NAT. NAT. HIST. BUL L. SIAM Soc. 46: 105-135 ， 1998

REVIEW OF THE TROPICAL ASIAN CYPRINID FISH GENUS POROPUNTIUS ，WITH DESCRIPTIONS OF NEW SPECIES AND TROPHIC MORPHS

η son R. Robertsl

ABSTRACT

Nominal Nominal species ref ，巴 rable th to 巴町opical Asian barbin cyprinid fish genus Poropuntius Smith ， 1931 (excluding Hypsibarbus Rainboth ， 1996) are reviewed ，and new species described from from the Salween ，Tenasserim ，Me k1 ong ，and Mekong basins. 百le cave fish Barbus speleops Roberts ， 1991 from Th am Phu Keao in Th ailand (Mekong basin) is referred to Poropuntius. At At least two species exhibit discrete trophic polymorphism ，including 出eriverine P. bolo νenensis new new species from streams on the Bolovens Plateau in southem Laos (Mekong basin) with four mo 中hs. Some of 出e 町ophic morphs of Poropuntius have generalized mouths and other trophic S汀uctures ，but others are so modified that that they would be assigned to different genera if they they were distinct species. Trophic polymorphism or phenotypic plasticity of the kind reported here here in Poropuntius is extremely widespread in cyprinoids and may have contributed greatly to to their speciation ，adaptive radiation ，and macroevolution ，including in Lake Lanao (Mindanao) and and Tana Lake (E 由iopia) (Mina ，et al.， 1996; Nagelkirke ，et al.， 1996) as well as over more extensive extensive geographical areas and longer time spans.

INTRODUCTION

Poropuntius Poropuntius comprises numerous m 巴dium-sized species (mostly 10-20 cm in standard length length as adu It) living mainly in mountain streams at moderate elevations (typically less than than 1，000 m) in mainland Asia ，from Assam and Yunnan to the Malay peninsula. Some lacustrine lacustrine and riverine species are significant ecologically and in artisan a1 wildcapture fisheries. fisheries. A number of the species 紅 e handsomely colored ， but they seldom if ever occur in in the aqu 紅白m trade. The majority have relatively generalized trophic s仕uctures ， with two well-developed pairs of barbels and moderately developed lips and homy jaw sheaths. 百le genus is ve 可 much in need of a comprehensive systematic revision but this is presently thwarted thwarted by paucity of material from many p訂 ts of the genus range ，especially in Assarn and and Yunnan. The present paper is based mainly on specimens collected by the author in areas areas where little or no collecting had been done previously ， but representing less than half the the range of the genus. Thus no freshly collect 巴d material has been exarnined for some of of the nominal species level taxa ，especially those described by Chinese ichthyologists. Some species exhibit substantial phenotypic plasticity in the form of discrete trophic mo 中hs ， which has not been previously reported or recognized as such in the genus. The new

Research IResearch Associate ，Smithsonian Tropical Research Institute; Department of Ic hthyology ，Califomia Academy of of Sciences ， San Francisco ，CA 94118 USA Received Received 27 July 1997; accepted 29 March 1998

105 105 106 106 TYSON R. ROBERTS

species species Poropuntius bolovenensis is described with fo 町 discrete trophic mo 中 hs having: 1) 1) norm a1 or gener a1 ized mouth ，1ヤ s and jaws (P. bolovenensis bolovenensis); 2) n町 ow head ，sh 紅 p snout ， elongate jaws and thickened lips (P. b. acuticeps); 3) short head and laminar laminar lower horny jaw sheath (P. b. glaridostoma); and 4) 1紅 ge ， broad head and wide jaws (P. b. laticeps). 官lis species and its mo 甲hs apparen t1 y is endemic to 出e Bolovens Plateau ，and may be threatened with extinction by the Xe Nam Noi ・Xe Pian hydropower project. project. h 血is paper intraspecific trophic mo 中hs 紅 'e formally described and named as species- level 住inomin a1 s. Many mo 中hologic a1 1y distinctive cypri 凶d fishes previously named as species species are really intraspecific morphs. Others may be full species ， but their status may never never be resolved because they are now extinct (e.g. the so-c a1 1ed cyprinid “species flock" 泊 Lake Lanao). Non-sexu a1 in 位'aspecific evolutionary divergence ，which may prime sympa 凶 c as well as a1 10pa 凶c speciation ， is gaining acceptance as am 司jor factor in mo 中hologic a1 dive 培ence ，speciation ，組 d generation ofbiodiversity (see WEST-EBE 周iARO， 1986; 1986; 1989; MEYER ， 1993; SMITH & SKULASON ，1996 ，and references cited 白erein). 百le sort sort of intraspecific trophic polymorphism reported here for P. bolovenensis ， in many instances instances with even greater differences among morphs ， occurs in many genera of the cyprinid cyprinid subfarnily Barbinae as we l1 as in some Schizothoracinae 佃 d Leuciscinae. The genetic genetic basis for such polymo 甲hism probably w 俗 present in the earliest cyprinoids ， or developed developed soon after 血eir divergence from other Ostariophys i. It eviden t1 y has played a major major role in their stunning adaptive radiation and dominance in temperate as well 邸 tropic a1 continen ta1 f詑sh waters ，especi al1 y in Asia. In modern faunas 仕lis ancient assortment of of genetic a1 1y-de 回 mined phenotypic differences is expr ，邸 sed 儲 in 回 specific morphs mainly 泊 species occupying lakes or streams on plateaus at higher elevations isolated by barriers such such as waterf a1 1s (as in Poropuntius bolovenensis on 出e Bolovens Plateau). Species not exhibiting exhibiting such morphs in their phenotypes may nevertheless c紅 ry al1 of the information 泊 their genome n民 essary to generate an array of discrete phenotypes. Ec ologic a1 conditions (in c1 uding biodiversity of food organisms and other fish species in 血e same habitat) may determine determine whether selection favors a phenotypically monomorphic or polymophic species. A larninar lower horny jaw shea 白 (as in P. b. glaridostoma) is one of the most 企'equen t1 y recurring 阻 d evolutionarily significant jaw 町田知res in Cyprinidae. It s presence often often involves a major dietary shift ， such as 企om feeding on insects or higher plants to feeding feeding on a1 gae 組 d associated org 白山ms which must be scraped 企om filamentous or gelatinous gelatinous mats. 百le genetic basis for shifting from an unculiferous or otherwise non- larninar larninar lower jaw sheath to a1 包凶nar shea 血 probably evolved very early in cyprinid evolution ， perhaps before divergence of Barbinae and Leuciscinae ，and perhaps evolved only only once. 百le larninar jaw shea 由 itself may be modified 泊 shape ，width ， hardness and other other characters to give rise to more 出an one in 回 specific trophic mo 中h. Di stinctive mo 中hs may evolve into full-fledged species or genera by speciation (whether a1 10pa 凶 c or sympa 凶 c) ，and by subsequent linear evolution or orthogenesis 凶 o separate genera. Monomorphic cypri 凶d genera with laminar horny jaw sheaths which may have evolved in in this fashion include but are not limited to Capoeta ，Cyprinion ，Onychostoma ， Scaphiodonichthys ，Semiplotus ， Varicorhinus. Highly specialized cyprinid genera characterized characterized by extremely broad or extremely n町 ow lower jaws typic a1 1y have laminar lower lower jaw sheaths. CYPRINID FISH GENUS POROPUNTIUS 107

The systematic portion of this paper is divided into two sections. In the first section all all of the nominal species known to me 出at are referable to Poropuntius (sensu Rainboth ， 1996a ，b) ， are listed in chronological order of their description beginning with P. deauratus (Valenciennes ，1842) ， the earliest described species. Some of these species are known only only from their original descriptions in Chinese. Most of them are probably valid species. The Th ai cavefish Barbus speleops Roberts 1991 is referred to Poropuntius. Puntius tawarensis tawarensis Weber and de Beaufort ，1916 ，is retained in Poropuntius following KoπELAT (1 989) but probably belongs elsewhere. The s巴cond section is devoted to descriptions of new speci 邸姐d of 住ophic mo 中hs. Materials.-Specimens Materials.-Specimens reported on in this paper are deposited in the fish collections of of the following institutions: ANSP ，Academy of Natural Sciences ，Philadelphia; CAS ， California California Academy of Sciences ，San Francisco; KlZ， Institute of Zo ology ， Chinese Academy of of Sciences ，Kunming; MCSNG ，Museo Civico di Storia Naturale ，Genoa; 阻沼町， Museum National National d' Histoire Naturelle ，Paris; TISTR ， Thailand Institute for Technical and Scientific Research ，B 組 gkok; USNM ， National Museum of Natural History ，Washington ， D.C.; and WIH ，Institute of Hydrobiology ， Chinese Academy of Sciences ，Wuhan.

Poropuntius Poropuntius Smith ， 1931

Poropuntius Poropuntius Smith ， 1931: 14 (勿 pe species by monotypy Poropuntius normani Smith ， 1931). 1931). Comparative Comparative osteological and other anatomical studies that might provide distinguishing characters characters for generic placement of the two dozen or more known species of Poropuntius and hundreds of other sp 巴cies of Asian barbine cyprinids remain largely undone. At present ， Poropuntius is definable mainly by characters present in many unrelated cyprinid species. species. Thus the last simple dorsal fin ray invariably is strongly serrated ，and dorsal fin soft soft rays almost invariably 紅巳 8 (or 8 11z). The main exception is the species Poropuntius bolovenensis ， in which a few specimens hav 巴 6 11z or 7 (or 7 11z). Most species have two pairs pairs of well developed barbels ，rostral and maxillary ， but some species have small maxill 訂 y barbels barbels only and one has no barbels. Mouth form tends to be generalized in species with well well developed barbels ， but at least two species have 住ophic mo 甲 hs with very different oral oral mo 中 hology. Some species with reduced barbels have reduced lower lips ， with trenchant cutting cutting edge on a broad ，homy lower jaw sheath. None of the species has a median chin barbel" “barbel" or mental lobe (mentum) like that in species of Tor ，Folifer ，and Parator. Pharyngeal Pharyngeal teeth have only been examined or reported on in a few species; these have three three rows with counts of 2，3 ，5. Such counts are characteristic of large numbers of barbin cyprinid cyprinid species. Most Poropuntius species have a longitudinal stripe extending the length of the body above above the lateralline. Usually this stripe is subdued ， and sometimes only the posterior half of of it is evident ， but in a few species the stripe is very bold. Many species have bold black or or dusky submarginal stripes on the upper and lower caudal fin lobes ， but in other species 出e 蛇 ipes are absen t. When present ， the stripe in the lower lobe tends to be bolder 白an that that in the upper lobe. The melanophores contributing to the stripes lie on the uppermost and lowermost (unbranched) principal caudal fin rays ， or on the lowermost two principal caudal caudal fin rays (i. e. the unbranched principal ray and the lowermost branched principal 108 108 TYSON R. ROBERTS

ray). ray). Juveniles 'O fm 姐 y 'O rm 'O st species have a midpeduncular dark round sp 'O t，but 由is usually usually disappe 紅 s at a small size 'O r is transf 'O rmed int 'O an 'O val 'O r 'O bl 'O ng bl 'O tch 'O ften c'O ntinu 'O us with the s凶pe 'O n the l'O wer caudal fin l 'O be. Such s凶pes ，h'O wever ， are n'O t unique unique t 'O Poropuntius ， but occ 町 als 'O in sp 配 ies currently assigned t 'O Acrossocheilus ， Barbodes and other genera. Sexual dimorphism is unkn 'O wn in m 'O st species. In a few species species males 'O r presumed males have been 'O bserved with extensive tuberculati 'O n 'O n 出 e head ，p 'O steri 'O r half 'O f body ，and 組 al fin. Poropuntius genyognathus new species has 抑制ng sexual dichr 'O matism. Poropuntius Poropuntius as rec 'O gnized here co 町'esp 'O nds essentially t 'O Poropuntius 'O f RAlN BOTH (1981) (1981) with rem 'O val 'O f 白 e species subsequently referred t 'O his new genus Hypsibarbus (RA 町 BOTH ，1996a ，b). A superficial character which may separate Poropuntius 合om ther 'O ther cypri 凶d genera is separati 'O n 'O f the r'O s住al cap fr 'O m the first infra 'O rbital 'O r lacrimal b'O ne by a deep groove. In tuberculate specimens ，白e ros 住um anteri 'O rt 'O血 is gr 'O'O ve is m 'O st heavily (sometimes exclusively) tuberculated. S'O me tuberculate specimens may als 'O have have tubercles 'O n 白 e sn 'O ut 'O r cheek posteri 'O rt 'O白 er 'O str 'O -lacrimal gro 'O ve ， but these 釘 e always always fewer in number and usually smaller than th 'O se on 血 er 'O柑 um itself. In many ther 'O ther barbin genera ，tuberculati 'O n tends to be more pr 'O n'O unced posteri 'O rt 'O血 e ros 佐um. Th e Malaysian and Ind 'O nesian species currently referred to as Puntius or Barbodes collingwoodi collingwoodi (Gunther ， 1868) bears a strong superficial resemblance t 'O generalized Poropuntius ， but lacks the rostr 'O・ lacrimal gro 'O ve 佃 d for 血 at reason may be c'O rrectly identified 出 a species 'O f Barbodes rather 由加 of Poropuntius. Th e tw 'O genera pr 'O bably are are cl 'O sely related. Poropuntius Poropuntius has a tr 'O pical and subtr 'O pical dis 凶buti 'O nm 'O stly in mainland S 'O utheast Asia. Asia. A single species occurs in 白e Ganges-Brahmapu 住a basin 'O f India. There 紅 e several several sp 即 ies in B 町 ma ，Th ailand ， s 'O uthern China ， and In do-China ，出en 'O ne 'O r tw 'O species species in the Malay peninsula and perhaps one in Sumatra. They occ 町 mainly in high gradient gradient f'O'O th i1l stre 釦ns at moderate elevati 'O ns (usually under 1，000 m). Species present in in large l'O wland rivers such as the Mek 'O ng mainstream tend t 'O occ 町 in habitats wi 血 swift current current such as rapids.

NOMINAL SPECIES OF POROPUNTIUS

Due mainly t 'O the large number 'O fn 'O minal species described 企'O m China f'O r which types types and 'O ther material are unavailable f'O r direct c'O mparis 'O n with other specimens it is imp 'O ssible t 'O do a th 'O r'O ugh systematic revision of the genus Poropuntius at this time. A number of early descripti 'O ns were based 'O n specimens with vague l'O cality data ， such as “Yunnan" ， which could have been c'O llected in the Irrawaddy ，Salween ，Mekong or Red Ri ver basins ，all 'O f which have endemic species of Poropuntius. F'O ra number 'O f the species species n 'O type material is available. Difficulties in unravelling the alpha tax 'O n'O my 'O f Poropuntius Poropuntius increased when a large number 'O f new species were described from China by Wu&L 町 (1977) without direct comparis 'O nt 'O t砿 a described earlier. M 組 Y if n'O t all 'O f 血 e species described by Wu & L 別 pr 'O bably 紅 e valid ， and all 紅 e reas 'O nably well figured ， but but general unavailability 'O f types or comp 紅 ative material makes their identificati 'O n pr 'O blematic and w 'O rk 'O n cl 'O sely related species difficul t. Th ese are listed here ，al 'O ng with CYPRINID FISH GENUS POROPUNTIUS 109

all all other known nominal species referable to Poropuntius ， in order of the date of publication.

In In some instances nominal species 紅 e also tentatively refe 町 ed to the synonymy of earlier described described species. Failure to assign a nominal species to synonymy is not an indication that that it is considered a valid species. For some nominal species about all that can be said is is the earlier the description ， and the more isolated the type locality ， the more likely that it it is a valid species.

Poropuntius Poropuntius deauratus (Valenciennes ， 1842)

Barbus deauratus Valenciennes ， in Cuvier and Valenciennes ， 1842: 188 (type locality Cochinchine") “Cochinchine") ?Poropuntius ?Poropuntius normani Smith ， 1931: 15 (type locality Pliew waterfall ，Kao Sabap ，ne 紅 Chantabun ， southeastem S即時. ?Li ssochilus smedleyi de Beaufort ， 1933: 34 (type locality Johore ， southem Malay peninsula). Poropuntius Poropuntius deauratus ，Kottelat ， 1989: 10. Material Material examined.-MNHN 2727 ， 94.5 mm ，Cochinchine ， Diard (holotype B. deauratus); deauratus); CAS 94470 ， 149 mm ，Nakom Phanom market ，8 May 1989 ，T. R. Roberts; CAS 94471 ， 7: 42.8-68.6 mm ，rapids in Mekong mainstream about 12 km S of That Phanom ，6 April 1991 ，T. R. Roberts; CAS 94472 ， 23: 40.5-80.3 mm ， rapids in lower Menam Mun at Tha Siao ， 12 km N of Phibun Mangsahan ，3 April 1991 ，T. R. Roberts; CAS 94473 ， 31: 20.7-113 mm ，Xe Nam Noi 26 km by road from Ban Nam Tang 叩 d1 km downstream from bridge over Xe Nam Noi on road down escarpment valley from top of of Bolovens Plateau to Attapeu town (elev. 270 m) ， 25 March 1995 ，T. R. Roberts; CAS 94474 ， 18: 22.8-105 mm ，Xe Nam Noi about 2 km upriver from Tat Faet waterfall ，27 ， March 1995 ，T. R. Roberts; CAS 94475 ， 69: 17.7-105 mm ，Xe Pian 1 km downstream from from Se Pa waterfall and 5-6 km upstream from Ban Hin Lat ， Sekong watershed ， southem Laos ，8 April 1995 ， T.R. Roberts; CAS 94476 ， 130: 15.5-116 mm ， small mountain stream where where it flows into Se Kaman just upstream from proposed Se Kaman 1 hydropower dam site ， 58 km by road from Attapeu town ， 14 Apri11995 ，T. R. Roberts; CAS 94477 ，2: 71. 5- 73.0 73.0 mm ，Se Kaman at Muang Saisetthirat ， 12 km E of Attapeu town ，8 April 1995 ，T. R. Roberts; Roberts; CAS 94478 ， 103 mm ，Nam Hinboun ， Feb. 1995 ，NINA; CAS 94479 ， 128 mm ， Ki nnak market ， southem Laos ， 7-15 Feb. 1994 ， T .J. W 紅 ren; CAS 94480 ，76 .3 mm ， Mekong at Ban H 叩 g Kh one ，Oc t. 1993-March 1994 ，I. G. Baird; CAS 94481 ，5: 43. 0-4 7.5 mm ，Mekong at Ban Hang Kh one ，March 1995 ，I. G. Baird; CAS 94482 ，2: 138-145 mm ， Stung Stung Treng market ， 2-18 Feb. 1994 ，T. R. Roberts; CAS 94483 ， 14: 21. 8-52.2 mm ，0 Champha ，4- 8 km upstream from its mouth into Tonle San ne 紅 Te Veng ，Ratanakiri province ，Cambodia ，1 4- 15 Feb. 1994 ， T.R. Roberts and K. E. Witte. Poropuntius Poropuntius deauratus is distinguished from all other species of Poropuntius with well well developed barbels ，組 d indeed ， from all other fish species in the Mekong basin ，by its its caudal fin coloration: bright lemon yellow ， with bold submarginal black stripes on the upper upper and lower lobes. Some individuals ，presumably male ， develop extensive tuberculation on the body ， but otherwise s民 ondary sexual dimorphism and dichromatism is unknown. Almost Almost no intraspecific variation in snout shape or mouth p訂 ts has been observed. Total gill gill rakers on first gill arch 11-13; lateralline scales 27-31; predorsal scales 9-11; and 110 110 TYSON R. ROBERTS

circumferential circumferential scale rows 22 (RAINBOTH ， 1996a: 98). The holotype has predorsal scales 10 ，transverse scale series 6. 1. 2，lateral scale series 28 ，circumpeduncular scales 14 ，and vertebrae vertebrae 24+15 = 39. This This is the species of the genus most commonly encountered in lowland p紅白 of 血e Mekong basin. It occ 町 s in the Mekong mainstream as well as in many 凶 butaries ，叩d is is strongly migratory. Large numbers formerly migrated between the Mekong mainstream and 出e Mun Ri ver in Th ailand ， but this migration has been blocked by 出e Pak Mun hydroelec 位ic d姐 1 (situated about 5 km up 出eMun Ri ver from its mouth into the Mekong). It It will be interesting to see whether the sp 巴cies survives in the Mun watershed. Although Poropuntius Poropuntius species inhabit headwaters in many places in Thailand and elsewhere where large large reservoirs have resulted from hydropower projects ， they apparently are unable to establish establish themselves in reservoirs ， and often disappear downstream from them. Examination Examination of individual specimens listed in the material examined reveals relative constancy constancy in mo 中hology of the mouth p紅 ts in P. deauratus. This should not be taken as an an indication that the species lacks inherent genetic v矧 ability the for mouth parts reported in in other species of Poropuntius ， but rather that such variabability is usually not expressed in in the phenotype. Th us P. deauratus can itself be considered as normally comprising a single single trophic mo 叩h，one with generalized mouth parts and s位ongly migratory behavior. Such a mo 中h is most likely to become widely dispersed without showing divergence of mouthparts mouthparts in the phenotypes until it establishes isolated populations in a remote basin or watershed watershed or lacustrine environment with less opportunity for dispersa l. Several other highly highly migratory riverine species of Poropuntius also appear to conserve generalized mouth p紅白. Comp 紅 ison of the 84.2 一mm holotype of P. normani Smi 白， 1931 (generic type species of of Poropuntius) with the 94.5-mm holotype and other specimens of P. deauratus reveals that 出ey are extremely similar and possibly conspecific. See also remarks under P. smedleyi. smedleyi.

Poropuntius Poropuntius clavatus (勘1c Clelland ， 1845) new combination

Barbus clavatus McClelland ， 1845: 280 ， p l. 21 ，fig. 2 (type locality “rivers at 白e foot of the the Sikkim Mountains on the northern 企ontier of Bengal"). Barbus clavatus ，Hora ， 1921: 185 ， p l. 9 ，fig. 1 (Senapati s仕eamne 紅 Kairong ，Naga Hills ， Assam). Assam). Material Material examined.-None. Original Original description evidently based on a single specimen 7 inches long. Location of type type material unknown. Hora' s description and much better figure ， presumably based on the the same species ，shows a generalized Poropuntius ， with two long barbels and apparently generalized generalized mouth parts. Lateral line scales 40-4 2，住組sverse scale rows 7-8/1 /3-4. Upper and lower caudal fin lobes with submarginal black stripes. This is the only species of of Poropuntius reported from the Ganges basin. CYPRINID Fl SH GENUS POROPUNTIUS 111

Poropuntius Poropuntius laoensis (Gunther ， 1868)

Barbus laoensis Gunther ， 1868: 115 (type locality “ Laos Mountains"). Poropuntius Poropuntius laoensis ，Kottelat ， 1989: 10. Poropuntius Poropuntius laoensis ，Rainboth ， 1996a: 99 (“ clear forested strearns of Myanm 紅， Laos ，

百 ailand ，叩dC 卸arn 巾bo 叫di 均a" 勺) Material Material exarnined.-None.

Poropuntius Poropuntius margarianus (Anderson ， 1878) new combination

Barbus margarianus An derson ， 1878: 867 ， p l. 79 ，fig. 1 (type locality “Narnpoung river ， Kakhyen hills" ， near Bharno ，Irr awaddy basin). Barbodes Barbodes (Barbodes) margarianus ，Wu and Li n， 1977: 238 ，242 (Yunn 組).

Material Material ex 創凶ned. -C AS 94247 ， 118 mm ， Mandalay market ，April1993 ，T. R. Roberts;

KIZ 781056 ・57 ，2: 128 同 157 mm ，Yinjiang ，Irr awaddy basin ，Yunnan ，Oc t. 1978.

百le 118 一mm Mandalay specimen agrees very well with Anderson's description and figure figure of P. margarianus except the s住ipe across the middle of the dorsal fin is very faint (血 is color feature may have been exaggerated by the illus 位ator). 百le spec 泊施n has gill rakers rakers on first gill arch 3+6 = 9; lateral scale series 32; predorsal scales 12; transverse scale rows rows 6/ 1/ 3; circumpeduncular scales 16. In fresh condition it was almost entirely white or or silvery ， with fins nearly colorless; dorsal and caudal fin with a little lack pigment distally; distally; middle of dorsal fin with black pigment on inteηadial membranes ， not forming the continuous continuous swath shown in Anderson' s figure of P. margarianus but in the sarne position; caudal caudal fin lobes without stripes.

Poropuntius Poropuntius hampaloides (Vinciguerra ， 1890)

Barbus hampaloides Vinciguerra ， 1890: 298 ， pl. 9 fig. 8 (type locality Meetan = a 凶 but ぽ y of of the Houngarao ， lower Salween basin). Poropuntius Poropuntius hampaloides ，Kottelat ， 1989: 10. Material Material examined.-MCSNG 17326 ， 89.3 mm ，Meetan (holotype B. hampaloides).

Observations. Observations. Snout 佐uncate or obtuse ，projecting only slightly beyon mouth. Horny jaw sheath of lower jaw broad and exposed ， lower lip broadly interrupted. Barbels absen t. Gill Gill rakers short ，3+ 01 = 13 on outer edge of frrst gill arch. Pharyngeal teeth in three rows ， 2，3 ，5. Scales in lateral series 29 ，transverse scale rows 5. 1. 2-3. Lateral 柑 ipe (not shown in in Vinciguerra's figure) visible under disecting microscope as group of melanophores on posterior posterior part of each scale in scale row above lateral line. Upper and lower caudal fin Iobes Iobes with bold marginal s凶 pes. Comparison with other species. Poropuntius hampaloides is known only from the holotype. holotype. In its specialized Iower jaw ， with wide transverse horny jaw sheath and broadly interrupted interrupted Iower lip ，it resembles wide-mouthed Poropuntius genyognathus new species from the Tenasserim basin. Direct comp 紅 ison of the holotype of P. hampaloides with an 112 112 TYSON R. ROBERTS

80.8 80.8 mm p釘 atype of P. scapanognathus new species from the middle Salween basin (MCSNG 48357) reveals different struc 伽re of the lower lip and lower homy jaw sheath. In In P. hampaloides the lower lip is res 凶cted to the comers of the mouth and the lower homy jaw shea 血 is broad; in P. scapanogna 幼us the lower lip is continuous ， and the lower jaw and lower homy jaw sheath are invariably n町 ow. Distribution. Distribution. Poropuntius hampaloides is known only from the Meetan ，a presumably high high gradient 凶 butary of the Houng 紅 ao in the lower Salween basin of southeastem Myanm 低

Poropuntius Poropuntius cogginii (Chaudhuri ， 1912) new combination

Barbus cogginii Chaudhuri ， 1912: 16 ， p l. 1 ，fig. 2 (type locality Lake Tali Fu [Er Hai or Lake Dali] ， Yunnan). ?Barbus ?Barbus gregorii Norman ， 1922: 562 (type locality Yunnan ，no other information). Barbus cogginii ，Rendahl ， 1928: 138 (Er Hai). Barbodes daliensis Wu & Li n， 1977: 251 ，fig. 7・11 (type locali 旬 Xiaguan [Er Hai]. Barbodes daliensis ，Chen ， 1998: 151 ，fig. 103. Material Material examined.-WIH 122090028 ，29:96.3-220 mm ， Xiaguan (l abelled syntypes of of B. daliensis) Diagnosis. Diagnosis. Poropuntius cogginii ， endemic to Lake Dali or Er Hai ， has long barbels and and well developed submarginal s凶pes on the caudal fin lobes. It is readily distinguished from all other Poropuntius except P. exigua by its relatively smaller fins ，p 紅白ularly 由e caudal caudal fin. Th us in a B. daliensis syntype of 188 mm SL 出e sum of the leng 血s of the dorsal ， one pectoral ， one pelvic ， the anal and the caudal fins is only 177 mm ， while in 183 and and 190 mm syntypes of P. carinatus 釦 d P. opisthoptera the total fm lengths 紅 e respectively 229 and 213 mm. Comparable relative differences in fm length and size and observed 白roughout the entire syntype series of these three nominal species. P. cogginii fu 凶 ler differs differs from P. carinatus 組 d P. opisthoptera in having a1 釘 ger head ， due mainly or entirely entirely to a greatly expanded opercular region. Discussion. Discussion. It is unclear whether Wu & LIN. 1978 intended to propose B. daliensis as as a new species ， or merely as a replacement name for Barbus gregorii Norm 叩， 1922. Since Since their account is accomp 組 ied by a full description and figure ， the specimens it is based based upon have a more definite locality (Xiaguan ， but almost certainly caught in Er Hai) ， 也e specimens 紅 e labelled syntypes in 白e WIH fish collection ，and 出e identification with B. B. gregorii is uncertain ，1 tentatively recognize B. daliensis as a validly described species in in its own righ t. As such ，it is clearly a species of Poropuntius ，佃 d is here identified for the the first time 部 a junior synonym of B. cogg 初ii. Wh ether B. cogginii is also a senior synonym of B. gregorii Norman ， 1922 requires further investigation.

The figure of a syntype of B. daliensis in Wu & L 町， 1977 (fig.7 ・ 11) ， with a scale bar of of 20 mm ，evidently represents a specimen of about 190 mm in standard leng 血， does not showp 釘 ticularly well either the smaller fms or the enlarged gill cover observed by me in all all 29 syntypes. The figure of B. daliensis in CHEN ， 1998 (fig. 103) ，however ， with a scale CYPRINID FISH GENUS POROPUNTIUS 113

bar bar of 50 mm indicating a standard leng 血 of about 210 mm ，shows both of these features very very well (it would be most helpful if 由e figure legends of type and non-type specimens figured figured in reports by Chinese ichthyologists included the standard leng 血， locality 叩 d sometimes sometimes also the catalog number).

Poropuntius Poropuntius tawarensis (Weber and de Beaufort ， 1916)

Punius Punius tawarensis Weber and de Beaufort ， 1916: 185 (type locality Lake Tawar ，Atjeh ， Suma 佐a). Poropuntius Poropuntius tawarensis Kottelat ， 1989: 42 ， p l. 14.

Material Material examined. 一-None. Poropuntius Poropuntius tawarensis ，known only from Lake Tawar in northwestem Sumatra ，is a large-scaled large-scaled species with long babels and apparently generalized mouth parts. Some individuals individuals develop a gibbosity or humeral hump ， a character seldom observed in other Poropuntius. Poropuntius. A faint longitudinal stripe above lateralline (Ko 百'E LAT ， 1989: 42). Caudal fin fin without marginal stripes. Scales in lateral series 25; 虻叩sverse scale rows 5. 1. 3; circumpeduncular circumpeduncular scales 12. It is doubtful whether 出is species is correctly placed in Poropuntius. Poropuntius.

Poropuntius Poropuntius bantamensis (Rendahl ， 1920)

Barbus bantamensis Rendahl ， 1920: 1 ，fig. 1 (type locality eastem foot of Doi Chieng Dao ， NW Siam). Poropuntius Poropuntius bantamensis ，Kottelat ， 1989: 10. Material Material examined. 一一 None.

Poropuntius Poropuntius gregorii (Norman ， 1923) new combination

Barbus gregorii Norman ， 1923: 562 (type locality Yunnan ，no fur 出er information). Pr eoccupied by Barbus gregorii Boulenger ， 1902. Barbus Barbus yunnanensis Fowler ， 1958 (unavailable substitute name for Barbus gregorii Norman ， 1923; 1923; preoccupied by Barbus yunnanensis Regan ， 1904. Barbus susanae Banister ， 1973 (substitute name for B. gregorii Norman ， 1923). Material Material examined.-None. Comment. If Barbus or Poropuntius gregorii sensu Norman ， 1923 is a valid species ， the the earliest available n創 ne is Poropuntius susanae (Banister ， 1973).

Poropuntius Poropuntius normani Smith ， 1931

Poropuntius Poropuntius normani Smith ， 1931: 15 (type locality Pliew waterfall ，Kao Sabap ， near Chantabun ， southeastem Siam). Acrossocheilus Acrossocheilus deauratus ，Smi 出， 1945: 200 (p 紅 tim). 114 114 TYSON R. ROBERTS

Material Material examined.-USNM 90297 ， 84.2 mm ， Pliew waterfall on Kao Sabap ，ne 紅 Chantabun ， southeastem Siam ，9 April 1925 ，H.M. Smi 出 (holotype). See remarks under P. deauratus and P. smedleyi.

Poropuntius Poropuntius smedleyi (de Beaufort ， 1933)

Lissochilus Lissochilus smedleyi de Beaufort ， 1933: 34 (type locality Johore ， southem Malay peninsula). Poropuntius Poropuntius smedleyi ，Kottelat ， 1989: 10.

Material Material examined. -C AS 63005 ， 8: 45. 6- 101 mm ， Sungai Tua ，Kel 佃 g drainage ，4 Feb ，1985 ，Mohd. Z 法制 a Ismail; CAS 63011 ，8: 10 4- 135 mm ， Sungai Marong ，29 Sept. 1985 ，Mohd. Zakaria Ismai l.

百us is another riverine species with phenotypically conservative and generalized mouth p紅 ts. It is probably also strongly migratory. Th e distinction between 出is species 組 dP. normani is unclear ， as is the distinction between both of these species and P. deauratus. One distinction noted between P. smedleyi and P. normani by DE BEAUFORT (1 933: 34) that that probably will not hold up is the difference in circumpeduncular scale counts.τ 'h us de de Beaufort reported 12 in P. smedleyi ，noting 白紙 P. normani has 14 according to SMITH (1931: (1931: 15). Ap 釘 tf 旨om these being counts based on single specimens ， there are problems with with the way circumpeduncular scales 紅 'e counted. Most workers report circumpeduncular scale scale counts only in even numbers ，i. e. ，12 ，14 ，16 ，indicating 血at they do not really complete complete the count all they way around caudal peduncle ， but count on one side 0 凶yand then then double the number. Thi s tends to exaggerate slight differences. An other problem is 血at it is possible to obtain different counts simply by a slight change in the starting point ， or or choosing to count from the starting point in one direction rather 出an in another. For ex 創 nple ， in the holotype of P. normani ，some scales on the caudal peduncle are missing. By counting where the scales are most complete ， one usually gets a count of 14. But coun 出19 in a slightly different way can give a count of 12. As in many other nominal species of Poropuntius ， P. smedleyi and P. normani 訂 e known mainly 仕om 血eir orig 也， al descriptions ， the authors of which compared their supposed new species only with written descriptions and not with actual specimens of other nominal t凱 a. Wh at is needed in such instances is examination of specimens from localities covering the the combined r佃，ge of the nominal forms to determine the number of species and their distribution ，組d 血en comp 組 son with the type specimens to deterr 凶ne which species names should be applied. Such ex 佃 unation is likely to be most 企ui 出 1 if examination of older older museum specimens is combined with observations of coloration and so on of freshly preserved preserved samples. In the c出 e of P. deauratus ， P. normani 佃 d P. smedleyi 出is should not not be difficult ， since there are many samples in museum holdings and their entire range is is in areas that ぽ e relatively accessible.

Poropuntius Poropuntius chondrorhynchus (Fowler ， 1934)

Barbus chondrorhynchus Fowler ， 1934: 123 ，fig. 81 (type locality Keng Tung ， Burma). Poropuntius Poropuntius chondrorhynchus ，Kottelat ， 1989: 10. CYPRINID FISH GENUS POROPUNTIUS 115

Material Material examined.-ANSP 58062 ， 185 mm ，Keng Tung market ，Southem Shan States ， Burma ， 17 Feb. 1933 ，R. M. deSchauensee (holotype); ANSP 58063 ， 156 mm ，collected with with the holotype (paratype). This This nominal species has two pairs of well-developed barbels and morphologically generalized generalized mouth p紅 ts. It is unknown whether the type specimens originated in 白e Mekong or Salween basins. Keng Tung is near the drainage divide.

Poropuntius Poropuntius kontumensis (Chevey ， 1934)

Cycocheilichthys Cycocheilichthys kontumensis Chevey ， 1934: 32 ，fig. 1 (type locality Kontum Lake near Pleiku ，central Vietnam). Poropuntius Poropuntius kontumensis ，Rainboth ， 1996a: 99 (“ clear mountain brooks and s住'eams in f白or 印es はte 吋d 紅 eas of Cambodia and Vi 記etna 凶am" 勺) Material Material examined.-CAS 94248 ， 4: 36.8-168 mm ，Vietnam ， Sai Gon river basin ， large large mountain stream tributary of Da Dung and Song Dong Nai S of Ban Ma Thuot at Cau Daktik 2 bridge 5.7 km on road from Gia Nghin to Dong Xoai ，Dac Lac province ， 7J加. 1994 ，K. E. Witte; CAS 424 ，10: 77 .3 -171 mm ，Cambodia ，Mekong basin ，0 Ch 組伊i， a small s位eam on road from Ann Long Mea to Ban Lung ，Ratanakiri province ， 12 Feb. 1994 ，T. R. Roberts and K. E. Witte. relatively A relatively generalized Poropuntius with two long pairs of barbels and usually generalized generalized mouth p紅 ts ，加 da relatively large ， thick head. Lateral line scales 35-37 ， predorsal predorsal scales 11-13 ，circumferential scale rows 24- 26; total gi1l rakers on first gi1l arch 12-16 12-16 (RA 別 BOTH ， 1996: 99). Tuberculation. Tuberculation. A 125-mm specimen from Saigon River is a presumed male with heavily heavily tuberculate rostrum ， body and anal fin. Large tubercles on snout ，rostrum 叩 d lacrima l. Dorsum of head with widely scattered very small tubercles. Tubercles on body confined confined to posterior half of body ，heaviest on scales above anal fin base ， each with 2-6 tubercles. tubercles. Scales on dorsum and sides of body with fewer and smaller tubercles. Dorsal ， pectoral pectoral and pelvic fins non-tuberculate ， caudal fin with a few minute tubercles on middle rays. rays. Anal fin with large tubercles on all rays. The Cambodian samples consists of 9 presumed presumed males ， 77.3-134 mm ，all with some anal fin tuberculation and several with some tubercles tubercles on the body ne 紅 the anal fin base ， and 171 一mm ripenning female ， with no tubercles tubercles on the body or anal fin. All of the specimens have well-developed tubercles on the the ros 住um and to a less extent on the lacrima l. The tubercles tend to be slightly larger but but no more numerous in the males than in the large female; the difference is not great.

Trophic Trophic polymorphism. All of the specimens in the Cambodian sample have similar mouth p紅 ts mo 甲hologically generalized for Poropuntius. The sample from the Sai Gon basin basin includes the four specimens listed here ， with generalized mouth parts ， and two additional additional specimens ，representing a trophic mo 中h with shovel-shaped lower jaw sheath described described below as P. kontumensis rasorius. 116 116 TYSON R. ROBER'τ3

Poropuntius Poropuntius krempfi (p el1 egrin and Chevey ， 1934) new combination

Barbus (Lissochilichthys) krempfi Pe l1 egrin and Chevey ， 1934a: 339 (type locality Nghia Lo ， Tonkin). Lissochilichthys Lissochilichthys krempfi Pe l1 egrin and Chevey ， 1934b: 467 ，fig. 2. Lissochilus Lissochilus krempfi ，Yen ， 1978: 97 ，fig. 4 1. Materi a1 examined.-None.

Poropuntius Poropuntius burtoni (Mukerji ， 1934) new combination

Barbus clavatus burtoni Mukerji ， 1934: 64 ，figs. 10- 11 (type 1∞ ality Phungkin Kh a，a tributary tributary of 由eM a1 i Kh a，Irr awaddy b部 in; see RA 町 BO 百五 1981: 37).

Materi a1 examined. 一-None.

This This nomin a1 species ，characterized by two pairs of we l1 developed b紅 bels and a short ， broad mouth ， horny jaw sheaths not described ，may be a synonym of another nomin a1 sp 田 ies described 企om the Irra waddy or other river system. It may represent an intrasp 民 ific trophic trophic morph of a species including a more generalized 位ophic mo 中h with an 紅 Tower mouth. mouth.

Poropuntius Poropuntius shanensis (Hora & Mukerji ， 1934)

Barbus shanensis Hora and Mukerji ， 1934: 362 ，fig. 3 (type locality Lawksawk ， southern Shan States ，Irr awaddy basin). Poropuntius Poropuntius shanensis ，Kottelat ， 1989: 10. Materi a1 examined. -C AS 94250 ，2: 175-176 mm ，In le Lake ，26 Feb. -4 March 1994 ， T. R. Roberts

Barbels Barbels short and 由in ，rostral barbel not extending posteriorly 邸 f釘 an anterior margin margin of eye ，maxillary barbel reaching posteriorly to below middle or posterior 213 of eye. eye. Later a1 sc a1 e series 33; pr 吋l ors a1 scales 12; transverse sc a1 e rows 6. 1. 3. Gill rakers on leading edge of first gill arch 2+1+5 = 8 (1) and 3+1+6 = 10(1).

Poropuntius Poropuntius ikedai (Harada ， 1943) new combination

Lissochilus Lissochilus ikedai Harada ， 1943: 23 ， p l. 7 ，fig. 23 (type locality Hain 阻). Acrossocheilus Acrossocheilus (Acrossocheilus) ikedai ，Wu and Li n， 1977: 292 ，fig. 7-39.

Material Material examined. ー鴨居H uncat a1 ogued ， 15: 99-193 mm ，Chang basin Jiang ，Hainan ， Feb. Feb. 1960.

Th is species ，despite its geographic a1 1y isolated distribution 企om other species of the genus ， undoubtedly belongs in Poropuntius. The origin a1 description ，住釘lslated from Japanese Japanese by Keiichi Matsuura and Atsushi Doi ，is as fo l1 ows: CYPR IN ID F IS H GENUS P O R O P UN TIU S 11 7

Fi glll 巴 1. P Oropll lllill S dea urOI Il S. M ekonga t Nakorn Pha nolll ，1 90 1ll1ll

Fi gut 巴 2. POropllllliu s 11 1([/ 苫a riallll S. Mancla lay. 11 8 111111

F igllr e 3. Poroplllll iu s kO /1 lwn ens is. Sai Go n ba sin . 125 111111 tllb 巴rCl1 lat 巴 ma le

Fig l1 re 4. P Orop lllllill Ss !l([lI ellsis . In le. 175 111111 11 8 T YSON R. R OsER TS

F igllr e 5. Poropullliu s bolove nen sis. X 巴 Na m No i，Meko ng ba sin . Upper ， P. b αcwi ce ps，J J6 mm (ho Joty p e); middJ 巴，P. b. bolovenensis ，J 24 mm (ho Jotyp 巴); Jowe r， P. b. glari dos lollla ，J 38 rnm (ho Jolyp e)

F ig llr e 6. Poro punliu s bo love nens is，ve ntr aJ vi 巴w o f h巴ad. Xe Nam No i，M eko ng bas in . L 巴ft P. b. aC Ll liceps ， J J6 rnm (ho Jo lyp 巴) ;m iddJ e， P. b. bolove nell sis，J 24 mm (ho Joty p e); ri ght P. b. glar idosloma ，J 38 mm (ho Jot y p巴) CY PR IN ID F IS H GENUS POROPU N Tl US 11 9

Fi gllr e 7. P orop lllllill S ge ll." og ll (/[l lII s. C hawa Kl oh. T enasse ri Jll ba sin . 11 2 JllJll (ho lotype) 。

Fi gllr e 8. P Oro pl ll ll ill S ge lりlog ll a llllls. Tlll 巴r K loh，T cn山間ri Jll ba sin . Na rr ow and br oad Jll Oll th 巴d grow th slag 巴S

( bOlh spec iJll ens abollt 8S Jll ITI )

Figw 巴 9. P orop unliu s ha l!t e. M 巴naJll M o巴i，Sa lw ee n ba sin ，6 1. 7 ITIITI (ho loty p e) 120 120 T YSO N R . R O Il ERT S

/ //

Fig ur e 10. P oro p l.ll1 li l.l s kO l1 lw l1 e川 is rasorills. Sai Gon bas in ‘ 87.5 111111 (ho lotype) 。

F igur e 1 . P o /口pll l1 li l.l s lII e!clI/O gra l1 ll1l1 /s . Hu ay Sa ngka lia ，M 巴klong ba sin ， 56.3 111111 (holotyp 巴).

Figurc Figurc 12. POI 口P IlI1 (/II :、¥.C {/PO l1 og l1 {/IIIIIS. Hu ay Kong. Sa l¥V cc nb a別 n， 76.2 111111 (ho lotyp c) CYPRINID FISH GENUS POROPUNTIUS 121

Dorsal Dorsal fin rays IV/8; anal fin rays: 3/5; pecωral fin rays 1/1 5-16; pelvic fin rays 1/ 8; lateral line line scales: 31-33; number of scales between dorsal fin origin and pelvic fin origin: 5/1 /3; circumference circumference scale count (around caudal peduncle) 14. Body length: head length =4 .1-4.4; body length: body depth = 3.0; predorsallength: body length = 52%; head length: caudal pe- duncle duncle depth = 1. 9-2.1; head length: head width = 1. 5- 1. 7; head length: eye diameter = 4. 0-- .1; 4.1; head length: interorbital width = 2.4 ー2.6; head length: snout length = 2. 6- 2.8; head length: postorbital postorbital leng 白=2. 4- 2.5; ph 町 ngeal teeth: 5，3， 2-2 ，3 ，5; giIl rakers: 6 + 13 (based on the specimens specimens of 150 ー177 mm body length). Body laterally compressed. Snout pointed ， covered with with homy tubercles; rostral fold of snout extending ventrally but not covering upper Ii p. Mou 白 inferior ，horseshoe shaped; lower jaw included in upper jaw when mouth closed. Upper and and lower Ii ps thin and soft ，separated from each jaw by a shallow groove; both Ii ps and chin covered covered wi 出 many pap iIl ae [=sensory pores?]. Surface of lower jaw homy. Two pairs of barbels; barbels; the barbel on 血e snout reaching below center of eye ， the other barbel originating from

comer of mouth extended posteriorly beyond posterior margin of eye (b 紅 belleng 出1. 8 times as as long as eye d.i ameter). Eye large; interorbital area distinctly convex. Dorsal fin origin slightly slightly closer ωcaudal fin base than to snout tip; last simple ray of dorsal fin hardened ，

covered covered wi 出 many se 町 'ae along the posterior margin. Pectoral fin equal to head length ，its tip not not reaching to pelvic fin. Pelvic fin origin slightly anterior to dorsal fin origin ，pelvic tip fin close close to but not reaching to anal fin origin. Ax ill 紅 y scales long and elongated. Anal fin tip not not reaching to caudal fin base. Caudal fin well developed and forked ，its length 1.5 times as

long long as head length; lower lobe longer th 釦 upper lobe. Lateral line complete ， running hori- zontally zontally but slightly lowered above pelvic fin. Pharyngeal teeth arranged in three rows ，their tips tips sharp and beak-like. Intestine long ， peritoneum black. Color in Ii fe: dorsal half dark green ，ven 仕組 half silver. Scales on side of body wi 白 semicircular black spots. AII fins dark. Ec ology: occ 町 s in rapid stre 釘ns in upper reaches of rivers. Reproductive periods probably in in December-January. The smallest specimen among specimens collected in May reaching 80 mm. Systematic notes and distribution. According to Li n (1933) ，血 is genus is relatively diversified diversified in China ，represented by 8 species. Nichols (1 927) described Barbus barbodon as a new species from Hainan Island ， but it clearly differs from L. ikedai. B. barbodon has 42 scales scales in lateralline but my examination of many specimens of L. ikedai shows 出at the largest lateralline lateralline scale count is not larger 也知 33. B. barbodon has short barbels (the barbel on 由e mouth comer is 1-1 .3 times as long as eye diameter) but L. ikedai has long barbels (the b訂 bel on on the mouth comer is 1.8 times as long as eye diameter). 百1is species was collected on1 y from from the Chang-jiang river system on Hainan Island. 百1I s species is named in honor of Navy General General Ki yoshi Ik eda. The species of Lissochilus found in Tokyo is L. krempfi and close to barbodon. L. barbodon.

Poropuntius faucis (Smith ， 1945)

Puntius faucis Smith ， 1945 (type locality Mechem Gorge ，Meping Ri ver ，Chao Phraya basin). basin). Poropuntius faucis ，Kottelat ， 1989: 10. Material Material examined.-USNM 119497 ， 40.6 mm ， Gorge of the Mechem ，tributary of the Meping Ri ver ，Chao Phraya basin ， July 1935 ， Buchanan and Harrison (holotype). Holotype Holotype has 25+15 = 40 vertebrae.

Poropuntius Poropuntius susanae (Banister ， 1973) new combination

Barbus Barbus gregorii Norman ， 1923: 562 (type locality Yunnan ，no further information). Preoccupied by Barbus gregorii Boulenger ， 1902. Barbus Barbus yunnanensis Fowler ， 1958: 12 (unavailable substitute name for Barbus gregorii 122 TYSON R. ROBERTS

Norm 如， 1923; preoccupied by Barbus yunnanensis Regan ， 1904). Barbus susanae Banister ， 1973: 143 (substitute name for B. gregorii Norman ， 1923). Barbodes (Barbodes) daliensis Wu and Li n， 1977: 251 ，fig. 7ー11 (substitute name for Barbus gregorii Norman ，1923 ， pre-empted by B. susanae Banister ， 1973). Material Material examined.-None. Comments.-Barbus yunnanensis Fowler ， 1958; Barbus susanae Banister ， 1973; and perhaps perhaps B. daliensis Wu and Li n，1977 ，were all proposed as substitute n創 nes for Barbus gregorii gregorii Norman ，1923 ， preoccupied in Barbus by B. gregorii Boulenger ， 1902. See remarks under under P. daliensis. Poropuntius Poropuntius carinatus (Wu and Li n， 1977) new combination

Barbodes Barbodes (Barbodes) shanensis carinatus Wu and Lin ， 1977: 240 ，fig. 7-3 (type locality Meng'a [Salween basin] ，Yunnan)

Material Material examined.- V，司 H 12209001 ，2: 165-183 mm ，Meng'a ，Salween basin ，Yunnan ， 1960 (syntypes).

Poropuntius Poropuntius carinatus supposedly differs 仕om other species in having a predorsal keel or or carina. It also has long barbels ，generalized mouth parts with elongate jaws and a pointed pointed head ，a curved stripe on body well above lateralline (possibly distinctive); caudal peduncle peduncle with a large black spot; submarginal stripes on upper and lower caudal fin lobes. The keel may be due to poorly fed condition of the syntypes or an artifact of preservation. A pronounced predorsal keel ， almost as pronounced ， is also present in syntypes of P. opisthoptera. opisthoptera. The syntypes of P. opisthoptera do not exhibit as むipe above the lateralline or or submarginal s凶pes on the caudal lobes ， but do have a faint midpeduncular triangular spo t. Poropuntius Poropuntius exigua (Wu & Li n， 1977) new combination

Barbodes Barbodes exigua Wu and Lin ， 1977: 249 ，fig. 7-9 (type locality Shi Cho ， flowing into northem end of Lake Dali). Barbodes Barbodes exigua ，Chen ， 1998: 150 ，fig. 102.

Material Material examined.-WIH 12209007 ， 17: 50 .4 -82.8 mm ， all from Shi Cho? ，1957 ・ 1964 (syntypes). Discussion. Discussion. Poropuntius exigua was originally distinguished from P. daliensis by Wu and Lin because of its much smaller size as sexually mature adults. This species ，apparently endemic to Lake Dali an d/ or streams flowing into it ，sh 紅白 with that species markedly smaller smaller fins (and particularly the caudal fin) 由加 any other Poropuntius. 百le specimens

紅 e also sexually mature at a remarkably small size ， smaller than any other species of Poropuntius Poropuntius seen by me in sexually mature condition with the likely exception of the very distinctive distinctive P. hathe new species from the Salween basin of Thailand. In addition ， the 12 largest largest specimens of the syntype series ， 58.5-82.8 mm ，have a more or less well developed nuchal nuchal hump ，a fature not observed in any other mainland Poropuntius including P. cogginii (of (of which P. daliensis is a junior synonym). 百le nuchal hump appears to be a normal CYPRINID CYPRINID FISH GENUS POROPUNTIUS 123

feature ， not an 紅 tifact or preservation.τ 'h e five sm a1 1est syntypes ，50.ι60.0 mm ， have no or a1 most no hump. 百le sm a1 1est of these is evidently 血e syntype figured by Wu & L 町， 1977 (fig. 7-9). In CHEN ， 1998 (fig. 102) one of 血el 紅 ger syntypes is figured. 百le sc a1 e bar of 30 mm indicates a standard length of about 110 mm ，much larger 出組曲e largest largest syntype. Be that as it may ，白 is figw 芭 shows very well the nuch a1 hump characteristic of 血is species ， reduced size of fins and especially of the caud a1 fin sh 釘 'ed only with P. cogginii ，and submargin a1 cau da1 stripes typically found in many species of Poropuntius including including P. cogginii.

Poropuntius Poropuntius opisthoptera (Wu ， 1977) new combination

Barbodes (Barbodes) opisthoptera Wu in Wu and Li n， 1977: 246 ，fig. 7-7 (type loc a1 ity Baoshan ， upper Sa1 ween basin ， Yunnan). Barbodes opisthoptera Chu and Chen ， 1989: 189 ，fig. 179.

Materi a1 ex 釘凶ned.-WIH 12209005 ， 15: 121-286 mm ， Huizen Cho ，Baoshan ，Nu Jiang Jiang basin ，Yunnan ，May 1958 (syntypes). Poropuntius Poropuntius opisthoptera supposedly differs from P. carinatus in having a somewhat sm a1 1er head and sm a1 1er eye ，佃d in lacking a stripe on body above the later a1 line and submargin a1 s凶pes on caud a1 fin lobes. 百le 286 一， mm syntype has 4+0+6=10 gill rakers on outer edge of frrst gill arch.

Poropuntius Poropuntius speleops (Roberts ， 1991) new combination

Barbus 司peleops Roberts ， 1991: 104 ，figs. 1-4 (type loc a1 ity 百lam Phu Kh ieo ，Phu Kh ieo Wildlife Wildlife Sanctuary [Mekong basin ，百 lailand]). Materi a1 examined. 一TISTR 2642 ， 57.9 mm ，Th am Phu Khi eo ，9 April 1989 ， P. Chapman (p 紅 'atype). Relationship Relationship to Poropuntius was not recognized when this species was first described. It It is the only known cave fish in the genus Poropuntius. Young fish to 60 mm have eyes nearly nearly norm a1 in position a1 though a bit sm a1 1er than in surface-dwelling species. Wi 白 grow 白白e eyes “retreat" deeper and deeper into the orbits ， which remain open but become slit-like. slit-like. Young fish have faint subm ぽ gin a1 s凶 pes on the caud a1 fin characteristic of Poropuntius Poropuntius (ROBERTS ，1991 ，fig. 1). Apart from reduction of eyes and pigmentation ， P. speleops speleops has not deviated greatly from riverine surface-dwelling species of the genus.

NEW SPECIES AND TROPHIC MORPHS OF POROPUNTIUS

During During three decades of field work on Southeast Asian freshwater fishes the author has has collected numerous samples of Poropuntius ， most of which have remained unidentified and u町 'eported until now. These include samples from a number of river basins or drainages 企om which few or no fish collections have b田 n made or reported upon previously. Am ong 血e species of Poropuntius ，sever a1 of which are undescribed ，釘e two species exhibiting exhibiting pronounced 住ophic polymorphism ，a phenomenon not previously reported in 血e genus. By 出is is me 釦 t 曲e occ 町 'ence ，wi 血in popula 討ons of a single species ， of two 124 124 TYSON R. ROBERTS

or or more discrete feeding mo 甲hs or phenotypes differing more or less strikingly 泊血e mo 中hology of the mouth ，lips ， and homy jaw sheaths. Such morphs ， with evident potential to to evolve into separate genera and sp 配 ies ， are here formally described as trinominal species-level species-level taxa.

Poropuntius Poropuntius bolovenensis new species

τ'h is species apparently is endemic to stre 創 ns on the summit of the Bolovens Plateau ， in 白e Sekong watershed of the lower Mekong basin in southem Laos. 1t is known 仕om two of the largest s住e創 ns on the plateau ， the Xe Nam Noi and the Xe Pian. It is app 紅 ently absent absent from the Houai Ho ，a somewhat smaller and more seasonal s佐'eam 泊出e southeastem comerof 由e Bolovens Plateau. The Xe Katam ，出 e only other s佐'eamon 血e plateau likely to to be inhabited by 血is species ， has not been ichthyologically s町 veyed. It is absent from the the lower Xe Nam Noi and lower Xe Pi an ， which have a relatively rich fish fauna including 血e common Mekong lowland species of Poropuntius ， P. deauratus. Poropuntius Poropuntius bolovenensis is the dominant cyprinid fish in the Xe Nam Noi and Xe Pian Pian on the Bolovens Pla 旬 au ， where the two river systems have a relatively depauperate fish fish fauna of less than 20 known species. As the main streams have been fairly well sampled sampled for fishes ， and local villages extensively interviewed about possible additional species ，it seems unlikely 白紙白is number w i1l be much increased by further exploration ， except except possibly in the Xe Katam ， where altogether different species might occ 町. Th e large series of P. bolovenensis collected at 白e proposed dam site for the Xe Nam Noi-Xe pi 組 hydropower project comprises four distinct forms or mo 中hs. 1 inte 中Irete these these forms as mo 甲hs of a species with pronounced trophic polymorphism. Most of the specimens specimens have mouth p副 s resembling those 泊 many other barbin species ，including most species species of Poropuntius with well-developed barbels ， such as P. deauratus. Th e species P. bolovenensis bolovenensis is based on this form. 百le four mo 中hs also have slight differences in body proportions proportions (head length ，body depth ， caudal peduncle length) and meristic characters (counts (counts of dorsal fin rays ，gill rakers ，佃 d scales). On the other hand ，their coloration in life life is virtually identical ， one reason for considering that they represent a single species. An other reason is 血at all of the smaller individuals have mouth p訂 ts of 白 e generalized type. 百lat is ， the smallest recognizable individuals of the 也ree divergent mo 中hs ，if 血ey are are present in 血， e sample ， are so similar to 血e most generalized morph 白紙 1 have been unable unable to distinguish them. Coloration Coloration of all individuals at the Xe Nam Noi site was nearly identical in life: a uniformly uniformly dark greenish body and ve 可 dark or dusky fins ， with almost no noticeable m 紅 ks. A 負er preservation 血em 紅 g泊als 凶peon 出， ecau 白 1fin lobes became more noticeable ， but but even then are subdued comp 紅 'ed to 血e marginal caudal lobe stripes in P. deauratus and and other species. Spec 加lens f旨om the upper Xe Pian locality had golden reflections not observed observed in other s創 nples. Specimens from the middle Xe Pi 組 locality were similar in coloration coloration to those of the Xe Nam Noi. Most of 白e Bolovens Plateau is many hundreds of meters above the surrounding lowlands. 百le Xe Nam Noi and Xe Pi 叩，出e two largest rivers on 白e plateau ，bo 白 flow into 白e Sekong watershed of southem Laos and northeastem Cambodia (the Sekong flows into into the Mekong mains 住eam at Stung Treng in Cambodia ，some 70 km south of the Lao Cyp 則 NID FISH GENUS POROPUNTIUS 125

border). 百le Upper Xe Nam Noi and Xe Pian where P. bolovenensis occurs are isolated from their lower reaches by high waterfalls. Several other species found in them also appear appear to be endemic. 百le only Poropuntius found in the lower Xe Nam Noi and lower Xe Pian is P. deauratus.

Poropuntius Poropuntius bolovenens お bolovenens お new morph

Holotype. -C AS 94251 ， 124 mm ，Xe Nam Noi 300 m downstream from prim 釘 ydam site site of Xe Nam Noi-Xe Pi 佃 hydropower scheme (elev. 730 m) ， Bolovens Plateau ，Sekong watershed ，southem Lao PDR ，24 March 1995 ，T. R. Roberts. P 鉱・ atypes. -C AS 94252 ， 110: 34.9-164 mm ，collected with the holotype; CAS 94253 ， 134 134 mm ，Xe Pian at secondary dam dam site of Xe Nam Noi ・XePi 組 hydropower scheme ， 8 km S of Ban Nam Tang ， Bolovens Plateau ，Sekong watershed ，southem Lao PSR ， 23 March 1995 ，T. R. Roberts; CAS 94254 ， 89: 22.1-114 mm ，Xe Pi 佃 about 3 km down gorge gorge from Ban Houay Chot (elev. about 400 m) ，1 April 1995 ，T. R. Roberts. Diagnosis. Diagnosis. Snout moderately elongate ， lips and homy jaw sheaths moderately developed ， margin of lower homy jaw sheath rounded with no 位enchant cutting edge. Two pairs of long barbels; rostral barbel reaching posteriorly to or beyond anterior m 紅 'gin of of eye ，maxillary barbel to or beyond posterior half of eye. Gill rakers on leading edge of first first gill arch 2-5+7-11 = 10 ー16. Frequencies of these counts in 76 specimens 10(3) ， 11(3) ，12(24) ，12(23) ，14(19) ，15(3) ， and 16(1). Pharyngeal teeth 2，3，5/5 ，3，2 (164 一.mm specimen). specimen). Dorsal fin soft rays almost invariably 8 (see comments below).

Secondary sexual dimorphism. The samples of P. bolovenensis were collected at 血e end of March 1995 ， towards the end of the 世y season. We were informed that there is often often a week of early rain during the month of April ，d町 ing which some fish spawn (e.g. Garra) ，but 出at the rain 白en stops 叩 d does not begin in e訂 nest until June ，when most fish fish spawn. Th us the specimens were not expected to be in reproductive condition. Nevertheless ， the large series of P. b. bolovenensis includes two presumed male specimens ， 88.8 釦 d 134 mm ， with pronounced tuberculation on 由 e posterior pぽ t of the body including the the anal fin. 百le last simple and frrst three branched anal fin rays each bear 3-8 small tubercles. tubercles. Most of the scales on the posterior pぽ t of the body ，particularly on the dorsum 叩 d on the lower p紅 t， each have a single tubercle near the middle of their exposed surface in in the 88.8 ・mm specimen ，and 1-3 tubercles in 白e 134 ・mm specimen.τbe entire rim of the the posterior nostrll (on both sides of the head) is covered with numerous small ，conical 町 ucωres which also seem to be tubercles ， but such tubercles are present in many other specimens. specimens. Rostral tubercles are particularly large and numerous in 血e 134 ・mm specimen ， but but not moreso 血an in some other large specimens lacking tuberculation on the body. Specimens Specimens with tuberculate bodies observed only in the nominate morph or subspecies of P. P. bolovenensis.

Poropuntius Poropuntius bolovenensis acuticeps new mo 中h

Holotype.-CAS 94255 ， 116 mm ，Xe Nam Noi 300 m downstream from main dam site site for Xe Nam Noi-Xe Pian hydropower scheme ， Bolovens Plateau ，Sekong watershed ， southem Lao Peoples Democratic Republic ，24 March 1995 ，T. R. Roberts. 126 126 TYSON R. ROBERTS

Paratype. -C AS 94256 ， 106 mm ，collected wi 出 the holotype.

Di agnosis. In 白is subspecies or morph ，represented by only two specimens ，mouth form form resembles that in the nominal morph except entire head ， snout and mouth are much narrower narrower and lips are hypertrophied. The lower lip is not continuous ， but interrupted near the the symphysis of the lower jaw. Dorsal fin with 7 branched rays. Lateral scale series 32- 33; 33; predorsal scales 13; transverse scales 7. 1. 3; circumpeduncular scales 16. Gill rakers on leading edge of first gill arch 3+1+5 = 9 (holotype) or 4+1+6 = 11 (p 紅 atype). Th e large series of P. b. bolovenensis includes a large number of smaller individuals some of which may be P. b. acuticeps. With the exception of six specimens ， 37.7-79.6 mm ， with only 7 (or ， in one specimen ，6 11z branched dorsal fin rays) all of these specimens have have 8 branched dorsal fin rays (usually 8 or 8 1ん but probably also some with 7 1/2). There is is some morphological variation in the mouth parts of these specimens but 1 could not discern discern any with clear-cut indication of acuticeps-like trophic s佐uc 加res. These specimens are are catalogued sep 紅 ately as CAS 94286 ， since they might represent the otherwise unrecognized unrecognized juveniles of P. b. αcuticeps.

Tuberculation. Tuberculation. Both specimens 紅 'e almost entirely non-tuberculate on the head as well well as on the body. Th e 106-mm p訂 atype has no tubercles at all on the rostrum ， and only a very few tiny tubercles on 血e side of the snou t. The 116-mm holotype has only a single tiny tiny tubercle on the left side of the rostrum and a few on the side of the snou t. Th e nearly complete complete absence of rostral tuberculation is striking compared to the more or less strongly tuberculate tuberculate ros 住um in P. b. bolovenensis of comparable size.

Poropuntius Poropuntius bolovenens おglaridos ωma new mo 中h

Holotype.-CAS 94257 ， 138 mm ，Xe Nam Noi 300 m downstream from main dam site site for Xe Nam Noi-Xe Pian hydropower scheme ， Bolovens Plateau ， Sekong watershed ， southern southern Lao Peoples Democratic Republic ，24 March 1995 ， T.R. Roberts. Paratypes.-CAS 94258 ， 10: 48.1-128 mm ，collected with the holotype; CAS 94259 ， 157 157 mm ，Xe Pian at secondary dam site for Xe Nam Noi-Xe Pi 組 hydropower scheme ， Bolovens Bolovens Plateau ， Sekong watβrshed ，southern Lao Pωples， Democratic Republic ， 23 March 1995 ，T. R. Roberts; CAS 94260 ，2: 76.1-83.3 mm ，Xe Pian about 3 km down gorge from Ban Huay Chot (elev. about 400 m) ，1 April 1995 ， T.R. Roberts. Di agnosis. Jaws extremely modified ，especially the lower jaw. Lower lip confined to to rictus (corner of mouth); lower horny jaw sheath greatly thickened and broad ， with a

sharp 仕組sv ぽ se cutting m 紅 gin. Upper jaw with well developed lip; upper homy jaw sheath sheath weakly developed ，continuous with upper lip ， and with very small transverse ridges. Head length 4 times in standard length. Body and caudal peduncle slender; standard length 4 times greatest depth; caudal peduncle twice as long as deep. Lateral scale series 35-39; predorsal predorsal scales 1ι17; transverse scales 7. 1. 3 -4; circumpeduncular scales 1ι16. Gill rakers rakers on leading edge of first gill arch 3-5+ 0- 1+8-12 = 13-17. Frequencies of these counts counts for 12 specimens 紅 e 13(1) ，15(6) ，16(3) ，17(2). Pharyngeal teeth 2，3，515 ，3，2 (157 ・ mm specimen). The entire series of 13 specimens of P. b. glaridostoma from the single Xe Nam Noi Cyp 町 NID FISH GENUS POROPUNTIUS 127

組 d tw 'O Xe Pi 叩 sampling sites exhibits little variati 'O n in 仕ophic s回 C旬 res and 'O ther characters. characters. Th e differences in the m 'O uth s町田知res between these specimens 佃 d P. b. bolovenensis bolovenensis is a1 m 'O st as much in the smallest specimens as in 出e larg 回 t，出 e 157 ・mm Xe Pian specimen. In the latter specimen the trenchant margin 'O f the l'O wer h'O my jaw shea 血is 10.6 mm wide. 百tis is dispr 'O p'O rti 'O nately wider 血佃 in 血e next largest specimens ， including including the 138 ・mm h'O l'O type ， in which the width 'O f the l'O wer h'O my jaw sheath is 'O nly 8.8 8.8 mm. In a1 1 but the smallest specimen the l'O w ぽ lヤ is definitely c'O nfinedωthe c'O mers 'O f the m 'O uth. In the sm a1 1est specimen ， 48.1 mm ， the lips 紅 ec 'O ntinu 'O us acr 'O ss the l'O wer h'O my jaw shea 出， but are n 'O l'O nger free fr 'O m it 阻 d have withdrawn t 'O am 'O re p'O steri 'O r p'O siti 'O n. 百le margin 'O f the l'O wer lip is indicated by pigmentati 'O n. In the next sm a1 lest specimen ， 52.7 mm ， the l'O wer lip is c 'O mpletely withdrawn fr 'O m the h'O my jaw sheath ，血 e ventr a1 surface 'O f which is theref 'O re entirely whitish. Specimens Specimens 'O f Poropuntius with a br 'O ad l'O wer h'O my jaw sheath with sharp cutting edge edge c'O mparable t'O白 at 'O f P. b. glaridostoma have n'O t been rep 'O rted by 'O ther auth 'O rs in any any species 'O f Poropuntius. It is 白 us 'O f interest t 'O n'O te 由創出ey 紅 'e rep 'O rted in 出is paper a1 s 'O in P. kontumensis rasorius new m 'O rph and P. genyognathus new species.

Secondary sexual dimorphism. N 'O t 'O bserved. Tubercles relatively sm a1 1，c'O nfined t 'O r'O s佐um in all specimens; h'O l'O type with small tubercles 'O n lacrim a1 p'O rti 'O n 'O f cheek bel 'O w n 'O s凶 ls.

Poropuntius Poropuntius bolovenens お laticeps new m 'O中 h

H 'O l'O type.-CAS 94261 ， 132 mm ，Xe Nam N 'O i 300 m d 'O wnstream fr 'O m main dam site site f'O r Xe Nam N 'O i・Xe Pian hydr 'O p'O wer scheme ，B 'O l'O vens Plateau ，Sek 'O ng watershed ， s'O uthern La 'O Pe 'O ple Dem 'O cratic Republic ，24 March 1995 ，T. R. R 'O berts.

Di agnosis. 百le single specimen representing 由is m 'O中 h is distinguished by a much br 'O ader head 仕lan any 'O f the 'O ther specimens 'O f P. bolovenensis. The m 'O uth p紅白a1 s'O are are distinctive. M 'O uth very sh 'O rt (ricturs 'O f jaws far an 旬ri 'O rt 'O vertical line at anteri 'O r m 紅 gin 'O f eye) and nearly as br 'O ad as in P. b. glaridostoma but with 'O ut the ex 住'eme reducti 'O n 'O f the l'O wer lip 'O r devel 'O pment 'O fa l 'O wer h'O rny jaw sheath with trenchant margin. margin. Upper and l'O wer h'O rny jaw sheaths well devel 'O ped; upper and l'O wer lips tightly adherent adherent t 'O h'O my jaw sheaths but c 'O mplete.

Poropuntius Poropuntius genyognathus new species

H 'O l'O type. -C AS 94485 ， 112 mm ，Chawa Kl 'O h，ups 住eam fr 'O m Ki ta 'O r Htee-tah ， Tenasserim ， 10 March 1992 ，T. R. R 'O berts.

P 訂 atypes. -C AS 94487 ， 39: 73.9-119 mm ，c 'O llected with the h'O l 'O type; CAS 94488 ， 5: 5: 40.2-58.8 mm ，rapids 'O n right bank 'O f Tenasserim Ri ver just upstream 企om Htee-tah ， 8 March 1992 ，T. R. R 'O berts; CAS 94489 ， 26: 48. 4- 101 mm ， rapids 'O n left bank 'O f Tenasserim Tenasserim mainstream just upriver fr 'O m Htee-tah ，9 March 1992 ，T. R. R 'O berts; CAS 94490 ， 9: 42. 4- 101 mm ，Kayeth 'O'O Kl'O h， 14 March 1992; CAS 94491 ， 14: 47. 4- 128 mm ， Tap 'O leh Kl'O h， 13 March 1992 ， T.R. R 'O berts; CAS 94492 ， 46: 37 .3-6 5.6 mm ， Tuler rapids 'O n Tenasserim mainstream ，ab 'O ut 2/3 'O f distance 企om Htee-tah t 'O Ba 'O washung ， 14 March 128 128 TYSON R. ROBERTS

1992 ，T. R. Roberts; CAS 94493 ， 129: 40 .3 149 mm ， Tuler Kl oh ，below Tuler rapids in Tenasserim Tenasserim mainstream ， about 2/3 of distance from Htee-tah to Baowashung ， 15 March 1992 ，T. R. Roberts; CAS 94495 ， 3: 95.3-122 mm ，Kaseh Kl oh ， 19 April 1992 ，T. R. Roberts. Roberts. This This very distinct new species is known only from the Tenasserim Ri ver basin in the southeastem southeastem part of Myanmar ， in the Indian ocean coast or westem of the upper Malay pe 凶nsula ，where it is one of the most abundant fish species. It differs from all other Poropuntius Poropuntius examined by me in having the last simple ray of the dorsal fin relatively thin (flexible (flexible distally) but st i1l strongly se 町 ated as in other Poropuntius. Only the max i1l訂 y barbel barbel is present ， and it is very small. 百1巴 lower jaw sheath is always heavily keratinized ， and and large specimens always have a very broad mouth. The species exhibits marked sexual dichromatism ， and there is considerable change in the mo 中hology of the lower jaw and homy jaw sheath with growth ， these parts starting out relatively narrow and becoming increasingly increasingly broader with age ， but at different rates depending on the individuals. Th us at at a given size it is possible to find specimens with s仕iking difference in the width of the mouth ， but all individuals st ぽ t out with n紅 row mouths and ， although some fairly large spe 岨 nens have a narrow mouth ，app 紅 'ently change to a broad or veηbroad mouth if 出ey grow to a large enough size. Th us while the feeding behavior may be quite different ，these forms 紅 e not considered to be discrete trophic mo 叩hs ，and 紅 e not formally described and named. named. Scales in lateral series 36-37; predorsal scales 10-11; transverse scale rows 7. 1. 3; circumpeduncular circumpeduncular scales 16. Coloratioo; Coloratioo; sexual dichromatism. Juveniles to about 40 mm have a well defined ， complete ，longitudinal black stripe just above the lateralline. Juveniles at somewhat larger sizes sizes tend to have no s回pe or only a faint one. In sexually ripe or ripening adults of both sexes ， but especially males ， the stripe is again evident ， but it is greatly thickened ，especially anteriorly ，and no longer confined to the body above the lateralline. Such a broadening of of the lateral s凶 pe has not been reported in other species of Poropuntius. In both sexes the the caudal fin ，especially the lower lobe ，is red or reddish; the upper lobe tends to be dusky. Neither Neither juveniles nor adults have marginal stripes on the caudal fiun lobes. A red caudal fin fin without marginal stripes is unique in the genus; the only other species observed by the author author with a red caudal fin ， P. hathe new species ， has both lobes red with well developed marginal marginal stripes. Coloration on body and fins apparently becomes intensified as fish approach approach spawning condition ， with the brightest and most intense colors in males. Broadening Broadening and darkening of the lateral stripe ， while it also occurs in sexually mature females ，is much more pronounced in males. The March samples in cI ude many mature males males and females with ripe or nearly ripe gonads ， and some males had flowing mil t. Rostral Rostral tuberculation is much more pronounced 泊 males 白血in females. Sexually dimorphic tuberculation tuberculation on body (as reported here in other species) not observed. Sexual dicromatism has has not been observed in other species ，possibly b巴cause they were not collected in spawning or or near-spawning condition.

Note 00 localities. The entire type series of P. genyognathus was collected by the author author on two trips to the Tenasserim Ri ver in southeast Burma ， one in March and one in April April of 1992. Most of the specimens were collected in the mainstream of the Tenasserim or or in the lower p紅 t of high gradient tributaries flowing into the Tenasserim mainstream CYPRINID FISH GENUS POROPUNTIUS 129

upstre 創 n from Htee-tah (sometimes spelled Ki ta) as far as the Tuler rapids (downstream from Baowashung). All of the tributaries are on the right side of the river (tributaries on the the left side ，which had been heavily logged ，were mostly dry ， while those on the right sid 巴， which had not been logged ，were flowing well). Some of the author' s notes ，including a map indicating the approximate location of each of the tributaries and the distance between between them ，were lost in the field. 百le position of the streams at increasing distances upstre 創 n from Hteet-tah follows the date of collection in March 1992. Kaseh K1 oh ， collected collected in April 1992 ，is near Mingaw (sou 白 of Htee-tah). Etymology. The name genyognathus is from the Greek ，genys ，jaw ，ax ， or ax-blade;

如 d gnathos ，jaw ， an allusion to the sharp cutting edge of the lower homy jaw sheath present present in all P. genyognathus.

Poropuntius Poropuntius hathe new species

Holotype. -C AS 94262 ，6 1. 7 mm ，rapids in Menam Moei next to highway 1035 129 km N of Mae Sot ，29 April 1991 ， T.R. Roberts. Paratypes.-CAS 94263 ， 11: 52.1-65.5 mm ，USNM 324210 ，6 1. 9 mm ，collected with the the holotype; CAS 94264 ，3: 5 1.1 -55.0 mm ， Hathe rapids in Menam Moei next to highway 1085 1085 132 km N of Mae Sot ，30 April 1991 ，T. R. Roberts. Di agnosis. Poropuntius hathe is a short ， deep bodied Poropuntius with two pairs of barbels ，gener a1 ized mouth pぽ ts ，and distinctive coloration. Standard length less than 3 times times body depth. Caud a1 fin red or orangish-red with bold black submarginal s凶 pes on upper upper and lower lobes. Gill rakers on first gill arch 3+ 1+7 = 11. Later a1 line scales 34 ， predorsal predorsal scales 14 ，transverse scale rows 6. 1. 3. Scales on posterior third of body noticeably smaller smaller th 加 those lying more anteriorly.

Etymology. The name “ hathe" is from Hathe Rapids ，one of the collecting sites of 白is species. A large and beautiful limestone formation lies in the middle of these rapids ， which extend for only a few hundred meters. Similar but less picturesque rapids occur in many other places in the Moei River.

Poropuntius Poropuntius heterolepidotus new species

Holotype.-CAS 94265 ， 138 mm ，Nam Kh ong near Nam Tok Susa ，Salween basin ， Thailand ， 19 April 1991 ， T.R. Roberts. Paratype.-CAS 94266 ， 89.9 mm ， Salween mainstream about 5 km upstream from mouth of Menam Moei ， 18-19 March 1989 ，T. R. Roberts. Diagnosis. Diagnosis. A moderately deep-bodied Poropuntius with two pairs of long barbels; sc a1 es in lateral series 39 -4 0， those on posterior half of body markedly sm a1 1er than those anteriorly; anteriorly; predorsal scales 15-17; transverse scale rows 7. 1. 3， circumpeduncular scales 17-18; 17-18; dors a1 fin origin elevated ，its origin considerably posterior to a vertic a1 line through pelvic pelvic fin origin; lower lip complete across lower jaw but tightly adherent to lower homy jaw shea 出; caudal fin with dark distal margin but caudal fin lobes without submargin a1 stnpes. stnpes. 130 130 'TY SON R. ROBERTS

A similar but less s凶king shift in sc a1 e size occ 町 s in P. hathe new species ，a1 so 合'O m the the Sa1 ween basin ，which has fewer sc a1 es ，pelvic fin origin a1 most direc t1 y below dors a1 fin fin origin ， and caud a1 fin lobes with black m 紅 gin a1 s住ipes. A similar and equ a1 1y s凶king shift shift in sc a1 e size occ 町 S 泊 some sp 回 ies of the related barbin genus Hypsibarb 附" especially 出e Irr awaddy endemic species H. myitkyinae (Pr ashad and Mukerji ， 1929) (RA 町 BO 叩， 1996a: 1996a: 86 ，fig. 23). Th ese are readily distinguished 企'O m P. heterolepidotus in having larger larger sc a1 es (therefore fewer in the later a1 scale series) and later a1 line tub u1 es with secondary secondary branches.

Etymology. Etymology. 泊施 name heterolepidotus is 仕'O m the Greek heteros ，different ，佃 d lepidotus ， scaly or sc a1 ed ， in reference to 由e abrupt change in sc a1 e size on the body.

Poropuntius Poropuntius konlumensis rasorius new morph

Holotype. -C AS 94267 ， 87.5 mm ，Vietnam ， Sai Gon river basin ，1紅 ge mountain s釘'eam 凶bu 旬ry of Da Dung and Song Dong Nai S of Ban Ma Th uot at Cau D 北 tik 2 bridge bridge 5.7 km on road from Gia Nghin to Dong Xoai ，Dac Lac province ，7 Jan. 1994 ，K. E. Witte. Witte.

P 紅 atype.-CAS 94268 ，5 1. 5 mm ，co l1 ected with the holotype.

Diagnosis. Diagnosis. Poropuntius kontumensis rasorius differs 合'O m P. kontumensis kontumensis in in having a wider lower jaw with lower lip broadly inte 町 upted rather 白血continuous ， and a shovel-shaped ，sharp-edged or laminar rather than rounded and non-larninar lower horny jaw shea 白. It differs 合'O m P. bolovenensis glaridostoma ， the onI y other Poropuntius from the the Mekong basin with a larninar lower jaw sheath ， in having a much larger head ， broader mouth ， less slender caud a1 pedun c1 e， and obvious subm 紅 gin a1 stripes on its upper 佃 d lower lower caud a1 fin lobes.

Poropuntius Poropuntius melanogrammus new species

Holotype. -C AS 94269 ， 56.3 mm ，Huay Sangk a1 ia on road to Chedi Sam Ong 7 km N of Sang k1 aburi ， 11 Feb. 1989 ， T.R. Rob 巴rt

P 紅 atypes. -C AS 94270 ， 6: 52.9-160 mm ，co l1 ected wi 由 the holotype; CAS 94271 ， 36.3 36.3 mm ，Huay M a1 ai on road to Chedi Sam Ong 21 km WNW of Sang k1 aburi ， 12 Feb. 1989 ，T .R. Roberts.

Diagnosis. Diagnosis. A moderately elongate species ，stand 紅 d length more 由加 three times greatest greatest body dep 血， distinguished from 叫1 other Poropuntius by its distinctive coloration 組 d (with possible exception of P. hampaloides) ωt a1 1ack of barbels (b oth pairs of barbels we l1 developed ， or ros 佐a1 barbels absent and we ak1 y developed maxi l1 ary barbels pr 田l ent in in a1 1 other species of Poropuntius); mouth narrow ， lower lip narrowly interrupted ne 紅 midline ， lower jaw with a very hard horny jaw sheath with s位'O ngly convex marg 泊. Poropuntius Poropuntius melanogrammus differs from a1 1 previously described species of Poropuntius ， with with possible exception of P. hampaloides ， in having a bold black s位ipe extending length of of body. This stripe ，persistent at a1 1 ages ， does not extend beyond scaled base of caudal fin. fin. Di st a1 half of dors a1 fin black; heavy concentration of melanophores on inter-radi a1 CYPRINID FISH GENUS POROPUNTIUS 131 membrane connecting last simple dors a1 fin ray with first branched dors a1 fin ray ， but a1 most no melanophores on last simple ray itself. Caud a1 fin with 出in black submarginal stripes stripes confined to upper- and lower-most simple principal caud a1 fin ray; dist a1 m 釘 'gin of caud a1 fin lobes a1 so black ，caud a1 fin otherwise yellow (出 in P. deauratus). Gill rakers 2+1+ι8 = 9(1)， 10(2) ， 11(2). Scales in lateral series 29-31; predors a1 sc a1 es 1ι11; transverse transverse sc a1 es 5，1 ，2; circumpeduncl 紅 sc a1 es 14. This This species is known only from swift ，clearwat 即位ibutaries of the Meklong basin in 官lailand. Etymology. Etymology. Th en 但 ne “melanogrammus" is 仕om the Greek “melanos" ， black or d紅 k，佃 d “grantmUS" ，line ， in reference to the bold midlater a1 stripe.

Poropuntius Poropuntius scapanognathus new species

Holotype. -C AS 94463 ， 76.2 mm ，Huay Kong just below mouth of Huay Long ，ne 紅 km 45 on highway 1126 NW of Mae Hong Son ，S a1 ween basin ，20 Feb. 1991 ， T.R. Roberts. Roberts. Paratypes. -C AS 94464 ， 2: 74.2-74.6 mm ，collected with the holotype; CAS 94465 ， 22: 22: 35.1 -6 6.2 mm ，Menam Moei at Kl erko village ， about 100 km N of Mae Sot ， 16 April 1989 ，T. R. Roberts; CAS 94466 ， 2: 52. 4-6 7 .4 mm ， Tiklo Melaah ， small stream flowing into into Menam Moei opposite Kl erko ，24 April 1989 ，T. R. Roberts; CAS 94467 ，33 .5 mm ， swift swift flowing ， sandy-botomed side channel of Menam Moei 121 km by road on highway 1085 1085 N of Mae Sot ，27 April 1991 ，T. R. Roberts; CAS 94468 ， 56: 23.0 ー75.5 mm ， and MCSNG 48357 ， 80.8 mm ，rapids in Menam Moei 129 km N ofMae Sot on highway 1085 ， 2ι30 April 1991 ，T. R. Roberts; CAS 94469 ， 10: 32 .3 -123 mm ，Menam Moei about 0.5 km upstream from confluence with Sa1 ween mainstream ， 18-19 March 1989 ，T. R. Roberts.

Poropuntius Poropuntius scapanognathus ， P. hampaloides ， and P. melanogrammus 紅 e 血e only three three known species of Poropuntius with a narrow midlateral stripe. Poropuntius scapanognathus scapanognathus differs from P. hampaloides ，合om the lower pぽ tof 血eS a1 ween basin ， in thes 加 cture of 山 lower lip and horny jaw she 拙. In P. scapanognathus lower jaw ，lip ，

組 d horny jaw sheath invariably narrow and lower lip continuous ， while in the 89.3 一， mm holotype holotype (and only known specimen) of P. hampaloides lower jaw ，lip ， and jaw shea 血 relatively relatively broader ， and lower lip restricted to corners of mouth. Coloration of P. hampaloides in in life unknown. Juvenile Juvenile P. scapanoganthus are very similar in coloration to P. melanogrammus ，a new species endemic to the Meklong basin ，esp 田 ia1 1y in having a continuous midlater a1 black black s住ipe. However ，this s佐ipe remains bold 由roughout life in P. melanogrammus ， whereas whereas in P. scapanognathus it is less bold in juveniles of 70 mm and absent in specimens over over 110 mm. At a1 1 sizes in P. melanogrammus the anterior and posterior portions of the longi 旬 din a1 s回pe 紅 e of nearly equ a1 pigmented ， while in P. scapanognathus 血 e anterior portion portion is more weakly pigmented. As the stripe disappears with growth in P. scapanognathus ，也 e posterior portion is the last to disappe 紅. In P. scapanognathus 白e leading leading edge of the dors a1 fin is darkened by melanophores concentrated along the last simple simple fin ray ， while in P. melanogrammus the distal margin of the dors a1 fm is darkened. 132 TYSON R. ROBERTS

Etymology. The name “ scapanognathus" is from the Greek ，“ skapane" ，a spade 'O r h'O e; and “gnathus" ， jaws.

DISCUSSION

Di screte tr 'O phic p'O lym 'O rphism like that f'O und in Poropuntius bolovenensis is much c'O mm 'O ner in cyprinids and 'O ther fishes than w 'O uld appear fr 'O m rep 'O rts in the scientific literature. literature. M 'O st ichthy 'O l 'O gists are unfamiliar with the phen 'O men 'O n 'O fn 'O n-sexual p'O lym 'O rphism and have n'O t been l'O'O king f'O ri t. Als 'O， much previ 'O us systematic w 'O rk has been based 'O ns 創 nples with relatively few individuals that lent themselves m 'O re readily t 'O inte. 中 retati 'O n as different species rather than as different f'O rms 'O f the same species. Wh en large samples were available ，出ey were n'O t adequately studied f'O r individual variati 'O n

'O fm 'O uth p釘 ts. Well d'O cumented examples 'O f tr 'O phic p'O lym 'O中 hism 紅 e available f'O r the ne 'O位 opical characin Saccodon wagneri (ROBERTS ， 1973) and f'O r the cichlid Cichlasoma minckleyi minckleyi endemic t 'O Cuatr 'O Cienagas in n'O rthem Mexic 'O (SAGE & SELANDER ， 1975; KORNFIELD & TA YLOR ， 1983). Experimental Experimental evidence that the f'O ur kinds 'O f Poropuntius f'O und in the Xe Nam N 'O i 'O n the B 'O l'O vens Plateau in s'O uthem La 'O s are intraspecific tr 'O phic m 'O中 hs 'O r subspecies 'O fa single species is n'O t available. N 'O direct 'O bservati 'O ns have been made 'O n 'O ne m 'O rph changing changing int 'O an 'O ther ，'O r 'O fm 'O rphs devel 'O ping from parents 'O fa different m 'O rph. The evidence 白紙 the variants 'O f P. bolovenensis 紅 'e in 住aspecific mo 中hs includes 1) their restricted restricted 'O ccurence in a limited ge 'O graphical area (i. e. is 'O lated streams 'O n the B 'O l'O vens Plateau); Plateau); 2) their simil 訂 ity t 'O intraspecific m 'O rphs in 'O ther species 'O f Poropuntius and 'O ther barbin genera; 3) their great similarity in c'O l'O rati 'O n and 'O ther characters t 'O each o出 er; 4) their distinctive features bec 'O me m 'O re and m 'O re pr 'O n'O unced with age but small specimens 紅 'e indistinguishable (as with smaller males and female individuals 'O fm 'O st sexually sexually dim 'O rphic species); and 5) their app 紅 'ent 'O cc 町 as 'ence p制'O fa single p'O pulati 'O n， with with all 'O f 白e mo 甲hs present in a single sample. In 住aspecific phen 'O typic variati 'O n 'O fm 'O uth pぽ ts has been d'O cumented in the S 'O uth Mrican cyprinid Barbus brucii by GROENEWALD (1958). Th is sp 町 ies has tr 'O phic m 'O中 hs with with greatly thickened lips and a mentall 'O be (a feature n'O t 'O bserved in Poropuntius); with m 'O derate lip devel 'O pment and n 'O mental l'O be; and with sect 'O riall 'O wer jaws. 官le greatly thickened thickened lips and mental l'O be 'O f wild-caught specimens regress when 出 ey are c'O nfined in in cement t創 lks. Tr 'O phic p'O lym 'O rphism is a significant mechanism f'O r phen 'O typic diversificati 'O n 'O f cyprinidae cyprinidae under certain circumstances ，especially when a few species 'O r individuals gain access access t 'O a new 'O r is 'O lated habitat ， such as am 'O untain tributary ab 'O ve a waterfall 'O ra newly f'O rmed crater lake. It may als 'O play ar 'O le in seemingly sudden appe 紅加ce 'O f “ev 'O luti 'O nary n 'O velties". Thus several distinctive cyprinid genera-Cyprinion ， Onychostoma ，Scaphestes ，Scaphiodon ，Scaphiodonichthys ，and Scaphognathops in Asia ， Varicorhinus Varicorhinus in Africa ，Acrocheilus in N 'O rth Am erica-are characterized by sh 'O vel-like 'O r 'O therwise m 'O dified l'O wer h'O my jaw sheaths simil 紅 t'O th 'O se that sh 'O w up in 'O ther cyprinid genera genera (e.g. Poropuntius) as intraspecific m 'O rphs. The rapid ev 'O luti 'O n 'O f spectacular endemic “species fl 'O cks" such as th 'O se 'O f the endemic cyprinids 'O f Lake Lana 'O and endemic hapl 'O chr 'O min cichlids in Lake Vict 'O ria may be based largely 'O n elab 'O rati 'O n 'O f CYPRINID CYPRINID FISH GENUS POROPUNTIUS 133 the the sort of intraspecific genetic and phenotypic variability documented earlier in Barbus brucii ，Saccodon wagneri ， Cichlasoma minckleyi ， and now in Poropuntius bolovenensis. A similar conclusion has been suggested for the Victoria cichlids by MEYER (1 993). Modification Modification of the pharyngeal jaws and teeth of cyprinoids undoubtedly has played a key role in their adaptive radiation ， but the exact nature of this role needs elucidation. In In large segments of cyprinids with varied feeding habits ， the mo 中hology of the ph 紅 yngeal jaws jaws and teeth changes remarkably little. Thus in the subfamily Barbinae ，species after species species has the standardized or fixed count of three rows of pharyngeal teeth with 2，3，5 teeth ，and the mo 中hology of these teeth is ，on the whole ， remarkably conservative. Th e modified modified ph 訂 yngeal jaws and teeth of cyprinoids are no doubt linked to the divergence of cyprinoids cyprinoids from their jaw-toothed ancestors and the loss of jaw teeth. Cyprinoids obviously have have not lost the genetic information involved in tooth development ， since all or almost all all of them still develop te 伯 on 恥 pharyngeal jaws. Absence of tee 出 on cyprinoid jaws is is directly linked ω 白ev 紅 iety of keratinized stuctures that develop on the jaws ， and the evolutionary evolutionary success of these s佐uctures in generating diverse oral feeding mechanisms ， including including unculiferous ridges ，grooves ，papillae ， and fimbriae on the lips and horny jaw sheaths sheaths and extra-oral structures such as the rostral cap and mental disc (MlN ZENMA Y， 1933; 1933; ROBERTS ， 1982). Keratinous formations such as unculiferous horny jaw sheaths which 紅 'e formed ， sloughed off and replaced very rapidly may generate variation for natural natural selection as effectively as multicuspid teeth ， and may lend themselves even more readily readily to trophic polymo 叩hism.

ACKNOWLEDGMENTS

Financial Financial suppo 此 for the author' s fieldwork in Asia has been provided by the Smithsonian Smithsonian Institution; Smithsonian Tropical Research Institute; Institute for Biological Exploration; Exploration; and Committee for Research and Exploration of the National Geographic Society Society (grant 5141-93). Permission for fieldwork in Th ailand was gr 組 ted by 白e Foreign Researcher Researcher Section of the Thai National Research Counci l. Permission for fieldwork in the the Peoples Democratic Republic of Laos was granted by the Lao Department of Forestry. Fieldwork Fieldwork in Cambodia was done in cooperation with FAO and the Cambodian Department of of Fisheries. A visit to examine specimens in the Museo Civico di Storia Naturale was greatly greatly facilitated by Guliano Doria ， Enrico Borgo ， and Mauro Brunetti. My visits to Chinese Chinese ichthyological collections in April-J une 1998 were coordinated by Zhang Chunguang of the Institute of Zoology ， Chinese Academy of Sciences ，Beijing; and He Shunping Shunping of the Institute of Hydr 油 iology ， Chinese Academy of Sciences ，Wuhan. Ho Shunping Shunping and Shan Xianghong greatly facilitated examination of specimens of Poropuntius in in Wuhan ， as did Y 姐 g Junxing and Cui Guihua in the Institute of Zoology ， Chinese Academy of Sciences ，Kunming. Th 創 lks 訂 e also due to J.-C. Hureau of MNI 王N ， B il1 Saul of of ANSP ， and Susan Jewett of USNM for sending specimens on loan ，and to Keiichi Matsuura Matsuura and Atsushi Doi for sending a photocopy of Harada (1 943) and translating the description description of Lissochilus ikeda i. Th e study was completed in the Department of Ich 血yology of of the California Academy of Sciences and in the Center for Conservation Biology ， Mahidol University ，B 叩 gkok (Salaya Campus). 134 134 TYSON R. ROBERTS

REFERENCES

ANDERSON ，J. 1878. Anatomical and 2o ological Researches ， Comprising an Account of the 2o ological Results of of the Two Expeditions to 】'unnan in 1868 and 1875. London ，2 vols BANIS 百 R， K. 1973. A Revision of 恥 large Barbus (Pi sces ，Cyprinidae) of east and central Af rica. Br. Bull. Mus. Mus. Nat. Hist. (zoo l.) 26 (1): 3-148. CHAUDHURI ，B. L. 1912. Contributions to 自己 fauna of Yunnan based on collections made by J. Coggin Brown ， B.Sc. ， 1909-1910. Part ll.-Fishes. Rec. lndian Mus. 6: 13-24 ， p l. 1. CHEN ， Y. 1998. 百le fishes of 血e Hengduan Mountains region. Science Press (B eijing) ，364 pp. CHEVEY ， P. 1934. Description de deux Cyprinidae nouveaux du lac de Kontum (Ann am). Bull. Mus. Nat. Hist. Nat. ， Paris (ser. 2) 6: 32-35 [published before 22 May 1934]. CHU ，X.L. ，AND Y. R. CHEN. 1989. The Fishes of Yunnan ， China. Part 1. Cyprinidae. Beijing. viii + 377 pp. CUVIER ，G. ，AND A. V 札 ENCIENNES. 1842. Histoire Naturelle des Poissons. Paris. 16. DE BEAUFORT ，L. F. 1933. On some new or r紅巳 species of Ostariophysi from the Malay peninsula and a new species species of Betta from Borneo. Bull. Raffles Mus. 8: 31-36. FOWLER ，H.W. 1934. Zoological results of 出E 白ird deSchauensee Siam 巴se expedition. Part I-Fishes. Proc. Acad. Acad. Nat. Sci. Philadelphia 85: 233-267. [date of publication 30 April 1934]. FOWLER ，H.W. 1958. Some new taxonomic names of fishlike vertebrates. Not. Nat. ，Philadelphia ， no. 310 ， 16 16 pp. GROENEWALD ，A. A. V .J. 1958. A revision of 血巴 genera Barbus and Varicorhinus (Pisces: Cyprinidae) in Trans- vaa l. Ann. Transvaal Mus. 23(3): 263-330 ， pls 59-70. GUNTHER ，A .L. 1868. Catalogue ofthe Fishes in the British Museum. 7. British Museum ，London xx + 512 pp. HARADA ，I. 1943. Freshwater Fishes of Hainan lsland [in Japanese]. Int ，巴 lligence Agency of 白e Japanese Navy in in Hainan ， 113 pp. ，23 pls. ，map. HORA ，S. L. 192 1. Fish and fisheries of Manipur with some observations 0 目白ose of the Naga H i1l s. Rec. lnd. Mus. Mus. 22(3): 165-124 ，pls. 9-12. HORA ，S. L. AND D.D. MUKERJ I. 1934. Notes on fishes in 出e Indian Museum. XXIII. On a collection of fish from the S. Shan states ，Burma. Rec. lndian Mus. 35(3): 353-370. KORNFIELD ，AND J.N. TAYLOR. 1983. A new species of polymorphic fish ， Cichlasoma minckleyi ，from Cuatro Ci 釦 agas ，Mexico (Teleostei ，Cic hI idae). Proc. Biol. Soc. Washington 96(2): 253-269. KoπELAT ，M. 1989. Zoogeography of 出e fishes from Indochinese inland waters with an annotated check-list ， Bull. Bull. Zoo l. Mus. ， Univ. Amsterdam ，12(1): 1-56. MEYER ， A. 1993. Trophic polymorphisms in cichlid fish: D 白血ey represent intermediate steps during sympa 凶c

speciation speciation and explain their rapid sp 巴ciation? pages 257-266 in Trends in /c hthyology ， J.H. Schroder ，J. Bauer ，M. Schartl (eds) ，G.S.F.-Bericht: Blackwell.

MINA ，M.V. ， N. MIRONOVSKY ，AND Y.Y. DGEBUADZE. 1996. Lake Tana large b紅 bs: phenetics ，growth ， and diversification. diversification. J. Fish Biol. 48: 383 -4 04. MCCLELLAND ，J. 1845. Description of fo 町 new species of fishes from rivers at the foot of the Boutan Mountains. 1. 1. Nat. Hist. ， Calcutta 5(1 8): 27 4- 282 ， p l. 2. MINZENMAY ， A. 1933. Die Mundregion der Cypriniden. 2001. lahrb. ， Anat. 57: 191-286. Mu 阻則1， D.D. 1934. Report on Burmese fishes colle 氾ted by Lt. Co l. R.W. Burton from the tributary streams of 也e Mali Kh a Ri ver of the Myitkyina di 町 ict (Upper Burma). Part 2. よ Bombay Nat. Hist.Soc. 37: 38- 80. 80. NAG 乱 KIRKE ，L.A. G. ，M.V. MINA ， T. WUDNEH ， F.A. SIBBING ，AND J.W.N. OSSE. 1996. In Lake Tana ，a unique fish fish fauna needs protection. Bioscience 45 (1 1): 772-775.

NOI 削 AN ， J.R. 1923. Thr ee new fishes from Yunnan ，collected by Pr ofessor J.W. Gregory. Ann. Mag. Nat. Hist. ， se れ 9， 11: 561-563. PELLEGRIN ，J. ，AND P. CHEVEY. 1934a. Poissons de Nghia-Lo (Tonkin) descriptions de qua 佐e esp とces nouvelles. Bull. Bull. Soc. 200 1. France 59: 337-343. PELLEGRIN ，J. ，AND P. CHEVEY. 1934b. Addendum a une note precedente sur les poisson de Nghia-Lo (Tonkin). Description Description de quatre esp 批 s nouvelles. Bull. Soc. 2001. France ， 59: 467 -4 68. RAINBOTH ，W .J. 198 1. Systematics of the Asian barbins. PhD dissertation ，University of Michigan ，Ann Arbor. 253 pp. RA 削 BO 澗， W .J. 1996a. Fishes of the Cambodian Mekong. FAO ，Rome. xii + 265 ，27 pls. CYPR 町 ID FISH GENUS POROPUNTIUS 135

RAINBOTH ，W .J. 1996b. 百le taxonomy ，systematics ，and zoogeography of Hypsibarbus ，a new genus of large barbs barbs (Pisces ， Cyprinidae) from the rivers of southeastem Asia. Univ. California Publ. Zo ol. 129 ， xiv + 199 199 pp. RENDAHL ， H. 1920. Eine neue Barbus- Art aus Siam. Ark. Zo ol. 12 (1 6): 1-3. RENDAHL ， H. 1928. Beitrage zur Kenntnis der Chinesische Susswasser Fische. 1. Systematischer Tei l. Ark. Zo ol. Stockholm 20A(I): 1-94.

ROBERTS ，T. R. 1973. Dental polymorphism 加 d systematics in Saccodon ， a neotropical genus of freshwater fishes

(P 紅 'odontidae ，Characoidei). J. Zo ol. Lo ndon 173: 303-32 1. ROBERTS ， T.R. 1982. Unculi (homy projections arising from single cells) ， an adaptive feature of the epidermis of of osta 巾 physan fishes. Zo ol. Sc 中 ta 11: 55-76. ROBERTS ，T. R. 199 1. Barbus speleops new species ，a blind cavefish from Th am Phu Kh ieo ，Mekong basin ， Th ailand. Nat. Bull. Hist. Siam Soc. 39: 103-109. SAGE ，R. D. AND R. K. SELANDER. 1975. Trophic radiation through trophic polymorphism in fishes. Proc. Nat. Acad. Acad. Sci. 72: 4669 -4 673.

SMI 叩， H.M. 193 1. Descriptions of new genera 如 d species of Siamese fishes. Proc. Uぷ Natl. Mus. 79: 1-4 8， l. pl. 1.

SMITH ，H.M. 1945. The fresh-water fishes of Siam ， or Th ailand 目 Bull. U.S. Nat l. Mus. 188 ， xii + 622 pp. SMITH ，T. B.， AND S. SKULASON. 1996. Evolutionary significance of resource polymorphisms in fishes ，amphib- ians ，and birds. Annu. Rev. Eco l. Syst. 27: 111-133. TCHANG ，T.- L. 1938. Notes on a new Barbus from Yunnan. Bull. Fan Mem. lnst. 7(2): 63 -6 5. VINCIGUERRA ， D. 1890. Viaggio di Leonardo Fea in Birmania e regione vicine. XXIV. Pesc i. Ann. Mus. Civ. Sι Nat.. Nat.. Genova. 129-362 ，pls. 7-1 1. WEST-EBERHARD ，M .J. 1986. Altemative adaptations ，speciation ，and phylogeny (a review). Proc. Nat l. Acad. Sci. Sci. USA 83: 1388-1392. W 邸 T-EBE 脳 IARD ，M .J. 1989. Phenotypic plasticity and the origins of diversity. Annu. Rev. Eco l. Syst. 20: 249- 278. 278. WU ，H .w.， ET AL. 1977. The cyprinid fishes of China [in Chinese]. Shanghai. 2: 229-598.

YEN ，M.D. 1978 [ldentification of the fresh-water fishes of North Vietnam] (in Vietnamese). H 組 oi ， 1-340.