Ichthyol Res (2011) 58:315–332
DOI 10.1007/s10228-011-0234-2

FULL PAPER

Validity of Scorpaena jacksoniensis and a redescriptionof S. cardinalis, a senior synonym of S. cookii(Scorpaeniformes: Scorpaenidae)
Hiroyuki Motomura • Carl D. Struthers •Mark A. McGrouther • Andrew L. Stewart

Received: 29 April 2011 / Revised: 14 June 2011 / Accepted: 14 June 2011 The Ichthyological Society of Japan 2011

Abstract The Scorpaena cardinalis complex, including
S. cardinalis, S. jacksoniensis and S. orgila, is defined. The
genus Ruboralga (type species: S. jacksoniensis) is regarded as a junior synonym of Scorpaena. Scorpaena jacksoniensis Steindachner 1866, previously treated as a juniorsynonym of Scorpaena cardinalis Solander and Richardson
1842, is regarded here as a valid species. Scorpaena cookiiGünther 1874, previously treated as a valid species, isregarded here as a junior synonym of S. cardinalis. Thus,
recent recognition of the two Australasian scorpionfishes,i.e., S. cardinalis and S. cookii, are re-identified in thisstudy as S. jacksoniensis and S. cardinalis, respectively.
Scorpaena plebeia Solander 1842 is regarded as a juniorsynonym of S. cardinalis. Scorpaena jacksoniensis is distributed along the east coast of Australia from southern
Queensland to eastern Victoria, whereas S. cardinalis
occurs around northern New Zealand, the Kermadec
Islands and offshore islands of the Tasman Sea. A neotype
is designated for S. cardinalis. Morphological changes withgrowth in the two species are described in detail.
Keywords Ruboralga  Scorpaena jacksoniensis Scorpaena cardinalis  Scorpaena cookii  Synonymy

H. Motomura (&)
The Kagoshima University Museum, 1-21-30 Korimoto,Kagoshima 890-0065, Japane-mail: motomura@kaum.kagoshima-u.ac.jp
C. D. Struthers  A. L. Stewart
Museum of New Zealand Te Papa Tongarewa,
P. O. Box 467, Wellington, New Zealand
M. A. McGrouther
Australian Museum, 6 College Street, Sydney,
NSW 2010, Australia

Introduction
During revisionary studies of the genus Scorpaena (Scorpaeniformes: Scorpaenidae) by the first author, examination of the holotype of Scorpaena jacksoniensis
Steindachner 1866a found this nominal species to be avalid species, although it has been treated as a juniorsynonym of Scorpaena cardinalis Solander and Richardson
in Richardson (1842) by numerous authors (e.g., Macleay
1881; Allen and Cross 1989; Allen et al. 2006). Subsequently, examination of the original drawing of S. cardinalis revealed it to be identical with Scorpaena cookiiGünther 1874, but distinct from S. jacksoniensis. Therefore, S. cookii, long regarded as a valid species by
numerous authors (e.g., Ogilby 1889; Poss 1999; Allenet al. 2006; Roberts et al. 2009), is placed here as a juniorsynonym of S. cardinalis. Accordingly, the two recently
recognized Australasian scorpionfishes, i.e., S. cardinalis
and S. cookii, are re-identified in this study as S. jacksoniensis and S. cardinalis, respectively. Scorpaena plebeia
Solander in Richardson (1842) is also considered as a
junior synonym of S. cardinalis. The genus Ruboralga(type species: S. jacksoniensis) is regarded as a juniorsynonym of Scorpaena.
The two species, S. jacksoniensis (as S. cardinalis) andS. cardinalis (as S. cookii), have been considered to be
distributed around both Australia and New Zealand waters(e.g., Paulin 1982; Poss 1999; Allen et al. 2006). Examination of a large number of specimens from the southwestern Pacific Ocean disclosed that the previous
distribution range of each species was incorrect owing tomisidentifications and that the two species in fact represented allopatric distributions, i.e., S. jacksoniensis in the
coastal water of eastern Australia; S. cardinalis in northern
New Zealand and offshore islands of the Tasman Sea.

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The two species are redescribed, and a neotype of S.
cardinalis is designated. Morphological changes withgrowth in the two species are also described in detail. The
S. cardinalis complex is defined in this study, and includes
S. cardinalis, S. jacksoniensis and Scorpaena orgila
Eschmeyer and Allen 1971 (endemic to Easter Island,
Chile).

Materials and methods
Measurements generally follow Motomura (2004a, b),
except head width (Motomura et al. 2005b, 2006a) andmaxillary depth (Motomura et al. 2006b). Counts follow
Motomura et al. (2005a, b, c) and Motomura and Johnson(2006), with predorsal scale counts following Motomura
et al. (2006b). The last two soft rays of the dorsal and anal
fins are counted as single rays, each pair being associatedwith a single pterygiophore. Standard and head lengths are
expressed as SL and HL, respectively. Terminology ofhead spines follows (Randall and Eschmeyer 2002: fig. 1)
and Motomura (2004b: fig. 1) with the following additions:the spine occurring at the base of the uppermost preopercular spine is referred to as the supplemental preopercularspine (Eschmeyer 1965); the spine occurring at the lateralsurface of the lacrimal bone is referred to as the laterallacrimal spine (Motomura and Senou 2008: fig. 2); thecoronal and pretympanic (as an extra spine) spines are
figured in Chen (1981: fig. 1) and Motomura et al. (2004:
fig. 14b), respectively. Institutional codes follow Leviton
et al. (1985), with the exception of the Australian National
Fish Collection at the Commonwealth Scientific andIndustrial Research Organisation’s Marine and Atmospheric Research laboratories, Hobart (CSIRO), Kagoshima University Museum, Kagoshima (KAUM) and Museumof New Zealand Te Papa Tongarewa, Wellington (NMNZ).
The following specimens of Scorpaena orgila were
examined for comparative purposes: CAS 24809, holotype,246.8 mm SL, off Ahu Akapu, Easter Island, Chile, spear,
J. Randall, 3 Feb. 1969; CAS 24810, 2 paratypes,
59.8–109.7 mm SL, off south end of Hanga Roa, Easter
Island, Chile, J. Randall and G. Allen, 10 Feb. 1969; CAS
24811, paratype, 75.5 mm SL, wreck about 20 m offshore
between Hanga Roa and Hanga Piko, Easter Island, Chile,
J. Randall and G. Allen, 27 Jan. 1969.

The Scorpaena cardinalis complex
Included species. Scorpaena cardinalis, S. jacksoniensis
and S. orgila.
Diagnosis. Lateral lacrimal spine with 2 spinous points(Fig. 1).

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Fig. 1 Ontogenetic changes of lacrimal bones of Scorpaena
cardinalis (a, b) and S. jacksoniensis (c, d). a AMS IB.5335,
110.0 mm SL; b NMNZ P.041669, 211.2 mm SL; c KAUM–I.
35882, 131.1 mm SL; d AMS I.30310-025, 173.6 mm SL. ALS
anterior lacrimal spine, PLS posterior lacrimal spine, LLS laterallacrimal spine, FP first spinous point, SP second spinous point. Bars
5 mm

Description. Dorsal fin with 12 spines and 9 soft rays
(rarely XIII, 8); fourth spine (rarely third spine) longest;
membrane of spinous portion of dorsal fin moderately
incised; second or third (rarely fourth) soft ray longest, its
length subequal to that of longest dorsal-fin spine; posterior
branch of last soft ray joined by membrane to caudal
peduncle for more than half its length. Anal fin with 3
spines and 5 soft rays; second spine longest; all soft rays
branched; first soft ray longest, its length longer than thatof longest dorsal-fin soft ray; posterior branch of last soft
ray joined by membrane to caudal peduncle for less than
one-tenth its length. Number of branched pectoral-fin rays
increasing with growth; lower unbranched rays thickened;
posterior margin of fin rounded. Pelvic fin with 1 spine and5 soft rays, all soft rays branched; second soft ray longest,
its length slightly longer than upper-jaw length; last soft
ray joined by membrane to abdomen for about half its
length. Caudal fin with 11 branched rays; posterior marginof fin slightly rounded. Gill rakers 14–19, relatively shortand spinous, longest raker on first gill arch shorter than gill
filaments around angle of gill arch; fourth gill slit closed bymembrane. Branchiostegal rays 7. Swimbladder absent.
Body moderately compressed anteriorly, progressively
more compressed posteriorly. Nape and anterior bodymoderately arched. Body depth slightly less than headlength. Numerous tiny tentacles on head. Posterior lacrimal
spine tentacle linked posteriorly to head by skin. A shorttentacle on posterior edge of low membranous tube associated with anterior nostril; the tentacle reaching posterior
margin of posterior nostril when laid back. SupraocularSynonymy of the Scorpaena cardinalis complex

tentacle usually present. Pectoral-fin axil without skinflaps. Exposed cycloid scales covering an area surroundedby opercular margin and tips of upper and lower opercularspines, other parts of head not covered with exposed scales.
Cycloid scales covering pectoral-fin base, some scales
exposed, some scales embedded by thin skin. Body scales
cycloid (often with weak cteni) in adults, ctenoid dorsallyin young; body scales extending onto basal rays or membranes of pectoral and caudal fins.
Mouth large, slightly oblique. Lower jaw with a moderate-sized symphysial knob. Width of symphysial gap
separating premaxillary teeth bands narrower than width ofeach band. Teeth on vomer and palatines. Underside oflower jaw smooth without ridges. Dorsal profile of snout
steep. Nasal spine simple, sharp, directed dorsally, its
length greater than anterior nostril diameter. Ascending
process of premaxilla not intruding into interorbital space,its posterior margin reaching level with posterior margin of
posterior nostril in dorsal view. Median interorbital ridge
well developed. Interorbital ridges well developed posteriorly, separated by a deep channel; interorbital ridges
initially diverging posteriorly in dorsal view and thenconjoined at occipital pit or joined with tympanic spine
base. Interorbital space deep, about one-third of orbit
extending above dorsal profile of head. Preocular spine
simple, directed dorsoposteriorly; tip of spine extending
beyond level with upper margin of pupil in lateral view.
Supraocular and postocular spines simple, well developed.
Tympanic spine simple, strongly pointed, directed posterodorsally. Interorbital, coronal and pretympanic spines
absent. Occipital pit moderately deep in young, becoming
shallower with growth, occiput nearly flat or slightly convex in large adults; longitudinal length of pit subequal towidth of pit. Occipital pit surrounded laterally by tympanic
and parietal spines and indistinct ridge between spines.
Parietal spine simple, its base curving strongly into
occipital pit. Nuchal spine simple; nuchal and parietal
spines joined at base. Sphenotic usually with two small
spines. Postorbital with tiny spines. Pterotic spine usuallysimple in young, with 2 or more points in adults in S.
jacksoniensis. Upper posttemporal spine simple, pointed,
its length shorter than that of lower posttemporal spine.
Lower posttemporal spine simple, its base length subequal
to that of pterotic spine. Supracleithral spine simple.
Cleithral spine flattened, with 2 spines.
Lateral lacrimal spine usually with 2 points. Anterior
lacrimal spine with 2 or more points, directed forward, its
tip reaching dorsal margin of upper lip. Posterior lacrimal
spine with 1 or 2 points, its tip not reaching upper lip;
length of posterior lacrimal spine subequal to that ofanterior spine. Suborbital with two ridges; first suborbital
spine at end of first ridge, located below pupil; second andthird suborbital spines on second ridge, located behind317

orbit. Space between ventral margin of eye and suborbital
ridge moderate. Suborbital pit present. Preopercle with fivespines; uppermost spine largest with a supplemental preopercular spine on its base. Preopercle, between uppermost
preopercular spine and upper end of preopercle, smooth
without serrae or spines. Upper opercular spine simple witha low median ridge; posterior tip of upper opercular spine
not reaching opercular margin. Lower opercular spine
simple with a distinct median ridge. No ridges between
upper and lower opercular spines.
Origin of first dorsal-fin spine above supracleithralspines. Posterior margin of opercular membrane reaching a
vertical through fourth dorsal-fin spine base. Posterior tipof pectoral fin not reaching a vertical through first anal-fin
spine base. Origin of pelvic-fin spine slightly posterior to
origin of pectoral fin. Posterior tip of depressed pelvic finreaching anus, but not reaching first anal-fin spine base.
Origin of first anal-fin spine slightly posterior to origin oflast dorsal-fin spine.
Remarks. The Scorpaena cardinalis complex is
known only from the South Pacific and comprises twoAustralasian species, S. cardinalis and S. jacksoniensis,
and an Easter Island endemic species, S. orgila. The
complex is characterized by having the lateral lacrimal
spine with two spinous points (Fig. 1), a character uniqueamong Indo-Pacific species of Scorpanea. Although S.
papillosa (Schneider and Forster in Bloch and Schneider
1801) has two upwardly directed spines on the lacrimalbone, the anterior spine arises from the anterodorsal
margin of the lacrimal bone and is not a part of laterallacrimal spine.
The posteroventrally directed posterior lacrimal spine
has been regarded as a diagnostic character for the IndoPacific Scorpaena (Poss 1999; Motomura et al. 2005c;
Motomura and Senou 2008). However, the posterior lacrimal spine of S. jacksoniensis is directed anteroventrally atleast in juveniles (posteroventrally in adults; see account of‘Morphological change with growth’), although, with the
exception of the posterior lacrimal spine condition, S.
jacksoniensis shared all the diagnostic characters of the
Indo-Pacific species of Scorpanea, i.e., 12 dorsal-fin spines,
9 dorsal-fin soft rays, teeth on the palatine, an occipital pit,
cranial bones strongly or moderately ossified, upper andlower opercular spines, some pectoral-fin rays branched,
the lateral surface of the body covered with cycloid or
ctenoid scales, and the lateral line with pored scales
continuing onto the caudal-fin base.

Scorpaena cardinalis Solander and Richardson 1842
(Standard English name: Sandy-Bay Cod; Australian name:Cook’s Scorpionfish; New Zealand name: Red Rockcod)(Figs. 1a, b, 2, 3, 4, 5, 6, 9, 10, 11; Tables 1, 3).

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Fig. 2 Preserved specimens of
Scorpaena cardinalis. a NMNZ
P.024541, 99.9 mm SL;
b BMNH 1855.8.16.88,
holotype of Scorpaena cookii,
216.2 mm SL; c NMNZ
P.034406, 440.6 mm SL

Scorpaena cardinalis Solander and Richardson in
Richardson 1842: 212 [original locality: Motuaro, Queen
Charlotte’s Sound, New Zealand (probably Motuaro, theBay of Islands, New Zealand; see Paulin 1982); type
locality: White Island, New Zealand, 37300 S, 177090 E,
based on neotype]; Whitley 1931: 326 (New Zealand);Paulin 1982: 439 (between Bay of Islands and White
Island, New Zealand); Doak 1984: 26, pls. 5, 59 (New
Zealand); Francis 1996: 24, pl. 30 (Three Kings Islands toEast Cape, New Zealand); Francis 2001: 33, pl. 38 (Kermadec Islands, New Zealand).
Scorpaena plebeia Solander in Richardson 1842: 214(type locality: Tolaga, New Zealand).
Scorpaena cookii Günther 1874: 78, pl. 55 [type locality:
Raoul Island (Sunday Island), Kermadec Islands, New Zealand]; Ogilby 1889: 155 (Lord Howe Island, Australia; newrecords from Australian water); Waite 1904: 220 (Lord Howe

123

Island); Eschmeyer and Allen 1971: 521 (Norfolk Island andKermadec Islands); Paulin 1982: 440 (Raoul Island, Kermadec Islands); Kuiter 1993: 119 (offshore reefs in NSW);Francis 1993: 159 (Lord Howe Island, Norfolk Island andKermadec Islands); Poss 1999: 2304, 2334 (in part, Lord
Howe Island, Coffs Harbour, and Middleton and Elizabeth
Reefs, NSW; Kermadec Islands); Paulin et al. 2001: 168 (New
Zealand); Francis 2001: 34, pl. 39 (Kermadec Islands, New
Zealand); Motomura et al. 2005c: 866 (Lord Howe Island andAuckland); Allen et al. 2006: 885 (in part, Lord Howe Island,
Coffs Harbour, Middleton and Elizabeth Reefs, and Kermadec Islands); Hirt-Chabbert 2006: 59, unnumbered fig.
(northern North Island, New Zealand).
Ruboralga cardinalis: Whitley 1968: 84 (New Zealand;
new combination; description also contains S. papillosa).
Ruboralga cookii: Whitley 1968: 84 (New Zealand, newcombination).

Synonymy of the Scorpaena cardinalis complex

319

Fig. 3 Fresh specimens of
Scorpaena cardinalis. a NMNZ
P.039245, 208.1 mm SL (photo:
R. McPhee/K. Parkinson);b NMNZ P.044152, neotype of
S. cardinalis, 308.1 mm SL(photo: C. Struthers)

Fig. 4 Distributional records ofScorpaena cardinalis (stars)
and S. jacksoniensis (circles)
based on collected specimens
examined in this study

Neotype. NMNZ P.044152, 308.1 mm SL, White Island,
New Zealand, 37300 S, 177090 E, 19–21 m, rod and line,
C. Struthers, 16 Mar. 2009.
Other material examined. 66 specimens, 25.3–472.5mm SL. NORFOLK ISLAND: AMS IB.5335, 110.0 mm
SL, off Kingston, 29030 S, 167570 E, B. Marsh and L.
Marsh, 16 Feb. 1961; AMS IB.5348, 306.2 mm SL, off
Kingston, 29030 S, 167570 E, 60 m, B. Marsh and L.
Marsh, 31 Mar. 1961; NMNZ P.014089, 256.9 mm SL,

29050 S, 167590 E, P. Roberts, 5 Sept. 1972; NMNZ
P.024541, 99.9 mm SL, Slaughter Bay, 29040 S, 167580 E,
2 m, M. Francis, 23 Nov. 1989; NMNZ P.024542, 2,
161.1–190.2 mm SL, Duncombe Bay, 29040 S, 167580 E,
15 m, M. Francis, 15 Nov. 1989; NMNZ P.026902,
136.4 mm SL, Crystal Pool, 29050 S, 167590 E, 4 m,
rotenone, M. Francis and J. Randall, 15 Feb. 1991.
LORD HOWE ISLAND: AMS I.5846, 236.4 mm SL, E.
Waite and A. McCulloch, 1903; AMS I.10623, 226.7 mm

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H. Motomura et al.

Fig. 5 Drawing by S.
Parkinson (404 mm total
length). Basis for originaldescription of Scorpaena
cardinalis by Solander andRichardson in Richardson(1842). The British Museumof Natural History, London

Fig. 6 Typical caudal-fin base color patterns of Scorpaena cardinalis. a NMNZ P.026902, 136.4 mm SL; b NMNZ P.039107,
272.2 mm SL. Arrows indicate white blotches on the caudal-fin base123

SL, Lord Howe Rise, 31310 S, 159050 E, P. Pedley, 1910;
AMS I.17359-013, 190.6 mm SL, Sylphs Hole, 31320 S,
159040 E, 2–4 m, spear, G. Allen et al., Feb. 1973; AMS
I.17364-020, 246.1 mm SL, 31320 S, 159040 E, 1–2 m,
D. Hoese, 8 Feb. 1973; AMS I.17369-003, 4, 67.5–228.3 mm
SL, Salmon Beach, 31320 S, 159040 E, 1–2 m, G. Allenet al., 14 Feb. 1973; AMS I.41368-001, 248.9 mm SL,
31330 S, 159050 E, Sept. 2001; AMS I.42728-004,
242.3 mm SL, Lord Howe Rise, 31480 S, 159200 E,
66–88 m, orange roughy trawl, NORFANZ Expedition,
RV Tangaroa, 23 May 2003; BMNH 1926.6.30.76,
141.8 mm SL, W. Baxter; BPBM 38878, 46.4 mm SL,
6 m, rotenone, J. Randall et al., 7 Feb. 1973; BPBM 38879,
46.1 mm SL, 25 m, rotenone, J. Randall et al., 24 Feb.
1973; CAS 109141, 3, 171.2–224.7 mm SL, AMS team;
NMNZ P.039107, 272.2 mm SL, off Balls Pyramid, Lord
Howe Rise, 31480 S, 159200 E, 66–88 m, orange roughy
trawl, NORFANZ Expedition, RV Tangaroa, 23 May
2003; NMNZ P.039245, 208.1 mm SL, 31300 S, 159040 E,
30–50 m, rod and line, A. L. Stewart and B. Wood, RV
Tangaroa, 22 May 2003; QM I. 22913, 279.3 mm SL,
Elizabeth Reef, Qld, 29550 S, 159020 E, R. McKay andJ. Short.
KERMADEC ISLANDS: BMNH 1855.8.16.88, holotypeof Scorpaena cookii, 216.2 mm SL, Raoul Island; NMNZ
P.005982, 462.0 mm SL, Curtis Island, 30300 S, 178340 W,
15 m, hand-line, H. Finlayson, 24 May 1974; NMNZ
P.007536, 67.2 mm SL, Macauley Island, 30100 S,
179300 W, J. Yaldwyn and W. Spiekman, 1975; NMNZ
P.007537, 215.2 mm SL, Raoul Island, 29130 S, 178100 W,
E. McGregor, 1975; NMNZ P.009350, 222.2 mm SL,
Napier Rock, 29140 S, 177520 W, hand-line, K. Bailey and
M. Francis, 18 Aug. 1985; NMNZ P.009559, 1 of 2,
328.0 mm SL, Raoul Island, 29100 S, 177500 W; NMNZ
P.013517, 151.7 mm SL, Raoul Island, 29100 S, 177500 W,
1961; NMNZ P.015666, 191.8 mm SL, Boat Cove, Raoul
Island, 29100 S, 177500 W, hand-line, Kingsford and Smith,Synonymy of the Scorpaena cardinalis complex

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Table 1 Meristics and morphometrics, expressed as percentages of standard length, of Scorpaena cardinalis

Standard length (SL, mm)

Neotype of
S. cardinalis
NMNZ P.44152

Holotype of
S. cookii
BMNH 1855.8.16.88

Non-types of
S. cardinalis(n = 65)

308.1

216.2

25.3–472.5

XII, 9i?7?x= 18/18

XII, 9i ? 6 ? xi
= 18/18

XII, 8–9
i–ii ? 3–12 ?
v–xvi = 16–18/16–18

Meristics
Dorsal-fin rays
Pectoral-fin raysa

Modes
XII, 9i ? 5 ? xi
= 17/17

Scale rows in longitudinal series

62

73

60–74

68

Pored lateral-line scales

24

24

23–25

24

Scales above lateral line

8

8

6–9

8

Scales below lateral line

22

21

21–26

22

Scale rows between 6th dorsal-fin spine baseand lateral line

8

9

8–9

9

Scale rows between last dorsal-fin spine baseand lateral line

10

10

8–11

10

Pre-dorsal scale rows

8

6

4–8

6

Gill rakersb

5 ? 11 = 16

5 ? 12 = 17

4–5 ? 9–13 = 14–16

5 ? 12 = 17

Body depth

33.7

28.6

28.6–36.7

Body width

23.2

21.0

18.7–26.0

22.7

Head length

43.6

43.3

41.7–47.6

44.7

Snout length

14.0

12.3

12.1–15.6

13.8

Orbit diameter

7.8

9.8

6.9–13.4

9.4

Interorbital widthc

6.5

4.9

4.9–7.1

5.9

Interorbital widthd

5.6

5.2

4.7–6.2

5.3

Head width
Upper-jaw length

13.8
22.2

13.1
21.7

12.7–14.1
20.4–24.1

13.4
22.2

Morphometrics (%SL)

Means
32.7

Maxillary depth

6.1

5.9

5.5–7.2

6.3

Suborbital space

3.1

3.1

2.2–4.4

3.3

Postorbital length

23.4

22.1

19.9–25.6

23.1

Between tips of opercular spines

6.2

5.4

4.7–7.1

5.9

Occipital pit length

5.6

6.0

5.5–6.6

6.0

Occipital pit width

5.9

6.0

5.6–6.9

6.0

Post-occipital pit length

8.5

9.7

7.7–10.2

8.9

Predorsal-fin length

36.1

37.6

34.7–41.1

37.3

Preanal-fin length

72.1

70.2

69.9–77.7

73.0

Prepelvic-fin length

40.8

38.0

36.2–52.1

41.5

1st dorsal-fin spine length

7.1

6.2

3.8–9.5

7.1

2nd dorsal-fin spine length

Damaged11.1

7.6–15.7

11.9

3rd dorsal-fin spine length

15.2

14.7

12.5–19.6

15.9

4th dorsal-fin spine length

15.1

14.2

12.4–20.5

16.2

5th dorsal-fin spine length
11th dorsal-fin spine length

14.9
5.7

14.4
5.0

13.1–19.6
4.2–7.1

15.8
5.6

12th dorsal-fin spine length

11.2

Damaged8.9–14.4

11.6

Longest dorsal-fin soft ray length

16.7

17.3

14.8–20.6

17.3

1st anal-fin spine length

10.2

10.3

6.1–12.1

9.3

2nd anal-fin spine length

Damaged

15.4

10.3–22.5

16.7

3rd anal-fin spine length

13.5

14.0

10.6–19.6

14.7

Longest anal-fin soft ray length

21.4

21.8

17.0–24.0

21.1

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H. Motomura et al.

Table 1 continueda

Neotype of
S. cardinalis
NMNZ P.44152

Holotype of
S. cookii
BMNH 1855.8.16.88

Non-types of
S. cardinalis(n = 65)

Pectoral-fin ray length

26.8

29.9

23.2–31.7

Pelvic-fin spine length

13.2

14.3

10.3–18.9

14.6

Longest pelvic-fin soft ray length
Caudal-fin length

24.2
25.8

24.1
27.3

19.7–26.6
20.9–30.4

23.4
26.1

Caudal-peduncle length

15.9

17.6

15.6–20.4

17.3

Caudal-peduncle depth

10.5

10.5

9.4–11.6

10.5

27.8

Upper unbranched ray ? branched ray ? lower unbranched rays = total rays on left side of body/total rays on right sideb

Rakers on upper limb ? rakers on lower limb = total rakerscAt vertical midline of eyedAt posterior end of preocular spine base

20 Mar. 1984; NMNZ P.017782, 2, 221.2–360.3 mm SL,
Meyer Island, 29100 S, 177500 W, hand-line, M. Francis
and K. Baile, 18 Aug. 1985; NMNZ P.028550, 4,
25.3–166.4 mm SL, Boat Harbour, Meyer Island, 29140 S,
177520 W, 0–3 m, rotenone, M. Francis and C. Ward, 3 June
1992; NMNZ P.028585, 76.6 mm SL, Denham Bay, Raoul
Island, 29160 S, 177570 W, 12 m, rotenone, M. Francis andC. Ward, 4 June 1992; NMNZ P.034404, 433.5 mm SL,
Raoul Island, 29350 S, 178050 W, 129 m, drop-line, N.
Mitchell, 7 June 1997; NMNZ P.034405, 369.2 mm SL,
Raoul Island, 29350 S, 178050 W, 116 m, drop-line, N.
Mitchell; NMNZ P.034406, 3, 334.8–440.6 mm SL, Raoul
Island, 29320 S, 178050 W, 117 m, drop-line, N. Mitchell, 9June 1997; NMNZ P.034409, 2, 343.8–359.2 mm SL, Raoul
Island, 29350 S, 178050 W, 146 m, drop-line, N. Mitchell, 8June 1997; NMNZ P.034412, 1 of 2, 454.2 mm SL, Raoul
Island, 29340 S, 178050 W, 114 m, drop-line, N. Mitchell, 8June 1997; NMNZ P.034423, 1 of 2, 446.3 mm SL, Raoul
Island, 29300 S, 178020 W, 154 m, drop-line, N. Mitchell, 6June 1997; NMNZ P.034424, 2, 416.2–472.5 mm SL, Raoul
Island, 29300 S, 178020 W, 144 m, drop-line, N. Mitchell, 6June 1997; NMNZ P.034428, 364.7 mm SL, Raoul Island,29350 S, 178050 W, 130 m, drop-line, N. Mitchell, 9 June
1997; NMNZ P.034429, 455.3 mm SL, Raoul Island,29340 S, 178050 W, 120 m, N. Mitchell, 9 June 1997;
NMNZ P.041669, 211.2 mm SL, 500 m off Smith’s Bluff,
Denham Bay, Raoul Island, 29160 S, 178570 W, 14–16 m,
rotenone, A. L. Stewart et al., 10 Nov. 2004.
NEW ZEALAND: BMNH 1886.11.18.17, 247.7 mm
SL, Auckland; NMNZ P.002190, 295.1 mm SL, White
Island, 37310 S, 177110 E, 7.3 m, A. Black, 25 Mar. 1957;
NMNZ P.005652, 336.5 mm SL, Oke Bay, Bay of Islands,
35140 S, 174160 E, 37 m, hand-line, A. Baker, Nov. 1971;
NMNZ P.005779, 213.2 mm SL, north of Auckland, ca.
36S, 175E, trawl, 1970; NMNZ P.005946, 197.5 mm SL,
Poor Knights Islands, 35300 S, 174440 E, 6–7 m, W.

123

Palmer, June 1973; NMNZ P.007712, 264.2 mm SL, White
Island, 37310 S, 177110 E, 100 m, hook and line, RV Tangaroa, 19 Jan. 1979; NMNZ P.018162, 1 of 3, 295.8 mm SL,
east side of Mayor Island, 37180 S, 176180 E, 37–91 m, set
net, FV Asterix, 19 Feb. 1986; NMNZ P.019957, 240.9 mm
SL, Oke Bay, Bay of Islands, 35140 S, 174160 E, 37 m, handline, A. Baker, Nov. 1971; NMNZ P.021469, 367.2 mm SL,
southeast of Ruamahuanui Island, Alderman Islands, 36580 S,
176050 E, 40 m, spear, G. Hardy, 7 Dec. 1987; NMNZ
P.037092, 276.9 mm SL, White Island, 37300 S, 177090 E,
19–21 m, rod and line, C. Struthers, 16 Mar. 2009. LOCALITY UNKNOWN: AMS I.43239-001, 213.5 mm SL, purchased at Pyrmont Fish Market, Sydney, 2004.
Diagnosis. A species of Scorpaena with the following
combination of characters: dorsal-fin soft rays 9 (8 rays in
one specimen); fourth (rarely third) dorsal-fin spine longest; pectoral-fin rays 16–18 (mode 17); scale rows inlongitudinal series 60–74 (68); pored lateral-line scales
23–25 (24); scales above lateral line 6–9 (8), below 21–26(22); scale rows between sixth dorsal-fin spine base andlateral line 8 or 9 (9); scale rows between last dorsal-fin
spine base and lateral line 8–11 (10); pre-dorsal scale rows
4–8 (6); gill rakers on upper limb 4–6 (5), lower limb 9–13
(12) [8–10 (9) and 1–4 (3) rakers on ceratohyal andhypobranchial, respectively], total rakers 14–18 (17);exposed cycloid scales covering pectoral-fin base and anteroventral surface of body (some scales covered by thinskin, especially in large adults); lateral surface of maxilla
without a longitudinal ridge; lateral lacrimal spine with 2
spinous points; anterior lacrimal spine with 1–3 small
spinous points on its posterior margin; posterior lacrimal
spine with 1–3 spinous points, directed posteroventraly
throughout life; median interorbital ridge present; coronalspines absent; occipital pit and supplement preopercularspine present; pterotic spine simple, smooth distally
throughout life; space between upper and lower opercularSynonymy of the Scorpaena cardinalis complex

spines covered by thin skin without sensory pores or
canals; a few skin flaps or tentacles on anterodorsal surface
of body; no large black blotch on spinous portion of dorsal
fin in both sexes; 2 large white blotches, their diameters
subequal to pupil diameter, on caudal-fin base; largest
recorded specimen, 472.5 mm SL.
Distribution. The species is distributed around northern
New Zealand (north of Tolaga Bay, including the Kermadec
Islands) and offshore islands of the Tasman Sea (Norfolk andLord Howe Islands, and Middleton and Elizabeth Reefs, NewSouth Wales, Australia) (Fig. 4). Although Poss (1999) andAllen et al. (2006) included Coffs Harbour on the northerncoast of New South Wales in range of S. cardinalis (as S.
cookii) on the basis of a single specimen (AMS I.19696-007,
64.0 mm SL), the specimen is re-identified here as S. jacksoniensis. Accordingly, S. cardinalis has not been recorded
from the coastal waters of Australian mainland.
Collection data for 87 specimens of S. cardinalis
deposited at NMNZ indicate that the species inhabits rocky
reefs in depths of less than 154 m.
Remarks. Scorpaena cardinalis Solander and Richardson in Richardson (1842) was originally described on thebasis of Solander’s unpublished manuscript and a line
drawing by S. Parkinson. The unfinished drawing, reproduced by Whitehead (1969: pl. 31), shows that the fish(404 mm total length) has a relatively elongate body, theposterior lacrimal spine with 4 points and a pair of distinct
blotches on the caudal-fin base (see Fig. 5). These characters agree with those of specimens from New Zealandconsidered here as S. cardinalis (see ‘Comparisons’).
Typical color patterns on the caudal-fin base of S. cardinalis are shown in Fig. 6, which is consistent with that ofthe drawing (Fig. 5).
No type specimen of S. cardinalis is known, and thespecimen on which the drawing is based is no longer
extant. The erection of a neotype for S. cardinalis is hereinjustified, owing to the similarity of S. cardinalis to S.
jacksoniensis and S. orgila and the confusion in the literature between the former two species. A specimen (NMNZ
P.044152, 308.1 mm SL; Fig. 3b) collected from White
Island, New Zealand, is proposed as the neotype for the
species. Accordingly, White Island becomes the type
locality of S. cardinalis under Article 76.3 [ICZN (TheInternational Commission on Zoological Nomenclature)1999]. Incidentally, Paulin (1982), discussing the original
type locality of S. cardinalis, regarded it as Motuaro(Motuarohia) in the Bay of Islands, although Richardson(1842) quoted it as Motuara Island in Queen CharlotteSound from Solander’s manuscript. Scorpaena cardinalis
is not known to occur south of about 38S, and MotuaraIsland in Queen Charlotte Sound is at 41S.
Richardson (1842) quoted the name Scorpaena plebeia,
also from Solander’s unpublished manuscript, based on the323

fish from Tolaga [Bay], New Zealand. Scorpaena plebeia
was not described as a new species by Richardson (1842),
who wrote ‘‘he [= Solander] describes merely its colors,
and there is no drawing of it extant, nothing is known of its
form. He distinguishes Scorpaena plebeia, probably fromthe contrast which its duller general tints make with his
cardinalis’’, indicating that Richardson did not believe that
S. plebeia was a new species. The name S. plebeia is most
likely unavailable (ICZN 1999: article 11.6), and we regard
S. plebeia as a junior synonym of S. cardinalis even if S.
plebeia is an available name.
Scorpaena cookii was originally described by Günther(1874) on the basis of a single specimen (BMNH
1855.8.16.88, 216.2 mm SL; Fig. 2b) from Raoul Island(as its old name, Sunday Island), Kermadec Islands, New
Zealand. It has long been regarded by numerous authors(see synonym list) as a valid species, distributed along theAustralian east coast, the Tasman Sea’s offshore islands
and around northern New Zealand. However, comparisonof the holotype of S. cookii with the neotype and a large
number of non-type specimens of S. cardinalis showed
them to be identical. Scorpaena cookii is therefore regarded here as a junior synonym of S. cardinalis. The meristics and morphometrics of the neotype and non-type
specimens of S. cardinalis and the holotype of S. cookii are
given in Table 1.
In his review of New Zealand scorpionfishes, Paulin(1982) described S. cardinalis (= S. jacksoniensis in thisstudy) and S. cookii (= S. cardinalis) on the basis of 6
(NMNZ P.002190, P.005652, P.005779, P.005982,
P.007712) and 7 specimens (NMNZ P.007536, P.007537),
respectively. Re-examination of those specimens during
this study showed them to be S. cardinalis. Paulin’s (1982)
figures 3 (S. cardinalis) and 4 (S. cookii) were also clearlyS. cardinalis.
Common names of S. cardinalis are as follows: SandyBay Cod (standard English name; Poss 1999), Cook’s
Scorpionfish (standard Australian name; Yearsley et al.
2006), and Red Rockcod (New Zealand name; Roberts
et al. 2009).
Scorpaena jacksoniensis Steindachner 1866 (Standard
English name: Northern Scorpionfish; Australian name:
Eastern Red Scorpionfish) (Figs. 1c–d, 4, 7, 8, 9, 10, 11;
Tables 2, 3).
Scorpaena jacksoniensis Steindachner 1866a: 50 (PortJackson, NSW, Australia; abstract of Steindachner 1866b);
Steindachner 1866b: 438, pl. 3, figs. 2, 2a (Port Jackson,
NSW, Australia; detailed description).
Scorpaena cardinalis (not of Solander and Richardson):Günther 1860: 116 (Australian Seas); Macleay 1881: 431(Port Jackson, NSW); McCulloch 1912: 17 (NSW);

123

324

H. Motomura et al.

Fig. 7 Preserved specimens of
Scorpaena jacksoniensis.
a KAUM–I. 35882, 131.1 mm
SL; b AMS I.30310-025,
173.6 mm SL; c NMW 75379,
holotype of Scorpaena
jacksoniensis, 182.9 mm SL

McCulloch 1922: 116 (NSW); McCulloch 1930: 383 (NSW,
Qld, Tas); Kuiter 1993: 119 (southern Qld to eastern Vic);
Hutchins and Swainston 1986: 42, 126, fig. 176 (Noosa Head,
Qld to Jervis Bay, NSW); Allen and Cross 1989: 447 (Moreton
Bay, Qld to Mallacoota, Vic); Kuiter 1996: 86, 2 unnumberedfigs. (southern Qld and NSW); Johnson 1999: 727 (north toFraser Island, Qld); Poss 1999: 2304, 2334 (southern Qld toTas); Yearsley et al. 1999: 192, unnumbered fig. (Australia);
Motomura et al. 2005c: 866 (Qld to NSW); Allen et al. 2006:
844 (Fraser Island, Qld to Mallacoota, Vic); Motomura et al.
2011: 11 (NSW).
Scorpaena cruenta (not of Solander in Richardson): Ogilby
1893: 63, pl. 20 (NSW); Stead 1908: 108, pl. 75 (NSW).

123

Ruboralga jacksoniensis: Whitley 1931: 326 (describedas new genus); Whitley 1964: 56 (Australia).
Holotype. NMW 75379, 182.9 mm SL, Port Jackson,
NSW, Australia, 22 Jan. 1866.
Other material examined. 60 specimens, 25.1–336.5mm SL. AUSTRALIA: AMS IA.246, 153.9 mm SL, off
North Head, NSW, 33170 S, 151360 E, 37–73 m, Mcneill
and Livingston, 18 June 1921; AMS IB.6850, 78.2 mm SL,
Minnie Water, NSW, 29500 S, 153180 E, AMS party, 2
Nov. 1963; AMS IB.7346, 3, 112.5–169.8 mm SL, near
Quarantine Beach, North Head, Sydney, NSW, 33500 S,
151180 E, rotenone, AMS party and D. Francois, 11 Jan.

Synonymy of the Scorpaena cardinalis complex

325

Fig. 8 Fresh specimens andlive individual of Scorpaena
jacksoniensis. a BPBM 27828,
141.0 mm SL (photo: J.
Randall); b AMS I.43647-001,
281.9 mm SL (photo: H.
Motomura); c ca. 42 cm total
length, off Ulladulla, 12 m
depth, 26 Mar. 2006 (photo:
S. Schulz)

1965; AMS IB.7406, 208.9 mm SL, Nelson Bay, Port
Stephens, 32430 S, 152080 E, T. Castle, 1965; AMS I.9538,
225.4 mm SL, near Sydney, NSW, Fisheries Dept., 1908;
AMS I.9539, 274.0 mm SL, same data as AMS I.9538;
AMS I.9684, 161.3 mm SL, Port Jackson, NSW, 33050 S,
151130 E, W. Macleay, 1908; AMS I.16890-003, 2,
58.7–128.4 mm SL, Jervis Bay, 35030 S, 150440 E,
13–15 m, trawl, D. Pollard, 22 Sept. 1971; AMS I.19696-

007, 65.0 mm SL, South Solitary Island, off Coffs Harbour,
NSW, 30120 S, 153160 E, 5–20 m, spear, B. Russell and R.
Kuiter, 27 Sept. 1976; AMS I.19848-002, 163.4 mm SL,
Broughton Island, NSW, D. Hoese, 18 Apr. 1976; AMS
I.30310-025, 3, 107.8–173.6 mm SL, southwest side insmall bay, North Solitary Island, NSW, 29560 S, 153230 E,
12–14 m, rotenone and derris, AMS and NSW Fisheries, 5
May 1977; AMS I.31124-002, 2, 138.9–163.7 mm SL,

123

326

Fig. 9 Relationship of number of branched pectoral-fin rays tostandard length in Scorpaena cardinalis (closed stars) and S.
jacksoniensis (open circles)

Fig. 10 Relationships of: a orbit diameter; b first dorsal-fin spine
length; c head length; d snout length; e upper-jaw length; andf postorbital length (all as % of standard length) to standard length in123

H. Motomura et al.

North Solitary Island, NSW, 29560 S, 153240 E, 15 m,
rotenone, A. Hogget and L. Vail, 9 Feb. 1982; AMS
I.31125-001, 264.9 mm SL, south of Cronulla Beach, Qak
Park, NSW, 34270 S, 151090 E, 2 m, hook and line,
P. Phillis, 17 Dec. 1990; AMS I.31253-033, 2, 74.0–79.3 mm
SL, Washhouse Beach, Camden Head, NSW, 31380 3000 S,
152500 2400 E, 4 m, rotenone, AMS party, 8 May 1991;
AMS I.32218-003, 94.5 mm SL, off Newcastle, NSW,
3253–540 S, 15200–010 E, 73 m, prawn trawl, RV Kapala,20 June 1991; AMS I.40829-016, 129.8 mm SL, northeast
corner of Clarke Island, Sydney Harbour, NSW,
33510 4300 S, 151140 2600 E, 1.5–11.8 m, rotenone, AMS
party, 27 Apr. 2001; AMS I.40840-043, 96.9 mm SL,
Sydney Harbour, NSW, 33500 5000 S, 151160 1700 E, 1–8 m,rotenone, AMS party, 4 May 2001; AMS I.41285-027,
63.8 mm SL, rock pool adjacent to Blue Pool, Angourie,
NSW, 29280 4400 S, 153210 5000 E, 2 m, rotenone andScorpaena cardinalis (closed stars, solid lines) and S. jacksoniensis(open circles, dotted lines)

Synonymy of the Scorpaena cardinalis complex

Fig. 11 Relationship of number of gill rakers to standard length in
Scorpaena cardinalis (closed stars) and S. jacksoniensis (opencircles)

dipnets, A. Gill et al., 25 Mar. 2002; AMS I.43420-019, 2,
108.4–111.9 mm SL, 100 m off Fairlight Beach, Sydney
Harbour, NSW, 33480 0400 S, 151160 2900 E, 2–8 m, rotenone and spear, J. Cooper et al., 19 Aug. 2004; AMS
I.43647-001, 281.9 mm SL, Avalon to Maroubra Beach,
NSW, 33300 –34000 S, 15115–180 E, 5–25 m, spear, NSW
Spear Fishing Association, 7 Aug. 2005; AMS I.43647003, 285.4 mm SL, same data as AMS I.43647-001; AMS
I.43647-027, 211.6 mm SL, same data as AMS I.43647001; BMNH 1870.11.30.69, 221.1 mm SL, Coral Sea,
A. Thomson; BMNH 1890.9.23.142, 142.8 mm SL, Evans
Reef, NSW; BMNH 1890.9.23.143, 172.1 mm SL, samedata as BMNH 1890.9.23.142; BPBM 27828, 141.0 mm
SL, Sydney Harbour, NSW, 33510 S, 151160 E, rotenone,
R. Kuiter, 6 Sept. 1981; CAS 13712, 3, 149.3–208.3 mm
SL, Evans Head (purchased at Sydney Fish Market),
W. Eschmeyer, 4 Nov. 1969; CAS 60395, 87.7 mm SL,
Sydney Harbour, NSW, 0–1.8 m, B. Collette; CAS 120760, 2,
193.5–203.0 mm SL, Port Hacking, NSW, D. Stead;
CSIRO H 4313-11, 201.0 mm SL, Sydney Fish Market,
NSW, G. Yearsley, 6 Aug. 1996; KAUM–I. 35882,
131.1 mm SL, same data as AMS I.43420-019; KAUM–I.
35883, 111.7 mm SL, same data as AMS I.43420-019;
NMNZ P.014679, 48.0 mm SL, Clovelly Pool, Sydney,
NSW, 33550 S, 151440 E, J. Moreland and J. Yaldwyn, 30
Mar. 1967; QM I. 2417, 226.3 mm SL, Moreton Bay, Qld,
27150 S, 153150 E; QM I. 3612, 141.4 mm SL, Cowan
Cape, Moreton Bay, Qld, 27080 S, 153220 E, J. Palmer;
QM I. 3613, 135.5 mm SL, same data as QM I. 3612; QM
I. 12348, 131.3 mm SL, off Peel Island, Moreton Bay, Qld,
27290 S, 153210 E, W. Bindley, 20 Apr. 1951; QM I.
22141, 5, 25.1–58.6 mm SL, Cook Island, NSW, 28120 S,
153350 E, 12–15 m, J. Johnson, 2 Feb. 1986; QM I. 29253,
150.9 mm SL, Tangalooma Wrecks, Moreton Bay, Qld,

327

27010 S, 153220 E, 3–6 m, J. Johnson and J. Short, 6 June
1994; QM I. 29740, 152.4 mm SL, Amity Point, Qld,
27240 S, 153260 E, 2–9 m, J. Johnson, 14 Mar. 1995; QM
I. 34187, 114.9 mm SL, east of Stradbroke Island, Qld,
27350 S, 153370 E, 71 m, Qld Fisheries Service, 16 Oct.
2001; QM I. 36484, 62.0 mm SL, Manta Ray Bommie,
Point Lookout, north of Stradbroke Island, Qld, 27250 S,
153330 E, 8–9 m, J. Johnson, 23 Feb. 2005; QM I. 37688,
251.0 mm SL, Manta Ray Bommie, Point Lookout,north of Stradbroke Island, Qld, 27250 S, 153330 E,
7–11 m, J. Johnson and M. Ekins, 14 Dec. 2005; QM I.
37689, 175.6 mm SL, same data as QM I. 37688.
LOCALITY UNKNOWN: NMW 12176, 285.8 mm SL.
Diagnosis. A species of Scorpaena with the following
combination of characters: dorsal-fin soft rays 9; fourth(rarely third) dorsal-fin spine longest; pectoral-fin rays
16–18 (mode 17); scale rows in longitudinal series 52–61(55); pored lateral-line scales 22–24 (23); scales above
lateral line 4–6 (5), below 20–25 (22); scale rows between
sixth dorsal-fin spine base and lateral line 6–8 (7); scalerows between last dorsal-fin spine base and lateral line 7–9(9); pre-dorsal scale rows 1–4 (2); gill rakers on upper limb
4–6 (5), lower limb 10–13 (11) [7–9 (8) and 2–5 (3) rakers
on ceratohyal and hypobranchial, respectively], total rakers
14–19 (16); embedded cycloid scales covering pectoral-finbase (some scales exposed); embedded cycloid scales onanteroventral surface of body (covered by thick skin, notvisible without dissection); lateral surface of maxilla
without a longitudinal ridge; lateral lacrimal spine with 2(rarely 3) spinous points; anterior lacrimal spine with 1 or 2
small spinous points on its posterior margin; posterior
lacrimal spine simple, directed anteroventrally in juveniles
and young less than 100 mm SL, ventrally or posteroventrally in adults; median interorbital ridge present; interorbital and coronal spines absent; occipital pit andsupplement preopercular spine present; pterotic spine
usually simple in young, with 2 or more points in large
adults usually more than 280 mm SL; space between upperand lower opercular spines covered by thick skin withsensory pores and canals; numerous skin flaps and tentacles
on anterodorsal surface of body; a large black blotch distally on posterior spinous portion of dorsal fin in males; nodistinct white blotches on caudal-fin base; largest recorded
specimen, 336.5 mm SL.
Distribution. Scorpaena jacksoniensis is distributed
along the coastal water of eastern Australia (Fig. 4) fromsouthern Queensland to Victoria. The species is common
on rocky coastal reefs of New South Wales (Stewart andHughes 2010, reported as S. cardinalis) in depths of
1–73 m (based on collection data for specimens examinedin this study).
It is rare in Victoria; only two old specimens (NMV A
15898, dry, Victoria, collected in 1890 and NMV R 13360,

123

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H. Motomura et al.

Table 2 Meristics and morphometrics, expressed as percentages of standard length, of Scorpaena jacksoniensis
Holotype of S. jacksoniensis
NMW 75379

Non-types of S. jacksoniensis(n = 60)

182.9

25.1–336.5

Dorsal-fin rays

XII, 9XII, 9XII, 9Pectoral-fin raysa

i ? 6 ? xi = 18/18

i–ii ? 3–7 ? ix–xiii= 16–18/16–18

i ? 5 ? xi = 17/17

Standard length (SL, mm)
Meristics

Modes

Scale rows in longitudinal series

57

52–61

55

Pored lateral-line scales

23

22–24

23

Scales above lateral line

6

4–6

5

Scales below lateral line

22

20–25

22

Scale rows between 6th dorsal-fin spine base andlateral line

7

6–8

7

Scale rows between last dorsal-fin spine base andlateral line
Pre-dorsal scale rows

9

7–9

9

4

1–4

2

Gill rakersb

6 ? 13 = 19

4–6 ? 10–13 = 14–19

Morphometrics (%SL)
Body depth

5 ? 11 = 16
Means

35.6

34.0–38.5

35.6

Body width

23.8

21.2–29.0

24.2

Head length

43.3

40.7–46.7

43.9

Snout length

11.9

11.9–14.6

13.0

Orbit diameter

10.1

8.6–13.3

10.6

Interorbital widthc

6.2

5.2–7.1

6.4

d

5.7

Interorbital width

6.2

5.1–6.4

Head width–

13.3–16.0

14.4

Upper-jaw length

20.7

19.5–23.9

21.7

Maxillary depth

5.4

5.3–7.0

6.2

Suborbital space

2.7

1.6–2.8

2.2

Postorbital length
Between tips of opercular spines

23.0
6.8

20.9–24.9
5.6–8.0

22.7
7.1

Occipital pit length–

4.9–7.0

5.8

Occipital pit width–

5.5–7.0

6.1

Post-occipital pit length–

6.8–9.6

8.4

Predorsal-fin length

36.2

34.9–40.8

37.1

Preanal-fin length

72.4

67.6–75.6

71.2

Prepelvic-fin length

36.2

36.2–43.7

39.2

1st dorsal-fin spine length

8.1

6.1–10.7

8.7

2nd dorsal-fin spine length

Damaged8.2–17.7

13.7

3rd dorsal-fin spine length

17.4

11.6–21.0

17.4

4th dorsal-fin spine length

18.8

13.1–21.7

18.0

5th dorsal-fin spine length

18.3

12.5–20.4

16.4

11th dorsal-fin spine length

5.5

3.3–8.7

5.7

12th dorsal-fin spine length

12.6

7.8–15.3

11.8

Longest dorsal-fin soft ray length

18.2

15.9–21.3

18.1

1st anal-fin spine length
2nd anal-fin spine length

8.2
16.3

5.9–12.5
12.2–22.4

8.9
17.5

3rd anal-fin spine length

14.8

11.1–19.3

15.2

Longest anal-fin soft ray length

21.0

19.2–24.8

21.9

Pectoral-fin ray length

26.5

24.7–33.2

28.8

123

Synonymy of the Scorpaena cardinalis complex

329

Table 2 continuedaHolotype of S. jacksoniensis
NMW 75379

Non-types of S. jacksoniensis(n = 60)

Pelvic-fin spine length

13.9

11.1–19.1

14.6

Longest pelvic-fin soft ray length

22.4

22.3–28.5

24.9

Caudal-fin length

27.1

23.6–33.3

28.2

Caudal-peduncle length

18.2

15.7–20.9

18.0

Caudal-peduncle depth

11.3

10.3–17.3

11.6

Upper unbranched ray ? branched ray ? lower unbranched rays = total rays on left side of body/total rays on right sideb

Rakers on upper limb ? rakers on lower limb = total rakerscAt vertical midline of eyedAt posterior end of preocular spine base–, data not taken

Table 3 Frequency distribution of selected meristics of Scorpaena cardinalis and S. jacksoniensis
Pectoral-fin rays

S. cardinalis
S. jacksoniensis

16/16

16/17

17/17

17/18

18/18

2

4

35

8

17N

1

H

4

5

47

Pored lateral-line scales

Scales above lateral line

22

4

23
9

3

2

24

25

56N

1

H

3

52

5

6
12

2

7

8

9

23

25N

4

H

36

17

Scale rows in longitudinal series
52

53

54

55

56

57

14

H

58

59

S. cardinalis
S. jacksoniensis

1

4

7

N

7

3

3

61

62

63

64

65

66

67

68

69

70

71

72

73

74

2

2

4N

5

3

8

8

8

13

3

3

1

1

2

1

1

1

5

6

7

8

11

37

13

3N

Scale rows between 6th DS and LL

Scale rows between last DS and LL

Pre-dorsal scale rows

6

7

1

7

S. cardinalis
S. jacksoniensis

17

60

8

H

44

8

9

6N

17

3

2

8

9

1

5

24

34

10

11

58N

2

H

2

3

4
1

1

25

21

7

H

and H include neotype of S. cardinalis and holotype of S. jacksoniensis, respectively

DS dorsal-fin base, LL lateral line

293 mm SL, Western Port, Victoria, 1933) from Victorianwater have been deposited at the Museum Victoria (D.
Bray, personal communication). Accurate localities of thetwo specimens are unknown because the latter specimen
was probably purchased from a fish market in Western
Port, Victoria. The southernmost record of S. jacksoniensis
based on voucher specimens is Jervis Bay, New South
Wales (35030 S).
A specimen (QM I. 16624, otoliths only, off Mooloolaba, Queensland, 26410 S) represents the northernmost
record of the species on the basis of a voucher specimen (J.
Johnson, personal communication). However, S. jacksoniensis is regularly caught off Fraser Island, Queensland(25130 S) (Johnson 1999), and occasionally reported as
taken on the southern end of the Great Barrier Reef (J.

Johnson, personal communication), although no museum
specimens have been retained from there.
Biological notes. Biological information on S. jacksoniensis (as S. cardinalis) was reported by Stewart andHughes (2010), who estimated the oldest individual to be
33 years old. They also found that the spawning peak in
New South Wales waters was March, and the species had ahighly specialized ovary adapted for the production of afloating gelatinous egg mass.
Remarks. Scorpaena jacksoniensis was originallydescribed by Steindachner (1866a) on the basis of a single
large specimen (NMW 75379, 182.9 mm SL; Fig. 7c) fromPort Jackson, New South Wales, Australia. Since its original description, numerous authors (see synonym list)
have regarded S. jacksoniensis as a junior synonym of S.

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330

cardinalis. However, the nominal name, S. cardinalis,applies to a species that has long been regarded as S. cookii(see ‘Remarks’ of S. cardinalis), and S. cardinalis does notoccur around the Australian mainland (see ‘Distribution’ of
S. cardinalis). Examination of the holotype of S. jacksoniensis found it to be identical with species nominallyidentified as Scorpaena ‘cardinalis’ from the coastal waterof the Australian mainland. Accordingly, S. jacksoniensis
is herein restored as a valid species. The meristics andmorphometrics of the holotype and non-type specimens of
S. jacksoniensis are given in Table 2.
Scorpaena cruenta reported by Ogilby (1893: pl. 20)
and Stead (1908: pl. 75) from NSW is re-identified here as
S. jacksoniensis. Scorpaena cruenta is currently known as a
junior synonym of S. papillosa (Paulin 1982; Motomura,
unpublished data).
Whitley (1931) proposed a new genus, Ruboralga, for S.
jacksoniensis and included three other Australian species,
S. ergastulora (as ergastulorum), S. sumptuosa and S.
bellicosa, and a New Zealand species, S. cardinalis. Subsequently, Whitley (1968) added S. cookii to Ruboralga.
Whitley’s (1931, 1968) R. cardinalis is most likely toinclude S. papillosa and S. cardinalis. Although Whitley(1931) stated that ‘‘Scorpaena from the Mediterranean Sea
seems to have a very different facies… [from] the Australian form’’ and described general characters of scorpionfishes, e.g., ‘‘orbital crests and bony stay of cheek withseveral spines’’, ‘‘second anal spine longer than third’’, and‘‘general coloration mainly reddish’’, he did not give
diagnostic characters to separate Ruboralga from Scorpanea. In addition, S. bellicosa that Whitley (1931) includedin Ruboralga was recently described by Motomura et al.
(2009) as a junior synonym of Parascorpaena aurita(Rüppell 1838). Moreover, we found no generic charactersto justify distinguishing Ruboralga from Scorpaena,
although all valid species of Scorpanea were examined(Motomura, unpublished data). Therefore, Ruboralga isregarded as a junior synonym of Scorpaena.
Common names of S. jacksoniensis are as follows:
Northern Scorpionfish (standard English name; Poss 1999)
and Eastern Red Scorpionfish (standard Australian name;Yearsley et al. 2006).
Morphological changes with growth. In common withmost members of the genus Scorpaena (see Motomura
et al. 2005b), the number of the branched pectoral-fin raysin S. cardinalis and S. jacksoniensis tend to increase withgrowth until they attain ca. 200 mm SL (Fig. 9).
The scales enclosed by the posterior tips of the upperand lower opercular spines and opercular margin alsochange with growth from cycloid to ctenoid in Scorpaena
bulacephala and S. onaria (see Motomura et al. 2005b, c,
respectively). However, those of S. cardinalis and S.
jacksoniensis remain cycloid throughout life. Furthermore,

123

H. Motomura et al.

whereas the occipital pit of S. bulacephala becomes deeper
with growth, those of S. cardinalis and S. jacksoniensis
become shallower.
The distal margins of the pterotic spines of young S.
jacksoniensis are smooth (a typical condition in Scorpaena), and those of large adults (usually more than 280 mm
SL) have two or more points, whereas those of S. cardinalis remain smooth throughout life.
Juveniles and young S. jacksoniensis less than 100 mm
SL have an anteroventrally directed posterior lacrimal
spine (Fig. 1c), the direction of the spine changes to ventrally or posteroventrally directed with growth (Fig. 1d).
The direction of the posterior lacrimal spine of S. cardinalis does not change throughout life (Fig. 1a, b).
Changes in relative lengths of some body parts withgrowth, especially fin spines and soft rays, are well knownin Scorpaenidae. However, except for studies of S. bulacephala and S. onaria by Motomura et al. (2005b: fig. 4;
2005c: fig. 7 respectively), no detailed descriptions of suchchanges, including accurate measurement data, have beenreported for other species of Scorpaena. Analyses of 37
measurements in S. cardinalis and S. jacksoniensis foundthat relative lengths of the orbit, head, snout, upper jaw,
postorbit and all fin rays of the two species changed significantly with growth. The orbit diameter and lengths ofall fin rays as a percentage of SL (only first dorsal-fin spine
length is given in Fig. 10 as an example) decrease withgrowth (Fig. 10a, b) as found in other species of Scorpaena. However, the head, snout, upper-jaw and postorbital
lengths become significantly longer as a percentage of SL
with growth (Fig. 10c–f). These changes have not beenpreviously reported in Scorpaenidae. As a result of thesechanges, the heads of the two species appear to become
more elongate with growth (see Figs. 2, 7).
There is considerable variation in the number of gillrakers in S. cardinalis and S. jacksoniensis, i.e., 14–19
rakers in both species (Tables 1, 2). Although the numberof gill rakers in several fish species generally decreases
with growth due to changes in diet, those of the two speciesof Scorpaena do not represent the growth-related change(Fig. 11). Moreover, the differences in number of gillrakers are not reflected by capture localities. The wide
range in the number of gill rakers in the two species istherefore regarded as an individual variation of eachspecies.
Comparisons. Ontogenetic differences (posterior lacrimal and pterotic spines) between S. cardinalis and S.
jacksoniensis are noted above. Scorpaena cardinalis isreadily distinguished from S. jacksoniensis by having
exposed cycloid scales covering on the anteroventral surface of the body (some scales covered by thin skin, especially in large adults) (vs. scales embedded in thick skin,not visible without dissection in S. jacksoniensis), and thinSynonymy of the Scorpaena cardinalis complex

skin without sensory pores or canals covering on a space
between the upper and lower opercular spines (vs. thickskin with sensory pores and canals covering the area).
Scorpaena cardinalis has smaller scales than S. jacksoniensis, having higher counts of pored lateral-line scales[23–25 (mode 24) in S. cardinalis vs. 22–24 (23) in S.
jacksoniensis], scales above the lateral line [6–9 (8) vs. 4–6(5)], scale rows in the longitudinal series [60–74 (68) vs.
52–61 (55)], scale rows between the sixth dorsal-fin spinebase and lateral line [8 or 9 (9) vs. 6–8 (7)], and scale rows
between the last dorsal-fin spine base and lateral line [8–11(10) vs. 7–9 (9)] (Table 3). The space between the dorsalfin origin and occipital pit of S. jacksoniensis is broadly
covered by thick skin, and only 1–4 scale rows (pre-dorsal
scale rows) are exposed, whereas that of S. cardinalis lacks
the thick skin and 4–8 scale rows are exposed (Table 3).
In addition, S. cardinalis differs from S. jacksoniensis in
having the posterior lacrimal spine with 1–3 (usually 2)
spinous points (vs. simple in S. jacksoniensis; Fig. 1), andthe anterodorsal surface of the body with a lower numberof skin flaps or tentacles than S. jacksoniensis.
The two species can also be distinguished by coloration;
S. cardinalis has two large white blotches, each diameter
subequal to pupil diameter, on the upper and lower caudal
peduncle at the caudal-fin base (Figs. 3, 5, 6), whereas S.
jacksoniensis lacks the distinct blotches. Males of S.
jacksoniensis have a large, elongate, black blotch distallyon the posterior spinous portion of the dorsal fin (Fig. 8b),whereas S. cardinalis lacks the blotch in both sexes.
Scorpaena cardinalis grows larger than S. jacksoniensis(largest recorded size, 472.5 vs. 336.5 mm SL).
Scorpaena orgila, the third species of the S. cardinalis
complex, is similar to S. cardinalis in overall body
appearance, having exposed cycloid scales covering theanteroventral surface of the body (some scales covered bythin skin), thin skin without sensory pores or canals covering a space between the upper and lower opercularspines, 24 pored lateral-line scales and 8–10 scales above
the lateral line. However, S. orgila differs from S. cardinalis in having a lower number of scale rows in the longitudinal series [50–59, based on data for specimens
examined in this study and given by Eschmeyer and Allen(1971) vs. 60–74 (mode 68); Tables 1, 3]. Because S.
orgila is restricted to Easter Island and known only fromtype specimens, more specimens are required to clarify
these taxonomic characters.
Acknowledgments We are especially grateful to A. Hay and S.
Reader (AMS), J. Maclaine (BMNH), W. Eschmeyer, T. Iwamoto, D.
Catania and M. Hoang (CAS), P. Last, A. Graham and W. White(CSIRO), H. Wellendorf and C. Prenner (NMW) and J. Johnson (QM)
for their kind assistance during the first author’s visits to their institutions. We thank J. Randall, L. O’Hara and A. Suzumoto (BPBM)
for the specimen loan, and D. Bray (NMV) and J. Johnson (QM) for

331
providing distributional information on S. jacksoniensis in Victoria
and Queensland, respectively. We are grateful to L. Moore and all
staff of the Fish Section of NMNZ for their generous support during
this study, and S. Schulz (Sydney, Australia), J. Randall (BPBM) and
R. McPhee (NMNZ) for providing some photographs of S. jacksoniensis and S. cardinalis. This work was also supported (in part) by
the NZ Foundation for Research Science and Technology through the
Te Papa Biosystematics of NZ EEZ Fishes subcontract within
NIWA’s marine Biodiversity and Biosecurity OBI programme (contract C01X0502) and a Grant-in-Aid for Young Scientists (B) from
the Ministry of Education, Science, Sports and Culture, Tokyo(19770067).

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