Revision of the Indo-West Pacific genus Brachypterois (Scorpaenidae: Pteroinae), with description of a new species from northeastern Australia

Abstract

A taxonomic revision of the scorpaenid genus Brachypterois resulted in three valid species. Brachypterois serrulifer Fowler 1938, previously regarded as ajunior synonym of Sebastes serrulatus Richardson 1846, can be distinguished from its congeners by having usually 16 pectoral-fin rays (vs. 15 in the congeners) and 0-22 spines on the median lateral ridge of the maxilla (vs. almost always absent), in addition to longer dorsal- and anal-fin soft rays, relatively fewer large dark spots on the caudal fin, and the posterior lacrimal spines usually directed ventrally. Brachypterois curvispina sp. nov., described on the basis of 32 specimens, can be distinguished from B. serrulata (Richardson 1846) by having fewer number of scale rows in the longitudinal series 41-45 (vs. 44-51 in B. serrulata); relatively long head, its length 41.6-44.5 (mean 43.1)% SL [vs. 38.3-43.1 (mean 40.2)% SL]; posterior lacrimal spines usually directed posteriorly, with the tip curved strongly upward in large males (vs. simply directed posteroventrally); and posteriorly directed spine(s) on the posterior corner of the outer angular ridge (vs. directed ventrally). Brachypterois serrulifer is distributed widely in the Indo-West Pacific, whereas distributional ranges of B. curvispina sp. nov. and B. serrulata are restricted to northeastern Australia and the northwestern Pacific, respectively. Intraspecific variation, including sexual dimorphism and ontogenetic changes in the three species are described in detail. A neotype, collected from southern Taiwan, is designated for S. serrulatus Richardson 1846.

Full text

Accepted by M.R. de Carvalho: 2 Jul. 2013; published: 31 Jul. 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
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1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3693 (4): 401–440
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http://dx.doi.org/10.11646/zootaxa.3693.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:3DF2C9DF-23A1-4659-B118-A2697ADEBFB0
Revision of the Indo-West Pacific genus Brachypterois (Scorpaenidae: Pteroinae),
with description of a new species from northeastern Australia
MIZUKI MATSUNUMA
1, 4
, MAKOTO SAKURAI
2
& HIROYUKI MOTOMURA
3
1
The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-30 Korimoto, Kagoshima 890-0065, Japan.
E-mail: k1139853@kadai.jp
2
The Kagoshima Immaculate Heart College, 4-22-1 Toso, Kagoshima 890-8525, Japan. E-mail: sakurai@juntan.k-junshin.ac.jp
3
The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan. E-mail: motomura@kaum.kagoshima-u.ac.jp
4
Corresponding author
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
Genus Brachypterois Fowler 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404
Key to the species of Brachypterois . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406
Brachypterois curvispina sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407
Brachypterois serrulata (Richardson 1846) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416
Brachypterois serrulifer Fowler 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 426
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Abstract
A taxonomic revision of the scorpaenid genus Brachypterois resulted in three valid species. Brachypterois serrulifer
Fowler 1938, previously regarded as a junior synonym of Sebastes serrulatus Richardson 1846, can be distinguished from
its congeners by having usually 16 pectoral-fin rays (vs. 15 in the congeners) and 0–22 spines on the median lateral ridge
of the maxilla (vs. almost always absent), in addition to longer dorsal- and anal-fin soft rays, relatively fewer large dark
spots on the caudal fin, and the posterior lacrimal spines usually directed ventrally. Brachypterois curvispina sp. nov., de-
scribed on the basis of 32 specimens, can be distinguished from B. serrulata (Richardson 1846) by having fewer number
of scale rows in the longitudinal series 41–45 (vs. 44–51 in B. serrulata); relatively long head, its length 41.6–44.5 (mean
43.1)% SL [vs. 38.3–43.1 (mean 40.2)% SL]; posterior lacrimal spines usually directed posteriorly, with the tip curved
strongly upward in large males (vs. simply directed posteroventrally); and posteriorly directed spine(s) on the posterior
corner of the outer angular ridge (vs. directed ventrally). Brachypterois serrulifer is distributed widely in the Indo-West
Pacific, whereas distributional ranges of B. curvispina sp. nov. and B. serrulata are restricted to northeastern Australia
and the northwestern Pacific, respectively. Intraspecific variation, including sexual dimorphism and ontogenetic changes
in the three species are described in detail. A neotype, collected from southern Taiwan, is designated for S. serrulatus Rich-
ardson 1846.
Key words: Brachypterois curvispina, Brachypterois serrulata, Brachypterois serrulifer, taxonomy, redescription, valid-
ity, sexual dimorphism

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402 · Zootaxa 3693 (4) © 2013 Magnolia Press
Introduction
The scorpaenid genus Brachypterois Fowler 1938, previously regarded as monotypic, represented only by
Brachypterois serrulata (Richardson 1846), is widespread in the Indo-West Pacific ranging from the Red Sea and
Persian Gulf east to northern Australia, and north to southern Japan (Kanayama & Amaoka 1981; Poss 1999). The
genus is characterized by having 3 anal-fin spines, and branched rays on pectoral fin, maxilla covered with scales
and spinous ridges on mandible (Fowler 1938; Kanayama & Amaoka 1981; Poss 1999).
Richardson (1846) described Sebastes serrulatus based on a single specimen found in an insect box sent from
China and the type locality was thus entered as Sea of China. Subsequently, Günther (1860), in his catalogue of fish
type specimens in the British Museum (= Natural History Museum, London; NHM), listed S. serrulatus, but
provided neither a registration number nor information on the holotype. A search of the type specimen in the NHM
has failed to locate it, and it is apparently lost (J. Maclaine, 2012, pers. comm.). According to the original
description, S. serrulatus is best assigned to Brachypterois as in previous studies (Chu 1962; Kanayama & Amaoka
1981).
Fowler (1938) described Brachypterois serrulifer as a new genus and species based on a single specimen
collected from the Philippines. However, recent researchers have regarded B. serrulifer as a junior synonym of S.
serrulatus (Chu 1962; Kanayama & Amaoka, 1981; Poss 1999).
Recent examinations of a large number of Brachypterois specimens from throughout the Indo-West Pacific has
resulted in three valid species, viz., B. serrulata, B. serrulifer and B. curvispina sp. nov. Brachypterois serrulifer is
distinguishable from its congeners on the basis of several morphological aspects, including counts of pectoral-fin
rays and gill rakers, proportional measurements, head spine shape and coloration. Brachypterois curvispina sp.
nov. is described herein on the basis of 32 specimens collected from northeastern Australia. A neotype of S.
serrulatus is designated. Morphological variation in Brachypterois, including sexual dimorphism and ontogenetic
changes, are also described in detail.
Materials and methods
Measurements generally followed Motomura (2004b, c), except for head width that followed Motomura et al.
(2005b, 2006a), maxillary depth that followed Motomura et al. (2006b), and body depth at anal-fin origin
measured from the vertical distance between dorsal- and anal-fin bases at level of first anal-fin spine. Counts
generally followed Motomura et al. (2005a, b, c) and Motomura & Johnson (2006), predorsal scale count followed
Motomura et al. (2006b), plus the following additions: scales above lateral line, vertical scale rows between first
pored lateral-line scale and dorsal-fin base; suborbital scale rows, scale rows between ventral margin of orbit and
suborbital ridge at mid-orbit level. The last two soft rays of the dorsal- and anal-fins were counted as a single ray,
each pair is supported by a single pterygiophore. The formula for configuration of the supraneural bones, anterior
neural spines and anterior dorsal fin pterygiophores follows Ahlstrom et al. (1976). Spine counts on the median
lateral ridge of the maxilla do not include those on the lateral portion of the articular head of the maxilla (Fig. 1).
Counts and measurements are made on the left side wherever possible, except for pectoral-fin rays which are
counted on both sides when possible.
Terminology of head spines generally follows Randall & Eschmeyer (2001: fig. 1) and Motomura (2004c: fig.
1); the spine on the lateral surface of the lacrimal bone is termed the lateral lacrimal spine (Motomura & Senou
2008: fig. 2). The coronal spine is figured in Chen (1981: fig. 1). Spines and ridges on the mandible (Fig. 2) are
defined as follows: spines around mandibular symphysis (SMS), spines on the outer edge of the dentary anterior to
the point of lower lip attachment, surrounding the 1st sensory pore of the dentary; spines on outer dentary ridge
(SODR),a row of spines on the outermost ridge or edge of the dentary; spines on inner dentary ridge (SIDR), a row
of spines on the innermost ridge of the dentary; spines on central dentary ridge (SCDR), row(s) of spines on
ridge(s) occurring along one or both sides of the sensory canal, running along the mid portion of the dentary; spines
on outer angular ridge (SOAR), a row of spines on the outermost ridge or edge of the angular; spines on inner
angular ridge (SIAR), a row of spines on a ridge following the sensory canal, running along the mid portion of the
angular.

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REVISION OF BRACHYPTEROIS
FIGURE 1. Lateral (top) and dorsal (bottom) views of maxilla of Brachypterois serrulifer, KAUM–I. 22033, 82.1 mm SL. 1
Spines on anterior portion of articular head; 2 Spines on lateral portion of articular head; 3 Spines on median lateral ridge; 4
Dorsal ridge. Left to anterior. Bar indicates 3 mm.
The mitochondrial DNA (mtDNA) cytochrome b gene is examined for three specimens of each B. serrulata
(KAUM–I. 12313, KAUM–I. 22034, Malaysia; KAUM–I. 33225, Thailand) and B. serrulifer (KAUM–I. 22033,
KAUM–I. 22035, Malaysia; KAUM–I. 39187, Taiwan). DNA extraction was performed with the Gentra Puregene
Tissue Kit (QIAGEN) following the manufacturer’s protocols. Using total DNA and the primer pair AJG15
(Akihito et al. 2000) and Simt12-Fb (Kuriiwa et al. 2007), approximately 2.3 kbp fragments, including the
cytochrome b gene and control region of mtDNA, were amplified under the following PCR conditions. Thirty
amplification cycles were performed at 94ºC for 45 sec, 56ºC for 30 sec and 72ºC for 2 min. Sequencing reactions
were carried out using the BigDye Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems) with the primer
L15369-CYB (Miya & Nishida 2000), following the manufacturer’s protocols and utilizing the automated genetic
analyzer models 3130 and 3500 (Applied Biosystems). The obtained DNA sequences were subjected to multiple
alignment using Clustal X (Thompson et al. 1997). All sequences are available on DDBJ/EMBL/GenBank
accession numbers AB793293–AB793298.The sequence difference (p-distance) was calculated using MEGA 5
(Tamura et al. 2011).

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404 · Zootaxa 3693 (4) © 2013 Magnolia Press
FIGURE 2. Diagram of mandible of Brachypterois serrulata, showing series of spines and ridges. SMS spines around
mandibular symphysis; SODR spines on outer dentary ridge; SIDR spines on inner dentary ridge; SCDR spines on central
dentary ridge; SOAR spines on outer angular ridge; SIAR spines on inner angular ridge; SP sensory pore. Left to anterior. Arrow
shows position of lower lip attachment to dentary.
Sex was determined by examination of gonad sections embedded in paraffin and stained with Mayer’s
haematoxylin-eosin or direct examination of the gonads by light microscope, and is indicated where applicable in
lists of specimens examined. Specimens with meristic data taken only are indicated by asterisk in front of the
registration number. Radiographs were taken from 7 specimens of B. curvispina, 17 specimens of B. serrulata and
13 specimens of B. serrulifer, indicated by “(X)” in lists of specimens examined. Standard length is abbreviated as
SL. The holotype of Brachypterois curvispina sp. nov. is deposited at the Australian National Fish Collection,
Commonwealth Scientific and Industrial Research Organisation, Division of Marine and Atmospheric Research,
Hobart. Institutional codes follow Sabaj Pérez (2012), with the following additions: OMNH, Osaka Museum of
Natural History, Osaka; NSMT, National Museum of Nature and Science, Tsukuba; SNFR, Seikai National
Fisheries Research Institute, Nagasaki; URM, Churashima Research Center, Okinawa Churashima Foundation,
Motobu (formerly fish collection at University of the Ryukyus). Curatorial procedures for newly collected
specimens followed Motomura & Ishikawa (2013).
Genus Brachypterois Fowler 1938
Brachypterois Fowler 1938: 79, fig. 35 (type species: Brachypterois serrulifer Fowler 1938; type by original designation and
monotypy)
Ranipterois Whitley 1950: 407 (type species: Brachypterois serrulifer Fowler 1938; unnecessary replacement name for
Brachypterois Fowler 1938; objective synonym of Brachypterois)
Type species. Brachypterois serrulifer Fowler 1938
Definition. A genus of the family Scorpaenidae, subfamily Pteroinae, defined by following combination of
characters: dorsal fin usually with 13 (rarely 12 or 14) spines; anal fin with 3 spines; pectoral fin with 5–8 branched
rays; caudal fin usually with 3 unsegmented procurrent rays in upper and lower series; maxilla covered by scales;
mandible with distinct ridges; occiput flat with scales; dorsal, anal and pelvic fin spines with developed notches
with poison glands; palatine teeth absent; orbit and suborbital ridge distinctly closed; 5 preopercular spines.
Description. Characters given in the generic definition are not repeated here, except for counts of fin rays.
Dorsal fin with 13 spines and 10 soft rays (rarely XII, 11; XIII, 9 or 11; XIV, 9 or 10). Anal fin with 3 spines and 5
soft rays (rarely III, 4 or 6). Pelvic fin with 1 spine and 5 soft rays. Gill rakers simple, short and robust, longest
raker on first gill arch shorter or subequal to gill filaments around angle of gill arch; gill rakers on lower limb
around angle of first gill arch more or less branched into two; fourth gill slit narrow (partially closed by
membrane). Branchiostegal rays 7. Vertebrae 10 + 14 (rarely 9 + 14, 10 + 15 or 11 + 13). Formula for configuration
of supraneural bones, anterior neural spines and anterior dorsal pterygiophores usually //2+1/1/1/1/1/1/1/1/1/1+1/.
Swimbladder present.

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REVISION OF BRACHYPTEROIS
Body oblong, moderately compressed anteriorly, strongly compressed posteriorly. Head large, its length
greater than body depth. Low membranous tube associated with anterior and posterior nostrils; posterior edge of
tube on anterior nostril slightly expanded. Numerous small papillae sometimes present on frontal surface of bulge
of snout and chin. Ctenoid scales covering on head, including cheek, lacrimal, suborbital pit, interopercle,
subopercle, opercle, posterodorsal pit of supraocular, channel between interorbital ridges, occiput and
suprapostorbital area surrounded by parietal, pterotic, lower posttemporal and cleithral spine bases; scales absent
on both lips and bulge of snout. Maxilla with weakly ctenoid scales; posterior portion of maxilla almost entirely
scaled or partly scaled as one-third maxilla depth. Mandible usually with exposed weakly ctenoid scales;
sometimes with embedded cycloid scales or lacking scales in large adults. Ctenoid scales covering on body and
extending onto base of caudal fin; scales on ventral body surface relatively weak; pectoral-fin base covered with
weakly ctenoid scales; a row of elongated cycloid scales on bases of soft-rayed portions of dorsal and anal fin.
Lateral line complete, weakly downward sloping posteriorly. Three sensory pores on underside of each dentary; a
small pore on each side of symphysial knob.
Mouth moderately large, oblique, forming an angle of ca. 40–50° to horizontal axis of head and body; upper
edge of maxilla swollen laterally forming a low ridge; usually with a median lateral ridge; posterior end of maxilla
just reaching a vertical through mid-orbit. Width of symphysial gap separating premaxillary teeth bands about
twice that of each band; upper jaw with a band of small slender conical teeth; about 4–6 tooth rows at front of upper
jaw; about 4 tooth rows at front of lower jaw. Vomer with a V-shaped patch of 1–5 rows of small teeth.
Dorsal profile of snout steep, forming an angle of ca. 40° to horizontal axis of head and body. Nasal spine on
relatively long ridge-like base, its length subequal to distance between anterior and posterior nostrils; with 1–2
rows of 1–15 small spines. Interorbital ridges moderately developed, with a row of 0–19 spines; separated by a
shallow channel, initially divergent posteriorly; sometimes conjoined with origin of coronal spine base. Interorbital
space shallow, about one-seventh of orbit diameter extending above dorsal profile of head. Coronal spine with 1–5
small spinous points on short base; base conjoined with origin of parietal spine base; 0–6 small spines forming a
pair of small patches between origins of coronal spine base. Tympanic with 2 spinous ridges; upper ridge shorter
than lower ridge with 2–8 spines; lower ridge with 2–20 spines. Preocular with 3–28 spines on its frontal surface
and margin of orbit. Supraocular with a transverse ridge located between interorbital ridge and outer edge of
supraocular, with 3–15 spines; 9–35 spines on margin of orbit; posterodorsal edge of supraocular expanded
posteriorly, with a small inner pit. Postocular with 3–16 spines on margin of orbit. Occipital area lacking anterior
ridge; a pair of short low ridges with 0–9 spines on posterior margin between parietal spine bases. Parietal spine
with 7–23 spinous points on a relatively long ridge diverging posteriorly; a short ridge with 0–7 spines just under
posterior portion of parietal ridge. Sphenotic spine with 5–32 spinous points. No postorbital spine. Pterotic spine
with 5–27 spinous points. Small patch of 0–4 spines behind pterotic spine. One or two short oblique ridges with 0–
13 spines between parietal spine and lower posttemporal spine. No upper posttemporal spine. Lower posttemporal
spine with 1–13 spinous points on a relatively short ridge; a short oblique ridge with 0–5 spines just above posterior
portion of spine base; one or two small patches of 0–7 spines in front of spine. Supracleithral spine with 0–6
spinous points on a relatively short ridge. Cleithral spine with 2–14 spinous points on a relatively long ridge.
Anterior portion of articular head of maxilla with 0–8 spines; lateral portion of articular head exposed when
mouth closed, with 0–5 spines. Inner edge of dentary elevated as a low ridge; central dentary ridge running along
sensory canal, comprising 2 ridges posteriorly in large adults; outer margin of dentary a low ridge; tip of ventral
surface of dentary slightly expanded ventrally. Outer edge of angular a low ridge; inner angular ridge running along
sensory canal. Retroarticular with 0–3 spines. Lacrimal with 2 lateral ridges (single in juveniles less than about 40
mm SL); upper ridge with 4–7 spines, its anterior origin conjoined with ridge on dorsal articular process of
lacrimal; lower ridge with 1–6 spines. Anterior lacrimal spine with 1–6 spinous points, directed downward.
Posterior lacrimal spine with 2–8 spinous points; all ridges on lacrimal strongly spinous. Suborbital with 2 ridges;
both ridges with supplementary upper and lower ridges; posterior upper ridge branched into two posteriorly; 2
small arched ridge below anterior and posterior ridges, sometimes covered with tiny skin with numerous small
pores; all ridges strongly spinous (21–89 spines in total). Poorly developed suborbital pit present. Preopercle spines
with one or more spinous points; lateral ridge of uppermost spine with 0–5 spines. A ridge of preopercle submargin
strongly spinous. Interopercle with 0–10 spines on distal margin. Upper opercular spine with 1–4 spinous points,
somewhat covered with skin; a simple lower opercular spine sometimes present, but usually under skin.

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406 · Zootaxa 3693 (4) © 2013 Magnolia Press
Origin of first dorsal-fin spine above cleithral spine; bases of first and second dorsal-fin spines closer than
those of succeeding adjacent spines; length of longest spine about half maximum body depth; penultimate (usually
twelfth) spine usually shortest; membranes of spinous portion of dorsal fin strongly incised, especially in anterior
portion. Dorsal-fin soft rays all branched; length of longest ray subequal to or slightly longer than that of longest
fin spine; posteriormost ray not joined by membrane to caudal peduncle. Origin of first anal-fin spine below base
of penultimate (usually twelfth) dorsal-fin spine; third spine longest. Anal-fin soft rays all branched; second (rarely
first or third) soft ray longest, its length longer than that of longest dorsal-fin soft ray; posteriormost ray not joined
by membrane to caudal peduncle. Pectoral fin long; usually fifth (branched ray) and tenth (unbranched ray) rays
longest in relatively small and large specimens, respectively, but highly variable; tip of longest ray extending well
beyond a vertical through base of first anal-fin spine, sometimes reaching a vertical through base of last anal-fin
soft ray, but not reaching caudal-fin base; lower unbranched rays weakly thickened. Pelvic-fin soft rays all
branched; third (rarely second) ray longest, much longer than upper jaw; tip of longest ray not reaching to anus
when depressed; last ray joined by membrane to abdomen for about one-fourth to one-third its length. Caudal fin
rounded, usually with 3 procurrent, 2 segmented unbranched, and 5 segmented branched rays in dorsal series, and
3 procurrent, 2 segmented unbranched, and 5 segmented branched rays in ventral series.
Remarks. Fowler (1938) proposed Brachypterois as a new monotypic genus for his new species, B. serrulifer,
stating that Brachypterois was characterized by having the following characters: short dorsal-fin spine, its length
shorter than that of dorsal-fin soft ray; pectoral fin long, its posterior tip reaching near caudal-fin base; ridges on
head serrated, including mandible; no dermal flap or cirri on head and body. Subsequently, the presence of spinous
ridges on the mandible has been considered by several researchers (e.g., Kanayama & Amaoka 1981; Eschmeyer
1986; and Poss 1999) as the main diagnostic feature of Brachypterois. However, it is now known that
Brachypterois is sexually dimorphic for this character (see under Sexual dimorphism).
Whitely (1951) proposed Ranipterois as a replacement name for Brachypterois Fowler 1938, with B. serrulifer
as the type species, regarding Brachypterois Fowler to be preoccupied by “Brachypterois Günther” as stated in
Jordan & Seale (1906: 189). However, that usage by Jordan & Seale (1906) was a lapsus calumni for the genus
Bathypterois Günther 1878 (Ipnopidae). Therefore, Ranipterois should be treated as an unnecessary replacement
name and an objective synonym of Brachypterois Fowler 1938 (Kanayama & Amaoka, 1981; this study).
Some previously publishes illustrations of Brachypterois [e.g., B. serrulifer in Fowler (1938: 80, fig. 35); B.
serrulata in Poss (1999: 2310, unnumbered fig. on bottom); and Nakabo (2002: 567, unnumbered fig. on top)] have
shown broad membranes in the spinous portion of the dorsal-fin. However, underwater photographs of live
individuals of Brachypterois have shown strongly incised spinous dorsal fin membranes (see Allen et al. 2003;
Allen & Erdmann 2012), supporting Barnhart & Hubbs (1946), who pointed out the inconsistency in dorsal fin
membrane shape between the figure of B. serrulifer given by Fowler (1938) and the holotype of the species.
Key to the species of Brachypterois
1a. Pectoral-fin rays usually 16 (rarely 15 or 17); 0–22 (mean 3.9) spines on median lateral ridge of maxilla in specimens >60 mm
SL; caudal fin with relatively few large dark spots, 2–10 spots on longest (middle) ray; posterior lacrimal spine usually
directed downward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. serrulifer (Indo-West Pacific)
1b. Pectoral-fin rays usually 15 (rarely 14 or 16); spines usually absent on median lateral ridge of maxilla (rarely 1–3 spines pres-
ent in specimens > 80 mm SL in B. serrulata); caudal fin with relatively large number of small sized dark spots, 5–26 spots on
(middle) longest ray; posterior lacrimal spine usually directed posteroventrally to posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Body with 44–50 (modally 47) scale rows in longitudinal series; 4–6 (5) scales above lateral line; 4–7 (6) scale rows between
last dorsal-fin spine base and lateral line; relatively short head, its length 38.3–43.1 (mean 40.2)% SL; relatively small orbit, its
diameter 11.1–14.5 (12.4)% SL; relatively short caudal fin, its length 30.4–44.7 (35.7)% SL; posterior lacrimal spine usually
directed posteroventrally, with no upward distal curve; spine(s) at corner of outer angular ridge developed only in adults, of
similar size to adjacent spines on ridge, directed ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . B. serrulata (northwestern Pacific)
2b. Body with 41–45 (modally 44) scale rows in longitudinal series; 4–5 (4) scales above lateral line; 4–5 (4) scale rows between
last dorsal-fin spine base and lateral line; relatively long head, its length 41.6–44.5 (mean 43.1)% SL; relatively large orbit, its
diameter 13.0–15.4 (14.3)% SL; relatively long caudal fin, its length 37.8–46.4 (42.3)% SL; posterior lacrimal spine usually
directed posteroventrally to posteriorly, with an upward distal curve in large adults; spine(s) at corner of outer angular ridge
developed in all stages, of greater size than adjacent spines on ridge, strongly directed posteriorly . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. curvispina sp. nov. (northeastern Australia)

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REVISION OF BRACHYPTEROIS
Brachypterois curvispina sp. nov.
[New English name: Australian Sawcheek Scorpionfish]
(Figures 3–5, 6A, 7, 8A–C, 9A–C, 10–11, 19, 20A; Tables 1–2)
Holotype. CSIRO H7204-02 (X), male, 78.5 mm SL, southeast of Weary Bay, Queensland, Australia (15°55.36′S,
145°39.60′E–15°55.15′S, 145°40.14′E), 40 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 18 Nov. 2003.
Paratypes. 29 specimens, 44.0–86.0 mm SL (all from Queensland, Australia)—CSIRO H6707-04 (9
specimens), 45.8–86.0 mm SL, southeast of Swain Reefs (22°33.63′S, 152°12.88′E–22°34.18′S, 152°13.05′E), 96
m, D. Gledhill, R/V Gwendoline May, demersal trawl, 24 Nov. 2005; CSIRO H6784-02 (X), female, 75.2 mm SL,
east of Cairns (16°54.41′S, 146°07.43′E–16°53.84′S, 146°07.63′E), 40 m, D. Gledhill, R/V Gwendoline May,
demersal trawl, 23 Nov. 2003; CSIRO H6923-02 (2), 66.1–68.1 mm SL, east-northeast of Shoalwater Bay
(22°04.15′S, 152°09.74′E–22°03.83′S, 152°09.20′E), 88 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 23
Nov. 2005; CSIRO H7024-03 (X), 70.3 mm SL, north of Cairns (16°33.46′S, 145°52.17′E–16°33.20′S,
145°52.69′E), 37 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 22 Nov. 2003; CSIRO H7097-02 (6),
44.0–70.8 mm SL, south of Swain Reefs (22°25.25′S, 152°09.84′E–22°25.78′S, 152°10.05′E), 98 m, D. Gledhill,
R/V Gwendoline May, demersal trawl, 24 Nov. 2005; CSIRO H7277-04 (X), 84.9 mm SL, Swain Reefs
(22°46.94′S, 152°31.86′E–22°43.19′S, 152°38.22′E), 130 m, C. Rigby, R/V Benjamin, prawn trawl, 24 June 2011;
CSIRO H7377-01, 67.5 mm SL, northeast of Gladstone (22°41.29′S, 152°07.01′E–22°40.90′S, 152°07.52′E), 86
m, D. Gledhill, R/V Gwendoline May, demersal trawl, 23 Apr. 2004; CSIRO H7378-01 (2; 1 female, 61.0 mm SL),
50.7–61.0 mm SL, northeast of Gladstone (22°39.46′S, 151°52.98′E–22°39.25′S, 151°52.44′E), 71 m, D. Gledhill,
R/V Gwendoline May, demersal trawl, 22 Nov. 2005; CSIRO H7379-01, female, 59.6 mm SL, northeast of Cairns
(16°32.08′S, 146°02.02′E–16°32.09′S, 146°01.40′E), 54 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 22
Nov. 2003; CSIRO H7380-01 (X), 74.8 mm SL, east of Cairns (16°48.15′S, 146°05.85′E–16°48.56′S,
146°05.49′E), 45 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 22 Nov. 2003; QM I.19656, 73.3 mm SL,
Flora Passage (17°08′S, 146°15′E), 43.9 m, Queensland Fisheries Service, 16 Oct. 1979; QM I.35150, 52.6 mm
SL, southeast of Endeavour Reef (15°49′30″S, 145°40′30″E), Seabed Biodiversity Study Team, dredge, 3 Oct.
2003; QM I.36322 (X), 63.5 mm SL, Capricorn Channel (22°32′06″S, 152°09′18″E), 95 m, Seabed Biodiversity
Study Team, dredge, 14 May 2004; QM I.36386, 68.0 mm SL, southwest of Swain Reefs (22°15′18″S,
152°07′30″E), 100 m, Seabed Biodiversity Study Team, dredge, 15 May 2004.
Non-type specimens. 2 specimens, 57.4–68.2 mm SL (both from Queensland, Australia)—CSIRO H6750-01,
57.4 mm SL, east of Dalrymple Island (9°40.61′S, 143°31.21′E–9°39.96′S, 143°31.31′E), 35 m, D. Gledhill, R/V
Gwendoline May, demersal trawl, 28 Jan. 2004; CSIRO H7381-01 (X), 68.2 mm SL, north of Bramble Cay
(9°03.70′S, 143°47.47′E–9°03.57′S, 143°48.03′E), 32 m, D. Gledhill, R/V Gwendoline May, demersal trawl, 27
Jan. 2004.
Diagnosis. A species of Brachypterois with the following combination of characters: pectoral-fin rays 14–16
(modally 15); scale rows in longitudinal series 41–45 (44); scales above lateral line 4–5 (4); scale rows between
last dorsal spine base and lateral line 4–5 (4); pre-dorsal scale rows 3–5 (4); suborbital scale rows 0–2 (1) (Fig.
6A); total gill rakers 16–18 (16); head length 41.5–44.5 (mean 43.1) %SL (Fig. 7); orbit diameter 13.0–15.4 (14.3)
%SL; upper-jaw length 20.2–23.0 (21.6) %SL; longest dorsal-fin soft ray length 19.5–24.5 (22.5) %SL; longest
anal-fin soft ray length 23.6–28.4 (25.3) %SL; caudal-fin length 37.8–46.4 (42.3) %SL; posterior lacrimal spine
usually directed posteriorly, the tip being strongly curved upward in large adults (Fig. 8A–C); spine(s) on corner of
outer angular ridge longer and stouter than others on ridge, directed posteriorly (Fig. 9A–C); number and size of
caudal fin spots moderate, 5–13 (mean 9.3) spots on longest caudal-fin ray (Figs. 3–5, 10).
Description. Meristics and morphometrics of specimens examined are shown in Tables 1–2. Characters stated
in the generic definition and specific diagnoses are not repeated here. Data for the holotype are presented first,
followed by paratype and non-type data (if different) in parentheses. Gill rakers simple, short and robust; longest
raker on first gill arch shorter or subequal to gill filaments around angle of gill arch; fourth gill slit narrow, partially
closed by membrane. Branchiostegal rays 7. Vertebrae 10 + 14. Formula for configuration of supraneural bones,
anterior neural spines and anterior dorsal pterygiophores usually //2+1/1/1/1/1/1/1/1/1/1+1/. Swimbladder present.
Body oblong, moderately compressed anteriorly, strongly compressed posteriorly. Head large, its length
greater than body depth. Low membranous tube associated with anterior and posterior nostrils; posterior edge of
tube on anterior nostril slightly expanded. Ctenoid scales covering head, including cheek, lacrimal, suborbital pit,

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interopercle, subopercle, opercle, posterodorsal pit of supraocular, channel between interorbital ridges, occiput and
suprapostorbital area surrounded by parietal, pterotic, lower posttemporal and cleithral spine bases; scales absent
on lips and bulge on snout. Maxilla with weakly ctenoid scales; posterior portion of maxilla entirely scaled.
Mandible with exposed weakly ctenoid scales. Ctenoid scales covering body, extending onto basal caudal fin;
weakly ctenoid scales on ventral body surface and pectoral-fin base; a row of elongated cycloid scales on bases of
soft-rayed portions of dorsal and anal fins. Lateral line complete, weakly sloping downward posteriorly. Three
sensory pores on underside of each dentary; a small pore on each side of symphysial knob.
TABLE 1. Frequency distribution of selected meristics of Brachypterois curvispina sp. nov., B. serrulata and B.
serrulifer.
H
and
N
indicate holotype and neotype, respectively; DS and LL indicate dorsal-fin spine base and lateral line,
respectively.
Mouth moderately large, oblique, forming an angle of ca. 40° (40–50°) to horizontal axis of head and body;
upper edge of maxilla swollen laterally forming a low ridge; median lateral ridge of maxilla lacking spines;
posterior end of maxilla extending slightly beyond a vertical through mid-orbit. Symphysial gap separating
premaxillary teeth bands about twice width of each band; upper jaw with a band of small slender conical teeth;
about 4–5 (4–6) tooth rows at front of upper jaw; about 4 tooth rows at front of lower jaw. Vomer with a V-shaped
patch of 2–4 (1–4) rows of small teeth.
Dorsal profile of snout steep, forming an angle of ca. 40° to horizontal axis of head and body. Nasal spine base
relatively long, its length subequal to distance between anterior and posterior nostrils; with a row of 7 (2–8) small
spines. Interorbital ridges moderately developed, with a row of 1 (1–8) spines; separated by a shallow channel,
initially diverging posteriorly; conjoined with origin of coronal spine base (not in small specimens). Interorbital
space shallow, about one-seventh of orbit diameter extending above dorsal profile of head. Coronal spine with 2
(2–6) small spinous points on short base; its base conjoined with origin of parietal spine base; 5 (2–5) small spines
forming a pair of small patches between origins of coronal spine base. Tympanic with 2 spinous ridges; upper ridge
shorter than lower ridge with 3 (2–5) spines; lower ridge with 7 (2–7) spines. Preocular with 10 (3–11) spines on its
frontal surface and margin of orbit. Supraocular with a transverse ridge located between interorbital ridge and outer
edge of supraocular, with 3 (3–7) spines; 12 (9–16) spines on margin of orbit; posterodorsal edge of supraocular
expanded posteriorly, with a small inner pit. Postocular with 8 (3–8) spines on margin of orbit. Occipital area
lacking anterior ridge; a pair of short low ridges with 7 (1–7) spines on posterior margin between parietal spine
bases. Parietal spine with 8 (7–18) spinous points on a relatively long ridge; its base diverging posteriorly; a short
ridge with 2 (0–7) spines just under posterior portion of parietal ridge. Sphenotic spine with 12 (7–12) spinous
Pectoral-fin rays Total gill rakers
14/14 14/15 15/15 15/16 16/16 16/17 17/17 15 16 17 18 19
B. curvispina 1
8
H
21 1
15
H
12 5
B. serrulata 48
90
N
95 127
77
N
71
B. serrulifer 511
88
H
41 22
70
H
14 1
Scale rows in longitudinal series
41 42 43 44 45 46 47 48 49 50 51
B. curvispina 2
4
H
5132
B. serrulata 3 5 25 29 19
17
N
71
B. serrulifer 1 4 20 19 26
19
H
131
Scales above lateral line Scale rows between last DS and
LL
Pre-dorsal scale rows
4567 4567 345678
B. curvispina 23
7
H
16
H
14 2
18
H
9
B. serrulata 1
81
N
29 1 35
59
N
116
68
N
21 1 1
B. serrulifer 272
29
H
31
53
H
45 1 20 52
32
H
1

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REVISION OF BRACHYPTEROIS
points. No postorbital spine. Pterotic spine with 17 (7–20) spinous points. Small patch of 2 (0–2) spines behind
pterotic spine. A short oblique ridge with 2 (0–2) spines between parietal spine and lower posttemporal spine. No
upper posttemporal spine. Lower posttemporal spine with 5 (1–5) spinous points on a relatively short ridge; a short
oblique ridge with 2 (0–2) spines just above posterior portion of spine base; a small patch of 3 (0–3) spines in front
of spine. Supracleithral spine with a spinous point on a relatively short ridge. Cleithral spine with 5 (3–11) spinous
points on a relatively long ridge.
TABLE 2 Meristics and morphometrics, expressed as percentages of standard length, of Brachypterois curvispina sp.
nov.
Holotype
CSIRO H7204-02
Paratypes
(n = 29)
Non-types
(n = 2)
SL (mm) 78.5 44.0–86.0 57.4–68.2
Meristics Modes
Dorsal-fin rays XIII, 10 XIII–XIV, 8–11 XIII, 10 XIII, 10
Anal-fin rays III, 5 III, 5 III, 5 III, 5
Pectoral-fin rays
a
ii + 7 + vi = 15/14 i–ii + 6–8 + v–viii =
14–15/14–16
ii + 6–7 + vi–vii =
15/15
ii + 6 + vi = 15/15
Scale rows in longitudinal series 42 41–45 44 44
Pored lateral-line scales 25 24–25 — 25
Scales above lateral line 5 4–5 4–5 4
Scales below lateral line — 8–10 10 10
Scale rows between 6th DS and LL 4 4–5 5 5
Scale rows between last DS and LL 4 4–5 5 4
Pre-dorsal scale rows 4 3–5 4–5 4
Suborbital scale rows 2 0–2 1 1
Gill rakers
b
5 + 11 = 16 5–6 + 11–12 = 16–18 5 + 12–13 = 17–18 5 + 12 = 16
Morphometrics (% SL) Means
Body depth 35.8 34.8–40.4 36.5–36.6 37.0
Body depth at anal-fin origin 28.8 25.9–31.4 29.4–29.9 29.2
Body width 22.2 19.1–26.5 22.0–22.9 22.6
Head length 41.5 41.6–44.5 42.0–42.2 43.1
Head width 14.6 14.5–16.6 15.0–16.1 15.5
Snout length 8.0 7.7–9.0 8.1–9.1 8.4
Orbit diameter 13.2 13.0–15.4 13.8–14.4 14.3
Interorbital width
c
5.5 5.5–6.7 5.7–6.2 6.0
Interorbital width
d
4.1 4.3–6.0 4.7–5.1 5.1
Upper jaw length 21.1 20.2–23.0 21.6 21.6
Maxillary depth 7.3 7.4–9.3 7.8–8.7 8.3
Suborbital depth 0.8 0.3–1.3 0.7–0.9 0.8
Postorbital length 21.7 20.1–22.1 20.7–20.8 21.1
Predorsal-fin length 34.1 33.6–37.4 35.4–35.5 35.5
Preanal-fin length 76.1 73.6–78.5 74.5–80.0 75.8
Prepelvic-fin length 37.3 35.5–40.8 38.0–40.1 38.2
1st dorsal-fin spine length — 10.5–14.0 10.3–12.2 11.5
2nd dorsal-fin spine length 14.9 14.0–17.9 13.5 15.2
3rd dorsal-fin spine length 16.4 15.6–17.8 14.7–17.6 16.6
......continued on the next page

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a
Upper unbranched + branched + lower unbranched rays = total rays on left side of body/ total rays on right side.
b
Rakers on upper limb + rakers on lower limb = total rakers.
c
At vertical midline of eye.
d
At posterior end of preocular spine base.
DS and LL indicate dorsal-fin spine base and lateral line, respectively.
Anterior portion of articular head of maxilla with 3 (0–4) spines; lateral portion of articular head lacking spine,
exposed when mouth closed. Inner edge of dentary elevated as a low ridge; central dentary ridge running along
sensory canal, comprising 2 ridges posteriorly; outer margin of dentary a low ridge; tip of ventral surface of dentary
slightly expanded ventrally. Outer edge of angular a low ridge; inner angular ridge running along sensory canal.
Retroarticular lacking spines. Lacrimal with 2 lateral ridges; upper ridge with 4 (3–5) spines, its anterior origin
conjoined with ridge on dorsal articular process of lacrimal; lower ridge with 3 (1–4) spines. Anterior lacrimal
spine with 3 (1–4) spinous points, directed downward. Posterior lacrimal spine broad, with 4 (2–4) spinous points;
all ridges on lacrimal strongly spinous. Suborbital with 2 ridges; both ridges with supplementary upper and lower
ridges; posterior upper ridge branched into two posteriorly; 2 small arched ridges below anterior and posterior
ridges; all ridges strongly spinous [60 (32–60) spines in total]. Poorly developed suborbital pit present. Preopercle
spines with one or more spinous points; lateral ridge of uppermost spine with 1 (0–2) spines. A ridge on preopercle
submargin strongly spinous. Two poorly developed spinous vertical ridges on postorbital. Interopercle lacking
spine. Upper opercular spine simple with a spinous point; a simple lower opercular spine present but usually under
skin (sometimes absent).
TABLE 2. (Continued)
Holotype
CSIRO H7204-02
Paratypes
(n = 29)
Non-types
(n = 2)
4th dorsal-fin spine length 17.5 15.9–19.3 — 17.7
5th dorsal-fin spine length 19.1 16.9–19.0 — 18.1
6th dorsal-fin spine length 18.9 17.4–20.0 16.0–18.8 18.4
7th dorsal-fin spine length — 17.0–20.5 19.5 18.5
8th dorsal-fin spine length — 16.1–20.0 15.8–17.9 17.4
9th dorsal-fin spine length 15.7 13.3–17.9 16.2 15.4
10th dorsal-fin spine length 12.0 9.7–14.1 9.1–12.2 11.3
11th dorsal-fin spine length 8.7 5.7–9.4 5.3–8.0 7.0
12th dorsal-fin spine length 5.2 3.0–4.9 4.4–5.3 4.2
13th dorsal-fin spine length 10.7 9.7–13.2 11.6–12.4 11.5
1st dorsal-fin soft ray length 17.2 16.8–21.0 17.7–20.6 18.9
Longest dorsal-fin soft ray length 22.5 19.5–24.5 21.1–23.5 22.5
1st anal-fin spine length 7.5 6.1–8.2 7.2 7.0
2nd anal-fin spine length 12.7 11.5–13.9 12.0–13.1 12.7
3rd anal-fin spine length 15.9 14.6–17.2 15.2–16.9 15.9
1st anal-fin soft ray length 24.3 22.2–27.9 22.3–25.8 24.3
Longest anal-fin soft ray length 25.7 24.0–28.4 23.6–27.0 25.3
Pectoral-fin ray length 56.4 46.6–58.2 52.8–60.6 53.0
Pelvic-fin spine length 17.1 15.4–18.0 15.1–17.9 16.7
Longest pelvic-fin soft ray length 29.6 25.8–32.7 26.8–33.3 29.1
Caudal-fin length 41.3 37.8–46.4 39.9–42.0 42.3
Caudal-peduncle length 17.5 14.9–18.8 17.4 17.1
Caudal-peduncle depth 11.0 10.3–11.6 10.6–11.3 10.8

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REVISION OF BRACHYPTEROIS
FIGURE 3. Preserved specimens of Brachypterois curvispina sp. nov. (A) CSIRO H6707-04 (1 of 9 specimens), 48.8 mm SL,
paratype; (B) CSIRO H6923-02 (1 of 2), 66.1 mm SL, paratype; (C) CSIRO H7024-03, 70.3 mm SL, paratype; (D) CSIRO
H6784-02, female, 75.2 mm SL, paratype; (E) CSIRO H7204-02, male, 78.5 mm SL, holotype.

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FIGURE 4. Preserved specimens of Brachypterois curvispina sp. nov. (A) CSIRO H7381-01, 68.2 mm SL; (B) CSIRO
H7380-01, 74.8 mm SL, paratype.
Origin of first dorsal-fin spine above cleithral spine; bases of first and second dorsal-fin spines closer than
those of succeeding adjacent spines; fifth (usually sixth or seventh, rarely fifth) spine longest; length of longest
spine about half of maximum body depth; penultimate (twelfth) spine shortest; membranes of spinous portion of
dorsal fin strongly incised, especially in anterior portion. Dorsal-fin soft rays all branched; fourth (sometimes third
or fifth) soft ray longest; length of longest ray subequal to or slightly longer than longest fin spine; posterior
branched ray not joined by membrane to caudal peduncle. Origin of first anal-fin spine below base of penultimate
(twelfth) dorsal-fin spine; third spine longest. Anal-fin soft rays all branched; second (sometimes first or third) soft
ray longest, its length greater than that of longest dorsal-fin soft ray; posterior branch of last soft ray not joined by
membrane to caudal peduncle. Pectoral fin long; fifth ray [fifth or sixth (branched) in relatively large specimens of
45.8–84.9 mm SL; ninth or tenth (unbranched) in relatively small specimens of 44.0–70.8 mm SL] longest; tip of
longest ray reaching a vertical through base of last anal-fin soft ray, but not reaching level with caudal-fin base;
lower unbranched rays weakly thickened. Pelvic-fin soft rays all branched; third (rarely second) ray longest, much

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REVISION OF BRACHYPTEROIS
longer than upper jaw; tip of longest ray not reaching to anus when depressed; last ray joined by membrane to
abdomen for about one-fourth (one-fourth to one-third) its length. Caudal fin with 14 segmented rays, 10 rays
branched, 4 remaining rays unbranched; 3 unsegmented and spinous procurrent rays in dorsal and ventral series;
posterior margin of fin rounded.
Color of fresh specimen (based on Fig. 5). Head and body reddish, darker dorsally, paler ventrally; a poorly
defined large black blotch (subequal to orbit diameter) on opercle. Five poorly defined reddish-brown bands on
head; anteriormost band running from anterior margin of eye over snout to tip of lower jaw; second from
anteroventral margin of eye extending onto lacrimal and both jaws; third just below eye reaching posterodorsal
corner of maxilla; fourth from posteroventral margin of eye to posterior portion of opercle; fifth from posterior
margin of eye to blotch on opercle. Five poorly defined vertical brown bands on body (not clearly observed due to
missing scales in most specimens examined). Ground color of dorsal and anal fins similar to that of head and body.
Numerous dark red spots, forming a more or less dotted line, on soft-rayed portion of dorsal fin. Anal fin with
about 6 similarly dotted dark red lines. Pectoral fin membrane dark brown; numerous creamy-white spots on rays,
forming about 6 dotted bands. Pelvic fin pinkish-white, with about 8 poorly defined dark brown bands. Caudal fin
membrane translucent; about 60 small dark red spots on rays. Eye dull yellow; iris black.
FIGURE 5. Fresh specimen of Brachypterois curvispina sp. nov., CSIRO H7381-01, 68.2 mm SL (photo: D. Gledhill).
Color of preserved specimens. Head and body creamy-white to brownish-white, darker dorsally, paler
ventrally; a poorly defined large brown to black blotch (subequal to orbit diameter) on opercle. Bulge on snout, tips
of jaws, cheek and postorbital area partly blackish. A blackish band obliquely crossing eye to interopercle; a
blackish band saddling nape and reaching center of opercle. Five poorly defined vertical blackish bands on body;
anteriormost band below between first and third dorsal-fin spine bases; second below between fourth and sixth
spine bases; third below between eighth and penultimate spine bases; fourth Y-shaped, upper margin of anterior
branch below last spine and third soft ray bases, upper margin of posterior branch below sixth and ninth soft ray
bases, both branches connected at level of lateral line or just below dorsal-fin base; fifth on caudal peduncle just in
front of caudal-fin base. Dorsal fin with translucent to dusky membrane; a large black blotch on spinous portion
between about sixth and ninth spines; about 6 (5–6) poorly defined black bands on soft-rayed portion. Anal fin with
translucent to dusky membrane and about 6 (5–8) poorly defined narrow black bands. Pectoral fin membrane
blackish, with about 6 (4–6) poorly defined narrow black bands. Pelvic fin membrane blackish, with about 5 (4–7)
poorly defined narrow black bands. Caudal fin membrane translucent with about 90 (80–90) small black spots on
rays; maximum spot diameter 2.6 [1.8–3.7 (mean 2.6) based on 11 specimens] times in caudal-fin ray width at
position where spot present on ray. Eye dark blue; iris white.

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FIGURE 6. Relationships of number of suborbital scale rows to standard length in (A) Brachypterois curvispina sp. nov., (B)
B. serrulata and (C) B. serrulifer.
Distribution. Brachypterois curvispina is currently known only from northern and northeastern coasts of
Queensland, Australia, from the east entrance of Torres Strait south to Swain Reefs and the Capricorn Islands (Fig.
11). The specimens of the 15 lots examined were collected from depths 32–130 m (average 70 m).
Etymology. The specific name “curvispina” refers to one of the diagnostic characters of this species, the
dorsally curved posterior lacrimal spine present in large adults.
Remarks. Two specimens (CSIRO H7381-01, 68.2 mm SL and CSIRO H6750-01, 57.4 mm SL) collected
from the extreme northern end of its range near the Australia/New Guinea border (off Bramble Cay in 32 m depth
and off Dalrymple Island in 35 m depth, respectively), differed from the typical form of B. curvispina in having a
relatively darker body and fin coloration in preserved specimens, and shallower sampling depths. Accordingly,
they were excluded from the type series. Additional material from the northern distributional range is necessary to
determine the extent of intraspecific variations in B. curvispina.

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REVISION OF BRACHYPTEROIS
FIGURE 7. Relationships of: (A) head length; (B) orbit diameter; (C) upper jaw length; (D) longest dorsal-fin soft ray length;
(E) longest anal-fin soft ray length; and (F) caudal-fin length to standard length in Brachypterois curvispina sp. nov. (stars,
solid lines), B. serrulata (circles, dotted lines) and B. serrulifer (triangles, chain lines). H1 holotype of B. curvispina; H2
holotype of B. serrulifer; N neotype of Sebastes serrulatus.

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FIGURE 8. Spines on lacrimal bones of Brachypterois curvispina sp. nov. (A–C), B. serrulata (D–F) and B. serrulifer (G–I).
(A) CSIRO H7204-02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H6784-02, female, 75.2 mm SL, paratype of
B. curvispina; (C) CSIRO H6750-01, sex undetermined, 57.4 mm SL; (D) KAUM–I. 24056, male, 97.9 mm SL; (E) KAUM–I.
24054, female, 76.5 mm SL; (F) KAUM–I. 24050, female, 63.9 mm SL; (G) KAUM–I. 22033, male, 83.0 mm SL; (H) CAS
60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. LLS lateral lacrimal
spine; ALS anterior lacrimal spine; PLS posterior lacrimal spine. Left to anterior. Bars indicate 5 mm.
Brachypterois serrulata (Richardson 1846)
[English name: Sawcheek Scorpionfish]
[Standard Japanese name: Nokogiri-kasago]
(Figures 2, 6B, 7, 8D–F, 9D–F, 10–13, 14A, 15, 18–19, 20B; Tables 1, 3)
Sebastes serrulatus Richardson 1846: 215 [original locality: “Sea of China”; type locality, based on neotype designated herein:
off Dong Kang, Pingtung, southern Taiwan]; Günther 1860: 106 (type catalog of BMNH)
Scorpaenodes serrulatus: Chu 1931: 142 (in part?; Sea of China; listed)
Brachypterois serrulatus: Chu 1962: 861, fig. 692 (South China Sea; short description); Chu et al. 1963: 462, fig. 348 (East
China Sea coast of China; short description); Chen 1981: 47, figs. 9-29, 32 (Taiwan; short description); Kanayama &
Amaoka 1981: 181, figs. 1, 2 (in part; Tosa Bay, Kochi, Japan; short description); Shen 1984: 185, unnumbered fig. on p.
192 (in part?; Taiwan; key); Amaoka 1984: 302, pl. 360-G [in part?; Japan; figure taken from Fowler 1938]; Eschmeyer
1986: 465 (in part; Indo-West Pacific; key); Jean & Kuo 1988: 55 (in part?; Taiwan; listed); Wang 1993: 112 (in part?;
South China Sea; listed); Ishida 1994: 5 (as phylogenetic material); Huang 1994: 739 (in part?; China, Taiwan and South
China Sea; listed); Qingwen et al. 1995: 549, fig. 736 (in part?; Japan to South China Sea; short description); Allen 1997:
76, pl. 19, fig. 7 (in part; Indo-West Pacific; ecological note and diagnosis); Chen et al. 1997: 168, unnumbered fig. on top
(in part?; Spratly Islands to China; short description); Poss 1999: 2299, 2310, unnumbered fig. on bottom on p. 2310 (in
part; Indo-West Pacific; key and short description; ); Randall & Lim 2000: 604 (in part?; South China Sea; listed);
Matsuura et al. 2001: 106 (Sanya Bay, Hainan Island, China; listed; specimen examined herein); Adrim et al. 2004: 119 (in
part?; Anambas Islands, South China Sea; listed)
Brachypterois serrulata: Nakabo 1993: 493, unnumbered fig. on top [Kochi, Japan; key; figured specimen from Gulf of
Thailand (T. Nakabo, 2012, pers. comm.) examined herein]; Ishida 1997: 209, unnumbered fig. on p. 209 (Tosa Bay,
Kochi, Japan; short description; Indo-West Pacific; figured specimen examined herein); Nakabo 2000: 567, unnumbered
fig. on top (Kochi, Japan; figure taken from Nakabo 1993); Murai 2001: 172, unnumbered fig. on middle (Iburi, Kochi,
Japan; short description;); Shinohara et al. 2001: 314 (in part; Tosa Bay, Kochi, Japan; listed; based on Amaoka 1984 and

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REVISION OF BRACHYPTEROIS
Nakabo 1993); Mandrytsa 2001:276 (as phylogenetic material; key); Nakabo 2002: 567, unnumbered fig. on top (Kochi,
Japan; figure taken from Nakabo 1993); Ide et al. 2003: 24 (Tosa Bay, Kochi, Japan; listed); Xinbo 2006: 270, fig. 156
(China and Taiwan; description); Shao et al. 2008: 245 (in part ?; Taiwan; listed; based on Chen 1981)
Dendrochirus bellus (not of Jordan & Hubbs): Rainboth et al. 2012: 80, pl. 55, fig. 1137 (Gulf of Thailand; listed)
FIGURE 9. Spines on angular bones of Brachypterois curvispina sp. nov. (A–C), B. serrulata (D–F) and B. serrulifer (G–I).
(A) CSIRO H7204-02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H6784-02, female, 75.2 mm SL, paratype of
B. curvispina; (C) CSIRO H6750-01, sex undetermined, 57.4 mm SL; (D) KAUM–I. 24056, male, 97.9 mm SL; (E) KAUM–I.
24054, female, 76.5 mm SL; (F) KAUM–I. 24050, female, 63.9 mm SL; (G) KAUM–I. 22033, male, 83.0 mm SL; (H) CAS
60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. Arrows indicate
spine(s) on posterior corner of outer angular ridge. Left to anterior. Bars indicate 1 mm.
Neotype. ASIZP 62399 (X), 92.8 mm SL, Dong-kang, Pingtung, southern Taiwan (22°31′N, 120°22′E), 100 m, J.-
H. Wu, bottom trawl, 1 Mar. 2001.
Non-type specimens. 112 specimens, 48.0–99.3 mm SL—JAPAN (following BSKU specimens all collected
by bottom trawl from Tosa Bay, Kochi): BSKU 12876, 59.2 mm SL, obtained at Usa Port, Sept. 1968; BSKU
42455, male, 84.2 mm SL, obtained at Mimase Port, 26 Nov. 1985; BSKU 52300, male, 91.8 mm SL, BSKU
52301, male, 93.2 mm SL, BSKU 52302, male, 72.9 mm SL, obtained at Irino Port, 10 July 2000; BSKU 52359,
male, 73.2 mm SL, obtained at Irino Port, 2 Aug. 2000; BSKU 52411, male, 71.4 mm SL, obtained at Kochi City
Central Fish Market, 19 Jan. 1995; BSKU 54828, 95.2 mm SL, obtained at Irino Port, 3 July 2000; BSKU 57572,
67.2 mm SL, obtained at Irino Port, 22 Feb. 2002: BSKU 57736, male, 69.3 mm SL, obtained at Irino Port, 12 Sept.
2001; BSKU 57874, male, 96.7 mm SL, obtained at Irino Port, 20 May 1999; BSKU 59821, female, 70.7 mm SL,
obtained at Kamikawaguchi Port, 31 July 1997; BSKU 62168, 64.8 mm SL, obtained at Irino Port, 3 Oct. 2002;

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BSKU 62457, male, 92.0 mm SL, obtained at Susaki Port, 10 Oct. 2003; BSKU 67433, 71.5 mm SL, obtained at
Susaki Port, 4 July 2002; BSKU 67592, male, 99.3 mm SL, BSKU 67593, 91.1 mm SL, obtained at Irino Port, 20
Mar. 1999; BSKU 70509, 71.3 mm SL, BSKU 70510, 86.5 mm SL, BSKU 70511, male, 65.7 mm SL, obtained at
Kamikawaguchi Port, 20 Feb. 1997; BSKU 86119, male, 91.4 mm SL, BSKU 86121, female, 72.5 mm SL,
obtained at Irino Port, 12 Mar. 1999; BSKU 86123, male, 73.4 mm SL, BSKU 86124, female, 67.2 mm SL,
obtained at Irino Port, 12 Mar. 1999; BSKU 82651, 48.0 mm SL, 75 m, R/V Toyohata-maru, 2 Nov. 1989; BSKU
82652, 54.6 mm SL, BSKU 82653, female, 67.2 mm SL, BSKU 82654, male, 74.6 mm SL, obtained at
Kamikawaguchi Port, H. Endo, 20 Mar. 1997; BSKU 86120, 76.6 mm SL, obtained at Irino Port, 12 Mar. 1999;
BSKU 86150, 63.1 mm SL, obtained at Kamikawaguchi Port, 27 Apr. 1999; BSKU 86476 (2), 62.1–63.4 mm SL,
obtained at Kamikawaguchi Port, 27 Apr. 1999; BSKU 89187, 52.1 mm SL, obtained at Saga Port, 27 Oct. 2006;
BSKU 106470, 85.7 mm SL, BSKU 106471, 82.4 mm SL, BSKU 106472, female, 68.0 mm SL, BSKU 106473,
74.9 mm SL, obtained at Irino Port, 11 June 1999; FAKU 132868 (5), 68.9–81.2 mm SL, Irino, Tosa Bay, Kochi,
26 June 1997;
*
HUMZ 36292, 72.1 mm SL, HUMZ 36563, 71.7 mm SL,
*
HUMZ 36638, 61.3 mm SL, Tosa Bay,
Kochi, K. Amaoka, trawl; KAUM–I. 6876 (X), male, 66.9 mm SL, 1 km north off Izashiki Port, Sata,
Minamiosumi, Kagoshima (31°05′N, 130°41′E), 40 m, M. Yamada, set net, 27 June 2007; KAUM–I. 44569, 80.9
mm SL, KAUM–I. 44570, 89.4 mm SL, KAUM–I. 44571, 78.6 mm SL, KAUM–I. 44572, 75.6 mm SL, KAUM–I.
44573, female, 76.0 mm SL, KAUM–I. 44574, 71.2 mm SL, Tosa Bay, Kochi (obtained at Irino Port), bottom
trawl, 11 June 1999;
*
OMNH 24502, 67.1 mm SL, Tosa Bay, Kochi, H. Hanazaki, trawl, 20 May 1997; SNFR
11585, 86.1 mm SL, Tachibana Bay, Nagasaki, 17 Nov. 1987. EAST CHINA SEA: FAKU 12094, 83.0 mm SL,
1948–1949. CHINA: MNHN A-4310 (2), 74.5–84.6 mm SL; NSMT-P 60871, 69.1 mm SL, Sanya Bay, Hainan
Island, 16 m, G. Shinohara, 22 Nov. 1997. TAIWAN: ASIZP 62365, 74.0 mm SL, Dong-kang, Pingtung (22°31′N,
120°22′E), 100 m, J.-H. Wu, 16 May 2001; ASIZP 62400, 65.7 mm SL, Fong Kang, Pingtung (22°26′N,
120°38′E), 100 m, J.-H. Wu, 28 Feb. 2001;
*
ASIZP 62992, 60.5 mm SL, water entrance way of 1st Nuclear Power
Plant, Shihmen, Taipei (25°03′N, 121°56′E), Y.-C. Liao, 1 Apr. 2005; KAUM–I. 39283 (X), female, 60.2 mm SL,
KAUM–I. 39284 (X), male, 70.1 mm SL, off Tashi, Yilan, KAUM Fish Team, bottom trawl, 6 July 2011; KAUM–
I. 46474, female, 75.6 mm SL, off Tashi, Yilan, H.-C. Ho, bottom trawl, 18 Nov. 2007; NTOU 20071117-3, 76.2
mm SL, Hsiao-liou-ciou, Pingtung, 17 Nov. 2007; NMMB-P 4115, 79.3 mm SL, Fong-kang, Pingtung, J.-H. Wu,
23 Aug. 2001; NMMB-P 5181, 83.1 mm SL, Dong-kang, Pingtung, M.-C. Yu, 16 Feb. 1960;
*
NTUM 3854, 51.2
mm SL, off Tashi, Yilan, 18 July 1977. VIETNAM: FRLM 31479, 57.6 mm SL, FRLM 31647, 65.7 mm SL, Nha
Trang, S. Kimura et al., 7 Oct. 2004; HUMZ 190502, 60.7 mm SL, HUMZ 190503, 66.3 mm SL, HUMZ 190539,
62.8 mm SL, Nha Trang, H. Imamura, 8 Oct. 2004; HUMZ 190610, 61.6 mm SL, Nha Trang, H. Imamura, 7 Oct.
2004; NMMB-P 12648 (2), 69.6–70.1 mm SL, Nha Trang, H.-C. Ho, 16 Apr. 2009; NSMT-P 68863, 57.0 mm SL,
Nha Trang, G. Shinohara et al., 6 Dec. 2003; NSMT-P 69054, 55.4 mm SL, Nha Trang, K. Matsuura, 6 Dec. 2003;
NSMT-P 70619, 59.7 mm SL, Nha Trang, K. Shibukawa, 7 Oct. 2004. MALAYSIA: KAUM–I. 12313, male, 77.9
mm SL, off Kota Kinabalu, Sabah, Borneo, G. Ogihara, 24 Sept. 2008; KAUM–I. 22034, male, 82.4 mm SL, off
Kota Kinabalu, Sabah, Borneo, G. Ogihara, 11 Aug. 2009; KAUM–I. 49319, female, 67.1 mm SL, off Kota
Kinabalu, Sabah, Borneo, H. Nishiyama, 4 Aug. 2012. THAILAND (following specimens all collected from Gulf
of Thailand): KAUM–I. 23300, male, 72.5 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M.
Matsunuma, trawl, 10 Sept. 2009; KAUM–I. 23436 (X), female, 62.5 mm SL, KAUM–I. 23990 (X), male, 88.2
mm SL, KAUM–I. 23991 (X), male, 86.0 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M.
Matsunuma, trawl, 24 Sept. 2009; KAUM–I. 23882, male, 74.0 mm SL, KAUM–I. 23883 (X), female, 71.9 mm
SL, KAUM–I. 23884 (X), female, 86.3 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M.
Matsunuma, trawl, 26 Sept. 2009; KAUM–I. 24049 (X), female, 53.2 mm SL, KAUM–I. 24050 (X), female, 63.9
mm SL, KAUM–I. 24051 (X), female, 66.0 mm SL, KAUM–I. 24052 (X), female, 72.7 mm SL, KAUM–I. 24053
(X), female, 76.2 mm SL, KAUM–I. 24054 (X), female, 76.5 mm SL, KAUM–I. 24055 (X), male, 81.2 mm SL,
KAUM–I. 24056 (X), male, 97.9 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M.
Matsunuma, trawl, 5 Oct. 2009; KAUM–I. 33225, male, 86.2 mm SL, obtained at fish market in Samut Sakhon
Province, T. Yoshida, trawl, 30 Nov. 2010; KAUM–I. 47467, 81.5 mm SL, KAUM–I. 47468, male, 94.5 mm SL,
obtained at Mahachai fish market, Samut Sakhon Province, T. Yoshida, trawl, 20 June 2012; KAUM–I. 47472,
99.2 mm SL, KAUM–I. 47473, female, 82.5 mm SL, KAUM–I. 47474, 82.2 mm SL, KAUM–I. 47475, 93.3 mm
SL, KAUM–I. 47476, 90.3 mm SL, KAUM–I. 47477, 87.2 mm SL, KAUM–I. 47478, male, 87.1 mm SL, KAUM–
I. 47479, 80.0 mm SL, KAUM–I. 47480, female, 70.0 mm SL, obtained at Mahachai fish market, Samut Sakhon

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REVISION OF BRACHYPTEROIS
Province, T. Yoshida, trawl, 20 June 2012; KAUM–I. 47799, female, 68.4 mm SL, obtained at Klong Wan fish
landing port, Prachuab Khirikhan Province, T. Yoshida, trawl, 24 July 2012; URM 12150, 69.4 mm SL, obtained at
Song Khla fish market, H. Senou et al., 21 Oct. 1983. INDONESIA: NCIP 5002 (3), 43.2–64.0 mm SL, off Seraya
Island, Natuna Islands, trawl, 16 Apr. 2011.
FIGURE 10. Relationships of number of spots on longest caudal-fin ray to standard length in Brachypterois curvispina sp.
nov. (stars), B. serrulata (circles) and B. serrulifer (triangles; dotted symbols indicate specimens from the Indian Ocean,
remaining specimens from the Pacific Ocean). H holotype of B. serrulifer. Note: counts unable to be made in holotype of B.
curvispina and neotype of Sebastes serrulatus.
FIGURE 11. Distributional records of Brachypterois curvispina sp. nov. (stars), B. serrulata (circles) and B. serrulifer
(triangles) based on collected specimens examined in this study. Closed symbols indicate type localities.

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FIGURE 12. Preserved specimens of Brachypterois serrulata. (A) NCIP 5002 (1 of 3 specimens), 43.2 mm SL; (B) KAUM–I.
44571, 78.6 mm SL; (C) ASIZP 62399, 92.8 mm SL, neotype of Sebastes serrulatus.
Diagnosis. A species of Brachypterois with the following combination of characters: pectoral-fin rays 14–16
(modally 15); scale rows in longitudinal series 44–51 (47); scales above lateral line 4–6 (5); scale rows between
last dorsal spine base and lateral line 4–7 (6); pre-dorsal scale rows 4–8 (5); suborbital scale rows 0–4 (2) (Fig. 6B);
total gill rakers 15–19 (17); head length 38.3–43.1 (mean 40.2) %SL (Fig. 7); orbit diameter 11.1–14.5 (12.4) %SL;
upper-jaw length 18.6–21.5 (20.2) %SL; longest dorsal-fin soft ray length 18.0–23.0 (20.2) %SL; longest anal-fin
soft ray length 21.0–27.0 (23.7) %SL; caudal-fin length 30.4–44.7 (35.7) %SL; posterior lacrimal spine usually
directed posteriorly, not curved upward distally (Fig. 8D–F); spine(s) on corner of outer angular ridge not
developed, size similar to other spines on ridge, directed ventrally (Fig. 9D–F); spines usually absent on median
lateral ridge of maxilla, rarely 1–3 spines present in large adults (>80 mm SL) (Fig. 14A); large number of small
spots on caudal fin, 5–26 (mean 13.3) spots on longest caudal-fin ray (Figs. 10, 12–13).

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REVISION OF BRACHYPTEROIS
FIGURE 13. Fresh specimens of Brachypterois serrulata. (A) NCIP 5002 (1 of 3 specimens), 55.9 mm SL (photo: Fahmi); (B)
FRLM 31479, 57.6 mm SL (photo: S. Kimura); (C) ASIZP 62992, 60.5 mm SL (photo: Y.-C. Liao); (D) KAUM–I. 6876, male,
66.9 mm SL; (E) KAUM–I. 23882, male, 74.0 mm SL; (F) KAUM–I. 24053, female, 76.2 mm SL; (G) BSKU 42455, male,
84.0 mm SL (photo: BSKU); (H) KAUM–I. 47468, 94.5 mm SL.
Color of fresh specimens. Head and body pinkish to reddish-brown, darker dorsally, paler ventrally; a poorly
defined large black blotch (subequal to orbit diameter) on opercle. Five dark red to black bands on head;
anteriormost band from anterior margin of eye running over snout to tip of lower jaw; second extending from
anteroventral margin of eye onto lacrimal and both jaws; third from just below eye to posterodorsal corner of
maxilla; fourth from posteroventral margin of eye to posterior portion of opercle; fifth from posterior margin of eye
to opercle, more or less continuous with blotch on opercle. Five poorly defined dark or black vertical bands on
body; anteriormost band extending from between first and third dorsal-fin spine bases; second from between fourth

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and sixth spine bases; third from between eighth and tenth spine bases; fourth Y-shaped, upper margin of anterior
branch below last spine and third soft ray bases, upper margin of posterior branch below sixth and penultimate soft
ray bases, both branches connected at level of lateral line or just below dorsal-fin base; fifth on caudal peduncle
just in front of caudal-fin base. Ground color of fins similar to that of head and body. Dorsal fin with a large dark
red to black blotch on spinous portion between approximately sixth and ninth spines; about 4–5 more or less dotted
dark red lines on soft-rayed portion. Anal fin with about 6–8 somewhat dotted dark red lines. Pectoral fin darker
distally, with about 6–10 oblique narrow dark red bands. Pelvic fin with about 6–7 narrow dark red to black bands.
Caudal fin membrane translucent with dark red band marginally (relatively wider in small specimens); about 60–
110 small dark red spots [maximum diameter 1.3–3.0 (mean 2.4) times in caudal-fin ray width at position where
spot present on ray; based on 13 preserved specimens] on rays. Eye vivid yellow; iris dark blue.
FIGURE 14. Relationships of number of spines on median lateral ridge of maxilla to standard length in (A) Brachypterois
serrulata and (B) B. serrulifer. H holotype of B. serrulifer. Note: all specimens examined, including type series, of B.
curvispina sp. nov. and neotype of Sebastes serrulatus lacked spines.
Color of preserved specimens. Head and body creamy-white to brown, darker dorsally, paler ventrally;
formerly reddish markings brownish and indistinct. Eye dark blue to gray; iris white to dark blue.
Distribution. Brachypterois serrulata is distributed in the northwestern Pacific, from the South China Sea
north to southern Japan, including the Natuna Islands, Borneo, the Gulf of Thailand, Ng Tran Bay, Vietnam and

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REVISION OF BRACHYPTEROIS
Taiwan. The northernmost records from southern Japan are from Tosa Bay, Shikoku Island (Fig. 11). The 5
specimens were collected at depths of 40–100 m (most from 100 m).
Remarks. Meristics and morphometrics of specimens of B. serrulata examined are shown in Tables 1, 3.
Brachypterois serrulata was originally described by Richardson (1846) as Sebastes serrulatus on the basis of a
single specimen (type locality stated as “Sea of China”) which is apparently lost (see Introduction). This species is
regarded herein as the valid name for this common species in the northwestern Asian waters based on our specimen
examinations in museum collections and field surveys in Japan, Taiwan, Thailand and Malaysia. The original
description by Richardson (1846) did not include distinct diagnostic characters, although the detailed description of
head spines and ridges were given. However, he did not mention any spine on the median lateral ridge of the
maxilla which might suggest an absence of spines on the ridge in the type specimen of S. serrulatus, meeting the
condition of B. serrulata recognized herein. According to our examination, specimens of B. serrulifer with similar
size (4 inches length) to the holotype of S. serrulatus have 0–12 (mean 3.1) spines on the median lateral ridge on
the maxilla [33 specimens examined of ca. 10 cm total length (65.4–75.2 mm SL)].
TABLE 3. Meristics and morphometrics, expressed as percentages of standard length, of Brachypterois serrulata.
Neotype
ASIZP 62399
Non-types from
Malaysia and north
of Vietnam
(n = 82
a
)
Non-types from
Thailand and
Indonesia
(n = 32)
SL (mm) 92.8 48.0–99.3 43.2–99.2
Meristics Modes
Dorsal-fin rays XIII, 11 XII–XIV, 9–11 XII, 9–11 XIII, 10
Anal-fin rays III, 5 III, 4–6 III, 5–6 III, 5
Pectoral-fin rays
b
ii + 6 + vii = 15/15 i–vi + 5–8 + v–viii =
14–16/ 14–16
i–iii + 6–8 + v–vii
= 14–16 / 14–16
ii + 7 + vi = 15/15
Scale rows in longitudinal series 49 44–51 44–49 47
Pored lateral-line scales — 24–26 24–25 25
Scales above lateral line 5 4–6 5–6 5
Scales below lateral line 11 9–12 9–11 10
Scale rows between 6th DS and LL 6 5–6 5–6 5
Scale rows between last DS and LL 6 5–7 4–6 6
Pre-dorsal scale rows 5 4–6 4–8 5
Suborbital scale rows 2 0–3 0–4 2
Gill rakers
c
5 + 12 = 17 5–6 + 11–13 = 16–
19
4–6 + 11–13 =
15–18
5 + 12 = 17
Morphometrics (% SL) Means
Body depth 34.4 33.2–39.9 31.8–38.8 36.3
Body depth at anal-fin origin 29.5 27.4–32.3 27.5–33.6 30.0
Body width 21.4 19.1–26.5 18.9–25.2 22.8
Head length 38.4 38.3–41.9 38.7–43.1 40.2
Head width 14.0 13.8–15.9 13.7–15.4 14.8
Snout length 7.8 7.0–9.0 7.3–9.2 8.1
Orbit diameter 11.7 11.4–13.6 11.1–14.5 12.4
Interorbital width
d
5.9 4.6–6.7 4.6–6.6 5.7
Interorbital width
e
5.4 4.1–5.9 3.9–5.7 5.0
Upper jaw length 19.2 19.1–21.5 18.6–21.5 20.2
Maxillary depth 7.4 6.4–8.2 6.3–8.3 7.4
......continued on the next page

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a
Morphometric data from 78 specimens.
b
Upper unbranched ray + branched ray + lower unbranched rays = total rays ton left side of body/ total rays on right side.
c
Rakers on upper limb + rakers on lower limb = total rakers.
d
At vertical midline of eye.
e
At posterior end of preocular spine base.
DS and LL indicate dorsal-fin spine base and lateral line, respectively.
Because the similarity of S. serrulatus to its congeners and the confusion in the literature between the identities
of S. serrulatus and B. serrulifer, a neotype for S. serrulatus is necessary to represent the species. A specimen
(ASIZP 62399, 92.8 mm SL; Fig. 12C) is therefore proposed as the neotype for S. serrulatus. Accordingly,
southern Taiwan becomes the type locality of S. serrulatus under the Article 76.3 of the International Code on
Zoological Nomenclature (ICZN, 1999).
TABLE 3. (Continued)
Neotype
ASIZP 62399
Non-types from
Malaysia and north
of Vietnam
(n = 82
a
)
Non-types from
Thailand and
Indonesia
(n = 32)
Suborbital depth 0.9 0.4–2.1 0.3–2.3 1.1
Postorbital length 20.0 18.8–21.6 19.0–21.7 20.3
Predorsal-fin length 30.3 29.9–35.6 30.6–34.4 32.1
Preanal-fin length 72.2 67.7–76.4 69.8–77.1 72.6
Prepelvic-fin length 35.1 35.3–40.0 36.4–39.7 37.6
1st dorsal-fin spine length 9.4 9.8–13.4 9.1–12.7 11.2
2nd dorsal-fin spine length 15.6 14.2–19.5 14.5–20.6 16.6
3rd dorsal-fin spine length 15.5 14.9–20.9 16.0–20.9 17.6
4th dorsal-fin spine length — 15.0–21.5 16.3–20.3 17.8
5th dorsal-fin spine length — 14.4–20.6 16.7–21.9 17.9
6th dorsal-fin spine length — 14.7–20.2 15.1–19.7 17.8
7th dorsal-fin spine length — 14.5–19.0 14.2–20.0 17.3
8th dorsal-fin spine length 14.3 14.3–19.4 15.0–18.7 16.5
9th dorsal-fin spine length — 12.3–16.8 13.4–15.8 14.3
10th dorsal-fin spine length 8.6 8.9–14.0 8.4–12.5 10.9
11th dorsal-fin spine length 7.8 4.0–9.8 4.1–8.9 7.1
12th dorsal-fin spine length 2.8 2.6–5.4 3.4–5.4 3.9
13th dorsal-fin spine length — 7.6–14.2 9.2–13.4 10.9
1st dorsal-fin soft ray length — 15.4–21.2 16.2–19.7 17.7
Longest dorsal-fin soft ray length 18.0 18.1–21.4 19.9–23.0 20.2
1st anal-fin spine length — 5.2–8.0 5.9–8.9 6.7
2nd anal-fin spine length — 9.6–13.7 10.4–14.8 11.9
3rd anal-fin spine length 14.3 12.9–17.4 14.1–18.1 15.4
1st anal-fin soft ray length 20.9 20.1–26.5 22.2–27.0 23.0
Longest anal-fin soft ray length 21.4 21.0–25.9 23.7–27.0 23.7
Pectoral-fin ray length 44.4 44.0–56.7 49.6–62.8 51.6
Pelvic-fin spine length 15.3 13.5–19.2 14.9–19.3 16.4
Longest pelvic-fin soft ray length 27.2 25.7–32.4 28.7–32.0 29.5
Caudal-fin length 32.0 30.4–38.7 34.9–44.7 35.7
Caudal-peduncle length 21.2 17.5–21.2 17.5–21.4 19.5
Caudal-peduncle depth 11.0 10.0–11.7 10.3–11.9 10.9

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Although Kanayama & Amaoka (1981) first recorded B. serrulata from Japan based on four specimens
collected from Tosa Bay, Kochi, the figured specimen (HUMZ 36733) was identified here as B. serrulifer. Another
specimen (HUMZ 35083) examined by them, not examined in this study, was examined and dissected by Ishida
(1994) during a phylogenetic study. The remaining two specimens (BSKU 2270 and BSKU 7262) are apparently
missing (H. Endo, 2012, pers. comm.).
Rainboth et al. (2012) reported B. serrulata from the Gulf of Thailand but provided a figure (pl. 55, fig. 1136)
taken from Gloerfelt-Tarp & Kailola (1984: unnumbered fig. on p. 108). The photographed specimen (NTM S.
11016-003), collected from Indonesia, was identified as B. serrulifer by us. On the other hand, a figure identified as
Dendrochirus bellus (Jordan & Hubbs 1925) (Scorpaenidae) in Rainboth et al. (2012: pl. 55, fig. 1137) is clearly a
B. serrulata.
Biological notes. Among specimens with sex determined, an individual of 60.2 mm SL (KAUM–I. 39283),
collected from off southern Taiwan in July 2011, was the smallest female examined in the present study that
possessed ripe eggs approaching terminal stage (about 0.4 mm in diameter). Two specimens (BSKU 52359, 73.2
mm SL; BSKU 59821, 70.7 mm SL) from Tosa Bay, Kochi, Japan and three specimens (KAUM–I. 23436, 62.5
mm SL; KAUM–I. 23883, 71.9 mm SL; KAUM–I. 23884, 86.3 mm SL) from the Gulf of Thailand, all five
specimens with developed gonads were collected between July and September, inclusive (Fig. 15A). Two males
(KAUM–I. 23882, 74.0 mm SL, Gulf of Thailand; KAUM–I. 12313, 77.9 mm SL, off Borneo) collected in
September had developed testes at the late spermatogenic stage (Fig. 15B). These data appear to indicate a
breeding season of B. serrulata between July and September. Stomach content analyses (four specimens only)
indicated that B. serrulata feeds on small shrimps and crabs.
FIGURE 15. Photographs of sections from gonads of Brachypterois serrulata. (A) ovary from KAUM–I. 24053, 76.2 mm SL;
(B) testis from KAUM–I. 12313, 77.9 mm SL.
Geographical variation. Some measurement variations were noted between specimens from Indonesia
(Natuna Islands) and Thailand (Gulf of Thailand), and those from other localities, including Malaysia (Borneo) and
north of Vietnam (see list of specimens examined). The former tended to have slightly greater proportions for head
length [38.7–43.1 (mean 40.7)% SL in Indonesia and Thailand specimens vs. 38.3–41.9 (40.0)% SL in other
specimens], and the greatest length of the longest dorsal-fin soft ray [19.9–23.0 (21.1)% SL vs. 18.1–21.4 (19.8)%
SL], longest anal-fin soft ray [23.7–27.0 (24.9)% SL vs. 21.0–25.9 (23.3)% SL], longest pectoral-fin ray [49.6–
62.8 (55.0)% SL vs. 44.0–56.7 (50.2)% SL], pelvic-fin spine [14.9–19.3 (17.3)% SL vs. 13.5–19.2 (16.0)% SL],
longest pelvic-fin soft ray [28.7–32.0 (30.5)% SL vs. 25.7–32.4 (29.1)% SL] and longest caudal-fin ray [34.9–44.7
(37.7)% SL vs. 30.4–38.7 (34.9)% SL] (Table 3). However no other significant morphological differences were
found between the two groups. In addition, all of the above characters change significantly with growth in

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Brachypterois (see under Morphological changes with growth). Such minor morphometric differences were
therefore regarded here as representing geographic variations of B. serrulata.
Brachypterois serrulifer Fowler 1938
[New English name: Sawmaxilla Scorpionfish]
[New Standard Japanese name: Yokoba-nokogiri-kasago]
(Figures 1, 6C, 7, 8G–I, 9G–I, 10–11, 14B, 16–17, 19, 20C; Tables 1, 4)
Sebastes serrulatus (not of Richardson): Alcock 1889: 298, pl. 22, fig. 2 (off Ganjam, Bay of Bengal, India; description);
Alcock 1896: 312 (off Ganjam, Bay of Bengal, India; based on Alcock 1889)
Brachypterois serrulifer Fowler 1938: 79, fig. 35 [type locality: 8.4 miles (ca. 13.5 km) off San Fernando Point Light, west
coast of Luzon Island, Philippines (16°39′36″N, 120°11′06″E), 45 fathoms (ca. 82.3 m) depth]; Herre 1945: 401 (Bay of
Bengal, India; comments on distribution in Bay of Bengal; based on 17 specimens, 7 of which examined herein); Herre
1952: 432 (Manila Bay and Ragay Gulf, Luzon, Philippines and Bay of Bengal, India; description; based on 14 and 15
specimens from Philippines and India, respectively, 14 and 7 of which examined herein); Herre 1953: 563 (Philippines and
Bay of Bengal, India; listed; based on Herre 1945, 1952); Aprieto & Villoso 1977: 43 (Visayan Sea, Philippines; listed);
Fourmanoir 1981: 97 (south off Lubang Island, Philippines; listed); Fourmanoir 1985: 52 (Philippines; listed; based on
Fourmanoir 1981)
Ranipterois serrulifer: Whitley 1950: 407 (new combination; see Remarks under Brachypterois)
Brachypterois serrulatus (not of Richardson): Kotthaus 1979: 25, fig. 476 (off Bombay, India; short description; 3 specimens
examined herein); Gloerfelt-Tarp & Kailola 1984: 108, unnumbered fig. on top left (Indonesia; short description; 2
specimens examined herein); Blaber et al. 1994: 392 (Gulf of Carpentaria, Australia; listed); Sujatha 1995: 57 (Bay of
Bengal, India; listed); Martin et al. 1995: 917 (Gulf of Carpentaria, Australia; listed); Manilo & Bogorodsky 2003: S102
(Arabian Sea; listed)
Brachypterois serrulata (not of Richardson): Dor 1984: 85 (Red Sea; listed); Russell & Houston 1989: 81 (Arafura Sea; listed);
Randall 1995: 106, fig. 227 (Gulf of Oman; short description); Krupp et al. 2000: 325, fig. 1 (off Iran, Arabian Gulf; short
description; 4 specimens examined herein); Allen et al. 2003: 372, unnumbered fig. on top left [in part; Indo-West Pacific;
diagnosis and ecological note; underwater photograph from North Sulawesi (G. Allen, pers. comm.)]; Allen et al. 2006:
877 (Timor and Arafura seas, Australia; review of synonymy and distribution in Australia); Kuiter & Debelius 2006: 187,
unnumbered fig. on top left (in part; Indo-West Pacific; listed); Bijukumar & Deepthi 2009: 149 (off Kerala, India; listed);
Al-Jufaili et al. 2010: 21 (Oman; listed); Golani & Bogorodsky 2010: 22 (Red Sea; listed); Allen & Erdmann 2012: 217,
unnumbered fig. on top (Indonesia; short description and ecological note); Rainboth et al. 2012: 80, pl. 55, fig. 1136 (Gulf
of Thailand; listed; figure taken from Gloerfelt-Tarp & Kailola 1984)
? Brachypterois serrulifer: Fourmanoir & Nhu-Nhung 1965: 94 (Nha Trang Bay, Viet Nam; short description)
Holotype. USNM 98886, 78.7 mm SL, 8.4 miles (ca. 13.5 km) off San Fernando Point Light, west coast of Luzon,
Philippines (16°39′36″N, 120°11′06″E), 45 fathoms (ca. 82.3 m), R/V Albatross, 10 May 1909.
Non-type specimens. 109 specimens, 40.4–94.5 mm SL—JAPAN: HUMZ 36733 (X), 82.3 mm SL, Tosa
Bay, Kochi (obtained at Mimase Port), K. Amaoka, 1956. TAIWAN: KAUM–I. 39187 (X), female, 61.3 mm SL,
off Kaohsiung, H.-C. Ho, trawl, 27 June 2011; NMMB-P 2808 (X), 70.9 mm SL, Tong Gang, Pingtung, 100 m, J.-
H. Wu, 8 Nov. 2001; NMMB-P 12647 (3 specimens; X), 61.2–85.1 mm SL, Tong Gang, Pingtung, M.-C. Yu, 16
Feb. 1960. PHILIPPINES: CAS 29406 (4), 59.2–64.8 mm SL, Nasipit, Agusan del Norte, Mindanao, F. B.
Steiner, 15 Apr. 1973; CAS 49547 (14), 51.0–79.3 mm SL, Manila Bay, Luzon, A. W. Herre, 25 May 1948; NSMT-
P 109949, 74.7 mm SL, E. Fujii; USNM 168172, 40.4 mm SL, Manila Bay to Lingayen Gulf, west coast of Luzon,
1–4 m, R/V Albatross, 11 May 1909. MALAYSIA: KAUM–I. 22033 (X), male, 83.0 mm SL, KAUM–I. 22035
(X), female, 63.9 mm SL, off Kota Kinabalu, Sabah, Borneo, G. Ogihara, 11 Aug. 2009; KAUM–I. 49286, 76.0
mm SL, off Kota Kinabalu, Sabah, Borneo, H. Nishiyama, 2 Aug. 2012. THAILAND (Andaman Sea coast):
KAUM–I. 47580, 73.3 mm SL, KAUM–I. 47581, 64.1 mm SL, off Ranong Province (obtained at Kuraburi Fishing
Port), T. Yoshida & H. Nishiyama, 27 June 2012. INDONESIA: NTM S. 11016-003 (2), 64.0–68.7 mm SL,
Mentawai Strait, Sumatra (0°04′S, 99°01′E), 50 m, T. Gloerfelt-Tarp, 6 May 1983. AUSTRALIA: AMS I. 21842-
009 (2; X), 68.3–85.6 mm SL, Arafura Sea (10°35′S, 133°48′E), R/V Soela; AMS I. 26288-001, 79.8 mm SL,
south east corner of Gulf of Carpentaria, 64 m, R/V Laakanuki, bottom trawl, 29 Apr. 1964; CSIRO A2507 (X),
80.6 mm SL, Arafura Sea (13°33′S, 136°57′E), 34.2 m, 4 Sept. 1963; NTM S. 10031-075 (4), 63.1–84.9 mm SL,
north of Smith Point, Cobourg Peninsula (12°58′S, 132°10′E), 27 m, H. Larson, 18 Oct. 1981; NTM S. 11656-001,
92.1 mm SL, north west of Cape Wessel (10°25′S, 136°55′E), 64 m, R. Williams, 1 Feb. 1985; NTM S. 11659-006

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REVISION OF BRACHYPTEROIS
(2), 84.6–93.9 mm SL, northern Cape Wessel (10°25′S, 136°40′E), 60 m, R. Williams; NTM S. 11895-010, 88.7
mm SL, northern Cape Wessel (10°15′S, 136°40′E), 55–60 m, M. Baron, 20 Feb. 1986; NTM S. 11896-004 (X),
92.0 mm SL, northern Cape Wessel (10°14′S, 136°34′E), 50–55 m, NT Fisheries, 20 Feb. 1986; NTM S. 12262-
001, 53.2 mm SL, north east of Goulburn Islands (10°20′S, 134°29′E), 59 m, NT Fisheries, 11 June 1987; NTM S.
12474-003, 89.2 mm SL, north of Groote Eylandt (13°16′S, 136°51′E), 37 m, R. Williams, 24 Aug. 1987; NTM S.
13038-001 (2), 70.6–77.0 mm SL, south east of Cape Grey, Gulf of Carpentaria (13°05′S, 136°45′E), 39 m, NT
Fisheries, 23 Aug. 1987; NTM S. 13253-001 (X), 84.6 mm SL, northern Cape Wessel (10°14′S, 136°53′E), 53 m,
R. Williams, 21 Nov. 1991; NTM S. 13260-002 (2), 55.4–78.8 mm SL, Gulf of Carpentaria (13°02′S, 139°22′E),
58 m, R. Williams, 25 Nov. 1991; NTM S. 13270-002 (3), 54.3–75.2 mm SL, Gulf of Carpentaria (11°04′S,
139°56′E), 57 m, R. Williams, 29 Nov. 1991. MYANMAR: CAS 64830 (4; 1 female, 61.2 mm SL), 52.7–78.0 mm
SL, Bay of Bengal (15°08′N, 94°54′E), 35 m, R/V Anton Bruun, shrimp trawl, 1 Apr. 1963; CAS 211494 (4; 1
female, 57.3 mm SL), 54.5–70.1 mm SL, Bay of Bengal (15°08′N, 94°54′E), R/V Anton Bruun, 1 Apr. 1963.
INDIA: CAS 207661 (5; 1 female, 60.8 mm SL), 60.8–66.6 mm SL, Neendakara, Kerala (8°56′N, 76°30′E),
bottom trawl, 8 Oct. 1966; CAS 29590, 92.2 mm SL, 10 miles off Bombay, 5–10 m, F. B. Steiner, trawl, 4–10 Nov.
1973; SU 14649 (7; 1 female, 55.0 mm SL), 51.8–77.7 mm SL, mouth of Hugli River, Bay of Bengal, A. Herre, R/
V Lady Fraser; ZMH 5602 (3), 77.9–66.5 mm SL, approximately 200 m off Mumbai, 70–84 m, H. Kotthaus, R/V
Meteor, bottom trawl, 27 Feb. 1965. PERSIAN GULF: SMF 26000, 45.9 mm SL, 28°56.494′N, 49°43.812′E, 45
m, H. Zetzsche, bottom trawl, 14 Dec. 1991; SMF 26001 (3), 56.1–73.7 mm SL, 29°13.587′N, 49°53.955′E, 41 m,
H. Zetzsche, bottom trawl, 10 Nov. 1991. OMAN: BPBM 21416, male, 76.0 mm SL, 5.6 km north of Khalil, Gulf
of Oman (23°38′N, 58°21.7′E), 27.4 m, G. R. Allen, 10 Mar. 1977; CAS 60396 (19; 3 females, 65.9–67.9 mm SL),
58.7–86.4 mm SL, Gulf of Oman (26°10–13′N, 57°02′E), 64–65 m, R/V Anton Bruun, trawl, 30 Nov. 1963; WAM
P. 25996.001, 72.5 mm SL, 10 km north of Khalil, Gulf of Oman (23°46′N, 58°20′E), 30–66 m, G. R. Allen, 10
Mar. 1977. ERITREA: CAS 209665 (2), 76.9–88.0 mm SL, Massawa Bay (15°40′N, 39°36′E), 79 m, L. W.
Knapp, otter trawl, 20 Sept. 1971; CAS 64825, 63.6 mm SL, Massawa Bay (15°40′N, 40°23′E), 79 m, L. W.
Knapp, otter trawl, 21 Sept. 1971. MADAGASCAR: MNHN 2004-0028, 94.5 mm SL, 23°36′0″S, 43°31′59″E,
340–360 m, 27 Feb. 1973.
Diagnosis. A species of Brachypterois with the following combination of characters: pectoral-fin rays 15–17
(modally 16); scale rows in longitudinal series 43–51 (47); scales above lateral line 4–7 (5); scale rows between
last dorsal spine base and lateral line 4–7 (5); pre-dorsal scale rows 3–6 (4); suborbital scale rows 0–2 (1) (Fig. 6C);
total gill rakers 15–17 (16); head length 38.8–44.6 (mean 41.7) %SL (Fig. 7); orbit diameter 11.8–15.1 (13.0) %SL;
upper-jaw length 19.1–22.2 (20.7) %SL; longest dorsal-fin soft ray length 20.5–26.2 (23.4) %SL; longest anal-fin
soft ray length 23.3–29.0 (26.2) %SL; caudal-fin length 36.1–45.1 (40.1) %SL; posterior lacrimal spine usually
directed ventrally (Fig. 8G–I); spine(s) on corner of outer angular ridge not especially developed, size similar to
that of other spines on ridge, directed ventrally (Fig. 9G–I); 0–22 (mean 3.9) spines on median lateral ridge on
maxilla in specimens >58 mm SL (Fig. 14B); relatively few large spots on caudal fin, 2–10 (mean 5.2) spots on
longest caudal-fin ray (Figs. 10, 16–17).
Color of fresh specimens. Head and body pinkish-brown, darker dorsally, paler ventrally; a poorly defined
large black blotch (subequal to orbit diameter) on opercle. Five reddish-brown to brown bands on head;
anteriormost band from anterior margin of eye running over snout to tip of lower jaw; second extending from
anteroventral margin of eye onto lacrimal and both jaws; third from just below eye to posterodorsal corner of
maxilla; fourth from posteroventral margin of eye to posterior portion of opercle; fifth from posterior margin of eye
to opercle, more or less continuous with blotch on opercle. Five poorly defined reddish-brown to brown vertical
bands on body; anterior-most band extending from between first and third dorsal-fin spine bases; second from
between fourth and sixth spine bases; third from between eighth and penultimate spine bases; fourth Y-shaped,
upper margin of anterior branch below last spine and fourth soft ray bases, upper margin of posterior branch below
seventh and penultimate soft ray bases, both branches connected at level of lateral line or just below dorsal-fin
base; fifth on caudal peduncle just in front of caudal-fin base. Ground color of fins similar to that of head and body.
Dorsal fin with a large dark red to brown blotch on spinous portion between about sixth and ninth spines; about 4–
5 more or less dotted dark red lines on soft-rayed portion. Anal fin with about 5 more or less dotted dark red lines.
Pectoral fin membrane dark brown, with about 6–8 poorly defined narrow dark red bands. Pelvic fin darker
marginally, with about 5–8 poorly defined narrow dark red bands. Caudal fin membrane translucent with dark red
narrow band marginally; about 20–60 small dark red to brown spots [maximum diameter 2.1–6.0 (mean 3.4) times

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FIGURE 16. Preserved specimens of Brachypterois serrulifer. (A) USNM 168172, 40.4 mm SL; (B) CAS 64830 (1 of 4
specimens), 62.3 mm SL; (C) CAS 29406 (1 of 4), 64.8 mm SL; (D) USNM 98886, 78.7 mm SL, holotype of B. serrulifer; (E)
NTM S. 10031-075 (1 of 4), 84.9 mm SL.

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REVISION OF BRACHYPTEROIS
FIGURE 17. Fresh specimens of Brachypterois serrulifer. (A) KAUM–I. 39187, female, 61.3 mm SL; (B) KAUM–I. 22033,
male, 83.0 mm SL; (C) KAUM–I. 47580, 73.3 mm SL; (D) size unknown, off Sonmiani, Pakistan, Oct. 2009 (photo: H. B.
Osmany).

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430 · Zootaxa 3693 (4) © 2013 Magnolia Press
in caudal-fin ray width at position where spot present on ray; based on 39 preserved specimens] on rays. Eye vivid
yellow; iris dark blue. Color photographs of live individuals from Indonesia available in Allen et al. (2003), Kuiter
& Debelius (2006), and Allen & Erdmann (2012) as B. serrulata.
Color of preserved specimens. Head and body creamy-white to brown, darker dorsally, paler ventrally;
formerly reddish markings brownish and indistinct. Eye dark blue to gray; iris white to dark blue.
Distribution. Brachypterois serrulifer is widely distributed in the Indo-west Pacific region, including
Madagascar, Red Sea and Persian Gulf, east to the Timor Sea and the Philippines, South and East China seas, and
north to Tosa Bay, Kochi, southern Japan (Fig. 11). The specimens of the 18 lots examined were from depths of 1–
360 m (mostly deeper than 50 m). According to underwater photographs of B. serrulifer (as B. serrulata), included
in Allen et al. (2003) and Allen & Erdmann (2012), the species also occurs off Bali and North Sulawesi, Indonesia.
Brachypterois serrulifer and B. serrulata occur sympatrically off Borneo, southern Taiwan and Tosa Bay, southern
Japan. However, the occurrence of B. serrulifer in Japan is rare, with only a single specimen recorded (vs. 51
specimens of B. serrulata recorded).
Brachypterois serrulifer is apparently not distributed off the northwestern coast of Western Australia, despite
occurring off the northern coast. Allen (1997) and Allen et al. (2006) recorded B. serrulata from northwestern and
northern coasts of Western Australia and northern coast of Northern Territory. However, according to distributional
records of B. serrulifer we examined (Fig. 11), the records of B. serrulata from the northern coasts of both states
are believed to be misidentifications of B. serrulifer. Among the specimens examined only a single putative
example of Brachypterois (NMV A24244, 24.4 mm SL), collected from northwest of Dampier, WA, and identified
and registered as B. serrulata, was collected from Western Australia. However, that specimen is now re-identified
here as Dendrochirus brachypterus (Cuvier in Cuvier & Valenciennes 1829) (Scorpaenidae).
Remarks. Meristics and morphometrics of specimens of B. serrulifer examined are shown in Tables 1, 4. The
species was originally described by Fowler (1938) on the basis of a single specimen (USNM 98886, 78.7 mm SL;
Fig 16D) collected off San Fernando Point Light, west coast of Luzon, Philippines (16°39′36″N, 120°11′06″E) at a
depth of 45 fathoms (ca. 82.3 m). It has been regarded by subsequent researchers (e.g., Chu 1962; Kanayama &
Amaoka 1981; and Allen et al. 2006) as a junior synonym of S. serrulatus Richardson 1846. However, examination
of the holotype of B. serrulifer shows it to have the same diagnostic characters as specimens included herein under
that name (see Comparisons). B
rachypterois serrulifer is therefore regarded as valid. Fourmanoir & Nhu-Nhung
(1965), Fourmanoir (1981, 1985) and several other authors also referred to B. serrulifer as a valid name (see
synonym list), although there is no evidence that they recognized both nominal species, B. serrulata and B.
serrulifer. Therefore, the record of B. serrulifer from Vietnam given by Fourmanoir & Nhu-Nhung (1965) is
doubtful and probably applicable to B. serrulata.
Alcock (1889) first recorded B. serrulifer from the Bay of Bengal (as B. serrulata), on the basis of a single
specimen not examined in this study. However, the detailed description and figure of the specimen given by Alcock
(1889) clearly shows it to be B. serrulifer. Brachypterois sp. reported by Oyugi (2005) from Kilifi, Kenya, but
lacking information for positive identification, is most likely to represent B. serrulifer, as there are no other
congeners distributed in the western Indian Ocean.
Biological notes. A female of 63.9 mm SL (KAUM–I. 22035) collected off Borneo in August 2009 had
developed eggs approaching the terminal stage (about 0.5 mm in diameter). A stomach content analysis (1
specimen) showed that B. serrulifer feeds on small shrimps and fishes.
Geographical variation. A difference in the number of caudal fin spots in B. serrulifer was noted between
specimens collected from the Indian (Figs. 16B, 17B, D) and Pacific oceans, including northern Australia (Figs.
16A, C–E, 17A, C). The former specimens had relatively fewer and larger spots on the caudal fin, compared with
their Pacific Ocean counterparts (Fig. 10) [maximum diameter of a basal spot on a middle caudal-fin ray 2.8–4.0
(mean 3.5), 2.1–6.0 (3.5), 2.6–6.0 (3.6), 2.3–3.8 (2.9) and 2.7–4.7 (3.3) times the caudal-fin ray width at the
position where spot present on ray in specimens from the Persian Gulf (3 specimens), Arabian Sea (13), Bay of
Bengal and Andaman Sea (8), northern Australia (8) and Philippines (7), respectively]. Since no other significant
differences were found between the two groups, the difference is regarded herein as representing geographical
variation in B. serrulifer.

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TABLE 4. Meristics and morphometrics, expressed as percentages of standard length, of Brachypterois serrulifer.
Holotype
USNM 98886
Non-types
(n = 109)
SL (mm) 78.7 40.4–94.5
Meristics Modes
Dorsal-fin rays XIII, 10 XII–XIII, 9–11 XIII, 10
Anal-fin rays III, 5 III, 4–5 III, 5
Pectoral-fin rays
a
ii + 8 + vi = 16/16 i–iii + 6–8 + vi–viii =
15–17/15–17
ii + 7 + vii = 16/16
Scale rows in longitudinal series 48 43–51 47
Pored lateral-line scales 25 23–26 24
Scales above lateral line 6 4–7 5
Scales below lateral line 11 9–12 10
Scale rows between 6th DS and LL 5 4–6 5
Scale rows between last DS and LL 5 4–7 5
Pre-dorsal scale rows 5 3–6 4
Suborbital scale rows 1 0–2 1
Gill rakers
b
5 + 11 = 16 4–6 + 10–12 = 15–18 5 + 11 = 16
Morphometrics (% SL) Means
Body depth 35.5 32.0–40.0 36.6
Body depth at anal-fin origin 30.1 25.8–33.1 29.4
Body width 22.2 18.1–27.4 22.1
Head length 42.7 38.8–44.6 41.7
Head width 14.7 13.7–16.1 14.7
Snout length 8.1 7.2–9.2 8.2
Orbit diameter 13.5 11.8–15.1 13.0
Interorbital width
c
5.0 4.7–6.9 5.8
Interorbital width
d
4.6 4.4–6.5 5.3
Upper-jaw length 21.0 19.1–22.2 20.7
Maxillary depth 7.9 6.1–8.4 7.5
Suborbital depth 1.5 0.3–1.9 0.8
Postorbital length 20.6 19.1–23.1 20.9
Predorsal-fin length 34.3 31.3–36.1 33.6
Preanal-fin length 72.7 68.2–77.5 72.8
Prepelvic-fin length 38.6 35.3–41.3 38.4
1st dorsal-fin spine length — 7.4–13.6 10.3
2nd dorsal-fin spine length — 12.4–24.5 18.2
3rd dorsal-fin spine length — 14.8–26.4 19.4
4th dorsal-fin spine length — 15.1–23.4 18.9
5th dorsal-fin spine length — 14.3–23.5 18.7
6th dorsal-fin spine length — 15.3–22.6 18.7
7th dorsal-fin spine length — 14.5–22.5 18.6
8th dorsal-fin spine length — 15.1–21.7 17.9
9th dorsal-fin spine length — 12.7–18.0 15.4
10th dorsal-fin spine length — 8.8–15.7 11.6
......continued on the next page

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a
Upper unbranched + branched + lower unbranched rays = total rays on left side of body/ total rays on right side.
b
Rakers on upper limb + rakers on lower limb = total rakers.
c
At vertical midline of eye.
d
At posterior end of preocular spine base.
DS and LL indicate dorsal-fin spine base and lateral line, respectively.
Discussion
Sexual dimorphism. Sexual dimorphism has been reported for a number of scorpaenid species. The males of
Parascorpaena mcadamsi (Fowler 1938) and several species of Scorpaenopsis have a black blotch on the posterior
spinous portion of the dorsal fin (vs. lacking in females) (Poss 1999; Randall & Eschmeyer 2001; Motomura et al.
2005c, 2011), and two species of Ebosia, E. bleekeri (Döderlein 1884) and E. falcata Eschmeyer & Rama-Rao
1977 have a distinct sexual difference in the shape of the parietal spine (Eschmeyer & Rama-Rao 1977). Moreover,
sexual and size-related differences in body coloration are known in Sebastes owstoni (Jordan & Thompson 1914)
(Yagishita et al. 2007). In the present study, sexual differences in the number of spines on ridges on the mandible
and the shape of the posterior lacrimal spine were recognized in Brachypterois. An analysis of 27 male (63.9–99.3
mm SL; Fig. 13D, E, G), 20 female (53.2–86.3 mm SL; Fig. 13F) and 51 sex-undetermined specimens (48.0–99.2
mm SL) of B. serrulata indicated that males tend to have a larger number of spines on the mandible ridges (Fig.
18). Specimens less than 63 mm SL, including three females and 10 sex-undetermined specimens, lacked spines on
the mandible (rarely 1–3 spines on the dentary ridges). Of specimens exceeding 65 mm SL, males had a greater
number of spines around the mandibular symphysis (1–10 vs. 0–2 in females), spines on the outer dentary ridge (1–
14 vs. 0–3), spines on the inner dentary ridge (4–22 vs. 0–7), spines on the central dentary ridge (3–25 vs. 0–1),
spines on the outer angular ridge (0–14 vs. 0–3) and spines on the inner angular ridge (4–14 vs. 0–5). In all of the
specimens examined, males had 1–58 (mean 31.9) spines on the dentary ridges, 0–24 (14.3) spines on the angular
ridges and 1–82 (47.0) spines in total, whereas females had 0–21 (4.0), 0–8 (2.1), and 0–29 (5.4) spines on the
dentary, angular and in total, respectively. Furthermore, males tend to have a better developed posterior lacrimal
spine, such being a strongly species-specific condition (Fig. 8D, E). Brachypterois curvispina and B. serrulifer
appeared to have similar sexually dimorphic features, although the numbers of sex-determined specimens of those
species were limited in the present study.
Morphological changes with growth. The numbers of all head spines tend to increase with growth in all
TABLE 4. (Continued)
Holotype
USNM 98886
Non-types
(n = 109)
11th dorsal-fin spine length 7.5 4.0–10.5 7.2
12th dorsal-fin spine length 4.6 2.9–5.9 4.0
13th dorsal-fin spine length 9.8 8.8–14.1 11.5
1st dorsal-fin soft ray length 16.8 14.5–23.8 18.8
Longest dorsal-fin soft ray length 24.4 20.5–26.2 23.4
1st anal-fin spine length — 5.1–8.8 7.1
2nd anal-fin spine length 12.6 10.0–14.9 12.6
3rd anal-fin spine length 17.4 13.4–18.3 16.2
1st anal-fin soft ray length 24.7 22.3–28.0 25.0
Longest anal-fin soft ray length 25.4 23.3–29. 0 26.2
Pectoral-fin ray length 58.7 44.8–65.9 55.8
Pelvic-fin spine length 17.0 14.2–19.4 17.3
Longest pelvic-fin soft ray length 30.6 24.8–34.6 29.4
Caudal-fin length 41.7 36.1–45.1 40.1
Caudal-peduncle length 18.7 15.3–21.1 18.2
Caudal-peduncle depth 11.2 10.4–12.7 11.5

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REVISION OF BRACHYPTEROIS
species of Brachypterois (Figs. 14, 18). For example, specimens of B. serrulata smaller than 68.4 mm SL have 1–8
(mean 3.8) spinous points on the nasal spine forming a single spinous ridge, whereas specimens larger than 69.3
mm SL have 3–15 (7.3) spinous points on the spine sometimes forming two spinous ridges, including a short outer
ridge with 1–5 spinous points. Similar changes of the nasal spine are known in several species of Scorpaenidae and
Neosebastidae [e.g., Parapterois (see Motomura, 2004a), and Maxillicosta meridianus Motomura, Last & Gomon
2006 (see Motomura et al. 2006a)]. The anterior portion of the central dentary ridge between the second and third
sensory pores reduced with growth in B. serrulata in specimens larger than 86 mm SL, a change which appeared to
be more obvious in males rather than females.
FIGURE 18. Relationships of: (A) number of spines on dentary and (B) number of spines on angular to standard length in male
(triangles), female (circles) and sex undetermined (crosses) specimens of Brachypterois serrulata.
Changes in the relative lengths of some body parts with growth, especially fin spines and soft rays, are well
known in Scorpaenidae (e.g., Motomura et al. 2005b, 2005c, 2006b, 2011). Analyses of 42 measurements in B.
curvispina, B. serrulata and B. serrulifer indicated that the relative lengths of the head and orbit, and all fin rays of
the three species tended to decrease with growth (Fig. 19A–C, caudal fin length only). In contrast, suborbital depth
became significantly greater with growth (Fig. 19D). As a consequence of these changes, the head and orbit in each
species appears to become shorter with growth (see Figs. 3–5, 12–13, 16–17).

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434 · Zootaxa 3693 (4) © 2013 Magnolia Press
Some variations in the number of branched pectoral-fin rays in Brachypterois were also noted (Fig. 20), but
were not significantly related to growth, although the number of those fin rays tends to increase with growth in
most members of Scorpaenidae, which is especially well known in Scorpaena (e.g., Motomura et al. 2005b, 2011).
FIGURE 19. Relationships of: (A) head length; (B) orbit diameter; (C) caudal-fin length; and (D) suborbital depth (all as % of
standard length) to standard length in Brachypterois curvispina sp. nov. (stars, solid lines), B. serrulata (circles, dotted lines)
and B. serrulifer (triangles, chain lines).

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REVISION OF BRACHYPTEROIS
FIGURE 20. Relationships of number of branched pectoral-fin rays to standard length in (A) Brachypterois curvispina sp.
nov., (B) B. serrulata and (C) B. serrulifer.
Genetic divergence. The genetic analysis indicated a substantial degree of genetic divergence between B.
serrulata and B. serrulifer. Comparisons on the basis of 526 base pairs of the mtDNA cytochrome b gene showed
that differences ranged from 3.0% to 3.2% between the two species, intraspecific sequence differences ranging
from 0.2% to 0.4%.
Species comparisons. Brachypterois serrulifer differed from B. curvispina and B. serrulata in having a higher
count of pectoral-fin rays [15–17 (modally 16) in B. serrulifer vs. 14–16 (15) in both B. curvispina and B.
serrulata] and fewer spots on the longest caudal-fin ray [2–10 (mean 5.2) vs. 5–13 (9.3) and 5–26 (13.3)] (Fig. 10;
Table 1). Furthermore, B. serrulifer had 0–22 (mean 3.9) spines on the median lateral ridge of the maxilla in
specimens larger than 60 mm SL, whereas B. curvispina and B. serrulata almost always lacked such spines

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436 · Zootaxa 3693 (4) © 2013 Magnolia Press
(rarely1–3 spines present in B. serrulata specimens larger than 80 mm SL) (Figs. 1, 14). Moreover, B. serrulifer
could be distinguished from B. serrulata by the lower counts of scale rows between the last dorsal fin spine base
and lateral line [4–7 (modally 5) in B. serrulifer vs. 4–7 (6) in B. serrulata], pre-dorsal scale rows [3–6 (4) in vs. 4–
8 (5)], suborbital scale rows [0–2 (1) vs. 0–4 (2)] and gill rakers [15–17 (16) vs. 16–19 (17)] (Fig. 6; Table 1).
Brachypterois curvispina had larger scales than B. serrulata and B. serrulifer, and lower counts of scale rows
in longitudinal series [41–45 (modally 44) in B. curvispina vs. 44–51 (47) in B. serrulata and 43–51 (47) in B.
serrulifer], scales above the lateral line [4–5 (4) vs. 4–6 (5) and 4–7 (5)] and scale rows between the last dorsal fin
spine base and lateral line [4–5 (4) vs. 4–7 (6) and 4–7 (5)] (Table 1). In addition, B. curvispina was distinguished
from the other species by the greater length of the head [41.6–44.5 (mean 43.1)% SL in B. curvispina vs. 38.3–43.1
(40.2)% SL in B. serrulata and 38.8–44.6 (41.7)% SL in B. serrulifer], orbit diameter [13.0–15.4 (14.3)% SL vs.
11.1–14.5 (12.4)% SL and 11.8–15.1 (13.0)% SL] and upper jaw [20.2–23.0 (21.6)% SL vs. 18.6–21.5 (20.2)% SL
and 19.1–22.2 (20.7)% SL] (Fig. 7A–C). Among the three species, B. curvispina was unique in having the
posterior lacrimal spine usually directed posteriorly, but with the tip strongly curved upward in large adults (vs.
usually directed posteriorly, but not curved upward in B. serrulata and usually directed ventrally in B. serrulifer)
(Fig. 8), and the (posteriorly directed) spine(s) around the posterior corner of the outer angular ridge longer and
stouter than the adjacent spines on the ridge (vs. ventrally directed and of similar size to adjacent spines in B.
serrulata and B. serrulifer) (Fig. 9).
In addition, B. serrulata differed from B. curvispina and B. serrulifer in having the shortest longest dorsal-fin
soft ray
[18.1–23.0 (mean 20.2)% SL in B. serrulata vs. 19.5–24.5 (22.5)% SL in B. curvispina and 20.5–26.2
(23.4)% SL in B. serrulifer], longest anal-fin soft ray [21.0–27.0 (23.7)% SL vs. 24.0–28.4 (25.3)% SL and 23.3–
29.0 (26.2) %SL] and caudal-fin [30.4–44.7 (35.7)% SL vs. 37.8–46.4 (42.3)% SL and 36.1–45.1 (40.1)% SL]
(Fig. 7D–F).
Acknowledgments
We are especially grateful to M. McGrouther, A. Hay and S. Reader (AMS), K.-T. Shao, Y.-C. Liao and M.-Y. Lee
(ASIZP), J. Maclaine and O. Crimmen (BMNH), H. Endo, T. Yamakawa and N. Nakayama (BSKU), E. Katayama
(NSMT; formerly BSKU), A. Graham (CSIRO), T. Nakabo, Y. Kai, N. Muto and A. Tokairin (FAKU), S. Kimura,
T. Hibino and D. Sasaki (FRLM), M. Yabe, H. Imamura, T. Yamanaka and S. Ohashi (HUMZ), Z. Arifin, R.
Pratiwi, M. Adrim, I. Alhakim and K. Wibowo (Indonesian Institute of Sciences, Jakarta), P. Musikasinthorn and S.
Arbsuwan (Kasetsart University, Bangkok), R. Causse (MNHN), H.-C. Ho (NMMB), M. Gomon and D. Bray
(NMV), K. Matsuura, G. Shinohara, S. Chiba, K. Kuriiwa, M. Nakae and S. Sato (NSMT), I.-S. Chen (NTOU), K.-
Y. Wu (NTUM), K. Hatooka (OMNH), T. Alpermann and H. Zetzsche (SMF), F. Krupp (formerly SMF), T.
Yoshino (formerly URM), J. Williams (USNM), S. Morrison (WAM) and R. Thiel and I. Eidus (ZMH) for their
kind hospitality during the first author’s visits to their institutions. We thank A. Suzumoto (BPBM), D. Catania
(CAS), G. Dally (NTM), K. Hoshino (SNFR) and J. Johnson (QM) for providing opportunities to examine
specimens. We also thank H. Endo, Fahmi, A. Graham and CSIRO Australian National Fish Collection, M. M.
Khan and H. B. Osmany (Marine Fisheries Department, Government of Pakistan), S. Kimura and Y.-C. Liao for
providing photographs of specimens when fresh; J. Johnson, Y.-C. Liao and H. Nishiyama (KAUM) for providing
data of specimens; S. Chiba for instructing the first author in molecular analysis; and G. Hardy (Ngungru, New
Zealand) and an anonymous referee for reading the manuscript and providing helpful comments; T. Hashimoto
(KAUM) and S. Arbsuwan for taking the radiographs; G. Allen (formerly WAM) for providing information on
underwater photographs of B. serrulifer taken in Indonesia; K. Miyamoto (Okinawa Churashima Foundation,
Okinawa) for providing data for URM specimen; E. Katayama and M. Meguro (KAUM) for their assistance during
the first author’s research in USA; G. Ogihara, T. Yoshida and other students and M. Ito, Y. Haraguchi, M.
Takayama and other volunteers of KAUM for their assistance and sampling specimens. This study was supported
in part by Grants-in-Aid for Scientific Research (B, 24370041 and C, 23580259) from the Japan Society for the
Promotion of Science, Tokyo, Japan (JSPS), a Grant-in-Aid for Young Scientists (B, 19770067) from the Ministry
of Education, Science, Sports and Culture, Tokyo, Japan, JSPS Asian Core Program “Establishment of Research
and Education Network on Coastal Marine Science in Southeast Asia”, JSPS International Training Program
“Protect Diversity of Bioresources in the Tropical Area”, and the Coastal Area Capability Enhancement in
Southeast Asia Project of the Research Institute for Humanity and Nature, Kyoto, Japan.

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