Pempheris mangula Cuvier, 1829

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 : 309-315

Pempheris mangula Cuvier, 1829

Fig. 4 ; Tables 1–3, 7 , 8

Pempheris mangula Cuvier, 1829: 195 ; Cuvier, in Cuvier & Valenciennes, 1831; Fricke, 1999: 320; Laith and Koeda, 2013. Pempheris rhomboidea Kossmann & Räuber, 1877: 398 , pl. 3 (Type locality: Massawa or Dahla, Eritrea; Fig. 4 b). Pempheris erythraea Kossmann & Räuber, 1877: 398 (Red Sea).

Pempheris russellii Day, 1888 (Type locality: Sind, Pakistan; Fig. 4 c).

Pempheris vanicolensis (not of Cuvier, in Cuvier & Valenciennes): Randall, 1983: 85.

Diagnosis. Pored lateral-line scales 49–60; scale rows above lateral line 4 1/2–5 1/2; circumpeduncular scales 14–18, usually 16; no blackish spot on pectoral fin base; usually only the tip of dorsal fin blackish; huge eye, the diameter 13.0–15.3% SL, 43.6–51.5% HL; body very deep 44.6–49.1% SL; usually irregular faint longitudinal light stripes on the body side.

Description. Body strongly compressed, body width 6.5–7.0 in SL; body very deep, body depth 2.0– 2.4 in SL; head length 2.7–3.4 in SL; snout very short, rounded, snout length 3.9–4.8 in HL; interorbital very narrow, 3.2–3.7 in HL; dorsal outline weakly 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 keeled, but sometimes weakly or not at all.

Mouth large and strongly oblique, upper jaw length 1.7–2.0 in HL; the maxilla extending to anterior pupil or slightly beyond, but reaching middle of pupil; maxilla broadly expanded posteriorly; lower jaw projecting beyond upper jaw; narrow band of villiform teeth in jaws.

Scales ctenoid, thin, semicircular in shape, wider than long, and deciduous; 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 or anterior half of anal fin rarely blackish, the rest brownish; paired fins pink or brown; caudal fin brown, rarely with dusky posterior margin ( Randall, 1983).

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

P. adusta P. adusta P. adusta P. adusta P. flavicycla Holotype (Indian Ocean) (Andaman Sea) (Pacific Ocean) Holotypes Number of Individuals 1 19 50 84 2

Standard length (mm) 130.4 100.5‒143.2 83.2‒135.9 62.4‒145.7 102.4, 137.0 As % of standard length

Head length 26.2 26.4‒30.3 (29.3) 27.7‒30.1 (28.9) 26.3‒31.6 (28.1) 28.3, 29.8 Head depth 31.5 31.1‒36.1 (33.1) 31.0‒33.9 (32.3) 29.8‒39.5 (32.5) 33.7, 34.2 Snout length 6.2 6.2‒7.0 (6.6) 5.7‒7.1 (6.3) 5.2‒7.9 (6.2) 6.7 Eye diameter 12.3 11.2‒14.0 (12.5) 10.9‒13.6 (12.5) 10.0‒13.2 (11.7) 11.7, 12.5 Interorbital width 9.2 8.3‒9.5 (8.9) 7.1‒8.4 (8.0) 7.2‒9.6 (8.0) 8.7, 9.2 Upper jaw length 14.6 14.2‒15.6 (15.0) 14.9‒16.7 (15.4) 13.1‒16.7 (14.5) 14.2, 15.4 Predorsal length 38.5 37.3‒41.6 (38.6) 36.0‒38.6 (37.3) 35.9‒48.2 (38.6) 39.5, 40.0 Prepelvic length 36.9 35.8‒41.2 (38.4) 34.9‒36.9 (36.4) 34.0‒44.7 (36.5) 37.5, 38.5 Preanal length 51.5 48.1‒55.5 (52.7) 47.1‒53.2 (49.3) 46.5‒53.7 (49.8) 50.0, 52.9 Body depth 40.0 41.4‒45.4 (43.6) 40.2‒44.3 (42.1) 40.2‒47.3 (43.8) 44.2, 45.2 Dorsal-fin length damaged 22.9‒26.1 (24.7) 22.9‒25.0 (24.2) 21.3‒26.4 (24.1) 24.0, 25.0 Anal-fin length damaged 12.7‒16.0 (14.7) 13.5‒17.0 (15.6) 10.9‒16.7 (13.8) 13.5, 14.2 Pectoral-fin length damaged 23.6‒26.3 (25.2) 23.8‒26.1 (25.0) 22.7‒27.4 (24.7) 25.0, 26.0 Pelvic-fin length damaged 12.7‒16.0 (14.3) 12.9‒15.1 (14.2) 10.4‒16.0 (13.1) 13.5 Dorsal-fin base 16.9 14.3‒16.8 (15.8) 15.1‒16.1 (15.7) 13.9‒17.6 (15.9) 15.4, 15.8 Anal-fin base 53.1 51.4‒56.4 (53.3) 52.4‒57.1 (55.6) 50.4‒61.3 (54.9) 51.9, 55.8 Caudal-peduncle length 8.5 7.6‒10.1 (9.0) 7.9‒9.1 (8.2) 7.3‒11.2 (8.8) 8.7, 9.2 Caudal-peduncle depth 9.2 7.6‒9.3 (8.6) 8.0‒8.9 (8.3) 7.2‒9.7 (8.4) 8.7, 9.2 As % of head length

Snout length 23.5 21.1‒25.0 (22.7) 20.0‒25.0 (21.9) 17.4‒27.3 (22.3) 22.6, 23.5, Eye diameter 47.1 38.1‒45.5 (42.2) 39.3‒46.2 (43.1) 36.1‒47.1 (41.5) 41.9, 41.2 Interorbital width 35.3 27.3‒32.4 (30.1) 24.0‒29.2 (27.6) 25.0‒32.4 (28.6) 29.0, 32.4 Upper jaw length 55.9 50.0‒53.6 (51.1) 52.0‒56.0 (53.3) 48.1‒57.1 (51.5) 50.0, 51.6 Genetics. Molecular analysis shows the base sequences of P. mangula between the Red Sea (Jaffa and Elat, Israel) and the Indian Ocean ( Muscat, Oman; Cabo Delgado, Mozambique) have only slight genetic divergences ( Fig. 2 ), showing the species as widely distribute in the Red Sea and western Indian Ocean, and the Mediterranean Sea.

Comparisons. Russell (1803, vol. 1: 10, pl. 111) described and illustrated a species of Pempheris from the Visakhapatnam coast of eastern India under the native name Mangula-kutti ( Fig. 5 ). Cuvier (1829: 195) noted that Russell's fish did not correspond to any of the species he knew, so he proposed the name P. mangula for it. Absence of type specimens precipitated much subsequent confusion. Day (1876: 175) described the characters for which he thought was '' P. mangula '', but these characters are mostly consistent with P. molucca Cuvier 1829 , especially in counts of anal-fin rays, pored scales above lateral line, and lack of coloration on any of the fins. Day also concluded that the P. mangula of Cuvier (1829) and Günther (1874) did not correspond with Russell's fish, and named it Pempheris russellii Day, 1888 , designating as type specimen a fish collected from Pakistan (ZSI 910, Fig. 4 c). However, this specimen has similar characters to P. mangula ( Tables 6 , 7 ), and means that P. russellii is a junior synonym of P. mangula . Kossmann & Räuber (1877) designated three syntypes collected from the Red Sea for P. rhomboidea . However, only one syntype (ZMB 9850, Fig. 4 b) remains; the other two are lost. Therefore, we herein establish the remaining syntype as lectotype of P. rhomboidea . The designated lectotype has completely overlapping characters with P. mangula ( Tables 7 , 8), and our molecular analysis shown only very slight genetic differences between Red Sea and Indian Ocean specimens, meaning that P. rhomboidea is junior synonym of P. mangula . Kossmann & Räuber (1877) claimed P. mangula of Klunzinger (1870) did not match any other species in the Red Sea, and proposed the new scientific name, P. erythraea . However, Klunzinger (1870) described P. mangula with the following characters: dorsal-fin rays: VI, 9–10; anal-fin rays: III, 37–38; pored lateral-line scales: 45–60; faint blackish spot on pectoral fin base, but sometimes not. These combined characters, especially the last, indicate that Klunzinger might have included two species in their reassessment of P. mangula . Because the black spot on the pectoral fin base is a specific character for P. adusta in the Red Sea, with pored lateral-line scales 45–60 less so for the species, and as we examined five specimens (SMNS 3498, 26606) of '' P. mangula '' identified by Klunzinger and confirms these as true P. mangula . Thus we conclude P. erythraea is also a junior synonym of P. mangula .

Remarks. Five Indian species, P. molucca and four species from the Red Sea, were collected from Southern India and/or Sri Lanka, but no specimens have been reported from the Visakhapatnam coast where Russell reported ''Mangula-kutti''. Pempheris malabarica Cuvier in Cuvier & Valenciennes, 1831, described from Southern India, is considered to be a junior synonym of P. molucca in our taxonomical revision on whole genus (Koeda et al., unpublish data). Our analyses based on the illustration yields the following characters: high [large] body depth; dorsal-fin rays V, 10; anal-fin rays III, 39; pectoral-fin rays I, 17; pelvic-fin rays I, 5; caudal-fin rays 19; pored lateral-line scales 59; scale rows above lateral line 4 1/2–5 1/2; scale rows below lateral line 14. The shaded parts on each fin might indicate branched rays. The specimens here, listed above as P. mangula , do not exactly match the characters in Russell's illustration, but do match some of the significant characters from the illustration, in having 39 anal-fin soft rays (34–42), pored lateral-line scales 59 (49–60), and scale rows above lateral line 4 1/2–5 1/2 (4 1/2–5 1/2). Additionally, Russell (1803) specified ''eyes enormously large'' in the text, which corresponds with the diagnosis of the present species.

The Visakhapatnam coast is located on southeastern Indian, and no specimens have been collected from this coast. However, many specimens of P. molucca , P. nesogallica , P. tominagai sp. nov., and P. mangula have been collected from Kerara and/or Tuticorin, southern India. Pempheris adusta also has similar characters to Russell's species, and has been collected from the Maldives and Sri Lanka, but has never been collected from the coast of India. Compared to Russell's description, P. nesogallica has a significantly smaller eye, and is not abundant along the Indian coast. Pempheris tominagai sp.nov. has lower scale counts, a shallower body depth, and a smaller eye. From these facts, we conclude that the present species correspond with the species described by Russell (1803), and our genetic results supported that this species widely distribute between the Indian Ocean to the Red Sea. Thus, the specimen collected at the southern India (BPBM 41095; 129.7 mm SL) are here designated the neotype of P. mangula .

As % of head length

Snout length 22.5 20.9–25.6 (22.9) 21.9 21.2 Eye diameter 45.0 43.6–48.1 (45.6) 46.9 51.5 Interorbital width 27.5 26.8–30.8 (28.9) 28.1 27.3 Upper jaw length 52.5 51.2–55.6 (53.2) 56.3 60.6 Fowler (1931) described the distribution of P. mangula as the Red Sea, South Africa, Madagascar, India, East Indies, Philippines, Vietnam, and New South Wales. However, the characters mostly align with P. schwenkii Bleeker, 1855 , and a new species, P. tominagai sp. nov., based on the low pored lateral line scale counts, number of scale rows above and below lateral line, and anal fin counts.

Pempheris mangula has been reported from South China Sea ( Randall & Lim, 2000), Indonesia ( Allen & Adrim, 2003), Mauritius ( Gudger, 1929), western coast of India ( Manilo & Bogorodsky, 2003), Réunion, Mauritius, Comoros, Madagascar, and the Seychelles ( Fricke et al., 2009). These reports are faunal lists with no species descriptions, but, as P. mangula is only known from the Indian Ocean, and is not in the Pacific Ocean, the report from South China Sea and Indonesia is likely a misidentification of other species. Randall (1983) reported P. vanicolensis from the Red Sea, and provided a color photograph. However, the photographed specimen (BPBM 18188) has been re-identified as P. mangula .

Distribution. Pempheris mangula is widely distributed in the western Indian Ocean, the Red Sea and the eastern Mediterranean Sea. This species is also distributed along the northwestern coast of Australia, but not along the coasts and islands of Indonesia. This species is the only species that has migrated to the Mediterranean Sea from the Red Sea. 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 5. Illustration of '' Mangula-kutti' ' in Russell (1803).

FIGURE 4 a. Neotype of Pempheris mangula (BPBM 41095), 122 mm SL, Tuticorin, India. b: lectotype of Pempheris rhomboidea (ZMB 9850), 105 mm SL, Masawa or Dahla, Eritrea; c: holotype of P. russellii (ZSI 910), 115 mm SL, Sind, Pakistan.

Tables

TABLE 7. Counts of Pempheris mangula, lectotype of P. rhomboidea (ZMB 9850), and holotype of P. russellii (ZSI 910).

	P. mangula Neotype	P. mangula Non-types	P. rhomboidea P. russellii Lectotype Holotype
Number of Individuals	1	53	1 1
Standard length (mm)	129.7	58.9–141.6	104.5 114.7
Counts
Dorsal-fin rays	VI, 9	VI, 9–10 rarely 10	VI, 9 VI, 9
Anal-fin rays	III, 37	III, 34–42	III, 39 III, 41
Pectoral-fin rays	I, 18	I, 15–18 usually 16–17	I, 16 I, 17
Pelvic-fin rays	I, 5	I, 5	I, 5 I, 5
Caudal-fin rays	9 + 8	9 + 8	9 + 8 9 + 8
Proccurent caudal rays	7 + 5	5–8 + 4–5	7 + 5 N/A
Left pored lateral-line scales	54	49–59	54 54
Right pored lateral-line scales	53	49–60	54 52
Scale rows above lateral line	5 1/2	4 1/2–5 1/2	4 1/2 damaged
Scale rows below lateral line	12	11–14	12 13
Predorsal scales	33	26–39	30 damaged
Circumpeduncular scales	16	14–18 usually 16	damaged damaged
Gill rakers	8 + 19	8–11 + 19–22	8 + 20 N/A
Vertebrae	10 + 15	10 + 15	10 + 15 N/A

TABLE 6. Factor loadings for the principal components (PC 1, PC 2) analysis of morphometric characters of the Pempheris adusta from Indian Ocean, Andaman Sea, and Pacific Ocean complex in specimens examined with all characters available for multivariate analyses (see Fig. 3).

	PC1	PC2
Standard length	0.24	‒0.12
Head length	0.24	0.06
Head depth	0.24	‒0.10
Snout length	0.22	0.34
Eye diameter	0.22	0.09
Interorbital width	0.23	‒0.02
Upper jaw length	0.23	0.17
Predorsal length	0.24	‒0.10
Prepelvic length	0.24	‒0.01
Preanal length	0.24	0.01
Body depth	0.23	‒0.18
Dorsal-fin length	0.24	0.04
Anal-fin length	0.22	0.40
Pectoral-fin length	0.23	‒0.12
Pelvic-fin length	0.23	0.02
Dorsal-fin base	0.23	‒0.24
Anal-fin base	0.21	0.54
Caudal-peduncle length	0.21	‒0.40
Caudal-peduncle depth	0.23	‒0.32