Zootaxa 4629 (4): 589–599 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4629.4.7 http://zoobank.org/urn:lsid:zoobank.org:pub:2BBBD373-C998-4581-828B-1EE3DBC65669

Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy & Selvanayagam, 2013, a junior synonym of S. vosmeri (Bloch, 1792) (: Nemipteridae)

BARRY C. RUSSELL1, 4, MOHAMMAD EUSUF HASAN2 & JEAN-DOMINIQUE DURAND3 1Museum & Art Gallery of the Northern Territory, GPO Box 4646 Darwin NT 0801, Australia 2Institute of Marine Sciences, University of Chittagong, Chattogram 4331, Bangladesh 3MARBEC, IRD-Ifremer-CNRS—Université de Montpellier, cc093, Place E. Bataillon, 34095 Montpellier Cedex 05, France 4Corresponding author: [email protected]

Abstract

Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy & Selvanayagam, 2013 was described from specimens collected from coastal waters of southern India and Sri Lanka. A comparison of recently collected specimens from Bangladesh, initially identified as S. igcarensis, with Scolopsis vosmeri (Bloch, 1792) showed morphological differences between the two species are minor, and that specimens of S. igcarensis in fact represent juvenile and subadult colour forms of S. vosmeri. Underwater and aquarium observations, as well as molecular data based on the COI barcode region, support this conclusion. Accordingly, S. igcarensis is regarded as a junior synonym of S. vosmeri, which is redescribed herein. Phylogenetic analysis of COI barcodes of Scolopsis specimens produced in this study, together with those available from GenBank, indicate S. vosmeri is part of a species complex which includes two additional cryptic sister species that require further taxonomic investigation.

Key words: Synonymy, , morphology, phenotypes, COI, molecular phylogeny

Introduction

Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy & Selvanayagam, 2013 was described from specimens col- lected from coastal waters of southern India and Sri Lanka. The species was distinguished from Scolopsis vosmeri (Bloch, 1792) by having a more robust suborbital spine; bony ridge beneath eye less prominent, covered with skin; pectoral fins not reaching to level of anus; and its lack of a white cheek band (Mishra et al. 2013). Recent collection of fresh specimens from the Bangladesh coast, northern Bay of Bengal, as well as underwater and aquarium observations, however, showed morphological differences with Scolopsis vosmeri are minor and the characters distinguishing S. igcarensis and S. vosmeri are somewhat variable, thus casting doubt on the taxonomic validity of the former species. In this paper we present morphological evidence and molecular data, based on the COI barcode region, that S. igcarensis in fact represents juvenile and subadult forms of S. vosmeri. Accordingly, we redescribe S. vosmeri and recognise S. igcarensis as a junior synonym.

Material and methods

Underwater observations of Scolopsis vosmeri undertaken by the first author in Myanmar during biodiversity sur- veys of the Myeik Archipelago in December 2014 (Russell 2015) and February 2016 (Russell 2016), were docu- mented using high definition underwater video (Sony Action Cam). Aquarium observations by the second author were made at Saint Martin’s Island, Bangladesh, and were supplemented by the collection of a range of different sized individuals, which were deposited in the collections of the United States National Museum (USNM) and University of Chittagong, Institute of Marine Sciences, Bangladesh (CUMS). Counts and measurements of speci- mens follow Russell & Gloerfelt-Tarp (1984) and were made on the left side wherever possible. All measurements

Accepted by W. Holleman: 23 May 2019; published: 9 Jul. 2019 589 are expressed in millimeters rounded to the nearest 0.1 mm. Proportional measurements of specimens are given as percentage of standard length (SL) or head length (HL) unless otherwise indicated. Institutional abbreviations for specimens examined follow Sabaj (2016). Species delineation as well as the intraspecific genetic diversity were estimated using the nucleotide variation of a portion the cytochrome c oxidase I (COI) region. This gene fragment was chosen since it is used as a DNA barcode for fish species in the worldwide barcoding initiative, FishBOL. Genomic DNA was extracted with the G-Spin Total DNA extraction mini kit (iNtRON biotechnology, Korea) from fins or muscle of four young individuals of Scolopsis vosmeri (initially identified as S. igcarensis) collected at Saint Martin’s Island, Teknaf, Cox’s Bazar, Bangladesh. Additional specimens of Scolopsis taenioptera (Cuvier in Cuvier & Valenciennes), S. vosmeri and S. monogramma (Cuvier in Cuvier & Valenciennes) collected from the Andaman Sea coast of Thailand were also analysed. Frag- ments of the COI were amplified by polymerase-chain reaction (PCR) using primers FishF1+FishF2 and FishR1; following PCR conditions described in Ward et al. (2005). Sequencing was performed by Macrogen Inc. (Seoul, South Korea; http://dna.macrogen.com). All new sequences obtained in this study were deposited in GenBank (Ac- cession numbers: MK757264–MK757266 and MK779327–MK779330) and added to a large Scolopsis species COI library available in GenBank, and previously used in the study by Hung et al. (2016) to highlight OTUs in the Scolopsis. Sequences of S. vosmeri from the Bay of Bengal and Andaman Sea were mined from GenBank (until December 2018) and used in the Neighbour Joining analysis. COI sequences of Parascolopsis melanophrys Russell & Chin, P. tosensis (Kamohara), Scaevius milii (Bory de Saint-Vincent), Pentapodus emeryii (Richardson) and P. setosus (Valenciennes in Cuvier & Valenciennes) were used as outgroups to root the tree. Phylogenetic relationships among Scolopsis species were depicted with a phylogenetic analysis based on the Neighbour-Joining (NJ) method implemented in MEGA 7 (Kumar et al. 2016). Evolutionary distances were computed using the Kimura 2-parameter method (Kimura 1980). Nodal support was assessed by bootstrapping, based on 500 pseudo-replicates. Specimens examined (n = 62, 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, Kara- chi. 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, India; CMFRI GB.31.98.5.6 (paratype of S. igcarensis), 148.1 mm SL, Tuticorin, Tamil Nadu, India; NTM S.11136-001, 116.1 mm SL, Madras (= Chen- nai); ZMB 8729* (holotype of Anthias vosmeri Bloch, 1792 left skin, in 3 parts), indeterminate SL, ‘japonisches Meer’ (erroneous, probably Tranquebar [= Tharangambadi], India: See Distribution below); ZSI F-331–333*, 3: 69.0–153.0 mm SL, Madras (= Chennai), (specimens figured in Day 1875); ZSI F-401*, c.57 mm SL, Bombay (= Mumbai) (specimen figured as Scolopsis leucotaenia (Bleeker) in Day 1875); 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, India; ZSI F-10624/2 (paratypes of S. igcarensis), 5: 75.9–115.6 mm SL, Kalpakkam, Tamil Nadu, India; ZSI F-10625/2 (paratypes of S. igcarensis), 3: 67.1–82.4 mm SL, Kalpakkam, Tamil Nadu, India. 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. MK779330), 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. MK779329), 96 mm SL, Saint Martin’s Island Jetty, Teknaf, Cox’s Bazar; USNM 443789 (GenBank No. MK779328), 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. MK779327), 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 13178, 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, 160 mm SL. Pulau Punyit.

590 · Zootaxa 4629 (4) © 2019 Magnolia Press RUSSELL et al. Results

Counts and proportional measurements of Scolopsis vosmeri and the type series of S. igcarensis are shown in Table 1. There are no clear differences between counts and proportional measurements of S. vosmeri and the types of S. igcarensis, and the main distinguishing features of S. igcarensis—a more robust suborbital spine; bony ridge beneath eye less prominent, covered with skin; and pectoral fins not reaching to level of anus (Mishra et al. 2013) appear to be characters that are largely those of juveniles of S. vosmeri. Moreover, the lack of a white cheek band in S. igcarensis, stated by Mishra et al. (2013) as always present in S. vosmeri, is a feature of the juvenile colouration of specimens of S. vosmeri observed by us, and a character that changes with growth, as detailed in the following redescription of S. vosmeri.

Taxonomy

Family Nemipteridae

Genus Scolopsis Cuvier, 1814

Scolopsis vosmeri (Bloch, 1792) English name: White-banded monocle bream Table 1, Figs 1–11

Anthias vosmeri Bloch, 1792: 120, Pl. 321 (type locality: ‘japonisches Meer’, but probably Tranquebar, India; see Remarks, below). Scolopsis leucotaenia (non Bleeker)—Day 1875: 88, Pl. XXIII, fig. 4. Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy & Selvanayagam, 2013: 444 (India: coasts of Tamil Nadu, Kerala, and Mumbai; Sri Lanka). Scolopsis vosmeri—Day 1875: 87 (in part, Eastern Indian Ocean Specimens), Pl. XXIII, fig. 3; Russell 1984: NEMIP Scol 2 (in part, Eastern Arabian Sea Specimens); Russell 1990: 122, Pl. VIII, fig. g (in part, Eastern Indian Ocean specimens, from Pakistan to Malacca strait, West of Borneo); Russell 2001: 3089 (in part, Andaman Sea, Malacca Strait, and lower South China Sea); De Bruin et al. 1995: Pl. XXIII, Fig. 159 (Sri Lanka Coast); Barman & Mishra 2009: 36, 50 (India); Psomadakis et al. 2015: 272, Pl. XXV, fig. 199 (Pakistan Coast).

Diagnosis. A species of Scolopsis with the following combination of characters: temporal region scaly, scales on top of head reaching forward to just in front of anterior nostrils; suborbital naked; lower margin of eye tangent to or just above a line from snout tip to upper pectoral-fin base; exposed bony ridge below eye bearing an antrorse and retrorse spine; scales on preopercle extending to posterior margin, 5–6 rows of scales behind suborbital and 3–4 rows behind eye; lower limb of preopercle usually with single row of embedded scales; 4–7 rows of scales on opercle; 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, these features not present in juveniles and smaller subadults; upper limb of opercular margin narrowly brown, lower limb narrowly reddish-orange; a small dark spot on upper base of pectoral fin; pelvic and anal fins crimson or orange-red. Description. Counts and proportional measurements as percentage of SL are provided in Table 1. Morphomet- ric characters given in the diagnosis above are not repeated. Dorsal-fin rays X,9; anal-fin rays III,7; pectoral-fin rays 18–19; pelvic-fin rays I,5; lateral line scales 40–44; 3½–4½ transverse scale rows above lateral line, 13–15½ rows below; gill rakers on first gill arch 9–13 (usually 11); vertebrae 22. Body moderately deep, laterally compressed, depth 36.9–51.1 % SL; dorsal profile of head more convex than ventral contour, head length 29.3–37.1 % SL; snout short, 24.5–33.2 % HL; eye diameter about equal to or a little more than snout length, 0.6–1.1 in snout; interorbital width 1.0–1.9 in eye diameter; suborbital depth 1.8–4.2 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; posterior margin of preopercle serrated or den- ticulate on posterior margin; posterior margin of opercle mostly smooth, a short, flat, pointed or rounded spine at

Scolopsis igcarensis junior synonym of S. vosmeri Zootaxa 4629 (4) © 2019 Magnolia Press · 591 upper angle; teeth in jaws conical, pointed, and slightly recurved; in several bands anteriorly on upper jaw; sides of lower jaw with a single series of teeth anteriorly that forms a band of 3 or 4 rows; lateral surface of maxilla smooth; gill rakers short and stubby, 9–13 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 1.4–2.5 times that of first and 1.1–1.5 times that of third; pectoral fins reaching to or just before of level of anus, 24.0–29.2 % SL; pelvic fins moderately long, reaching beyond anus, 22.4–32.1 % SL; caudal fin forked, lobes slightly rounded; lower lobe a little shorter than upper; body covered with ctenoid scales.

FIGURE 1. Scolopsis vosmeri, juvenile c.45 mm SL, underwater, Myeik Archipelago, Myanmar (from video clip, B.C. Rus- sell).

FIGURE 2. Scolopsis vosmeri, subadult, USNM 443789, 77 mm SL, Bangladesh (photo: M.E. Hasan).

592 · Zootaxa 4629 (4) © 2019 Magnolia Press RUSSELL et al. TABLE 1. Selected counts and morphometric data for type series of Scolopsis igcarensis (from Mishra et al. 2013) com- pared with other specimens of S. vosmeri examined here. Scolopsis igcarensis (Type series, n=13) Scolopsis vosmeri (n=37) SL 67.1–148.1 mm 77.0–173.1 mm Counts Lateral line scales 40–44 40–44 Scales above/below lateral line 3½ /13–15½ 3½–4½ /13–15½ Gill rakers (total) 10–13 9–12 Scale rows on opercle 5 4–7 Scale rows on preopercle behind eye 3–4 3–6 Pectoral fin rays 15–18 16–19 Proportions Body depth (% SL) 36.9–50.9 39.8–51.1 Head length (% SL) 29.7–34.1 29.3–37.1 Snout length (% HL) 24.5–30.2 24.9–33.2 Eye diameter (% HL) 29.9–40.9 28.2–37.1 Eye in snout 0.6–0.9 0.8–1.1 Interorbital width in eye 1.0–1.9 1.0–1.7 Suborbital depth in eye 2.0–4.4 1.8–3.0 Caudal-peduncle depth in peduncle length 0.8–1.3 0.7–1.7 Base of dorsal fin (% SL) 54.9–62.7 56.9–62.8 Base of anal fin (% SL) 16.0–20.0 15.9–21.0 Pectoral fin length (% SL) 24.6–29.2 24.0–28.1 Pelvic fin length (% SL) 25.1–31.1 22.4–32.1 Pectoral length in pelvic length 0.8–1.1 0.8–1.2 First dorsal spine in longest dorsal spine 1.6–2.3 1.2–2.3 Longest dorsal spine in longest dorsal ray 1.0–1.3 1.0–1.2 First anal spine in second anal spine 1.7–2.5 1.4–2.3 Third anal spine in second anal spine 1.2–1.5 1.1–1.5

FIGURE 3. Scolopsis vosmeri juvenile c.40 mm SL (above) and subadult c.80 mm SL (below) underwater, Myeik Archipelago, Myanmar (from video clip, B.C. Russell).

Life colours, juveniles (< c.60 mm SL): reddish brown, darker on back; paling light pinkish to white on lower part of body; a narrow white band along base of dorsal fin; a distinctive white lateral band from behind upper margin

Scolopsis igcarensis junior synonym of S. vosmeri Zootaxa 4629 (4) © 2019 Magnolia Press · 593 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 pinkish; dorsal fin with traces of dusky medial band along its length (Figs. 1, 3) Subadults (c. 60–80 mm SL): colour as for juveniles, but dark bands bordering white lateral band becoming less distinct; white bar on nape incompletely formed, not extending onto operculum; caudal peduncle becoming uniformly pale whitish; fins reddish (Figs. 2, 3, 4, 6).

FIGURE 4. Paratype of Scolopsis igcarensis, ZSI F-10625/2, 79.7 mm SL, Kalpakkam, India (photo from Mishra et al. 2013).

Adults (> c.80 mm SL):Reddish brown to dark brown, with a black spot basally on each scale except for breast and caudal peduncle, shading to white on caudal peduncle; margins of scales on side of body with greenish hue; a white or yellowish white lateral band from origin of lateral line to below posterior part of dorsal fin (this band occasionally faded or absent); head reddish brown with a curving white bar extending from nape onto operculum, broader ventrally; posterior edge of opercle above opercular spine narrowly reddish brown; opercular membrane below opercular spine narrowly orange-red; posterior edge of preopercle pale orange; suborbital broadly whitish with white adjacent area behind angle of mouth; eye red suffused, with yellow blotch present on upper part; a small black spot on upper base of pectoral fin; dorsal fin reddish brown, usually with brighter red margin; pectoral, pelvic and anal fins crimson or orange-red; anal spines whitish; caudal fin pale reddish, upper and lower edges darker, (Fig. 5, 7).

FIGURE 5. Scolopsis vosmeri, adult, BPBM 27195, 155.1 mm SL, Hikkaduwa, Sri Lanka (photo: J.E. Randall).

594 · Zootaxa 4629 (4) © 2019 Magnolia Press RUSSELL et al. Distribution. Scolopsis vosmeri is known from the northern and eastern Indian Ocean from Pakistan, India, Sri Lanka, Bangladesh, Myanmar and Thailand (Andaman Sea), to western Indonesia and Borneo. It inhabits shallow inshore rock and coral reefs, often in turbid water, in depths to about 30 m.

FIGURE 6. Scolopsis vosmeri, subadults c.80 mm SL underwater, showing ability to ‘flash’ lateral band on (red arrow, upper left) and off (red arrow, lower right). Myeik Archipelago, Myanmar (from video clip, B.C. Russell).

Comparisons. Scolopsis vosmeri is distinguishable from all other species of Scolopsis by having a distinctive white or yellowish-white lateral band. This band extends from behind the eye to the upper caudal peduncle and is margined above and below by darker bands in juveniles and subadults; in adults, it extends from behind the upper part of the opercle to below the posterior part of dorsal fin. Underwater observations of this species in Myanmar (Fig. 6) and in aquarium in Bangladesh (Fig. 7) show that subadults and adults can actively ‘flash’ the white lateral band on and off while swimming, perhaps as a signal to conspecifics.

FIGURE 7. Scolopsis vosmeri, adults c.150 mm SL (upper) and c. 100 mm SL (lower) showing ability to ‘flash’ lateral band on and off. Photographed in aquarium, St Martins Island, Cox’s Bazaar, Bangladesh (photo: M.E. Hasan).

Scolopsis igcarensis junior synonym of S. vosmeri Zootaxa 4629 (4) © 2019 Magnolia Press · 595 Genetics. Genetic analysis of juvenile specimens initially identifiable as Scolopsis igcarensis, collected from Bangladesh, show them to be indistinguishable from some adult specimens of S. vosmeri from Bangladesh (MK779330 and MH311294), Western Thailand (ACRSD-F174) and Myanmar (MH235710) (mean within clade divergence of 0.1%). Hung et al. (2016) recognised two genetically and geographically distinct (West Pacific vs Indian Ocean) lineages for the species assigned to Scolopsis vosmeri. These specimens, identified as S. vosmeri, uploaded from GenBank, form two sister clades that present 12.7% and 13.9% of divergence from S. vosmeri from Bangladesh, Myanmar and Thailand (Fig. 8). These two cryptic sister Scolopsis species, separated by 4.2% of divergence, are provisionally named here Scolopsis sp. A and Scolopsis sp. B, form part of a species complex that has been gener- ally referred to as S. vosmeri (Russell 1984, 1990, 2001) and will be dealt with separately in a forthcoming paper (Russell et al. in prep.).

FIGURE 8. Neighbour-Joining (NJ) tree showing phylogenetic relationships among Scolopsis species. The Scolopsis vosmeri complex comprises three species: S. vosmeri sensu stricto and a pair of sister species, from the NW Pacific–S. China Sea (Scol- opsis sp. A) and Indian Ocean (Scolopsis sp. B) respectively.

Remarks. Bloch’s (1792: 120, Plate 321) description and figure of Anthias vosmeri (Fig. 9) clearly shows the reddish body and fins, pale band extending from the nape across the opercle (incompletely shown in Bloch’s fig- ure) and yellowish band beneath the lateral line, that are characteristic of this species. However, the type locality, “japonischen Meers”, given by Bloch is almost certainly wrong. In many cases Bloch incorrectly reported fishes from India and the East Indies as coming from Japan (Cuvier & Valenciennes 1830; Paepke 1999, 2001), but later corrected some of his localities (Paepke 2001), with Bloch & Schneider (1801) subsequently reporting the locality

596 · Zootaxa 4629 (4) © 2019 Magnolia Press RUSSELL et al. of Anthias vosmeri, as “Habitat cum antecedente”, referring to that of a previously described species, Anthias mac- rophthalmus Bloch [= Priacanthus hamrur (Forsskål)], whose locality was given as Tranquebar (= Tharangambadi) in the Indian state of Tamil Nadu. Thus, it is likely that the type specimen of Scolopsis vosmeri was collected from south-eastern India, where this species also is known to occur today.

FIGURE 9. Anthias vosmeri Bloch (from Bloch 1792: Pl. 321).

Day (1875: Plate XXIII, Fig. 1, 2, 3) provided three figures of Scolopsis vosmeri indicating three life stages: young, semi-adult, and adult respectively. The adult stage (Fig. 10) shows the horizontal white band and thus conforms to S. vosmeri, while the others represent different species. Day (1875: Plate XXIII, fig. 4) also figured a specimen from Bombay (= Mumbai) as Scolopsis leucotaenia (Bleeker) (Fig. 11) that closely resembles the colour pattern of juveniles of S. vosmeri described here. As pointed out by Mishra et al. (2013), the true S. leucotaenia is the juvenile form of (Cuvier in Cuvier & Valenciennes) and differs in colour from S. vosmeri in having a distinct black spot anteriorly on the dorsal fin between the first three rays (absent in Day’s Plate XXIII, fig. 4), and the uppermost dark band on the body above the lateral line (versus band mostly on or below the lateral line in Day’s figure and juveniles of S. vosmeri). Scolopsis leucotaenia (= margaritifera) also differs from S. vosmeri in having the head scales reaching forward only to the anterior margin of the eye (versus head scales extending forward to or in front of anterior nostrils in S. vosmeri); fewer lateral-line scales (34–39, versus 42–43); longer pelvic fins, that reach almost to or beyond the anal fin origin (versus pelvics reaching only to just beyond anus); centre of the scales on the body with pearly or whitish spots (versus body scales dark brown in middle); and the second anal spine not longer and stronger than the third spine. Day’s figured specimen (ZSI F-401), c.57 mm SL, is damaged but has the head scaled to the front of anterior nostril, a prominent suborbital ridge beneath the eye (right side), at least 40 lateral-line scales, and 17 pectoral-fin rays—characters that are consistent with S. vosmeri. Thus, we consider Day’s record of S. leucotaenia to also represent the juvenile form of S. vosmeri.

Acknowledgements

We thank Dr. John E. Randall, (Bernice P Bishop Museum, Honolulu) for providing photographs and specimens; S. Biswas and K.K. Satpathy, (Indira Gandhi Centre for Atomic Research, Kalpakkam), S.S. Mishra (Zoological Sur- vey of India, Kolkata), K.K. Bineesh (Zoological Survey of India, Port Blair), and K.V. Akhilesh (Central Marine Fisheries Research Institute, Mumbai) for access to collections and specimens in India; M. McGrouther (Australian Museum, Sydney), S. Bogorodsky and T. Alpermann (Senckenberg Naturmuseum, Frankfurt), G. Moore and G.R.

Scolopsis igcarensis junior synonym of S. vosmeri Zootaxa 4629 (4) © 2019 Magnolia Press · 597 Allen (Western Australian Museum, Perth), and P. Bartsch (Museum für Naturkunde, Berlin) for access to collec- tions and specimens in their care. We are grateful to R. Howard (Flora & Fauna International, Yangon) for providing opportunity for the first author to participate in diving surveys in Myanmar. Sahat Ratmuangkhuang (Andaman Coastal Research Station for Development, Kasetsart University, Ranong) kindly provided tissue samples of several Andaman Sea Scolopsis specimens. The second author thanks M.Z.R. Chowdhury, Director, Institute of Marine Sci- ences, University of Chittagong, Chattogram, for his cooperation.

FIGURE 10. Scolopsis vosmeri, adult, figured in Day (1875: Pl. XXIII, fig. 3); note the horizontal white stripe characteristic of S. vosmeri.

FIGURE 11. Scolopsis vosmeri, juvenile, figured in Day (1875: Pl. XXIII, fig. 4, as S. leucotaenia).

References

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598 · Zootaxa 4629 (4) © 2019 Magnolia Press RUSSELL et al. Bloch, M.E. (1792) Naturgeschichte der ausländischen Fische. Vol. 6. Auf Kosten des Verfassers und in Commission bei dem Buchhändler Hr. Hesse, Berlin, xii + 126 pp., pls. 289–323. Bloch, M.E. & Schneider, J.G. (1801) M.E. Blochii, Systema Ichthyologiae Iconibus cx Ilustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo. Sumtibus Auctoris Impressum et Bibliopolio Sanderiano Commissum, Berolini, lx + 584 pp., 110 pls. https://doi.org/10.5962/bhl.title.5750 Cuvier, G. (1814) Observations et recherches critiques sur différens poissons de la Méditerranée et, à leur occasion, sur des poissons des autres mers plus ou moins liés avec eux. [In a report by A.D. = A. G. Desmarest.] Bulletin des Sciences, par la Société Philomathique de Paris, Série 3, 1, 80–92. Cuvier, G. & Valenciennes, A. (1830) Histoire naturelle des poissons. Tome cinquième. Livre cinquième. Des Sciénoïdes. Vol. 5. Chez F. G. Levrault, Paris, xxviii + 499 + 4 pp., pls. 100–140. Day, F. (1875) The Fishes of India, being a natural history of the fishes known to inhabit the seas and freshwater of India, Burma and Ceylon. William Dawson & Sons, London. Part 1, 168 pp., 1–40 pls. [Dates of publication of parts of this work from Menon & Rama Rao (1974), see below] De Bruin, G.H.P., Russell, B.C. & Boguscb, A. (1995). FAO Species identification field guide for fishery purposes. The Marine fishery resources of Sri Lanka. FAO, Rome, 400 pp., 32 pls. Hung, K.-W., Russell, B.C. & Chen, W.-J. (2016) Molecular systematics of threadfin breams and relatives (Teleostei, Nemip- teridae). Zoologica Scripta, 46 (5), 536–551. Kimura, M. (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120. Kottelat, M. (2011) Pieter Bleeker in the Netherlands East Indies (10 March 1842—ca. 21 September 1860): new biographical data and a chronology of his zoological publications. Ichthyological Exploration of Freshwaters, 22 (1), 1–94. Kumar, S., Stecher, G., Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7 for bigger dataset. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054 Menon, A.G.K. & Rama Rao, K.V. (1974) On the dates of the parts of Day’s `Fishes of India’. Journal of the Society for the Bibliography of Natural History, 7 (1), 143. https://doi.org/10.3366/jsbnh.1974.7.1.143 Mishra, S.S., Biswas, S., Russell, B.C., Satpathy, K.K. & Selvanayagam, M. (2013) A new species of the genus Scolopsis Cuvier, 1830 (Perciformes: Nemipteridae) from southern India and Sri Lanka. Zootaxa, 3609 (4), 443–449. https://doi.org/10.11646/zootaxa.3609.4.7 Paepke, H.-J. (1999) Bloch’s fish collection in the Museum für Naturkunde der Humboldt Universität zu Berlin: an illustrated catalog and historical account. Ruggell (Liechtenstein). Theses Zoologicae, 32, 1–216, pls. 1–32. [unpaginated] Paepke, H.-J. (2001) Comments on the old Japanese fish collections in the Museum of Natural History of the Humboldt University of Berlin. Ichthyological Research, 48 (3), 329–334. https://doi.org/10.1007/s10228-001-8155-0 Psomadakis, P.N., Osmany, H.B. & Moazzam, M. (2015) Field identification guide to the living marine resources of Pakistan. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, x + 386 pp., 42 colour pls. Russell, B.C. (1984) Family Nemipteridae. In: Fisher, W. & Bianchi, G. (Eds.), FAO Species Identification Sheets for Fisheries Purposes. Western Indian Ocean. Fishing Area 51. Vol. 4. FAO, Rome, pp. 1–6. [“NEMIP Nem 2” to “NEMIP Scol 8”] Russell, B.C. (1990) FAO species catalogue. Vol. 12. Nemipterid Fishes of the World. (Threadfin breams, Whiptail breams, Monocle breams, Dwarf monocle breams, and Coral breams). Family Nemipteridae. An Annotated and Illustrated Catalogue of Nemipterid Species known to Date. In: FAO Fisheries Synopsis. No. 125. Vol. 12. Rome, FAO, pp. 1–149, VIII pls. Russell, B.C. (2001) Family Nemipteridae. In: Carpenter, K.E. & Niem, V.H. (Eds.), FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Vol. 5. Bony fishes part 3 (Menidae to Pomacentri- dae). FAO, Rome, pp. 3051–3089. Russell, B.C. (2015) Survey of coral reef fishes of the Myeik Archipelago, Myanmar. In: Report No. 13 of the Tanintharyi Conservation Programme, a joint initiative of Fauna and Flora International (FFI) and the Myanmar Forest Department. FFI, Yangon, pp. 1–37. Russell, B.C. (2016) Survey of coral reef fishes of the Myeik Archipelago, Myanmar. In: Report No. 38 of the Tanintharyi Conservation Programme, a joint initiative of Fauna and Flora International (FFI) and the Myanmar Forest and Fisheries Departments. FFI, Yangon, pp. 1–57. Russell, B.C. & Gloerfelt-Tarp, T. (1984) A new species of Parascolopsis (Pisces: Nemipteridae) from Indonesia. Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences, 1, 111–114. Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5. American Society of Ichthyologists and Herpetologists, Washington, D.C. Available from: http:// www.asih.org/ (accessed 16 August 2016) Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R. & Hebert, P.D.N. (2005) DNA barcoding Australia’s fish species. Philosophi- cal transactions of the Royal Society of London, Series B, 360 (1462), 1847–1857. https://doi.org/10.1098/rstb.2005.1716

Scolopsis igcarensis junior synonym of S. vosmeri Zootaxa 4629 (4) © 2019 Magnolia Press · 599