The taxonomic identity of the monocle bream Scolopsis vosmeri species complex (Perciformes: Nemipteridae), with comments on molecular phylogenetic relationships within the genus Scolopsis Author Russell, Barry C. Museum & Art Gallery of the Northern Territory, PO Box 4646, Darwin NT 0801, Australia. & School of Environmental and Life Sciences, Charles Darwin University, PO Box 40146, Casuarina NT 0811, Australia. Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a. M., Germany. Tilman. Alpermann @ senckenberg. de; https: // orcid. org / 0000 - 0002 - 8723 - 4576 Author Bogorodsky, Sergey V. 0000-0002-8723-4576 Station of Naturalists, Omsk, Russia. lpermann@senckenberg.de Author Mal, Ahmad O. Marine Biology Department, Faculty of Marine Sciences, King Abdulaziz University, Jeddah, Saudi Arabia. Marine Biology Department, Faculty of Marine Sciences, King Abdulaziz University, Jeddah Saudi Arabia. Author Bineesh, K. K. 0000-0001-9775-018X Andaman and Nicobar Regional Centre, Zoological Survey of India, Port Blair- 744102, Andaman Islands, India. kkbineesh @ gmail. com; https: // orcid. org / 0000 - 0001 - 9775 - 018 X kkbineesh@gmail.com Author Alpermann, Tilman J. 0000-0002-8723-4576 lpermann@senckenberg.de text Zootaxa 2022 2022-03-07 5105 4 501 538 journal article 20321 10.11646/zootaxa.5105.4.3 75a36a22-3d52-4649-8f77-25a47110f692 1175-5326 6333936 09027BC7-79FA-4D0A-B7DE-B9965A7887A6 Scolopsis vosmeri ( Bloch, 1792 ) Figures 1–3 ; Table 1 . New English name: Vosmaer’s monocle bream Anthias vosmeri Bloch, 1792: 120 , pl. 321 (‘japanischen Meers’ , probably Tranquebar , Tharangambadi, Tamil Nadu , India ; holotype : ZMB 8729 ). Anthias vosmaeri — Forster 1795: 16 (unjustified emendation—the subsequent spelling of the species name as vosmaeri by Forster (1795) , Lacepède (1802) , Shaw (1803) and Bleeker (1873 , 1876–1877) is more correct, but under the rules of nomenclature ( ICZN 1999 : Article 32.2) the original spelling of the name is the “correct original spelling” and the name vosmaeri is thus an “unjustified emendation”). Anthias vosmari — Bloch & Schneider 1801: 304 (misspelling). Lutjanus vosmaeri — Lacepède 1802: 213 (list, unjustified emendation). Pomacentrus enneadactylus Lacepède, 1802: 505 , 508 (‘Mer des Indes’). Sparus vosmaeri — Shaw 1803: 441 ( Japan , unjustified emendation). Sparus vosmerianus — Shaw in Shaw & Nodder 1810 : no page number, pl. 939 (‘Indian Seas’, latinisation of vernacular ‘Vosmerian Sparus’). Scolopsides vosmeri — Cuvier 1829: 178 (the name “ Anth. Vosmeri ” first referred to the genus Scolopsides in a footnote); Cuvier in Cuvier & Valenciennes 1830: 333 ( Java ); Bleeker 1849: 27 (Batavia= Jakarta ). Scolopsis argyrosomus — Kuhl & van Hasselt in Cuvier & Valenciennes 1830: 333 (name in synonymy of Scolopsides vosmeri , not available). Scolopsis vosmaeri — Bleeker 1873: 361 (Sumatra, Penang, Singapore , Thousand Is (= Kepulauan Seribu), Bangka, Java, Celebes, Sumbawa; unjustified emendation); Bleeker 1876-77: 8 (ibid.). Scolopsis vosmeri — Day 1878: 87 (in part, pl. XXIII, fig. 3); Wongratana 1978: 31 , fig. 5 (Gulf of Thailand and Andaman Sea, Thailand ); Russell 1990: 122 (in part); Shibukawa in Matsuura & Kimura 2005: 47 (Libong Island, west. coast of southern Thailand ); Barman & Mishra 2009: 36 ( India ); Shibukawa in Kimura et al. 2009: 160 (Andaman Sea); Shibukawa in Matsunuma et al . 2011: 131 ( Malaysia , off Terengganu ); Mishra et al . 2013: 447 , fig. 3; Yoshida et al . 2013: 147 (Gulf of Thailand ); Psomadakis et al . 2015: 272 , pl. XXV, fig. 199 ( Pakistan ); Psomadakis et al . 2019: 487 ( Myanmar ). Scolopsis torquata (non Cuvier)— Allen & Erdmann 2012: 489 ( Brunei , juvenile; Thailand , Phuket , adult). Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy & Selvanayagam, 2013: 444 ( India —coasts of Tamil Nadu and Kerala , Mumbai , and Sri Lanka ). Material examined (n=66, 56.1–173.1 mm SL, * = morphometric data not taken): Pakistan : NTM S.17856-025, 148.5 mm SL, Karachi ; NTM S.17856-026, 141.8 mm SL, Karachi ; NTM S.17856-027, 129.7 mm SL, Karachi. India : AMS I.21013-009, 112.1 mm SL, Madras (= Chennai ) ; BPBM 27712 ( paratype of Scolopsis igcarensis Mishra et al., 2013 ), 74.0 mm SL, off Mulloor Point , Vizhinjam , Kerala ; CMFRI GB.31.98.5.6 ( paratype of S. igcarensis ), 148.1 mm SL, Tuticorin , Tamil Nadu ; MNHN A-8089*, 2: 141.9–147.1 mm SL ( syntypes Pomacentrus enneadactylus Lacepède, 1802 ) ‘mer des Indies’ ; MNHN A-8093*, 137.6 mm SL, Pondicherry (= Puducherry ) ; MNHN A-8094*, 155.8 mm SL, Pondicherry ; NTM S.11136-001, 116.1 mm SL, Madras (= Chennai ) ; ZMB 8729 * ( holotype of Anthias vosmeri Bloch, 1792 left skin, in 3 parts), c. 128.9 mm SL, ‘japonischen Meers’ (erroneous, probably Tranquebar [= Tharangambadi ], Tamil Nadu , India : See Distribution below) ; ZSI F-331-333*, 3: 69.0–153.0 mm SL, Madras (= Chennai ), (specimens figured in Day 1878 ) ; ZSI F-401*, c. 57 mm SL, Bombay (= Mumbai ) (specimen figured as Scolopsis leucotaenia (Bleeker) in Day 1878 ) ; ZSI F-2475* & 2476*, 2: 124.0–129.0 mm SL, Madras (= Chennai ) ; ZSI F-10623/2 ( holotype of S. igcarensis ), 118.2 mm SL, Kalpakkam ( 12°33´N , 80°11´E ), Tamil Nadu ; ZSI F-10624/2 ( paratypes of S. igcarensis ), 5: 75.9–115.6 mm SL, Kalpakkam , Tamil Nadu ; ZSI F-10625/2 ( paratypes of S. igcarensis ), 3: 67.1–82.4 mm SL, Kalpakkam , Tamil Nadu . ZSI F-10626/ 2, 137 mm SL, Tharuvaikullam ; ZSI F-10627/2*, 5: 56.1–133.8 mm SL, Kalpakkam. Sri Lanka : BPBM 18765 ( paratype of S. igcarensis ), 74.0 mm SL, off Negombo ; BPBM 19031 , 150.0 mm SL, off Negombo ; BPBM 19054 ( paratype of S. igcarensis ), 75.4 mm SL, off Negombo ; BPBM 27195, 155.1 mm SL, Hikkaduwa ; NTM S.13160-007, 162.5 mm SL, Chilaw. Bangladesh : USNM 443785 ( Genbank No. MK 779330 ), 138 mm SL, Teknaf Beach , Cox’s Bazar ; USNM 443786, 106 mm SL, Teknaf Beach , Cox’s Bazar ; USNM 443787, 125 mm SL, Teknaf Fish Market , Cox's Bazar ; USNM 443788 ( Genbank No. MK 779329 ), 96 mm SL, Saint Martin's Island Jetty, Teknaf , Cox's Bazar ; USNM 443789 ( Genbank No. MK 779328 ), 77 mm , Saint Martin's Island Jetty, Teknaf , Cox's Bazar ; CUMS F505.002 , 128 mm SL, Teknaf Beach , Cox’s Bazar ; CUMS F505.005 , 93 mm SL, Saint Martin's Island , Teknaf , Cox's Bazar ; CUMS F505.007 ( Genbank No. MK 779327 ), 85 mm SL, Teknaf Fish Market , Cox's Bazar ; CUMS F505.008 , 134 mm SL, Teknaf Fish Market , Cox's Bazar ; CUMS F505.009 , 135 mm SL, Teknaf Fish Market , Cox's Bazar . Thailand : NTM S.10674-017, 126.4 mm SL, Phuket , Hat Kata and Ko Pu islet. Indonesia : MNHN 7837, 154 mm SL, Java , Batavia; NTM S.10733-011, 123.6 mm SL, Bali , Jimbaran; NTM S.16686-008, 152.2 mm SL, Bali , Jimbaran; NTM S.10664-001, 133.3 mm SL, Sumatra , Bengkulu ; USNM 183178 , 2 : 151.0– 153.2 mm SL, Sulawesi , Makassar. Singapore : ZRC 4502, 134.2 mm SL, Singapore Fish market ; ZRC 3400 , 3 : 121.6–144.1 mm SL, Ellenborough Market. South China Sea: ZRC 4169, 173.1 mm SL. East Malaysia : NTM S.14230-001, 2: 97.5–139.8 mm SL, Sabah , Sandakan; USNM 183179 , 2 : 151.6–155.0 mm SL, Sabah , Sandakan. Brunei : WAM P.33122-011, 1 of 13: 131.6 mm SL, Pelumpong I .; WAM P.33125-001, 160 mm SL, Pulau Punyit . FIGURE 1. Scolopsis vosmeri , live individuals. A : Brunei; B : Sri Lanka; C : eastern Thailand. Photos by G.R. Allen (A & C), J.E. Randall (B). FIGURE 2. Scolopsis vosmeri : A : juvenile, Brunei; B : subadult, BPBM 18765, paratype of S. igcarensis , 74 mm SL, Sri Lanka; C : adult, BPBM 27195, 155.1 mm SL, Hikkaduwa, Sri Lanka. Photos by G.R. Allen (A & B), J.E. Randall (C). Diagnosis. A species of Scolopsis with the following combination of characters: head scaly except for subopercle which is naked with numerous small pores; scales on top of head reaching forward to or just in front of anterior nostrils; lower margin of eye tangent to or just above a line from snout tip to upper pectoral-fin base; suborbital (second infraorbital) with a large retrorse spine, and a small antrorse spine on third infraorbital immediately above suborbital spine; posterior margin of preopercle serrate or denticulate, lower edge smooth; second anal-fin spine longer and more robust than first or third anal-fin spines; most body scales, except for caudal peduncle, with black spot; caudal peduncle white; a white band usually present beneath lateral line, from origin of lateral line to below posterior part of dorsal fin; a broad white bar from nape onto operculum, and suborbital with distinctive white patch; upper limb of opercular margin edged brown, lower limb edged reddish orange; no wedge-shaped dark spot on upper base of pectoral fin (sometimes small dark spot present); dorsal, anal and pelvic fins crimson or orangered; juveniles and smaller subadults lacking white bar on nape and with dark-edged white band from behind eye to upper part of caudal peduncle. Description. Counts and proportional measurements are provided in Table 1 . Dorsal-fin rays X,9; anal-fin rays III,7; pectoral-fin rays 17–19 (usually 18-19); pelvic-fin rays I,5. Body moderately deep, laterally compressed, depth 2.0– 2.7 in SL; dorsal profile of head more convex than ventral contour, head length 2.7–3.4 in SL; snout short, 3.0-4.0 in HL; eye moderately large, 2.4–3.5 in HL; lower margin of eye tangent to or just above a line from snout tip to upper pectoral-fin base. Snout length about equal to or less than eye, 0.9–1.6 in eye diameter. Interorbital width 1.0– 1.9 in eye diameter. Suborbital depth 1.8–4.4 in eye diameter; posterior margin of suborbital with a moderately large backwardly-directed spine at upper edge and 3 smaller spines or several serrations below this, lower edge smooth; a small antrorse spine on infraorbital just above suborbital spine. Posterior margin of preopercle serrate or denticulate, lower edge smooth. Opercle with smooth, low bony ridge anteriorly, running parallel to edge of preopercle; posterior margin of opercle with short flat spine just above level of upper base of pectoral fin. Exposed portions of posttemporal and supracleithrum with serrations. Teeth in jaws small, conical, curved, in 5–6 bands anteriorly, in a single series laterally. Premaxilla and dentary smooth, a little fleshy. Gill rakers short and stubby, 8–13 on first arch. Dorsal-fin membranes slightly incised, fourth and/or fifth spines longest. Second anal-fin spine longer, and more robust than other two spines, its length 1.1–2.5 times that of first and 1.1–1.5 times that of third; pectoral fins reaching to or just before of level of anus, 3.4–4.2 in SL; pelvic fins moderately long, reaching to between anus and origin of anal fin, 3.1–4.5 in SL; caudal fin forked, lobes slightly rounded; lower lobe a little shorter than upper. Body covered with ctenoid scales. Temporal region of head scaly, scales extending forward to or just in front of anterior nostrils; suborbital naked, with numerous small pores; scales on preopercle extending to margin, 3–4 (usually 3) rows behind eye, 5–8 (usually 8) transverse scale rows on preopercle; lower limb of preopercle with 1–2 rows of embedded scales; opercle with 4–9 (usually 5) transverse scale rows. Lateral-line scales 40–44 (usually 42); 3½–4½ (usually 3½) transverse scale rows above lateral line, 13½–15½ (usually 14½) rows below; vertebrae 22. Largest specimen examined 173 mm SL. Color of live fish (based on underwater photographs, Fig. 1 ). Adult body color reddish brown, shading to white on caudal peduncle; a blackish spot basally on each scale, except for breast and caudal peduncle; a white band usually present beneath lateral line from origin of lateral line to below posterior part of dorsal fin; a curving white bar extending from nape onto operculum, broader ventrally; posterior edge of opercle above spine narrowly edged dark reddish brown, opercular membrane below opercular spine narrowly edged orange-red; posterior edge of preopercle pale orange-red; suborbital broadly whitish; lips and area behind angle of mouth whitish; indistinct red-suffused dark brown bar across eye, with yellow blotch present on upper part; upper base of pectoral fin sometimes with a small black spot (usually faint or absent); spinous portion of dorsal fin pale reddish brown with red margin, soft portion semi-translucent with red margin on anterior half; pectoral, pelvic and anal fins reddish; anterior edge of pelvic fins suffused pale blue; anal-fin spines pale; caudal fin translucent greyish, upper and lower edges pale red, posterior margin narrowly blue. Juveniles ( Fig. 2A ) reddish brown dorsally, darker on back, a light pinkish to white on lower part of body; base of dorsal fin with narrow white band; distinctive white lateral band from behind upper margin of eye to upper caudal peduncle, this band bordered above and below by dark reddish brown or blackish band; body scales, particularly those on upper sides and back with darker centres; suborbital and lower part of preopercle pale whitish or silvery; fins greyish or pinkish with bluish hue; dorsal fin with traces of dusky medial band along its length. Color of fresh specimens (based on photographs, Fig. 2B & C ). Similar to those while fish alive but pectoral fins usually become reddish orange. TABLE 1. Selected counts and morphometric data for Scolopsis curite , S. japonica and S. vosmeri . Modal counts and mean proportions are in brackets.

Scolopsis vosmeri	Scolopsis japonica	Scolopsis curite
n=52	n=69	n=49
SL (mm)	74.0–173.1	61.1–160.9	63.2–159.9
Counts
Lateral line scales	40–44 (42.0)	39–44 (42)	39–45 (42)
Scales above lateral line	3½–4½ (3½)	3½–4 (3½)	3½–4 (3½)
Scales below lateral line	13½–15½ (14)	13½–16½ (15½)	12½–15½ (14½)
Gill rakers (total)	8–13 (11)	8–12 (11)	9–12 (11)
Scale rows behind eye	3–4 (4)	2–6 (4)	3–4 (3)
Scale rows on preopercle	5–9 (6)	5–10 (8)	5–8 (8)
Scale rows on opercle	4–7 (5)	4–10 (7)	4–9 (5)
Pectoral fin rays	17–20 (19)	16–19 (18)	17–19 (18)
Proportions
Body depth (% SL)	36.9–51.1 (45.7)	38.6–50.9 (44.9)	38.9–49.2 (44.6)
Head length (% SL)	29.3–37.1 (32.0)	30.5–35.7 (33.2)	30.5–36.9 (34.0)
Base of dorsal fin (% SL)	55.4–62.8 (59.1)	55.2–63.9 (59.3)	54.5–62.2 (59.1)
Base of anal fin (% SL)	15.9–21.0 (18.0)	16.1–20.7 (18.7)	17.1–21.0 (18.3)
Pectoral fin length (% SL)	24.0–29.2 (26.3)	21.3–30.4 (26.1)	20.0–29.0 (26.5)
Pelvic fin length (% SL)	22.4–32.1 (25.9)	22.8–30.1 (26.2)	22.5–29.6 (25.9)
Snout length (% HL)	24.9–33.2 (29.1)	22.8–36.2 (28.7)	22.0–33.6 (27.9)
Eye diameter (% HL)	28.2–40.9 (33.7)	32.8–46.6 (38.5)	30.6–43.2 (36.9)
Interorbital width (% HL)	19.6–34.0 (25.3)	21.8–33.8 (27.0)	19.6–29.0 (24.6)
Snout length in eye	0.9–1.6 (1.2)	1.0–2.0 (1.4)	0.9–1.9 (1.3)
Interorbital width in eye	1.0–1.9 (1.4)	1.0–2.1 (1.4)	1.1–2.1 (1.5)
Suborbital depth in eye	1.8–4.4 (2.5)	1.3–4.5 (2.9)	1.7–3.9 (2.7)
Caudal–peduncle depth in peduncle length	0.7–1.7 (1.1)	0.5–1.2 (0.9)	0.7–1.2 (1.0)
First dorsal spine in longest dorsal spine	1.2–2.3 (1.9)	1.5–2.3 (1.8)	1.5–2.1 (1.8)
Longest dorsal spine in longest dorsal ray	0.9–1.3 (1.1)	0.8–1.2 (1.0)	0.9–1.1 (1.0)
First anal spine in second anal spine	1.4–2.5 (2.0)	1.4–2.4 (1.8)	1.5–2.2 (1.9)
Third anal spine in second anal spine	1.1–1.5 (1.3)	1.0–1.4 (1.2)	1.1–1.5 (1.3)

Molecular analysis . In the phylogenetic analysis of the partial mitochondrial COI gene ( Fig. 12 ), Scolopsis vosmeri specimens form a highly supported and well divergent monophyletic clade, which is the sister clade to the clade composed of S. curite and S. japonica . Intra-specific genetic variation in S. vosmeri COI barcoding sequences seems comparatively low although the only specimen from the Pacific Ocean (i.e., voucher DOS06041 (GenBank number MK777772 ) with collection locality in southern Vietnam ) is somewhat divergent from the residual specimens in our analysis, which all come from locations in the northern Indian Ocean (ranging from the west coast of India to the west coast of Thailand ). The sequence of the specimen from Vietnam is distinguished by 9 unique base differences in 527 bp, accounting for about 1.7 percent. Unfortunately, only this single published sequence of S. vosmeri is currently available from the western Pacific Ocean for phylogenetic analysis and therefore no firm conclusions on the intra-specific diversity with respect to Indian Ocean versus Pacific populations of this species can be drawn from this observation. A set of unpublished COI sequences of conspecifics collected from Brunei (G. Allen, M. Erdman & A. Sembiring, unpublished) fall into the large clade with sequences from the Indian Ocean, indicating that this clade extends widely onto the Sunda Shelf. Our hypothesis that genetic divergent lineages have evolved during periods of isolation due to sea level low stands during glacial cycles and subsequently expanded/changed their distribution ranges is supported by broad evidence collated in more detail for other species (see e.g., Gaither & Rocha 2013 ). Habitat and distribution. Scolopsis vosmeri inhabits shallow inshore rock and coral reefs, often in turbid water, in depths to about 30 m . Known from the northern Indian Ocean from Pakistan , western India , Sri Lanka , Bay of Bengal , and Andaman Sea to western Indonesia , East Malaysia and Brunei . Not recorded from the Red Sea , Arabian Gulf , east African coast, or islands of the Western Indian Ocean . Remarks. The holotype of Anthias vosmeri Bloch (ZMB 8729) consists of a partially damaged dried skin (left side; operculum and anal fin missing; dorsal fin incomplete; other fins broken, incomplete), 151.7+ mm TL, c. 128.9 mm SL ( Fig. 3A ). Although Bloch’s description and plate of Anthias vosmeri omit any detail of the suborbital and its distinctive spine, these can be plainly seen in the holotype specimen ( Fig. 3A ) and were also mentioned by Cuvier in his description of Scolopsides vosmeri , the type of which was examined in Berlin by Valenciennes ( Cuvier & Valenciennes 1830 ) during visits in 1826 and 1829 ( Karrer 1978 ; Bauchot et al. 1997 ). Bloch’s color plate also shows a reddish body and fins, a pale band extending from the nape across the opercle (incompletely shown), yellowish band beneath the lateral line, and lack of a dark spot on the upper base of the pectoral fin ( Fig. 3B ), that are characteristic of and clearly distinguish Scolopsis vosmeri from all other species in the genus. FIGURE 3. Anthias vosmeri Bloch. A : holotype, ZMB 8729, c.128.9 mm SL; B : Plate 321 from Bloch (1792) . The type locality given by Bloch as “japonischen Meers” is almost certainly wrong, and records of this species from Japan (e.g., Masuda et al. 1984 ) are misidentifications of Scolopsis japonica . Bloch sometimes incorrectly reported fishes from India and the East Indies as coming from Japan ( Cuvier & Valenciennes 1830 ; Paepke 2001 ), and in the case of S. vosmeri evidence points to the type being collected from India , where this species also is known to occur today. Bloch later corrected some of his localities ( Paepke 2001 ), and for Anthias vosmari [sic] Bloch & Schneider (1801) reported the collection locality as “Habitat cum antecedente”, referring to that of the previously described species, Anthias macrophthalmus Bloch, 1792 [= Priacanthus hamrur ( Forsskål, 1775 ) ], whose locality was given as Tranquebar (= Tharangambadi ) in the Indian state of Tamil Nadu . Bloch’s naming of his species Anthias vosmeri in honour of Arnout Vosmaer (1720–1799) suggests his specimen could have come from Vosmaer, who was keeper of the Dutch Stadtholder’s collection of William V, Prince of Orange, in Voorburg, Holland ( Pieters 1980 ) and was one of several scientists known to have contributed specimens for Bloch’s Allgemeine Naturgeschichte der Fische ( Wells 1981 ). Other specimens of Scolopsis vosmeri are known to have existed in this important Dutch collection at the time. Lacepède (1802) based his description of Pomacentrus enneadactylus on two dried skins (MNHN A-8089), type locality ‘Mer des Indes’, which formed part of the Stadtholder’s collection of dried fish (acquired by the Muséum national d'histoire naturelle following the French conquest of Holland in 1795, along with other natural history collections of the Batavian Republic — Lipkowitz 2014 ). Although largely uniform brown in color, Lacepède’s specimens lack the black wedge-shaped spot on the upper pectoral fin-base and are identifiable as Scolopsis vosmeri ( Bauchot et al . 1983 ) . Specimens described by Cuvier in Cuvier & Valenciennes (1830) as Scolopsides kate Cuvier, 1829 (= Scolopsis japonica —see Remarks under that species below), based on specimens collected from Pondicherry (= Puducherry ), India , by Leschenault are also identifiable as Scolopsis vosmeri . Cuvier (1830) gave the fresh color (provided by Leschenault) as “gris legérèment vineuse” [grey, slightly wine colored]. He also mentions the presence of a silvery band below the lateral line: “et l’on y voit la trace d’un bande plus argenté” [one sees traces of a silvery band] which is still faintly visible in one of the specimens (MNHN A.8093). The specimens of Scolopsides kate from Pondicherry were also recognized by Bleeker (1873: 362) as Scolopsis vosmeri . Scolopsis igcarensis Mishra et al ., 2013 , collected from coastal waters of southern India and Sri Lanka , has recently been shown to represent juvenile and subadult forms of S. vosmeri ( Russell et al. 2019 ) and also is included in the synonymy of S. vosmeri . The English common name Vosmaer’s monocle bream is proposed for this species.