Pempheris adusta Bleeker, 1877

Koeda, Keita, Yoshino, Tetsuo, Imai, Hideyuki & Tachihara, Katsunori, 2014, A review of the genus Pempheri s (Perciformes, Pempheridae) of the Red Sea, with description of a new species, Zootaxa 3793 (3), pp. 301-330 : 303-307

Pempheris adusta Bleeker, 1877

Fig. 1 , Tables 1–5

Pempheris adusta Bleeker, 1877: 50 , table 383, fig. 1: Fricke, 1999: 320: Koeda et al., 2013a, b: Koeda et al., in press.

Pempheris mangula (not of Cuvier): Günther, 1874: 102, table 59, fig. B: Klunzinger, 1870: 469.

Pempheris molucca (not of Cuvier): Carcasson, 1977: 147.

Pempheris oualensis (not of Cuvier, in Cuvier & Valenciennes): Snyder, 1912: 497: Okada, 1938: 179: Okada & Matsubara, 1938: 179: Matsubara, 1955: 590: Tominaga, 1963: 289.

Pempheris vanicolensis (not of Cuvier, in Cuvier & Valenciennes): Allen & Steene, 1987: 55, pl. 36, figs. 5, 6: Randall, 1992: 72, fig. 147: Thamrongnawasawat & Saisaeng, 2009: 115, upper fig.

Pempheris flavicycla flavicycla Randall et al., 2013: 1 –23.

Pempheris flavicycla marisrubri Randall et al., 2013: 1 –23.

Pempheris sp.: Hatooka, 2002: 879: Senou et al., 2006: 77: Senou et al., 2007: 56: Koeda et al., 2010: 74: Koeda et al., 2012 a, b.

Diagnosis. Faint blackish spot on pectoral fin base; pored lateral-line scales 56–64 (52–58 in Andaman Sea; 51–62 in the Pacific); scale rows above lateral line 4 1/2–6 1/2 (usually 5 1/2; 4 1/2–5 1/2, usually 4 1/ 2in Andaman Sea and the Pacific Ocean); circumpeduncular scales 14–18, usually 16; distinct blackish band on outer edge of anal fin (rarely faint or only anterior half prominent; usually none in Pacific); blackish band on posterior edge of caudal fin (caudal fin whole yellowish in Andaman Sea); upper-jaw length 14.2–15.6% SL; body depth 41.4–45.4% SL. Pempheris adusta in the Pacific is distinguished from the other members of the genus by the following combination of characters: scales ctenoid and deciduous, thin, semicircular in shape; pored lateral-line scales 51–62; scale rows above lateral line 4 1/2–5 1/2 (usually 4 1/2); faint blackish spot on pectoral fin base; usually no blackish outer margin on anal fin; dusky band on anal fin base.

Description [based on Indian Ocean specimens]. Body strongly compressed, body width 5.8–8.9 in SL; body deep, body depth 2.2–2.4 in SL; head length 3.3–3.8 in SL; snout very short, rounded, snout length 4.0– 4.4 in HL; eye large, the diameter 2.2–2.6 in HL; interorbital very narrow, 3.1–3.7 in HL; dorsal outline loosely convex to dorsal fin base, extending somewhat linearly to caudal peduncle; ventral outline convex to anal fin base, extending somewhat linearly to caudal peduncle; chest usually not keeled, but sometimes weakly keeled.

Mouth large and strongly oblique, upper jaw length 1.9–2.0 in HL; the maxilla extending to middle of pupil; maxilla broadly expanded posteriorly, its surface with level; lower jaw projecting beyond upper jaw; narrow band of villiform teeth in jaws.

Scales ctenoid and deciduous, thin, semicircular in shape; concealed small scales under surface scales; scales of ventral area larger than those of dorsal area; almost all parts of head scaled, except for just anterior to orbit; about one-third of basal part of anal fin covered with small scales; scales above lateral line more deciduous; pored lateral-line scales adherent, running parallel to dorsal outline to middle of caudal fin.

Coloration when fresh. Body scales shiny reddish brown; tip of dorsal fin broadly black, the rest hyaline; paired fins pink; posterior margin of caudal fin distinctly blackish, remainder reddish brown; entire caudal fin yellow to orange in the Andaman Sea fishes ( Randall, 1992; Allen & Steene, 1987).

Coloration when preserved. Body scales dark brown to reddish brown; areas missing scales light tan to pale brown with irregular vertical dark brown lines marking scale pockets; black pigmentation on dorsal, anal, and caudal fins usually persistent, but sometimes not; paired fins hyaline.

Genetics. Molecular analysis reveals the base sequences between the Red Sea (Rabigh, Saudi Arabia) and the Andaman Sea (Phuket, Thailand) showed no genetic difference ( Fig. 2 ). Furthermore, only 0.4 % difference was shown on mitochondrial 16S ribosomal DNA between Indian Ocean (Rabigh, Saudi Arabia; Phuket, Thailand) and Pacific Ocean (Thao I., Thailand; Taitung, Taiwan; Okinawa I., Japan), and the groups did not comprise different monophyletic groups. In this paper, discussion is limited to P. adusta from the Indian Ocean; Indo-Pacific Pempheris will be treated in a revision of the entire genus.

Comparisons. Pempheris adusta is widely distributed in the Indian Ocean, Andaman Sea and southern and western Pacific, and is abundant in these regions. Most of the counts characters were overlapped between the western Pacific fish and the Indian Ocean, and Andaman Sea fishes, and some modal differences could found in following characters: pored lateral-line scales 51–62 (vs. 56–64 in Indian Ocean; 52–58 in Andaman Sea; Table 2); scale rows above lateral line 4 1/2–5 1/2, usually 4 1/2 (vs. usually 5 1/2; usually 4 1/2; Table 3); usually no blackish band on anal fin (distinct blackish band on outer edge of anal fin in Indian Ocean and Andaman Sea); blackish band on anal fin base (no band on anal fin base in Indian Ocean and Andaman Sea); anterior margin of dorsal fin blackish (vs. tip of dorsal fin blackish in Indian Ocean and Andaman Sea); posterior margin of caudal fin dusky (vs. distinct blackish in Indian Ocean; dusky or more faint than Pacific in Andaman Sea). Although, some of the specimens in Indian Ocean have the same coloration as Pacific specimens, and some of the specimens from the Pacific have have the same coloration as fish from the Indian Ocean. In the PCA of 19 measured characters, PC1 explained 90.3% of the variance and all of the 19 characters loaded on the same level ( Table 6 ), and PC2 explained only 2.2%. On the PC1 axis, plots of Indian Ocean, Andaman Sea, and western Pacific fishes were almost completely overlapped ( Fig. 3 ).

The molecular analyses revealed sequences of specimens (listed as P. adusta ) collected from the Red Sea and the Andaman Sea have exactly the same DNA sequence, and have only a 0.4% difference between these and Pacific Ocean fish for mitochondrial 16S ribosomal DNA. This is a minimal divergence compared to the other species in this genus ( Fig. 2 ). For example, P. oualensis , the most diversified [diverse/different?] species in this genus, had 2.8% divergence in the single species. The Red Sea and the Andaman Sea groups do not comprise different monophyletic groups from the Pacific Ocean group. This means that the variations between the Indian, Andaman and the Pacific are regional, and that the species cannot be considered as more than a single species. Randall et al. (2013) described two subspecies, P. flavicycla flavicycla Randall, Satapoomin and Alpermann, 2013 from the Indian Ocean and Andaman Sea, and P. flavicycla marisrubri Randall, Bogorodsky and Alpermann, 2013 from the Red Sea, but they did not compare these species with P. adusta , or with most of the other Pacific species. In the present study, the counts and measurement of two holotypes ( BPBM 17633: P. flavicycla flavicycla ; SMF 33630: P. flavicycla marisrubri ) and most of the paratypes completely overlap those of characters to P. adusta as determined in this study ( Tables 4 , 5). In addition, the specimens ( SMF 33631, 33636) collected in same time with their holotype of P. flavicycla marisrubri have exactly same DNA sequences as our P. adusta . This evidence and our results confirm that the two subspecies are junior synonyms of P. adusta .

Anal-fin soft rays

34 35 36 37 38 39 40 41 42 43 44 45

P. adusta (Indian Ocean) 1 2 2 5 2 7 3 1 P. adusta (Andaman Sea) 1 7 20 8 9 5 P. adusta (Pacific Ocean) 2 1 6 15 22 23 12 4 3 P. mangula 2 2 3 7 9 8 5 4 2 P. nesogallica 1 8 4 4

P. tominagai 1 1 5 3 4 3

Pored lateral-line scales (Left)

47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 P. adusta (Indian Ocean) 1 7 6 11 5 7 1 2 P. adusta (Andaman Sea) 1 7 11 7 14 9 1

P. adusta (Pacific Ocean) 1 5 7 19 41 51 39 30 23 12 4 3 P. mangula 2 3 1 4 10 11 6 5 4 1 2 P. nesogallica 1 6 10 10 7 2 1 P. tominagai 3 8 6 16 4 5

Scale rows above lateral line 4 1/2 5 1/2 6 1/ 2 P. adusta (Indian Ocean) 12 52 4 P. adusta (Andaman Sea) 45 5

P. adusta (Pacific Ocean) 210 24

P. nesogallica 43

P. mangula 24 28

P. tominagai 35

10+15 10+15 10+15 10+15 10+15 Remarks. Fishes of the genus Pempheris from the Red Sea and Mediterranean Sea were mostly recorded as Pempheris vanicolensis in several publications ( Ben-Tuvia, 1985; Diamant et al., 1986; Golani & Ben-Tuvia, 1989; Golani & Diamant, 1991; Golani, 1998; Goren & Galil, 2001). These reports are faunal lists with no species descriptions, but, as P. vanicolensis is only known from the western Pacific (Koeda et al., unpublish data), and not from the Indian Ocean (except the Andaman Sea). What might account for the misidentifications of P. adusta as P. vanicolensis from the Red Sea, is the blackish band on the anal fin that both species share. Several studies have reported P. vanicolensis from the Indian Ocean by photograph ( Randall, 1992; Thamrongnawasawat & Saisaeng, 2009). However, these images appear to be of P. adusta because of the blackish band along the anal fin margin and blackish spot on the pectoral fin base. Pempheris vanicolensis can clearly distinguish from the species here considered in having no blackish spot on pectoral fin base; a bright yellow pectoral fin; pored lateral-line scales 57–65; scale rows above lateral line 5 1/2–6 1/2.

This species spawn mainly during April to June, but spawning occurred year round in Okinawa I., Japan (Koeda et al., in press). The spawning occurred after sunset, and only individuals larger than 110 mm SL join to the spawning group.

Distribution. Pempheris adusta is widely distributed in the Indian Ocean and Red Sea, but not in the Mediterranean Sea. This species also widely distribute in the Pacific Ocean and Andaman Sea, and has widest distribution all species in the genus. The specimens were collected from 0–20 m depth.

Figures

FIGURE 2. Phylogenetic tree of the 15 Pempheris species recovered from mitochondria 16 S, estimated by the maximum likelihood (ML) method, using as out group Parapriacanthus ransonetii (Pempheridae). Only bootstrap values based on 1000 replications higher than 50 % by ML (left) and neighbor-joining (right) are displayed, respectively.

FIGURE 3. Plot of first and second principal components scores (PC 1 and PC 2) based on 19 body measurements of specimens of Pempheris adusta from Indian Ocean (solid triangle), Andaman Sea (gray square), and Pacific Ocean (open circle). Proportional eigenvalues of PC 1 and PC 2 are 17.21 and 0.41, respectively. Eigenvectors are presented in Table 6.

FIGURE 1 a – c. a: Holotype of Pempheris adusta (RMNH 6161), 130 mm, Ambon I., Indonesia. b: P. adusta (USNM- 402278), 122 mm, Anjouan I., Comoros, Indian Ocean. c: P. adusta (URM-P 45863), 101 mm, Phuket, Thailand, Andaman Sea.

Tables

TABLE 4. Counts of Pempheris adusta, holotype (RMNH 6161), specimens from Indian Ocean, Andaman Sea, Pacific Ocean, and two holotypes of P. flavicycla (P. flavicycla flavicycla: BPBM 17633; P. flavicycla marisrubri: SMF 33630).

	P. adusta Holotype	P. adusta (Indian Ocean)	P. adusta (Andaman Sea)	P. adusta (Pacific Ocean)	P. flavicycla Holotypes
Number of Individuals	1	51	50	526	2
Standard length (mm)	130.4	66.9‒148.8	83.1‒135.9	40.4‒158.4	137.0
Counts
Dorsal-fin rays	VI, 9	VI, 8‒10 usually 9	VI, 8‒9 usually 9	VI, 8‒10 usually 9	VI, 9, 10
Anal-fin rays	III, 40	III, 36‒44	III, 37‒43	III, 37‒45	III, 41, 42
Pectoral-fin rays	I, 16	I, 15‒18 usually 16‒17	I, 15‒17	I, 15‒18 usually 16‒17	I, 17
Pelvic-fin rays	I, 5	I, 5	I, 5	I, 5	I, 5
Caudal-fin rays	9+8	9+8	9+8	9+8	9+8
Proccurent caudal rays	6+4	6‒9+4‒6	5‒8+4‒6	5‒8+4‒5	7+4
Left pored lateral-line scales	57	56‒64	52‒58	51‒62	60, 61
Right pored lateral-line scales	57	56‒63	53‒58	51‒62	60
Scale rows above lateral line	4 1/2	4 1/2‒6 1/2 usually 5 1/2	4 1/2‒5 1/2 usually 4 1/2	4 1/2‒5 1/2 usually 4 1/2	5 1/2
Scale rows below lateral line	13	11‒16	11‒13	11‒15	12, 13
Predorsal scales	29	27‒38	27‒34	26‒38	30, 37
Circumpeduncular scales	16	14‒18 usually 16	16	14‒16 usually 16	16
Gill rakers	8+21	8‒10+20‒24	8‒9+21‒22	7‒10+20‒23	9+22, 10+21