Ichthyol Res
DOI 10.1007/s10228-014-0445-4

FULL PAPER

A new species of scorpionfish, Ebosia saya (Scorpaenidae:
Pteroinae), from the western Indian Ocean and notes on freshcoloration of Ebosia falcata
Mizuki Matsunuma • Hiroyuki Motomura

Received: 14 September 2014 / Revised: 12 October 2014 / Accepted: 13 October 2014
Ó The Ichthyological Society of Japan 2014

Abstract A new scorpionfish, Ebosia saya sp. nov.
(Scorpaenidae: Pteroinae), is described on the basis of 10
specimens collected from the Saya de Malha Bank in depthsof 95–126 m. Comparisons with two valid Indo-Pacificspecies, Ebosia bleekeri (Döderlein in Steindachner andDöderlein 1884) (western Pacific Ocean) and Ebosia falcata
Eschmeyer and Rama-Rao 1978 (northern and eastern Indian
Ocean), showed E. saya to be most similar to the latter, withboth species sharing 8 (usually) anal-fin soft rays, 17 (usually) pectoral-fin rays, the elongated parietal spine of males
relatively narrow and strongly falcate posterodorsally, andthe pectoral fin predominantly yellow in males, whereas
E. bleekeri has 7 (usually) anal-fin soft rays, 16 (usually)
pectoral-fin rays, the elongated parietal spine of males relatively broad and weakly falcate, and the pectoral fins entirely
red in males. However, E. saya differs from E. falcata inhaving 5 scale rows above the lateral line [vs. 4 or 5 (modally4) in E. falcata], slightly greater total numbers (5–20, mean
12.3) of preocular, supraocular, and postocular spines at
60–90 mm SL (vs. 4–18, 8.4), shorter postorbital length

This article was registered in the Official Register of Zoological
Nomenclature (ZooBank) as B264ADE4-84F9-42A4-AFB68EA791FF671C.
This article was published as an Online First article on the online
publication date shown on this page. The article should be cited byusing the doi number.
M. Matsunuma (&)
Seikai National Fisheries Research Institute, 1551-8 Taira,Nagasaki 851-2213, Japane-mail: k1139853@kadai.jp
H. Motomura
The Kagoshima University Museum, 1-21-30 Korimoto,Kagoshima 890-0065, Japane-mail: motomura@kaum.kagoshima-u.ac.jp

18.1–19.2 (mean 18.7) % SL [vs. 19.8–23.5 (21.5) % SL],and the blotch above the pectoral-fin base and blotches on the
pectoral-fin membrane all relatively smaller. Fresh coloration of E. falcata is described for the first time, based on
specimens from the Andaman Sea (easternmost record forthe species).
Keywords Ebosia bleekeri  sexual dimorphism 
distribution  Saya de Malha Bank

Introduction
The Indo-West Pacific scorpionfish genus Ebosia Jordan
and Starks 1904 (Scorpaenidae: Pteroinae) is characterizedby having the coronal, parietal, and nuchal spine bases
continuous, and the parietal spine elevated, being a thinbony crest in males (Eschmeyer and Rama-Rao 1978; thisstudy). Two valid species, Ebosia bleekeri (Döderlein in
Steindachner and Döderlein 1884) (western Pacific Ocean)and Ebosia falcata Eschmeyer and Rama-Rao 1978
(northern and eastern Indian Ocean), are currently
recognized.
During a revisionary study of Ebosia, 10 specimens ofan unidentified species from the Saya de Malha Bank,
western Indian Ocean, were found in several museum
collections. The specimens were closely related to E. falcata in having usually 8 anal-fin soft rays, usually 17
pectoral-fin rays, and the elongated parietal spine in males
relatively narrow and strongly falcate posterodorsally.
However, examination subsequently revealed that the Sayade Malha Bank specimens could be distinguished from
E. falcata on the basis of several characters, including
meristics, morphometrics, and coloration, thereby representing an undescribed species of Ebosia. Moreover, newly

123M. Matsunuma, H. Motomura

collected specimens of E. falcata from the Andaman Sea
enabled detailed morphological and genetic comparisonswith its congeners, and photographs of the specimens taken
prior to preservation provided fresh coloration of the species for the first time. Morphological changes with growth,and sexual dimorphism in head spine condition and coloration of the species of Ebosia are herein described indetail.

Materials and methods
Measurements generally followed Motomura (2004b, c),with head width, head depth, maxillary depth, and body
depth at anal-fin origin following Motomura et al. (2005b,
2006a), Motomura (2004a), Motomura et al. (2006b) andMatsunuma et al. (2013), respectively. Counts generally
followed Motomura et al. (2005a, b, c) and Motomura andJohnson (2006); predorsal scale counts followed Motomura
et al. (2006b). The diameter of the blotch above the pectoral-fin base was measured as the least distance between
the upper and lower margins of the blotch. The diameter of
blotches on the pectoral-fin membrane was represented as
the mean of diameter measurements of 3–4 selected blotches on the anterodorsal portion of the pectoral fin (measured as horizontal width of blotch with fin closed). The
last two soft rays of the dorsal and anal fins were counted
as single rays, each pair being associated with a single
pterygiophore. Counts and measurements were made on
the left side wherever possible, except for pectoral-fin rays
that were counted on both sides.
Head spine terminology, shown in Fig. 1, generally follows Randall and Eschmeyer (2002: fig. 1) and Motomura(2004c: fig. 1). Supplemental preopercular spine and laterallacrimal spine terminology follows Eschmeyer (1965) andMotomura and Senou (2008: fig. 2), respectively. Sex wasdetermined by examination of gonad sections embedded inparaffin and stained with Mayer’s haematoxylin–eosin, or
direct examination of gonads by light microscope wheneverpossible. In lists of specimens examined, (G) indicates sex
determined by gonad examination; (E), sex determinedfrom parietal spine morphology.
The mitochondrial DNA (mtDNA) cytochrome b gene
was examined following DNA extraction from two specimens of E. falcata (KAUM–I. 47529, KAUM–I. 47530,
Andaman Sea coast of Thailand) and three specimens ofE. bleekeri (KAUM–I. 35803, KAUM–I. 35817, East
China Sea; KAUM–I. 44331, Kagoshima Prefecture,
Japan), 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 and123

control region of mtDNA, were amplified under the following PCR conditions. Thirty amplification cycles were
performed at 94 °C for 45 sec, 56 °C for 30 sec and 72 °C
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 andNishida 2000), following the manufacturer’s protocols andutilizing the automated genetic analyzer models 3130 and3500 (Applied Biosystems). The obtained DNA sequenceswere subjected to multiple alignments using Clustal X(Thompson et al. 1997). All sequences are available on
DDBJ/EMBL/GenBank accession numbers AB971745–
971749. The sequence difference (p-distance) was calculated using MEGA 6 (Tamura et al. 2013).
Standard length is abbreviated as SL. In the description,
features for the holotype are presented first, followed byparatype data (if different) in parentheses. A distributional
map was prepared using Quantum GIS 2.2 (Quantum GIS
Development Team 2014), with data from Natural Earth.
Institutional abbreviations follow Sabaj Pérez (2014), withthe following addition: NSMT: National Museum of Nature and Science, Tsukuba. Specimens formerly depositedat KSHS (Kochi Prefectural Kochi Nishi Senior HighSchool, Kamobe, Kochi) have been transferred to NSMT
(G. Shinohara, pers. comm.).

Ebosia saya sp. nov.
(New English name: Saya Lionfish) (Figs. 1–2, 3a–c, 4a, 5,8c–f, 10a–c, 12c, 13–16; Tables 1–2)
Holotype. USNM 307910, female (G), 87.0 mm SL, off
Madagascar (10°300 S, 61°100 E), 110–115 m, RV Vityaz,
Cruise 17, Station 2809, 8 January 1989.
Paratypes. Nine specimens, 64.1–89.7 mm SL: HUMZ
73685, female (G), 82.6 mm SL, Saya de Malha Bank
(10°590 S, 61°020 E), 126 m, 5 September 1977; HUMZ
73838, male (G), 78.0 mm SL, HUMZ 73840, male (G),78.2 mm SL, KAUM–I. 70134 (formerly HUMZ 73839),
male (G), 85.2 mm SL, Saya de Malha Bank (10°300 S,
61°320 E), 95 m, 3 September 1977; LACM 44793-3, three
specimens [one male (G) and two females (G)],65.3–75.7 mm SL, collected with holotype; USNM 407273(formerly USNM 307910), two specimens, female (G),63.0–76.6 mm SL, collected with holotype.
Diagnosis. A species of Ebosia distinguished from othermembers of the genus by the following combination ofcharacters: anal-fin soft rays 8; pectoral-fin rays 17–18(modally 17); scale rows above lateral line 5; scale rows
between last dorsal-fin spine base and lateral line 4–5 (5);scale rows between sixth dorsal-fin spine base and lateralline 4–6 (5); scales on preopercular region 11–22 (meanNew Ebosia from western Indian Ocean
Fig. 1 Lateral (top) and dorsal(bottom) views of head ofEbosia saya sp. nov., USNM
307910, holotype, 87.0 mm SL.
Shaded areas indicate skin flaps.
1 Preocular spine; 2 Supraocularspine; 3 Postocular spine; 4Coronal spine; 5 Parietal spine;
6 Nuchal spine; 7 Sphenotic
spine; 8 Pterotic spine; 9 Lower
posttemporal spine; 10Supracleithral spine; 11 Upper
opercular spine; 12 Preopercularspine; 13 Supplemental
preopercular spine; 14
Suborbital ridge/spine; 15
Lateral lacrimal ridge / spine;
16 Anterior lacrimal spine; 17
Posterior lacrimal spine. Bars
indicate 5 mm15.4); postorbital length 18.1–19.2 (18.7) % SL; a blotch
above pectoral-fin base and blotches on pectoral-fin
membrane relatively small; yellowish median fins andyellow pectoral fin in males; elongated parietal spine inmales relatively narrow, strongly falcate posterodorsally,
its tip reaching third to seventh dorsal-fin spine bases.
Description. Morphometrics and selected meristics areshown in Tables 1–2. Meristics given in Table 1 are notrepeated here. Pectoral-fin with 1 (1–3) upper unbranched? 10 (9–11) middle branched ? 6 (5–7) lower unbranched
rays; 0 or 1 branched ray divided into more than three(based on paratypes; unclear in holotype). Pelvic fin with 1

spine and 5 soft rays. Branchiostegal rays 7. Swimbladderpresent.
Body oblong, moderately compressed anteriorly, extensively compressed posteriorly; depth moderate, maximum
body depth slightly less than longest dorsal-fin spine length.
Head large, its length greater than body depth. A short flap
with a median ridge on posterior edge of low membranous
tube associated with anterior nostril; its tip extending
slightly beyond posterior margin of posterior nostril when
depressed posteriorly (based on paratypes; tip of flap broken
in holotype). Supraocular with a small flap, its length about
half of iris diameter (subequal to iris diameter in small

123

M. Matsunuma, H. Motomura
Fig. 2 Fresh specimens of male(a) and female (b) of Ebosia
saya sp. nov. a HUMZ 73840,
78.5 mm SL (parietal spine
broken); b HUMZ 73685,
82.6 mm SL (right side;
reversed), both from Saya de
Malha Bank. Photos: HUMZ

specimens). Two extremely small flaps on preopercle
margin below third preopercular spine base; its tip notreaching posterior margin of interopercle when laid flat. An
extremely small skin flap anterodorsally on orbit surface; its
length less than one-third of posterior nasal pore diameter.
Snout tip with two extremely short barbels; length slightly
shorter than that of orbit surface flap. No other skin flaps onhead or body, including ventral margin of lacrimal.
Relatively well-developed ctenoid scales possessing 3–7
spinules on postorbital and suborbital regions; suborbital
scale area not (or slightly) extending anteriorly beyond
level of mid-orbit; suborbital pit always without scales.
Preopercular region bordered by posterior margin of
maxilla, suborbital ridge and posterior marginal ridge ofpreopercular, covered with relatively well-developed ctenoid scales possessing 3–9 spinules. Opercle covered witha few ctenoid scales possessing 1–6 spinules on upper
portion; remaining portion possibly with cycloid scales(most scales lost in all specimens examined; condition
estimated from the few remaining scales). Anterior regionof occipital area with a few [ca. 7 (7–8)] weakly ctenoid

123

scales; large scales with 3–6 spinules, small scales with asingle spinule or without spinules; spinules mostly
embedded with tip only exposed. Interorbital region,
including frontal surface of preocular, dorsal surface ofsupraocular and postocular, and interorbital canal withoutscales. A small patch of weakly ctenoid scales with 1–3
spinules (all cycloid in small specimens) on suprapostorbital region bordered by sphenotic, pterotic, and posttemporal spine bases, and coronal, parietal, and nuchal ridges.
Other regions of head, including snout, lacrimal, maxilla,mandible, and interopercle without scales. Weakly developed ctenoid scales with 1–7 spinules anterodorsally on
body above lateral line and anterior to level of fifth dorsalfin spine base; ventral surface of chest with a few ctenoid
scales with 3–6 (1–6) spinules; remainder of body withcycloid scales. Basal regions of dorsal- and anal-fin bases
without scales; caudal-fin base with small cycloid scales;
basal scale morphology on pectoral-fin base unknown(scales lost in all specimens examined).
Mouth set relatively low on head, protruding moderately
downward; moderately large, slightly oblique, forming an

New Ebosia from western Indian Ocean

Fig. 3 Preserved specimens of Ebosia saya sp. nov. (male: a–b;
female: c) and E. falcata (male: d–e; female: f) at different stages.
a KAUM–I. 70134, 85.2 mm SL, Saya de Malha Bank; b LACM
44793, 73.5 mm SL (one of three specimens), Saya de Malha Bank;c USNM 307910, holotype of E. saya, 87.0 mm SL, Saya de Malha
Bank; d ZMH 5582, 88.5 mm SL, off India (one of 21 specimens);
e ZMH 5582, 70.9 mm SL, off India (one of 21 specimens); f USNM
218411, holotype of E. falcata, 93.4 mm SL, off Somalia

angle of ca. 20° to horizontal axis of head and body.
Anterior region of maxilla with a poorly developed medianlateral ridge; upper edge of posterior maxilla swollen laterally, forming a low ridge; posterior margin of maxilla justreaching mid-orbit level. Lips thickened; lower lips broadly
extending onto mandible. Lower jaw with a small symphyseal knob. Symphyseal gap separating premaxillaryteeth bands distinctly broader than width of each band (gapwidth approximately twice teeth band width); both jaws
with a relatively narrow band of small, slender conicalteeth; about 5–8 tooth rows at front of upper jaw; about 3–6
tooth rows at front of lower jaw; tooth band narrowing
posteriorly. Eight to ten rows of small conical teeth forming
blunt V-shaped patch on vomer; no palatine teeth. Underside of dentary with three sensory pores on each side; twosmall pores on each side of symphysial knob of lower jaw

on each side. Gill rakers on first gill arch short, tip expanded; longest raker on first gill arch about one-third length of
longest gill filament; a small fourth gill slit present.
Dorsal profile of snout steep, forming an angle of ca. 60°
to horizontal axis of head and body. Nasal bone without
spines. Ascending process of premaxilla intruding slightly
into interorbital space, its posterior margin just reaching
beyond level of posterior margin of posterior nostril. Preocular with 6 (2–6) spinous points along orbit margin.
Supraocular with 3 (1–6) spinous points on outer margin.
Postocular with 7 (2–7) small spines along orbit margin.
Interorbital ridge weakly developed, reduced anteriorly,
lacking spines; diverging anteriorly and posteriorly indorsal view, distance between interorbital ridges narrowest
at vertical midline of eye; interorbital canal moderately
broad and deep, its width about one-third of iris diameter.

123

M. Matsunuma, H. Motomura

Fig. 4 Preserved specimens of Ebosia saya sp. nov. (a) and E. falcata(b), showing black markings on specimens and relatively longsupraocular skin flap. a USNM 407273 (one of two specimens),
63.0 mm SL, Saya de Malha Bank; b ZMH 5582 (one of 21
specimens), 48.8 mm SL, India

Coronal with a single ridge continued to interorbital andparietal rides, with 2 (1–3) minute spinous points. Parietal
with a single ridge; in females, a low elevated ridge withspinous margin, with 3 (3–4) spinous points (based on
holotype and female paratypes); in males, posteriormost
parietal spine elongated as a thin bony crest, curved posterodorsally, its tip reaching level of third to seventh dorsal-fin spine bases (based on male paratypes). Nuchal witha single ridge completely fused to parietal ridge with asingle spinous point on posterior end, directed posterodorsally. Coronal–parietal–nuchal ridges of both sides
running almost parallel in dorsal view (Fig. 1). Tympanic

without spines or ridges. Anterior margin of occipital area
sloped transversely from between origins of coronal ridges,slightly pointed posteriorly in dorsal view; posterior margin of occipital area bordered by net-like sensory canal.
Postorbital spine and exposed sensory canal absent. Sphenotic with 9 (4–8) small spines surrounding a short sensory
canal. Pterotic with a short ridge with 3 (2–5) small spines.
Lower posttemporal with 1 (1 or 2) small spine. Cleithrumwith two low ridges; upper short ridge without spinous
point; lower long ridge curved posteroventrally with 0 (0 or
1) spinous points posteriorly.
Lateral lacrimal ridge with 2 (2–4) small spines; otherridges on lacrimal spinous. Suborbital with a single ridgewith 17 (9–15) spines; 5 (1–18) associated small spines
below ridge. Anterior lacrimal spine with 2 (0–2) spinous
points directed ventrally. Posterior lacrimal spine with 3(2–4) spinous points. Preopercle with 5 spines; upper 3
spines of similar length; lower 2 spines skin covered; nosupplemental spine on first (uppermost) preopercular spinebase; 0 (0–2) supplemental spine on second preopercularspine base; 1 (0–3) supplemental spine on third preopercular spine base. A single exposed upper opercular spine
directed posterolaterally. Lower opercular spine absent.
Doral-, anal- and pelvic-fin spines with deep grooves(most likely associated with venom glands). Origin of firstdorsal-fin spine above pterotic spine base; bases of first andsecond dorsal-fin spines closer than those of subsequentadjacent spines; fifth or sixth spine likely longest (unclear
in all specimens examined); penultimate (twelfth) spine
shortest, its length 0.7 (or 0.6) and 0.8–0.9 (based onparatypes) times that of antepenultimate (eleventh) andposteriormost (thirteenth) spines, respectively; membrane

Fig. 5 Distributional map of Ebosia saya sp. nov. (stars), E. falcata (circles), and E. bleekeri (triangles) based on specimens examined in thisstudy. H indicates holotype

123

New Ebosia from western Indian Ocean
Table 1 Frequency distribution of selected meristics of Ebosia saya sp. nov., E. bleekeri, and E. falcata
Dorsal-fin rays

Anal-fin rays

XIII, 8E. bleekeri

1

XIII, 9

XIII, 10106

3

E. falcata

29H8

E. saya8H

2

XIII, 11

III, 6

III, 7

III, 8

6

100

3

2

34H

1

III, 92

10H

Pectoral-fin rays (one/other sides)

E. bleekeri
E. falcata

15/15

15/16

16/16

16/17

17/17

1

7

91

8

2

1

4

a

E. saya

17/18

18/18

25

4

1H

H

1

1

8
Scale rows in longitudinal series
41

42

43

E. bleekeri
E. falcata

1

3

44

45

46

47

48

49

50

51

1

3

7

15

16

9

2

3

2

1

4

E. saya

3

1

1H

1
Scales above LL
4

E. bleekeri
E. falcata

34

H

Scales below LL

5

6

7

8

9

10

11

12

31

57

2

6

12

22

1

1

2

1

1

2

E. sayab

9H
SR between 6th DS and LL
3

E. bleekeri
E. falcata

1

E. saya

4

5

1

36

30

H

2

1

6H

SR between last DS and LL

4

5

18

85
H

E. falcata

135

E. saya

3

7H

6

4

25

1

Pre-dorsal-fin scales

5

6

2

3

40

9

18

65

H

18

14

1H

1

Upper gill rakers

E. bleekeri

H

10

4

11

2

15

H

3

1

Total gill rakers

11

12

14

15

86

10

2

24

20

10
H

7H

4

Lower gill rakers
6

20

4

3

7H

a

One specimen had 17/19 pectoral-fin raysb

No data for scales below lateral line; all examined specimens lacking scales in this region

Hindicates holotype

1

13
6

H

16

17

69

13

18

5

4H

SR and DS indicate scale rows and dorsal-fin spine, respectivelyof spinous portion of dorsal fin strongly incised. Dorsal-fin
soft rays all branched (based on paratypes); fourth (or fifth)
ray longest, its length distinctly less than that of longest
dorsal-fin spine; posteriormost ray free from caudal
peduncle. Origin of first anal-fin spine below twelfth dorsal-fin spine base; third spine longest; length of first spine
0.7 (or 0.6) and 0.4 (based on paratypes) times those of

second and third spines, respectively. Anal-fin soft rays all
branched (based on paratypes); forth ray longest (based onparatypes), its length subequal to that of longest dorsal-fin
soft ray [longest anal-fin soft ray length 1.0 times that of
dorsal fin (based on paratypes)]; posteriormost ray freefrom caudal peduncle. Pectoral fin long, sixth ray (branched ray) [or eleventh or twelfth ray (unbranched)] longest;

123M. Matsunuma, H. Motomura

its tip just reaching level with anal-fin base posterior end(or reaching penultimate anal-fin soft ray); lower rays
weakly thickened. First pelvic-fin spine base below third
dorsal-fin spine base; all pelvic-fin soft rays branched;usually second soft ray longest, its tip just reaching firstanal-fin spine base (or anus); posteriormost soft ray withmembranous connection to abdomen for approximately
one-fourth of ray length. Caudal fin long, its length 1.2(1.2–1.6) times head length; profile of posterior margin of
fin rounded; usually with 2 procurrent rays, 2 segmented
unbranched rays, and 4 segmented branched rays in dorsal
and ventral series (a single specimen of USNM 407273
with 3 segmented branched rays in dorsal series). Caudal
peduncle relatively short and low, its depth 0.7 (0.6–0.7)
times caudal-peduncle length.
Color of fresh specimens. Based on two paratypes(Fig. 2). Ground color of head and body cream-white topale pink, darker dorsally, whitish ventrally. Three relatively narrow dark red bands (width slightly less than iris
diameter) with narrow brown margin on sides of head;anteriormost band below eye to preopercular region; middle band from posteroventral margin of orbit reaching
obliquely to ventral margin of subopercle; posteriormost
band saddling nape level with central region of parietal
ridge, reaching mid posterior margin of opercle at level ofupper origin of pectoral-fin base. Five moderately broad
pale brown bands (two or three vertical scale rows width)
on sides of body; anteriormost band below first or second
dorsal-fin spine bases, running posteroventrally to first
pored lateral line scale, with an indistinct connection with a
blotch above pectoral-fin base; second band below fourth
to sixth dorsal-fin spine bases, running vertically andapparently reaching abdomen; third and fourth bands
below eight to tenth dorsal-fin spine bases and posteriormost (thirteenth) dorsal-fin spine to second dorsal-fin softray bases, respectively, almost vertical and appearing toreach to abdomen and anal-fin base, respectively; fifth band
below last three dorsal-fin soft ray bases, indistinct anteroventral orientation. A relatively small brown to black
blotch just above pectoral-fin base (slightly smaller thaniris). Eyes yellow, iris black, without radial markings.
Spinous portion of dorsal fin apparently similar to
ground color of body (skin mostly lost); soft-rayed portionof dorsal fin translucent and pale yellow in male [Fig. 2a;
broken in female (Fig. 2b)]. Anal-fin spines translucent;
soft-rayed portion of anal fin translucent and pale yellow inmale (reddish in female). Pectoral fin bright yellow,
somewhat reddish basally in male (mostly broken, but
entirely reddish in remaining portion in female); about 17
small black blotches (smaller than or subequal to iris)
sparsely scattered, apparently forming four or five rows of
blotches; broadly blackish posteriorly in female (noblackish portion in male). Pelvic-fin rays reddish in both123

sexes, tips yellow in male. Caudal fin pale yellow without
markings (broken in female).
Color of preserved specimens. Overall creamy white (orpale orange), slightly darker dorsally (Figs. 3a–c, 4a).
Three relatively narrow pale brown bands on sides of head;anteriormost band below eye; middle band from posteroventral margin of orbit, reaching obliquely to ventral
margin of subopercle; posteriormost band saddling nape atlevel of parietal ridge, reaching opercle at level of upper
origin of pectoral-fin base. Five moderately broad pale
brown bands (two or three vertical scale rows width) on
sides of body (two posteriormost bands obscured in holotype); anteriormost band below first or second dorsal-fin
spine bases, running posteroventrally; second band below
fifth or sixth dorsal-fin spines bases; third band below
eighth or ninth dorsal-fin spine bases; fourth band below
first or second dorsal-fin soft ray bases; fifth band below
sixth or seventh dorsal-fin soft ray bases. A relatively small
dark brown blotch just above pectoral-fin base [slightly (ormuch) smaller than iris]. Eyes darkish blue with silvery
sheen (or entirely dark blue), iris black.
Dorsal and anal fins translucent without markings [ca. 6
small brown spots on dorsal soft rays in 63.0 mm SL specimen (USNM 407273; Fig. 4a)]. Pectoral-fin rays andmembrane translucent [broadly blackish posteriorly in small
specimens, USNM 407273 (one of two), 63.0 mm SL and
LACM 44793-3 (one of three), 65.3 mm SL]; about threeextremely small brown spots (diameter about one-tenth ofiris diameter) scattered on membrane [ca. 16–25 and ca.
6–18 blotches in males and females, respectively; blotch size
larger in small specimens]. Pelvic fin translucent without
markings. Caudal fin translucent without markings [with ca.
5–8 small brown spots scattered on rays in small specimens].
Distribution. Ebosia saya is currently known only fromthe Saya de Malha Bank, western central Indian Ocean, in
depths of 95–126 m (Fig. 5).
Etymology. The specific name saya refers to the type
locality of the species, Saya de Malha Bank. It is treated asa noun in apposition.

Ebosia falcata Eschmeyer and Rama-Rao 1978(English name: Falcate Lionfish) (Figs. 3d–f, 4b, 5, 6, 8b–e, 10d–f, 12b, 13–16; Tables 1–2)
Ebosia falcata Eschmeyer and Rama-Rao 1978: 64,
figs 1D–F, 2, 3 (type locality: off Somalia); Kotthaus 1979:
20, figs 470, 472a, b (in figure legends; as Ebosia sp. in text;
off west coast of India and Somalia; short description);
Eschmeyer 1986: 465 (western Indian Ocean; list); Mandrytsa 2001: 276 (as phylogenetic material; key); Manilo and
Bogorodsky 2003: S102 (western Indian Ocean; list)

New Ebosia from western Indian Ocean
Table 2 Morphometrics of Ebosia saya sp. nov., E. bleekeri, and E. falcata, expressed as percentages of standard length
Ebosia saya sp. nov.

Ebosia falcata

Holotype(USNM 307910)

Paratypes(n = 9)Mean

SL (mm)87.0

63.0–85.2

Body D (% SL)

35.2

34.2–38.3

36.0

Body Da

24.8

23.1–29.0

26.1

Ebosia bleekeri

Holotype(USNM 218411)

Non-types(n = 37)

Mean

Non-types(n = 111)

Mean

93.4

48.8–121.1

39.2

31.0–41.5

35.0

29.4–42.7

37.2

27.2

21.3–28.4

23.6

22.6–33.0

27.7

9.4–159.6

Body W

21.7

20.8–25.1

22.3

24.6

18.6–26.6

21.8

16.9–27.0

22.1

Head L

40.2

39.0–42.4

41.1

41.6

38.1–46.9

43.3

35.6–44.5

40.3

Head W

14.7

14.1–15.2

14.5

14.6

13.5–16.8

14.8

12.7–15.4

14.1

Head D

17.8

15.8–17.3

16.7

16.8

15.2–19.7

17.8

13.9–17.8

15.9

Snout L–

13.2–16.5

14.6

15.3

13.3–15.7

14.3

11.0–17.7

15.5

Orbit diameter

12.1

11.1–12.9

12.1

11.5

9.9–12.9

11.7

9.0–13.6

11.0

Interorbital Wb

8.0

8.3–10.2

9.2

9.6

8.1–10.7

9.5

7.6–11.0

9.3

Interorbital Wc

6.9

7.5–8.9

8.0

7.4

6.5–9.6

8.0

6.9–10.4

8.6

Upper-jaw L

14.7

14.8–16.7

15.6

15.1

14.8–17.4

16.2

14.2–18.0

15.9

Maxillary D

5.3

5.2–5.9

5.5

5.0

5.0–6.0

5.4

4.7–7.1

5.7

Suborbital D

3.6

2.6–3.7

3.2

4.1

2.6–4.7

3.5

0.4–4.6

3.2

Postorbital L

19.2

18.1–19.2

18.7

19.8

19.8–24.2

21.9

14.7–19.2

17.1

Pre-dorsal-fin L

36.6

36.2–38.2

37.3

38.0

34.9–39.6

37.1

33.2–39.5

36.9

Pre-anal-fin L

69.0

65.7–70.7

68.1

70.6

63.1–70.8

67.6

65.0–74.6

69.5

Pre-pelvic-fin L

38.2

35.2–37.1

36.3

35.1

34.8–39.6

37.0

33.0–39.8

36.1

1st DS L

24.4

23.4–27.2

24.7–

20.6–24.7

22.8

19.5–31.3

26.2

2nd DS L

29.8

33.6–35.4

33.4–

24.0–35.4

30.3

23.6–50.1

33.1

3rd DS L–

36.7–39.9

38.3–

31.7–41.7

35.6

27.4–49.4

37.7

4th DS L

35.5

37.1–42.8

38.5–

34.4–43.0

38.5

31.3–48.2

40.1

5th DS L

36.1

35.2–43.7

39.3–

35.6–45.4

39.3

31.7–49.4

40.4

6th DS L

–

–

–

–

32.4–41.5

38.0

31.8–48.7

39.7

7th DS L

36.2

44.3

40.2

–

35.1–41.0

37.3

31.5–50.8

41.0

8th DS L

–

–

–

–

27.2–38.1

34.0

30.1–48.9

40.6

9th DS L

29.9

30.1–36.9

32.5–

25.1–35.3

30.7

31.5–47.3

38.8

10th DS L–

26.9–32.9

29.9–

21.9–31.1

27.2

26.7–41.7

33.3

11th DS L

19.3

21.8–25.8

23.6–

16.3–24.3

20.8

17.7–35.2

24.3

12th DS L

13.6

12.2–16.3

14.0–

9.6–17.9

13.6

11.6–22.7

14.6

13th DS L–

15.5–19.0

16.9

13.0

13.0–20.9

17.1

13.7–24.2

17.6

1st DSR L–

23.1–28.6

25.3

27.5

22.5–30.7

26.2

23.2–29.9

25.7
32.6

Longest DSR L

32.1

33.9–36.8

34.3

35.5

30.1–40.4

34.1

28.4–37.8

1st AS L

6.9

6.5–8.2

7.3

7.0

6.0–9.4

7.5

6.0–10.3

7.8

2nd AS L

10.6

9.9–13.5

11.8

11.3

10.3–16.0

12.2

10.1–16.7

12.2

3rd AS L–

17.3–19.1

18.0

17.6

14.6–20.0

17.5

13.5–24.9

16.7

1st ASR L–

23.2–31.9

26.7–

25.8–32.8

28.7

23.7–32.9

27.6

Longest ASR L–

32.4–38.6

34.3

37.4

29.5–41.2

35.5

29.0–39.8

34.2

1st P1 L

42.6

43.0–50.4

45.2

45.7

40.6–54.9

49.7

37.8–63.0

47.5

Longest P1 L

52.2

55.9–57.8

56.5

66.8

51.0–73.5

63.7

50.9–91.1

63.3

P2S L

17.0

16.6–18.9

17.7

17.7

14.5–19.5

17.5

13.4–26.3

17.6

Longest P2SR L

31.7

30.8–37.9

33.3

38.9

29.4–38.9

34.8

27.2–48.4

32.8

Caudal-fin L

49.3

50.2–64.6

54.3–

48.3–61.1

54.5

42.2–65.0

55.3

Caudal-peduncle L

15.6

14.1–16.9

15.4

14.9

12.7–16.8

14.6

12.9–17.6

15.7

Caudal-peduncle D

10.2

9.1–10.4

9.9

9.6

8.5–10.3

9.4

9.2–11.4

10.2

aat anal-fin originb

at vertical midline of eyecat posterior end of preocular spine base

AS: anal-fin spine; ASR: anal-fin soft ray; D: depth; DS: dorsal-fin spine; DSR: dorsal-fin soft ray; L: length; P1: pectoral-fin ray; P2S: pelvic-fin spine; P2SR:
pelvic-fin soft ray, W: width123

M. Matsunuma, H. Motomura

Holotype. USNM 218411, female (G), 93.4 mm SL, off
Somalia (11°040 N, 51°150 E), 76–80 m, H. A. Fehlmann onboard of RV Anton Bruun, bottom trawl, 17 December
1964.
Other specimens examined. Thirty-seven specimens,
48.8–121.1 mm SL: THAILAND (ANDAMAN SEA
COAST): KAUM–I. 33281, male (G), 116.4 mm SL, off
Ranong Province (obtained at fishing port at Pak Nam), T.
Yoshida and G. Ogihara, 7 December 2010; KAUM–I.
47529, male (G), 116.4 mm SL, KAUM–I. 47530, male(G), 121.1 mm SL, KAUM–I. 47531, male (G), 118.0 mm
SL, off Ranong Province (obtained at fishing port at Pak
Nam), T. Yoshida and H. Nishiyama, 23 June 2012.
INDIA: ZMH 5582, 21 specimens [six males (E), nine
females (E), and seven sex undetermined], 48.8–88.5 mm
SL, west-southwest of Kochi (9°400 –450 N, 75°380 –390 E),
138–210 m, RV Meteor, bottom trawl, 10 February 1965.
PAKISTAN: KAUM–I. 63511, female (G), 72.7 mm
SL, KAUM–I. 63512, male (G), 93.1 mm SL, KAUM–I.
63513, male (G), 92.0 mm SL, KAUM–I. 63514, male (G),90.5 mm SL, KAUM–I. 63515, male (G), 89.4 mm SL,
KAUM–I. 63516, female (G), 87.4 mm SL, KAUM–I.
63517, male (G), 76.8 mm SL, KAUM–I. 63518, female(G), 74.5 mm SL, KAUM–I. 63519, male (G), 76.2 mm
SL, KAUM–I. 63520, female (G), 63.3 mm SL, KAUM–I.
63521, female (G), 63.9 mm SL, off Pakistan (obtained atWest Wharf, Karachi), 50–60 m, bottom trawl, 1999–2000.
SOMALIA: ZMH 5581, female (E), 99.9 mm SL, off
Somalia (1°180 –200 S, 41°530 –560 E), 177–243 m, RV
Meteor, bottom trawl, 15 January 1965.
Diagnosis. A species of Ebosia distinguished from othermembers of the genus by the following combination ofcharacters: anal-fin soft rays 7–9 (modally 8); pectoral-fin
rays 16–19 (modally 17); scale rows above lateral line 4–5(modally 4); scale rows between last dorsal-fin spine baseand lateral line 4; scale rows between last dorsal-fin spinebase and lateral line 4; scale rows between sixth dorsal-fin
spine base and lateral line 3–5 (modally 4); scales onpreopercular region 7–14 (mean 10.6); postorbital length
19.8–24.2 (mean 21.9) % SL; a blotch above pectoral-finbase and blotches on pectoral-fin membrane relatively
large; yellowish median fins and broadly yellow pectoral
fin in males; elongated parietal spine in males relatively
narrow and strongly falcate posterodorsally, its tip reaching
fourth to eighth dorsal-fin spine bases.
Color of fresh male specimens. Ground color of headand body pale red, darker dorsally, whitish ventrally(Fig. 6). Three relatively narrow (width subequal to iris
diameter) dark red bands with narrow brown inner andwhite outer margins on sides of head; anteriormost bandcrossing eye, reaching posterodorsal corner of maxilla;
middle band from posteroventral margin of orbit, reaching
obliquely to ventral margin of subopercle; posteriormost

123band saddling nape at level of parietal ridges, reaching
posteroventral margin of opercle at level of upper origin of
pectoral-fin base. Five vertical moderately broad dark red
bands (about three vertical scale rows width) saddling
body; anteriormost band below first or second dorsal-fin
spine bases, extending posteroventrally and continuouswith a blotch above pectoral-fin base; second band below
fifth dorsal-fin spine base; third band below eighth to tenthdorsal-fin spine bases; fourth band below first to third
dorsal-fin soft ray bases; fifth band below sixth to eighth
dorsal-fin soft ray bases. Chest without markings. A relatively large black blotch above upper origin of pectoral-finbase (subequal to or slightly larger than iris). Eyes brightyellow to orange, iris black, without radial markings. Skin
flap on supraocular reddish; flaps on posterior margin of
preopercular white.
Spinous dorsal fin coloration similar to ground color ofbody, with 1–4 poorly defined red bands on each spine;
soft-rayed portion of dorsal fin pale yellow, somewhat
reddish basally. Anal fin pale yellow. Pectoral fin broadly
bright yellow, somewhat reddish basally and dorsally(Fig. 6a–b) or about two-thirds of upper portion red(Fig. 6c); with about 12–20 relatively large dark brown to
black blotches (subequal to iris) having reddish-brownmargins, sparsely scattered, and forming four or fiveindistinct rows of blotches. Pelvic fin yellowish to reddish
without markings. Caudal fin pale yellow, with several
indistinct small reddish-yellow to red spots scattered dorsally or basally.
Color of preserved specimens. Overall creamy-white tobrown (Figs. 3d–f, 4b); red markings retained as brown;
yellow coloration diffuse. Soft-rayed portion of dorsal fin
with ca. 10–15 brown spots on rays; soft-rayed portion ofanal fin with ca. 5–10 brown spots on rays; caudal fin withnumerous (ca. 30–40) brown spots on rays (number ofspots on all fins apparently greater in small specimens).
Distribution. Ebosia falcata is distributed off Somalia,
Pakistan and the west coast of India in the northwestern
Indian Ocean, and off the Andaman Sea coast of Thailand(Eschmeyer and Rama-Rao 1978; Kotthaus 1979; thisstudy; Fig. 5). The species has been collected by bottom
trawl in depths of 47–243 m (mostly [100 m) (Eschmeyer
and Rama-Rao 1978; this study).
Remarks. The present non-type specimens agreed wellwith the holotype and original description of E. falcata,with the exception of minor differences in some meristics
and morphometrics, including dorsal-fin rays [XIII, 9–11 inthis study vs. XIII, 9–10 in Eschmeyer and Rama-Rao(1978)] and snout length (13.1–16.2 % SL vs. 12–15 %SL), which may simply reflect intraspecific variations.
Ebosia falcata was originally described by Eschmeyer andRama-Rao (1978) on the basis of 26 specimens collected
off Somalia and the west coast of India, but has notNew Ebosia from western Indian Ocean
Fig. 6 Fresh male specimens ofEbosia falcata. a KAUM–I.
47529, 116.4 mm SL;
b KAUM–I. 47531, 118.0 mm
SL; c KAUM–I. 47530,
121.1 mm SL, all fromAndaman Sea

123

M. Matsunuma, H. Motomura

Discussion

Fig. 7 Relationships of body width (triangles), orbit diameter(circles), and pelvic-fin spine length (crosses) (all as % of standard
length) to standard length (mm) in Ebosia bleekeri

previously been recorded from the other localities listed
above (Eschmeyer and Rama-Rao 1978; Kotthaus 1979;
Manilo and Bogorodsky 2003). Accordingly, the specimens taken off Pakistan and in the Andaman Sea represent
the first records of E. falcata in those areas and suggest awidespread distribution of the species in the northeastern
Indian Ocean. Although Eschmeyer and Rama-Rao (1978)
provided a detailed description of E. falcata, it was based
solely on preserved specimens and therefore lacked any
description of fresh coloration. Photographs of the fourAndaman Sea specimens revealed that males of E. falcata
have a reddish head and body with yellowish fins whenfresh. Fresh female coloration remains unknown.
Genetic divergence. The genetic analysis, based on 555
base pairs of the mtDNA cytochrome b gene from two
examples of E. falcata (Andaman Sea) and three E. bleekeri (East China Sea), indicated a substantial degree ofgenetic divergence between the species, with differences
ranging from 2.5 % to 3.4 % in contrast to intraspecificdifferences of 0 % to 0.9 %.

123Morphological changes with growth. Analyses of 44
measurements taken from 38 specimens (48.8–121.1 mmSL) of Ebosia falcata indicated an absence of any positively growth-correlated morphological changes. However,relative lengths of the orbit diameter, head, thirteenth (last)dorsal-fin spine, first dorsal-fin soft ray, second and thirdanal-fin spines, and pectoral, pelvic, and caudal fins tended
to decrease as a percentage of standard length with growth,with similar changes, except for head length, also occurringin E. bleekeri [111 specimens (9.4–159.8 mm SL) (seeFig. 7)]. Additionally, body width and suborbital depth
became greater with growth in E. bleekeri. Since thenumbers and size range of available specimens of E. saya
were limited (n = 10, 63.0–87.0 mm SL), any growthrelated changes in relative lengths of body parts were notapparent. Decreasing length of fin rays with growth has
already been reported for several members of Scorpaenidae, e.g., Scorpaena spp. (Motomura et al. 2005c, 2011b)
and Pterois brevipectoralis (Mandrytsa 2002) (Matsunuma
and Motomura 2013).
The supraocular skin flap tends to become shorter(reduced in size) with growth in Ebosia, as in several
members of Pterois (Pteroinae) (Matsunuma and Motomura 2011, 2013). In E. saya, a small specimen (63.0 mm
SL, one of two specimens of USNM 407273; Fig. 4a) had arelatively large supraocular flap, its length slightly greaterthan the pupil diameter, whereas larger specimens ([ ca.
70 mm SL) had a small supraocular flap (length about half
pupil diameter). No other significant growth-related changes were found for skin flaps.
As in most members of Scorpaenidae, the number ofbranched pectoral-fin rays in both species of Ebosia
increases with growth. In E. bleekeri, all pectoral-fin raysof a 28.1 mm SL specimen were unbranched. Subsequently, all rays remained unbranched (or 6 or 7 rays
became branched at the same time) between 47.6 and61.1 mm SL, with 6–9 (rarely 5 or 11) rays branched in
specimens exceeding 64.8 mm SL (Fig. 8a). Among thebranched pectoral-fin rays, 0–8 rays tended to divide into
more than three (remaining rays branched into two) withgrowth, but this was highly variable (Fig. 8d). The numberof pelvic-fin soft rays with more than three branches alsotended to increase with growth in E. falcata and E. saya. In
E. falcata, all pelvic-fin rays of all specimens less than
63.6 mm SL had only two branches, with more than threebranches for 1–4 (rarely 0) rays in specimens exceeding
63.9 mm SL.
Sexual dimorphism and dichromatism. Sexual
dimorphism of parietal spine morphology in Ebosia,
already well documented, e.g., Schmidt (1931), Matsubara(1943), Eschmeyer and Rama-Rao (1978), was reconfirmed

New Ebosia from western Indian Ocean
Fig. 8 Relationships of numberof branched pectoral-fin rays(a–c) and pectoral-fin rays withmore than three branches (d–f)
to standard length (mm) in
Ebosia bleekeri (a, d),E. falcata (b, e), and E. saya. sp.
nov. (c, f). Pectoral-fin rays
counted on both sides wheneverpossiblehere on the basis of specimens for which sex had been
determined by gonad examination, e.g., four males and six
females of E. saya (Figs. 2, 3a–c), 10 males and sevenfemales of E. falcata (Figs. 3d–f, 6), and 11 males and nine
females of E. bleekeri (Fig. 9). The parietal spine of malesof the three species becomes a thin elongated bony crest,
curved posteriorly with growth (Figs. 10a–b, d–e, 11a–c).
By contrast, females have a low parietal ridge with a spinous margin (Figs. 10c, f, 11d).
Matsubara (1943) noted that the supraocular spine of male
E. bleekeri tends to be stronger than that of females. Chanand Chilvers (1973) also reported that male E. bleekeri differed from females in having stronger and a greater numberof spines on the preocular, supraocular, and suborbital ridgeand lower preopercular spine base (spines on the preopercular spine base refer to supplemental preopercular spines inthis study). Among the head spines, significant differences in
the number of supplemental spines on the third preopercularspine base were noted here between the sexes in E. bleekeri
and E. falcata (Fig. 12a–b), but not in E. saya, probably due
to the limited material (Fig. 12c). Moreover, in E. bleekeri,
males tended to have a single spine larger than adjacentspines on the supraocular (Fig. 11a–c), whereas all supraocular spines of females were small and similarly sized(Fig. 11d). No other significant sex-related differences inhead spine condition were apparent in Ebosia.
Sexual dichromatism has been reported for several
species in the scorpaenid genera Parascorpaena, Scorpaena, and Scorpaenopsis, wherein males only possess alarge black blotch on the spinous portion of the dorsal fin(e.g., Poss 1999; Randall and Eschmeyer 2002; Motomura
et al. 2005c, 2006b, 2007). Differences in fin coloration
between the sexes were also recognized in E. saya andE. bleekeri. Photographs of two paratypes of E. saya taken
prior to preservation showed the males with yellowish
median fins and entirely yellow pectoral fins (Fig. 2a), in
contrast to females (reddish median fins and red pectoralfins) (Fig. 2b). Moreover, male E. bleekeri had yellowish toorange median fins (Fig. 9a–b), such fins being red infemales (Fig. 9c). Although male E. falcata had yellowish
median fins and broadly yellow (about one-third or almost
entirely) pectoral fins (Fig. 6), the fresh coloration of
females remains unknown. However, considering themorphological similarities between E. saya and E. falcata,
similar sexual dichromatism in the latter is not unlikely.
The elongated parietal spine of males is an importantdiagnostic character for Ebosia species, but is not useful for
females and young males. For example, Pterois lunulata
Temminck and Schlegel 1843, reported by Iwamoto and
McCosker (2014; pl. 17, fig. 98, CAS 235752) from thePhilippines, was identified here as a female of E. bleekeri.
In fact, females and young males of the latter have infrequently been misidentified as species of Pterois, especiallyas P. lunulata or Pterois volitans (Linnaeus 1758), basedon our examinations of museum collections during thisstudy. However, Ebosia can be readily distinguished from
Pterois by the continuous coronal and parietal ridges (vs.
separated in Pterois); branched pectoral-fin rays (vs. all
unbranched); the nasal spine usually absent (vs. present); a
skin flap on the posterior lacrimal spine tip absent (vs.
present) (M. Matsunuma, unpublished data). It was further
noted that Pterois antennata (Bloch 1787), also reported by
Iwamoto and McCosker (2014; pl. 17, fig. 97, CAS
235573), was in fact P. volitans.
Species comparisons. Of the three nominal species
previously assigned to the genus Ebosia, e.g., Pterois
bleekeri Döderlein in Steindachner and Döderlein 1884(= E. bleekeri), Ebosia kagoshimae Oshima 1956, andEbosia falcata Eschmeyer and Rama-Rao 1978, E. kagoshimae was originally described on the basis of two specimens from Shibushi Bay, Kagoshima Prefecture,southeastern coast of Kyushu Island, southern Japan.
Although the present location of those specimens is
unknown, making them unavailable for the present study,123

M. Matsunuma, H. Motomura
Fig. 9 Fresh specimens ofmales (a–b) and a female (c) of
Ebosia bleekeri. a BSKU
38437, 119.0 mm SL, Tosa
Bay, Japan (Photo: BSKU);
b KAUM–I. 35804, 118.0 mm
SL, East China Sea; c KAUM–I.
35803, 108.2 mm SL, East
China Sea

photographs of them plus the description of E. kagoshimae
given by Oshima (1956), with the exception of dorsal- andanal-fin ray counts (made by different methods), agreed

123

closely with the original description of P. bleekeri.
Although Oshima (1956) stated that E. kagoshimae differed from E. bleekeri in having the relatively long parietal

New Ebosia from western Indian Ocean
Fig. 10 Lateral views of headof Ebosia saya sp. nov. (male:a–b; female: c) and E. falcata(male: d–e; female: f), showing
ontogenetic changes and sexual
differences of parietal andsupraocular spines. a KAUM–I.
70134, 85.2 mm SL (right side;
reversed); b LACM 44793-3,
73.5 mm SL (right side;
reversed); c USNM 307910,
holotype of E. saya, 87.0 mm
SL (right side; reversed);d KAUM–I. 33281, 116.4 mm
SL; e ZMH 5582, 70.9 mm SL;
f ZMH 5582, 82.2 mm SL. Bars
indicate 5 mmspines of male and lacking a vertical black band on thenape, the parietal spine length is changed with growth andband markings sometimes become defused in preserved
specimen. Ebosia kagoshimae is therefore regarded as a
junior synonym of P. bleekeri and conspecific with specimens identified as E. bleekeri in this study, as in Eschmeyer and Rama-Rao (1978). Accordingly, only two
nominal species, E. bleekeri and E. falcata, are currently
regarded as valid in the genus Ebosia.
Ebosia saya is readily distinguished from E. bleekeri byhaving usually one more anal-fin soft ray [8 vs. 6–8(modally 7) in E. bleekeri] (Table 1), one more pectoral-fin
ray [17–18 (modally 17) vs. 15–17 (16)], the elongatedparietal spines of males relatively narrow (width of elongated portion clearly less than pupil diameter) and strongly
falcate posterodorsally, the posteriormost tip reaching tolevel with the sixth or seventh dorsal-fin spine base at ca.
90 mm SL [vs. relatively broad (width ca. 1.0–1.5 times
pupil diameter) and less falcate posterodorsally, its tip
reaching to level with the first to fourth dorsal-fin spinebase at ca. 90 mm SL] (Figs. 10a–b, 11a–c), and the pectoral fin of males yellow (vs. red in both sexes) (Figs. 2, 9).
Ebosia saya is most similar to E. falcata in overall body
appearance, with both species sharing the aforementioned
characters separating E. saya from E. bleekeri. However,the former can be distinguished from E. falcata by having
usually one more scale row above the lateral line [5 vs. 4–5(modally 4) in E. falcata] (Table 1), 4–6 (modally 5) scalerows between the sixth dorsal-fin spine base and the lateralline [vs. 3–5 (4)], 4–5 (modally 5) scale rows between thelast dorsal-fin spine base and the lateral line (vs. 4), aslightly higher number of scales on the preopercular region[11–22 (mean 15.4) vs. 7–14 (10.7)] (Fig. 13), and aslightly shorter postorbital length [18.1–19.2 (mean
18.7) % of SL vs. 19.8–24.2 (21.9) % of SL] (Table 2;
Fig. 14). Moreover, E. saya tended to have a relatively
higher number of spines on the preocular, supraocular, andpostocular bones compared to E. falcata (Fig. 15),
although the spine number increased with growth in both
species. At 60–90 mm SL, E. saya (n = 10) had 1–6 (mean
3.1) spines on the preocular, 1–6 (3.7) spines on the supraocular, and 2–9 (5.2) spines on the postocular bones,
giving a total number of 5–20 (12.3) spines, whereas
E. falcata at the same stage (n = 15) had 1–7 (2.7), 1–6(2.9), 2–6 (3.3), and 4–18 (8.4) spines, respectively. Such
differences in head spine number between the two species
may be indicative of a more rapid growth rate and smaller
maximum body size in E. saya, compared with E. falcata(largest examined specimens of E. saya and E. falcata
85.2 mm SL and 121.1 mm SL, respectively). Ebosia saya
also differed from E. falcata in having the blotch above thepectoral-fin base and blotches on the pectoral fin smaller(Figs. 3–4, 16) [maximum diameter of blotch above pectoral-fin base 32.6–56.2 (mean 41.2) % of orbit diameter(OD) in E. saya (n = 6) vs. 26.6–95.2 (66.7) % of OD in
E. falcata (n = 25); that of blotches on pectoral fin
13.5–34.5 (mean 21.7) % of OD in E. saya (n = 10) vs.
27.1–62.1 (42.0) % of OD in E. falcata (n = 23)].

123

M. Matsunuma, H. Motomurab Fig. 11 Lateral views of head of male (a–c) and female (d) Ebosia

bleekeri, showing ontogenetic changes and sexual differences ofparietal and supraocular spines. a URM-P 3733, 123.3 mm SL;
b MNHN2008-1399, 91.4 mm SL; c KAUM–I. 39277, 100.5 mm SL;
d KAUM–I. 35803, 108.2 mm SL. Arrowheads indicate largest
supraocular spines. Bars indicate 5 mm

As with E. saya, E. falcata is readily distinguishable
from E. bleekeri by having usually one more anal-fin softray (7–9, modally 8) (Table 1), usually one more pectoralfin ray (16–18, modally 17), and males with a narrowstrongly falcate parietal spine (Fig. 9d–e) and a broadly
yellow pectoral fin (Fig. 6). Moreover, E. falcata differedfrom E. bleekeri in having usually one fewer scale rows
above the lateral line [5 vs. 5–7 (modally 6) in E. bleekeri](Table 1), 3–5 (modally 4) scale rows between the sixth
dorsal-fin spine base and lateral line [vs. 4–6 (5)], 4 scalerows between the last dorsal-fin spine base and the lateralline [vs. 4–6 (5)], slightly fewer scales on the preopercularregion (Fig. 13), and a greater postorbital length (Table 2;
Fig. 14).
Comparative materials. Ebosia bleekeri: 11 specimens, 9.4–159.6 mm SL. JAPAN: BSKU 198, 101.1 mm
SL; BSKU 250, 110.6 mm SL; BSKU 1717, 92.7 mm SL;
BSKU 2420, 115.4 mm SL; BSKU 8215, 127.7 mm SL;
BSKU 9463, 90.8 mm SL; BSKU 9467, 68.8 mm SL;
BSKU 12840, 47.6 mm SL; BSKU 13843, 96.0 mm SL;
BSKU 36277, 61.1 mm SL; BSKU 38437, 119.0 mm SL;
BSKU 52412, 91.6 mm SL; BSKU 55594, 28.1 mm SL;
BSKU 68780, 109.0 mm SL; BSKU 69892, 69.9 mm SL;
BSKU 70979, 108.3 mm SL; BSKU 74743, 74.4 mm SL;
BSKU 78142, 58.1 mm SL; BSKU 86197, 136.7 mm SL;
BSKU 97577, 127.4 mm SL; BSKU 103849, 121.0 mm
SL; BSKU 103850, 128.9 mm SL; BSKU 103851,
122.0 mm SL; BSKU 109563, 105.9 mm SL; FAKU 1342,
94.6 mm SL; FAKU 3115, 41.6 mm SL; FAKU 4539,
62.5 mm SL; FAKU 4940, 80.6 mm SL; FAKU 28066,
124.0 mm SL; FAKU 59336, 93.3 mm SL; FAKU 97045,
140.1 mm SL; HUMZ 35842, 88.2 mm SL; HUMZ 36256,
86.2 mm SL; HUMZ 177808, 67.9 mm SL; KAUM–I.
44331, 105.9 mm SL; KSHS 22472, 147.1 mm SL; KSHS
22494, 159.6 mm SL; KSHS 22495, 139.1 mm SL; KSHS
22496, 134.3 mm SL; MUFS 5290, 106.4 mm SL; NSMTP 66383, 82.5 mm SL; USNM 148915, 78.2 mm SL;ZUMT 4608, 122.5 mm SL; ZUMT 47014, 124.4 mm SL;ZUMT 47228, 63.0 mm SL; ZUMT 49647, 47.6 mm SL;ZUMT 50927, 74.1 mm SL.
EAST CHINA SEA: FAKU 69245, 105.5 mm SL;
FAKU 81698, 107.7 mm SL; FAKU 81699, 86.6 mm SL;
HUMZ 199824, 103.3 mm SL; HUMZ 199871, 115.9 mm
SL; HUMZ 199872, 116.0 mm SL; HUMZ 199992,
109.0 mm SL; HUMZ 199993, 123.7 mm SL; HUMZ
200059, 117.5 mm SL; KAUM–I. 35803, 108.2 mm SL;

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New Ebosia from western Indian Ocean

Fig. 12 Relationships of number of supplemental spines on third
preopercular spine to standard length (mm) in male (diamonds) andfemale (circles) Ebosia bleekeri (a), E. falcata (b), and E. saya. sp.

Fig. 13 Relationships of the number of scales on the preopercularregion to standard length (mm) in Ebosia saya sp. nov. (stars),
E. bleekeri (triangles), and E. falcata (circles). H indicates holotype

KAUM–I. 35804, 120.2 mm SL; KAUM–I. 35817,
106.6 mm SL; KAUM–I. 35818, 105.3 mm SL; KAUM–I.
57753, 64.8 mm SL; KAUM–I. 57771, 68.8 mm SL;
SNFR 362, 117.1 mm SL; SNFR 543, 119.3 mm SL;
SNFR 545, 120.1 mm SL; SNFR 1280, 94.7 mm SL;
SNFR 11119, 2 specimens, 117.5–125.5 mm SL; SNFR

nov. (c). Symbols possessing cross indicate sex estimated fromparietal spine morphology. Spines counted on both sides wheneverpossibleFig. 14 Relationships of postorbital length (as % of standard length)
to standard length (mm) in Ebosia saya sp. nov. (stars), E. bleekeri(triangles), and E. falcata (circles). H indicates holotype

13098, 5 specimens, 94.3–145.3 mm SL; SNFR 13251,
42.7 mm SL; URM-P 3733, 123.3 mm SL; URM-P 3734,
109.2 mm SL; URM-P 7134, 127.6 mm SL; URM-P
23694, 134.5 mm SL; URM-P 23695, 124.9 mm SL;
URM-P 23696, 114.6 mm SL; URM-P 23697, 78.7 mm

123

M. Matsunuma, H. Motomura

Fig. 15 Relationships of the total number of preocular, supraocular,
and postocular spines to standard length (mm) in Ebosia saya sp. nov.
(stars; solid line) and E. falcata (circles; dashed line). Curved lines
indicate polynomial trendlines. Spines counted on both sides
whenever possible. H indicates holotype

SL; URM-P 25719, 127.8 mm SL; URM-P 25720,
109.8 mm SL; URM-P 43724, 105.2 mm SL; ZUMT
52112, 111.7 mm SL; ZUMT 52120, 118.4 mm SL.
TAIWAN: ASIZP 63066, 120.8 mm SL; ASIZP 61433,
2 specimens, 52.7–57.6 mm SL; KAUM–I. 39276,
136.2 mm SL; KAUM–I. 39277, 100.5 mm SL; KAUM–I.
39278, 89.6 mm SL; KAUM–I. 39279, 94.0 mm SL;
KAUM–I. 39280, 121.1 mm SL; KAUM–I. 39281,
93.8 mm SL; KAUM–I. 52578, 115.0 mm SL; KAUM–I.
52579, 68.5 mm SL; NMMB-P 9558, 105.5 mm SL;
NMMB-P 9560, 107.6 mm SL; NTUM 3854, 76.1 mm SL.
SOUTH CHINA SEA: BSKU 17282, 79.2 mm SL.
VIETNAM: NMMB-P 12649, 81.3 mm SL.
PHILIPPINES: MNHN 2004-1629, 30.8 mm SL;
MNHN 2005-827, 56.5 mm SL.
INDONESIA: NCIP 5003, 79.8 mm SL; WAM
P. 30720-002, 9.4 mm SL.
VANUATU: MNHN 2008-1399, 91.4 mm SL.

123

Fig. 16 Relationships of maximum diameter of blotch on pectoral-fin
membrane (mm) to maximum diameter of blotch above pectoral-finbase in Ebosia saya sp. nov. (stars) and E. falcata (circles).
H indicates holotype

AUSTRALIA:
NMV
A4410,
2
specimens,
60.9–72.6 mm SL.
NO DATA: FAKU 103332, 129.3 mm SL; ZUMT
16705, 77.8 mm SL (probably from Japan).
Acknowledgments We are deeply grateful to M. McGrouther, A.
Hay, and S. Reader (AMS); K.-T. Shao, Y.-C. Liao, and M.-Y. Lee(ASIZP); H. Endo, T. Yamakawa, N. Nakayama (BSKU), and E.
Katayama (formerly BSKU); T. Nakabo, Y. Kai, A. Tokairin(FAKU), and N. Muto (formerly FAKU); M. Yabe, H. Imamura, and
T. Kawai (HUMZ); R. Causse (MNHN); Y. Iwatsuki (MUFS), K.
Hidaka, and K. Miyamoto (formerly MUFS); Z. Arifin, R. Pratiwi, M.
Adrim, I. Alhakim, and K. Wibowo (NCIP); H.-C. Ho (NMMB); M.
Gomon and D. Bray (NMV); K. Matsuura and G. Shinohara (NSMT);
K.-Y. Wu (NTUM); T. Yoshino (formerly URM); K. Hoshino and M.
Okamoto (SNFR); J. Williams (USNM); S. Morrison (WAM); R.
Thiel and I. Eidus (ZMH); and K. Sakamoto (ZUMT) for their kind
hospitality during the first author’s visits to their institutions. We arealso grateful to R. Feeney (LACM) for providing opportunities to
examine specimens, H.-C. Ho (MNHB) for donating specimens, H.
Endo and T. Kawai for providing photographs of fresh E. bleekeri andE. saya, respectively, and Y. Haraguchi, students, and other KAUM
volunteers for curatorial assistance and sampling specimens. The firstauthor is especially grateful to S. Chiba (NSMT) for advice withmolecular analysis and M. Sakurai (The Kagoshima Immaculate
Heart College) for advice with gonad examinations. G. Hardy

New Ebosia from western Indian Ocean(Ngunguru, New Zealand) read the manuscript and provided help
with English. Specimens from Pakistan were donated to KAUM from
Marine Fisheries Department, Government of Pakistan, and P. Psomadakis (FAO, Rome). This study was supported in part by Grantsin-Aid for Scientific Research (A: 26241027, B: 24370041, and C:
23580259 and 26450265) from the Japan Society for the Promotion of
Science, Tokyo, Japan (JSPS), 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’’, the Coastal
Area Capability Enhancement in Southeast Asia Project of the
Research Institute for Humanity and Nature, Kyoto, Japan, and the‘‘Biological Properties of Biodiversity Hotspots in Japan’’ project of
the National Museum of Nature and Science, Tsukuba, Japan, and by
the grant awarded to the first author from the Mikimoto Fund forMarine Ecology.

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