Zootaxa 3936 (4): 575–583 www.mapress.com /zootaxa / ISSN 1175-5326 (print edition) Article Copyright © 2015 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3936.4.7 http://zoobank.org/urn:lsid:zoobank.org:pub:9F14082D-3680-4D75-B3F3-398F912ECCDD Redescription of Pethia melanomaculata (Teleostei: Cyprinidae) from Sri Lanka SUDESH BATUWITA1, KALANA MADUWAGE2,3 & HIRANYA SUDASINGHE4,5 1 Postgraduate Institute of Agriculture, University of Peradeniya, Sri Lanka Department of Biochemistry, Faculty of Medicine, University of Peradeniya, Sri Lanka 3 School of Medicine and Public Health, University of Newcastle, Australia 4 Department of Zoology, University of Peradeniya, Sri Lanka 5 Corresponding author. E-mail: hsudasinghe@gmail.com 2 Abstract The name Pethia melanomaculata (Deraniyagala) is available for the Sri Lankan fish previously referred to P. ticto, being distinguished from its Indian congeners by the combination of the following characters; having ½4/1/3½ scales in transverse line on body; body depth 32.4–41.5% of standard length (SL); head length (HL) 26.1–29.2% of SL; snout length 25.3–35.6% of HL; eye diameter 24.4–31.9% of HL; a small black humeral spot on lateral-line scales 3 or 4; a black spot on caudal peduncle, on scales 16–18 of the lateral line series; 3 unbranched dorsal-fin rays, the last one strongly serrated, with 8–11 serrae. Key words: Pethia ticto, Pethia punctata, Puntius, India, taxonomy Introduction Pethia ticto was described from southern Bengal (now in Bangladesh and West Bengal, India) by Hamilton (1822). This species, commonly identified from its small, black humeral spot and the larger black spot or blotch on the caudal peduncle, has been reported from India, Myanmar and Sri Lanka (Hora et al. 1939; Jayaram 1991, 1999; Talwar & Jhingran 1991). Several studies have shown, however, that the fishes until recently identified as P. ticto in many parts of South Asia in fact represent distinct species (Linthoingambi & Vishwanath 2007; Mercy & Jacob 2007; Knight et al. 2012; Dishma & Vishwanath 2013; Katwate et al. 2014a,b; Lalramliana et al. 2014). Deraniyagala (1956, 1958) proposed the subspecies name Puntius ticto melanomaculatus (type locality Kantalai Reservoir, Eastern Province, Sri Lanka) for Sri Lanka’s ‘Ticto barb’, since uncritically synonymized with P. t. ticto by most authors (e.g., Mendis & Fernando 1962; Senanayake 1980; Pethiyagoda 1991, 2006; Jayaram 1999), possibly because of the very brief description provided by Deraniyagala (1956), who did not have access to comparative material from India. Molecular data presented in Meegaskumbura et al. (2008) and Pethiyagoda et al. (2012), however, suggest that the Sri Lankan fish may be distinct from P. ticto. Here we redescribe Pethia melanomaculata based on the examination of fresh material from Sri Lanka and distinguish it from its Sri Lankan and peninsular-Indian congeners. Material and methods Specimens referred to in this work are deposited in the National Museum of Sri Lanka, Colombo (NMSL); the collection of the Wildlife Heritage Trust of Sri Lanka (WHT) now at NMSL; the Southern Regional Station of the Zoological Survey of India, at Chennai (ZSI-SRS); and Department of Zoology, University of Peradeniya, Sri Lanka (DZ). Measurements were made with digital Vernier callipers to the nearest 0.1 mm. Drawings were made using a Motic dissecting stereomicroscope with camera lucida. Methods for taking counts and measurements follow Accepted by R. Pethiyagoda: 23 Feb. 2015; published: 23 Mar. 2015 575 Pethiyagoda & Kottelat (2005), except that the insertion of a fin is the posterior extremity of its base; and scales in transverse line on body were counted downwards and backwards from the dorsal-fin origin to the mid-ventral scale row anterior to the anal-fin origin. Head Length (HL) and measurements of the body are presented as proportions of standard length (SL), while subunits of the head are indicated as proportions of head length. Values in parenthesis after a count represent the frequency of that count. Osteological descriptions are based on specimens cleared and single-stained using the method of Taylor & Van Dyke (1985); osteological nomenclature follows Conway (2011). Pethia melanomaculata (Deraniyagala, 1956) (Figures 1A, 2, 3A) Puntius ticto melanomaculatus Deraniyagala, 1956: 34; 1958: 129. Barbus ticto (not Hamilton, 1822): Day, 1878: 576; Senanayake, 1980: 118. Puntius ticto (not Hamilton, 1822): Deraniyagala, 1952: 35; Munro, 1955: 46; Pethiyagoda, 1991: 118. Puntius melanomaculatus (Deraniyagala, 1956): Meegaskumbura et al., 2008: 148. Pethia melanomaculata (Deraniyagala, 1956): Pethiyagoda et al., 2012: 71. Diagnosis. Pethia melanomaculata is distinguished from all the other congeners in Sri Lanka and the peninsula of India by the combination of the following characters: barbels absent; lateral line incomplete, with 7–10 perforated scales; 20–23 scales in lateral series; 3 unbranched dorsal-fin rays, the last one with 8–11 strong serrae; ½4/1/3½ scales in transverse line on body; ½2/1/2½ scales in transverse line on caudal peduncle; predorsal scales 8–9; body depth 32.4–41.5% SL; head length 26.1–29.2% SL; snout length 25.3–35.6% HL; eye diameter 24.4–31.9% HL; a small black humeral spot on the 3rd or 4th lateral-line scales; a black spot on caudal peduncle, on scales 16–18 of the lateral-line series; no prominent projection at postero-lateral border of premaxilla; superior angle of palatine process of maxilla blunt; dorsal border of anguloarticular curved, with a deep concavity; lateral border of 5th ceratobranchial with a deep notch. Description. See Fig. 1A for general appearance and Table 1 for morphometric data. Body laterally compressed. Dorsal profile of head gently notched above posterior orbital margin. Predorsal profile convex, rising gradually to dorsal-fin origin. Postdorsal profile sloping down to hypural notch. Ventral profile convex up to pelvic-fin base, then almost flat to anal-fin origin, rising markedly between anal-fin origin and insertion, then approximately level to hypural notch. Caudal peduncle short, its depth 1.0–1.4 times its length. Head small, laterally compressed. Snout blunt, rounded, subequal to eye diameter. Mouth small, subterminal, angle of gape not reaching vertical from anterior margin of eye. Barbels absent. Eyes large, dorsolaterally oriented, closer to tip of snout than to margin of operculum. Dorsal-fin with its origin posterior to pelvic-fin origin, approximately equidistant from tip of snout and caudalfin base; its distal margin concave; 3 unbranched and 8 branched rays (18), last one strong with 8 (8), 9 (4), 10 (4) or 11 (1) serrae posteriorly. Pectoral fin with 1 unbranched and 11 (2) or 12 (15) branched rays, its tip reaching pelvic-fin base when adpressed. Pelvic fin with 1 unbranched and 8 (16) branched rays, its distal margin rounded, not reaching anal-fin origin when adpressed. Anal fin with 3 unbranched and 5 (17) branched rays, its distal margin concave. Caudal fin forked, with i,8,8,i (1), i,8,9,i (5), i,9,8,i (4), i,9,9,i (7) principal rays. Lateral line incomplete, with 7–10 pored scales; 20 (4), 21 (9), 22 (3) or 23 (2) scales in lateral series. Scales in transverse line on body ½4/1/3½ (18), predorsal scales 8 (16) or 9 (1), prepelvic scales 9 (3) or 10 (2), scales in transverse line on caudal peduncle ½2/1/2½ (18). Precaudal vertebrae 16; caudal vertebrae 14. Supraneurals 4. No prominent projection at postero-lateral border of premaxilla. Superior angle of palatine process of maxilla blunt. Lengths of anterior and superior articular borders of infraorbital 3+4 less than half of its width. Tip of coronoid process of dentary blunt. Dorsal border of anguloarticular curved, with deep concavity. Lateral arm of 5th ceratobranchial narrow, pointed, with a small foramen at its base. Fifth ceratobranchial with a deep notch on its lateral border and 5+3+2 small, conical teeth. Coloration. In preservative, body overall pale brown. A small black humeral spot above pectoral fin, level with mid eye, confined to 3rd or 4th lateral-line scale. Black spot on caudal peduncle larger than humeral spot, somewhat smaller than eye diameter, originating posterior to anal-fin insertion, on scales 16–18 of lateral series. All fins hyaline, without any distinct marks. 576 · Zootaxa 3936 (4) © 2015 Magnolia Press BATUWITA ET AL. FIGURE 1. A, Pethia melanomaculata, colour in life (not preserved); B, copy of iconotype of Cyprinus ticto Hamilton, 1822 (Courtesy Linnean Society of London); C, illustration of Puntius ticto, from Day (1878, pl. 144, fig. 7); D, distribution of Pethia species in Sri Lanka: ● Pethia melanomaculata (location of holotype in red); other known locations (black); {■}, Pethia bandula (purple); P. cumingii (yellow); P. nigrofasciata (green); P. reval (blue). REDESCRIPTION OF PETHIA MELANOMACULATA Zootaxa 3936 (4) © 2015 Magnolia Press · 577 FIGURE 2. A, left dentary (buccal view; CP, coronoid process; AA, anguloarticular); B, left premaxilla (dorsal view); C, left maxilla (buccal view; PP, palatine process); D, left infraorbitals 3+4 (lateral view); E, 5th ceratobranchial: of Pethia melanomaculata (WHT 11066, 32.8 mm SL), P. punctata (WHT 11078, 34.9 mm SL) and P. ticto (WHT 11054, 37.0 mm SL). Scale bar: 1 mm. 578 · Zootaxa 3936 (4) © 2015 Magnolia Press BATUWITA ET AL. FIGURE 3. Position of caudal spot in A, Pethia melanomaculata; B, P. punctata; and C, P. ticto. Scale bar: 1 mm. TABLE 1. Morphometric data for Pethia melanomaculata, holotype, NMSL FF 728/A; and WHT 7719 and WHT 7748 (n=13). Pethia melanomaculata holotype mean (s.d) range 37.4 33.8 (2.3) 31.0–37.4 Total length – 132 (2.3) 128–136 Predorsal length 45.5 55.0 (3.0) 45.5–57.4 Postdorsal length 50.8 54.5 (2.4) 50.8–58.0 Standard length (mm) In percent of standard length Preanal length 65.5 69.9 (2.0) 65.5–72.6 Prepelvic length 46.8 47.5 (1.6) 44.5–50.3 Body depth 32.4 38.3 (2.3) 32.4–41.5 Head length 28.1 27.8 (0.9) 26.1–29.2 Dorsal-fin base length 15.5 17.1 (0.8) 15.5–18.1 Dorsal-fin height 23.0 25.3 (1.5) 23.0–28.4 Anal-fin base length 10.2 11.8 (1.2) 9.7–14.0 Anal-fin depth 17.1 17.0 (1.0) 15.7–18.9 Pelvic-fin length 18.7 21.6 (1.4) 18.7–23.4 Pectoral-fin length 20.1 20.9 (1.5) 16.8–22.4 Caudal-peduncle length 18.7 19.0 (1.9) 16.3–22.5 Caudal-peduncle depth 13.1 15.3 (0.9) 13.1–17.0 Length of upper caudal-fin lobe – 32.4 (5.2) 27.2–46.8 Length of lower caudal-fin lobe – 33.0 (1.9) 29.2–36.7 Length of medial caudal-fin rays 10.4 14.7 (1.8) 10.4–16.5 In percent of head length Snout length 27.6 29.3 (3.1) 25.3–35.6 Orbital diameter 30.5 29.9 (1.9) 24.4–31.9 Interorbital width 33.3 37.6 (2.0) 33.3–39.6 Internarial width 19.0 18.8 (2.9) 12.8–22.0 In life, dorsally light golden brown. Sides of body iridescent silver with scattered melanophores on distal margin of anterior mid-body scales. Position of humeral and caudal spots same as in preserved specimens. Ventrally white. Prominent markings on dorsal fin absent. Fins hyaline to light yellow. Breeding males with a reddish body coloration. REDESCRIPTION OF PETHIA MELANOMACULATA Zootaxa 3936 (4) © 2015 Magnolia Press · 579 Distribution. Pethia melanomaculata is restricted to the northern and eastern dry zone (annual rainfall < 1,500 mm) of Sri Lanka where it is widely distributed in tanks and small rivers, primarily within the Mahaweli River basin (Fig. 1D). It is the only species of Pethia that occurs in the island’s dry zone. Discussion The ‘Puntius ticto’ of most 20th-century authors is now recognized as a species complex comprising of several distinct species, such as P.ticto; P. punctata (Day 1865); P. stoliczkana (Day 1871); P. manipurensis (Menon, Rema Devi & Vishwanath 2000); P. pookodensis (Mercy & Jacob 2007); P. nigripinna (Knight, Rema Devi, Indra & Arunachalam 2012); P. expletiforis (Dishma & Vishwanath 2013); P. lutea (Katwate, Katwate, Raghavan, Paingankar & Dahanukar 2014a); P. rutila (Lalramliana, Knight & Laltlanhlua 2014) and P. longicauda (Katwate, Paingankar, Raghavan & Dahanukar 2014b). Pethia melanomaculata is distinguished from P. ticto (sensu stricto, from southern Bengal) by having ½4/1/3½ (vs. ½4/1/4½) scales in transverse line on body, 3 (vs. 4) unbranched dorsal-fin rays, last unbranched dorsal-fin ray with 8–11 (vs. 15–21) serrae, 20–23 (vs. 22–25) scales in lateral series, and caudal-peduncle spot on scales 16–18 (vs. 17–20) of the lateral series (Fig. 3C). It further differs from P. ticto by the following osteological characters (Fig. 2): prominent projection at postero-lateral border of premaxilla absent (vs. present); superior angle of palatine process of maxilla blunt (vs. pointed); tip of coronoid process of dentary blunt (vs. posteriorly pointed); dorsal border of anguloarticular curved, with a deep (vs. shallow) concavity; lateral arm of 5th ceratobranchial narrow and pointed (vs. wide and blunt); and lateral border of 5th ceratobranchial with a deep (vs. shallow) notch. The identity of P. punctata was clarified by Katwate et al., 2014c. Pethia melanomaculata can be distinguished from P. punctata (sensu Katwate et al., 2014c) by having 20–23 (vs. 23–25) scales in lateral series, humeral spot located on 3rd or 4th (vs. beneath 4th) lateral-line scale, spot on caudal peduncle on scales 16–18 (vs. 19–22) of lateral series (Fig. 3B), and last unbranched dorsal-fin ray with 8–11 (vs. 12–18) serrae. It is further distinguished from P. punctata by the following osteological characters (Fig. 2): prominent projection at postero-lateral border of premaxilla absent (vs. present); superior angle of palatine process of maxilla blunt (vs. pointed); lengths of anterior and superior articular borders of infraorbitals 3+4 less than half of its width (vs. more than half of its width); tip of coronoid process of dentary blunt (vs. posteriorly pointed); dorsal border of anguloarticular curved, with a deep (vs. shallow) concavity; and lateral border of 5th ceratobranchial with a deep notch (vs. notch absent). Pethia melanomaculata differs from P. stoliczkana by having an incomplete (vs. complete) lateral line, 20–23 (vs. 25–26) scales in lateral series, and ½4/1/3½ (vs. ½4/1/5½) scales in transverse line on body (Lalramliana et al. 2014). Pethia melanomaculata differs from P. manipurensis by having ½4/1/3½ (vs. ½3/1/3½) scales in transverse line on body (Menon et al. 2000). It is distinguished from P. pookodensis by having a single black spot (vs. usually two) on the caudal peduncle (Mercy & Jacob, 2007). It can be distinguished from P. nigripinna by having the black caudal-peduncle spot on scales 16–18 (vs. 18–19, appearing as a band, forming a ring around the caudal peduncle) of the lateral series and having ½4/1/3½ (vs. ½4/1/2–2½) scales in transverse line on body (Knight et al. 2012). It differs from P. expletiforis by possessing an incomplete (vs. complete) lateral line and possessing a humeral spot (vs. humeral spot absent: Dishma & Vishwanath 2013). Pethia melanomaculata is distinguished from P. lutea by having the humeral spot located on the 3rd or 4th lateral-line scale (vs. on the 3rd or 4th lateral-line scale and extending one scale up and down, forming a short bar), and the black caudal-peduncle spot located on scales 16–18 (vs. 17–19) of the lateral series (Katwate et al. 2014b). It differs from P. rutila by having an incomplete (vs. complete) lateral line and humeral spot conspicuous (vs. inconspicuous) located on 3rd or 4th (vs. beneath 3rd and 4th) lateral-line scale (Lalramliana et al. 2014). It is distinguished from P. longicauda by having ½4/1/3½ (vs. ½3/1/3½) scales in transverse line on body and a single black spot (vs. usually two) on the caudal-peduncle, on scales 16–18 (vs.16–18 and 20–22) of the lateral series (Katwate et al. 2014b.). Four additional species of Pethia have been recognized from Sri Lanka: P. cumingii (Günther, 1868); P. nigrofasciata (Günther, 1868); P. bandula (Kottelat & Pethiyagoda 1991); and P. reval (Meegaskumbura, Silva, Maduwage & Pethiyagoda, 2008). Pethia melanomaculata is distinguished from all four species by having only two black spots on the side of the body: a humeral spot and a spot on the caudal peduncle (vs. two black bars, one behind the gill opening and another at the posterior extremity of the anal-fin base in P. bandula, P. cumingii and P. 580 · Zootaxa 3936 (4) © 2015 Magnolia Press BATUWITA ET AL. reval, and three black bars with the addition of a vertical bar beneath the dorsal-fin base in P. nigrofasciata), and ½4/1/3½ (vs. ½3/1/3½ in P. cumingii and P. reval; see: Meegaskumbura et al. 2008), and ½4/1/4½ in P. bandula (see: Kottelat & Pethiyagoda 1991) scales in transverse line on body. Deraniyagala (1956, 1958) distinguished the Sri Lankan population he named P. ticto melanomaculatus from P. t. ticto stating that “It differs from the ‘forma typica’ of India in being smaller usually possessing a longitudinal row of black spots, a black spot under origin of dorsal fin and in the dorsal fin lacking red pigment.” He added that he based his diagnosis on Day (1878) and Hamilton (1822). He evidently did not have specimens of P. ticto before him and relied entirely on these earlier descriptions for diagnostic information. In the original description of P. ticto (type locality “south east parts of Bengal”) Hamilton (1822) mentioned the two black spots “on the lateral line above each pectoral fin” and “near the end of the tail”, both clearly visible in his drawing (see Fig. 1B). The “black spot under origin of dorsal fin” mentioned by Deraniyagala (1956) is variable, as is the “row of black dots or dusting along the sides and a black spot under the origin of the dorsal”. These small black dots appear also on or beneath the dorsal fins of some specimens (see Fig.1A, and the photograph on p. 119 of Pethiyagoda 1991). The black spots on the dorsal fin of P. ticto mentioned by Hamilton (1822) are absent in the series of P. melanomaculata we have examined. The illustration of P. ticto in Hamilton (1822: pl. 8, fig. 87, reproduced here from the copy in the Linnean Society of London as Fig. 1B) is consonant with its description (e.g., colour pattern, serration of the last unbranched dorsal-fin ray), but the scalation appears to be incorrectly portrayed. The figure shows this fish as having not more than 15 scales in lateral series, which is extremely low for any species of Pethia, and is likely to be the result of an artist’s or copyist’s error. The transverse scale count (½5+1+3) in the figure is however plausible. In his material from the northeast of India, Day (1878: 577) counted 23–26 lateral-line scales in P. ticto, and 5–6/4–5 scales in transverse line on body (Fig. 1C). Linthoingambi & Vishwanath (2007), too, redescribed P. ticto from the Brahmaputra drainage as having 22–26 lateral-line scales and ½5/1/5½ scales in transverse line on body. The specimens identified as P. ticto by us from Bengal (n = 6) possess 22–25 scales in lateral series and ½4/1/4½ scales in transverse line on body. Thus it is possible that even within the Ganges- Brahmaputra drainage there exists more than a single species referable to P. ticto, stressing the need to stabilize this name through a redescription of the species based on topotypes (see also Lalramliana et al. 2014; Katwate et al., 2014c). Comparative material Pethia cumingii: WHT 0074, 12, 25.7–33.5 mm SL; Sri Lanka; Dombagaskanda, Ingiriya; WHT 1794, 9, 19.5– 37.7 mm SL; WHT 7518, 35.0 mm SL; Sri Lanka, Mawanana, Neluwa. Pethia melanomaculata: holotype, NMSL FF 728/A, 37.4 mm SL; Sri Lanka, Kantalai Reservoir, 08°22’N, 80°59’E, 100 m; WHT 7719 A–E, F–J, 10, 31.2–49.5 mm SL; WHT 7748 A, B, 2, 31.0–33.0 mm SL: Sri Lanka, Hasalaka, near Mahiyanganaya, 07°21’N, 80°57’E; 150 m. DZ 1500, DZ 1501, DZ 3018, DZ 3019, DZ 3020, 27.9–30.2 mm SL; Sri Lanka, Ulhitiya oya, Ulhitiya. Pethia nigrofasciata: WHT 7679, 4, 39.8–33.5 mm SL; Sri Lanka, Athwelthota; WHT 9203, 6, 25.9–32.9 mm SL; Sri Lanka, Walandura, Kuruwita. Pethia punctata: AMS B.7746, 41.5 mm SL, India, Kerala, Kurriavanoor [River]; WHT 7717 G, 35.7 mm SL; WHT 7717 A–F, H–J, 9, 34.3–47.5 mm SL; WHT 7747 A–G, 7, 47.3–50.3 mm SL; India, Netravadi River 1 km downstream of Uppinangadi, Karnataka, 12.83°N, 75.25° E; 16 March 1996. Data from Katwate et al. 2014c. Pethia ticto:WHT 7718 A, B, F–I, 5ex., 30.8–34.6 mm SL; India, Nadia WB Pond, Bengal, WHT 7720, 34.1 mm SL; India, Boncron, West Bengal. Pethia cf. punctata: ZSI/ SRS F. 4310 A–F, 6, 22.5–28.1 mm SL; India, Tamil Nadu: Villianur, near Pondicherry, 11°55’N, 79°50’E, 10 m. REDESCRIPTION OF PETHIA MELANOMACULATA Zootaxa 3936 (4) © 2015 Magnolia Press · 581 Acknowledgments We thank two anonymous reviewers for valuable comments that helped improve the manuscript; Gina Douglas (Linnean Society of London) for permission to reproduce a copy of the iconotype of Cyrpinus ticto Hamilton, 1822, in the Library of the LSL; and Sudath Nanayakkara (WHT) for hospitality. SB thanks Nanda Wickramasinghe, Manori Goonatilake and Chandrika Munasinghe (all of NMSL) for permission to access material in their care. HS thanks the Department of Wildlife Conservation of Sri Lanka for permits to carry out field work. References Conway, K.W. (2011) Osteology of the South Asian genus Psilorhynchus McClelland, 1839 (Teleostei: Ostariophysi: Psilorhynchidae), with investigation of its phylogenetic relationships within the order Cypriniformes. 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