Zoological Studies 46(1): 35-45 (2007)

Scorpaena pepo, a New Species of Scorpionfish (Scorpaeniformes:
Scorpaenidae) from Northeastern Taiwan, with a Review of S. onaria
Jordan and Snyder
Hiroyuki Motomura1,*, Stuart G. Poss2, and Kwang-Tsao Shao3
1The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan
224552 Arcadia Farm Road, Pass Christian, MI 39571, USA
3Reseach Center for Biodiversity, Academia Sinica, Nankang, Taipei 115, Taiwan(Accepted February 14, 2006)

Hiroyuki Motomura, Stuart G. Poss, and Kwang-Tsao Shao (2007) Scorpaena pepo, a new species of scorpionfish (Scorpaeniformes: Scorpaenidae) from northeastern Taiwan, with a review of S. onaria Jordan and
Snyder. Zoological Studies 46(1): 35-45. A new species of scorpionfish, Scorpaena pepo, is described on thebasis of 4 specimens collected from off the northeastern coast of Taiwan in the northwestern Pacific Ocean, at
a depth of ca. 200 m. The new species is closely related to S. onaria with which co-occurs in Taiwan. The 2
species are distinguished from other Indo-Pacific species of Scorpaena by the following combination of characters: the lateral surface of the lacrimal with a single spine, the posterior margin of the anterior lacrimal spinewith an additional spine, the median interorbital ridge well developed, embedded cycloid scales covering theanteroventral surface of the body and the pectoral-fin base, a high body depth (36.4-45.1% of SL), 43-48 longitudinal scale series, and the lateral line sloping steeply downward above the anterior 1/2 of the pectoral fin.
The new species can be distinguished from S. onaria by having 16 pectoral-fin rays; 46-48 longitudinal scaleseries; the posterior margin of the maxilla extending well beyond a vertical through the posterior margin of theorbit; the highest portion of the median interorbital ridge greater in height than the interorbital ridges; the posterior end of the base of the 1st suborbital spine located below the eye, between the posterior margins of thepupil and orbit; the supraocular tentacle being shorter than the pupil diameter, its length 1.7-3.9% (mean 3.1%)
of SL; and the head and body when fresh yellowish-orange, with numerous scattered small distinct black spots,
except on ventral surface. Scorpaena pepo sp. nov. is a large species (attaining at least 245.1 mm SL), whereas S. onaria from the northwestern Pacific Ocean is relatively small (the largest specimen, 187.9 mm SL). Two
geographical populations regarded as S. onaria are compared in detail with S. pepo.
http://zoolstud.sinica.edu.tw/Journals/46.1/35.pdf
Key words: Scorpaenidae, New species, Scorpaena pepo, Taiwan, Scorpaena onaria.

F rom 16 to 21 May 2005, the 7th Indo-

onfish were tinged with bright yellowish-orange
and had numerous small black spots scattered on
the entire head, body, and fins. Of over 30 individuals of the species found at the market, 3 smaller
specimens (223.1-245.1 mm standard length)were purchased by the 1st author on 18 and 19
May and registered at Academia Sinica, Taipei
and the Australian Museum, Sydney. On 19 May
2005 another specimen (172.9 mm standard
length) apparently conspecific with those from thefish market, collected from the same locality, wasPacific Fish Conference, which was organized by
the Ichthyological Society of Taiwan, the National
Museum of Marine Biology and Aquarium, andAcademia Sinica, was held in Taipei, Taiwan.
During the conference, the 1st author found alarge number of large-sized individuals (thelargest, over 40 cm total length) of scorpionfish,
collected from off the northeastern coast of Taiwan
at a depth of ca. 200 m, at Taipei Fish Market on
Wan-Da Road. In the market, these fresh scorpi-*To whom correspondence and reprint requests should be addressed. E-mail:motomura@kaum.kagoshima-u.ac.jp

35

36

Zoological Studies 46(1): 35-45 (2007)

found in the collection of the Department of
Zoology, National Taiwan University, Taipei.
These 4 specimens were identified as being
attributable to Scorpaena by having the following
diagnostic characters for the Indo-Pacific speciesof Scorpaena: 12 dorsal-fin spines, teeth on thepalatines, an occipital pit, the posterior lacrimal
spine directed posteroventrally, some pectoral-fin
rays branched in adults, and the pored lateral-line
scales continuing onto the caudal-fin base (Poss
1999, Motomura et al. 2005b). The specimens
were most similar in overall body appearance to S.
onaria Jordan and Snyder, 1900 which co-occurs
with the former in Taiwan. However, the former differed from S. onaria in several aspects, including
the numbers of the pectoral-fin rays and longitudinal scale series, head morphology, and coloration.
Thus, the specimens from northeastern Taiwan are
described herein as a new species, S. pepo. In
addition to a description of S. pepo, a review of therelated species, S. onaria, is also given.

MATERIALS AND METHODSCounts and measurements follow Motomura(2004a b), with additional counts following
Motomura et al. (2005b). Standard length isexpressed as SL. The last 2 soft rays of the dorsal
and anal fins are counted as single rays, as eachpair are associated with a single pterygiophore.
Pectoral-fin ray counts begin with the uppermost
element. Terminology of the head spines follows
Randall and Eschmeyer (2001) and Motomura etal. (2005b). Institutional codes follow Leviton et al.
(1985) with additional institutional abbreviations asfollows: Research Center for Biodiversity,
Academia Sinica, Taipei, Taiwan (ASIZP); Divisionof Fisheries Sciences, University of Miyazaki,
Japan (MUFS); and Museum of New Zealand Te
Papa Tongarewa (NMNZ).
Scorpaena pepo sp. nov.
New English name: Pumpkin Scorpionfish(Figs. 1, 2A)
Holotype: ASIZP 65020, male, 244.3 mm SL,
off NE coast of Taiwan, hook and line fishing, ca.
200 m depth; purchased at Taipei Fish Market by
H. Motomura, 19 May 2005.
Paratypes: AMS I. 43631-001, male, 223.1mm SL, same data as for holotype except date, 18
May 2005; ASIZP 65021, female, 245.1 mm SL,
same data as holotype; NTUM 4555, male, 172.9mm SL, Nanfangao, Ilan, NE Taiwan, 25 Dec.
1981.
Diagnosis: An Indo-Pacific species of
Scorpaena with the following combination of characters: pectoral-fin rays 16; longitudinal scalesseries 46-48; embedded cycloid scales covering
anteroventral surface of body and pectoral-finbase; lateral line sloping steeply downward above
anterior 1/2 of pectoral fin; posterior margin of
maxilla extending well beyond a vertical throughposterior margin of orbit; median interorbital ridge
well developed, its highest portion greater in height
than interorbital ridges; lateral surface of lacrimal
with a single spine; anterior lacrimal spine with anadditional spine; posterior lacrimal spine simple; 3suborbital spines, posterior end of 1st suborbital
spine base located below eye between posterior
margins of pupil and orbit; space between upperand lower opercular spines not covered with fleshyskin; supraocular tentacle shorter than pupil diameter, its length 1.7-3.9% (mean 3.1%) of SL; underside of lower jaw without tentacles; no skin flap on
pectoral-fin axil; relatively deep body (depth 36.442.6% of SL); head and body yellowish-orange
when fresh, with numerous small distinct blackspots, averaging about 1/10 body scale diameterin size, scattered on entire head and body (except
for ventral surface of head and body), dorsal, analand caudal fins, and upper 1/2 of pectoral fin; alarge black blotch at the margin of membrane
between 6th and 10th or 11th dorsal-fin spines inmales.
Description: Proportional measurements aspercentages of SL of the specimens of Scorpaena
pepo are given in table 1. Frequency distributionsof selected meristic characters are presented intable 2. Features for the holotype are presented
first, followed by paratype data (if different) inparentheses.
Dorsal fin with 12 spines and 9 soft rays (lastsplit to base); 1st spine length more than 1/2 thatof 2nd spine; 3rd (4th in 2 paratypes) spine
longest, subequal to length of 2nd anal-fin spine;
4th to 11th spines becoming progressively shorter;
length of 11th spine 1.7 in last spine; interspinous
membranes moderately incised; all soft rays
branched; 1st (2nd in 2 paratypes) soft ray longest,
length subequal to length of longest anal-fin softray; membrane between soft rays weakly incised;posterior branch of last soft ray joined by membrane to caudal peduncle for more than 3/4 its
length. Anal fin with 3 spines and 5 soft rays (lastsplit to base); 1st spine 2.1 (1.9-2.1; mean 2.0) in
2nd spine; 2nd spine longest; all rays branched;37

Motomura et al. -- New Species of Scorpaena from Taiwan

Table 1. Morphometric characters, expressed as percentages of standard length, of Scorpaena pepo sp.
nov. and the southwestern Pacific and northwestern Pacific populations of S. onaria. Means in parentheses
include type data
Scorpaena pepo sp. nov.

Scorpaena onaria
Northwestern PacificSouthwestern Pacific

Holotype
ASIZP 65020

Paratypes
n=3

Holotype
USNM 49405

Paratypes
n=2

Non-typesn = 27

Non-typesn = 24Standard length (mm)244.3

172.9-245.1

157.7

136.6-144.8

81.1-187.9

56.5-265.2

Body depth
Body width
Head length
Snout length
Orbit diameter
Interorbital width1
Interorbital width2
Head width
Upper-jaw length
Maxilla depth
Suborbital space
Postorbital length
Between tips of opercular spines
Occipital pit length
Occipital pit width
Post-occipital pit length
Predorsal-fin length
Preanal-fin length
Prepelvic-fin length
1st dorsal-spine length
2nd dorsal-spine length
3rd dorsal-spine length
4th dorsal-spine length
5th dorsal-spine length
6th dorsal-spine length
7th dorsal-spine length
8th dorsal-spine length
9th dorsal-spine length
10th dorsal-spine length
11th dorsal-spine length
12th dorsal-spine length
Longest dorsal-ray length
1st anal-spine length
2nd anal-spine length
3rd anal-spine length
Longest anal-ray length
Pectoral-ray length
Pelvic-spine length
Longest pelvic-ray length
Caudal-fin length
Caudal-peduncle length
Caudal-peduncle depth

40.1
26.1
46.7
13.3
10.1
6.9
6.5
16.1
23.7
7.3
4.1
25.2
6.8
6.4
6.4
8.9
38.3
76.6
45.7
6.5
12.6
16.9
15.6
13.2
13.8
13.0
11.1
8.8
6.8
11.5
19.5
7.0
15.1
12.3
20.7
27.8
13.3
23.2
24.1
16.0
10.5

36.4-42.6 (39.6)
23.3-29.4 (26.7)
46.8-49.3 (48.0)
13.1-13.8 (13.4)
10.4-12.3 (10.8)
6.6-7.0 (6.8)
5.8-6.4 (6.2)
16.7-16.9 (16.6)
23.7-25.2 (24.3)
7.2-8.1 (7.5)
3.4-4.6 (4.1)
23.7-26.4 (25.4)
6.9-8.1 (7.3)
6.6-6.7 (6.6)
6.4 (6.4)
8.8-10.1 (9.3)
38.9-39.5 (38.9)
74.5-79.0 (76.3)
43.6-46.7 (45.6)
7.0-8.4 (7.5)
13.4-13.9 (13.3)
15.2-17.2 (16.4)
16.8 (16.2)
16.3-17.6 (17.0)
13.9-15.2 (14.1)
12.6-14.3 (13.6)
11.9-13.7 (13.0)
9.8-11.2 (10.6)
7.6-9.4 (8.6)
6.7-7.3 (6.9)
11.1-12.5 (11.9)
18.0-19.7 (19.1)
7.7-8.6 (7.9)
14.7-17.1 (15.9)
12.3-16.6 (14.0)
17.9-22.7 (20.2)
26.9-29.4 (28.1)
12.8-16.1 (14.3)
23.9-26.1 (24.4)
25.4-27.5 (25.8)
15.8-16.9 (16.4)
10.5-10.9 (10.7)

38.2
23.3
48.1
12.5
12.9
7.5
6.2
16.1
24.2
8.9
3.6
25.1
8.9
6.5
6.4
11.5
42.2
75.6
44.6
-3
-3
19.5
18.8
17.9
16.2
14.9
12.3
9.9
8.1
14.3
19.2
8.5
16.9
14.5
21.3
26.5
14.2
23.5
29.0
16.6
10.1

37.7-39.0
21.4-23.1
47.2-50.4
11.8-12.3
13.1-14.4
5.4-5.6
23.1-24.2
7.1-8.4
3.7-4.2
23.9-25.8
40.3-42.5
71.2-73.0
44.4-45.4
9.2-9.7
15.6-16.7
20.0-21.2
7.9-8.9
14.4-14.6
20.6
9.1-10.0
17.8
15.3-16.1
19.8-21.2
26.1-29.8
15.2
22.1-22.3
28.9-30.6
15.7-16.1
9.5-10.5

36.6-41.8 (39.4)
20.0-26.5 (22.7)
45.7-50.4 (48.0)
11.8-14.4 (12.9)
11.0-15.5 (12.9)
5.4-8.0 (6.5)
4.7-6.6 (5.7)
14.2-16.1 (15.5)
22.5-24.5 (23.5)
6.2-8.4 (7.2)
1.8-4.8 (3.1)
21.9-26.1 (23.9)
7.5-9.3 (8.5)
5.5-6.7 (6.1)
5.7-6.8 (6.3)
7.6-10.5 (9.0)
36.5-42.8 (39.9)
71.2-78.0 (74.3)
42.8-48.3 (45.1)
7.8-10.9 (9.1)
14.2-18.3 (15.7)
19.0-22.8 (20.4)
17.4-22.0 (19.5)
16.4-21.8 (18.7)
16.2-19.6 (17.6)
14.7-18.7 (16.3)
13.2-16.7 (14.8)
10.6-14.4 (12.5)
8.5-12.0 (9.8)
6.6-10.2 (8.3)
12.7-16.0 (14.5)
18.2-22.0 (19.7)
7.9-10.5 (9.2)
16.7-20.2 (18.8)
14.1-18.0 (15.7)
19.1-23.5 (20.6)
23.4-31.6 (27.5)
14.0-19.6 (15.8)
20.4-25.5 (22.9)
26.1-31.2 (28.7)
15.1-18.0 (16.2)
9.5-11.2 (10.3)

36.5-45.1 (40.9)
17.7-28.6 (24.1)
44.8-50.1 (47.6)
12.4-14.5 (13.5)
9.7-14.2 (11.8)
4.8-7.5 (6.3)
4.1-7.0 (5.8)
15.6-16.5 (16.0)
22.1-26.3 (23.8)
6.5-8.4 (7.4)
2.5-5.7 (4.2)
21.8-26.2 (24.0)
6.9-9.7 (8.6)
6.1-7.6 (6.8)
6.2-7.1 (6.6)
6.2-9.6 (8.3)
37.0-42.3 (39.4)
71.2-78.7 (74.3)
37.7-48.0 (44.9)
6.5-11.4 (8.8)
11.7-18.1 (15.1)
16.7-23.3 (20.0)
16.9-21.8 (19.4)
16.3-22.5 (18.8)
14.7-20.2 (17.4)
14.5-18.5 (16.5)
12.6-17.3 (14.8)
10.8-14.4 (12.7)
7.5-11.8 (10.0)
7.1-9.8 (8.7)
11.1-16.3 (14.1)
17.7-23.1 (21.0)
6.0-10.1 (8.8)
14.5-21.2 (17.8)
13.6-17.9 (16.1)
19.3-23.6 (21.7)
25.8-30.2 (28.2)
12.5-18.0 (15.7)
22.3-27.1 (24.5)
27.7-33.0 (29.7)
14.8-18.4 (16.4)
9.2-12.4 (11.2)

1At vertical midline of eye; 2at posterior end of preocular spine base; 3probable regenerated spine; -no data.

38

Zoological Studies 46(1): 35-45 (2007)

between base of last dorsal-fin spine and lateralline 8 (7-9). Gill rakers on upper limb 5; lower limb
10 (11 in 1 paratype), including 2 (3 in 1 paratype)rakers on hypobranchial; gill rakers short and spinous, longest raker on 1st gill arch shorter thanlongest gill filament; 4th gill slit closed by membrane. Vertebrae 24. Swimbladder absent.
Body moderately compressed anteriorly, progressively more compressed posteriorly; nape andanterior part of body moderately arched; bodymoderately deep, but less than head length.
Numerous small papillae on head, especially on
dorsal surface, snout, and lateral surface above
suborbital ridge; numerous tiny tentacles and cirrion head, including dorsal margin of eye; a very
short tentacle at posterior end of supraocular spine
base, its length about 1/5 (1/3 in 1 paratype;absent in 1 paratype) pupil diameter; a large fimbriate flap at posterior lacrimal spine, its length
greater than supraocular tentacle, joined posteriorly

posterior branch of last soft ray not joined by membrane to caudal peduncle; membrane between softrays weakly incised. Pectoral fin with 16 rays, anuppermost ray and lower 10 rays (9 in 2
paratypes) unbranched, remaining rays branched;
8th ray longest, length less than head length; lower
unbranched rays thickened; distal margin of fin
slightly bilobed. Pelvic fin with 1 spine and 5 softrays, all soft rays branched; 2nd soft ray longest,
length longer than longest dorsal-fin spine; last soft
ray joined by membrane to body for more than 1/2
its length. Caudal fin with 14 segmented rays,
most rays branched; posterior margin of fin slightly
rounded.
Longitudinal scale series 47 (46-48); pored
lateral-line scales 23 (22 in 2 paratypes); scales
above lateral line 7 (6 in 2 paratypes), below 16(17 in 1 paratype); predorsal scale rows 6 (5 in 1
paratype); scale rows between base of 6th dorsalfin spine and lateral line 8 (6 in 1 paratype),

Table 2. Frequency distribution of selected meristic characters in Scorpaena pepo
sp. nov. and the southwestern Pacific (S) and northwestern Pacific (N) populations of
S. onaria
Pectoral-fin rays (one side/other side)

Longitudinal scale series

16/16 16/17 17/17 17/18 18/18
S. pepo sp. nov.
S. onaria (S)S. onaria (N)S. pepo sp. nov.
S. onaria (S)S. onaria (N)

3

19
25

4
2H

1

45

46

47

48

6
11H

9
6

2
4
7

1H
2
1

1

2
4

22

23

24

15

16

17

2
2
1

2H
20
26H

1
2

7
19H

3H
13
8

1
3

Scales above / below lateral line
4

5

6

7

2

2
13
1

2H

10
26H

/

Predorsal scale rows

Scales between base of lastdorsal-fin spine and lateral line

3

4

5

6

7

8

9

13
3

1
7
19

3H
1
8H

1
1
18H

2H
22
10

1

3

4

5

4
4

4H
20
25H

HIndicates the holotype.

1

Gill rakers
Lower limb

Upper limb

S. pepo sp. nov.
S. onaria (S)S. onaria (N)44

4H

Pored lateral-line scales

S. pepo sp. nov.
S. onaria (S)S. onaria (N)

43

6

10

11

1

3H
4
19H

1
11
11

Total
12

9

14

15

16

17

2
3

3H
3
17H

1
11
9

8
1

Motomura et al. -- New Species of Scorpaena from Taiwan

to head by fringed skin; a few small tentacles
associated with pored lateral-line scales; pectoralfin axil without a skin flap; no distinct tentacles onlateral surface of body, except at pored lateral-line
scales, on fin rays and membranes, on maxilla,
and on underside of lower jaw.
Ctenoid scales covering an area between
posterior tips of upper and lower opercular spines,
but not on opercular margin; cycloid scales, covered with thin skin, covering an area surrounded
by upper preopercle, lower posttemporal, andupper opercular spines; embedded cycloid scales
covering an area adjacent to postorbital, sphenotic, pterotic, and upper preopercular spines; otherparts of head not covered with scales; wellexposed ctenoid scales covering entire lateral surface of body, but not extending onto rays or membranes of fins, except at base of caudal fin;embedded cycloid scales covering ventral surface
of body, including breast and between pelvic fins,and pectoral-fin base; scales on pectoral-fin basecovered by epidermis. Lateral line sloping steeply
downward above proximal 1/2 of pectoral fin.
Mouth large, slightly oblique, forming an angleof about 40 (30 in 1 paratype) to horizontal axisof head and body; posterior margin of maxilla
extending well beyond a vertical through posterior
margin of orbit; lateral surface of maxilla smooth,
without distinct ridges; lower jaw with a symphysialknob. Width of symphysial gap separating premaxillary teeth bands slightly greater than width ofeach band; premaxilla with a band of short,incurved, conical teeth; tooth band narrowing posteriorly; tooth band on premaxilla wider than that oflower jaw; lower jaw with a band of villiform teeth,most teeth shorter than those of upper jaw; small
teeth on vomer and palatines. Underside of dentary with 3 small sensory pores on each side, 1stpore below anterior lacrimal spine, 2nd pore below
and between anterior and posterior lacrimal
spines, 3rd pore located on posterior margin of
dentary; a pair of small pores behind symphysial
knob of lower jaw; a pair of small pores on side ofsymphysial knob; underside of lower jaw smooth,
without ridges.
Dorsal profile of snout steep, forming anangle of about 50 to horizontal axis of head andbody. Nasal spine simple, conical, directed
upwards, its length slightly greater than anterior
nostril diameter. Anterior nostril with a distinct tentacle, its length greater than nasal spine.
Ascending process of premaxilla not intruding into
interorbital space, its posterior margin not extending beyond level with posterior margin of posterior

°°

°

39

nostril. A well-developed median interorbital ridge
running from posterior margin of ascending
process of premaxilla to posterior end of supraocular spine base (slightly posterior to posterior end of
preocular spine base in 2 paratypes); medianinterorbital ridge higher than interorbital ridges inlateral view. Interorbital ridges well developed,
separated by a deep channel, beginning posterior
to a line through posterior margins of posterior
nostrils and then conjoined level with, and between
origins of, tympanic spines, forming a distinct ridge
to anterior angular edge of occipital pit; interorbital
ridges diverging anteriorly and posteriorly in dorsal
view, with least distance between interorbital
ridges narrowest at level of anterior margin of
pupil; interorbital space moderately deep, about
1/4 of orbit extending above dorsal profile of head.
Preocular spine simple (with an additional small
spine laterally in 1 paratype), directed nearly dorsally. Supraocular spine simple, its tip locatednear vertical midline of eye, spine shorter than preocular, postocular, and tympanic spines.
Postocular spine simple (with 2 points in 1
paratype), its length slightly less than tympanic
spine. Tympanic spine simple, pungent; bases of
tympanic spines joined with interorbital ridges anda transverse ridge in front of occipital pit.
Interorbital, coronal, and pretympanic spines
absent. A distinct transverse ridge (formed fromposterior part of interorbital ridges) anterior to
occipital pit slightly curved posteromedially in dorsal view; occipital pit relatively shallow, center of
pit usually slightly convex; a distinct transverse
ridge in rear of occipital pit between bases ofnuchal spines; occipital pit surrounded laterally by
tympanic and parietal spines, and low ridges
between tympanic and parietal spines on eachside. Parietal spine simple, its base curving
strongly into occipital pit. Nuchal spine simple(with 2 points in 1 paratype); nuchal and parietal
spines joined at base. Sphenotic with 2 (3 in 1
paratype) small, sharp spines. Postorbital smooth,
embedded in skin (with 1 or 2 slender spines in 3
paratypes). Pterotic spine simple, pointed. Upper
posttemporal spine simple, pointed, small, directed
posteriorly (ventroposteriorly in 2 paratypes), its
length much shorter than lower posttemporalspine. Lower posttemporal spine simple, its baselength approximately equal to that of pterotic
spine. Supracleithral spine simple, not strongly
pointed. Cleithral spine flattened, pointed.
Lateral surface of lacrimal with a strongly
pointed spine, spine located below anterior marginof orbit. Anterior lacrimal spine pointed, directed40

Zoological Studies 46(1): 35-45 (2007)

forward, its tip reaching dorsal margin of upper-jaw
lip; an additional short spine near posterior base ofanterior lacrimal spine, the additional spine directed anteriorly. Posterior lacrimal spine simple,
directed ventroposteriorly, its tip not reaching
upper-jaw lip; posterior lacrimal spine larger thananterior spine. Suborbital ridge with 3 spines, posterior end of 1st spine base below and between
posterior margins of pupil and orbit, 2nd spine
extending beyond orbit, 3rd spine at end of suborbital ridge. Broad space between ventral margin ofeye and suborbital ridge. Suborbital pit absent.
Preopercle with 5 spines, uppermost largest, witha supplemental preopercular spine on its base;
2nd to 5th spines without a median ridge, directed
posteriorly; 2nd spine smallest, with narrow base;

3rd to 5th spines with broad base. Dorsal ramus
of preopercle without serrae or spines. Upper
opercular spine simple, with a low median ridge.
Lower opercular spine simple with a distinct median ridge. Space between upper and lower opercular spines without distinct ridges (with a low ridgein 1 paratype) and not covered with fleshy skin.
Posterior tips of upper and lower opercular spines
not reaching opercular margin.
Origin of 1st dorsal-fin spine anterior to baseof supracleithral spine. Posterior margin of opercular membrane extending beyond a verticalthrough origin of 4th dorsal-fin spine. Posterior tipof pectoral fin not reaching a vertical through tip ofdepressed pelvic fin and origin of last dorsal-fin
spine. Posterior tip of depressed pelvic fin not

(A)

(B)

Fig. 1. Color photographs of Scorpaena pepo sp. nov. (A) ASIZP 65020, holotype, male, 244.3 mm SL, off the northeastern coast of
Taiwan; (B) ASIZP 65021, paratype, female, 245.1 mm SL, collected with holotype. Note that the body color is slightly faded in photographs taken 2 days after collection. Photos by H. Motomura.

Motomura et al. -- New Species of Scorpaena from Taiwan

reaching anus. Origin of pelvic-fin spine posterior
to origin of pectoral fin. Origin of 1st anal-fin spine
posterior to origin of last dorsal-fin spine.
Color when fresh: Head and body bright yellowish-orange (Fig. 1; yellow and light orange pigments not evident in photos taken 2 days after collection), mottled with very pale gray blotches dorsally; small distinct black spots scattered on headand body, except for ventral surface, median finsand upper 1/2 of pectoral fin. Underside of lower
jaw bright orange, mottled with distinct white
blotches. One or 2 irregular narrow white bands
across spinous portion of dorsal fin medially; membrane of spinous portion of dorsal fin yellowishorange, mottled with poorly defined whitish andblackish blotches; a large black blotch, greaterthan orbit diameter, on outer margins of membranes between 6th and 11th (tenth in 1 paratype)spines in males (absent in females). Soft-rayed
portion of dorsal fin orange proximally, grayish
subdistally, with poorly defined whitish blotches
and blackish spots over fin; distal margin of the fin
whitish. Pectoral-fin base whitish; pectoral fin yellowish-orange with small black spots scattereddorsally, poorly defined whitish blotches ventrally.
Pelvic and anal fins yellowish-orange, with poorly
defined whitish blotches. Caudal fin yellowishorange basally, gray subdistally, whitish distally,

41

with irregular white markings arranged into weakly
organized transverse bars, and with scatteredsmall indistinct black spots. Color when alive
unknown.
Color of preserved specimens: Head andbody dark yellow dorsally, mottled with large poorly
defined gray blotches, becoming pale yellow ventrally (Fig. 2A). Small distinct black or gray spots,
averaging about 1/10 body scale diameter in size,
scattered over head and body, except for on ventral surfaces, median fins, and dorsal 1/2 of pectoral fin. An oblique, broad, blackish band fromventral margin of orbit to behind posterior marginof maxilla. Lips and underside of mandible paleyellow with poorly defined gray blotches anteriorly.
Dorsal fin yellowish-white with a large dense black
blotch, greater than orbit diameter, on outer margins of membranes between 6th and 11th (10th in
1 paratype) spines in males (absent in females).
Pectoral fin yellowish with black spots on upper 1/2
of fin; inner surface of pectoral fin yellowish withindistinct grayish blotches centrally; axil of fin yellowish, without markings. Pelvic fin uniformly yellowish with 2 or 3 scattered gray spots. Anal fin
yellowish, with a few scattered pale black spots.
Caudal fin translucent yellow with small black

(A)

(A)

(B)
(B)

Fig. 2. Preserved specimens of Scorpaena pepo sp. nov. (A)and S. onaria (B). (A) AMS I. 43631-001, paratype, male,
223.1 mm SL off the northeastern coast of Taiwan (photo by C.
Bento); (B) MUFS 14402, male, 164.6 mm SL, off thesoutheastern coast of Kyushu I. southern Japan (photo by H.
Motomura).

Fig. 3. Color variation in Scorpaena onaria. (A) MUFS 12919,
male, 142.2 mm SL, off the southeastern coast of Kyushu I.
southern Japan (photo by H. Motomura); (B) FRLM 6927,
male, 178.5 mm SL, off Mie, central Japan (photo by S.
Kimura).

42

Zoological Studies 46(1): 35-45 (2007)

spots basally and centrally.
Distribution: Currently known only from off thenortheastern coast of Taiwan, probably on deepwater rocky reefs.
Etymology: Derived from the Latin pepo
meaning“pumpkin”, alluding to the yellowishorange body color of this species when fresh.

DISCUSSION
Scorpaena pepo sp. nov. and S. onaria
Jordan and Snyder, 1900 resemble each other inoverall appearance and are distinguished from allother congeners in the Indo-Pacific Ocean by thefollowing combination of characters: the lateral surface of the lacrimal with a single spine, the posterior margin of the anterior lacrimal spine with anadditional spine, and the median interorbital ridge
well developed, embedded cycloid scales covering
the anteroventral surface of the body and thepectoral-fin base, a high body depth (36.4-45.1%
of SL), 43-48 longitudinal scale series, and the lateral line sloping steeply downward above the anterior 1/2 of the pectoral fin (Motomura et al. 2005b,
this study). Incidentally, these 2 species appear to
be closely related to a Tahitian species, S. lacrimata Randall and Greenfield, 2004, and a Hawaiian
species, S. pele Eschmeyer and Randall, 1975,
but the 2 can be distinguished by 43-48 longitudinal scale series (vs. 61 in S. lacrimata; Randall
and Greenfield 2004) and the well-developed
median interorbital ridge (vs. absent in S. pele;
Motomura et al. 2005b).
Scorpaena onaria, originally described from
Misaki, Japan, was redescribed by Motomura et al.
(2005b), with discussions of its distribution, morphological changes with growth, sexual dichromatism, and biology. The species is found in thenorthwestern Pacific (southern Korea, Japan, andTaiwan), southwestern Pacific (eastern Australia
and northern New Zealand), and eastern Indian
Oceans (off northwestern Australia and in the
Andaman Sea), but has not been reported fromthe western central Pacific Ocean (Motomura et al.
2005b). A fish reported as Scorpaena ballieui by
FSFRL (1976) from the North Pacific about midway between Japan and the Hawaiian Is. at a
depth of 77 m is probably S. onaria, judging from a
photograph (M.Pac-23) of the fish published by
FSFRL (1976). However, the whereabouts of thespecimen is currently unknown, and the locality
cannot be verified.
Motomura et al. (2005b) recognized 2 geo-

graphically distinct populations for S. onaria: anorthern population from the northwestern Pacificand a southern population from the southwestern
Pacific. Motomura et al. (2005b) regarded specimens of S. onaria from northwestern Australia
probably assignable to the southern population,
but the status of S. onaria from the Andaman Sea
remains unknown because specimens from theAndaman Sea are no longer available.
Scorpaena pepo can be distinguished from S.
onaria by having 16 pectoral-fin rays (vs. usually
17 rays on each side of the body (ca. 82% of 54
specimens examined), rarely asymmetrically 16
and 17 rays (ca. 6%) or 17 and 18 rays (ca. 10%),and rarely 18 rays (ca. 2%) on each side in S.
onaria; see Table 2). Five larger individuals of S.
pepo (not retained), examined by the 1st author at
the market in Taipei, also had 16 pectoral-fin rays
on each side of the body. Mean values of thenumber of the pectoral-fin rays are 16.0 for S.
pepo, 16.9 for the southern population of S. onaria
and 17.0 for the northern population. In S. pepo,
the number of scale rows counted in longitudinal
series seems to be slightly greater (46-48, mean
47.0) than that of S. onaria (43-47, mean 44.8,
based on 52 specimens; see Table 2), althoughexamination of more specimens of S. pepo isrequired.
The morphology of the head, including thespine and ridge structures, of S. pepo is very similar to that of S. onaria (head of the latter is illustrated in Motomura et al. 2005b: fig. 3). However, theposterior margin of the maxilla in both sexes of S.
pepo extends well beyond a vertical through theposterior margin of the orbit, whereas that in both
sexes of the northern population and males of thesouthern population of S. onaria is located below
the eye between the center of the pupil and theposterior margin of the orbit. In mature females
from the southern population of S. onaria it
extends slightly beyond a vertical through the posterior margin of the orbit (Motomura et al. 2005b).
The posterior end of the 1st suborbital spine base(on infraorbital 2, not the lateral lacrimal spine) in
S. pepo is located below the eye between the posterior margins of the pupil and orbit, whereas thatof S. onaria is located below the eye between thecenter and posterior margin of the pupil (Motomura
et al. 2005b; this study). The median interorbital
ridge of S. pepo is well developed, with its highest
part higher than the interorbital ridges. In S. onaria
this ridge is lower than the interorbital ridges
throughout its length.
In the field, the yellowish-orange coloration

Motomura et al. -- New Species of Scorpaena from Taiwan

easily distinguishes S. pepo from S. onaria, whosecoloration is primarily red (Fig. 3; color variationsof southern population of S. onaria were figured byMotomura et al. 2005b: fig. 1). In some near-shore
scorpaenids, body coloration varies with depth.
Individuals from shallow waters have brown or
black spots on the fins, while in specimens fromdeeper water, these same markings are often red(Motomura and Shinohara 2005, Motomura et al.
2005a). However, this does not appear to be true
in S. onaria, in which coloration of specimens
taken from 8 to 1000 m were usually all primarily
red (Fig. 3). Scorpaena pepo has numerous small
distinct black (sometimes gray in preservative)
spots scattered over the head, and body, andmedial fins (Figs. 1, 2A). Such spots are also present on the upper 1/2 of the pectoral fin. Although
S. onaria also has similar spots on its body andfins, they are usually absent from the head, except
rarely for a few that may be present on the opercle, and are usually fewer in number than seen in
S. pepo (see Figs. 2B, 3; Motomura et al. 2005b:
figs. 1, 2). Also in S. pepo, the spots are relatively
small, their size averaging about 1/10 of a bodyscale diameter. In S. onaria, the diameter of an
average spot is about 1/3 that of a body scale(Motomura et al. 2005b; Figs. 2B, 3).
Scorpaena pepo and the southern populationof S. onaria have a short supraocular tentacle,
which is usually shorter than the pupil diameterand the tentacle on the posterior lacrimal spine(the supraocular tentacle is absent on the rightside of the head in 1 paratype of S. pepo). In contrast, the supraocular tentacle of the northern population of S. onaria is well developed and greaterthan the pupil diameter (often greater than the orbit
diameter) and the length of a tentacle on the posterior lacrimal spine (Motomura et al. 2005b). The
length of the supraocular tentacle of S. pepo is
1.7-3.9% (mean 3.1%) of SL, compared to 4.911.3% (mean 7.6%) of SL for the northern population of S. onaria (based on 19 specimens). As
Motomura et al. (2005b) pointed out, the ventral
margin of the dentary of individuals of the southern
population of S. onaria is more convex, and thehead is more rounded than for individuals of thenorthern population. The condition of the dentaryin S. pepo is similar to that of the southern population of S. onaria.
The largest specimen of S. pepo collected(ASIZP 65021, northeastern Taiwan) and thelargest individual from the southern population of
S. onaria (NMNZ P. 40691, northern New Zealand;
Motomura et al. 2005b) are 245.1 and 265.2 mm

43SL, respectively. In contrast, the largest recorded
size of the northern population of S. onaria is187.9 mm SL (MUFS 14395, southern Japan; thisstudy). Scorpaena pepo reaches at least 40 cm
total length (Motomura, pers. obs., see“Introduction”). Thus, S. pepo is more similar to the southern population of S. onaria rather than the northern
population, even though S. pepo co-occurs with
the northern population of S. onaria, at least in
Taiwan. However, with the exception of the presence or absence of a large black blotch on the
spinous portion of the dorsal fin, the sexual dichromatism exhibited in the southern population of S.
onaria (see Motomura et al. 2005b: 878-879) was
not found in either S. pepo or the northern population.
The following additional characters of S. pepo
appear to be diagnostic (but examination of more
specimens is needed): the median interorbital
ridge ending posterior to midway between the posterior ends of the preocular spine bases in dorsal
view, sometimes ending midway between the posterior ends of the supraocular spine bases (vs.
ending anterior to midway between the posterior
ends of the preocular spine bases in S. onaria;
Motomura et al. 2005b, this study); and the postorbital with 1 or 2 slender, sharp spines on bothsides of the head (only the holotype lacks thespine), which are longer than the sphenotic spines(vs. the postorbital smooth, covered with skin,without the slender spines in S. onaria; Motomura
et al. 2005b, this study).
There are several minor morphometric differences, e.g., orbit diameter, and spine and soft ray
lengths, between S. pepo and the northern population of S. onaria (Table 1). However, these proportions change with growth (Motomura et al. 2005b:
877-878, fig. 7), and as mentioned above, there isa size difference between specimens of S. pepo
(172.9-245.1 mm SL) and of the northern population of S. onaria (81.1-187.9 mm SL), suggestingthat these morphometric differences are probably
not reliable for species discrimination. Smaller
specimens of S. pepo need to be analyzed for
growth-related changes of the species, as done forthe southern population of S. onaria by Motomura
et al. (2005b).
Comparative material examined: 54 specimens of Scorpaena onaria. Data for 19 specimens(121.0-265.2 mm SL; deposited at AMS, CAS,
MNHN, and NMNZ) from the southwestern Pacific
Ocean were listed in Motomura et al. (2005b) andadditional 5 specimens (56.5-164.3 mm SL) fromthe area are as follows: CSIRO CA 3172, 120.5

44

Zoological Studies 46(1): 35-45 (2007)

mm SL, Tasman Sea (33 08'S, 156 11'E), 142 m
depth, coll. by FRV Soela, 28 Jan 1982; CSIRO H
643-18, 160.1 mm SL, northeast of Whitsunday
Group (19 37-38'S, 150 30-32'E), Queensland,
312-318 m depth, coll. by FRV Soela, 15 Nov.
1985; CSIRO H 2466-02, 153.1 mm SL, Lord
Howe Rise, Tasman Sea, 1000 m depth, coll. by
A. Hinson; CSIRO H 6007-15, 56.5 mm SL, southof Norfolk I. (29 41-42'S, 168 01-02'E), Norfolk
Ridge, Tasman Sea, 322-337 m depth, coll. by
FRV Tangaroa, 14 May 2003; CSIRO H 6011-01,
164.3 mm SL, north of Norfolk I. (28 53-54'S, 167
41-42'E), Norfolk Ridge, Tasman Sea, 110-441 m
depth, coll. by FRV Tangaroa, 15 May 2003. Data
for 30 specimens (81.1-187.9 mm SL) from thenorthwestern Pacific Ocean are as follows: CAS
106275 (2 paratypes), 136.6-144.8 mm SL, Tokyo,
Japan, coll. by K. Otaki; CAS 125248, 81.1 mm
SL, Oki-shima Island, Sea of Japan, coll. by U.S.
Fish Commission, 2 Aug. 1906; FRLM 6927, 178.5mm SL, Koshika, Shima, Mie, Japan, 110 m depth,line-fishing, coll. by S. Kimura, 22 Dec. 1987;
FRLM 8518, 117.3 mm SL, same data as FRLM
6927 except depth, 60-100 m, and date, 25 Jan.
1989; FRLM 21012, 126.2 mm SL, Wagu, Shima,
Mie, Japan, 60-100 m depth, gill net, coll. by T.
Yamashita and T. Sado, 22 Oct. 1997; FRLM
21879, 124.5 mm SL, Goza, Shima, Mie, Japan,
30-60 m depth, line-fishing, coll. by T. Yodo, 31
Mar. 1998; MUFS 12916-12920 (5 specimens),
120.3-153.3 mm SL, off Nango, Miyazaki, Japan,
purchased at Meitsu Fish Market by H. Motomura,
19 Apr. 1997; the following 4 lots with the samecollecting data as MUFS 12916 except for thedate: MUFS 12993, 181.2 mm SL, 20 Dec. 1997;
MUFS 13180, 157.4 mm SL, 17 May 1997; MUFS
14402, 164.6 mm SL, 21 Dec. 1997; MUFS
16396, 147.3 mm SL, 14 Sept. 1998; MUFS
13407-13409 (3 specimens), 144.1-180.9 mm SL,
off Kawaminami, Miyazaki, Japan, purchased at
Kawaminami Fish Market by H. Motomura, 5 June
1997; MUFS 14395-14400 (6 specimens), 110.2187.9 mm SL, off Amami-oshima, Kagoshima,
Japan, purchased at Meitsu Fish Market by H.
Motomura, 20 Dec. 1997; NSMT-P 19211, 114.0mm SL, Yawatano, Izu, Japan, 100 m depth, coll.
by S. Tomotake, 12 Apr. 1968; NSMT-P, 48664,
157.7 mm SL, off Akajima Island, Goto Islands,
Nagasaki, Japan, coll. by A. Morota, 4 Aug. 1995;
NTUM 1435, 141.9 mm SL, off Hualien, Taiwan;
OMNH 13888, 144.7 mm SL, off Himi, Toyama,
Sea of Japan, purchased at a fish market by K.
Hatooka, 22 Apr. 2000; USNM 49405 (holotype),157.7 mm SL, Misaki, Japan, coll. by K. Otaki.

°

°

°

°

°

°

°

°

Acknowledgments: We are grateful to S. Kimura(FRLM), K. Hatooka (OMNH) and J. T. Williams
and S. J. Raredon (USNM) for specimen loans of
Scorpaena onaria and Y. C. Liao (ASIZP) for
transport of the type specimens of S. pepo. We
wish to thank W. N. Eschmeyer, T. Iwamoto, D.
Catania and J. Fong (CAS), P. R. Last, A.
Graham, G. K. Yearsley, T. Fountain, L. Conboy
and W. T. White (CSIRO), P. Pruvost and N. Bailly(MNHN), Y. Iwatsuki, K. Hidaka, A. Chakraborty,
D. Kanbayashi, M. Taniguchi and T. Fukuhara(MUFS), A. L. Stewart and C. Roberts (NMNZ), K.
Matsuura and K. Shibukawa (NSMT), and K. Y.
Wu (NTUM) for their kind assistance during theperiod when the 1st author was examining specimens of S. onaria and S. pepo at their institutions.
,
The 1st author s visits to CSIRO; MNHN; andNMNZ were supported by the National Oceans
Office (Hobart) and CSIRO Marine Research(Hobart); the Food and Agriculture Organization of
the United Nations (Rome), the Instituto Españolde Oceanografia (Madrid) and W. N. Eschmeyer(CAS); and the Biosystematics of New ZealandExclusive Economic Zone Fishes Programme(New Zealand Foundation for Research Scienceand Technology contract MNZX0203), respectively. We thank S. Tsukawaki (Kanazawa University,,
Japan) for his kind support of the 1st author s trip
to Taipei. We are grateful to M. McGrouther, T.
Trnski, J. M. Leis, S. Reader, A. Hay, and all staffof the Fish Section, Australian Museum, Sydney
and Y. Motomura for their generous support during,
the 1st author s stay in Australia. This study was
supported in part by the Visiting Collection
Fellowship grant awarded to the 1st author by theAustralian Museum (Sydney, Australia).

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