Zoological Studies 43(1): 1-7 (2004)

Morphological Comparison of a Poorly Known Scorpionfish,
Parapterois macrura, with a Related Species, P. heterura(Scorpaenidae: Pteroinae)
Hiroyuki Motomura
Ichthyology, Division of Vertebrate Zoology, Australian Museum, 6 College Street, Sydney, New South Wales 2010, Australia
Tel: 61-2-93206310. E-mail: motomurah@austmus.gov.au(Accepted September 9, 2003)

Hiroyuki Motomura (2004) Morphological comparison of a poorly known scorpionfish, Parapterois macrura,with a related species, P. heterura (Scorpaenidae: Pteroinae). Zoological Studies 43(1): 1-7. The taxonomic
status of a scorpionfish Parapterois macrura (Scorpaenidae: Pteroinae), originally described as Pterois macrurafrom the Malabar Coast off Calicut, west coast of India by Alcock (1896), has long been in doubt. Comparisonof P. macrura collected from Indian waters with a related congener P. heterura (Bleeker, 1856), originallydescribed from Ambon, Indonesia and subsequently collected from the southeast coast of Africa and western
Pacific Ocean, revealed that the former was distinguished from P. heterura by the following characters: a scaleless interorbital space, lower counts of longitudinal scale series, greater head and snout lengths and head
depth, a shorter tentacle on the posterior spine of the lacrimal, the upper margin of the orbit higher than or levelwith the base of the first dorsal spine (in lateral view) in adults and top of a bulge on the snout much lower thanthe ventralmost margin of the orbit (in lateral view) in adults. Morphological changes with growth for both
species and populations in P. heterura are also discussed. http://www.sinica.edu.tw/zool/zoolstud/43.1/1.pdf
Key words: Comparison, Scorpaenidae, Parapterois macrura, Parapterois heterura.

Tand Yazdani 1968, Menon and Rama-Rao 1975),
no reports of Pterois (= Parapterois) macrura have,been published since Alcock s (1896) originaldescription and subsequent report (Alcock 1898),
although Parapterois heterura is well known.
Thus, the taxonomic status and characters of P.
macrura have remained obscure.
Examination of Parapterois specimens collected from India showed them to represent P.
macrura, which could be distinguished from P.
heterura on the basis of several morphologicalcharacters recognized in this study. The present
paper includes confirmation of the validity of P.
macrura, assessments of diagnostic characters of
P. heterura and P. macrura proposed in the key by
Mandrytsa (2001), descriptions of newly recognized diagnostic characters and morphologicalchanges with growth for both species, and comments on populations in P. heterura.

he genus Parapterois Bleeker, 1876
(Scorpaenidae: Pteroinae), proposed as a new
genus for Pterois heterurus Bleeker, 1856, originally described from Ambon, Indonesia, is characterized by high pectoral fin ray counts (18-21 vs.
12-17 in the related genus Pterois Oken), a low
anal fin spine counts (2 vs. 3), some branchedpectoral fin rays (vs. unbranched), and a truncatedcaudal fin with the upper and lower rays longest inlarge adults (vs. a rounded caudal fin without longrays) (Eschmeyer and Rama-Rao 1978,
Eschmeyer 1986). Although Parapterois has long
been considered a monotypic genus (Eschmeyer
1986, Ishida 1997), Mandrytsa (2001) treated apoorly known scorpionfish, Pterois macrura
Alcock, 1896, originally described from the
Malabar Coast off Calicut, west coast of India, asa second species of Parapterois and provided a
key to the two species. With the exception of
Mandrytsa (2001) and some type catalogs (Menon
1

2

Zoological Studies 43(1): 1-7 (2004)

Institutional codes follow Leviton et al. (1985),
with additional institutional abbreviations as follows:
Division of Fisheries Sciences, Faculty ofAgriculture, Miyazaki University, Japan (MUFS);South African Institute for Aquatic Biodiversity,
South Africa (SAIAB, formerly RUSI); and Raffles
Museum of Biodiversity Research, Department of
Biological Sciences, National University ofSingapore (ZRC, formerly NMS). The following
specimens of Parapterois species were examined
during this study: Parapterois heterura from western Indian Ocean (14 specimens, 39-116 mm SL) SAIAB 802 (holotype of Pterois natalensis vonBonde, ,1923),,, 66 mm, SL,,, Natal coast, South Africa(29 26 30 S, 31 25 40 E), 53 m, 30 July 1920;
SAIAB 3023, 1 specimen, 79 mm SL, off
Mozambique, May 1973, coll. by B. Andrews;
SAIAB 48157, 4 specimens,,,87-116 mm SL, off
Mozambique (19 51 S, 36 18 E), bottom trawl, 66
m, 18 June 1994, coll. by R. W. Leslie; SAIAB
62730, 1 specimen, 64 mm SL, mouth of St., Lucia,,
River, ,KwaZulu-Natal,
South Africa (28 23 30 S,,,
32 26 10 E), 35 m, 22 Sept. 1999, coll. by S.
Fennessy; SAM 11855 (syntypes of Pterois nigripinnis Gilchrist, 1904), 3 specimens, 39-51 mm SL,
4 km off mouth of Umhlanga River, Natal, South
Africa, 40-48 m, 26 Mar. 1901, coll. by P. Faure;
SAM 34263, 2 specimens, , 87-112 mm, SL, off
Beira, Mozambique (19 51 S, 36 18 E), 66 m;
SAM 34264, 2 specimens,
70-79, mm SL, off Beira,,Mozambique (19 49 S, 36 05 E), 54 m. Parapterois heterura from western Pacific Ocean (12specimens, 57-139 mm SL) - MUFS 16696, 1
specimen, 62 mm SL, off Meitsu, Nango, Miyazaki,
Japan, 23 Nov. 1998, coll. by H. Motomura; MUFS
17718, 1 specimen, 67 mm SL, off Meitsu, Nango,
Miyazaki, Japan, 5 June 1999, coll. by H.
Motomura; MUFS 17730-17731, 2 specimens, 6574 mm SL, off Meitsu, Nango, Miyazaki, Japan, 12June 1999, coll. by H. Motomura; NSMT-P 34409,

MATERIALS AND METHODSSpecimen lengths are given as standard
length (expressed as SL), measured from theanteriormost tip of the upper lip to the posterior
end of the hypural plate. Head and snout lengths
were measured from the anteriormost tip of theupper lip to the posterior end of the opercular flap
and to the anterior margin of the eye, respectively.
Head depth was taken obliquely from the uppermost margin of the orbit (frontal) to the posterior
corner (near the preopercle) of the suborbital ridge(see Fig. 1). Pored lateral line scales could not becounted in this study because all of the specimens
examined were in poor condition, owing to theirhaving been caught by trawl nets. Scales in thelongitudinal series, being nearly vertical to theoblique scale rows above the lateral line from theposterior point of the supracleithral spine base to
the caudal fin base, were usually counted on thebasis of deciduous scale traces. Gill raker counts
included all rudiments.

°

°

°

°

°

°

°
°

°

Fig. 1. Head of Parapterois macrura (MUFS 16910, 110 mm
SL, off Malpe, Karnataka, west coast of India) in lateral view.
Head depth is defined in this study as a straight line from theuppermost orbit margin to the suborbital ridge. Numerous
scales and tiny spines are not illustrated.

°

Table 1. Frequency distribution of selected counts in Parapterois species
Anal fin spines Anal fin rays

P. heterura
P. macrura

Pectoral fin rays

2

7

8

18

19

20

43

44

45

46

47

48

49

50

51

21
11

20
11

1
-

2
-

11
7

10
4

1

1

1
2

3
2

3
1

3
1

3
1

1
-

1
-

Upper gill rakers

P. heterura
P. macruraLongitudinal scale series

Lower gill rakers

Total gill rakers

4

5

6

10

11

12

13

14

15

16

17

18

19

1
-

23
5

2
6

2
-

10
4

8
7

5
-

1
-

2
-

10
1

7
7

6
3

1
-

3

Motomura -- Comparison of Parapterois spp.

1 specimen, 88 mm SL, Futo, Ito, Sagami Bay,
Shizuoka, Japan, 2-3 m, 11 July 1991, coll. by M.
Aizawa; NSMT-P 34604, 1 specimen, 89 mm SL,
Tosa Bay, Kochi, Japan, bottom trawl, 30 m, 1 Oct.
1989, coll. by O. Okamura; NSMT-P 46604, 1
specimen, 57 mm SL, off Osezaki, Suruga Bay,
Shizuoka, Japan, bottom trawl, 150-300 m, 26 Oct.
1994, coll. by H. Saito; NSMT-P 63542, 1, specimen,
86, mm SL, East China Sea (27 57-59 N, 125 5659 E), bottom trawl, 117-118 m, 20 Oct. 2001, coll.
by T. Saito and H. Namikawa; RMNH 5872 (holotype of Pterois heterurus Bleeker, 1856), 78 mm
SL, Ambon, Maluku Is., Indonesia; URM-P 17403,
1 specimen, 139 mm SL, Chinen, Okinawa,
Ryukyu Is., 30 Apr. 1986, coll. by T. Yoshino; URMP 17420, 1 specimen, 114 mm SL, Chinen,
Okinawa, Ryukyu Is., 5 May 1986, coll. by T.
Yoshino; ZRC 728, 1 specimen,
96 mm,, SL, South
China Sea (05 22-25 N, 107 36-47 E), 70-73 m,
11 June 1956, coll. by M. F. V. Manihine. Parapterois macrura (11 specimens, 77-111 mm SL) MUFS 16908-16910, 3 specimens, 78-110 mm SL,
off Malpe, Karnataka, west coast of India, bottom
trawl, 80 m, 30 Mar. 2000, coll. by H. Motomura
and Y. Iwatsuki; ZMH 5598, 7 specimens, 77-111mm SL, off near
Karwar,,, Karnataka, west coast ofIndia (14 42 N, 73 54 E), bottom trawl, 75 m, 15
Feb. 1965, coll. by A. Kotthaus; ZMH 5599, 1 specimen, 81 mm SL, off near Cochin, west coast of

°

°

°

°

°

°

,
,
India (9 40 N, 75 38 E), bottom trawl, 138-210 m,
10 Feb. 1965, coll. by A. Kotthaus. Localities of
specimens of both species examined during thisstudy are indicated in figure 2.

°°

RESULTS AND DISCUSSION
Validity of Parapterois macrura
Parapterois macrura was originally describedas Pterois macrurus from the Malabar Coast off
Calicut, west coast of India by Alcock (1896).
Since the original description and a subsequent
report (Alcock 1898), there have been no further
references to that species, except in type catalogs(Menon and Yazdani 1968, Menon and Rama-Rao
1975), until Mandrytsa (2001) recognized both P.
heterura (see Figs. 3A, 4A) and P. macrura (seeFigs. 3B, 4B) as valid species of Parapterois andprovided a key to the species. In his key,
Mandrytsa (2001) briefly noted that P. macrura differed from P. heterura in having the upper marginof the orbit higher than the level of the first dorsal
fin spine base (vs. slightly lower in the latter), theinterorbital space without scales (vs. covered withscales), a tentacle on the lacrimal not reaching theposterior margin of the maxilla (vs. extending●Parapterois heterura★P. macrura

N
30°

0°
S

30°
N

30°
E

60°
E

90°
E

120°
E

Fig. 2. Collection localities of specimens of Parapterois species examined in this study. Circles and stars indicate P. heterura and P.
macrura, respectively.

4

Zoological Studies 43(1): 1-7 (2004)

beyond it), and a flap with small black spots present on the tip of the first dorsal fin spine (vs. flap
absent).
A flap on the tip of the first dorsal fin spine,
was not apparent in Alcock s (1898) original figureof Pterois macrura, nor in examples of that species
examined during this study (owing to their poor
condition, as the dorsal fin spines and membranesof most were damaged). This character still needs
confirmation. Furthermore, careful examination of
the orbit/dorsal fin spine character and lacrimaltentacle length revealed that these were not,always reliable for species discrimination (seebelow).
Examination during this study of specimens(11 specimens, 77-111 mm SL) of Parapterois collected from the west coast of India (Fig. 2) showed
them to lack scales on the midline of the narrowinterorbital space. On the other hand, the interorbital space of all specimens (26 specimens, 39139 mm SL) of P. heterura, collected from South
Africa, Mozambique, Indonesia, South China Sea,
East China Sea, and Japan (Fig. 2), was covered
with scales. Accordingly, the Indian and otherspecimens of Parapterois could be readily distinguished from each other by this single character.
Parapterois heterura (type locality: Ambon,
Indonesia), including 6 known nominal species,
viz. Ebosia pavo Schmidt, 1931 (Nagasaki,
Japan), E. starksi Franz, 1910 (Kanagawa,Japan), Pterois jordani Regan, 1905 (Inland Sea,
Japan), P. natalensis von Bonde, 1923 (Natal,
South Africa), P. nigripinnis Gilchrist, 1904 (Natal,
South Africa), and P. tanabensis Tanaka, 1918
(A)

(Wakayama, Japan), which have been treated as
junior synonyms of P. heterura (see Eschmeyer
and Rama-Rao 1978, Eschmeyer 1998), has at notime been reported from Indian waters, including
the west coast of India. Although the type specimens of P. macrura (registered in the Zoological
Survey of India, Calcutta) could not be examined
during this study, both the type locality and,Alcock s (1896) description of a scaleless interorbital space in that species, support the identification of the Indian west coast specimens examined
here as P. macrura being a valid species.
Parapterois macrura is currently known only
from the west coast of India (Alcock 1896; thisstudy: Fig. 2), whereas P. heterura is widely distributed in scattered localities of the Indo-West
Pacific, viz. east coast of South Africa, Mozambique, Indonesia (Ambon), South China Sea,
Taiwan, East China Sea and southern Japan(Eschmeyer 1986, Poss 1999; this study: Fig. 2).
Comparisons
Meristic characters: Dorsal fin spines andrays, and anal fin spines of Parapterois macrura
numbered the same (viz. XIII, 9, and II, respectively) as those of P. heterura, with no apparent variations. Although Eschmeyer (1986) described P.
heterura as having 2 anal fin spines in adults and 3
in young, all of the specimens examined (39-139mm SL) had 2 anal fin spines (Table 1). In fact,
the number of anal fin spines is unlikely to changewith growth in either P. heterura or P. macrura.
The numbers of anal and pectoral fin rays,
(A)

(B)
(B)

Fig. 3. Adults of Parapterois heterura (A) and P. macrura (B).
(A) SAIAB 48157, 104 mm SL, off Mozambique; (B) ZMH 5598,
104 mm SL, off Karwar, west coast of India.

Fig. 4. Young of Parapterois heterura (A) and P. macrura (B).
(A) ZRC 728, 96 mm SL, South China Sea; (B) MUFS 16909,
83 mm SL, off Malpe, west coast of India.

5Motomura -- Comparison of Parapterois spp.

and total gill rakers also showed no significant differences between the 2 species (Table 1).
However, the number [mean 45.9 (range 43-49)] ofscales in the longitudinal series in P. macrura tended to be slightly greater than those [47.7 (45-51)]
in P. heterura (Table 1). Furthermore, P. heterura
had a strong modal count of 5 gill rakers on theupper limb, whereas P. macrura had equally 5 or 6
gill rakers on that element (Table 1).
Morphometric characters: Parapterois macrura is very similar to P. heterura in overall body
appearance (see Figs. 3-4). Parapterois macrura,
however, significantly differed from P. heterura inhaving greater head length [mean 42.4% of SL(range 39.7-44.1% of SL) vs. 38.2% (34.9-39.7%)
in the latter; Fig. 5A], snout length [mean 15.8% of
SL (range 12.9-19.1% of SL) vs. 12.6% (11.214.1%); Fig. 5B] and head depth [mean 23.0% of
SL (range 21.0-25.1% of SL) vs. 19.0% (17.620.4%); Fig. 5C]. With the exception of those proportions, no other significant differences in proportional measurements were found between the 260
Head length (mm)

(A)
50
40
30

Snout length (mm)

10
25

(A)

(B)

(B)

20
15
10
5
0
30

Head depth (mm)Parapterois heterura
P. macrura

20

species.
Head morphology: Parapterois species havea tentacle on the posterior spine of the lacrimal.
The tip of the tentacle in P. macrura usually failed
to extend beyond the level of the posterior marginof the maxilla (see Fig. 1), although those in 2
smaller specimens (77-78 mm SL), which comprised 18.2% of the total number of specimens
examined (11 specimens, 77-111 mm SL), did soslightly. Contrary to P. macrura, P. heterura had along tentacle, usually extending beyond the levelof the posterior margin of the maxilla, althoughthose of 2 larger specimens (114-139 mm SL),
which comprised 7.7% of the total number of specimens examined (26 specimens, 39-139 mm SL),
failed to reach the posterior margin.
The upper margin of the orbit of larger P.
macrura (7 specimens, 79-111 mm SL) was levelwith or higher than the base of the first dorsal fin
spine (in lateral view), whereas that of smaller
ones (4 specimens, 77-83 mm SL) was below thelevel of the first dorsal fin spine base (Fig. 6B). In
P. heterura, the upper margin of the orbit was considerably below the level of the first dorsal fin spinebase (in lateral view) throughout life (Fig. 6A).
Therefore, although larger P. macrura could bereadily distinguished from P. heterura by the orbit/
dorsal fin spine character, smaller examples of the
2 species were difficult to distinguish from eachother solely by that character. Accordingly, the 2

(C)

25
20
15
10
5
20

40

60

80
100
120
Standard length (mm)

140

160

Fig. 5. Relationships of head length (A), snout length (B) andhead depth (C) to standard length in Parapterois heterura (circles) and P. macrura (stars).

Fig. 6. Heads of Parapterois heterura (A) and P. macrura (B)in lateral view. Upper and lower drawings indicate larger andsmaller specimens of each species, respectively. Horizontal
lines indicate levels of the uppermost margin of the orbit andtop of a bulge on the snout. (A, upper), URM-P 17403, 139
mm SL; (A, lower), MUFS 17718, 67 mm SL; (B, upper), MUFS
16910, 110 mm SL; (B, lower), MUFS 16908, 78 mm SL.
Numerous scales and tiny spines are not illustrated.

6

Zoological Studies 43(1): 1-7 (2004)

above-mentioned characters are not always reli,able for species discrimination, although Mandry,tsa (2001) used them as species diagnostic characters.
The top of a bulge on the snout of larger P.
macrura (6 specimens, 101-111 mm SL) was considerably below the level of the ventralmost marginof the orbit (in lateral view), whereas that of smaller specimens (5 specimens, 77-83 mm SL) was
located between the levels of the ventralmost margins of the pupil and orbit (Fig. 6B). The top of asimilar bulge on the snout level with the middle ofthe pupil was found in all specimens examined of
P. heterura, except for 2 larger specimens (114139 mm SL) in which the top of the bulge was
located between the levels of the ventralmost margins of the pupil and orbit (Fig. 6A).
Change of nasal spines with growth: Exami
nation of different life stages of P. heterura and P.
macrura showed that the number of nasal spines
in both species increased to 2 or more with growth.
In P. macrura, young (77-79 mm SL) and adults(81-114 mm SL) had a single and 3 nasal spines
on one side, respectively, 3 other adults (110 mm
SL, 105 mm SL and 104 mm SL) having 5, 6 and 7
nasal spines on one side, respectively. On theother hand, P. heterura young (39-97 mm SL) andadults (87-139 mm SL) had a single and 2 nasal
spines on one side, respectively, with the exception of a single adult (116 mm SL) which had 3
nasal spines on one side.
Populations of P. heterura: A pit between theposterior nostrils of all P. macrura specimens
examined was not covered by scales. Although P.
heterura collected from the western Indian Ocean,viz. South Africa and Mozambique, had a naked pit
as in P. macrura, P. heterura collected from thewestern Pacific Ocean, viz. South China Sea, East
China Sea, and Japan, had a scaly pit. Additional
differences between western Indian and western
Pacific Ocean populations of P. heterura were notfound during this study. The 2 populations might
be different species. Future studies, including molecular analyses, will help elucidate the taxonomic
status of these 2 populations.
Key to species and populations of Parapterois
1a.

Scales absent on midline of interorbital space throughout
life; longitudinal scale series 43-49 (mean 45.9); headlength 39.7-44.1% (42.4%) of SL; snout length 12.919.1% (15.8%) of SL; head depth 21.0-25.1% (23.0%) of
SL; tip of tentacle on posterior spine of lacrimal notextending beyond level of posterior margin of maxilla

1b.

2a.
2b.

(sometimes extending slightly beyond in young); upper
margin of orbit higher than or level with base of first dorsal
spine (in lateral view) in adults, lower than in young; top ofbulge on snout well below ventralmost margin of orbit (inlateral view) in adults, located between levels of ventralmost margins of pupil and orbit in young ...........................
........................................ P. macrura (west coast of India)
Scales present on midline of interorbital space throughout
life; longitudinal scale series 45-51 (mean 47.7); headlength 34.9-39.7% (38.2%) of SL; snout length 11.214.1% (12.6%) of SL; head depth 17.6-20.4% (19.0%) of
SL; tip of tentacle on posterior spine of lacrimal usuallyextending beyond level of posterior margin of maxilla(sometimes not reaching it in adults); upper margin oforbit well below level of first dorsal spine base (in lateralview); top of bulge on snout usually level with midline ofpupil (in lateral view), sometimes located between levelsof ventralmost margins of pupil and orbit in adults .......... 2
Pit between posterior nostrils covered with scales ............
............................ P. heterura (western Pacific population)
Pit between posterior nostrils naked ..................................
............... P. heterura (southeast African coast population)

Acknowledgments: I am grateful to the following
persons and institutions for specimen loans: Y.
Iwatsuki (MUFS); K. Matsuura and G. Shinohara(NSMT); M. J. P. van Oijen and J. van Egmond(RMNH); P. C. Heemstra, M. E. Anderson and V.
Mthombeni (SAIAB); M. G. van der Merwe and L.
Hoenson (SAM); T. Yoshino (URM); H. Wilkens(ZMH); and P. K. L. Ng and K. K. P. Lim (ZRC). I
extend my gratitude to M. N. Venugopal (Department of Fishery Biology, College of Fisheries,
Mangalore) and A. Chakraborty (MUFS) for theirassistance in field work at Mangalore, India and H.
,
Kohno (MTUF) for his translation of Mandrytsa s(Russian) paper into Japanese. I also thank Y.
Motomura (Sydney, Australia) for her assistanceand G. S. Hardy (Ngunguru, New Zealand), whoread the initial manuscript and offered helpful comments. This study was supported in part by a
postdoctoral fellowship grant awarded to theauthor by the Japan Society for the Promotion of
Science for Research Abroad (Tokyo, Japan).
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