Species Diversity 18: 163–173
25 November 2013
DOI: 10.12782/sd.18.2.163

Diagnosis of Pterois brevipectoralis163

Newly Recognized Diagnostic Characters of the Poorly
Known Lionfish Pterois brevipectoralis(Scorpaenidae: Pteroinae), with Notes on
Fresh Coloration
Mizuki Matsunuma1,3 and Hiroyuki Motomura2
1 The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-24 Korimoto, Kagoshima 890-0065, Japan

E-mail: k1139853@kadai.jp

2 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp
3 Corresponding author(Received 8 May 2013; Accepted 17 November 2013)
Pterois brevipectoralis (Mandrytsa, 2002), previously known only from the holotype and two paratypes collected fromthe Saya de Malha Bank (western Indian Ocean), in depths of 57–90 m, is redescribed on the basis of 10 additional specimens collected from the type locality and the Cargados Carajos Shoals, Mascarene Islands. The species is characterized by
the presence of usually XIII, 10 dorsal-fin rays, usually 16 pectoral-fin rays, a relatively low number (6–10) of dark blotcheson the pectoral-fin membrane, a relatively short pectoral fin [longest ray 44.5–66.4 (mean 53.4)% SL; not extending beyond
the level of the posterior end of the dorsal-fin base in specimens over 91.3 mm SL], ctenoid scales covering the head anddorsolateral body, and the posterior portion of the pectoral-fin rays (not associated with fin membrane) with several bands.
In addition, extremely large (greater than orbit diameter), fan-like skin flaps on the tip of the posterior lacrimal spine andthe posterior margin of the preopercle in large adults are recognized herein as diagnostic characters for the species. The firstreport of fresh coloration, based on a specimen from the Saya de Malha Bank, is included, and ontogenetic changes in various features are described in detail.
Key Words: Pterois brevipectoralis, Pteropterus, diagnosis, distribution, Cargados Carajos Shoals, Mascarene Islands, Sayade Malha Bank, Indian Ocean.

Introduction
The Indo-Pacific scorpaenid genus Pterois Oken, 1817 has
been characterized within the subfamily Pteroinae as having
three anal-fin spines, all pectoral-fin rays unbranched, theparietal spine of males not elongated, and no spine or ridgeon the mandible (Eschmeyer and Randall 1975; Poss 1999).
Smith (1957) and Mandrytsa (2001, 2002) regarded Pteropterus Swainson, 1839 as valid, being distinguished from
Pterois by having more than 15 pectoral-fin rays (vs fewer
than 15 in Pterois), the head and dorsolateral body covered
with ctenoid scales (vs all scales cycloid), less than 65 scalerows in the longitudinal series (vs fewer than 65), and theventral margin of the orbit close to the suborbital ridge (vsgreatly separated). In his phylogenetic analysis of the Scorpaenoidei, based on morphological characters, Mandrytsa(2001) included the following five species in Pteropterus:
Scorpaena antennata Bloch, 1787, Pterois radiata Cuvier in
Cuvier and Valenciennes, 1829, Pterois sphex Jordan andEvermann, 1903, Pteropterus mombasae Smith, 1957, andPteropterus brevipectoralis Mandrytsa, 2002 (as Pteropterussp.). However, most recent studies have treated Pteropterus
as a junior synonym of Pterois (e.g., Herre 1952; Eschmeyer
and Randall 1975; Poss 1999). In addition, Kochzius et al.
(2003) suggested Dendrochirus Swainson, 1839 to also be ajunior synonym of Pterois on the basis of a molecular phylogenetic analysis. Although Dendrochirus had previously
been considered distinguished from Pterois by a single pectoral fin character, i.e., several rays branched in adults, compared with all rays unbranched throughout life in Pterois(Smith 1957; Poss 1999; Mandrytsa 2001), such separationof the two genera was questioned by Eschmeyer and Randall(1975). However, a comprehensive phylogenetic study of
Pteroinae has yet to be made, and the relationships between
Pterois, Pteropterus, and Dendrochirus remain unclear. Accordingly, we tentatively follow the widespread practice ofregarding Pteropterus as a junior synonym of Pterois, as previously done by Matsunuma and Motomura (2011).
Pterois brevipectoralis (Mandrytsa, 2002) was originallydescribed as Pteropterus brevipectoralis on the basis of three
specimens (65.5–147 mm SL) collected from the Saya de
Malha Bank, western Indian Ocean, in depths of 57–90 m(Mandrytsa, 2002). The species has not been reported since.
Because Mandrytsa’s (2002) detailed description of the species was based solely on preserved specimens, it lacked any
description of fresh coloration. The recent location of several additional specimens (49.2–117.9 mm SL), collectedfrom the Saya de Malha Bank and the Mascarene Islands,
representing first records from the latter locality, and the© 2013 The Japanese Society of Systematic Zoology

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M. Matsunuma and H. Motomura

Fig. 1.
Lateral (top) and dorsal (bottom) views of head of Pterois brevipectoralis, HUMZ 73848, 125.7 mm SL. Shaded areas indicate skinflaps. 1, nasal spine; 2, preocular spine; 3, supraocular spine; 4, postocular spine; 5, coronal spine; 6, tympanic spine; 7, parietal spine; 8,nuchal spine; 9, pterotic spine; 10, lower posttemporal spine; 11, supracleithral spine; 12, sphenotic spine; 13, supplemental preopercularspine; 14, preopercular spine; 15, suborbital ridge/spine; 16, lateral lacrimal ridge/spine; 17, posterior lacrimal spine. Barbel on anteroventral
lacrimal based on right side in lateral view; nasal spines and right supraocular skin flap not illustrated in dorsal view.

Diagnosis of Pterois brevipectoralis165

Fig. 2.

Fresh specimen of Pterois brevipectoralis, HUMZ 73846, 115.8 mm SL, Saya de Malha Bank. Photo: HUMZ.

recognition of additional diagnostic features and ontogenetic changes, has led to the redescription of Pterois brevipectoralis given herein. Furthermore, a photograph of a specimen
from the Saya de Malha Bank, taken prior to preservation,
has enabled a description of the fresh coloration of the species.

base of the uppermost preopercular spine is termed the supplemental preopercular spine (Eschmeyer 1965) and that on
the lateral surface of the lacrimal bone is called the laterallacrimal spine (Motomura and Senou 2008: fig. 2; Moto­mura et al. 2011: fig. 1). Standard length is abbreviated as
SL. Institutional codes followed Sabaj Pérez (2012).

Materials and Methods

Pterois brevipectoralis (Mandrytsa, 2002)(Figs 1–6; Tables 1–2)

Measurements generally follow Motomura (2004b, c),with head width and maxillary depth following Motomura
et al. (2005b, 2006a) and Motomura et al. (2006b), respectively. The following additional measurements are given:body depth at anal-fin origin—body depth at level of firstanal-fin spine base; 1st–9th pectoral-fin ray length—distance from the uppermost pectoral-fin ray origin to each
ray tip. Lengths of skin flaps on the tips of supraocular andposterior lacrimal spines were measured as the distance between the flap base and the extreme end of the flap. Counts
generally follow Motomura et al. (2005a–c) and Motomura
and Johnson (2006), with predorsal scale counts following
Motomura et al. (2006b). The following additional count isgiven: scales above lateral line—vertical scale rows between
the first pored lateral-line scale and the dorsal-fin base. The
last two soft rays of the dorsal and anal fins were counted as
single rays, each pair being associated with a single pterygiophore. Counts and measurements were made on the left
side wherever possible, except that pectoral-fin rays werecounted on both sides.
Head spine terminology (shown in Fig. 1) generally follows Randall and Eschmeyer (2002: fig. 1) and Motomura(2004c: fig. 1), with the following additions: the spine at the

Pteropterus sp.: Mandrytsa 2001: 16 (phylogenetic material;
key).
Pteropterus brevipectoralis Mandrytsa 2002: 129, unnumbered fig. on p. 129 (type locality: Saya de Malha Bank).
Examined specimens (10 specimens, 49.2–117.9 mm
SL). MASCARENE ISLANDS: USNM 392069, 49.2 mm
SL, lagoon on west side of Frigate Island, Cargados Carajos
Shoals, Mauritius (16.6°S, 59.5°E), 0–1 m, V. Springer et al.,
1 April 1976; USNM 400514, 2 specimens, 59.2–69.3 mm
SL, ca. 2 miles (i.e., ca. 3.2 km) east of Raphael Island, Cargados Carajos Shoals, Mauritius (16°S, 59°31′E), 0.2–1.1 m,
V. Springer et al., 3 April 1976. SAYA DE MALHA BANK:HUMZ 73841, 110.1 mm SL, HUMZ 73843, 111.7 mm SL,
HUMZ 73844, 91.3 mm SL, HUMZ 73845, 101.0 mm SL,
HUMZ 73846, 115.8 mm SL, HUMZ 73847, 117.9 mm SL,
HUMZ 73848, 125.7 mm SL, 10°30′S, 61°32′E, 95 m, 3 September 1977.
Diagnosis. A species of Pterois with the following
combination of characters: dorsal-fin rays usually XIII, 10;
anal-fin soft rays 6; pectoral-fin rays usually 16 (rarely 15);
scale rows in longitudinal series 46–51; pectoral-fin rays
relatively short, not extending beyond level of posterior

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M. Matsunuma and H. Motomura

end of dorsal-fin base in large specimens of >91.3 mm SL,
longest ray length 44.5–66.4% (mean 53.4%) SL; large, fanlike skin flaps (length greater than half of orbit diameter inlarge specimens of >91.3 mm SL) on tip of posterior lacrimal spine and posterior margin of preopercular, relative
size increasing with growth; head and dorsolateral parts ofbody covered with ctenoid scales; 6–10 large, black blotcheson pectoral-fin membrane; posterior parts of pectoral-finrays, not associated with fin membrane, with several reddish(dark in preserved specimens) bands.
Description. Characters included in specific diagnosis
not repeated here. Morphometrics and selected meristics
as shown in Tables 1–2; in particular, anal fin with 3 spines,
pelvic fin with 1 spine and 5 soft rays, scales below lateralline 11, branchiostegal rays 7.
Body oblong, moderately compressed posteriorly; depth
moderate, greater than length of longest dorsal-fin spine.
Three barbels on snout tip, their length subequal to orslightly longer than nasal tentacle length. Short tentacle(simple in juveniles, with a lateral branch in large adults)
on posterior edge of low, membranous tube associated withanterior nostril, its tip extending beyond posterior marginof posterior nostril when depressed posteriorly. Supraocular with long tentacle with 2–4 lateral branches in juveniles,
low skin flap in large adults. Anteroventral part of lacrimal
with short tentacle, its length subequal to barbels on snout
tip. Large fan-like skin flap on tip of posterior lacrimal
spine, becoming larger with growth, tip of flap extending
beyond margin of preopercle when laid along maxilla inadults of >91.3 mm SL (not reaching margin in juveniles of<69.3 mm SL). Two relatively large (but smaller than posterior lacrimal spine flap) fan-like flaps on margin of preopercle, their tips extending beyond posterior margin ofinteropercle when laid flat. Usually also a large, fan-like flap
present on ventral portion of third preopercular spine. Smallskin flap present on anterodorsal portion of orbit surface, its
length less than diameter of posterior nasal pore. No otherskin flaps on head or body.
Ctenoid scales covering head and anterodorsal surface of body; remainder of body with cycloid scales. Basal
scales absent on dorsal and anal fins. Relatively small cycloid scales extending onto bases of pectoral and caudal fins.
Occipital area entirely covered with ctenoid scales. Dorsal
surface of postocular covered with weakly ctenoid scales,
mostly possessing 1 or 2 spines. Cheek with ctenoid scales,
but suborbital pit lacking scales. Preopercle generally covered with cycloid scales, several larger scales being weakly
ctenoid. Basal part of posterior lacrimal spine partly covered
with small, weakly ctenoid scales, most possessing a single
spine. Dorsal surfaces of supraocular and postocular, andposterior portion of interorbital area partly covered withrelatively small, weakly ctenoid scales with usually 1 but
up to 3 spines. Central and anterior portions of interorbital
area lacking scales. Interopercle lacking scales. Opercle withmostly cycloid and some large, weakly ctenoid scales.
Mouth moderately large, slightly oblique, forming angleof ca. 30–40° to horizontal axis of head and body; upperedge of posterior maxilla swollen laterally, forming low

Table 1. Morphometrics, expressed as percentages of standard
length, of Pterois brevipectoralis.
Mascarene
Islands(n=3)

Saya de Malha
Bank
(n=7)

SL (mm)49.2–69.3

91.3–125.7

Body D (% SL)
Body D1
Body W
Head L
Head W
Snout L
Orbit diameter
Interorbital W2
Interorbital W3
Upper-jaw L
Maxillary D
Suborbital D
Postorbital L
Pre-dorsal-fin L
Pre-anal-fin L
Pre-pelvic-fin L
1st DS L
2nd DS L
3rd DS L
4th DS L
5th DS L
6th DS L
7th DS L
8th DS L
9th DS L
10th DS L
11th DS L
12th DS L
13th DS L
1st DSR L
Longest DSR L
1st AS L
2nd AS L
3rd AS L
1st ASR L
Longest ASR L
1st P1 L
2nd P1 L
3rd P1 L
4th P1 L
5th P1 L
6th P1 L
7th P1 L
8th P1 L
9th P1 L
Longest P1 L
P2S L
Longest P2SR L
Caudal-fin L
Caudal-peduncle L
Caudal-peduncle D

36.6–40.4
30.7–31.9
24.2–26.5
39.6–43.6
13.6–15.3
9.6–11.3
13.6–15.2
7.7–9.1
7.5–8.1
18.5–20.3
7.3–8.3
0.8
18.9–19.9
33.5–36.8
69.4–73.7
35.6–40.3
17.1
24.8–28.7
28.2–28.9
32.9–34.9—
38.2–40.3
36.0–43.0
35.6–42.1
34.3–39.0
30.3–36.9—
11.2–18.2
17.7–18.8
23.8–25.7
26.1–28.5
10.2–10.6
17.6–19.3
19.1–23.6
28.7–31.6
32.2–34.5
51.7–65.4
61.0–71.6
64.5–72.6
66.4
66.1–68.9
62.5–64.4
61.7
56.6
38.7–52.0
63.0–66.4
23.4–25.7
43.4–47.3
35.0–38.3
16.6–17.3
11.3–11.7

38.3–41.7
26.6–31.0
24.5–28.9
40.9–45.1
15.3–17.0
10.5–11.8
10.5–11.8
7.2–9.2
6.4–8.6
21.6–23.4
6.4–8.6
0.9–1.7
19.6–20.6
34.9–37.0
70.2–75.1
39.2–42.8
—
—
26.0
—
—
—
—
—
37.7
34.4–36.1
17.5–30.2
13.1–15.3
11.5–15.8
19.4–21.4
25.8–26.0
8.2–10.3
14.2–18.1
15.8–18.4
22.9–26.6
27.2–28.5
31.7–41.7
38.4–49.4
44.4–50.7
41.0–52.2
44.5–56.0
43.7–55.5
40.1–50.0
40.5–46.8
38.6–54.9
44.5–46.8
17.9–19.8
35.8–40.2
34.8–36.2
14.5–16.8
10.4–11.2

39.4
29.5
26.1
43.5
15.8
11.0
14.8
8.4
7.5
21.6
8.4
1.1
19.9
36.0
72.6
40.4
17.1
26.8
27.7
33.9
—
39.2
40.1
39.8
37.0
34.4
24.0
14.6
15.0
21.9
26.5
9.9
16.7
19.5
26.5
29.4
44.4
51.0
54.3
50.7
52.8
51.4
48.5
46.4
45.1
53.4
21.5
39.7
36.0
15.8
11.0

Membrane above 6th
P1 extending up to

85.9–93.0%
of 6th P1 L

76.7–91.2% of
6th P1 L

86.1% of 6th
P1 L

LocalityMeans

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: width.
1 at anal-fin origin; 2 at vertical midline of eye; 3 at posterior end of
preocular spine base.

Diagnosis of Pterois brevipectoralis167

Table 2.

Number of specimens of Pterois brevipectoralis displaying selected meristics.
Pectoral fin rays (one/other sides)

Dorsal fin soft rays

Pored LL scales

15/15

15/16

16/16

9

10

24

25

26

1

3

6

1

9

1

1

1

SR in longitudinal series

Scales above LL

46

47

48

49

50

51

7

8

1

1

1

1

1

1

4

5

SR between LL and last DSB

SR between LL and 6th DSB

Predorsal-fin SR

6

7

8

5

6

7

5

6

7

2

6

2

5

4

1

5

4

1

Upper gill rakers

Lower gill rakers

Total gill rakers

4

5

10

11

12

14

15

16

17

3

7

5

3

2

2

3

4

1

DSB: dorsal-fin spine base, LL: lateral line, SR: scale rows.

ridge; posterior margin of maxilla extending beyond midorbit level, but not reaching level of posterior orbit margin.
Symphyseal gap separating premaxillary teeth bands distinctly wider than width of each band; upper jaw with bandof small, slender conical teeth; about 9 tooth rows at front ofupper jaw; about 4–6 tooth rows at front of lower jaw; small
teeth in 4–5 rows forming blunt V-shaped patch on vomer;
no palatine teeth. Three sensory pores on underside of eachdentary, and one small pore on each side of symphysial
knob.
Dorsal profile of snout relatively steep, forming angle ofca. 40–50° to horizontal axis of head and body. Nasal spinewith 1–3 spinous points. Preocular spine with 2–10 spinous
points. Supraocular with 0–3 small spines. Postocular with
4–15 small spines. Interorbital ridge moderately developed,
reduced posteriorly. Coronal spine with single point (rarely
2 points in large adults). Tympanic spine with a single point(rarely 2 points in large adults). Anterior margin of occipital area sloped transversely from between bases of coronalspines, curved posteriorly in dorsal view. Parietal spinewith 1–7 spinous points, its base relatively long, diverging
posteriorly. Nuchal spine with 1–2 spinous points, its basecompletely conjoined with that of parietal spine. Postorbital spine absent. Sphenotic with 2–14 small spines. Pterotic with 1–8 small spines. Lower posttemporal spine with
1–4 spinous points. Cleithrum with upper and lower ridges;
upper ridge with 1 flattened spine posteriorly (rarely 2
spines in large adults); lower ridge with 1–4 spines.
Lateral lacrimal spine with 1–4 spinous points; all lacrimal ridges strongly spinous in adults. Suborbital with upper
main and lower supplemental ridges; upper ridge with 4–19
spines; lower ridge with 4–12 spines. Anterior lacrimal spine
absent. Posterior lacrimal spine broad, plate-like, with 1–4
spinous points on distal margin. Preopercle with 3 spines,
all of similar length; uppermost spine directed posterodorsally, other 2 directed posteriorly to posteroventrally; basesof uppermost and middle spine with 1–3 and 0–2 supplemental spines, respectively; lowermost spine broad, platelike, with 2–4 spinous points on distal margin. Exposed

Fig. 3.
Preserved specimens of Pterois brevipectoralis. A, USNM
392069, 49.2 mm SL, Frigate Island, Cargados Carajos Shoals,
Mascarene Islands; B, USNM 400514, 1 of 2 specimens, 69.3 mm
SL, Raphael Island, Cargados Carajos Shoals, Mascarene Islands;
C, HUMZ 73844, 91.3 mm SL, Saya de Malha Bank.

opercular spine absent.
Origin of first dorsal-fin spine above lower posttemporalspine; bases of first and second dorsal-fin spines closer thanthose of subsequent adjacent spines; seventh or eighth spine
longest; twelfth spine usually shortest, its length 49–84%

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M. Matsunuma and H. Motomura

Fig. 4.
Skin flaps on lacrimal (A and C) and preopercle (B and D) of Pterois brevipectoralis at different growth stages. A, B, USNM
392069, 49.2 mm SL; C, D, HUMZ 73848, 125.7 mm SL. Arrows indicate third preopercular spine.

and 90–114% of that of antepenultimate and posteriormost
spines, respectively; membrane of spinous portion of dorsal
fin strongly incised. Dorsal-fin soft rays all branched; fourth
ray longest, but distinctly shorter than longest dorsal-fin
spine; posteriormost ray not joined by membrane to caudal
peduncle. Origin of first anal-fin spine just below posteriormost dorsal-fin spine base; third spine longest; length offirst spine 53–70% and 45–65% of that of second and third
spines, respectively. Anal-fin soft rays all branched; secondor third ray longest, its length slightly greater than that oflongest dorsal-fin soft ray; posteriormost ray not joined bymembrane to caudal peduncle. Pectoral fin relatively long,
tip of longest ray extending beyond vertical drawn throughfirst anal-fin spine base in all juvenile/adult stages, andjust reaching or extending beyond caudal-fin base in juveniles of <69.3 mm SL, just reaching or not reaching vertical drawn through posteriormost dorsal-fin soft ray basein large adults of >91.3 mm SL; all rays unbranched, lower

rays weakly thickened. First pelvic-fin spine base below
third dorsal-fin spine base; all pelvic-fin soft rays branched;usually second (rarely third) soft ray longest, its tip extending beyond first anal-fin spine base when depressed, but notreaching posteriormost anal-fin soft ray base; posteriormost
soft ray joined by membrane to abdomen for less than about
one-fifth to one-fourth of ray length. Caudal fin usually with
4 procurrent rays, 2 segmented unbranched rays, and 5 segmented branched rays in dorsal and ventral series (rarely 3,
2 and 5 rays, respectively, in each or either series); profile
of posterior margin of fin moderately rounded. Caudal peduncle moderately short, deep, its depth 65–74% of caudalpeduncle length.
Color of fresh specimen. Based on color photographof HUMZ 73846, taken before preservation (Fig. 2). Groundcolor of head and body white, somewhat pinkish dorsally.
Jaws and skin flap on posterior lacrimal spine tip pinkish.
Reddish band crossing eye, reaching posterodorsal cornerDiagnosis of Pterois brevipectoralis169of preopercle. Large blackish blotch on ventral corner of
opercle. Broad reddish band saddling nape at level of parietal spine, numerous narrow reddish bands on body sides.
Dorsal-fin membrane translucent; 2–3 reddish bands oneach dorsal-fin spine; ca. 25 small black spots on soft rays altogether. Anal-fin membrane translucent; ca. 11 small blackspots on soft rays altogether. Pectoral fin dark red, with several relatively large black blotches on membrane and narrow
black band on distal margin; free portion of rays creamywhite with 1–3 reddish bands. Pelvic fin dark red; strongly
blackish marginally. Caudal-fin membrane translucent withca. 50 small black spots on rays altogether. Eye yellowish;
iris black.
Color of preserved specimens (based mostly on Mascarene Islands specimens, the coloration of Saya de MalhaBank specimens having disappeared almost completely, except for black markings) (Fig. 3). Head and body creamywhite. Five brown bands on head; anteriormost band on
snout reaching from anteroventral margin of orbit to anteroventral margin of lacrimal; second band extending frombelow eye (from ventral margin) across posterodorsal corner of maxilla to ventral corner of preopercle; third band
relatively broad from supraocular spine (supraocular tentacle base), obliquely crossing eye and reaching subopercle;
fourth band just behind eye, saddling nape at level of coronal spine; fifth band saddling nape at level of parietal spine,reaching opercle. Large brown blotch on subopercle, numerous brown bands on lateral body, including caudal peduncle. Dorsal-fin membrane translucent; 1–3 brown bands oneach dorsal-fin spine; 21–40 small black spots on soft rays
altogether. Anal-fin membrane translucent; 16–32 small
black spots on soft rays altogether. Pectoral-fin membranetranslucent, blackish marginally, with 1–10 relatively large
black blotches on membrane, numerous small, black spots
on border between posterior rays and membrane; free portion of rays creamy-white with 1–4 brown bands. Pelvic-fin
membrane blackish, rays creamy-white. Caudal-fin membrane translucent with 49–62 small black spots on rays altogether. Supraocular tentacle with 2–4 brown bands (basedon USNM 392069, USNM 400514; Fig. 3A, B).
Remarks. Although the holotype of P. brevipectoralis was not available for this study, the present specimens(91.3–125.7 mm SL) collected from the Saya de Malha Bank
agreed with the description of Pteropterus brevipectoralis(=Pterois brevipectoralis) given by Mandrytsa (2002), withthe exception of the longitudinal series scale row counts[46–50 in the present specimens vs 53–55 in Mandrytsa(2002) as “scale vertical rows”]. This difference is likely due
to different counting techniques, for although Mandrytsa(2002) stated that his counts generally followed Eschmeyer(1969), the longitudinal series scale rows thus being thoseabove the lateral line from the row over the posterior endof the supracleithral spine base to the row at the end of thehypural (as in the present study), he may have included several scale rows behind the hypural plate, thereby resulting
in higher numbers. Although Mandrytsa (2002) provided
a drawing of the holotype of P. brevipectoralis, an accurate
count of longitudinal series scale rows based on this draw-

Fig. 5.
Supraocular tentacle of Pterois brevipectoralis, USNM
392069, 49.2 mm SL.

ing is not possible due to an apparent inaccuracy in thenumber of scale rows depicted. Although the drawing indicates at least 29 pored lateral line scales, despite the lateral line being partly hidden by the pectoral fin, Mandrytsa(2002) described the type series of P. brevipectoralis as having 25–26 pored lateral line scales. Moreover, the present
specimens have 23–26 pored lateral line scales. In addition,
the drawing indicates ca. 45 scale rows in the longitudinal series, including a row above the first pored lateral line
scale, to the posteriormost row, but this is fewer than the
figures of 53–55 given by Mandrytsa (2002) in his description. Minor differences in morphometrics, regarded here
as intraspecific variation, were found between the present
specimens and the original description, including a slightly
shorter head length (39.6–45.1% of SL vs 42.3–44.9% of SL
in the original description), slightly longer upper-jaw length(18.5–23.4% of SL vs 17.6–20.2% of SL), and slightly deeper
caudal-peduncle depth (10.4–11.7% of SL vs 9.5–10.2% of
SL). Although Mandrytsa (2002) wrote that the species hasa single nasal spine, the present specimens had 1–3 spines,

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M. Matsunuma and H. Motomura

Fig. 6.
Relationships of (A) body depth at anal-fin origin, (B) second pectoral-fin ray length, (C) longest pelvic-fin soft ray length, and(D) upper jaw length (as % of standard length), and (E) length of skin flap on supraocular and (F) length of skin flap on posterior lacrimal
spine tip [as % of orbit diameter (OD)] to standard length in Pterois brevipectoralis.

with the number of spines tending to increase with growth(see “Morphological changes with growth” below). The present specimens (49.2–69.3 mm SL) collected from the Mascarene Islands also conformed to the diagnosis of P. brevipectoralis. Morphological differences apparent between the
Mascarene Islands and Saya de Malha Bank specimens, including head spine condition and relative lengths of bodyparts, are regarded here as intraspecific ontogenetic changes(see “Morphological changes with growth” below). The Mascarene Islands specimens represent the first records for the
species from that region, although they are relatively smaller(49.2–69.3 mm SL) and from a shallower depth (<1.1 m in
the Cargados Carajos Shoals) than the three type specimens(65.5–147 mm SL) collected at 57–90 m depth from the Sayade Malha Bank (Mandrytsa 2002).
Morphological changes with growth. The numberof head spines increases with growth in P. brevipectoralis.
Whereas juvenile and young specimens of less than 91.3 mm
SL have a simple nasal spine with a spinous point, large
adults of more than 101.0 mm SL usually have 2 (rarely 3)
points, as in Parapterois (Scorpaenidae: Pteroinae) (Motomura 2004a).
Growth-related changes in the relative lengths of somebody parts are well documented for several members of the
Scorpaenidae (e.g., Motomura et al. 2005b, c, 2006b, 2011).
Analyses of 51 measurements in P. brevipectoralis indicatesthat body depth at the anal-fin origin, caudal-peduncle
length and depth, and the lengths of almost all fin rays tend
to decrease as a percentage of SL with growth (Fig. 6A–C).
In addition, the relative length of the upper jaw and thehead width both become significantly greater with body size
(Fig. 6D). Notably, the relative lengths of the 1st to 8th pectoral-fin rays all become markedly shorter with growth (Figs
6B, 7; Table 1), small specimens (49.2–69.3 mm SL) having
relatively long pectoral-fin rays, with the tip of the longest
ray just reaching or extending slightly beyond the caudal-finbase (Fig. 3A, B), and large adults (>91.3 mm SL) having
relatively short pectoral-fin rays, with the tip of the longest
ray not reaching the caudal-fin base (Figs 2, 3C).
In contrast, the skin flap on the tip of the posterior lacrimal spine becomes larger with growth (Figs 4A, C, 6E).
The smallest specimen (49.2 mm SL) examined has a small
skin flap on the spine tip, with the posterior tip of the flap
not reaching the posterior margin of the maxilla when
depressed posteriorly (Fig. 4A). In contrast, large adults(>91.3 mm SL) have a large, fan-like skin flap on the spineDiagnosis of Pterois brevipectoralis171

Fig. 7.
Relationship of position of longest pectoral-fin ray tip to
standard length (mm) in Pterois brevipectoralis. A, D, and C represent the anal-fin base end, the dorsal-fin base end, and the caudalfin base, respectively; =, >, < indicate just reaching or extending
slightly beyond, extending distinctly beyond, and not reaching,respectively.

tip, extending considerably beyond the posterior marginof the maxilla when depressed (Fig. 4C). The skin flaps on
the posterior margin of the preopercle also tend to becomelarger with growth (Fig. 4B, D), small specimens (<59.2 mmSL) having 2 small skin flaps on the margin below thethird preopercular spine (Fig. 4B) whereas large specimens(>69.3 mm SL) usually have 2 large, fan-like skin flaps inthe same region and an additional large, fan-like flap on theventral part of the third preopercular spine (Fig. 4D). Skin
flap lengths in the largest specimen examined (125.7 mmSL) exceed the orbit diameter (111.5% of orbit diameter).
The skin flap on the supraocular becomes reduced withbody growth (Figs 3, 6D). The smallest specimen (49.2 mmSL) has a long supraocular tentacle (167% of orbit diameter)with three pairs of lateral branches (Fig. 5) whereas large
specimens of >91.3 mm SL have a small skin flap (length
9–38% of orbit diameter) (Fig. 1). In addition, large adults
have a lateral branch on the anterior nasal flap (Fig. 1)
whereas juveniles have a simple flap (Fig. 5). No other significant growth-related change in head skin flaps was found.
The numbers of dark spots on the median fins tend to decrease with body growth in P. brevipectoralis (Figs 2–3, 8).
A juvenile specimen (49.2 mm SL) has a total of 40, 29, and57 spots on the dorsal, anal, and caudal fins, respectively,
whereas a large specimen (117.9 mm SL) has 21, 17, and 50
spots, respectively. Such changes are most apparent on thedorsal and anal fins.
Species comparisons. Mandrytsa (2002) stated that P.
brevipectoralis could be distinguished from a related con-Fig. 8.
Relationships of number of spots on dorsal (circles), anal(crosses), and caudal fins (triangles) to standard length in Pterois
brevipectoralis.

gener, Pterois sphex Jordan and Evermann, 1903 (as Pteropterus sphex), which shares similar dorsal and pectoral fin ray
counts, by having relatively short pectoral fins (not extending beyond a vertical through the end of the dorsal-fin basein the former vs extending farther in P. sphex); however, as
mentioned above, relative lengths of the pectoral-fin rays
change significantly with growth in P. brevipectoralis (Fig. 7).
This character, therefore, is available for species discrimination only in large adults. Mandrytsa (2002) also stated thatP. brevipectoralis differs from P. sphex in having relatively
shallower notches of the pectoral-fin membrane, the depthof the notch between the fifth and sixth rays not exceeding
one-fifth of those ray lengths (vs over two-fifths in P. sphex).
This character was confirmed as diagnostic by our examination of nine present specimens (49.2–125.7 mm SL) of P.
brevipectoralis and two specimens (55.8–102.6 mm SL) of P.
sphex. The membrane above the sixth pectoral-fin ray extends to up to 76.7–93.0% (mean 86.1%) of the ray length in
P. brevipectoralis (Table 1) but only 55.6–70.5% in P. sphex.
In addition, P. brevipectoralis is easily distinguished from P.
sphex in the coloration of the pectoral fin, which has 6–10
large blotches in P. brevipectoralis, but 3–13 narrow verticalbands in P. sphex (Eschmeyer and Randall 1975: fig. 2; thisstudy).
Pterois brevipectoralis is also similar to Pterois mombasae (Smith, 1957), occurring sympatrically in the western
Indian Ocean, in overall body appearance. Both species
have XIII, 10 dorsal-fin rays, a relatively short dorsal-fin
spine and pectoral-fin rays, dark blotches on the pectoralfin membrane, and bands on the posterior portions of the172

M. Matsunuma and H. Motomura

pectoral-fin rays (not associated with fin membrane). Pterois
brevipectoralis differs from P. mombasae in having fewer
pectoral-fin rays [15–16 (mode 16) vs 17–20 (18) in P. mombasae], generally fewer blotches on the pectoral fin (6–10
vs 6–24), and shorter pectoral-fin rays [longest ray length
44.5–66.4 (mean 53.4)% SL vs 64.9–84.5 (75.2)% SL] (Smith
1957; Matsunuma and Motomura 2011; this study). Moreover, P. brevipectoralis is uniquely characterized by having
extremely large (greater than half the orbit diameter), fanlike skin flaps on the tip of the posterior lacrimal spine andthe posterior margin of the preopercle in large adults, suchflaps becoming extremely large with growth. In contrast,
similar flaps in P. mombasae are relatively small (less thanhalf the orbit diameter) in large adults and become reducedwith growth (this study).
Comparative material. Pterois mombasae (Smith,1957): BSKU 61061, 84.2 mm SL, BSKU 61062, 90.2 mm SL,
Kashiwa-jima island, Kochi, Japan; BSKU 72569, 79.3 mm
SL, BSKU 86402, 59.4 mm SL, BSKU 86404, 56.3 mm SL,
Tosa Bay, Kochi, Japan; BSKU 100149, 23.2 mm SL, Okino-shima island, Kochi, Japan; CAS 75355, 136.5 mm
SL, off east coast of Sri Lanka; CAS 75358, 102.0 mm SL,
Trincomalee, Sri Lanka; CMNH–ZF 15450, 30.9 mm SL,
Yaku-shima island, Osumi Group, Japan; KAUM–I. 6582,
83.4 mm SL, KAUM–I. 6588, 106.9 mm SL, KAUM–I.
28824, 52.3 mm SL, Okinawa-jima island, Ryukyu Islands,
Japan; KPM-NI 26919 (formerly IOP 1103), 26.2 mm SL,
KPM-NI 26930 (formerly IOP 3120), 49.5 mm SL, Izu,
Shizuoka, Japan; MNHN 1995–737, 98.4 mm SL, Futuna
Island, Wallis and Futuna; NSMT-P 54352, 38.3 mm SL,
Susaki, Chiba, Japan; NSMT-P 110789 (formerly KSHS
22471), 112.1 mm SL, NSMT-P 110789 (formerly KSHS
22473), 83.6 mm SL, Kashiwa-jima island, Kochi, Japan;
PMBC (uncat.), 2 specimens, 70.3–75.0 mm SL, Phuket,
Thailand; SAIAB (formerly RUSI) 117, 128.8 mm SL, holotype of Pteropterus mombasae, Mombasa, Kenya; URM-P
4264, 136.9 mm SL, URM-P 4265, 103.5 mm SL, URM-P
4266, 92.3 mm SL, URM-P 4267, 92.7 mm SL, URM-P 4268,
85.0 mm SL, URM-P 4269, 63.3 mm SL, Sesoko-jima island,
Ryukyu Islands, Japan; URM-P 41467, 75.1 mm SL, Okinawa-jima island, Ryukyu Islands; USNM 265918, 54.9 mm
SL, Macclesfield Bank; USNM 382904, 67.3 mm SL, Santa
Cruz Island, Solomon Islands. Pterois sphex: AMS I.16203001, 42.9 mm SL, Waimea Bay, Oahu Island, Hawaiian Islands; AMS I.16205-001, 102.6 mm SL, Makua, Oahu Island,
Hawaiian Islands; LACM 37099-4, 40.6 mm SL, Kahe Point,
Oahu Island, Hawaiian Islands; LACM 37100-5, 2 specimens, 50.6–55.8 mm SL, Kahe Point, Oahu Island, Hawaiian Islands; USNM 50650, 111.7 mm SL, holotype of Pterois
sphex, Honolulu, Oahu Island, Hawaiian Islands.

Acknowledgments
We are especially grateful to M. Yabe, H. Imamura, T.
Kawai, T. Yamanaka, and S. Ohashi (HUMZ), and to J. Williams (USNM), for their kind hospitality during the firstauthor’s visit to their institutions and providing the photo-graph of P. brevipectoralis when fresh. We are grateful to M.
McGrouther (AMS), H. Endo (BSKU), D. Catania (CAS),
H. Kawase (CMNH), H. Senou (KPM), K. Matsuura and
G. Shinohara (NSMT), U. Satapoomin (PMBC), O. Gon(SAIAB), T. Alpermann (SMF), and T. Yoshino (formerly
URM) for providing opportunities to examine comparative
material. G. Hardy (Ngungru, New Zealand) read the manuscript and provided helpful comments. This study was supported in part by Grants-in-Aid for Scientific Research (B,24370041 and C, 23580259) from the Japan Society for the
Promotion of Science, Tokyo, Japan (JSPS), and a Grant-inAid for Young Scientists (B, 19770067) from the Ministry of
Education, Science, Sports and Culture, Tokyo, Japan, JSPS
Asian Core Program “Establishment of Research and Education Network on Coastal Marine Science in Southeast Asia”,
JSPS International Training Program “Protect Diversity of
Bioresources in the Tropical Area”, and the Coastal Area Capability Enhancement in Southeast Asia Project of the Research Institute for Humanity and Nature, Kyoto, Japan.

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