. Chiba(NSMT) for instructing the first author in molecular analysis and also
T. Yoshida (KAUM) for providing specimen data, Y. Sakurai (Okinawa Prefecture, Japan), S. Koizumi (URM), K. Koeda (formerly
URM) and H.-C. Ho (MNHB) for donating specimens, M. Aizawa(BLIH) and H. Endo for providing photographs of fresh P. paucispinula, Y. Haraguchi, students and other KAUM volunteers for
curatorial assistance and sampling specimens, and G. Hardy(Ngunguru, New Zealand) for reading the manuscript and providing
help with English. We also thank the two anonymous reviewers andeditors for their helpful suggestions. This study was supported in partby 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 Asian Core

New Pterois from western Pacific Ocean
Program ‘‘Establishment of Research and Education Network on
Coastal Marine Science in Southeast Asia’’, JSPS International
Training Program ‘‘Protect Diversity of Bioresources in the TropicalArea’’, the Coastal Area Capability Enhancement in Southeast Asia
Project of the Research Institute for Humanity and Nature, Kyoto,
Japan, and the ‘‘Biological Properties of Biodiversity Hotspots inJapan’’ project of the National Museum of Nature and Science,
Tsukuba, Japan, and by grant awarded to the first author from the
Mikimoto Fund for Marine Ecology.

References
Akihito, Iwata A, Kobayashi T, Ikeo K, Imanishi T, Ono H, Umehara
Y, Hamamatsu C, Sugiyama K, Ikeda Y, Sakamoto K, Fumihito
A, Ohno S, Gojobori T (2000) Evolutionary aspects of gobioid
fishes based upon a phylogenetic analysis of mitochondrial
cytochrome b genes. Gene 259:5–15
Allen GR (1997) Marine fishes of tropical Australia and south-east
Asia. 3rd revised edition. Western Australian Museum, Perth
Allen GR, Adrim M (2003) Coral reef fishes of Indonesia. Zool Stud
42:1–72
Allen GR, Cross NJ (1989) Scorpaenidae. In: Paxton JR, Hoese DF,
Allen GR, Hanley JE (ed) Zoological catalogue of Australia.
Vol. 7. Pisces. Petromyzontidae to Carangidae. AustralianGovernment Publishing Service, Canberra, pp 438–452
Allen GR, Cross NJ, Hoese DF (2006) Scorpaenidae. Lionfishes,
rockfishes, scorpionfishes, stingfishes, stonefishes, waspfishes.
In: Hoese DF, Bray DJ, Paxton JR, Allen GR (ed) Zoologicalcatalogue of Australia. Vol. 35, parts 1–3: fishes. CSIRO
Publishing, Collingwood, pp 876–892
Allen GR, Erdmann MV (2012) Reef fishes of the East Indies. Vols.
I–III. Tropical Reef Research, Perth
Allen GR, Steene R, Humann P, DeLoach N (2003) Reef fish
identification. Tropical Pacific. New World Publications,Jacksonville
Bloch ME (1787) Naturgeschichte der ausländischen Fische. Berlin.
Naturgeschichte der Ausländischen Fische 3:i–xii ? 1–146
Cuvier G, Valenciennes A (1829) Histoire naturelle des poisons, vol
4. Chez FG Levrault, Paris
Eschmeyer WN (1965) Western Atlantic scorpionfishes of the genus
Scorpaena, including four new species. Bull Mar Sci 15:84–164
Eschmeyer WM (1986) Family No. 149: Scorpaenidae. In: Smith
MM, Heemstra PC (ed) Smith’s sea fishes. Macmillan South
Africa, Johannesburg, pp 463–478
Eschmeyer WN, Randall JE (1975) The scorpaenid fishes of the
Hawaiian Islands, including new species and new records (Pisces:
Scorpaenidae). Proc California Acad Sci Ser 4 40:265–333
Fricke R, Kulbicki M, Wantiez L (2011) Checklist of the fishes of
New Caledonia, and their distribution in the southwest Pacific
Ocean (Pisces). Stuttg Beitr Naturk A New Ser 4:341–463
Gloerfelt-Tarp T, Kailola PJ (1984) Trawled fishes of southernIndonesia and northwestern Australia. Australian Development
Assistance Bureau (ADAB), Directorate General of Fisheries,
Indonesia (DGF), and German Agency for Technical Cooperation (GTZ)
Herre AW (1952) A review of the scorpaenoid fishes of thePhilippines and adjacent seas. Philip J Sci 80:381–482
Hirata T (2010) Nettai-minokasago Pterois antennata. In: Takagi M,
Hirata T, Hirata S, Nakata C (ed) Fishes of Ainan, Ehime
Prefecture. Soufusha shuppan, Matsuyama, p 34
Hutchins JB (2001) Checklist of the fishes of Western Australia. Rec
West Aust Mus Suppl(63):9–50

Jones S, Kumaran M (1980) Fishes of the Laccadive Archipelago.
The Nature Conservation and Aquatic Sciences Service,
Trivandrum
Jordan DS, Evermann BW (1903) Descriptions of new genera andspecies of fishes from the Hawaiian Islands. Bull US Fish Comm
22:161–208
Kochzius M, Söller R, Khalaf MA, Blohm D(2003) Molecularphylogeny of the lionfish genera Dendrochirus and Pterois(Scorpaenidae, Pteroinae) based on mitochondrial DNA
sequences. Mol Phyl Evol 28:396–403
Kotthaus A (1979) Fische des Indischen Ozeans. Ergebnisse derichthyologischen Untersuchungen während der Expedition des
Forschungsschiffes ‘‘Meteor’’ in den Indischen Ozean, Oktober
1964 bis Mai 1965. a. Systematischer Teil, XXI. Diverse
Ordnungen. Meteor Forsch Ser D Biol(28):6–54
Kubota S, Tanase H, Muto N, Tokairin A, Kimura K, Nakabo T(2011) Numerous fishes stranding on the beach of Tanabe Bay,
Wakayama Prefecture, Japan in winter in 2011. Journal of Japan
Driftological Society 9:13–16
Kuiter RH (1996) Guide to sea fishes of Australia. A comprehensive
reference for divers and fishermen. New Holland Ltd, Sydney
Kuiter RH, Debelius H (2006) World atlas of marine fishes. IKANUnterwasserarchiv, Frankfurt
Kuriiwa K, Hanzawa N, Yoshino T, Kimura S, Nishida M (2007)
Phylogenetic relationships and natural hybridization in rabbitfishes (Teleostei: Siganidae) inferred from mitochondrial andnuclear DNA analyses. Mol Phyl Evol 45:69–80
Mandrytsa SA (2001) Lateral line system and classification of
scorpaenoid fishes (Scorpaeniformes: Scorpaenoidei). Perm
State Univ Press, Perm
Mandrytsa SA (2002) A new species of the genus Pteropterus(Scorpaenidae: Scorpaeniformes) from the Indian Ocean. J Ichthyol 42:129–130
Masuda H, Araga C, Yoshino T (1975) Coastal fishes of southern
Japan. Tokai Univ Press, Tokyo
Matsunuma M, Aizawa M, Sakurai Y, Motomura H (2011) Record ofa lionfish, Pterois mombasae (Smith, 1957), from Yaku-shimaIsland, southern Japan, and notes on distributional implicationsof the species and P. antennata in Japan. Nat Kagoshima 37:3–8
Matsunuma M, Jawad LA, Motomura H (2013a) New records of ascorpionfish, Parapterois macrura (Scorpaenidae: Pteroinae),
from Oman and Somalia, western Arabian Sea. Biogeography
15:49–54
Matsunuma M, Motomura H (2011) First records of a lionfish, Pterois
mombasae (Scorpaenidae: Pteroinae), from Japan, and morphological comparisons with P. antennata. Japan J Ichthyol
58:27–40
Matsunuma M, Motomura H (2013a) A new lionfish of the genus
Dendrochirus (Scorpaenidae: Pteroinae) from the Tuamotu
Archipelago, South Pacific Ocean. Spec Divers 18:1–7
Matsunuma M, Motomura H (2013b) Newly recognized diagnostic
characters of the poorly known lionfish, Pterois brevipectoralis(Scorpaenidae: Pteroinae), with notes on fresh coloration. Spec
Divers 18:163–173
Matsunuma M, Sakurai M, Motomura H (2013b) Revision of the
Indo-West Pacific genus Brachypterois (Scorpaenidae: Pteroinae), with description of a new species from northeastern
Australia. Zootaxa 3693:401–440
Miya M, Nishida M (2000) Use of mitogenomic information inteleostean molecular phylogenetics: a tree-based exploration
under the maximum-parsimony optimality criterion. Mol Phyl
Evol 17:437–455
Motomura H (2004a) Morphological comparison of a poorly known
scorpionfish, Parapterois macrura, with a related species,
P. heterura (Scorpaenidae: Pteroinae). Zool Stud 43:1–7

123

M. Matsunuma, H. Motomura
Motomura H (2004b) New species of scorpionfish, Scorpaenacocosensis (Scorpaeniformes: Scorpaenidae) from the Cocos
Islands, Costa Rica, eastern Pacific Ocean. Copeia 2004:818–824
Motomura H (2004c) Revision of the scorpionfish genus Neosebastes(Scorpaeniformes: Neosebastidae) with descriptions of five new
species. Indo-Pac Fish 37:1–75
Motomura H (2009) Scorpaenidae. Scorpionfishes. In: Kimura S,
Satapoomin U, Matsuura K (ed) Fishes of Andaman Sea, west
coast of southern Thailand. Natl Mus Nat Sci, Tokyo, pp 63–69
Motomura H, Fricke R, Eschmeyer WN (2005a) Redescription of apoorly known scorpionfish, Scorpaena canariensis (Sauvage),
and a first record of Pontinus leda Eschmeyer from the Northern
Hemisphere (Scorpaeniformes: Scorpaenidae). Stuttg Beitr Naturk Ser A (Biol) 674:1–15
Motomura H, Johnson JW (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 PR, Gomon MF (2006a) A new species of the
scorpionfish genus Maxillicosta from the southeast coast ofAustralia, with a redescription of M. whitleyi (Scorpaeniformes:
Neosebastidae). Copeia 2006:445–459
Motomura H, Last PR, Yearsley GK (2005b) Scorpaena bulacephala,a new species of scorpionfish (Scorpaeniformes: Scorpaenidae)
from the northern Tasman Sea. Zootaxa 1043:17–32
Motomura H, Last PR, Yearsley GK (2006b) New species of shallow
water scorpionfish (Scorpaenidae: Scorpaena) from the central
coast of Western Australia. Copeia 2006:360–369
Motomura H, Matsunuma M, Ho H-C (2011a) New records of three
scorpaenid fishes (Teleostei: Scorpaeniformes) from Taiwan.
J Fish Soc Taiwan 38:97–107
Motomura H, Paulin CD, Stewart AL (2005c) First records ofScorpaena onaria (Scorpaeniformes: Scorpaenidae) from thesouthwestern Pacific Ocean, and comparisons with the Northern
Hemisphere population. New Zealand J Mar Freshwater Res
39:865–880
Motomura H, Senou H (2008) A new species of the scorpionfish
genus Scorpaena (Scorpaenidae) from Izu Peninsula, Pacificcoast of Japan. J Fish Biol 72:1761–1772
Motomura H, Struthers CD, McGrouther MA, Stewart AL (2011b)
Validity of Scorpaena jacksoniensis and a redescription of
S. cardinalis, a senior synonym of S. cookii (Scorpaeniformes:
Scorpaenidae). Ichthyol Res 58:315–332
Poss SG (1999) Scorpaenidae. Scorpionfishes (also, lionfishes,
rockfishes, stingfishes, stonefishes, and waspfishes). In: Carpenter KE, Niem VH (ed) FAO species identification guide forfisheries purposes. The living marine resources of the western
central Pacific. Vol. 4. Bony fishes part 2 (Mugilidae to
Carangidae). FAO, Rome, pp 2291–2352

123

Poss SG, Rama-Rao KV (1983) Scorpaenidae. In Fischer W, Bianchi
G (ed) FAO species identification sheets for fishery purposes.
Western Indian Ocean. Fishing area 51, vol. 4. FAO, Rome,pp 1–13? ‘‘SCORP Apis 1’’ to ‘‘SCORP Syna 1’’
Quantum GIS Development Team (2014) Quantum GIS Geographic
Information System. Open Source Geospatial Foundation Project. http://www.qgis.org/. Accessed 11 June 2014
Randall JE (1995) Coastal fishes of Oman. Crowford House
Publishing Pty Ltd, Bathurst, Australia
Randall JE (2005) Reef and shore fishes of the South Pacific, New
Caledonia to Tahiti and the Pitcairn Islands. Univ Hawai’i Press,
Honolulu
Randall JE, Allen GR, Steene RC (1997) Fishes of the Great Barrier
Reef and Coral Sea. 2nd edition. Univ Hawai’i Press, Honolulu
Randall JE, Eschmeyer WN (2002) Revision of the Indo-Pacificscorpionfish genus Scorpaenopsis, with descriptions of eight new
species. Indo-Pac Fish 34:1–79
R Core Team (2014) R: A language and environment for statisticalcomputing. R Foundation for Statistical Computing, Vienna,
Austria. http://www.r-project.org/. Accessed 12 June 2014
Sabaj Pérez MH (2014) Standard symbolic codes for institutional
resource collections in herpetology and ichthyology: an online
reference. Version 5.0 (22 September 2014). American Societyof Ichthyologists and Herpetologists, Washington, DC. http://
www.asih.org/resources/standard-symbolic-codes-institutionalresource-collections-herpetology-ichthyology. Accessed 21
October 2014
Satapoomin U (2011) The fishes of southwestern Thailand, theAndaman Sea–are view of research and a provisional checklistof species. Phuket Mar Biol Cent Res Bull 70:29–77
Shimizu T(1984) Nettai-minokasago Pterois antennata. In: Masuda
H, Amaoka K, Araga C, Uyeno T, Yoshino T (ed) The fishes of
the Japanese Archipelago. Tokai Univ Press, Tokyo, p 302, pl
282C
Smith JLB (1957) The fishes of the family Scorpaenidae in thewestern Indian Ocean. Part 2. The subfamilies Pteroinae,
Apistinae, Setarchinae and Sebastinae. Ichthyol Bull JLB Smith
Inst Ichthyol Rhodes Univ 5:75–87
Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013)
MEGA6: molecular evolutionary genetics analysis version 6.0.
Mol Biol Evol 30:2725–2729
Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG(1997) The Clustal-X windows interface: flexible strategies for
multiple sequence alignment aided by quality analysis tools.
Nucl Acid Res 25:4876–4882