RAFFLES BULLETIN OF ZOOLOGY 2021
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 69: 109–117

Date of publication: 26 March 2021
DOI: 10.26107/RBZ-2021-0009
http://zoobank.org/urn:lsid:zoobank.org:pub:BE23B2D5-6216-4293-8FBA-33F565F2D7BF

Two new species of Stolephorus (Teleostei: Clupeiformes: Engraulidae)
from the western PacificHarutaka Hata1* & Hiroyuki Motomura2
Abstract. Stolephorus concursus, new species, and Stolephorus celsior, new species, both characterised by the
maxilla extending beyond the preopercle posterior margin and numerous black spots on the suborbital area, snout,and lower-jaw tip, are described on the basis of specimens from the southwestern Pacific Ocean and Manila Bay,
Luzon, Philippines, respectively. Although the former and Stolephorus bataviensis Hardenberg, 1933 both havethe pelvic fin extending to below the dorsal-fin origin, S. concursus, new species, has greater numbers of total gillrakers on the first and second gill arches (38–41 and 30–34, respectively vs. 33–38 and 27–32 in the latter), andlonger pectoral fin (16.5–18.1% SL vs. 14.7–16.4%) and snout (4.8–5.4% SL vs. 3.7–4.7%). Stolephorus celsior,new species, is similar to both Stolephorus babarani Hata, Lavoué & Motomura, 2020 and Stolephorus baweanensis
Hardenberg, 1933 in having the pelvic fin not extending to below the dorsal-fin origin, but differs from them ingenerally having a greater number of total gill rakers on the first gill arch (40 or more vs. 41 or fewer in the latter
two), in addition to longer pectoral and pelvic fins (> 16.6% SL and 9.5% SL, respectively vs. < 16.8% and 9.4%).
Key words. Actinopterygii, Stolephorus concursus, Stolephorus celsior, morphology, taxonomy, anchovy

INTRODUCTION

MATERIAL AND METHODSSpecies of the genus Stolephorus (Clupeiformes: Engraulidae)
with numerous spots on the suborbital area, snout,and lower-jaw tip were regarded as a single species(Stolephorus waitei Jordan & Seale, 1926) by Whitehead
et al. (1988). However, following molecular analyses andmorphological examinations, Hata et al. (2019) revealed
that S. waitei sensu Whitehead et al. (1988) in fact included
three species, Stolephorus bataviensis Hardenberg, 1933,
Stolephorus baweanensis Hardenberg, 1933, and true S.
waitei. Subsequently, Stolephorus babarani Hata, Lavoué &Motomura, 2020, a species also characterised by numerous
black spots, was described as a new species, based on
specimens collected from Panay, Philippines (Hata etal., 2020a). Recently, a number of specimens from thesouthwestern Pacific (Fiji and Tonga) and Manila Bay,
Luzon, Philippines, all having numerous black spots andrepresenting separate species in each locality, were identified
as new to science. Both new species, characterised by greatergill raker numbers than in other species of Stolephorus so
far known, are described herein.

Counts and proportional measurements followed Hata &Motomura (2017). All measurements were made with digitalcallipers to the nearest 0.01 mm. Standard and head lengths
are abbreviated as SL and HL, respectively. Osteologicalcharacters, including vertebral counts, were observed onradiographs of 17 specimens of S. celsior and 23 S. concursus.
Counts and measurements, expressed as percentages ofSL or HL, are given in Tables 1 and 2. Institutional codes
follow Sabaj (2019).
TAXONOMY
Stolephorus celsior, new species[New English name: Rosario Anchovy](Fig. 1; Tables 1, 2)
Holotype. BMNH 1966.1.17.93, 60.0 mm SL, Rosario(formerly Salinas), Cavite, Manila Bay, Luzon, Philippines.
Paratypes. 15 specimens, 48.2–64.2 mm SL. BMNH
1966.1.17.94–97, 4 specimens, 56.1–60.4 mm SL, KAUM–I.
146837, 55.8 mm SL, KAUM–I. 146838, 61.0 mm SL,
NSMT-P 139183, 64.2 mm SL, NSMT-P 139184, 57.4 mm
SL, collected with holotype; BMNH 1966.1.17.135–142, 8
specimens, 48.2–57.5 mm SL, same locality as holotype.

Accepted by: Zeehan Jaafar
Center for Molecular Biodiversity Research, National Museum of Nature andScience, 4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan; Email: k2795502@
kadai.jp (*corresponding author)
1

Diagnosis. A species of Stolephorus with the following
combination of characters: long maxilla, 22.1–23.2% SL(mean 22.6%), its posterior tip slightly short of or justreaching posterior margin of opercle; no predorsal scute;

The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan;
Email: motomura@kaum.kagoshima-u.ac.jp
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Hata & Motomura: Two new species of the genus Stolephorus
Table 1. Meristics of two new species of the genus Stolephorus.
Stolephorus celsior, new species
Holotype(BMNH
1966.1.17.93)

Paratypes(n = 15)

Manila Bay, Luzon,
Philippines
Standard length (SL; mm)

Stolephorus concursus, new species
Holotype(USNM
330901)Paratypes(n = 23)

Viti Levu
Island, Fiji

Fiji and
Tonga

60.0

48.2–64.2

Modes

72.8

68.7–84.7

Modes

Dorsal-fin rays (unbranched)

3

3

3

3

3

3

Dorsal-fin rays (branched)

13

12–14

13

13

11–14

13

Anal-fin rays (unbranched)

3

3

3

3

3

3

Anal-fin rays (branched)

19

16–20

18

18

17–20

19

Pectoral-fin rays (unbranched)

1

1

1

1

1

1

Pectoral-fin rays (branched)

14

12–13

12

13

12–15

13

Pelvic-fin rays (unbranched)

1

1

1

1

1

1

Pelvic-fin rays (branched)

6

6

6

6

6

6

Gill rakers on 1st gill arch (upper)

19

17–19

18

18

16–18

17

Gill rakers on 1st gill arch (lower)

24

23–25

23

23

21–23

23

Gill rakers on 1st gill arch (total）

43

40–44

42

41

38–41

39

Gill rakers on 2nd gill arch (upper)

13

11–13

12

13

11 or 1212Gill rakers on 2nd gill arch (lower)

22

20–22

21

21

19–22

20

Gill rakers on 2nd gill arch (total)

35

32–35

32

34

30–34

32

Gill rakers on 3rd gill arch (upper)

10

9–11

10

10

9 or 109Gill rakers on 3rd gill arch (lower)

12

12 or 13

12

12

10–12

12

Gill rakers on 3rd gill arch (total)

22

21–23

22

22

19–22

21

Gill rakers on 4th gill arch (upper)

10

8 or 9

8

9

7–9

8

Gill rakers on 4th gill arch (lower)

11

10 or 11

10

10

9–11

10

Gill rakers on 4th gill arch (total)

21

18–20

18

19

16–19

18

Gill rakers on posterior face of 3rd gill arch

6

5–7

6

5

4–6

5

Prepelvic scutes

6

4–6

6

6

3–6

6

Scale rows in longitudinal series

34

34 or 35

34

35

35–37

36

Transverse scales

8

8

8

8

8 or 98

Pseudobranchial filaments

18

17–20

19

22

19–25

22

Abdominal vertebrae

19

18–20

19

20

19 or 20

20

Caudal vertebrae

20

19 or 2019

20

20 or 21

20

Total vertebrae

39

37–39

39

40

40 or 41

40

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Table 2. Morphometrics of two new species of the genus Stolephorus.
Stolephorus celsior, new species
Holotype(BMNH
1966.1.17.93)

Stolephorus concursus, new species

Paratypes(n = 15)

Manila Bay, Luzon, Philippines
Standard length (SL; mm)

Holotype(USNM 330901)

Paratypes(n = 23)

Viti Levu
Island, Fiji

Fiji and
Tonga

60.0

48.2–64.2

Means

72.8

68.7–84.7

Means

Head length (HL)

26.6

26.0–27.5

26.6

27.0

26.4–28.5

27.4

Body depth

22.0

20.3–22.2

21.2

20.3

19.4–21.7

20.5

Pre-dorsal-fin length

56.6

54.6–58.0

56.0

54.6

53.7–57.2

55.2

Snout tip to pectoral-fin insertion

29.3

27.2–29.8

28.5

29.1

27.8–30.8

29.6

Snout tip to pelvic-fin insertion

46.7

44.5–48.0

46.0

47.9

46.1–48.8

47.7

Snout tip to anal-fin origin

65.3

62.4–66.9

64.4

65.5

63.0–66.6

64.9

Dorsal-fin base length

14.8

14.1–16.7

15.6

15.9

15.0–17.3

16.0

Anal-fin base length

21.2

18.0–23.3

20.6

19.5

18.9–21.7

20.7

Caudal-peduncle length

17.0

15.0–18.7

17.3

16.5

14.1–18.4

16.5

Caudal-peduncle depth

10.4

10.1–11.1

10.5

9.3

8.8–10.8

9.4

Orbit diameter

9.0

8.7–9.3

9.0

8.9

7.8–9.7

8.9

Eye diameter

7.3

6.8–8.1

7.3

7.5

6.4–7.8

7.4

Snout length

4.3

4.0–4.5

4.3

5.0

4.8–5.4

5.0

D–P1

33.7

32.5–36.0

34.4

33.8

33.6–34.7

34.0

D–P2

24.3

23.5–25.3

24.4

21.6

21.0–24.1

22.5

D–A

21.8

20.5–23.3

22.2

22.1

21.8–23.6

22.6

P1–P2

17.8

16.5–21.8

18.1

20.0

17.3–20.6

19.3

P2–A

18.7

16.6–21.0

18.2

17.5

15.9–19.2

17.4

Pectoral-fin length

17.0

16.6–17.9

17.2

17.0

16.5–18.1

17.0

Pelvic-fin length

10.4

9.5–10.2

9.8

10.2

9.5–10.6

10.1

Maxilla length

22.4

22.1–23.2

22.6

23.3

22.5–25.2

23.7

Mandibular length

18.3

17.9–18.5

18.1

19.6

19.1–20.7

19.9

Supramaxilla end to maxilla end

6.1

5.1–6.1

5.5

5.1

4.8–5.6

5.2

Orbit diameter

33.9

33.0–34.6

33.8

32.8

29.4–36.6

32.7

Eye diameter

27.4

24.9–29.6

27.3

27.9

24.2–29.4

26.9

Snout length

16.2

14.8–16.5

16.1

18.6

17.0–19.4

18.4

Interorbital width

22.1

21.2–24.1

22.3

20.8

18.4–21.0

20.1

Postorbital length

50.7

46.9–52.8

50.3

49.4

48.9–58.1

51.7

D–P1

126.8

118.1–138.2

129.2

125.3

117.8–131.3

124.4

D–P2

91.2

85.6–95.2

91.5

80.0

77.6–87.4

82.2

D–A

82.0

75.7–87.7

83.2

81.8

78.9–87.4

82.7

P1–P2

66.8

61.5–83.8

68.0

74.3

61.9–77.7

70.6

P2–A

70.1

63.6–76.9

68.7

64.7

57.3–71.8

63.7

As % HL

Abbreviations: D–P1 (distance from dorsal-fin origin to pectoral-fin insertion); D–P2 (distance from dorsal-fin origin to pelvic-fin insertion);D–A (distance between origins of dorsal- and anal fins); P1–P2 (distance between insertions of pectoral- and pelvic fins); P2–A (distance
between pelvic-fin insertion to anal-fin origin).

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Hata & Motomura: Two new species of the genus Stolephorus

Fig. 1. (A) Lateral view of body and (B) dorsal and (C) ventral
view of head of holotype of Stolephorus celsior, new species,
BMNH 1966.1.17.93, 60.0 mm SL, Rosario, Cavite, Manila Bay,
Luzon, Philippines.

pelvic scute without spine; gill rakers 17–19 (modally 18)
in upper series on first gill arch, 23–25 (23) in lower series,
40–44 (42) in total; gill rakers 11–13 (12) in upper series on
second gill arch, 20–22 (21) in lower series, 32–35 (32) intotal; gill rakers 9–11 (10) in upper series on third gill arch,
12 or 13 (12) in lower series, 21–23 (22) in total; gill rakers
8–10 (8) in upper series on fourth gill arch, 10 or 11 (10) inlower, 18–21 (18) in total; gill rakers 5–7 (6) on hind faceof third gill arch; prepelvic scutes 4–6 (6); transverse scales
8; pseudobranchial filaments 17–20 (19); paired dark patcheson parietal and occipital regions without a following pair ofdark lines; numerous black spots on snout and lower-jaw tip,
rarely on suborbital area; depressed pelvic fin not reaching tovertical through dorsal-fin origin; snout rather long, 4.0–4.5%
SL (mean 4.3%); pectoral fin rather long, 16.6–17.9% SL(17.2%); pelvic fin long, 9.5–10.4% SL (9.8%); head large,
26.0–27.5% SL (26.6%); orbit large, 8.7–9.3% SL (9.0%);
lower jaw rather short, 17.9–18.5% SL (18.1%).

Fig. 2. Relationships of total gill-raker numbers (TGR) on (A) firstgill arch (1GA), (B) second gill arch (2GA), (C) third gill arch,and (D) fourth gill arch relative to standard length in Stolephorus
celsior, new species (triangles), S. concursus, new species (circles),S. babarani (inverted triangles), S. bataviensis (squares), and S.
baweanensis (diamonds).

insertion, thereafter nearly straight and parallel to body axis
and ventral profile of body from anal-fin origin to caudalfin base slowly rising. Abdomen rounded, covered with six(four to six) spine-like scutes. Postpelvic and predorsal scutes
absent. Anus just anterior to anal-fin origin. Snout tip rounded;
snout length less than eye diameter. Mouth large, inferior,
ventral to body axis, extending backward beyond posterior
margin of eye. Maxilla long, its posterior tip pointed, justshort of or reaching posterior border of opercle. Lower jaw
slender. Single row of conical teeth on each jaw and palatines.
Small conical teeth patch on pterygoids. Several small conical

Description. Data for holotype presented first, followedby paratype data in parentheses (if different). Counts andmeasurements, expressed as percentages of SL or HL, given
in Tables 1 and 2.
Body laterally compressed, elongate, deepest at dorsal-fin
origin. Dorsal profile of head and body gently elevated fromsnout tip to dorsal-fin origin, thereafter gradually lowering touppermost point of caudal-fin base. Ventral profile of headand body slightly convex from lower-jaw tip to pelvic-fin
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Fig. 3. Morphometrics and meristic of Stolephorus celsior, new species (triangles), S. concursus, new species (circles), S. babarani (inverted
triangles), S. bataviensis (squares), and S. baweanensis (diamonds) (all vs. standard length; SL). A, ratio of caudal-peduncle depth toSL; B, pectoral-fin length to SL; C, mandibular length to SL; D, pelvic-fin length to SL; E, distance from dorsal-fin origin to pelvic-fin
insertion to SL; F, head length to SL; G, orbit diameter to SL; H, snout length to SL; I, maxilla length to SL.

pupil diameter, from just behind upper opercular margin to
caudal-fin base. Pairs of dark patches on parietal and occipitalregions. No dark lines on dorsum. Numerous black spots
on snout and lower-jaw tip (sometimes on suborbital area).
Melanophores scattered on posterior margin of dorsal scale
pockets. All fins whitish, semi-transparent. Melanophores
scattered along fin rays of dorsal and caudal fins. Live andfresh colouration unknown.

teeth on vomer. No teeth on dorsal surface of hyoid. Eyelarge, round, covered with adipose eyelid, positioned laterally
on head dorsal to horizontal through pectoral-fin insertion,
visible in dorsal view. Pupil round. Orbit elliptical. Nostrils
close to each other, anterior to orbit. Posterior margins ofpreopercle and opercle rounded, smooth. Subopercle withrounded posterior margin. Posterior margin of preopercle
slightly concave (moderately concave in some paratypes),
indented. Gill membrane without serrations. Interorbital space
flat, width less than eye diameter. Pseudobranchial filaments
present, longest filament shorter than eye diameter. Gill rakers
long, slender, rough, visible from side of head when mouth
opened. Isthmus muscle long, reaching anteriorly to posterior
margin of gill membranes. Urohyal hidden by isthmus muscle(visible only following dissection). Gill membrane on eachside joined distally, most of isthmus muscle exposed (notcovered by gill membrane). Head scales and lateral line
absent. Fins scaleless, except for broad triangular sheath ofscales on central part of both lobes of caudal fin. Dorsal-fin
origin posterior to vertical through base of last pelvic-fin ray,slightly posterior to midpoint of body. Dorsal and anal finswith three anteriormost rays unbranched and closely spaced,first ray of both fins minute. Anal-fin origin just below baseof eighth (seventh to ninth) dorsal-fin ray. Posterior tip ofdepressed anal fin not reaching caudal-fin base. Uppermost
pectoral-fin ray unbranched, inserted below midline of body.
Posterior tip of pectoral fin not reaching vertical throughpelvic-fin insertion; pelvic fin shorter than pectoral fin;pelvic-fin insertion anterior to vertical through dorsal-finbase. Posterior tip of depressed pelvic fin not reaching tovertical through dorsal-fin origin.

Distribution. Currently known only from the southern partof Manila Bay, Luzon, Philippines (Rosario, Cavite).
Etymology. The specific name celsior, a Latin adjective,refers to the higher gill raker counts in the new species
compared to similar species.
Comparisons. The new species is assignable to the genus
Stolephorus, as defined by Whitehead et al. (1988) and
Wongratana et al. (1999), having a long isthmus muscle
reaching anteriorly to the posterior margin of the gillmembrane, the urohyal covered by muscle, and prepelvic
scutes, in addition to the absence of postpelvic scutes.
Stolephorus celsior, new species, differs from all othercongeners, except for S. babarani, S. baweanensis, S.
bataviensis, and S. concursus, new species (described herein)in having a long upper jaw, its posterior tip beyond thepreopercle posterior margin (slightly short of or extending
beyond the opercle posterior margin) and numerous dusky
spots on the snout and lower-jaw tip (Whitehead et al.,
1988; Wongratana et al., 1999; Kimura et al., 2009; Hata
& Motomura, 2018a–d; Hata et al., 2019, 2020a, b, 2021;
Gangan et al., 2020; this study). However, S. celsior has agreater number of total gill rakers on each gill arch (e.g.,
40–44 on first gill arch in S. celsior vs. 41 or fewer in the

Colour of preserved specimens. Body uniformly pale ivory
with whitish longitudinal band, width slightly less than113

Hata & Motomura: Two new species of the genus Stolephorus
Table 3. Frequency distribution of total vertebrae in Stolephorus celsior, new species, S. concursus, new species, S. babarani, S. bataviensis,
and S. baweanensis.
Total vertebrae

Stolephorus celsior

n = 17

Stolephorus concursus

n = 23Stolephorus babaranin = 23Stolephorus bataviensis
Stolephorus baweanensis

37

38

39

1

7

9

1

40

41

22

1

21

1

n = 19

4

14

n =9

8

1

1

other four species; Table 1; Fig. 2A) and a generally deeper
caudal peduncle (10.1–11.1% of SL vs. < 10.3%; Table 1;
Fig. 3A). Moreover, the new species differs distinctly from S.
bataviensis and S. concursus in having the posterior tip of thedepressed pelvic fin not reaching to below the dorsal-fin origin(vs. beyond vertical through dorsal-fin origin). Stolephorus
celsior further differs from S. bataviensis in having a longer
pectoral fin (16.6–17.9% SL vs. 14.7–16.4%; Table 2; Fig.
3B) and mandible (17.9–18.5% SL vs. 18.0–20.2%; Table
2; Fig. 3C), and lower vertebral number (total vertebrae
18–20 + 19 or 20 = 37–39 vs. 20 or 21 + 19–21 = 39 or
40; Tables 1, 3). Detailed comparisons between S. celsiorand S. concursus are given under Comparisons of the latter.
Although S. celsior, S. babarani, and S. baweanensis all havethe pelvic fin extending to below the dorsal-fin origin, thepectoral fin (16.6–17.9% SL) is longer in S. celsior (vs. <16.8% in the other species; Table 2; Fig. 3B). Additionally,
the pelvic fin (9.5–10.4% SL) is longer (vs. < 9.4%; Table
2; Fig. 3D), and the dorsal-fin origin to pelvic-fin insertion
distance (23.5–25.3% SL vs. < 24.0%; Table 2; Fig. 3E)
generally greater in S. celsior. The latter also differs fromS. babarani in its larger head (26.0–27.5% SL in S. celsior
vs. 23.9–25.5% in S. babarani; Table 2; Fig. 3F) and orbit(8.7–9.3% SL vs. 7.6–8.4%; Table 2; Fig. 3G), and longer
snout (4.0–4.4% SL vs. 3.6–3.9%; Table 2; Fig. 3H) andmaxilla (22.1–23.2% SL vs. 20.8–22.3%; Table 2; Fig.
3I). Moreover, the vertebral number in S. celsior tends
to be lower than in S. baweanensis (18–20 + 19 or 20 =37–39 in S. celsior vs. 19 or 20 + 19 or 20 = 39 or 40 in
S. baweanensis; Tables 1, 3).

Fig. 4. (A) Body and (B) head of holotype of Stolephorus concursus,
new species, USNM 330901, 72.8 mm SL, Nadi Bay, Viti Levu
Island, Fiji.

NSMT-P 139186, 73.0 mm SL, collected with holotype;
USNM 330906, 13 specimens, 69.7–74.6 mm SL, Fiji.
Diagnosis. A species of Stolephorus with the following
combination of characters: long maxilla, 22.5–25.2% SL(mean 23.7%), its posterior tip slightly short of or extending
beyond posterior margin of opercle; no predorsal scute;pelvic scute without spine; gill rakers 16–18 (modally 17)
in upper series on first gill arch, 21–23 (23) in lower series,
38–41 (39) in total; gill rakers 11–13 (12) in upper series
on second gill arch, 19–22 (20) in lower series, 30–34 (32)in total; gill rakers 9 or 10 (9) in upper series on third gillarch, 10–12 (12) in lower series, 19–22 (21) in total; gillrakers 7–9 (8) in upper series on fourth gill arch, 9–11 (10)in lower, 16–19 (18) in total; gill rakers 4–6 (5) on hind faceof third gill arch; prepelvic scutes 3–6 (6); transverse scales
8 or 9 (8); pseudobranchial filaments 19–25 (22); paired dark
patches on parietal and occipital regions without a following
pair of dark lines; numerous black spots on suborbital area,
snout, and lower-jaw tip; depressed pelvic fin extending

Stolephorus concursus, new species[New English name: Concourse Anchovy](Fig. 4; Tables 1, 2)
Holotype. USNM 330901, 72.8 mm SL, Nadi Bay, Viti
Levu Island, Fiji, 4 July 1973.
Paratypes. 23 specimens, 68.7–84.7 mm SL. BPBM 38338,
2 specimens, 82.2–84.7 mm SL, Pangaimotu, Vava’u, Tonga;
URM-P. 33477, 84.7 mm SL, Vava’u Bay, Vava’u Island,
Vava’u Islands, Tonga; USNM 451515, 3 specimens, 68.9–
73.4 mm SL, KAUM–I. 146839, 75.8 mm SL, KAUM–I.
146840, 79.4 mm SL, NSMT-P 139185, 75.1 mm SL,
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Colour of preserved specimens. Body uniformly paleivory with whitish longitudinal band, slightly less than pupil
diameter, from just posterior to upper opercular margin to
caudal-fin base. Pairs of dark patches on parietal and occipitalregions. No dark lines on dorsum. Numerous black spots
on snout, lower-jaw tip, and suborbital area. Melanophores
scattered on posterior margin of dorsal scale pockets. All
fins whitish, semi-transparent. Melanophores scattered along
fin rays of dorsal and caudal fins. Posterior margin of caudal
fin dark. Live and fresh colouration unknown.

posteriorly beyond vertical through dorsal-fin origin; snout
long, 4.8–5.4% SL (mean 5.0%); pectoral fin rather long,
16.5–18.1% SL (17.0%); pelvic fin long, 9.5–10.6% SL(10.1%); head large, 26.4–28.5% SL (27.4%); lower jaw
long, 19.1–20.7% SL (19.9%).
Description. Data for holotype presented first, followedby paratype data in parentheses (if different). Counts andmeasurements, expressed as percentages of SL or HL, given
in Tables 1 and 2.

Distribution. Currently known only from Tonga and Fiji.

Body laterally compressed, elongate, deepest at dorsal-fin
origin. Dorsal profile of head and body gently elevated fromsnout tip to dorsal-fin origin, thereafter gradually lowering touppermost point of caudal-fin base. Ventral profile of headand body slightly convex from lower-jaw tip to pelvic-fin
insertion, thereafter nearly straight and parallel to body axis
and ventral profile of body from anal-fin origin to caudal
fin base slowly rising. Abdomen rounded, covered with six(three to six) spine-like scutes. Postpelvic and predorsal
scutes absent. Anus just anterior to anal-fin origin. Snout
tip rounded; snout length less than eye diameter. Mouth
large, inferior, ventral to body axis, extending backward
beyond posterior margin of eye. Maxilla long, its posterior
tip pointed, just short of or reaching posterior border of
opercle. Lower jaw slender. Single row of conical teethon each jaw and palatines. Small conical teeth patch on
pterygoids. Several small conical teeth on vomer. Eye large,
round, covered with adipose eyelid, positioned laterally onhead dorsal to horizontal through pectoral-fin insertion,
visible in dorsal view. Pupil round. Orbit elliptical. Nostrils
close to each other, anterior to orbit. Posterior margins ofpreopercle and opercle, smooth. Subopercle with rounded
posterior margin. Posterior margin of preopercle indented,
convex (rounded, concave in one paratype). Gill membranewithout serrations. Interorbital space flat, width less than eye
diameter. Pseudobranchial filaments present, longest filament
shorter than eye diameter. Gill rakers long, slender, rough,visible from side of head when mouth opened. Isthmus
muscle long, reaching anteriorly to posterior margin of gillmembranes. Urohyal hidden by isthmus muscle (visible only
following dissection). Gill membrane on each side joined
distally, most of isthmus muscle exposed (not covered bygill membrane). Scales cycloid, thin, deciduous, except
for prepelvic scutes. Numerous vertical grooves on bodyscales. Head scales and lateral line absent. Fins scaleless,except for broad triangular sheath of scales on caudal fin.
Dorsal-fin origin posterior to vertical through base of last
pelvic-fin ray, slightly posterior to midpoint of body. Dorsal
and anal fins with three anteriormost rays unbranched andclosely spaced, first ray of both fins minute. Anal-fin origin
just below base of eleventh (ninth to twelfth) dorsal-fin ray.
Posterior tip of depressed anal fin not reaching caudal-finbase. Uppermost pectoral-fin ray unbranched, inserted below
midline of body. Posterior tip of pectoral fin not reachingvertical through pelvic-fin insertion; pelvic fin shorter thanpectoral fin; pelvic-fin insertion anterior to vertical throughdorsal-fin base. Posterior tip of depressed pelvic fin reaching
to vertical through third (first to sixth) dorsal-fin ray origin.

Etymology. The specific name concursus is derived from
Latin meaning “crowd”, in reference to the schooling habitof the species.
Comparisons. Included in the genus Stolephorus sensu
Whitehead et al. (1988) and Wongratana et al. (1999)(see “Comparisons” under S. celsior), the new species iseasily distinguishable from all other congeners—except
for S. babarani, S. bataviensis, S. baweanensis, and S.
celsior, new species—by the long upper jaw, its posterior
tip beyond the preopercle posterior margin (slightly shortof or extending beyond the opercle posterior margin), andnumerous dusky spots on the suborbital area, snout, andlower-jaw tip (Whitehead et al., 1988; Wongratana et al.,
1999; Kimura et al., 2009; Hata & Motomura, 2018a–d;
Hata et al., 2019, 2020a, b, 2021; Gangan et al., 2020; thisstudy). However, the former is easily distinguished from S.
babarani, S. baweanensis, and S. celsior by the posterior
tip of the depressed pelvic fin extending beyond verticalthrough the dorsal-fin origin (vs. not reaching to below thedorsal fin origin in the other three species), a longer snout(4.8–5.4% SL in S. concursus vs. < 4.6% SL; Table 2; Fig.
3H) and mandible (19.1–20.7% SL vs. < 19.0%; Table 2;
Fig. 3C), and higher vertebral number [total vertebrae 40
or 41 (modally 40) vs. 39 or 40 (39) in S. baweanensis,
39 or fewer in S. babarani and S. celsior; Tables 1, 3].
Stolephorus concursus further differs from S. babarani andS. baweanensis in having a longer head (26.4–28.5% SL vs.
23.9–25.5% in S. babarani; 23.9–27.1% in S. baweanensis;
Table 2; Fig. 3F), pectoral fin (16.5–18.1% SL vs. 15.0–16.3%
in S. babarani; 14.0–16.8% SL in S. baweanensis; Table 2;
Fig. 3B), and pelvic fin (9.5–10.6% SL vs. 8.1–9.4% in S.
babarani; 6.5–9.4% in S. baweanensis; Table 2; Fig. 3D),
and from S. babarani in maxilla length (22.5–25.2% SL in
S. concursus vs. 20.8–22.3% in S. babarani; Table 2; Fig.
3I). Stolephorus concursus is distinguished from S. celsior
by higher longitudinal series scale row numbers (35–37 in
S. concursus vs. 34–35 in S. celsior; Table 1).
Stolephorus concursus differs from S. bataviensis in having
higher gill raker numbers on the first [16–18 (modally 17)+ 21–23 (23) = 38–41 (39) vs. 14–17 (15) + 19–22 (20) =33–38 (35) in S. bataviensis; Fig. 2A] and second gill arches[11–13 (modally 12) + 19–22 (20) = 30–34 (32) vs. 9–12
(11) + 17–20 (18) = 27–32 (29); Fig. 2B], and a longer snout[4.8–5.4% SL (mean 5.0%) vs. 3.7–4.7% (4.3%); Fig. 3H]

115

Hata & Motomura: Two new species of the genus Stolephorus
and pectoral fin [16.5–18.1% SL (17.0%) vs. 14.7–16.4%(15.5%); Fig. 3B].

20H03311; the JSPS Core-to-Core Program: B Asia-Africa
Science Platforms; the “Biological Properties of Biodiversity
Hotspots in Japan” project of the National Museum of Natureand Science, Tsukuba, Japan; and “Establishment of Glocal
Research and Education Network in the Amami Islands”
project of Kagoshima University adopted by the Ministry of
Education, Culture, Sports, Science and Technology, Japan.

Key to species previously identified as Stolephorus waitei“Spotty-face Anchovy” by Whitehead et al. (1988)
Stolephorus waitei sensu Whitehead et al. (1988) was
diagnosed by a long maxilla, extending posteriorly at least
beyond the preopercle anterior margin; dusky spots on thesuborbital area (in adults), snout, and lower-jaw tip; andfewer gill rakers on the lower limb of first gill arch (25 or
fewer). The included species can be separated as follows[data for S. babarani, S. bataviensis, S. baweanensis, andS. waitei based on Hata et al. (2019, 2020a)]:

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1. Maxilla short (< 19.2% SL), not reaching preopercle posterior
margin; a few spots on suborbital area and lower-jaw tip;
pseudobranchial filaments 28 or more.......................................
..............................................S. waitei (Queensland, Australia)– Maxilla long (> 19.9% SL), just short of or extending beyond
preopercle posterior margin; numerous spots on suborbital
area (adults) and lower-jaw tip; pseudobranchial filaments 25
or fewer....................................................................................2
2. Posterior tip of depressed pelvic fin beyond vertical throughdorsal-fin origin........................................................................3– Posterior tip of depressed pelvic fin not reaching vertical throughdorsal-fin origin........................................................................4
3. Gill rakers on first and second gill arches 14–17 + 19–22 =33–38 and 9–12 + 17–20 = 27–32, respectively; snout short(3.7–4.7% SL); pectoral fin short (14.7–16.4% SL).................
........................................ S. bataviensis (Taiwan to Indonesia)– Gill rakers on first and second gill arches 16–18 + 21–23 =38–41 and 11–13 + 19–22 = 30–34, respectively; snout long
(4.8–5.4% SL); pectoral fin long (16.5–18.1% SL)..................
............................. S. concursus, new species (Tonga and Fiji)
4. Gill rakers on first gill arch 40 or more; pectoral and pelvic
fins long (> 16.6% SL and 9.5% SL, respectively)..................
.......S. celsior, new species (Manila Bay, Luzon, Philippines)– Gill rakers on first gill arch 41 or less; pectoral and pelvic finsshort (< 16.8% SL and 9.4% SL, respectively)......................5
5. Gill rakers on first and second gill arches 14–17 + 19–22 =33–38 and 9–12 + 17–21 = 26–32, respectively; snout long
(3.8–4.6% SL)............................................................................
................... S. baweanensis (India to Vietnam and Indonesia)– Gill rakers on first and second gill arches 16–18 + 21–23 =38–41 and 10–13 + 18–21 = 30–33, respectively; snout short(3.6–3.9% SL).......................S. babarani (Panay, Philippines)

ACKNOWLEDGEMENTS
We thank O. Crimmen, J. Maclaine, and D. Nicholson(BMNH), A. Suzumoto (BPBM), K. Matsuura, G. Shinohara,
and M. Nakae (NSMT), K. Miyamoto (OCF), and J.
Williams, K. Murphy, S. Raredon, and D. Pitassy (USNM)
for opportunities to examine specimens of Stolephorus.
We also thank G. Hardy (Ngunguru, New Zealand), forchecking an early version of the manuscript. This study was
supported in part by a Grant-in-Aid from the Japan Society
for the Promotion of Science for JSPS Fellows (DC2: 296652); the Sasakawa Scientific Research Grant from theJapan Science Society (28-745); JSPS KAKENHI Grant
Numbers 19K236910001, JP23580259, JP26450265, and116

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