Japan. J. Ichthyol. 40 (4): 405-412, 1994 魚 類 学 雑 誌 40 (4): 405-412, 1 9 94

A New Cyprinid Fish, Hampala salweenensis , from the Mae Pai River System, Salween Basin, Thailand

Atsushi Doi and Yasuhiko Taki

Laboratory of Ichthyology,Tokyo Universityof Fisheries, 4-5-7 Konan, Minato-ku,Tokyo 108, Japan (ReceivedApril 14, 1993;in revisedform September16, 1993; acceptedSeptember 18 , 1993)

Abstract A new cyprinid fish, Hampala salweenensis , is described from three specimens collected from the Mae Pai River system, Mae Hong Son , northwestern Thailand. The species is distinguishable from its congeners by the combination of the following characters: total lateral line scales 26-27; two roundish black blotches on body side; upper and lower edges of caudal fin with a black marginal band .

The cyprinid genus Hampala Bleeker, 1859-1860 were defined as those without haemal spines; the has been known to be represented by five species all second and third vertebrae were counted separately . occurring in Southeast Asia, i.e., Hampala macro- The specimens used for the present study belong to lepidota (Valenciennes, 1842 in Cuvier and Valen- the following institutions: California Academy of ciennes, 1842; including H. m. sabana Inger et Chin , Sciences (CAS); Institute for Breeding Research , 1962 as a subspecies), H. ampalong (Bleeker, 1852), Tokyo University of Agriculture (IBRP); Institute H. bimaculata (Popta, 1905), H. lopezi Herre, 1924 of Taxonomic Zoology, University of Amsterdam and H. dispar Smith, 1934 (Weber and de Beaufort , (ZMA); Museum of Fisheries, Kasetsart University, 1916; Herre, 1924; Smith, 1945). They are close to Bangkok (KUMF); National Museum of Natural each other in external morphological characters but History, Leiden (RMNH); National Science distinguishable primarily by their color patterns . Museum, Tokyo (NSMT); The Natural History During our field surveys in Thailand in 1989 and Museum, London (NHM). 1991, we obtained three specimens of Hampala which resembled insular Southeast Asian H. ampa- long and H. bimaculata in color patterns rather than Hampala salweenensis sp. nov. continental Southeast Asian H. macrolepidota and (Figs. 1, 2) H. dispar. Subsequent comparisons with the five known species have warranted the description of a Holotype. NSMT-P 35838, 200.6 mm standard length new species. (SL), Mae Surin River (a branch of the Mae Pai River), at Ban Huei Phan, Mae Hong Son, northwestern Thailand, ec. 1991. Methods Paratypes. NSMT-P35839, 127.6mmSL, locality and data as for the holotype; KUMF 2959, 197 Methods for counts and measurements largely fol- .6mm SL, Mae Pai River, near Mae Hong Son Fisheries Station, Mae lowed Hubbs and Lagler (1947). For counting the ong Son, northwestern Thailand, 6 Aug. 1989. scale counts above and below the lateral line, the scale immediately in front of the insertion of the Diagnosis. A species of Hampala distinguishable dorsal, anal and pelvic fins was counted as one-half. from its congeners by the combination of the follow- Length of the upper and lower caudal fin lobes was ing characters: lateral line scales 26-27; two roundish measured as a straight distance from the base of the black blotches on both body sides, one on flank middle caudal ray to the tip of the upper and lower beneath dorsal fin origin and above lateral line, the lobes. Total lateral line scale count was represented other on caudal peduncle crossing lateral line; upper by the total number of pored scales. Radiographs and lower edges of caudal fin with a black marginal were used to count vertebrae. Abdominal vertebrae band.

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Fig. 1. Hampala salweenensis sp. nov., NSMT-P 35838, holotype, 200.6mm SL, from the Mae Surin River (a branch of the Mae Pai River), at Ban Huei Phan, Mae Hong Son, northwestern Thailand.

Fig. 2. Above-Hampala salweenensis sp. nov., 200.6mm SL (NSMT-P 35838); middle-H. ampalong, 111.5 mm SL (NHM 1866.5.2:112); below-H. bimaculata, 152.3mm SL (NHM 1983.6.20:4).

Description. In the following description, mean occiput 20.4% (20.4-20.5%) SL. Snout pointed, its values for the three specimen are given first, followed length 36.7% (35.6-37.3%) of head length (HL) in parenthesis by ranges. and slightly dented around nostril region. Eyes Body moderately compressed; width 17.2% (16.2- situated in dorsal half of head. Orbit diameter 17.8%) of SL; greatest depth at origin of dorsal fin 19.7% (17.9-22.9%) of HL. Postorbital length ray, 30.5% (29.7-31.6%) of SL. Head large; its 49.3% (48.0-50.7%) of HL. Interorbital width length 31.0% (30.8-31.2%) of SL and its depth at 34.0% (33.0-34.8%) of HL. Mouth large, deeply

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Fig. 3. Above-, 191.8 mm SL (NSMT-P 31975); middle-H. macrolepidota , 180.0 mm SL (NSMT-P 35841); below -H. lopezi, 100.6mm SL (CAS 138090).

Fig. 4. Geographic distribution of genus Hampala. •£, H. ampalong; •¡ H. bimaculata; •› , H. dispar; •¥, H. lopezi; •œ H. macrolepidota; •š, H. salweenensis. Data source from the examined specimens, Day (1876-

1878), Weber and Beaufort (1916), Herre (1924), Fowler (1934, 1935 , 1937, 1941), Pellegrin and Fang (1940), Smith (1945), Inger and Chin (1962), Taki (1974), Kottelat (1985), Mai and Nguyen (1988) and Roberts (1989).

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Table 1. Meristic

Table 2. Morohometric

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characters of six Hampala species

characters of six Hampala species

― 4 0 9 ― A. Doi & Y. Taki cleft, its corner extending to a vertical line from tion, encompassing the Indochinese Peninsula, anterior border of eye. A pair of maxillary barbels Malay Peninsula and Greater Sunda Islands (Fig. present; length of the barbel 81.2% (52.7-108.9%) 4a). H. dispar and H. salweenensis are continental in of orbital diameter. distribution, but their distributions are confined to Dorsal fin size moderate, with four simple rays and small ranges, the middle Mekong for the former and eight branched rays. Anal fin with three simple rays the Salween basin for the latter (Fig. 4b). The dis- and five branched rays. Pectoral fin with pectoral fin tribution of H. dispar falls within that of H. macro- rays counts 15.3 (15-16). Pelvic fin with 9 rays. lepidota, whereas H. salweenensis is not sympatric Proximal half of last simple dorsal fin ray osseous, with either of these two species, though its range with small denticles, its distal half not osseous, seg- adjoins that of H. macrolepidota. H. ampalong and mented; last simple dorsal fin ray length 21.1% H. bimaculata are distributed in the Greater Sunda (20.5-21.7%) of SL. Last simple anal fin ray non- Islands: the former in Sumatra and western Borneo, osseous, its length 15.6% (14.5-16.5%) of SL. and the latter in western, northern and eastern Origin of dorsal fin about midway between tip of Borneo (Fig. 4b). H. lopezi has been recorded solely snout and base of caudal fin and slightly in advance from Busuanga Island, Philippines, an island in the of insertion of pelvic fins. Predorsal length 53.0% island-chain connecting north Borneo and the Philip- (52.9-53.2%) of SL, prepelvic length 53.1% (53.0- pine Archipelago (Fig. 4b). 53.1% ) of SL and preanal length 76.2% (75.7- These six species are very similar in meristic and 76.8%) of SL. morphometric characters, no character alone clearly Lateral line complete; decurved downward, run- distinguishing them from each other unless com- ning on lower side of trunk and on middle of caudal bined with color patterns (Figs. 2 and 3; Tables 1 peduncle. Scales large. Lateral line scales 24.3 (24- and 2). 25), two scales on caudal fin base. Predorsal scales When the adult coloration of H. salweenensis is 10.3 (10-11). Scales above lateral line to dorsal compared with continental Southeast Asian species, origin 4.2 (3.5-4.5); scales below lateral line to anal i.e., H. dispar and H. macrolepidota, H. salweenensis origin 3.5; scales below lateral line to pelvic insertion approaches H. macrolepidota in having black 2.5. Circumpeduncular scales 12. marginal stripes along the upper and lower edges of Gill rakers small, canine-like, coarsely set, number the caudal fin. However, H. salweenensis differs from 1.3 (1-2) on upper limb, 10 on lower limb. the two species in having two blotches on the body Color in alcohol. -Back dark, grayish brown; belly side, one under the dorsal fin and the other on the brownish white. Scales with black margin except for caudal peduncle. During the juvenile stage, H. abdominal surface. Two large black blotches on dispar and H. macrolepidota have an ill-defined cross both body sides; one on flank (above 9th–lOth lateral bands on the caudal peduncle and a thin cross band line scale in NSMT-P 35838; above 10th-1 lth in along the caudal fin base, but these markings disap- NSMT-P 35839 and KUMF 2959), the other on pear with growth (Fowler, 1937; Taki and Kawa- caudal peduncle (on 20th-21st scale in NSMT-P moto, 1977). The smallest specimen of H. macro- 35838; on 21st-22nd in NSMT-P 35839 and KUMF lepidota and of H. dispar examined in the present 2959). Pectoral, pelvic and anal fins pail whitish study, 71.8mm SL and 100.4mm SL respectively, brown. Frontal edge of dorsal fin black, dorsal fin had no markings on the posterior part of the body. rays pail black. Upper and lower edges of caudal fin H. dispar has no distinct marginal bands on the with a black band, caudal fin rays pale black. caudal fin throughout its life history. In addition to the differences in coloration, H. Etymology. The name salweenensis refers to the salweenensis differs from H. macrolepidota in lower distribution of this species, the only member of the limb gill-raker count (10 in the former and 8-9 in the genus known from the Salween basin. latter), and from H. dispar in the length of the last simple anal ray (14.5-16.5% of SL vs. 11.2-14.7% of SL), snout length (35.6-37.3% of HL vs. 29.9- Comparison 31.7% of HL) and postorbital length (48.0-50.7% of HL vs. 52.4-56.7% of HL). Of the six species of Hampala (Figs. 2 and 3), H. Hampala salweenensis is similar to the two species macrolepidota has the largest geographic distribu- from Sunda Islands, i.e., H. ampalong and H. bimac-

― 4 1 0 ― A New Cyprinid from Thailand ulata, in having two blotches on the body side. 112, 1, 111.5,East Indian Archipelago;ZMA 120.777,6, However, H. salweenensis has fewer total lateral line 79.6-125.9, Batang Hari, Djambi, Sumatra, 23 Apr. scales than H. ampalong (26-27 in the former and 1909. RMNH 7613, 3, 99.8-110.9, 28-31 in the latter). Further, the following mor- Howong, Borneo, Aug. 1898; RMNH 7614, 4, 72.4- phometric characters show differences between H. 113.9, Bo, Borneo, May-Aug. 1900; RMNH 7615, 3, salweenensis and H. ampalong: length of last simple 73.5-101.4, Kajan, Borneo, Sept.-Oct. 1900; NHM dorsal ray (20.5-21.7% of SL vs. 22.4-24.8% of 1978.3.20:66,1, 247.0, S. Tutoh, S. Uoong, Sarawak, SL), length of lower caudal fin lobe (28.7-31.2% of Borneo, 20 Mar. 1978; NHM 1978.9.5:14, 1, 122.4, Camp 5. Meliau, Sarawak,Borneo, 5 Sept. 1978;NHM SL vs. 31.7-34.1% of SL) and snout length (35.6- 1978.9.5:15,1, 91.0, S. Lansar, Sarawak,Borneo, 5 Sept. 37.3% of HL vs. 31.8-34.1% of HL). In the de- 1978;NHM 1982.4.21:1,1, 230.0, S. Lansar, Sarawak, scription of IL ampalong, Weber and de Beaufort Borneo, 21 Apr. 1982; NHM 1982.4.21:2, 1, 115.6, Meliau R., Meliau Gorge, Sarawak, Borneo, 21 Apr. (1916) stated that "upper and lower margin of 1982;NHM 1983.6.20:4,1, 152.3,Batang R caudal with a faint narrow black streak," but such a ., Sarawak, Borneo, 20 June 1983; NHM 1983.6.20:13-14, 1, 79.8, streak was not shown in Bleeker (1852, original Batang R., Sarawak, Borneo, 20 June 1983; NHM description of the species, and 1863-1864), nor was 1983.6.20:97-98,1, 87.0, BatangR, Sarawak,Borneo, 20 it observed in the specimens examined in the present June 1983. study. Hampala dispar NSMT-P 31975, 1, 191.8, Tha Bo, Thai- land, 29 Mar. 1989; NSMT-P 35850, 5, 107.9-123.8, The color patterns of H. salweenensis and H. bi- Market at Sung Noen, Khorat, Thailand, 4 Dec. 1991; maculata, although agreeing in possessing two NSMT-P 35851, 1, 100.4,Haui Mong R. at Ban Dong blotches on the body side, differ in the shape and Bung, about 7 km NW from Tha Bo, Nongkai, Thailand, position of the blotches. The blotches in H. bimacu- 11 July 1990. lata are saddle-shaped, which are forming a circular Hampala lopezi CAS 138090, 5, 58.9-189.7, Barrio San Nicolas,Wayan R., Busuanga,Philippines, 21 June 1940. blotch on each side in large specimens. The anterior Hampala macrolepidotaNSMT-P 35852, 1, 140.0,Market blotch is situated beneath the dorsal fin origin (above at Phun Phin, Surat Thani, Thailand, 1 Aug. 1990; 10th-11th or less frequently 9th-10th lateral line NSMT-P 35855, 1, 179.4, Boeng Boraphet Channel, in scales) in H. salweenensis, whereas it is positioned front of Nakhon Sawan Fisheries Station, Nakhon below the posterior half of the dorsal fin (on 11th-13 Sawan,Thailand, 16 Jan. 1992;NSMT-P 35853, 1, 244.6, Market at Pak Nam Pho, Nakhon Sawan, Thailand, th or 12th-14th lateral line scales) in H. bimaculata. 15 Jan. 1992; NHM 1978.3.20:65,1, 140.9,S. Tutoh, The two species differ in many proportional charac- Sarawak, Borneo, 20 Mar. 1978; NSMT-P 35854, 1, ters, i.e. head depth (more than 20.4% of SL in H. 283.3, Boeng Boraphet Channel, in front of Nakhon salweenensis vs. less than 19.3% in H. bimaculata), Sawan Fisheries Station, Nakhon Sawan, Thailand, 16 Jan. 1992; NSMT-P 35841, 1, 180.0, Boeng Boraphet predorsal length (less than 53.2% of SL vs. more Channel, in front of Nakhon Sawan Fisheries Station, than 53.5%), length of the dorsal and anal fin base Nakhon Sawan,Thailand, 16 Jan. 1992;IBRP 4643, 10, (more than 16.2% and 9.8% of SL vs. less than 71.8-102.8,Nam Kem Stream near its conjunctionto the 15.9% and 9.7%) and interorbital width (more than Nam Khon, at Tha Ngon, Laos, 12 Sept. 1970. 33.0% of HL vs. less than 31.7%). Hampala lopezi is the only Hampala species with a longitudinal band on the body side and is hence Acknowledgments readily distinguishable from other species. Further, the following differences were observed between H. We thank Dr. Hiroshi Kohno, Tokyo University salweenensis and H. lopezi: head length (less than of Fisheries, for his comments during the investiga- 31.2% of SL in H. salweenensis vs. more than 31.7% tion. Grateful thanks are also due to Dr. Kitjar in H. lopezi), predorsal length (less than 53.2% of SL Jayen, Dr. Sompong Hiranyawat, Mrs. Jiamjit vs. more than 54.7%) and interorbital width (more Boonsom of the National Inland Fisheries Institute, than 33.0% of HL vs. less than 31.4%). Thailand, Mr. Somwang Pimonbutr, Mr. Paibool Roongpiboolsopit and staff members of the Mae Hong Son Fisheries Station, Thailand, for their co- Comparative Materials operation in field collection. We are indebted to the following persons and institutions for the loan of Hampala ampalongRMNH 4968, 1 specimen,89.2mm SL, specimens: Messrs G.J. Howes and P. Campbell Palembang, Sumatra, date unknown; NHM 1866.5.2: (NHM); Dr. M. J.P. van Oijen (RMNH); Dr. T.

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Iwamoto and Mr. D. Catania (CAS); Dr. I.J.H. provisory annotated check-list. Hydrobiologia, 121: Isbrucker (ZMA); Drs. K. Matsuura and R. Arai 249-279. Mai, D.Y. and V.T. Nguyen. 1988. Species composition (NSMT). and distribution of the freshwater fish fauna of southern Vietnam. Hydrobiologia, 160: 45-51. Pellegrin, J. and P.W. Fang. 1940. Poissons du Laos Literature Cited recueillies par MM. Delacour, Greenway, ed. Blanc. Description d'un genre, de cinq especes et d'une variete. Bleeker,P. 1852. Diagnostischebeschrijvingen van nieuwe Bull. Soc. Zool. France, 65: 111-123. of weinigbekende vischsoortenvan Sumatra. Tiental I- Popta, C.M.L. 1905. Suite des descriptions preliminaires IV. Nat. Tijds. Ned. Ind., 3: 569-608. des nouvelles especes de poissons recueillies au Borneo Bleeker,P. 1859-60. Conspectussystematis Cyprinorum. Central par M. le Dr. A.W. Nieuwenhuis en 1898 et en Nat. Tijds. Ned. Ind., 20: 421-441. 1900. Notes Leyden Mus., 25: 171-186. Bleeker,P. 1863-64. Atlas ichthyologiquedes Indes Orien- Roberts, T.R. 1989. The freshwater fishes of western tales Neerlandaises.3. Cyprins. 150pp.,pls. 102-144. Borneo (Kalimantan Barat, Indonesia). Mem. Calif. Cuvier, G. and A. Valenciennes.1842. Histoire naturelle Acad. Sci., 14: xii+210pp. des poissons,Vol.16. Paris-Strasbourg.xx+472pp., pls. Smith, H.M. 1934. Contributions to the chthyology of 456-487. Siam. J. Siam Soc., Nat. Hist. Suppl., 9: 287-325. Day, F. 1876-1878. The fishes of India; being a natural Smith, H.M. 1945. The fresh-waterfishes of Siam, or history of the fishesknown to inhabit the seas and fresh Thailand. Bull. U.S. Nat. Mus., 188: xi+622pp. waters of India, Burma, and Ceylon. London. xx+778 Taki, Y. 1974. Fishes of the Lao Mekong basin. USAID pp. Mission to Laos, Agric. Div., vi+232pp. Fowler, H.W. 1934. Zoological results of the third de Taki, Y. and A. Kawamoto. 1977. Differentiation of the Schauensee Siamese expedition.Part 1. Fishes. Proc. cyprinids, and H. dispar. Japan. Acad. Nat. Sci. Phila., 86: 67-163. J. Icthyol., 24: 61-65. Fowler, H.W. 1935. Zoological results of the third de Weber, M. and L.F. de Beaufort. 1916. The fishes of the SchauenseeSiamese expedition.Part 6. Fishesobtained Indo-Australian Archipelago. 3. Ostariophysi: II. Cypri- in 1934. Proc. Acad. Nat. Sci. Phila., 87: 89-163. nidae, etc. E.J. Brill, Leiden. xv+455pp. Fowler, H.W. 1937. Zoological results of the third de SchauenseeSiamese expedition.Part 8. Fishesobtained in 1936. Proc. Acad. Nat. Sci. Phila., 89: 125-264. タ イ 北 西 部 サ ル ウ ィ ン河 水 系 パ ー イ 川 か ら採 集 さ れ た コ Fowler, H.W. 1941. Contributions to the biologyof the イ 科 の1新 種Hampala salweenensis PhilippineArchipelago and adjacent regions. Bull. U.S. Nat. Mus., 100(13): x+879. 土 井 敦 ・多 紀 保 彦 Herre, A.W.C.T. 1924. Distribution of the true fresh- water fishes in the Philippines.1. The PhilippineCyprin- タイ 北 西 部 サ ル ウ ィ ン河 水 系 パ ー イ 川 か ら採 集 され た3個 体 idae. Philipp. J. Sci., 24: 249-306,pls. 2. の 標本 に も とづ いて,コ イ科 の新 種Hampala salweenensisを 記 載 Hubbs, C.L. and K.F. Lagler. 1947. Fishes of the Great した.本 種 はHampala属 の 他 種 か ら全側 線 鱗 数26-27枚,体 側 Lakes region. Univ. Mich. Press, Ann Arbor, Michigan. の2つ の 黒 斑 の 存 在,尾 鰭 上 下 葉 端 の 黒 色 バ ン ドの 存 在 の組 合 213pp. せ に よ り識 別 され る. Inger, R.F. and P.K. Chin. 1962. The fresh-waterfishes of North Borneo. Fieldiana Zool., 45: 1-268. (〒108東 京 都 港 区 港 南4-5-7東 京 水 産 大 学 魚 類 学 研 Kottelat, M. 1985. Fresh-water fishes of Kampuchea. A 究 室)

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