Ichthyol Res
DOI 10.1007/s10228-015-0479-2

FULL PAPER

A new species of scorpionfish, Ebosia vespertina (Scorpaenidae:
Pteroinae), from the southwestern Indian Ocean
Mizuki Matsunuma1 • Hiroyuki Motomura2

Received: 7 May 2015 / Revised: 26 June 2015 / Accepted: 26 June 2015
Ó The Ichthyological Society of Japan 2015

Abstract A new species of scorpaenid fish, Ebosia vespertina sp. nov., is described on the basis of 19 specimensfrom off Mozambique, South Africa, and Madagascar,southwestern Indian Ocean. The new species is most similar to Ebosia falcata and Ebosia saya, known from thenorthern and eastern Indian Ocean and Saya de Malha
Bank, respectively, in having usually 8 anal-fin soft rays,
usually more than 17 pectoral-fin rays, and the elongatedparietal spine in males narrow and strongly curved posterodorsally. However, E. vespertina can be distinguishedfrom the two latter species by the following combination ofcharacters: pectoral-fin rays 17–18 (modally 18); scalerows above the lateral line 4–6 (5); scale rows between thelast dorsal-fin spine base and lateral line 4–5 (5); scale rows
between the sixth dorsal-fin spine base and lateral line 4–5(5); postorbital length 17.4–20.3 (mean 18.7) % of standard
length (SL); longest pectoral-fin ray length 54.6–73.9(65.1) % of SL; longest pelvic-fin soft ray length 32.4–44.0(37.6) % of SL. The remaining congener, Ebosia bleekeri,

This article was registered in the Official Register of Zoological
Nomenclature (ZooBank) as 685CE299-1C44-45A2-B99CF4C9E6313EAE.
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.
& Mizuki Matsunuma
k1139853@kadai.jp
Hiroyuki Motomura
motomura@kaum.kagoshima-u.ac.jp
1

Seikai National Fisheries Research Institute, 1551-8 Taira,Nagasaki 851-2213, Japan

2

The Kagoshima University Museum, 1-21-30 Korimoto,Kagoshima 890-0065, Japan

known from the western Pacific Ocean, usually has 7 analfin soft rays and 16 pectoral-fin rays, and the elongatedparietal spine in males relatively broad and not so strongly
curved.
Keywords South Africa  Mozambique  Madagascar 
Sexual dimorphism  Ontogenetic change

Introduction
The Indo-West Pacific scorpionfish genus Ebosia Jordan
and Starks 1904 (Scorpaenidae: Pteroinae) inhabits sandyor muddy bottoms in depths usually less than 100 m andhas therefore been collected mainly by bottom trawl. The
genus is characterized by having the coronal, parietal andnuchal spines bases continuous, and the parietal spine
elevated, being a thin bony crest in males (Jordan andStarks 1904; Eschmeyer and Rama-Rao 1978; Matsunuma
and Motomura 2014). Three valid species are currently
recognized in the genus, viz. Ebosia bleekeri (Döderlein in
Steindachner and Döderlein 1884) (western Pacific Ocean),Ebosia falcata Eschmeyer and Rama-Rao 1978 (northernand eastern Indian Ocean) and Ebosia saya Matsunuma andMotomura 2014 (Saya de Malha Bank, western Indian
Ocean).
During a revisionary study of Ebosia, 19 specimens of apreviously unrecognized species of Ebosia, collected off
Mozambique, South Africa and Madagascar, were found inSouth African museums, all having been primarily identified and catalogued as E. falcata or a species of a different
scorpaenid genus. Following careful examination, it wasdetermined that the southern African specimens represented a distinct species of Ebosia, differing from all other123

M. Matsunuma, H. Motomura

known congeners. They are herein described as a new
species.

Materials and methods
Measurements generally followed Motomura (2004b, c),with head width, head depth, maxillary depth and body
depth at the anal-fin origin following Motomura et al.
(2005b, 2006a), Motomura (2004a), Motomura et al.
(2006b) and Matsunuma et al. (2013), respectively.
Counts generally followed Motomura et al. (2005a–c) andMotomura and Johnson (2006); predorsal scale and cheekscale counts followed Motomura et al. (2006b) andMatsunuma and Motomura (2013), respectively. Measurements of the diameter of the blotch above the pectoral-fin base and blotches on the pectoral-fin membrane
followed Matsunuma and Motomura (2014). 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 (counted on both sides). Head spine terminology
generally follows Randall and Eschmeyer (2002: fig. 1),
Motomura (2004c: fig. 1), and Matsunuma and Motomura(2014: fig. 1). Supplemental preopercular spine and laterallacrimal spine terminology follows Eschmeyer (1965) andMotomura and Senou (2008: fig. 2), respectively. Sex wasdetermined by direct examination of gonads by lightmicroscope whenever possible. In the lists of specimens
examined, (G) indicates sex determined by gonad examination; (P), sex estimated from parietal spine morphology. Standard length is abbreviated as SL. In thedescription, features for the holotype are presented first,
followed by paratype data (if different) in parentheses.
The sample sizes for meristic characters, including the

Fig. 1 Fresh specimen of
Ebosia vespertinus sp. nov.,
Mozambique (not retained).
Photo: M. Lee

123holotype, are shown in parentheses. Institutional abbreviations follow Sabaj Pérez (2014).

Ebosia vespertina sp. nov.
(New English name: Western Falcate Lionfish) (Figs. 1–7;
Tables 1–2).
Holotype. SAM 41190 (formerly one of SAM 34266),
male (G), 99.1 mm SL, off Mozambique (18°200 S,
37°200 E), 68 m, RV Algoa, bottom trawl, 17 June 1994.
Paratypes. Eighteen specimens, 50.0–109.9 mm SL(seven males, eight females and three sex undetermined).
MOZAMBIQUE: SAM 34265, four specimens, two males
and two females (G), 58.9–106.1 mm SL, off Mozambique(19°490 S, 36°050 E), 54 m, RV Algoa, bottom trawl, 14 June
1994; SAM 34266, six specimens, three males and threefemales (G), 70.5–108.3 mm SL, collected with holotype.
SOUTH AFRICA: KAUM–I. 75290 (formerly one of
SAIAB 76118), female (P), 82.9 mm SL, 9 km off Richards
Bay, KwaZulu-Natal (28°500 1600 S, 32°110 2100 E), 50 m, A.
Connell, 7 May 2003; SAIAB 76118, male (P), 86.0 mm SL,
collected with KAUM–I. 75290; SAIAB 186409, two
specimens, one female (G) and one sex undetermined,
50.0–86.8 mm SL, off Thukela, KwaZulu-Natal(29°21.8170 S, 31°48.7610 E), 70 m, S. Fennessy, bottom
trawl, 19 March 2010; SAIAB 186466, male (P), 109.9 mm
SL, off KwaZulu-Natal (28°42.3800 S, 32°22.3060 E),
108–110 m, ACEP Trawl 3.2, S. Fennessy, bottom trawl, 18
August 2010; USNM 435892 (formerly one of SAIAB
76118), female (P), 65.0 mm SL, collected with KAUM–I.
75290. MADAGASCAR: SAIAB 189715, two specimens,
sex undetermined, 57.9–63.1 mm SL, off Tolagnaro(25°22.40 S, 47°02.80 E), bottom trawl, 3 May 2010.
Diagnosis. A species of Ebosia distinguished from othermembers of the genus by the following combination of

New Ebosia from southwestern Indian Ocean
Table 1 Frequency distribution of selected meristics of Ebosia vespertina sp. nov., E. falcata and E. saya
Pectoral-fin rays (one/other sides)

E. falcataa

16/16

16/17

17/17

17/18

18/18

4

1

4

26

4

1H

33H

H

1

1

E. saya8

E. vespertina

1

E. falcata

18H

SR in longitudinal series

3

4

41

1

5

30

6

3

18

H

H

E. saya

1

6

4

10H

1

Upper gill rakers
4

5

1

35

E. saya3

H7

E. vespertina

12

5

1

H

1
4

2
2H

10
15

14H

42
3

43

44
4
1

46

47

48

3

1

H

2

1

1

H

1

2

49

50

51

4

4H

1

Total gill rakers

11
H

1

45

6H

2

Lower gill rakers
6

H

5

H

6

3

3

SR between last DS and LL

4

5

9H

SR between 6th DS and LL

E. vespertina

E. falcata

Scales above LL

20

12
3

14
1

H

3

7

12

7H

10

15

16

17

14

18

5

6

4

H

3

5

H

1H

DS dorsal-fin spine base, LL lateral line, SR scale rowsaOne specimen had 17/19 pectoral-fin rays

H

indicates holotype

characters: anal-fin soft rays 7–8 (modally 8); pectoral-fin
rays 17–18 (18); scale rows above lateral line 4–6 (5); scalerows between last dorsal-fin spine base and lateral line 4–5(5); scale rows between sixth dorsal-fin spine base andlateral line 4–5 (5); postorbital length 17.4–20.3 (mean
18.7) % SL; longest pectoral-fin ray length 54.6–73.9(65.1) % SL; longest pelvic-fin soft ray length 32.4–44.0(37.6) % of SL; elongated parietal spine in males relatively
narrow, strongly falcate posterodorsally, its tip reaching thelevel of second to fifth dorsal-fin spine bases; a blotch
above pectoral-fin base and blotches on pectoral-fin
membrane relatively large; soft-rayed portions of dorsal
and anal fins, and caudal and pectoral fins yellowish inmales when fresh.
Description. Selected meristics and morphometrics
shown in Tables 1–2. Meristics given in Table 1 not
repeated here. Dorsal-fin rays XIII, 9; anal-fin rays III, 8[III, 7 (2 specimens) or III, 8 (17)]; pectoral fin with 1 (leftside) or 2 (right side) upper unbranched rays ? 10 (9–12 inparatypes) branched rays ? 7 (left side) or 6 (right side)(5–8) lower unbranched rays; pelvic-fin rays I, 5. Poredlateral-line scales lost in all specimens examined; scales
below lateral line uncountable in holotype [9 (2) or 10 (1)(based on only paratypes)]; pre-dorsal-fin scale rows 3 [2(6) or 3 (13)]; oblique cheek scale rows 2 [2 (13) or 3 (4)];horizontal cheek scale row 1; vertical cheek scale row 0.
Lower limb of gill arch with 2 [1 (3), 2 (15) or 3 (1)] gillrakers on hypobranchial. Branchiostegal rays 7. Swim
bladder present.
Body oblong, moderately compressed anteriorly, extensively compressed posteriorly; depth moderate, maximum
body depth less than longest dorsal-fin spine length. Headlarge, its length greater than body depth. A short simple,
pointed 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. Supraocular with a small
flap, its length 44.0 % of orbit diameter (18.0–72.9 % oforbit diameter; becoming shorter with growth). Two small
flaps on preopercle margin below third preopercular spinebase; tips not reaching posterior margin of interopercle whenlaid flat. An extremely small skin flap anterodorsally on orbit
surface (usually 1 or 2, rarely 3) [an abnormal 106.1 mm SL
specimen (one of SAM 34265) with 18 cirri in left orbit(uncountable in right orbit)]; its length less than one-third of
posterior nasal pore diameter. Snout tip with two extremely
short barbels; its length subequal to that of orbit surface flap.
No other skin flaps on head or body, including ventral marginof lacrimal.
Relatively well-developed ctenoid scales possessing 3–7
spinules on postorbital and suborbital regions; suborbital
scale area not extending anteriorly beyond level of midorbit; suborbital pit always without scales. Preopercularregion bordered by posterior margin of maxilla, suborbital123

M. Matsunuma, H. Motomura
Table 2 Morphometrics ofEbosia vespertina sp. nov.,
expressed as percentages ofstandard length

Holotype (SAM 41190)

Paratypes (n = 18)

SL (mm)99.1

50.0–109.9

Body depth (% SL)

36.5

30.8–39.2

Body depth at anal-fin origin

25.9

23.5–28.7

25.7

Body width

24.4

15.4–25.5

22.2

Head length

39.9

37.6–41.1

39.4

Head width

14.8

12.8–15.2

14.2

Head depth

16.8

14.4–17.5

16.4

Snout length

14.6

12.2–15.8

13.8

Orbit diameter

11.0

10.3–12.8

11.2

Interorbital width at mid-orbit

8.9

8.0–10.5

9.4

Interorbital width at preocular spine base

7.8

6.6–9.5

8.3

Upper-jaw length

15.7

13.9–16.5

15.7

35.7

Maxillary depth

5.7

4.8–6.8

5.5

Suborbital depth

4.2

1.7–4.4

3.4

Postorbital lengthPre-dorsal-fin length

18.9
36.6

17.4–20.3
31.8–39.5

18.7
35.9

Pre-anal-fin length

68.3

64.4–70.2

67.4

Pre-pelvic-fin length

36.6

32.4–38.1

34.8

1st dorsal-fin spine length

26.4

25.4–30.1

27.4

2nd dorsal-fin spine length

33.4

29.6–40.2

35.0

3rd dorsal-fin spine length

38.5

38.9–46.5

41.2

4th dorsal-fin spine length

42.2

38.3–47.9

43.4

5th dorsal-fin spine length–

44.0–51.0

45.7

6th dorsal-fin spine length

43.1

43.1–51.2

45.5

7th dorsal-fin spine length

43.8

41.7–51.2

45.8

8th dorsal-fin spine length–

40.7–49.0

43.6

9th dorsal-fin spine length

37.8

35.9–47.6

38.8

10th dorsal-fin spine length–

30.6–37.9

34.1

11th dorsal-fin spine length

19.7

22.0–28.6

24.6

12th dorsal-fin spine length

12.2

12.0–22.0

15.3

13 dorsal-fin spine length
1st dorsal-fin soft ray length

16.8
25.9

15.6–20.9
24.4–30.4

18.1
27.2
34.5

Longest dorsal-fin soft ray length33.5 (4th)31.0–37.1 (4th or 5th)

1st anal-fin spine length

7.3

6.6–10.8

8.6

2nd anal-fin spine length

10.6

10.1–16.0

13.1

3rd anal-fin spine length

16.0

14.7–22.0

17.6

1st anal-fin soft ray length

27.5

25.3–31.5

28.1

Longest anal-fin soft ray length35.2 (4th)31.1–41.2 (3rd or 4th)

36.0

1st pectoral-fin ray length

49.7

39.8–59.7

51.0

Longest pectoral-fin ray length62.1 (7th)54.6–73.9 (4–7th)

65.1

Pelvic-fin spine length

15.7

13.3–20.6

17.4

Longest pelvic-fin soft ray length33.9 (3rd)32.4–44.0 (2nd or 3rd)

37.6

Caudal-fin length

54.4

46.4–59.0

53.8

Caudal-peduncle length

16.0

13.4–17.0

15.1

Caudal-peduncle depth

10.3

8.8–11.1

9.9

ridge and posterior marginal ridge of preopercular, covered
with relatively well-developed 16 (14–16) ctenoid scalespossessing 3–9 spinules. Opercle covered with a few

123

Mean

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

New Ebosia from southwestern Indian Ocean

the remaining scales). Anterior region of occipital areawith a few [ca. 7 (7–8)] weakly ctenoid scales; large scaleswith 3–6 spinules, small scales with a single spinule orwithout spinules; scales mostly embedded with tip of
spinule only exposed. Interorbital region, including frontal
surface of preocular, dorsal surface of supraocular andpostocular, and interorbital canal without scales. A small
patch of weakly ctenoid scales with 1–3 spinules (allcycloid 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 dorsal-fin
spine base; ventral surface of chest with a few ctenoid
scales with 3–6 (1–6) spinules; reminder of body withcycloid scales. Basal regions of dorsal and anal fins withoutscales; caudal-fin base with small cycloid scales; basal
scale morphology on pectoral-fin base unknown (fewremaining scales cycloid; most scales lost in all specimens
examined).
Mouth set relatively low on head, protruding moderately
downward; moderately large, slightly oblique, forming anangle 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
extending slightly beyond mid-orbit level. Lips thickened;
lower lips broadly extending onto mandible. Lower jaw
with a small symphyseal knob. Symphyseal gap separatingpremaxillary teeth bands distinctly broader than width ofeach band (gap width approximately twice teeth band
width); both jaws with a relatively narrow band of small,
slender conical teeth; about 5–8 tooth rows at front ofupper jaw; about 3–6 tooth rows at front of lower jaw; teeth
bands narrowing posteriorly. Eight to ten rows of small
conical teeth forming blunt V-shaped patch on vomer; nopalatine teeth. Underside of dentary with three sensory
pores on each side; two small pores on each side of symphysial knob of lower jaw on each side. Gill rakers on firstgill arch short, tips expanded; longest raker on first gill archabout 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 7 (1–10) spinous points along orbit margin.
Supraocular with 7 (1–6) small spines on outer margin.
Postocular with 7 (2–12) small spines along orbit margin.
Interorbital ridge not developed, lacking spines; diverging

anteriorly and posteriorly in dorsal view, distance between
interorbital ridges narrowest at vertical midline of eye;interorbital canal moderately broad and deep, its widthabout one-third of iris diameter. Coronal with a single ridge
continued to interorbital and parietal aspects, with a single(1–3) minute spinous point. Parietal with a single ridge; infemales, a low elevated ridge with spinous margin, with
1–3 blunt spinous points (based on female paratypes); in
males, parietal ridge elongated as a narrow thin bony crest,
curved posterodorsally, its tip reaching level of second to
fifth dorsal-fin spine bases (based on holotype and maleparatypes). Nuchal with a single ridge completely fused toparietal ridge with a single spinous point on posterior end,
directed posterodorsally. Coronal–parietal–nuchal ridges ofboth sides running almost parallel in dorsal view. Tympanic without spines or ridges. Anterior margin of occipitalarea sloped transversely from between origins of coronalridges, 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 region with 10 (3–13) small spines surrounding ashort sensory canal. Pterotic with a short ridge with 5 (1–4)
small spines. Lower posttemporal with a short ridge with 2(1 or 2) small spines. Cleithrum with two low ridges; upperridge short without spinous points; lower ridge long,
curved posteroventrally with 2 (1 or 2) spinous points
posteriorly.
Lateral lacrimal ridge short, with 3 (1–4) small spines;
other ridges on lacrimal strongly spinous (not spinous insmall specimens). Suborbital with a single ridge seemingly
divided into anterior and posterior portions, with 6 (4–10)
and 7 (3–9) small spines on anterior and posterior ridges,
respectively; 22 (0–13) small associated spines below
ridge; 35 (7–29) spines in total. Anterior lacrimal spinewith a single (or absent) spinous point directed ventrally.
Posterior lacrimal portion plate-like with 3 (2–5) spinous
points distally. Preopercle with 5 spines; upper 3 spines ofsimilar length; lower 2 spines reduced and skin-covered; nosupplemental spine on first (uppermost) preopercular spinebase; a single (or 2) supplemental spine on second preopercular spine base; third preopercular spine base with asingle blunt ridge with 3 (1–3) spinous points. 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; fourth or fifth spine longest (based onparatypes; unknown in holotype); penultimate (twelfth)[rarely posteriormost (thirteenth)] spine shortest, its length
62 (47–67) and 73 (68–96) % of that of antepenultimate(eleventh) and posteriormost (thirteenth) spines,123M. Matsunuma, H. Motomura

respectively (data on specimen with shortest thirteenthspine not included); membrane of spinous portion of dorsal
fin strongly incised. Dorsal-fin soft rays all branched (firstray unbranched in small specimens); fourth (or fifth) ray
longest, its length distinctly less than that of longest dorsalfin spine (71–82 % of longest dorsal-fin spine length; basedon paratypes); 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 69(60–69) and 46 (42–55) % of that of second and third
spines, respectively. Anal-fin soft rays all branched; fourth(third or fourth) ray longest, its length subequal to that oflongest dorsal-fin soft ray [105 (97–113) % of longest
dorsal-fin soft ray length]; posteriormost ray free from
caudal peduncle. Pectoral fin long with rounded contour,
seventh ray (fourth to seventh) longest; its tip slightly
extending beyond level of anal-fin base posterior end (extending beyond caudal-fin base in small specimens); lower
five rays weakly thickened. Pelvic-fin spine base below
third dorsal-fin spine base; all pelvic-fin soft rays branched;third (second or third) soft ray longest, its tip extending
slightly beyond first anal-fin spine base; posteriormost soft
ray with membranous connection to abdomen for approximately half (one-fourth to half) of ray length. Caudal fin
long with rounded contour, its length 136 (119–148) % ofhead length, with 2 (rarely 3) procurrent rays, 2 segmented
unbranched rays, and 4 segmented branched rays in dorsal
and ventral series. Caudal peduncle relatively short, low, its
depth 65 (60–76) % of length.
Color of fresh specimen. Based on photograph of a malespecimen (Fig. 1). Ground color of head and body pale red,whitish ventrally. Three relatively narrow (width subequal
to iris diameter) dark red bands with narrow whitish margins on sides of head; anteriormost band from ventral
margin of orbit, running ventrally; middle band fromposteroventral margin of orbit, reaching obliquely ventral
margin of subopercle; posteriormost band saddling nape atlevel of central portion of parietal ridge, reaching posteroventral margin of opercle at level of upper origin of
pectoral-fin base. Both jaws yellowish; mandible andbranchiostegal membrane bright yellow. Eyes orange, iris
black, seemingly lacking radial markings. Flaps on posterior margin of preopercular yellow. Five vertical, moderately broad dark red bands (about four or five vertical scalerows width) saddling body; anteriormost band below firstto second dorsal-fin spine bases, extending posteroventrally
and continuous with a black blotch above pectoral-fin base;second band below fourth to fifth dorsal-fin spine bases;
third band below eighth to tenth dorsal-fin spine bases;
fourth band below first to third dorsal-fin soft ray bases;
fifth band below sixth to seventh or eighth dorsal-fin softray bases. A relatively large black blotch (slightly larger
than iris) above upper origin of pectoral-fin base.

123

Spinous dorsal fin coloration similar to ground color ofbody; soft-rayed portion of dorsal fin pale yellow, somewhat reddish basally. Anal fin pale yellow, somewhat
reddish basally. Pectoral fin dark yellow, somewhat reddish
basally, with numerous relatively large black blotches(larger than iris). Pelvic fin red, broadly blackish. Caudal
fin broadly bright yellow, paler basally, with ca. 6 small
black spots (distinctly smaller than iris) scattered basallyand dorsally.
Color of preserved specimens. Overall brownish white(or creamy white), slightly darker dorsally (Figs. 2–3);
posterior margin of scales or scale pockets blackish. Three
relatively narrow brown bands on sides of head; anteriormost band just below eye; middle band from posteroventral
margin of orbit, reaching obliquely ventral margin of
subopercle; posteriormost band saddling nape at level ofparietal ridge, reaching opercle at level of upper origin of
pectoral-fin base. Five moderately broad pale brown bands(ca. four vertical scale rows width) on sides of body (basedon paratypes; obscured in holotype); anteriormost band
below first and second dorsal-fin spine bases, runningposteroventrally, continuous with a dark brown or black
blotch above pectoral-fin base; second band below fourth
to sixth dorsal-fin spine bases; third band below eighth totenth dorsal-fin spine bases; fourth band below first to
second dorsal-fin soft ray bases; fifth band below sixth toeighth dorsal-fin soft ray bases. A relatively large darkbrown or black blotch just above pectoral-fin base (maximum diameter subequal to orbit diameter). Eyes darkish
blue, iris black.
In large males (including holotype) ([80 mm SL),dorsal and anal fin membrane translucent without spots (orentirely blackish in 82–86 mm SL specimens). Pectoral-fin
membrane translucent, blackish marginally, with ca. 9(6–11) relatively large, scattered black blotches (diameter
subequal to orbit diameter). Pelvic fin translucent, broadly
blackish, paler basally. Caudal-fin membrane translucent
without markings (or blackish posteriorly, with ca. 5 small
black spots on basal-dorsal portion in 82–86 mm SL
specimens). In large females ([70 mm SL), dorsal-fin
membrane translucent with ca. 19–23 small black spots
scattered on soft rays. Anal-fin membrane translucent with
0–5 small black spots scattered on soft rays. Pectoral-fin
membrane broadly blackish, paler dorsally and basally,
with ca. 10 indistinct relatively large scattered black blotches. Pelvic fin as in large males. Caudal-fin membranetranslucent with ca. 51–59 small black spots scattered overentire fin (or blackish posteriorly, with ca. 11 black spots
on basal-dorsal portion). In small specimens of both sexes(\70 mm SL), dorsal and anal fins entirely blackish without spots (or with ca. 25 and ca. 8–17 small black spots
scattered on soft rays of dorsal and anal fins, respectively).
Pectoral fin entirely blackish with ca. 5 indistinct blackNew Ebosia from southwestern Indian Ocean
Fig. 2 Preserved specimens ofEbosia vespertinus sp. nov.
a SAM 34265 (one of four
specimens), female, 93.5 mm
SL, Mozambique; b SAM
41190, holotype, male,
99.1 mm SL, Mozambique

blotches. Pelvic fin entirely dusky. Caudal fin blackish
posteriorly, with ca. 15–28 small black spots on basaldorsal portion.
Distribution. Ebosia vespertina has currently beenrecorded from off Mozambique and the east coast of South
Africa between latitudes 18° and 29° S, and southeastern
Madagascar in the southwestern Indian Ocean. Sampling
data accompanying six specimen lots examined indicatedcapture by bottom trawl at depths of 54–110 m. A photographed specimen (Fig. 1; not retained) was collected off
Tolagnaro(formerly
Fort-Dauphin),Madagascar(25°14.60 S, 47°09.10 E), from a depth of 79–80 m.
Etymology. The specific name, vespertina, derived from
Latin meaning western, alludes to the species having thewesternmost distribution within the genus.
Morphological changes with growth. Analyses of 44
measurements taken from 19 specimens (50.0–109.9 mmSL) of E. vespertina indicated that the relative body widthand depth, head width and depth, and snout length proportions tended to increase as a percentage of standard
length with growth. In contrast, orbit diameter and all fin
ray length proportions [except dorsal-fin spines (significantly damaged in most specimens examined)] became
significantly less with growth. The supraocular skin flap
became shorter (reduced in size) with growth in E. vespertina, as in congeners (Matsunuma and Motomura 2014).

Small specimens (\70 mm SL) had a relatively large
supraocular flap, 3.7–5.5 mm (5.8–8.4 % of SL), whereas
larger specimens ([90 mm SL) had a small flap,
2.1–5.0 mm (1.9–5.2 % of SL).
The number of pectoral-fin rays with more than threebranches tended to increase with growth in E. vespertina,
from usually absent (rarely 1) in small specimens (\80 mm
SL) to 4–9 (22–50 % of the total number of pectoral-finrays) in large specimens ([90 mm SL) (Fig. 4). However,an abnormal example of a relatively large specimen(93.5 mm SL; one of seven specimens of SAM 34265)
lacked such highly divided pectoral-fin rays. Nevertheless,the rate of increased pectoral-fin ray division in E. vespertina appeared to be greater than in E. falcata (Fig. 4).
The number of spinous points on many head spines
significantly increased with growth in E. vespertina (except
those associated with the coronal, parietal, pterotic, lower
posttemporal, supracleithral, opercular, preopercular, supplemental preopercular and anterior lacrimal spines). For
example, small specimens (\70 mm SL) possessed a single
spinous point on the preocular, 3–6 spines on the sphenotic
region, and 7–13 spines on the suborbital ridge, whereas
relatively large specimens ([90 mm SL) possessed 1–10,
9–13, and 18–35 spines, respectively. Furthermore, theparietal spine of males became more elongated with growth(Fig. 5b, c).

123M. Matsunuma, H. Motomura

Fig. 4 Relationships of pectoral-fin rays with more than threebranches as a percentage of number of total pectoral-fin rays tostandard length (mm) in Ebosia vespertina sp. nov. (red stars);
E. falcata (blue circles); and E. saya (green diamonds). Counted on
both sides whenever possible

Fig. 3 Preserved specimens of Ebosia vespertinus sp. nov., showing
black markings on specimens. a SAM 34265 (one of four specimens),
female, 58.9 mm SL, Mozambique; b KAUM–I. 75290, female,
82.9 mm SL, South Africa; c SAM 34265 (one of four), male,
106.1 mm SL, Mozambique (parietal spine broken; right side
reversed)

Sexual dimorphism and dichromatism. Sexual
dimorphism of parietal spine morphology in E. vespertina
was recognized on the basis of five male (70.5–106.1 mmSL) and six (58.9–96.0 mm SL) female specimens, their
sex having been determined by gonad examination (seeMatsunuma and Motomura 2014). In males, the parietal
spine becomes a thin elongated bony crest, strongly curved
posterodorsally with growth, its posterior margin reachinga vertical through the second to fifth dorsal-fin spine bases(Figs. 1, 2b, 5b, c). By contrast, all females examined had alow parietal ridge with a dull spinous margin (Figs. 2a, 5a).
Further sexual dimorphism included the third preopercularspine base, with females possessing a spineless ridge or asingle spine on the base and males usually a ridge with 1–5
small spines, and the number of supraocular spines, with 1
or 2 in females and 1–7 in males, although the number ofspines tended to increase with growth in both sexes.

123

Sexual dichromatism has been reported for E. bleekeri
and E. saya, wherein males possessed yellowish soft-rayed
portions of the dorsal and anal fins (plus a yellow caudal fin
in E. bleekeri and yellow pectoral fin in E. saya), whereas
females of both species possessed all red fins (Matsunuma
and Motomura 2014). Although life or fresh coloration of
female E. vespertina was undetermined in this study, males
possessed bright yellow soft-rayed portions of the dorsal
and anal fins, and yellow caudal and pectoral fins whenfresh (Fig. 1). Large preserved female specimens had
broadly blackish pectoral fins (Figs. 2a, 3a, b), which are
paler in males (membrane almost translucent; Figs. 2b, 3c),indicating that females also had darker pectoral fins thanmales in life. Considering the morphological similarities
between E. vespertina and E. saya, females of the former
likely possess red pectoral fins compared to yellow inmales. Moreover, the number of black spots on the softrayed portion of the dorsal and caudal fins also differs
between the sexes in E. vespertina. Large females ([80 mm SL) possessed 19–23 blackish spots on the dorsalfin soft-rayed portion and 11–59 blackish spots on thecaudal fin in contrast to males (0 and 0–8 spots, respectively). No difference was found between the sexes in thenumber of spots on the soft-rayed portion of the anal fin,tending to decrease with growth in both sexes.
Species comparisons. Among species of Ebosia,
E. vespertina is characterized by 17–18 (modally 18)
pectoral-fin rays, compared with 15–17 (modally 16) inNew Ebosia from southwestern Indian Ocean

Fig. 5 Lateral views of head of Ebosia vespertina sp. nov. (a:female; b–c: male), showing ontogenetic change and sexual difference of parietal spine. a SAM 34265 (one of four specimens),
93.5 mm SL; b SAM 34266 (one of six), 108.3 mm SL (right side
reversed); c SAIAB 76118, 86.0 mm SL. Bars and arrowheads
indicate 5 mm and parietal spine, respectively

E. bleekeri and 16–19 (modally 17) in E. falcata andE. saya (see Matsunuma and Motomura 2014; Table 1).
Although a single specimen of E. falcata (ZMH 5581)
possessed 19 and 17 pectoral-fin rays on the left and rightsides of the body, respectively, it was considered to represent an abnormal condition. Moreover, E. vespertina canbe distinguished from E. bleekeri by having 7–8 (modally8) anal-fin soft rays [versus 6–8 (modally 7) in E. bleekeri];
the elongated parietal spine in males narrow and strongly
curved posterodorsally, the width of the elongated portion
clearly less than the pupil diameter (versus less falcate
posterodorsally and relatively broad, its width ca. 1.0–1.5
times pupil diameter); and a yellowish pectoral fin in males(versus entirely red) (Matsunuma and Motomura 2014; thisstudy).
Ebosia vespertina is closely related to E. falcata andE. saya in sharing modally 8 anal-fin soft rays, and anarrow, strongly curved parietal spine and broadly yellowish pectoral fin in males (the latter when fresh) (Matsunuma and Motomura 2014; this study). However, in
addition to the above-mentioned difference in the numberof pectoral-fin rays, subtle differences exist in the numberof longitudinal series scale rows and gill rakers between
E. vespertina and the latter two species (Table 1), viz.,
scale rows 47–51 (modally 49 or 50) in E. vespertina
versus 41–48 (44) in E. falcata and 44 or 46 in E. saya
(only two specimens available); upper gill rakers 4–6 (4)versus 4–6 (5) and 4 or 5 (5); lower gill rakers 10 or 11 (10)versus 10–12 (11) and 10 or 11 (11); and total gill rakers
14–17 (14) versus 14–17 (16) and 15 or 16 (15). Ebosia
vespertina also differs from E. falcata by having usually
one more scale row above the lateral line [4–6 (modally 5)
in E. vespertina versus 4–5 (4) in E. falcata]; scale rows
between the last dorsal-fin spine base and lateral line [4–5(5) versus 4]; scale rows between the last dorsal-fin spinebase and lateral line [4–5 (5) versus 3–5 (4)]; and a shortpostorbital length [17.4–20.3 (mean 18.7) % of SL versus
19.8–24.2 (21.9) % of SL] (Table 1; Fig. 6a, d). Furthermore, E. vespertina can be distinguished from E. saya by
the longer pectoral fin [longest ray length 54.6–73.9 (mean65.1) % of SL in E. vespertina versus 55.9–57.8 (56.5) %of SL in E. saya]; longer pelvic fin [longest pelvic-fin soft
ray length 32.4–44.0 (37.6) % of SL versus 30.8–37.9(33.3) % of SL]; a larger blotch above the pectoral-fin baseand blotches on the pectoral fin membrane [maximum
diameter of blotch above pectoral-fin base 54.7–99.7 (77.7)% of orbit diameter (OD) in E. vespertina (n = 14) versus
32.6–56.2 (41.2) % of OD in E. saya (n = 6); that of
blotches on the pectoral-fin membrane 25.6–42.0 (34.6) %of OD versus 13.5–34.5 (21.7) % of OD in E. saya (n =10)] (Figs. 6b, c, e, f, 7). In addition, relationships between
body size and number of pectoral-fin rays with more than
three branches may be helpful for the identification of large
examples ([80 mm SL) of E. vespertina and E. falcata(Fig. 4).
Padate et al. (2014) recently recorded three specimens ofE. falcata from the Gulf of Manner (off southern India) andnoted the fresh coloration of one of the specimens. Since
the photograph included (Padate et al., 2014: fig. 2C)
showed an elongated parietal spine and yellowish pectoral,
soft-rayed portions of dorsal and anal, and caudal fins, thespecimen was here regarded to have been a male.
Comparative materials. Ebosia bleekeri (111 specimens), E. falcata (38) and E. saya (10) listed in Matsunuma
and Motomura (2014) with the following additions. Ebosia
bleekeri: KAUM–I. 65441, 90.9 mm SL, female (G), Tosa
Bay, Kochi Prefecture, Japan, BSKU, bottom trawl, 17
December 2012; MZS Pis892, female (P), 110.2 mm SL,
syntype of Pterois bleekeri, Tokyo, Japan; NMW 75648,
male (P), 109.3 mm SL, syntype of P. bleekeri, Tokyo,
Japan; SNFR 13580, male (G), 124.6 mm SL, East China
Sea, bottom trawl, 2 June 2008; SNFR 13589, male (G),96.3 mm SL, SNFR 13590, male (G), 93.7 mm SL, East
China Sea, bottom trawl, 31 May 2008; SNFR 14108, male123

M. Matsunuma, H. Motomura

Fig. 6 Relationships of (a, d) postorbital length, (b, e) longest
pectoral-fin ray length and (c, f) longest pelvic-fin soft ray (as % ofstandard length in d–f) to standard length (mm) in Ebosia vespertina

Fig. 7 Relationships of maximum diameter of blotch on pectoral-fin
membrane to maximum diameter of blotch above pectoral-fin base(mm) in Ebosia vespertina sp. nov. (red stars); E. falcata (bluecircles); and E. saya (green diamonds). Arrowheads indicate holotype(G), 98.2 mm SL, East China Sea, bottom trawl, 1 June
2008. Ebosia falcata: ZSI F 7400/2, sex undetermined,
56.0 mm SL, paratype of Ebosia falcata, off Somalia(11°040 N, 51°150 E), 76–80 m, bottom trawl, H. Fehlmann
onboard RV Anton Bruun, 17 December 1964.

123

sp. nov. (red stars; dotted lines); E. falcata (blue circles; solid lines);and E. saya (green diamonds; dashed lines). Arrowheads indicate
holotype

Acknowledgments We are deeply grateful to R. Bills, O. Gon and
E. Heemstra (SAIAB); M. Bougaardt and D. Clarke (SAM); M.
McGrouther, A. Hay and S. Reader (AMS); K.-T. Shao, Y.-C. Liaoand M.-Y. Lee (ASIZP); H. Endo, T. Yamakawa, N. Nakayama, R.
Asaoka (BSKU), E. Katayama and R. Misawa (formerly BSKU); Y.
Kai, A. Tokairin (FAKU), T. Nakabo 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);
M. Meister and M.-D. Wandhammer (MZS); Z. Arifin, R. Pratiwi, M.
Adrim, I. Alhakim and K. Wibowo (NCIP); H.-C. Ho (NMMB); A.
Palandacic (NMW); 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); K.
Gopi and S. Mishra (ZSI); and K. Sakamoto (ZUMT) for their kind
hospitality during the first author’s visits to their institutions. We areespecially grateful to E. Heemstra (SAIAB) for providing the photograph of the Madagascar specimen of E. vespertina which wastaken by M. Lee during the 2010 Atimo Vatae expedition to south
Madagascar [principal investigator, P. Bouchet (MNHN)]. We arealso grateful to R. Feeney (LACM) for providing opportunities to
examine specimens; H.-C. Ho (MNHB) and H. Endo, N. Nakayama,
R. Misawa and other students of BSKU for donating specimens; andstudents and volunteers of KAUM for curatorial assistance. G. Hardy(Ngunguru, New Zealand) read the manuscript and provided help
with English. This study was supported in part by Grants-in-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 ofResearch and Education Network on Coastal Marine Science inSoutheast Asia’’, JSPS International Training Program ‘‘Protect
Diversity of Bioresources in the Tropical Area’’, the Coastal Area
Capability Enhancement in Southeast Asia Project of the ResearchInstitute for Humanity and Nature, Kyoto, Japan the ‘‘Biological
Properties of Biodiversity Hotspots in Japan’’ project of the National
Museum of Nature and Science, Tsukuba, Japan, and by a grantawarded to the first author from the Mikimoto Fund for Marine
Ecology.

New Ebosia from southwestern Indian Ocean

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