A GENERICAL CLASSIFICATION OF NEMACHEILINAE WITH DESCRIPTION OF TWO NEW GENERA (TELEOSTEI: CYPRINIFORMES: COBJTIDAE)~

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1 Trav. Mus. Hist, nat, ctgrigore Antipa) Vol. XXXV pp A GENERICAL CLASSIFICATION OF NEMACHEILINAE WITH DESCRIPTION OF TWO NEW GENERA (TELEOSTEI: CYPRINIFORMES: COBJTIDAE)~ On discute la position des Nemacheilinae. l'historique du probleme des genres inclus, les caracteres servant a distinguuer les genres et groupes d'especes et on passe en revue les 23 genres consideres valides, dont deux nouveaux: Longischistlrra et Serninemachilus: les auteurs n'ont pas pu examiner trois genres dccrits rkcernrnent par des auteurs chinois: Paranemacheilus. Henrinoemacheilus (qu'ils considerent synonyrne du genre precident) et Sphaerophysa; la position de Eloposfoma et la validit6 de Barbucca semblent douteuses. On distingue plusieurs groupes d'especes dans le cadre du genre Schistura. Les auteurs discutent les relations possibles entre les genres. Nemacheilinae is the largest of the three subdivisions (usually ranked as subfamilies) of Cobitidae (or Cobitidinae: Kottelelat, 1986). Sawada (1982) excluded the nemacheilines from Cobitidae, ascribing them to Homalopteridae; he considers Cobitidae and Homalopteridae as sister taxa, placing them in the new superfamily Cobitoidea. His views have been adopted by Menon (1987) and by Kottelat (1989, 1990) who changed the family name Homalopteridae in Balitoridae. Roberts (1989) retains on the contrary the nemacheilines in Cobitidae, while Wu et a1 (1981) believe that Cobitidae and Homalopteridae are not closely related, the former (also including Nemacheilinae) being considered closest to Catostomidae and Gyrinocheilidae, the latter to Cyprinidae. Since the raising of the superfamily Cobitoidea would imply the raising to superfamily rank also of the four other families of cypriniform fishes, a compromise solution has been adopted by Banarescu (1990) by downgrading Homaleptoridae (= Balitoridae) and Gastromyzonidae as subfamilies of Cobitidae, ' The present contribution is dedicated to the memory of Hialmar Rendahl from the Stockholm Museum of Natural History, in appreciation of his fundamental contributions to the knowledge of Cobitidae.

2 430 PETRU M. BANARESCU. I'EODOR T. NALBANT alongside Nemacheline, Vaillantellinae, Cobitinae and Botiinae. The purpose of the present paper being the delimitation of genera within the nemacheiline lineage, no further references are made concerning the relationships of the various lineages of cobitid and homalopterid loaches. HISTORICAL REVEW OF THE PROBLEM Starting with 1839, numerous generical names have been proposed for nemacheiline loaches, besides Nemacheilus Van Hasselt, 1823; most of these have been considered by subsequent authors as synonyms or subgenera of the latter. Prior to 1964, the only genera accepted as valid, besides Nemacheilus (usually spelt Nemachilus or Noemacheilus) were: Vaillantella Fowler, 1904 from Indonesia and the Malay Peninsula; Aborichthys Chaudhuri, from northern India; Lefua Herzenstein, 1888 (= Elxis Jordan and Fowler, 1903) from the north of East Asia and from Kalimantan or Borneo Island; Oreonectes Giinther, 1868 from southern China. Weber and de Beaufort (1916); Rendahl (1933; 1948); Berg (1949); Nichols ( 1943) accepted as valid genus also "Barbatula" (actually with doubtful nomenclatorial status - see later), ascribing to it several Chinese species. In a series of preceding papers (Banarescu and Nalbant, 1964, 1966, 1968, 1974, 1975, 1976; Bilnarescu et al., 1978, 1982; Mirza et al., 1981) several generical names proposed by older authors have been resurrected and shown actually to be valid: Schistura, Acanthocobitis, Yunnanilus, Paracobitis, Orthrias, Oreias, Triplophysa and Hedinichthys. However, in the three former papers the species were still listed, provisionally, under Noemacheilus and some erroneous opinions have been advanced, for example that Nemachilichthys is a synonym of Noemacheilus s. str., Micronemachilus a synonym of Pogononemacheilus and that Hedinichthys includes all High Asian species with reduced and encapsulated air bladder. Four new genera (Turcinoernacheilus, Mesonoemacheilus, Nun and Physoschistura) and two new subgenera (Oxynoemacheilus and Indoreonectes) have also been described. Several more recent authors have accepted the new genera resurrected or described by us, either as valid subgenera (Jayaram, 198 1; Menon, 1987) or as full genera (Zhu and Guo, 1985; Zhu and Cao, 1987; Kottelat, 1989, 1990) while others (Zhu, 1982 a, 1982 b; Yang and Xie, 1983; Krupp, 1987; Krupp and Schneider, 1989) still use the generic name Nemacheilus (or Noemacheilus) in the old wide sense.

3 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 43 1 Other genera described in recent years are: Paranoemacheilus Zhu and Cao 1987 from the Xijiang (or Zhujiang, formerly Hsikiang or Pearl river) basin in southern China; Neonoemacheilus Zhu and Guo, 1985 from the Brahmaputra, Salween and Irrawaddy basins in northern India and Burma/Yunnan; Sphaerophysa Cao and Zhu, 1988 from the Xijiang River watershed of Yunnan; Barbucca Roberts, 1989 from Kapuas River, western slope of Borneo or Kalimantan Island; Sectoria Kottelat, 1990 fi-om the Chao Phraya River basin in Thailand; Tuberoschistura Kottelat, 1990 from the Mekong, Chao Phraya basins and the eastern slope of the Malay Peninsula and subgenera Qinghaichthys Zhu, 1981 (of Trilophysa) from the High Asian reach of the Changjiang or Yangtse River basin, Petruichthys Menon, 1987 from the Salween River basin of Burma and Troglocobitis Parin, 1983 from Turkmenistan. CHARACTERS USED IN DISTINGUISHING GENERA Many recent authors consider that the only phyletically significant characters in distinguishing higher taxa of vertebrates are osteological ones; others consider mainly karyological and/or biochemical characters. Actually all characters, including external morphology, rays, scales etc. have phyletical significance. Many fish genera distinguished initially only on the base of "classical" characters proved to be monophyletic. Since our studies were based exclusively on preserved specimens, most borrowed from various museums and which could not be dissected, only "classical" characters were used, namely: General shape of body and of head; position of mouth; relative position of fins; number of dorsal and anal rays; shape of dorsal and caudal fins; presencelabsence and disposition of scales; presencelabsence and relative length of lateral line; absencelpresence and degree of development of an adipose creast on the caudal peduncle; colour pattern (complete or incomplete transversal stripes; longitudinal stripe; irregular mannoration; presencelabsence and position of a blackish spot on the base of the caudal fin); shape of lips (almost smooth; slightly or deeply furrowed or covered by pappilae); length and shape of the intestine; presencelabsence and degree of development of a tooth-like prolongation ("processus dentiformis") on the upper jaw (Fig. 1 A, B); shape of the bony capsule of the anterior chamber of the air-bladder; shape and degree of development of the posterior, free chamber of the air bladder. A special mention deserves the sexual dimorphism. There is no sexual dimorphism (except body depth in females full with eggs and probably length of paired fins) in some genera (Paracobitis, Oreias) and species groups of Schistura. In ~VemachiIichthys the sexes differ in colour pattern. Two other types of sexual dimorphism are present in various genera:

4 432 PETRU M. BANARESCU, TEODOR T. NALBANT ( 1) The Noemacheilus-Schistura type: an osseous movable suborbital flap is present in males (Fig. 2 A) in which the pectoral rays are thickened, ossified and covered by breeding tubercles: breeding tubercles are often present on the body sides as well (Fig. 2 D). (2) The Orthrim-Triplophysa type: there is no suborbital flap, but the head sides of males, below the eyes, are covered by numerous breeding tubercles, either all over the head sides (Orthrias: Fig. 2 C) or grouped in two fields separated by a groove (Triplophysa: Fig. 2 B); the pectoral rays are broadened, thickened and covered by breeding tubercles, too; in some species these tubercles are present on the body sides as well. These characters are also useful in delimiting intra-generical groups of species. POSITION OF VAILLANTELLA This genus, traditionally placed in Nemacheilinae, has been ascribed by Nalbant and Banarescu (1977) to a subfamily of its own, characterised by an unusually long dorsal fin, the two pairs of rostral barbels confluent at their bases, gill opening oblique in its lower part (as in Botiinae; in Nemacheilinae it is vertical) and the capsule of the anterior chamber of the air-bladder unique, fibrous, with two distinct osseous plates. Sawada (1982) and later Kottelat (1990 do not accept our view-point, arguing that the fibrous nature of the air-bladder capsule is a plesiomorphic character. Actually, the capsule is not only fibrous, but also unique, as in Botiinae, while in true Nemacheilinae (and in Homalopterinae, too) it consists in a pair of lateral dilatations. This facts justifies Sawada's (op. cit.) assumption that Nemacheilinae and Homalopterinae are close relatives, as well as the subfamily status of Vaillantellinae. The latter shares characters both with Nemacheilinae (absence of a preorbital spine) and with Botiinae (air-bladder capsule, rostral barbels confluent, gill opening oblique), justifying the grouping of Nemacheilinae, Homalopterinae, Botiinae, Cobitinae and Vaillantellinae in a single family, as suggested by Banarescu (1990). The sister relationships of Vaillantellinae (whether closer to Nemacheilinae or to Botiinae) is not important in this paper which deals exclusively with Nemacheillinae s. str. SYSTEM OF NEMACHEILINAE 1. Acanthocobitis Peters, (Fig. 3) Type species: Acanthocobitis longipinnis Peters (= Cobitis pavonaceus Mc Clelland) by monotypy; feminine.

5 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 433 Body deeper than in most other nemacheilines, strongly compresed posteriorly; head slightly compressed; snout blunt; a slight indication of adipose keel; no barbel-like prolongation of the nasal comet; 10-18, rarely eight or nine branched dorsal rays; edge of dorsal fin straight or slightly covex; caudal fin slightly emarginate, straight or convex. Scales large, usually imbricated, distributed all over the body; their shape remembers that of the scales of Botiinae, having a reduced and central focal zone (Fig. 3 E). Lateral line complete or extending at least under the dorsal fin; its orifices perforate the scales. Upper lip covered by two-three rows of papillae; lower lip broadened on both sides of the median interruption and covered by numerous papillae (Fig. 3 B). Processus dentiformis of the upper jaw (premaxilla) strong. Posterior chamber of the air-bladder rudimentary. Intestine short, with a single loop (Fig. 3 C). Sexual dimorphism of Nemacheilus- Schistura type: either a suborbital flap (not as strong as in the species of Nernacheilus) or only a "suborbital groove" (Kottelat 1990), actually an indication of suborbital flap. Colour pattern: an irregular net of rnarrnorations and of more or less metamerically disposed spots which may represent the fragmentation of transversal stripes; a well marked black spot on the upper part of the caudal fin base (Fig. 3 A), not on the middle of the base, as in Nemacheilm, Micronemacheilus and Nemachilichthys. Range: the Indian subcontinent, Ceylon Island, the Burmanese province (Irradwaddy, Sittang and Salween rivers) and the western slope of the Malay Peninsula; two spedies also in the Mae Khlong River system, eastern slope of Peninsula Thailand (Kottelat, 1989, 1990) - but perhaps introduced there. Four species; the type of the genus, A. pavonaceus differs from the others in having the anus closer to the pelvics than to the anal fin insertion and the higher number of dorsal rays (14-18 branched ones); A. botia and A. rubidipinnis have a complete lateral line and differ fiom each other mainly in the number of branched dorsal rays (10-13 in the former, in the latter), while A. zonalternans is the only species of the genus with incomplete lateral line; it has branched dorsal rays. A. phuketenis is considered by Kottelat (1990) to be a synonym of A. zonalternans; this seems probable, but the length of the lateral line must be verified. 2. Nemacheilus Bleeker, 1863 (Fig. 4) Type species: Cobitis fasciata Valenciennes, by original designation; masculine.

6 434 PETRU M. BANKRESCU, TEODOR T. NALBANT Synonyms: Noemacheilus Van Hasselt 1823; nomem nudurn. Modigliania Perugia, 1893; type species: Modigiania papillosa Perugia = Cobitis fasciata Valencienne, by monotypy; feminine. Pogononemacheilus Fowler, 1937; type species: Nemacheilus masyae Smith, by original designation; masculine. Tuberoschistura Kottelat, 1990; type species: Nemacheilus baenzingeri Kottelat, by original designation; feminine; new synonymy. Body elongate, almost cylindrical; no adipose creast on caudal peduncle; no barbel-like prolongation of nostrils. Dorsal fin with 7-10 branched rays. Pectoral rays produced, extending beyond the margin of the fin membrane (Fig. 4 A). Insertion of the pelvic fins, as in most genera of nemacheilines (except Turcinomacheilus and the not described genus from Lake Tana, Ethiopia) slightly behind the vertical of the dorsal insertion. Scales present, well developed, in most species with reduced focal zone (Fig. 4 E), imbricate (at least posteriorly); lateral line complete, its orifices perforating the scales. Barbels usually long. Lips narrow, in most species deeply furrowed (Fig. 4 B), in fewer only slightly furrowed; lower lip usually with a slight median incision or interruption (except in N. olivaceus). Processus dentiformis of the upper jaw moderately developed; no corresponding median notch on the lower jaw. Sexual dimorphism: a well developed suborbital flap is present in males and the 2" to 6Ih rays of the pectoral fins are covered with breeding tubercles; these rays are only slightly thickened and not broadened, as in members of other genera. Breeding tubercles are often present on the body sides, too, especially in its posterior half. Posterior, fiee chamber of the air-bladder rudimentary; the two dilatations of the air-bladder capsule are, in most species, more or less isodiametric; the manubrium connecting them is in most species narrow (Fig. 4 D), while in N. selangoricus it is short and broad. The intestine is short, having in most species a single loop. Colour pattern variable; most species have a variable number of more or less complete crossbars, either narrow (Fig. 4 A) or broad; in others, the crossbars are interrupted, consisting it an upper and a lower spot or stripe; N. binotatus and the three south-westem Indian species have on the contrary a single longitudinal stripe, extending from the opercle or from the tip of the snout to the caudal fin base. Various combinations exist between these two extremes: species in which the crossbars are divided each in a dorsal and a lateral spot, the lateral one being more or less confluent into a longitudinal stripe (N. pallidus: illustrations in Kottelat, 1990, his Fig. 36) or in which the crossbars, reduced to their median

7 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 435 fragment, coexist with a well marked longitudinal stripe (N. ornutus: Kottelat, op. cit.., his Fig. 34). In other species there are only lateral spots, usually arranged in a longitudinal series, a dorsal and a lateral one: N. baezingeri (Kottelat, 1983; 1990, his Fig. 175). Finally, the troglobiontic N. cataractus (whose generical position seems us uncertain) is colourless (Kottelat and Gby, 1989). An important colour character, common to all or most species, is the occurrence of a dark spot on the middle of the caudal base. General remarks on the genus and on its species: The genus, as accepted by us, includes numerous species, some of which are not quite similar. Tuberoschistura Kottelat, 1990 (type species: Nemacheilus baezingeri Kottelat, 1983) is, in our opinion, a junior synonym of Nemacheilus. The type species has, like most other Nemacheilus, prolonged pectoral rays, large eyes, forked caudal fin, imbricate scales, a similar shape of the lips and intestine; also, its colour pattern is similar to that of other Nemacheili, U.O. N. corica and N. oxianus. It differs from the other members of the genus in having a slightly higher number of dorsal rays, a more anterior anus and the occurrence of elongate tubercles around the eye. We ascribe to this genus U.O. the North Indian N. corica and the Central Asian (Aral Sea basin) N. oxianus. Possibly, two species have been confounded under the latter name and it will be necessary to examine typical specimens. The species that actually is a Nemacheilus has prolonged pectoral rays, long barbels etc. and the same type of sexual dimorphism as the Indonesianhdochinese members of the genus. The colour pattern remembers that of N. corica and partially of N. baezingeri. We also included in Nemacheilus, contrary to Kottelat (1990), three littleknown species from south-western India (N. monilis, N. poonensis and N. anguilla), characterised by a rather short dorsal fin (7 or 8 branched rays, the latter number being rare in Indochines and Indonesian species), a longitudinal stripe, large eyes, etc.; there is no information about the sexual dimorphism and nature of the pectoral fin rays. Six species are listed by Kottelat (1984) from the western Indonesian Islands (besides N. obesus, now ascribed to another genus), four others (and two of "Tuber~schistura'~ from the Mekong and the Chao Phraya rivers basins (Kottelat, 1990), and another one, N. ornatus from the Malay Peninsula, one (M. masyae) from the same Peninsula and south-eastern Thailand and a third one, N. selagoricus, from the Malay Peninsula and two Indonesian islands (Kottelat, 1984; 1990). Two other species have been described by Roberts (1989) from the western slope of Borneo or Kalimantan Island in Indonesia: lactogeneus and

8 436 PETRU M. BANARESCU, TEODOR T. NALBANT maculiceps. The former apparently is a true Nemacheilus, but the latter is Schistura (first member of this genus found in Indonesia). This makes a total amount of 16 species in south-eastem Asia (another species, the hypogean N. cataractus Kottelat and GCry, 1989 may belong to another genus); these are accepted as valid species of the genus, alongside four Indian ones (three in the south-west, corica in the north) and the Middle Asian N. oxianus. 3. Neonoemacheilus Zhu and Guo, 1985 (Fig. 5) Type species: Noemacheilus Iabeosus Kottelat, 1982, by original designation; masculine. Synonym: Infundibulatus Menon, 1987; type species: Nemachilus peguensis Hora, 1929, by original designation; masculine. Similar in body shape with Nemacheilus. Dorsal fin with 8 or 9 branched rays; its distal edge slightly notched. Caudal fin forked. No adipose keel or creast on the caudal peduncle. Body almost completely scaled; lateral line complete; scales roundish, with a large central focal zone (Fig. 5 E). Lateral line complete. Lips very thick, hypertrophied, forming a preoral cavity which was described by Kottelat (1982; 1990) and by Zhu and Guo (1985). The air-bladder has not been described by earlier authors; in the single available specimen of N. labeosus (Fig. 5 D) the posterior chamber seems to be well developed and free (contrary to all species of Nemacheilus); the two dilatations of the air-bladder capsule are ovoid, the manubrium connecting them is comparatively long and narrow, resembling that of Nemacheilus binotatus (compare with Fig. 4 D). Lntestine short (Fig. 5 C). Sexual dimorphism: according to Kottelat (1982; 1990), the males have a suborbital flap and the rays of the pectorals are slightly thickened; breeding tubercles have not been recorded, but they are probably present. Colour pattern: N. peguensis seems, according to its original description (Hora, 1929) to be colourless; this feature may however be an artefact, due to the poor state of preservation of the only available specimen. N. labeosus has vertical crossbars, either complete and continuos (the holotype: Kottelat, 1982, his Fig. 1; 1990 his Fig. 43) or subdivided each in a dorsal and a median spot, the last six median spots being confluent into a longitudinal stripe (Fig. 5 A). n! mengdingensis has about the same number of crossbars, the anterior ones reaching only to or slightly below the lateral line, the posterior ones almost to the ventral side (Zhu, 1989). A more intensive bar is present on the caudal fin base.

9 NEMACHEILINAE (TELEOSTEI) GENERlCAL CLASSIFICATION. 2 NEW GENERA 437 Range: the Burmanese province as delimited by Banarescu (1992) and partially suggested by Kottelat (1989): N. peguensis ranges in the Irrawaddy and Sittang basins, N. labeosus in the Salween River basin of Burma, N. mengdingensis in the Salween basin of Yunnan, China. 4. Micronemacheilus Rendahl, 1944 (Fig. 6) Type species: Nemacheilus cruciatus Rendahl, by original designation; masculine. Body deeper than in most genera of nemacheilines and compressed: caudal peduncle short, deep and compressed, with a slight indication of adipose creast. Nasal cornets short. Dorsal fin in the middle of the body, with 8-12 branched rays; its distal edge slightly concave or straight. Caudal fin slightly to moderately emarginate. Anal with five branched rays; anus a short distance in front of anal fin insertion. Pectoral fin rays not produced. Body completely covered by scales; these are large, longer than deep, with reduced and excentrical focal zone. Lateral line complete. Mouth inferior. Lips fleshy; upper lip continuos, with three rows of papillae. Lower lip interrupted in the middle; in the median part either two pairs of comparatively large papillae (M. cruciatus: Rendahl, 1944), or two papillose fields, similar to those in Acanthocobitis (M. pulcher: Fig. 6 B). Processus dentiformis of the upper jaw moderately to well developed. Intestine short, usually with a single loop (Fig. 6 C). Posterior chamber of the air-bladder very large and free (Fig. 6 D), connected to the anterior encapsulated chamber by a very short ducts; in M. cruciatus this chamber is subdivided by a strangulation in two portions (Rendahl, 1944), while in M. pulcher it is unique. No sexual dimorphism. Colour pattern: there are both vertical crossbars - often restricted to the upper part of the body - and a longitudinal row of metameric spots, often confluent into a broad longitudinal stripe; a well marked blackish spot in the middle of the caudal fin base, as in the species of Nemacheilus and Nemachilichtys. Range: the south of East Asian subregion as delimited by Banarescu (1992); two species: pulcher in the basins of Xijiang (Hsikiang) and Song-Koi (Red) rivers and Hainan island, cruciatus in the coastal rivers of central Vietnam.

10 438 PETRU M. BhJ&ESCU, TEODOR T. NALBANT 5. Schistura Mc Clelland, 1838 (Figs 7-1 1) Type species: Schistura rupecula Mc Clelland, by subsequent designation of Jordan, 19 19; feminine. Synonym: Acoura Swainson, 1839; type species: Cobitis obscura Swaison = Cobitis savona Hamilton, by subsequent designation of Jordan, 19 19; feminine. Body elongate, of almost uniform depth, compressed posteriorly; head either depressed or compressed; snout usually blunt; adipose creast usually absent (in most species) or present only in the posterior part of the body; in a few species the posterior nostril is prolonged in a tube, which does never reach the same length as the "nasal barbel" of the species of Oreonectes. Dorsal fin short, with seven or eight, rarely ten branched rays; the free edge of the fin is slightly convex or straight, in some species slightly concave in the median part; caudal fin slightly to deeply emarginate, never straight or convex; anus immediately in front or slightly in advance of the anal fin insertion. Body either scaled or partially or totally naked; lateral line complete or incomplete. Upper lip slightly furrowed, continuos or with a narrow median interruption. Lower lip interrupted in the middle, moderately furrowed. Processus dentiformis of the upper jaw present, its degree of development differs in the various species groups of the genus; a correspondent incision on the lower jaw is present in many species, absent in others. Intestine short, with only an oddulation or a loop in its posterior part. The capsule of the anterior chamber of the air-bladder consists of two dilatations connected by a manubrium; they are never in direct contact. The posterior chamber of the air-bladder is rudimentary. Sexual dimorphism is absent in most species; in other species the male have a suborbital flap - as in Nemacheilus - and the rays of the pectorals are slightly ossified and covered by breeding tubercles, while in others there are breeding tubercles on the pectoral rays but no suborbital flap. Colour pattern: dark crossbars of variable form or number are present in all species of the genus; there is no longitudinal stripe; often a blackish or brownish spot at the base of the dorsal fin. The last transversal bar, at the base of the caudal fin, is, in many species, darker than more anterior ones; it is however always a vertical bar, never a roundish spot, as in Nemacheilus and Acanthoco bitis. Range: southern Chinatnorthern Vietnam, the Indochinese Peninsula, the western slope of Kalimantan or Borneo island ("Nemacheilus" maculiceps, Roberts, 1989, from Kapua river, is in fact a Schistura), the Indian subcontinent,

11 440 PETRU M. BANARESCU. TEODOR T. NALBANT S. acuticephala from the Irrawaddy basin and S. rnultifasciata from the Ganges and Brahmaputra could be included within this group, too; they have however a more deeply forked caudal fin. (3) the montana group The few species of this group differ from those of the rupecula group in having a not-depressed head (which is about as wide as high) and a more deeply forked caudal fin. Only three species: montana and beavani fiom Ganges River basin of the Himalayas and poculi from the Irrawaddy and Mekong rivers basins; the latter is one of the few fish species shared by these two river basins; in general, the fish fauna of the Salween has closer ties to those of the Irrawaddy, Brahmaputra and Ganges, while the fish fauna of the Mekong basin has closer ties with those of the Chao Phraya, of the Malay Peninsula and of westem Indonesia. (4) the scuturiginu group Head slightly depressed. Body almost completely scaled; the scales of the lateral line are often perforated. Lateral line complete or almost complete. Caudal fin rather deeply forked. Most species have sexual dimorphism: suborbital flap and breeding tubercles on the pectoral fin (scaturigina, bella, prashadi, mahnerti, maepaiemsis, desmotes, kohchangensis); in S. sikmaienis there is only a slight indication of suborbital flap, like a groove (Kottelat, 1990); S.devdevi seems not to have sexual dimorphism. The processus dentifomis is present in most species (except perhaps S. sikmaiensis) but is feebler than in the species of the rupecula group and there is no corresponding notch on the lower jaw. Crossbars, in variable number, are present in all species of the group, but in most of them these are complete, extending fiom the back to (or almost to) the belly, while in S. prashadi they are divided each in a few irregular dorsal spots and a short vertical bar or spot. Following species can be assigned to this group: scaturigina, zonata (conspecific according to Menon, 1987) and devdevi from the Irrawaddy basin, prashadi &om the Brahmaputra and Chindwin, mahnerti and maepaiensis from the Salween basin, bella from the Mekong, desmotes (= myrrnekia) from the Chao Phraya basin, kohchagensis from the coastal rivers of south-eastem Thailand. ' (5) the alepidota-pakistanica group Head depressed; caudal fin truncate or slightly emarginate; totally devoid of scales; lateral line long, reaching at least to below the dorsal fin; no sexual dimorphism; processus dentiformis is strongly developed and a corresponding

12 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 439 Ceylon island, Afghanistan and eastern Iran; absent from most of Indonesia, the Malay Peninsula, High Asia, the Middle and Near East, the north of East Asia and probably from the Changjiang (Yangtse) River basin, too. This is the largest and most variable genus of the subfamily; it probably is, in present acceptance, polyphyletic, since it includes several rather unsimilar groups of species, which can not, however, be clearly delimited. Following groups of species can be distinguished: (1) the rupecula group (Schistura s.str., since the group comprises the generotype). Head depressed, wider than deep; scales present at least in the median and posterior part of the body; lateral line complete or almost complete; caudal fin slightly to moderately emarginate (almost straight); processus dentiformis strongly developed, with a corresponding incision on the lower jaw. No suborbital flap in males, but in some species there are breeding tubercles on the enlarged rays of the pectoral fin. This is apparently the largest group of the genus, comprising most or all species from the south of East Asia (hingi, humilis, incerta, fmciolata, chapaensis and probably three more recently North Vietnam described species), many of those of the Mekong and Chao Phraya basins (spilota, waltoni, sexcaudata, breviceps, fowleriana, kengtungensis, nicholsi, schultzi, spiloptera, magnzjluvis), several species fiom the Burmanese rivers Irrawaddy and Salween (reidi, vinciguerrae, moeiensis, shanensis, paucifasciata, balteata) and several Indian ones (rupecula - Fig. 7 - inglisi, notostigma, altipedunculata, denisonii, dayi). To the same group may also belong S. alticrista fiom the Salween River basin (Kottelat, 1990) that seems however to have a true adipose creast on the caudal peduncle and no incision on the lower jaw. This group does not range in the Indus River basin and west of it. (2) the cincticauda group The species of the cincticauda group differ from those of the preceding group mainly in their much shorter lateral line. They usually have a depressed head, well developed processus dentiformis, scaled body; the males have no suborbital flap, but in some species the rays of the pectoral fin are enlarged. The group is confined to the Indochinese Peninsula; following species are ascribed to it: cicticauda, balteata, paucicincta (Salween River basin), paucifasciata, raoi, kunjupkhulensis, malaisei (Irrawaddy basin), menamensis, dubia (Chao Phraya or Menam basin), daubentoni, laterimaculata (Mekong River basin) and robertsi (Peninsular Thailand, being apparently the only species of Schistura present in the Malay Peninsula:

13 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 441 notch is present on the lower jaw; vertical crossbars; sometimes a slight adipose creast in the posterior part of the caudal peduncle. Restricted to the Indus river basin in Pakistan and India. Four or five species: alepidota, pakistanica, nalbanti, paludani and panjabensis (Mirza et all ). Menon (1987) considers nalbanti to be a junior synonyms of punjabenis, and alepidota a junior synonym of pakistanica; he may be right in respect of nalbanti (although this has much longer pectoral fins than punjabensis) but not in respect of alepidota, which differs very much from pakistanica by having a shorter lateral line, an almost truncate (versus slightly emarginate) caudal fin, a comparatively better developed adipose creast, more crossbars, numerous spots on the upper face of the head, etc. (6) the kessleri group Small size; rather compressed head; totally devoid of scales or with a quite reduced scaled area on the caudal peduncle; short lateral line, not reaching to the middle of the body; emarginate or slightly forked caudal fin; reduced processus dentiformis; no sexual dimorphism; seven branched dorsal rays; from Turkmenia and eastern Iran to the Indus River basin in Pakistan; two species, kessleri and lindbergi (Mirza et all., 198 1). (7) the naseeri-baluchiorum group Small size; rather compressed head; pointed snout, always narrowed anteriorly; body scaled, at least in its posterior part; lateral line of variable length, never complete; forked caudal fin; reduced or moderately developed processus dentiformis; sexual dimorphism present in most species, apparently absent (or not yet recorded) in two of them: S. prashari and S. naseeri. From Iran to the Indus River basin; 12 species, one of them, S. afasciata, is almost unique in the genus in having no crossbars (Mirza et all., 1987). This group may perhaps be lumped with the scaturigina group. Menon (1987) accepts only five of these species as valid. He may be right when synonymizing naseeri with prashuri; we also agree that S. anambarensis, harnaiensis and machensis are closely related to balachiorum, but they are at least subspecifically distinct; S. macrolepis may be a subspecies of S. arij (not a full synonym of it), but S. curtistigma and S. afasciata are in no case synonyms of arifi. (8) isolated species - S. savona (genotype of Acoura Swaison, 1939) (Fig. 8). Head about as wide as deep; caudal fin deeply forked; slight indication of processus dentiformis; no notch on the lower jaw; apparently mo sexual

14 442 PETRU M. BANARESCU, TEODOR T. NALBANT dimorphism. Body almost completely scaled; lateral line complete. Contrary to most other species of the genus, the dark crossbars are much wider than the light interspaces. Ganges River basin in the Himalayas. - S. manipurensis (Fig.9): similar to the cincticauda group but with reduced processus dentiformis and the males provided with a well developed suborbital flap. Brahmaputra and Chidwin basins. - S. semiarmata; head comparatively deep; caudal moderately forked; sexual dimorphism present; processus dentiformis strong, but only a slight notch on lower jaw; scales imbricate, present on most of the body; lateral line complete; crossbars extended to or slightly below the lateral line; numerous minute black spots on body sides. South India. - S. rendahli - similar to S. denisoni but with very reduced lips and a shorter lateral line; a species known from a single specimen (Bhfirescu and Nalbant, 1986) from Nadur Madhmeswar, southern India. - S. shadiwalensis: similar to the species of the S. montana group, but the crossbars are connected by a median longitudinal stripe and the peritoneum is silvery. Indus River basin in Pakistan. - S. microlabra: head depressed, snout blunt; body almost completely naked, scales being present only on the caudal peduncle; lateral line short, extending below the dorsal fin only; caudal fin slightly notched, the stripe on its base interrupted; mouth small; processus dentiformis moderately developed; Indus River basin in Pakistan. - S. cristata (Fig. 11): characterised by a short but high and thick adipose creast on the caudal peduncle which is much shorter and thicker than in the species of Paracobitis; crossbars only in the posterior part of the body; processus dentiformis well developed; no sexual dimorphism. Rivers of Afghanistan and Turkmenistan. 6. Oreias Sauvage, 1974 (Fig. 12) Type species: Oreias dabryi Sauvage, by monotypy; masculine. Body low, slightly compressed or subcylindrical, totally devoid of scales. Caudal peduncle long and low, with no trace of adipose keel or of thickening; its least depth larger or slightly smaller than its width at the level of the anal fin insertion. Caudal fin truncate or forked. Dorsal fin with 7-8, rarely 6 branched rays; its distal edge emarginate. Head depressed; eyes on its dorsal face. Lateral line complete, extending to the base of the caudal fin. Processus dentiformis strongly developed (Fig. 12 B); a corresponding notch is present on the lower jaw.

15 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 443 Lips moderately to slightly furrowed. Intestine short; posterior chamber of the airbladder rudimentary. No sexual dimorphism. Colour pattern variable; never complete (circular) crossbars. High Asia; four species: three in the basin of the upper Chanjiang (Yangtse), one in that of the Aral Sea. Some authors lump the three Changjiang species in a single one; they differ however in the shape of the caudal fin, the width of the head and of the caudal peduncle (Bhkescu and Nalbant, 1976). 7. Paracobitis Bleeker, 1963 (Fig. 13) Type species: Cobitis malapterura Valancienne, by original designation; feminine. Synonyms: Pseudodon Kessler, 1974; type species: Cobitis longicauda Kessler, by monotypy; nomem preocupatum. Adiposia Annadale and Hora, 1920; type species: Nemacheilus macmahoni Chauduri = N. rhadineus Regan, by original designation. Body elongate, compressed posteriorly; head strongly depressed. Caudal peduncle long and low; a long dorsal adipose creast, of uniform depth, extends from behind the dorsal fin to the caudal. Caudal fin slightly emarginate or truncate (rounded in P. boutanensis). Scales present; lateral line 'complete. No sexual dimorphism. Processus dentiformis strongly developed; the correspondent notch on the lower jaw is present, but reduced. Colour pattern variable: either incomplete crossbars or an irregular net of small spots or punctuation. The genus is exclusively western Asian: it comprises five described species: malapter-urus (the Tigris-Euphrates basin), rhadineus (= macmahoni) (rivers of Turkmenistan, northern Iran and the Halmand River), longicauda (the Aral Sea encatchment area), ghazniensis (Ghani River, Helamand River basin), boutanensis (a part of the Helmand basin in Afghanistan) and a few undescribed ones. The species Cobitis tigris Heckel and Nemacheilus cristatus Berg, once ascribed to Paracobitis (Bakescu and Nalbant, 1966) actually belong: the former to Orthrim, the latter to Schistura. 8. Homatula Nichols, 1929 Type species: Nemacheilus potanini Giinther, by original designation; feminine. The two species of.the genus closely resemble Parcobitis, having a similarly long (however somewhat lower) adipose creast of uniform depth; they

16 444 PETRU M. BANARESCU, TEODOR T. NALBANT remember however, in colour pattern (regular metameric crossbars) the Schistura rupecula - group of species. They also have a depressed head, complete lateral line, a strong processus dentiformis on the upper jaw, a coresponding incision on the lower jaw and no sexual dimorphism. We formely placed them in Schistura (Banarescu and Nalbant, 1974), while recent Chinese authors (e.g. Zhu, 1969) ascribe them to Pracobitis. We now prefer to include them in a distinct genus, mainly because of the great geographical distance between their range and that of the western Asian (not High Asian) Paracobitis. Hornatula comprises two species, potanini and variegata (Nernacheilus variegatus Sauvage and Dabry, 1874 = N. berezowskii Giinther, 1897 = N. oxygnathus Regan, 1908), both from the upper Chingjiang (= Yangtze) river basin in Sichuan, Yunnan and southern Gansu. Two other species and one subspecies of "Paracobitis" have been described more recently from China: P. anguillioides Zhu and Wang, 1985 from the Yunnan stretch of the Lankangjiang or Mekong River (that belongs to the South Asian subregion), P. wujiagensis Ding and Deng, 1990, from a tributary of the Changjiang in Sichuan, P. oligolepis Cao and Zhu, 1989 from the Xijiang basin of Yunnan and P. erhaiensis Zhu and Cao from Yunnan (Changjiang basin) (Zhu, 1989; Chu and Che, 1990). P. oligolepis is considered by Chu and Che (op. cit.) a subspecies of variegatus and may be a Homatula. The three other species have a much shorter adipose creast; possibly P. anguiloides and erhaiensis belong to Schistura, while P. wujiangenis, which also has a ventral adipose keel (beside the dorsal one) and a very small mouth, apparently of a pelicular type, may represent a distinct genus. 9. Longischistura new genus (Fig. 14) Type species: Nemacheilus striatus Day. Etymology: after longus, Latin for long, and Schistura; the name indicates that the genus is more elongate than Schistura and has a longer dorsal fin. Body elongate, of almost uniform depth; head depressed; a long adipose creast extends from a short distance behind the dorsal to the caudal base. Caudal fin rather deeply forked. Dorsal fin long, with ten branched rays; its edge straight or quite slightly convex. Anal fin with five branched rays. Origin of pelvics under the middle of the dorsal fin base. Processus dentiformis moderately developed; no corresponding incision on the lower jaw. Body almost completely scaled; lateral line complete or almost complete, but hardly distinct in its posterior part; its orifices perforate the scales. Lips moderately furrowed (Fig. 14 B). Intestine short. The two halves of the air-bladder capsule in direct contact, not connected by a

17 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 445 manubriurn. posterior chamber of the air-bladder strongly reduced, often totally absent (Fig. 14 D). No sexual dimorphism; narrow crossbars, extending from the dorsal almost to the ventral side. No species of Schistura, Parcobitis or Hornatula has such a long dorsal fin (never more than nine branched rays) and two halves of the air-bladder capsule in direct contact. A single species surely belongs to this genus: L. striata from southern India. Nemacheilus bhimashari Hora from the same area has a similarly long dorsal fin, but a much deeper body, fewer and broader crossbars. Menon (1987) considers it a junior synonym of striata; according to the description and illustration it is specifically distinct, its generical position being obscure. 10. Sectoria Kottelat, 1990 (Fig. 15) Type species: Nemacheilus atriceps Smith, by original designation; feminine. Similar to Schistura (especially to the species of the rupecula group), having a depressed head, short dorsal fin (with 8-9 branched rays) with almost straight distal edge, slightly emarginate caudal fin, almost complete lateral line; no sexual dimorphism. It differs however from Schistura having no processus dentiformis and very thin lips, the jaws being covered by a horny sheath with sharp edges; intestine longer than in all Schistura, with several loops (Fig. 15. C), The only species in the genus, S. atriceps, from the Chao Phraya drainage area, bears much similarity with S. bucculenta from the Mekong River Physoschistura Banarescu and Nalbant, 1982 (Fig. 16) Type species: Nemacheilus brunneanus Annadale, 19 18, by original designation; feminine. Elongate or rather deep nemacheiline loaches with scaled body; short lateral line not reaching beyond the dorsal fin; eight or nine branched dorsal rays; distal margin of the dorsal fin slightly convex; caudal fin forked. Processus dentiformis feebly to moderately developed; no corresponding incision on the lower jaw. The two halves on the air-bladder capsule joined and coalescent on their inner face (not connected by a manubriurn); posterior chamber of the airbladder well developed, not encapsulated, more or less conical, in direct contact with the capsule (Fig. 16 D). Brownish, irregular crossbars on the body.

18 446 PETRU M. BANARESCU, TEODOR T. NALBANT The original description of the genus mentions two rather unsimilar species (the deep-bodied P. brunneana and the more elongate P. elongata ) which have however almost identical air-bladders. Kottelat (1990) accepts the genus as valid, adding new diagnostic characters ("mouth strongly arched, times wider than long; lower lip with a median interruption, forming two lateral broadly triangular pads with deep fuiurow"), describing a new species, P. pseudobrunneana and ascribing to it three more species: Nemacheilus raoi Hora, N. rivulicola Hora and N. shanensis Hora. P, pseudobrunneana has the same type of air-bladder as P. brunneana and surely belongs to this genus; the generical position of the three other species (none of which has been available to us) remains uncertain; N. raoi is listed here as a possible member of the Schistwa cincticauda group of species. 12. Nun Bhaescu and Nalbant, 1982 (Fig. 17) Type species: Cobitis galilaea Giinther, 1864, by original designation; masculine. Elongate and slightly compressed body, totally devoid of scales; head slightly deeper than wide; incomplete lateral line, extending to above the last anal : ray; dorsal fin in the middle of the body, with branched rays; its distal margin is slightly convex; anal fin with constantly six branched rays; caudal fin feebly emarginate or almost truncate; insertion of pelvic fins slightly behind that of the dorsal fin; a slight indication of adipose keel on the caudal peduncle; lips slightly crenulated; processus dentiformis of the upper jaw strongly developed, but no corresponding incision on the lower jaw. No sexual dimorphism. Intestine short, with a single loop; peritoneum light brownish. Posterior chamber of the airbladder rudimentary. Colour pattern variable; in some specimens there are well delimited lateral spots and some blotches above them, in others the lateral spots are vaguely delimited and do not differ from the blotches above. The genus bears much similarity to Schistura, differing from it in the higher number of branched rays in the dorsal and mainly in the anal fin; the only other nemacheiline loach with six branched anal rays is the tropical Afiican "Nemacheilus" abyssinicus. Nun comprises a single species, N. galilaeus, endemic to the Jordan River basin. 13. Mesonoemacheilus Banarescu and Nalbant, 1982 (Fig. 18) Type species: Nemacheilus triangularis Day, designation; masculine , by original

19 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 447 Body elongate, slightly or moderately compressed anteriorly, strongly compressed posteriorly, covered by scales; head slightly depressed or compressed. Lateral line incomplete, reaching either far beyond the middle of the body or ending in front of the dorsal fin and interrupted. Scales with excentric focal zone. Dorsal fin with 7, 8 or 10 branched rays; its outer edge is slightly convex in four of the species, straight in M pulchellus: caudal fin forked. No nasal barbels. lips usually more deeply furrowed than in Schistura. In four of the five species, the two rostral barbels of each side are confluent at their bases; in M. reticulofmciatus they are not confluent. Processus dentiformis moderately developed; no correspondent incision on the lower jaw. Sexual dimorphism recorded in a single species: M. herrei in which the males have a movable suborbital flap and the pectoral rays are broadened, thickened and covered by breeding tubercles; the three other south Indian species of the genus may display sexual dimorphism, too, but not M. reticulofmciatus. The posterior, free chamber of the air-bladder is rudimentary; the two halves of the capsule are roundish and connected by a long and narrow manubriurn (Fig. 18 D). A net of irregular dark bars and spot on the body, often acquiring a large extension and reducing the fundamental whitish coloration to isolated spots. A black spot on the middle of the caudal fin base; this is roundish in three species, while in M. guentheri and in many specimens of M. reticulofmciatus it is vertical and deeper than long. The only other nemacheiline genus having a black spot on the middle of the caudal base is Nemacheilus. Four or five species of the genus also share one character with Schistura savona, type species of Acoura: the confluence of the rostral barbels of each side; the five Mesonoemacheilus species and S. savona also are the only nemacheilines having more brownish colour than whitish. Possibly they are interrelated, in which case Acoura may deserve generic rank and Mesonoemacheilus (with the possible exclusion of M. reticulofmciatus) may become its junior synonym. Mesonoemacheilus comprises five species: four in the Western Ghats (triangularis, guentheri, herrei and pulchellus, the latter being the only one with ten branched dorsal rays) and reticulifasciatus in the Brahrnaputra River basin, north-eastern India. Menon (1987) considers M. herrei as a synonym of guentheri. Actually, both differ in colour (the white coloration is reduced to V- and Y-shaped spots in herrei, to roundish ones in guentheri), size of scales (larger in herrei) etc. 14. Nemachilichthys Day, 1878 (Fig. 19) Type species: Cobitis rupelli Sykes, 1841, by monotypy; masculine.

20 448 PETRU M. BANKRESCU. TEODOR T. NALBANT Elongate, slightly compressed, almost cylindrical body, almost completely scaled (excepting the breast, belly, and a narrow stripe between anal fin and anus. Eye larger than in most nemacheilines; snout pointed. Anus some distance in front of the anal fin. Dorsal fin in the middle of the body, with branched rays; its distal edge notched. Caudal fin deeply forked, its lobes pointed. No trace of adipose creast. Lateral line complete or almost complete. Lips moderately furrowed; there is a median incision and 1-4 pairs of grooves on the upper lip, a broader incision and 2-4 pairs of grooves on the upper lip. Processus dentiformis moderate; no corresponding incision on the lower jaw. Intestine short, with a single loop. Posterior chamber of the air-bladder capsule rudimentary. A peculiar type of sexual dimorphism: there is no suborbital flap, nor are there breeding tubercles or thickened rays in the pectoral fin, but the colour pattern differs in two sexes: the males have complete and well-delimited crossbars, the females only , incomplete and unsharply delimited ones. The genus is endemic to the basin of Kistna River, southern India; it comprises two species with vicariant range: ruppelli in the upper Kistna proper, shimonogensis in its tributary, Thunga River. 15. Orthrias Jordan and Fowler, 1903 (Fig. 20) Type species: Orthrim oreas Jordan and Fowler, 1903 (= Cobitis toni Dybowski) by original designation; masculine. Body elongate, slightly compressed or almost cylindrical, completely or nearly completely covered by scales. Lateral line long, complete or at least reaching beyond the middle of the body; its tubes do not penetrate the scales. Head slightly depressed or compressed. Lips, especially the lower one, moderately furrowed. No nasal barbels. Dorsal fin with 7-9, rarely 10 branched rays; its edge usually slightly concave (slightly convex only in 0. panthera). Pelvic fins inserted slightly behind the origin of the dorsal. Caudal fin slightly emarginate to deeply forked. Usually no trace of dorsal creast in the posterior part of the peduncle. Processus dentiformis feebly or moderately developed; no corresponding incision in the lower jaw. Colow pattern variable; in some species there is a rather irregular net of spots and punctuations, in others more or less regular rows of metameric spots, while in others - regular metameric crossbars, similar to those in most species of Schistura, extending from the back almost to the ventral face. Sexual dimorphism present, but differing from that in Nemacheilus, many Schistura etc.: the rays of the pectoral fm in males are broadened, thickened and, at least during the spawning season, covered by breeding tubercles; breeding

21 NEMACHEILJNAE (TELEOSTEI) GENERICAL CLASSIFICATION. 2 NEW GENERA 449 tubercles also develop on the sides of the head; these are uniformly disposed, not separated by a groove in two fields. There is no subocular flap. A number of authors, mainly American and Chinese ones, consider that the valid generic name for the group of species here included in Orthrias is Barbatula Linck, 1789, type species Cobitis barbatula Linnaeus by tautonomy, Orthrias being its junior synonym (it is worth mentioning that Nichols (1943) includes in "Barbatula" mainly species now in Triplophysa, Schistura, Homatula and Oreias, a single one in Orthrias, and Zhu (1990) includes only species of Triplopphysa). However, as Rendahl (1933) remarked, in proposing the name Barbatula, Linck (1789) mentions "hierher gehoren Cobitis barbatula taenia", i.e. the type species is Cobitis taenia. Kottelat's (1990) objection that a coma has been intended between the names barbatula and taenia can not be proved. We retain therefore Orthrias as valid generical name for this group of species. Baarescu and Nalbant (1966) described the subgenus Oxynoemacheilus, type species Cobitis persa. All five species included by them in Oxynoemacheilus actually have the characters of Orthrias, only their body and especially caudal peduncle being lower and the caudal fin forked (in 0. barbatulus, 0. angorae etc. it is only slightly emarginate). Their drawing of a male of "Noemacheilus (Oxynoemacheilus)" persa (their Fig. 3) seems to indicate a subocular pad; actually. the species has no mobile subocular flap (like that in Nemacheilus and Schistura), but a small groove. Oxynoemacheilus is hence a synonym of Orthrias. Orthrias comprises about 18 valid described species, most in western Asia, two in Europe (0. barbatulus and the north-caucasian 0. merga besides the subspecies 0. brandti bureschi) and one in Siberia and the northern part of the East Asian subregion (0. toni). There are surely many others undescribed. 16. Seminemacheilus new genus (Fig. 2 1) Type species: Nemacheilus lendli Hanko, 1924 Etymology: the name is a combination of the Latin word semi = half and the name of the genus Nemacheilus; it indicates the short lateral line of the type species. Synonym: Heminoemacheilus Banarescu, 1977 (not of Zhu and Cao, 1987; nomem nudurn). Gender: masculine. Body deeper than in most other nemacheilines, moderately compressed, totally or almost totally devoid of scales. Lateral line very short, reaching only to or almost to below the middle of the dorsal fin. Snout blunt; eyes small,

22 450 PETRU M. BANARESCU, TEODOR T. NALBANT distant. A feebly developed dorsal adipose creast is present on the caudal peduncle. Dorsal fin with 7, rarely 8 branched rays; its distal edge is convex (almost straight in young specimens). Insertion of pelvic fins opposite that of the dorsal fin. Upper lip almost smooth; lower one interrupted in the middle and with one or two pairs of grooves. A slight indication of processus dentiformis. Free, posterior chamber of the air-bladder rudimentary; both halves of the airbladder capsule spherical, in direct contact, not connected by a manubriurn (a character shared with Physoschistura - especially P. brunneana - Longischistura and the Triplophysa dalaica/kulmanni group of species). Sexual dimorphism little known and probably feebly developed: the first simple ray of the pectoral fin in males of S. lendli is thickened; breeding tubercles have not been found but they may develop during the spawning season. There is no subocular flap. Colour pattern: irregular dark points and small spots, grouped in longitudinal belts. A single described species, S. lendli, in northern and central Anatolia; it is totally devoid of scales. An underscribed species, provided with a single row of scales has been found in Iran. (Nalbant and Bianco, in prep.). We initially intended to describe this genus as Heminoemacheilus: it has been mentioned under this name, without description, in a zoogeographical paper (Banarescu, 1977). In the meantime, this name has been proposed for another species of noemacheilines. 17. Triplopphysa Rendahl, 1933 (Figs 22-25) Synonyms: Diplophysa Kessler, 1974; type species: D. strauchii Kessler, 1874, by subsequent designation (Berg, 19 16); preoccupied in Coelenterata; Deuterophysa Rendahl, 1933; replacement name for Diplophysa; also preoccupied (in Lepidoptera); Triplophysa Rendahl, 1933: 2 1 ; type species: Nemacheilus hutjertjuensis Rendahl, 1993, by original designation; feminine (as subgenus); Tauphysa Rendahl, 1933: 22; type species: Nemacheilus kungessanus Kessler, by original designation (as subgenus); Hedinichthys Rendahl, 1933 : 26; type species: Nemacheilus yarkandensis Day. by original designation; masculine (as subgenus); Didymophysa Whiteley, 1950; replacement name for Diplophysa Kessler; Diplophsoides Fowler, 1958; replacement name for Diplophysa Kessler and Deuterophysa Rendahl.

23 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 451 Nemacheilines of variable size, among them are the largest members of the subfamily. Body elongate, either slightly to moderately compressed or almost cylindrical; caudal peduncle either compressed or cylindrical, in some species very slender. Totally devoid of scales (Zhu, 1981, mentions however the occurrence of minute scales in two species). Lateral line complete in most species, incomplete and short in a few ones. No adipose keel on the caudal peduncle, but in some species there is a small and short dorsal elevation in the posterior part of the peduncle (at the base of the caudal fin). Dorsal fin short, with 7-9 branched rays; its distal edge slightly notched or almost straight; caudal fin either slightly to deeply furcate or truncate (straight). Upper lip more deeply furrowed than in other genera, in many species it is divided in several lobes, which can be subdivided, too (Fig. 23 A-E); never with roundish papillae resembling those in Acanthocobitis. Lower lip strongly furrowed, too, at least in most species (Fig. 23 A-E). Neither processus dentiformis on the upper jaw, nor incision on the lower one. Length and shape of the intestine very variable: in some species it is short, having a single loop, in others it is long to very long, describing many convolutions (Fig. 24 A-G). The two chambers of the air-bladder capsule are either connected by a manubrium (in most species) or in direct contact, as in Longischistura, Physoschistura and Seminemacheilus (Triplophysa kullmanni, T. dalaica and T. posteroventralis) (Fig. 25 B). The shape and degree of development of the posterior, free chamber of the air-bladder is subject to a strong variation within the genus; one can distinguish four types of structures: (1) posterior chamber rudimentary (Fig. 25 A, B); (2) posterior chamber consisting in a distal wide dilatation and a long and very narrow tube connecting this dilatation to the anterior, encapsulated chamber (Fig. 25 C); (3) posterior chamber well developed, but elongate (not vesicular), in almost direct contact with the anterior chamber or connected to it by a short tube (Fig. 25 D); (4) posterior chamber very large, unique (Fig. 25 E), or subdivided in two parts and connected to the anterior chamber by a short tube (Fig. 25 F). The sexual dimorphism is strongly marked; in males, the outer 6th to 8th divided rays of the pectoral fin are thickened, broadened and covered by breeding tubercles; breeding tubercles are also present on the whole side of the head, being - in most species - arranged into two groups, separated by a deep groove extending fiom the eye almost to the insertion of the maxillary barbel. In T. yarkandensis there is no such groove and the breeding tubercles are restricted to a triangular area on the antero-inferior side of the eye (Zhu, 1981; we did not observe this character in the few available specimens). In some species breeding tubercles are present, at least during the spawning period, also on the body sides. There is no mobile subocular flap, as in Nemacheilus and other genera. The same type of

24 452 PETRU M. BANARESCU, TEODOR T. NALBANT sexual dimorphism occurs in Orthrias, but the subocular groove delimiting two groups of breeding tubercles on the sides of the head is absent in this genus. The colour pattern is variable, too: in some species there are more or less regular vertical crossbars, as in many Schistura species; in others a multitude of irregular dark small blotches, spots, points etc. cover the body sides; a fragmentation of the crossbars and a combination of irregular spots and crossbars is characteristic to other species. Zhu (1981) distinguishes four main types of coloration - channel colour, grass-like colour, stone-mottled colour and dusky colour - considering the each of them is characteristic to the species living a special habitat. Rendahl (1933), the author of the most thorough study of the High Asian loaches, and an excellent anatomist, payed much attention to the structure of the air-bladder and ascribed the species with well-developed posterior chamber of the air-bladder to three new subgenera: Triplophysa (posterior chamber very large and subdivided: Fig. 25 F), Tauphysa (posterior chamber large but relatively narrow and simple: Fig. 25 D) and Deuterophysa (new name for Diplophysa Kessler, preoccupied; later changed in Didymophysa Whitely; posterior chamber consisting in a distal, wide vesicle, connected to the anterior, encapsulated chamber by a very long and narrow tube: Fig. 25 C). He ascribed most scaleless High Asian nemacheilines with rudimentary posterior chamber to Nemacheilus s.str., with the exception of N. yarkandensis, characterised by a strong development of the anterior chamber and of the capsule, connected not only with 2nd and 4th vertebrae, but also with the fifth one. He included N. yarkandensis in a fourth new subgenus, Hedinichthys. In an earlier paper (Banarescu and Nalbant, 1975), we expressed the opinion that the degree of development of the posterior air-bladder chamber is an adaptative character, the species with well-developed posterior chamber being lacustrine; species with quite different types of air-bladder are very similar in other characters and vice-versa - species having similar or even identic types of airbladder often differ in other characters. We therefore lumped three of Rendahl's subgenera and the species with rudimentary posterior chamber in a single genus, adopting for them the name Triplophysa (that has page priority), while recognising Hedinichthys as distinct genus. Our view-point has also been adopted by Kottelat (1990) and Zhu (198 I), none of which mentions our 1975 paper. Zhu (op. cit.) considers Hedinichthys as subgenus of Triplophysa, described another new subgenus, Quinghaichthys, for N. alticeps Herzenstein. We presently accept Zhu's view-point concerning Hedinichthys, having momentary no own opinion about his new subgenus. A more thorough study of the several tens of species of Triplophysa which we had the opportunity to examine is in preparation.

25 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION. 2 NEW GENERA Oreonectes Giinther, 1966 (Figs 26-28) Type species: Oreonectes platycephalus Giinther, 1868, by monotypy; masculine. Synonyms: Octonema Martens 1869; type species: Homaloptera (Octonema) rotundicauda Martens, by monotypy; feminine. Octonema Herzenstein, 1887; type species: 0. pleskei Herzenstein (= Diplophysa costata Kessler) by subsequent designation; preoccupied. Lefua Herzenstein, 1888; type species: Octonema pleskei Herzenstein (= Dyplophysa costata Kessler) by monotypy; replacement name for Octonema Herzenstein, not of Martens; feminine. Elxis Jordan and Fowler, 1903; type species: E. nikkonis Jordan and Fowler, by original designation; masculine. Indoreonectes Rita and Banarescu, 1978: type species: Oreonectes keralensis Rita and Nalbant, 1978, by original designation; masculine. Sundoreonectes Kottelat, 1990; type species: Nemacheilus obesus Vaillant, 1902, by original designation; masculine. Body elongate, low and thick, sub-cylindrical, completely covered by well developed scales (larger than in most nemacheilines); these have an almost circular shape, are imbricate, their focal zone is small and excentric. The scales also cover the breast and throat. The caudal peduncle is high and compressed; a dorsal adipose creast is present, at least in the posterior part; sometimes also a ventral adipose creast. The head is depressed, wider than deep; the eyes are small, espaced, have a dorsal position. The anterior nostrils are prolonged in a long nasal barbel. The lips are fleshy, narrower than in most other nemacheilines; the anterior one is continuos and smooth; the posterior one with a narrow median interruption, slightly furrowed or smooth. Lateral line short or absent. Dorsal origin nearer caudal base than tip of snout; contrary to most of other genera of nemacheilines, the pelvic fins are in advance of the dorsal. Distal edge of the dorsal and anal fins rounded; caudal fin rounded or almost straight. Anus a short distance in front of anal fin. Shape and degree of development of the posterior chamber of the air-bladder variable within the genus. Kottelat ( 1984, 1990) contests the monophyly of Oreonectes as delimited by Banarescu and Nalbant (1968) and by Rita et al. (1979), considering that the three accepted subgenera are distinct genera, the only character common to them - presence of a nasal barbel - also occurring in some Schistura. Actually these subgenera share also other characters, for example size of scales, rounded caudal

26 454 PETRU M. B,~N&SCU, TEODOR T. NALBANT fin, anterior position of the pelvic fins comparatively to the dorsal. It is worth mentioning that Kottelat himself (1990), when describing a new genus for the species obesus (ascribed by us to Oreonectes) chosed the name Sundoreonectes, hence suggesting at least a similarity with the three other "genera". The genus can be divided into four clearly delimited subgenera, each with another distribution, the range of the genus being disjunct: Oreonectes Giinther s. str. (= Octonema Martens; Fig. 26): nostrils distant, anterior one just in front of eye; nasal barbels shorter than in the three other subgenera, reaching to about the anterior margin of eye; posterior chamber of the air-bladder large, in direct contact with the capsule of the anterior chamber; dots or a longitudinal stripe of body sides; no crossbars. No lateral line. Southeastern China; three described species. Lefua Herzenstein (= Octonema Herzenstein, not of Martens; = Elxis Jordan and Fowler); nostrils distant; nasal barbels reaching at least to the middle of the eye; posterior chamber at the air-bladder large, vesicular, connected to the capsule by a long and narrow duct; no lateral line; a longitudinal stripe and small dots on body sides; no crossbars. Northern areas of East Asia; three species. Indoreonectes Rita and Banarescu (Fig. 27): nostrils close to each other, the posterior one being distant from the eye; length of nasal barbels variable; posterior chamber of the air-bladder rudimentary (Fig. 27 C); lateral line present, short. Crossbars on the body sides. South-western India; two species. Sundoreonectes Kottelat, 1990 (Fig. 28): nostrils close to each other, the posterior one distant from the eye; nasal barbels long (as in Lefia): lateral line present, short, not reaching beyond the origin of the pelvics; posterior chamber of the air-bladder vesicular, connected to the anterior chamber by a long, narrow duct (Fig. 28 C); crossbars on body sides. South-eastern Asia: Malay Peninsula and Borneo (Kalirnantan) island; two species. 19. Aborichthys Chaudhuri, (Fig. 29) Type species: Aborichthys kempi Chaudhuri, 1913, by monotypy; masculine. Body elongate, of uniform depth, depressed or sub-cylindrical anteriorly, strongly compressed posteriorly. Caudal peduncle strongly compressed, provided with a dorsal adipose creast. Dorsal short, with 6-7 branched rays, placed in the middle or in the anterior half of the body; its distal edge convex or almost straight; pelvic fins slightly in advance of the dorsal; caudal fin rounded. Anus shifted anteriorly, much closer to the pelvics than to the anal fin insertion. Body

27 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 455 completely covered by large, imbricate, elliptic scales. Lateral line present but incomplete. There is a short outgrowth of the nasal membranes, but not nasal barbels as in Oreonectes. Lips fleshy, the upper one continuos, the lower one interrupted in the middle; the lips are not fringed; papillae have been mentioned in the literature on the upper lip, but could not be distinguished in the specimens we examined (Fig. 29 B). No processus dentiformis; no sexual dimorphism. Intestine with two or three loops. The shape of the posterior chamber of the air-bladder could not be distinguished in the specimens we examined. Complete or almost complete crossbars on the body sides, better marked in the posterior two thirds of the body. An intensive blackish dorsal spot on the base of the caudal fin, similar to that in Acanthocobitis. Southern slope of the Himalayas (encatchment area of Ganges and Brahmaputra rivers); four described species (one of them perhaps invalid). 20. Yunnanilus Nichols, 1925 (Fig. 30) Type species: Nemacheilus pleurotaenia Regan, 1904, by original designation; masculine. Synonyms: Eonemachilus, Berg, 1938; type species: Nemacheilus nigromaculatus Regan, 1904, by original designation; masculine. Petruichthys Menon; type species: Nemacheilus brevis Boulenger, 1893; by original designation; masculine. Body size variable; five of the nine described species till 1988 are among the deepest members of the subfamily; the four others have a more elongate body. Caudal peduncle short, deep. Mouth more anterior than in the other nemacheilines, in several species being even terminal and upwardly directed. Dorsal fin in the middle or behind the middle of the body, with 8-10 branched rays; its distal edge slightly convex. Pelvic fins inserted slightly behind the dorsal insertion; caudal fin slightly to moderately emarginate or even almost truncate. Scales usually present and embedded but in some species they are absent from different areas of the body. Lateral line incomplete or absent. Nostrils widely separated; anterior one prolonged into a short tube (Fig. 30 B). Kottelat and Chu (1988) mention that the upper lip has no or almost no furrows and the lower one has a medium interruption and usually two deep furrows on each side; we found, in Y. brevis, many short furrows on the upper lip and longer ones on the lower lip (Fig. 30 F). Intestine short. Posterior chamber of the air-bladder very large, vesicular, connected to the anterior, encapsulated chamber by a short duct ( at least in some species: Fig. 30 D). The genus is

28 456 PETRU M. BANARESCU. TEODOR T. NALBANT concentrated in the Yunnan Plateau of China: basins of the rivers Changjiang or Yangtse, Xijiang, Yuanjiang (upper Songkoi) and Lankangjiang (upper Mekong); one species in Inle Lake, Burma. Numerous species, many undescribed Paranemacheilus Zhu, 1983 Type species: Paranemacheilus genilipis Zhu, 1983, by original designation; masculine. Synonyme: Heminoemacheilus Zhu and Cao, 1987; type species: Heminoemacheilus zhengbaoshani Zhu and Cao, by original designation; masculine; new synonymy. Body deeper than in the most other genera of the subfamily (excepting Micronemacheilus and partially Yunnanilus), covered by minute scales; in one species the scales also extend on the sides of the head. Lateral line incomplete, extending only to above the top of the pectoral fin. Dorsal fin in the posterior half of the body, with 7-9 branched rays; its distal edge convex anteriorly, almost straight posteriorly; insertion of the pelvic fins slightly in front of that of the dorsal fin in one species, behind this insertion in the second described species. Caudal fin slightly notched. A dorsal adipose creast on the posterior part of the caudal peduncle. Lips furrowed, posterior one interrupted in the middle. All three pairs of barbels much longer than in any other genus of nemacheilines. Both halves of the air-bladder capsule connected by a broad manubriurn. Second chamber of the air-bladder wide, vesicular, not divided; it is connected to the anterior chamber by a duct that is longer than in Micronemacheilus and Yunnanlis (compare with Fig. 6 D and 30 D), but much shorter than in the species of the "Deuterophysa" - group of Triplophysa. Intestine short, with a single loop. Sexual dimorphism not mentioned, probably absent. Colour pattern: minute dark spots and points, uniformly distributed on the body sides in P. gentilkis. No specimens were available to us; the diagnosis above is based on the original descriptions of both nominal genera and of their type species. The two "genera" are extremely similar; the description of "Heminomacheilus" mentions only two differences between it and "Paranemacheilus": totally scaleless head (in P. genipilis the checks are covered by scales, a character unique in nemacheilines) and absence of postcleithrum. In our opinion, these are only specific differences. Both species, P. genipilis and P. zhengbaoshani were described from the Xijiang (formally Pearl River or Hsikiang) River basin in Guangxi Province, China, but in different localities. This genus bears a certain similarity with Micronemacheilus and Yunnanilus, having a deep body, short intestine and a very large second chamber

29 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 457 of the air-bladder which is, in all three genera, undivided, contrary to the airbladder of some Triplophysa species. 22. Turcinoemacheilus Banarescu and Nalbant, 1964 (Fig. 3 1) Type species: Turcinoemacheilus kosswigi Banarescu and Nalbant, by original designation; masculine. Body low, almost cylindrical, of uniform depth, totally devoid of scales. Lateral line short, not extending beyond the tip of the pectoral fin; its orifices are larger than in most other nemacheilines. No adipose creast on the caudal peduncle. Head depressed. short; eyes small, distant; barbels short; no nasal barbels; lips smooth, no processus dentiformis on the upper jaw. All fins small; caudal slightly notched, dorsal with seven branched rays, anal with five; pelvic fins inserted well in advance of the dorsal fin. The anus has a more anterior position than in most other genera of the subfamily (except in Aborichthys) being closer to the pelvic than to the anal fin. No sexual dimorphism. intestine short, straight. second chamber of the air-bladder rudimentary or absent. The two halves of the air-bladder capsule rounded, connected by a narrow manubrium; the capsule has irregular lateral expansions. A longitudinal brownish stripe along the body sides. Slightly delimited pale spots on the upper side of the body and above the longitudinal brownish stripe; fins unspotted. A single species in the drainage area of the upper Tigris and Euphrates rivers in Eastern Turkey, probably also in Irak. 23. Sphaerophysa Cao and Zhu, 1988 Type species: Sphaerophysa diachiensis Cao and Zhu, by original designation; feminine. Body rather elongate, slightly compressed. Dorsal fin long and low, with branched rays, its distal edge convex; caudal fin slightly notched, almost truncate; anal fin with six branched rays, as in Nun and in the undescribed Ethiopian genus of the subfamily, insertion of the pelvic fins slightly behind that of the dorsal. Insertion of the pelvic fins slightly behind that of the dorsal. The genus is unique in the subfamily in having very long and high adipose creast both on the dorsal and on the ventral sides of the caudal peduncle, extending from the end of the dorsal - and anal - fins to beyond the beginning of the caudal fin; the dorsal creast resembles the "dorsal adipose fin" of certain catfishes (especially sisorids and bagrids), but in few of these is the "adipose fin" as long as the

30 PETRU M. BANARESCU. TEODOR T. NALBANT adipose creast of Sphaerophysa. We do not know any genus of Ostariophysi having a ventral adipose creast. Sphaerophysa is also unique among Nemacheilinae (in the present acceptance) in having the two halves of the air- bladder capsule fused in a single, more or less spherical, piece with a pair of anterior "horns"; the posterior, free chamber of the air-bladder is rudimentary (Zhu, 1989, his Fig. 2/16). The comparison with other nemacheilines, especially with those in which the two halves of the capsule are joined, not connected by a manubrium (Physoschistura, Serninemacheilus, some Triplophysa such as T. kullmanni: Figs 16 D, 21 D, 25 B), suggests that the unique capsule of Sphaerophysa resulted through the fusion of two initially distinct pieces, representing the final stage of a gradual process of coalescence of these pieces. In Vaillantella, which is now excluded from Nemacheilinae (Nalbant and Banarescu, 1977), the air-bladder capsule is unique, too, but has quite another shape and is incomplete, consisting of two distinct lateral plates. We believe that the capsule of Vaillantella represents a primitive condition, that of Sphaerophysa a derived one. (Description after Cao and Zhu (1988) and Zhu (1989); no specimens have been available). The single described species of the genus inhabits the basin of Mekong or Lankanjiang River in Yunnan Province, China. The little-known Ellopostoma Vaillant, 1902 (type species: Aperioptus megalomycter Vaillant) is ascribed by some authors to Nemacheilinae; its position is however uncertain and Kottelat (1990) does not mention it among the members of the subfamily. Barbucca Roberts, 1989 (type species: Barbucca diabolica Roberts) may be a valid genus of nemacheilines; its illustration suggests close similarity with Schistura (or Acozlra) savona in respect of colour pattern, but the description mentions characters not met with in other members of the subfamily (occurrence of labial barbels; large breeding tubercles on the posterior part of body sides present also in females) while omitting the description of other important diagnosis characters; this is why the genus is not dealt with here. The only African member of the subfamily, "Nemacheilus" abyssinicus, known only after the holotype from lake Tama, Ethiopia (examined by the senior author of this paper) represents another genus, characterised by six branched anal rays (a character shared only with Nun and Sphaerophysa), the pelvic fins well in advance of the dorsal ( a character shared with Turcinoemacheilus; in Oreonectes,

31 NEMACHEILMAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 459 Aborichthys and Paranemacheilus the insertion of the pelvic fins only slightly in advance from that of the dorsal), distal edge of the dorsal fin convex, the caudal fin slightly notched, irregular spots on the body sides. The specimen being in a bad state of preservation, other characters could not be examined. Most of the 23 genera dealt with here (as well as Barbucca and the undescribed African genus) include only few (one to four) species; Acanthocobitis, Paracobitis, Physoschistura, Mesonoemacheilus, Oreonectes and especially Nemacheilus and Yunnanilus comprise more species (from five-six to more than 20) while Orthrias, Triplophysa and Schistura are the only large genera of the subfamily. Ln our opinion, Orthrias and Triplophysa are actually monophyletic while Schistura, in its present acceptance, is - almost surely - a provisional, polyphyletic assemblage; many of the species now ascribed to it may prove to be more closely related to species of Oreias, Parcobitis, Physoschistura etc. than to other "Schistura". The same may be true also for Nemacheilus in its present acceptance (see also Kottelat, 1990), possibly for Mesonoemacheilus and Physoschistura, too. Actually monophyletic are on the contrary Acanthocobitis, Yunnanilus (see also Kottelat and Chu, 1988 and Kottelat, 1990) and in our opinion also Oreonectes; there are marked differences between the four subgenera accepted in this paper, but the characters shared by all four suggests that they are more closely related to each other than to other nemacheilines. The monophyly of most small genera: Neonoemacheilinus, Nemachilichthys, Micronemacheilus does not raise problems. Turcinoemacheilus on the one hand, Sphaerophysa on the other hand, are the most aberrant genera of the subfamily, i.e. the most distantly related to the others. Orthrias and Triplophysa are obviously related, having the same type of sexual dimorphism; they represent a pair of sisters, the scaleless Triplophysa being the apomorphic member of the pair; they also represent the northern branch of the subfamily. Seminemacheilus differs much from them, but seems however to have a similar type of dimorphism (no sub-orbital flap, thickened pectoral ray, possibly also breeding tubercles on the sides of the head), while its range is northern, too; it may be the sister of the pair Orthrias-Triplophysa. Schistura, Oreias, Paracobitis, Homatula, Longischistura, Physoschistura, Nun and Sectoria are interrelated; they have a similar colour pattern, usually a well developed processus dentiformis, the same type of sexual dimorphism (or no sexual dimorphism at all). While Schistura is, in its presence acceptance, almost surely polyphyletic, this group of eight genera probably is monophyletic.

32 460 PETRU M. BANARESCU, TEODOR T. NALBANT Neonoemacheilus may be the apomorphic sister of Nernacheilus, being characterised by the pre-oral cavity. The type of sexual dimorphism and the colour pattern suggests that the pair Nemacheilus-Neonoemacheilus, and also Acanthocobitis and Mesonoemacheilus are closer to the Schistura than to the Orthrias group of genera. These 12 genera represent the southern branch of the subfamily, opposed to the northern branch (the Orthrias group); it is worth mentioning however that four genera of the southern branch (Oreias, Homatula, even Nun and Paracobitis) have a rather northern distribution. Micronernacheilus is very distinct; the presence of a black spot in the middle of the caudal fin base suggests some affinity with Nernacheilus, i.e. with the southern branch of the subfamily. Paranemacheilus bears a certain degree of similarity (possibly of kinship) with Micronemacheilus. Nernachilichthys has a dark spot on the middle of the caudal fin base, too, but quite another type of sexual dimorphism as Nernacheilus and Schistura. As mentioned previously, Oreonectes and Aborichthy bear much similarity; they represent a pair of sister genera. This pair, as well as Yunnanilus, differs sharply from both the northern and the southern groups of genera. It is suggested here that the main trunk of the subfamily (after the separation of Turcinoernacheilus and of Sphaeropphysa) divided into five branches: the northern (Orthrias-Triplophysa, Seminemacheilus), the southern (Schistura-Nemacheilus-Acanthocobitis) branches, the Oreonectes- Aborichthys pair, Yunnanilus and Nemachilichthys. ADDENDA The examination by the junior author of three adult females with ripe ovulae of Ellopostoma rnegalomycter (CAS , Indonesia: Borneo Island. Kapuas basin, close to Pontianak) revealed that this genus belongs to Nemacheili, very close to genera Nemacheilus (s.str.) and Tuberoschistura. Possibly Ellopostoma represents the sister lineage of both Nemacheilus and Tuberoschistura. 0 CLASIFICARE GENERICA A NEMACHEILINELOR CU DESCFUEREA A DOUA NO1 GENURI (TELEOSTEI: CYPRJNIFORMES: COBITIDAE) REZUMAT Autorii discutg pozitia nemacheilinelor, istoricul problemei genurilor subfamiliei, arata care sunt principalele caractere ce servesc la delimitarea

33 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 461 genurilor $i grupurilor de specii. Ei considera drept valide 23 genuri certe; au avut la dispozitie exemplare din 21 genuri. isi reafirma parerea anterioara ca Vaillentella reprezinta o subfamilie distincta, iar pozitia lui Ellostorne este incerta; de asemena, validitatea lui Barbucca este problematica. Aceste trei genuri nu sunt deci luate in consideratie. Se descriu doua genuri noi: Longischistura $i Seminemacheilus. in cadrul marelui gen Schistura se disting mai multe grupe de specii. se dau diagnozele genurilor.yi grupurilor de specii; se da ilustratia pentru 20 de genuri. Genul Schistura pare a nu fi monofiletic; in schimb alte doua genuri mari, Orthrias $i Triplophysa, sunt cert monofiletice. Se discuta relahile probabile dintre genuri; cele mai distincte sunt Turcinoemacheilus $i Sphaerophysa. Singura specie africana a familiei apartine cu certitudine unui gen inca nedescris. AKNOWLEDGEMENTS This contribution is based mainly on the study of a large number of specimens, belonging to tens of species of nemacheilines. received in exchange or on loan from various museums and ichthyologists, during the last three decades. The authors are grateful to the curators and the ichthyologists who facilitated the study of specimens under their care or sent specimens in exchange: MIne M.L. Bauchot. Paris; late J. Boehlke, Philadelphia; M. Boeseman, Leyden; H.D. Clausen, Kobenhavn; M. Goren, Tel-Aviv; P.H. Greenwood, London; Barbara Herzig, Wien; K.C. Jarayam, Calcutta; late P. Kiihsbauer, Wien; W. Klausewitz, Frankfurt; late C. Kosswig and late W. Ladiges, Hamburg; E. Lachner, Washington; A.G.K. Menon, Madras; M.R. Mirza. Lahore; H. Nijssen, Amsterdam; late G.V. Nikolski, Moscow; H.J. Paekpe, Berlin; F.D. Por. Jerusalem; late D.E. Rosen, New York; late A.N. Svetovidov, Sankt-Petersburg; late F.A. Turdakov, Frunze; Y.F. Wu, Xining. We are also thankful to Anca Bhwescu, who read the manuscript, improved it linguistically. made valuable remarks on some unclarified points and typed the whole manuscript. REFERENCES BANARESCU (P.), Position zoogkographique de I'ichthyofaune d'eau douce d'asie occidentale. C-vbium (3e serie). 2, 1: BANARESCU (P.), Zoogeography of Fresh Waters. I. General distribution and dispersal of freshwater animals: Aula-Verlag, Wiesbaden. BANARESCU (P.), Zoogeography of Fresh Waters. 11. Distribution and dispersal of freshwaters animals in North America and Eurasia: Aula-Verlag, Wiesbaden. BANARESCU (P.), NALBANT (T.), SiiDwasserfische der Tiirkei. 2 Teil. Cobitidae. Mitt. Hamburg Mus. Inst., 61: BANARESCU (P.). NALBANT (T.), Cobitidae (Pisces) from Afghanistan and Iran. Vidensk. Medd. Dansk. Natzrrh. Foren., 129: BANARESCU (P.), NALBANT (T.), Cobitidae (Pisces, Cypriniformes) collected by the German India Expedition. Mitt. Hamburg Mus. Inst., 65: BANARESCU (P.), NALBANT (T.), The species of Schistura (= Homatula) from the Upper Yangtze drainage (Pisces, Cobitidae). Rev. Roum. Biol., 19:

34 462 PETRU M. BANARESCU, TEODOR T. NALBANT BANARESCU (P.), NALBANT (T.), A collection of Cyprinoidei from Afghanistan and Pakistan with description of a new species of Cobitidae (Pisces, Cypriniformes). Mitt. Hamburg Zool. MUS. Inst., 72: BANARESCU (P.), NALBANT (T.), The genus Oreias Sauvage, 1974 (Pisces, Cobitidae). Nymphaea (Oradea), 4: BANARESCU (P.), NALBANT (T.), BALK (S.), SiiBwasserfische der Tiirkei, 11 Teil. die Gattung Orthrias in der Tiirkei und in Siidbulgarien. Mitt. Hamburg Zool. Mus. Inst., 75: BANARESCU (P.), NALBANT (T.), GOREN (M.), The noemacheiline loaches from Israel (Pisces: Cobitidae: Noemacheilinae). Israel J. Zool., 31: BERG (L.S.), Ryby presnykh vod S.S.S.R. i sopredelnykh stran, 2: Izd. Akad. Nauk, Moskva-Leningrad. CAO (W.-X.). ZHU (S.-Q.), A new genus and species of Nemacheilinae from Dianchi Lake, Yunnan Province in China. Acta Zootax. sinica, 13: (In Chinese, Engl. summ.). CHAUDHURl (B.L.), Description of a new species of Nemachilris from northern India. Rec. Ind. MIIS.. 5,3: CHU (X.), CHE (Y.), The fishes of Yunnan, China, 2 - Science Press, Beijing. (In Chines. Engl. summ.). DING (R.), DENG (Q.), The noemacheiline fishes of Sichuan with description of a new species. I. Paracobitis, Nemacheilsrs and Oreias. Zool. Research, 11, 4: (In Chinese, Engl. summ.). HORA (S.L.), Notes on fishes in the Indian Museum XVIII. Loaches of the genus Nemacheilus from Burma. Rec. Ind. Mus., 31: JAYARAM (K.C.), The freshwater fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka - A handbook: Zool. Survey of India, Calcutta. KOTTELAT (M.), A new noemacheiline loach from Thailand and Burma. Jap. J. Ichthyol., 29,2: KOTTELAT (M.), Noemacheilus baezingeri n.sp., a new noemacheiline loach from nothem Thailand (Osteichthyes: Cypriniformes: Cobitidae). Rev. Suisse Zool., 90: KOTTELAT (M.), Revision of the Indonesian and Malaysian loaches of the subfamily Noemacheilinae. Jap. Ichthyol., 31: KOTTELAT (M.) Cobitis Linneaeus (Osteichthyes: Cypriniformes): proposed designation of Cobitis taenia Linnaeus 1758 as type species and request for a rulling on the stem of the family-group name Cobitididae Swaison. Bull. Zool. Nom., 43: KOTTELAT (M.) Zoogeography of the fishes from Indochinese island waters with an annotated check-list. Bull. Zool. Museum Amsterdam, 12,l: KOTTELAT (M.) Indochines nemacheilines: a revision of nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and southern Vietnam: Verl. F. Pfeil, Miinchen. KOTTELAT (M.), CHU (X.-L.), Revision of Yunnanilus with description of a miniature species flock and six new species from China (Cypriniformes: Homalopteridae). Env. Biol. Fish., 23: KOTTELAT (M.), GERY (J.), Nemachei1u.s troglocataractus, a new blind cavefish from Thailand (Osteichthyes, Balitoridae). Spixiana, 11: KRUPP (F.), Ichthyogeography of the Levant. - Beihefte zum Tubinger Atlas des Vorder Orients A. (Nafunvissenchaften), 28:

35 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 463 KRUPP (F.), SCHEIDER (W.), The fishes of the Jordan River drainage basin and Azraq Oasis. Fauna of Saudi Arabia, 10: LINCK (H.F.), Versuch einer Eintheilung der Fische nach den Ztihnen. Magaz. f: das Neueste aus der Physic und Naturgeschichte, Gothe, 6,3: MENON (A.G.K.), The Fauna of India and adjacent countries, Pisces, 4, Teleostei - Cobitoidea, 1, Homalopteridae: Zoological Survey of India, Calcutta. MIRZA (M.R.), NALBANT (T.T), BANARESCU (P.M.) A review of the genus Schistura in Pakistan with description of new species and subspecies (Pisces, Cobitidae, Noemacheilinae). Bijdr. Djerk., 51.1: NALBANT (T.T), BANARESCU (P.), Vaillantellinae. a new subfamily of Cobitidae (Pisces. Cypriniformes). Zool. Meded., 52: NICHOLS (J.T.) The Freshwaters Fishes of China: The American Museum of Natural History, New York. PARIN (N.V.), Noemacheilus (7roglocobitis) starostini sp.n. (Osteichthyes, Cobitidae) a new blind fish from subterraneous waters of Kugitangtau (Turkemenia). Zool. Zh., 62: (In Russian, Engl. summ.). RENDAHL (H.), Studien uber innerasiatische Fische. Ark. Zool., 25A, 11 : RENDAHL (H.), Einige Cobitiden von Annam und Tonkin. Goteborgs K. Vetensk. Vitter. Samh. Handl., 6 3B, 3: RENDAHL (H.), Die SuBwasserfische Birmas. I. Die Familie Cobitidae. Ark. Zool., 40A, 7: RITA (S.D.), BANARESCU (P.M.), NALBANT (T.T), Oreonectes (Indoreonectes) keralensis a new subgenus and species of loach fiom Kerala, India (Pisces, Cobitidae). Trav. Mus. Hist. nut. "Gr. Antipa", 19: ROBERTS (T.R.0, The freshwater fishes of Western Borneo (Kalimantan, Barat, Indonesia) (Cyprinoidei, Cypriniformes). Mem. Fac. Fish. Hokkaido Univers., 28,2: SAWADA (Y.), Phylogeny and zoogeography of the superfamily Cobitoidea (Cyprinoidei, Cypriniformes). Mem. Fac. Fish. Hokkaido Univers., 28,2: WEBER (M.). DE BEAUFORT (L.F), The fishes of the Indo-Australian Archipelago Ostariophsyi: I-XV E.J. Brill, Leiden. ZHU (S.-Q.), Notes on the scaleless loaches (Nemachilinae, Cobitidae) from Quinghai-Xijang plateau and adjacent territories in China. Geological and ecological studies of Qignghai- Xijangplateau, 2: ZHU (S.-Q.) A new genus and species of Nemacheilinae (Pisces, Cobitidae) from China. Acta Zootax. Sinica, 8: (In Chinese, Engl. summ.). ZHU (S.-Q.), The loaches of the subfamily Nemacheilinae in China (Cypriniformes: Cobitidae): Jiangsu Science and Technology Publishing House, Nanjing, China. (In Chinese, Engl. summ.). ZHU (S.-Q.), CAO (W.X.), The noemacheline fishes fiom Guandong and Guangxi with description of a new genus and three new species (Cypriniformes: Cobitidae). Acta Zootax. Sinica, 12: (In Chinese, Engl. summ.). ZHU (S.-Q.), GUO (Q.-Z.), Description of a new genus and a new species of noemacheiline loaches fiom Yunnan Province, China (Cypriniformes: Cobitidae). Acta Zootax. Sinica, 10: (In Chinese, Engl. summ.).

36 464 PETRU M. BANARESCU, TEODOR T. NALBANT WU (HS.-W.), CHEN (Y.), CHEN (X.), CHEN (J.), The taxonomical system and phylogenetic relationship of the families of the suborder Cyprinoidea (Pisces). Scientia Sinica. 24,4: Received: May 1995 Accepted: August 1995 Institutu1 de Biologie Laboratonrl de Taxonornie Animal2 Str. Frumoasa 31 B Bucure~ti

37 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 465 Fig. 1 - Processus dentiformis in nemacheilines: A, well develop processus dentiformis (in Oreias furcotus), lateral view; B, same, ventral view; C, Moderately developed processus dentiformis, ventral view.

38 T.N.del. Fig. 2 - Types of sexual dimorphism in nemacheilines: A, suborbital flap in a male of Nemacheilus mevae; B, disposition of the breeding tubercles on the sides of the head in a male of Triplophysa sroliczkai; C, disposition of the breeding tubercles on the sides of the head in a male of Orthrias barbatulus; D. disposition of a breeding tubercles on the pectoral fin in a male of Triplophysa djaggastensis.

39 . Fig. 3 - Acanthocobitisphuketensis : A, lateral view; B, lips (ventral view of the mouth); C, intestine; D, air-bladder capsule and posterior chamber of the air-bladder; E, scale.

40 468 PETRU M. BANARESCU, TEODOR T. NALBANT

41 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 469

42 470 PETRU M. BANARESCU, TEODOR T. NALBANT T.N. del. Fig. 6 - Micronemacheilws pulcher: A, lateral view; B, lips (ventral view of the mouth); C, intestine; D, air-bladder capsule and posterior chamber of the air-bladder; E, scale.

43 NEMACHEILlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 471

44 PETRU M. BANARESCU, TEODOR T. NALBANT

45 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 473

46 474 PETRU M. BANARESCU, TEODOR T. NALBANT

47 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 475

48 476 PETRU M. BANARESCU, TEODOR T. NALBANT I

49 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 477

50 478 PETRU M. BANARESCU. TEODOR T. NALBANT

51 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 479

52 Fig Physoschis~ur-a br-unneana: A, lateral view; B, lips (ventral view of the mouth); C. intestine; D, air-bladder capsule and posterior chamber of the air-bladder; E, scale.

53 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 481

54 482 PETRU M. B,~N&SCU, TEODOR T. NALBANT Fig Mesonoemacheiluspulchellus: A, lateral view; B, lips (ventral view of the mouth); C, intestine; D, air-bladder capsule and posterior chamber of the air-bladder; E, scale.

55 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 483 Fig Nernachilichthys ruppelli: A, lateral view; B, ventral view of mouth and lips.

56 484 PETRU M. BANARESCU, TEODOR T. NALBANT

57 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 485

58 PETRU M. BANARESCU. TEODOR T. NALBANT

59 NEMACHElLlNAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 487

60 488 PETRU M. BANARESCU, TEODOR T. NALBANT w T.N. de~. Fig Shape of the intestine in seven species of Triplophysa: A, T. papillosolabiata; B, T. malloryi; C, T. rnicrophthalrna; D, T. kullrnanni; E, T. tenuicauda; F, T. djaggastaensis; G, T. dorsonotata.

61 NEMACHEILRVAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 489 T.N. Fig 25 - Shape of the air-bladder capsule and posterior chamber of the air-bladder in six species of Triplophysa: A, 7: niicrophthalma; B, 7: krrllmanni; C, 7: papillosolabiata (type "Deuterophysa"); D, T. tenuicauda (type "Tazrphysa"); E, T. malloiyi; F, 1: hutjertjuensis.

62 490 PETRU M. BANARESCU. TEODOR T. NALBANT

63 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 491

64 492 PETRU M. BANARESCIJ, TEODOR T. NALBANT T.N. de~. Fig Oreonectes (Sundoreonectes) obeszcs: A, lateral view; B, scale; C, air-bladder capsule and posterior chamber of the air-bladder.

65 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION, 2 NEW GENERA 493

66 494 PETRU M. BANARESCU, TEODOR T. NALBANT a c.d C) V1 P) u d.a 6i x 2 5" 2 La-

67 NEMACHEILINAE (TELEOSTEI) GENERICAL CLASSIFICATION. 2 NEW GENERA 495