Platax 20: 55-66, 2023
doi: 10.29926/platax.202312_18.0005

First record of Thrissina splendida (Clupeiformes, Engraulidae)
from Taiwan and a key to all Taiwanese species of Thrissina
Harutaka Hata
National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. NW, Washington, DC 20560,
USA*

Corresponding author. Email: k2795502@kadai.jp

Abstract
A single specimen of Thrissina splendida Hata, 2022, collected from Yun-Lin County,
Western Taiwan, was recently found in the fish collection of the University of Michigan
Museum of Zoology, representing the first record of this species from Taiwan. It had
previously been recorded only from the Korean Peninsula and the Chinese coast. The
Taiwanese specimen has been described in detail herein, and a key to all TaiwaneseThrissina spp. has additionally been provided.
Keywords: anchovies, biodiversity, ichthyofauna, new record, taxonomy, Thrissina adelae

Although Northwestern Pacific species ofIntroduction

Thrissina, which are characterized byThrissina Jordan & Seale, 1925, anpossessing a long maxilla posteriorly (justIndo-Pacific genus of marine and/or

reaching the pectoral fin insertion), hadbrackish water anchovies (Engraulidae),long been regarded as a single species,

presently

Thrissina

includes

34

valid

speciesadelae(Rutter,

1897)(Roberts, 1978; Wongratana, 1983, 1987;(Whitehead et al., 1988), Hata (2022)

Whitehead et al., 1988; Wongratana et al.,showed that “T. adelae” included true T.

1999; Gill et al., 2018; Hata & Motomura,adelae (distributed off Fujian Province,

2019; Hata & Nakae, 2019; Hata & Koeda,

China to Northern Vietnam) and a new

2020; Hata, 2020, 2022; Hata et al., 2020,

species Thrissina splendida Hata, 2022

2021, 2022a, 2023a-b). Some species of(Southeastern

the genus are caught abundantly and

Peninsula and Bohai Sea to Fujian

important for fisheries in Taiwan (Yang,

Province). Although both species occur

2013; Chiang et al., 2014; Hata, 2019).
55

F

coast

of

the

KoreanPlatax 20: 55-66, 2023
doi: 10.29926/platax.202312_18.0005

sympatrically off Fujian Province, theHistory, Chicago, USA (FMNH); National

identity of individuals in Taiwanese waters,

Museum of Marine Biology and Aquarium,

namely the Taiwan Strait located east of

Checheng, Taiwan (NMMB); University

Fujian Province, has not been confirmed toof Michigan Museum of Zoology, Ann

date. During a revisionary study of theArbor, USA (UMMZ); and Smithsonian

genus, a specimen collected off the west

Institution, National Museum of Natural

coast of Taiwan that had been deposited in

History, Washington, DC, USA (USNM).

the University of Michigan Museum ofNomenclature and authorship of the genus

Zoology was identified as T. splendida.

Thrissina Jordan & Seale, 1925 followed

Because the specimen represents the first

Kottelat (2013).

record of this species from Taiwan, it isdescribed in detail below, and a key for the

Results

six Taiwanese species of Thrissina has also

Family Engraulidae Gill, 1861

been provided.

Thrissina Jordan & Seale, 1925
Thrissina splendida Hata, 2022
Taiwanese species name: 多稜⾧頜稜鯷Materials and methods
Methods for counts and proportional(Fig. 1)

measurements (Tables 1-2) followed Hata
and Motomura (2019), all being made with

Specimen examined. UMMZ 194341,

digital calipers to the nearest 0.01 mm.

102.3 mm SL, 122.1 mm total length,

Abbreviations

Mailiao, Yun-Lin County, Taiwan.

are

as

follows—

SL=standard length; UGR, LGR, andgill rakers, respectively (with associatedDescription of the Taiwanese specimen.
Body compressed laterally, withnumbers indicating the specific gill arch);

greatest depth at dorsal fin origin. Dorsal

distance from dorsal fin origin to pectoralprofile gradually elevated from snout tip to

fin insertion=D–P1; distance from dorsal

dorsal fin origin, thereafter lowering to

fin origin to pelvic fin insertion=D–P2;

uppermost point of caudal fin base. Ventral

distance between dorsal and anal fin

contour lowering from snout tip to lower

origins=D–A; distance between pectoral

margin of mandibular articulation, paralleland pelvic fin insertions=P1–P2; distanceto body axis thereafter to anal fin origin,from pelvic fin insertion to anal fin

and subsequently rising to lowermost point

origin=P2–A. Institutions are abbreviatedof caudal fin base. Abdomen covered with

as follows: Field Museum of Natural

17 and 9 sharp, needle-like scutes from

TGR= upper limb, lower limb, and total

56

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Platax 20: 55-66, 2023
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Tab. 1. Counts of Thrissina splendida.
Hata (2022)

Standard length (mm)This study

Holotype

Paratypes

North Korea

TaiwanCAS-SU 26536

n = 18n =4

UMMZ 194341

129.8

62.9–142.7

102.5–145.4

102.3

Mode

Dorsal fin rays (unbranched)

3

3

3

3

3

Dorsal fin rays (branched)

11

10–11

10

10

10

Anal fin rays (unbranched)

4

3–4

4

4

4

Anal fin rays (branched)

40

37–43

40–42

41

41

Pectoral fin rays (unbranched)

1

1

1

1

1

Pectoral fin rays (branched)

12

11–12

11

10

11

Pelvic fin rays (unbranched)

1

1

1

1

1

Pelvic fin rays (branched)

6

6

6

6

6

Caudal fin rays

19

19

19

19

19

Gill rakers on 1st gill arch (upper)

12

11–13

11–12

13

12

Gill rakers on 1st gill arch (lower)

18

16–18

17–18

18

18

Gill rakers on 1st gill arch (total)

30

27–31

28–30

31

30

Gill rakers on 2nd gill arch (upper)

10

9–11

9–10

11

10

Gill rakers on 2nd gill arch (lower)

17

16–18

16–18

18

17

Gill rakers on 2nd gill arch (total)

27

25–29

26–27

29

27

Gill rakers on 3rd gill arch (upper)

7

7–9

7–8

7

7

Gill rakers on 3rd gill arch (lower)

10

9–10

9–10

10

10

Gill rakers on 3rd gill arch (total)

17

16–19

16–17

17

17

Gill rakers on 4th gill arch (upper)

5

5–7

6–7

6

6

Gill rakers on 4th gill arch (lower)

9

8–9

8

9

8

Gill rakers on 4th gill arch (total)

14

13–16

14–15

15

14

Gill rakers on posterior face of 3rd gill arch

5

4–6

4–5

5

5

Pre-pelvic scutes

19

16–20

17

17

17

Post-pelvic scutes

10

9–11

10–11

9

10

Total pelvic scutes

29

26–30

27

26

26

Scale rows in longitudinal series

46

44–46

44–46

44

45

Transverse scales

12

11–12

11–12

11

11

Vertebrae

46

46–49

47

45

47

57

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Platax 20: 55-66, 2023
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Tab. 2. Measurements of Thrissina splendida
Hata (2022)

This study

Holotype

Paratypes

North Korea

TaiwanCAS-SU 26536

n = 18n =4

UMMZ 194341

129.8

62.9–142.7

102.5–145.4

102.3

Mean

Head length

20.9

19.6–24.7

20.3–22.1

24.1

22.6

Body depth

25.7

20.5–25.1

23.2–25.2

22.7

22.7

Pre-dorsal fin length

52.3

49.1–52.3

52.1–53.8

52

51.4

Snout tip to pectoral fin insertion

23.3

21.9–27.0

22.8–24.6

26.4

25

Snout tip to pelvic fin insertion

38.5

38.2–43.1

38.3–42.7

40.7

40.5

Snout tip to anal fin origin

56.4

57.1–60.6

57.6–61.4

60

59.1

Dorsal fin base length

8.8

7.6–9.4

7.9–8.8

8.9

8.5

Anal fin base length

34.7

33.5–39.4

34.2–37.8

37.6

36.4

Caudal peduncle length

8.7

6.7–9.8

7.0–7.9

7.1

8.1

Caudal peduncle depth

broken7.4–9.0

7.7–9.1

8.4

8.2

D–P1

37.7

32.7–38.2

34.9–37.9

33.9

35.1

D–P2

27.5

22.6–29.2

26.5–29.0

26.1

25.7

D–A

26.4

20.3–24.9

23.1–246

23.6

22.6

P1–P2

16.7

14.6–18.4

16.0–18.8

14.9

16.4

P2–A

19.7

16.7–22.7

19.3–22.3

18.6

19.3

Pectoral fin length

broken17.7–20.4

18.0–19.9

20.2

19.1

Pelvic fin length

7.7

6.9–8.7

7.2–8.4

7.8

7.9

Maxilla length

24.5

24.6–28.1

23.4–25.5

27.1

26

Mandibular length

15.9

14.3–18.1

15.3–16.5

18

16.7

Supramaxilla end to maxilla end

10.7

9.3–12.0

9.1–10.7

10.2

10.8

4th anal fin ray length

broken11.7–13.3

11.3–12.5

13.3

12.7

5th anal fin ray length

broken11.2–13.1

10.1–12.3

13

12.2

Orbit diameter

5.8

5.1–7.0

5.6–6.6

6.5

6.1

Eye diameter

4.9

4.3–6.2

5.0–5.7

5.3

5.3

Snout length

3.1

3.1–4.0

3.4–3.7

3.8

3.6

Standard length (mm; SL)
As % of SL

Interorbital width

5.1

4.5–5.6

4.8–5.3

5.5

5.2

Postorbital length

12.5

11.7–13.8

12.1–12.5

13.9

12.9

58

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Platax 20: 55-66, 2023
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Fig. 1. (A) Lateral, (B) dorsal, and (C) ventral views of a preserved specimen ofThrissina splendida collected from Mailiao, Yun-Lin County, Western Taiwan:
UMMZ 194341, 102.3 mm SL, 122.1 mm total length.

59

FPlatax 20: 55-66, 2023
doi: 10.29926/platax.202312_18.0005

isthmus to pelvic scute, and pelvic scute to

Posteriormarginofpreopercle

anus, respectively. Anus just anterior toconvex and smooth. Subopercle withanalroundedfin

origin.

Caudalpeduncleposteriormargin.

compressed; its depth greater than orbit

membranediameter. Head compressed. Snout length

Pseudobranchial filaments present andmuch shorter than orbit diameter. Snout tip

covered with a fleshy membrane. Gillpointed, slightly above horizontal throughrakers long, slender, and with serrae

center of eye but lower than its upper rim.

similarly sized. Gill membrane on each

Eyes round, covered with thin, adipose

side joined distally, with most of theeyelids, lateral on head, dorsal toisthmus muscle exposed and not covered

horizontal through pectoral fin insertion,

by gill membrane. Branches of cephalicand visible in dorsal and ventral views.

lateralis sensory canal exposed at upper

Pupil round and orbit elliptical. Nostrils

part of gill opening. Pectoral fin insertion

close to each other, anterior to anteriorslightly posterior to opercle’s posterior tip.

margin of orbit and above horizontal

Uppermost ray of pectoral fin unbranched;through midline of body. Interorbital spaceother rays branched. Posterior tip offlat. Mouth large, inferior, below body axis,

pectoral fin pointed, slightly beyond pelvic

and extending backwards well beyond

fin insertion. Pectoral fin triangular, withposterior margin of eye. Mandible slender

upper, lower, and posterior margins nearlyand much shorter than maxilla. Anterior tip

straight. Pelvic fin insertion anterior toof mandible slightly dorsal to lower

dorsal fin origin. Posterior tip of depressed

margin of eye. Maxilla (broken on left side)

pelvic fin not reaching vertical throughlong, its posterior tip pointed and beyond

dorsal fin origin. Pelvic fin triangular, withposterior-most point of pectoral fin

upper, lower, and posterior margins nearly

insertion. First supramaxilla small andstraight. Anteriormost ray of pelvic fin

oval. Second supramaxilla elongated andunbranched; other rays branched. Dorsal

cylindrical anteriorly, posteriorly round,

fin origin anterior to anus. Anterior-mostand symmetrical. Small uniserial, conicalthree rays of dorsal fin unbranched; other

teeth on both jaws. Single row of fine teeth

rays branched. Single spine-like scuteon palatine. Several conical teeth on vomer.

located just anterior to dorsal fin origin.

Fine teeth patch on pterygoids. Small

Dorsal fin triangular; dorsal contour risingdense conical teeth on upper edges of

sharply from fin origin to third fin ray tip,basihyal and basibranchial. No teeth on

thereafter lowering to tip of posterior-mostdorsal surface of hyoid.

fin ray. Anal fin origin just below origin of60

F

without

Opercular
serrations.

Platax 20: 55-66, 2023
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8th dorsal fin ray. Lower contour of anal

Remarks.

The

specimenfromfin dropping from fin origin to 4th fin ray

Taiwan was assigned to the genus

tip, subsequently almost parallel to ventral

Thrissina, defined by Whitehead et al.

profile of body. Caudal fin forked. Body

(1988) and Wongratana et al. (1999; asscales completely lost, except for ventral

Thryssa) as having 17 and 9 strongly

scutes.

keeled pre-pelvic and post-pelvic scutes,

Coloration of the preserved specimen.

respectively, on the ventral edge, a spine-

Body uniformly pale ivory. Melanophores

like scute at the dorsal fin origin, dorsal

densely scattered on upper part of lateraland anal fins with 13 and 44 rays,

body surface. Numerous melanophores

respectively, its uppermost pectoral fin ray

densely scattered on dorsum, forming

not extended as a filament, and small

indistinct double lines from occipital area

conical teeth on both jaws. Moreover, theto dorsal fin origin and end of dorsal fin

specimen was identified by the followingbase to caudal fin base. Branches ofcombination of characters, which closely

cephalic lateralis sensory canal on uppermatched the diagnostic features of T.

part of gill opening black, forming asplendida (Hata, 2022): long maxilla

distinct black blotch. All fins pale andposterior to posterior end of pectoral fin

semi-transparent. Melanophores scatteredbase; small first supramaxilla; paired dark

along fin rays of dorsal and caudal fins. No

lines on dorsum; distinct black blotch on

melanophores on pectoral, pelvic, or anal

upper part of gill opening; 11 transverse

fins, nor on jaws, gill rakers, isthmus,

scales; 44 longitudinal series of scale rows;

mouth roof, or inner surface of opercle.

snout 3.8% of standard length (SL); lower

Melanophores scattered on upper branch

jaw 18.1% of SL; D–A 23.6% of SL; 4thof gill arch, gill filaments on upper part ofanal fin ray length 13.3% of SL; and 5th

first gill arch, and dorsal surfaces of theanal fin ray length 13.0% of SL. Otherhyoid and bathyhyal.

meristic and morphometric characters,

Distribution. Thrissina splendida isincluding gill-raker counts on each gill

distributed in the 1) Northwestern Pacificarch, of the specimen generally matchedalong the Chinese coast from the Bohai

those of T. splendida from Hata (2022),

Sea to Fujian Province, 2) northwestern

except for slight differences in branchedand southeastern coasts of the Korean

pectoral fin ray (10 in the Taiwanese

Peninsula, and 3) west coast of Taiwan

specimen vs. 11–12; Table 1) and(Hata, 2022; this study).

vertebral (45 vs. 46–49) counts. Detailed
comparisons of T. splendida with similar
61

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Platax 20: 55-66, 2023
doi: 10.29926/platax.202312_18.0005

congeners were given in Hata (2022), who

Becausedescribed the former from 19 specimens.

mistakenly regarded as a senior synonym

The present specimen, together with theof T. chefuensis in the past, TaiwaneseNorth Korean ones also discussed herein,

records of T. kammalensis are in fact

are the only examples of T. splendida

misidentifications of T. chefuensis, thereported since the original description. The

former now being recognized as differing

former represents the first record of thefrom T. chefuensis in having a longer

species from Taiwan. Additionally, four

maxilla that posteriorly reaches thecomparative specimens (FMNH 55401,

posterior margin of the opercle (vs. not

102.5–145.4 mm SL) represent the firstreaching the posterior margin of therecords from North Korea and the east

preopercle in T. chefuensis; Hata et al.,coast of the Yellow Sea.

2020). Differences between the twoSpecies of the genus Thrissina in Taiwan.

species were detailed by Whitehead et al.

According to Young et al. (1994), Hata

(1988), Wongratana et al. (1999), and Hata

(2019), and Hata & Koeda (2020), voucher

and Nakae (2019). Although Shen and Wu

specimens of five species of the genus(2011) showed a color photograph of anThrissina

chefuensis

individual named “Thryssa baelama” in(Günther, 1874), Thrissina dussumieri

their Taiwanese fish list, the individual is(Valenciennes,

1848),

Thrissina

probably T. encrasicholoides due to theencrasicholoides(Bleeker,1852),

absence of an orange spot on the cleithrum

Thrissina hamiltonii (Grey, 1835), and(Hata et al., 2023b). Consequently, the six

Thrissina setirostris (Broussonet, 1782)-

species included in the following key have

have been recorded from Taiwan. However,been confirmed from Taiwanese waters. In

the species previously called “T. hamiltonii”

addition, Taiwanese specimens in freshin the Northwestern Pacific, including

condition (except for T. splendida) are

Taiwan, was described as a separate newshown in Hata (2019), with T. katana

species: Thrissina katana Hata, Lavoué &having being labelled as T. hamiltonii.
Key to species of Thrissina recorded
from Taiwan
1a. Maxilla short, posteriorly not beyond
posterior margin of preopercle ….…..... 2
1b. Maxilla long, posteriorly beyond
opercular margin .……..………............. 3viz.-

Thrissina

Motomura, 2022. In addition, Thrissina
kammalensis (Bleeker, 1849), which has
sometimes been reported in Taiwanesewaters (e.g., Shen, 1984; Shen & Wu, 2011;
Shao, 2023), is actually limited to the Gulfof Thailand and Indonesia (Whitehead etal., 1988; Wongratana et al., 1999).
62

F

T.

kammalensis

had

been

Platax 20: 55-66, 2023
doi: 10.29926/platax.202312_18.0005

2a. 1LGR > 26; keeled ventral scutes> 23 ………………..… T. chefuensis a2b. 1LGR ≦ 26; keeled ventral scutes< 18 …………… T. encrasicholoides b102.5–145.4 mm SL, Namp'o-hang,North Korea (Yellow Sea; 38°42'14.9"N125°23'59.1"E).

Acknowledgements
3a. Posterior tip of maxilla just reachingor slightly extending beyond posterior
margin of opercle, not reaching pectoral
fin insertion ….…………… T. katana c
3b. Posterior tip of maxilla at least
reaching pectoral fin insertion ………. 4

I thank R. Singer (UMMZ) and C.
McMahan, S. Mochel, and K. Swagel(FMNH) for opportunities to examine
specimens of T. splendida. I also thank H.C. Ho (NMMB) for advice on the new
Taiwanese name of the species, T. Munroe,
K. Murphy, D. Pitassy, S. Smith, and A.
Reft (USNM) for curatorial assistance, and
G. Hardy (Ngunguru, New Zealand) forchecking an early version of themanuscript. This study was supported inpart by a Sasakawa Scientific Research
Grant from the Japan Science Society (28745); a Grant-in-Aid of research from theJapan Society for the Promotion of
Science (JSPS) for JSPS fellows (DC2:29-6652); a JSPS KAKENHI grant(19K23691); and a JSPS Overseas
Research Fellowship (202160519).

4a. 1LGR < 13; maxilla very long, its
posterior tip beyond pelvic fin insertion in
adults…………………… T. setirostris b
4b. 1LGR > 15; maxilla moderately long,its posterior tip not reaching pelvic fin
insertion ……………………………… 5
5a. ventral scutes < 25; posterior tip ofmaxilla reaching at least halfway along
pectoral fin and to the pelvic finbase …………….………. T. dussumieri b
5b. ventral scutes > 25; posterior tip ofmaxilla slightly beyond or slightly shortof posterior-most point of pectoral finbase ……………..………. T. splendida dReferences
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