NAT. HIST. BUL L. SIAM Soc. 45: 171-203 ， 1997

SYSTEMA TIC REVISION OF THE TROPICAL ASIAN LABEOIN CYP 悶 NID FISH GENUS CIRRHINUS ，WITH DESCRIPTIONS OF NEW SPECIES AND BIOLOGICAL OBSERV ATIONS ON C. C. LOBATUS

乃son R. Roberts*

ABSTRACT

Cirrhinus 訂 e lab ∞ in cyprinids of mainland 甘opic a1 Asia ， ranging from the In dus to southern southern China and the Mekong and the Tapi dr 出nage in peninsul 紅百四iland. Fourteen

species 紅 e recog 凶zed in 白e present revision. lndia has five species ，Myanm 訂 four (including

one endemic to Lake ln1 e) ，Th ail 加 d eight ，Cambodia ，Laos and Vietnam each have seven ，佃 d China only one. 百lree species are described as new: C. rubirostris frorn Tenasserim bωin ，

southeast 巴rn Myanrn 紅; C. inomatus from lrrawaddy and Si 仕組g basins; 組 d C. omatipinnis frorn frorn the middle Mekong basin of Thailand. Th e replacement name C. lu is proposed for the endemic Inle Lake species C. horai (Banarescu ，1986) ， preoccupied in Cirrhinus by C. horai Lakshamanan ，1966 (a subjective junior synonym of C. cirrhosus). Cyprinus cirrhosus Bloch ， 1795 1795 is identified as the most senior synonym of Cyprinus mrigala Hamilton ， 1822; Cyprinus ariza ariza Buch 組 an ， 1807 as 血e most senior synonym of Cirrhinus reba (Har 凶Iton ，1822); 飢 d Le uciscus molitorella V a1 enciennes ， 1844 as 曲e rnost se 凶or synonym of Cirrhina chinensis Gun 由民 1868.τbus the va1 id names for 白ese species are Cirrhinus cirrhosus ， C. ariza ，and C. C. molitorella. Cirrhinus jullieni is a relatively rare species ，a1rn ost inv 副 ably confused with C. C. siamensis by previous authors ， with a restricted dis 住ibution in the lower Mekong and Chao P恥aya (a1 1 previous records from Chao Phraya are based on C. siamensis or 0 血er species). Cirrhinus Cirrhinus cirrhosus is one of the most important cultured fish species in lndia ，B 加 gla- desh desh and Myanm 民 and C. molitorella is cultured widely in China. Both have been introduced

into into other coun 凶 es. Sever a1 other species 紅 'e impo 口組t in wild-capt 町 e fisheries ， including C. C. lobatus ， C. microlepis ，and C. siamensis in 出e Mekong basin. Most species of Cirrhinus probably probably have direct sexu a1 development ， with sex ratios approaching 1. 1: Cirrhinus lobatus ， however ， is a sequenti a1 (protogynous) hermaphrodite: all migrating individu a1 s develop fe- m a1 e gonads begin 凶ng at about 45 mm standard leng 出; at about 65 mm ，some individu a1 s develop develop m a1 e gonads. It is suggested 血at C. lobatus migra 也事 ups 佐eam for reproduction employ as 回 tegy of “ever-changing leadership" to find their way past physic a1 barriers such as as Kh one waterfalls in southern Laos. Wh at appears to be a massive long distance migration of of very large numbers of 血is species may represent numerous episodes of recruitrnent and falling falling out en route ，so 血at individual fish (especially the sm a1 ler and move numerous fem a1 es) travel travel only a fraction of 白e entire migratory rou 旬百lU S the so-called “ long distance" migra-

tions tions of C. lobatus 如 d some other cyprinid species may differ fundantally from the true long distance distance migrations of many birds ，mammals ，and 0 血er fishes.

* Research Associate ， Smithsonian Tropic a1 Research Institute Received Received 24 July 1997; accepted 20 October 1997.

171 171 172 172 TYSON R. ROBERTS

INTRODUCTION

The tropical Asian labeoin cyprinid genus Cirrhinus includes species of m 吋or importance importance in aquaculωre ， wild cap 阻 re fisheries ， and ecology. Th e most important cultured species 紅 e C. cirrhosus (mrigal) and C. molitorella ， both consequently transported beyond their their natural r佃 ges. Several other species are important objects of fisheries an d/ or are ecologically ecologically dominant species. Cirrhinus microlepis is 加 important species in wild capture fisheries fisheries of the middle and lower Mekong basin ，including the Great Lake-Tonle Sap system system in central Cambodia. Two of the smaller species ， C. siamensis and C. lobatus ， are perhaps perhaps the most abundant of all naturally occu 町 ing fish species in the Mekong basin. They are heavily fished during reproductive migrations occurring mainly in May-July and non-reproductive non-reproductive migrations occurring mainly in December-January (ROBERTS ， 1993; ROBERTS & BAIRD ， 1996) Despite Despite their importance in fisheries and ecology ， the systematics and biology of most Cirrhinus Cirrhinus species have not been extensively investigated. A systematic study of Cirrhinus species species of Th ailand by Sodsuk ， 1988 is deficient because the author did not study type specimens specimens of any of the described species ， and hence the species are not well identified. Th e specimens that study was based upon are all deposited in the National Inland Fisheries

Institute Institute in Bangko k. As these collections 紅 e not generally available for examination by researchers ，it has not been possible to re-exarnine or re-identify the specimens. A general review of the biology ，now somewhat out of date ，is available only for C. cirrhosus cirrhosus (JHINGRAN & KHAN ， 1979). Far less is known of other important species such as as C. microlepis. Migrations of Mekong species 訂 e just beginning to be documented (ROBERTS ， 1993; ROBERTS & BAIRD ， 1996). Samples from spawning migrations of C. lobatus lobatus indicate that it is a precocious protogynous hermaphrodite (ROBERTS & BAIRD ，

1996: 1996: 247-248 ，figs. 22-24). It also is an ecological keystone sp 配 ies ， and probably 出e most abundant fish species in the Mekong basin. This This paper is concemed mainly with alpha level systematics of species in the genus

Cirrhinus. Cirrhinus. Henicorhynchus Smi 由， 1945 ，reg 紅 ded as a distinct genus by BANARESCU ， 1972; 1972; 1983 and KoπELAT ， 1989 (but in any event a subjective junior synonym of Cirrhinichthys Cirrhinichthys BLEEKER ， 1863) is treated here as a subjective junior synonym of Cirrhinus.

民1ATERIALS AND METHODS

τ'h e specimens repo 民ed on are deposited in the following institutions: M 心ffi ，Am erican Museum ofNatural Hi story ，New York; ANSP ，Academy ofNatural Sciences ，Philadelphia; BMNH ，British Museum ofNatural Hi story ， London; CAS ，California Academy of Sciences ， San Francisco; MNHN ，Museum National d'Histoire Naturelle ，Paris; MSNG ，Museo Civico d'Storia d'Storia Naturale ， Genoa; USNM ，National Museum ofNatural History ，Washington ，DC; ZMA ， Zoologische Museum ，Universitet van Amsterdam; ZRCS ， Zoological Reference Collections ， National University of Singapore; and ZSIISRS ，Zoological Survey of India ， Southem Regional Station ， Madras. Species Species synonymies in this paper (as in my papers generally) are intended to provide basic basic systematic references ， not complete or even extensive bibliographies. They include SYSTEMATICS AND BIOLOGY OF CIRRHINUS 173

prim 紅 y synonymy (original descriptions and designations of scientific names) and secondary synonymy (subjective name changes due to matters of taxonomic judgment). Additional synonymy may be provided ，when the references themselves permit reasonably sure identification identification of the species ， or when 1 have been able to examine voucher specimens upon which which the identifications were based. Some important instances of “negative" synonymy are are included ，when species are reidentified as something else. Such entries are preceded with with the neo ・Latin term “ nec" meaning “ not" or “ not of'. Type localities are cited as direct quotations from the original publications. Further explanation explanation or interpretation of the localities ，if not provided in the original publications ， is is clearly indicated as additional ， by being placed in parentheses or brackets. Some detail has has gone into listing the material examined ，particularly in the matter of providing localities ， dates ， and name of collectors. The importance of such information will become increasingly evident evident as the ranges of species change more and more due to human impacts (including introductions). introductions). As in most of my systematic revisions of genera ，組 account of the genus and generic type type species is given first ， followed by accounts of the other species in alphabetical orde r.

Cirrhinus Cirrhinus Oken 1817

Cirrhinus Cirrhinus Oken ， 1817 : 1182a (type species by monotypy Cyprinus cirrhosus Bloch ，1795). Isocephalus Isocephalus Heckel ， 1843: 1029 (type species app 訂 ently never designated according to Eschmeyer ， 1990: 200; type species by present designation Cyprinus cirrhosus Bloch ， 1795). 1795). Mrigala Mrigala Bleeker ， 1859: 427 (type species Cyprinus mrigala Hamilton ，1822 ，by absolute tautonymy; tautonymy; or Cirrhina bengalensis Bleeker ， 1853 by subsequent monotypy of Bleeker Bleeker = Cyprinus mrigala according to Bleeker ， 1860: 226). Cirrhinichthys Cirrhinichthys Bleeker ， 1863: 202 (type species by original designation and monotypy Cirrhinus Cirrhinus dussumieri Valenciennes in Cuvier & Valenciennes ， 1842 [=Cirrhinus ariza Buchanan ， 1807] (senior synonym of Henicorhynchus). Henicorhynchus Henicorhynchus Smith ， 1945: 245 (type species by original designation and monotypy Henicorhynchus Henicorhynchus lobatus Srnith ， 1945).

Cirrhinus Cirrhinus are small to moderately large labeoin cyprinid fishes. Large adults of 出e species species range from around 100 mm (C. lobatus ， C. soi) to 1m (c. cirrhosus). Cirrhinus Cirrhinus either lack barbels entirely ， or have one or two pairs of very small barbels. When only one pair of barbels is present ，it usually is 由 e rostral pair. In species (or individuals) individuals) in which only maxillary barbels are present ，these 訂 e very small or minute.

When both pairs of barbels 訂 e present ， the rostral barbels 訂 e invariably larger than the maxillary maxillary barbels. Length of rostral usually barbels much less than eye diamete r. The soft mouth parts immediately associated with the jaws in Cirrhinus are a discrete rostral rostral cap ， upper and lower lips ， and upper and lower homy jaw sheaths; the upper an d/ or or lower lip may be very weakly developed or absent (nomenclature after ROBERTS ，1982). These These structures are relatively simple compared to their counte 中 arts in other labeoins ， for 出ey bear a rninimum of homy tubercles and of papillae ，fimbriae ，plicae or other unculiferous s位uctures. 174 174 TYSON R. ROBERTS

Wh en papillae do occ 町 on 血 e rostral cap ，血 ey 紅 'e small and numerous ， not arr 佃 ged on deeply grooved fJlll briae ， but sufficiently developed to suggest 恥 condition of 白e more highly developed papillose fimbriae on 由e rostral cap of the labeoin genera Crossocheilus ，Garra and their close relatives (see ROBERTS ，1982 ，fig. 8). 百lI s explains why Day and other nineteenth cenωry ichthyologists ，particularly those working on In dian and Burmese fishes ， tended to include species of Cirrhinus in 也， e genus Crossocheilus. τ'h e larger keratinous s佐山知res known 酪 tubercles or “breeding tubercles" (which presupposes 出eir function ，usually unknown) are found in many cyprini 白血d esp 民 ially in in labeoins. 百ley are relatively poorly developed 泊 Cirrhinus but most of the species do have have tubercles on the snout ，especially on 血e ros 住al cap. Tubercles are perhaps least developed developed (absent?) in the lacus 凶 le species C. lu ，佃 d most highly developed in C. rubirostris. rubirostris. In血 e la 枇:er sp 田 ies 白ey are present in juveniles and adults of bo 出 sexes. In Cirrhinus Cirrhinus fulungee ， C. lineatus ， and C. lobatus ，血e great majority of spec 泊lens have no tubercles tubercles or very weakly developed tubercles ， but occasional individuals are strongly tuberculate. tuberculate. Cyprinids Cyprinids of the labeoin group 紅 e distinguished by the presence of a soft gular or vomero ・pala ta1 org 佃.百 lis usually consists of a paired series of fleshy lamellae ，釘med with with posteriorly-directed fingerlike processes or fimbriae ，on a discrete fleshy pad in the roof roof of 血emou 血.百 le vomero ・pala ta1 organ ，possibly a diagnostic characters of labeoins ， is is present in all species of Cirrhinus (Fig. 1). In some species the lamellae 紅 'e reduced but but large fimbriae project downwards and backwards 金om the roof of the mouth. Cirrhinus Cirrhinus have long ，血 in ， highly coiled intestines. Th e species 釘 e probably all iliophagous iliophagous (fi 田 ding on mud and de 凶旬s) an d/ or suctorial feeders on small invertebrates. V 註旬 ally no 仕ling ，however ， has been recorded on the feeding behavior of naturally-occurring Cirrhinus. Cirrhinus. Dorsal Dorsal fm with 3 or 4 simple rays and 8 to 16 branched rays. Last simple ray slender ， without without serrae on its posterior border. Th e anterior one or two simple rays usually are very small small and difficult to detect without or dissection radiography. Th e number of simple rays conveys conveys relatively little or no information useful in distinguishing species. 百le number of branched branched rays ，however ，is importan t. Th e smaller species of Cirrhinus (1 紅 g回 t specimens about about 10 0- 120 mm stan 伽 rd len 例。 and some moderately 1紅 'ge species almost inv 制 ably have have 8 branched dorsal fin rays ， whereas larger species have 合om 9 to 16 (with last branched branched ray usually divided to its base). Anal fm branched rays 5. Pelvic fin rays 9. Prin cipal caudal fin rays 10/9. None of the species previously recognized as Cirrhinus were known to have 9 or 10 branched branched dorsal fin rays ，and 血is was used by Banarescu and Kottelat as 血 e principal basis for for distinguishing Henicorhynch ω(wi 出 8 branched dorsal fm rays) 企om Cirrhinus (with 11 11 or more branched dorsal fin rays). Such a division ，which sep 紅 ates almost all of 血e larger larger species 企om all of the smaller ones ， does not seem phylogenetically valid. At least until until phylogenetic relationships are better understood ，it seems preferable to have one 1紅 ge (and (and perhaps polyphyletic) genus Cirrhinus 白血two smaller (and probably polyphyletic) genera. genera. An other problem with recognizing two generic groups on the basis of the supposed gap gap in number of branched dorsal fin rays is 白紙白e gap does not exist. 百lU S Cirrhinus ariza ariza c阻 have 8 or 9 branched dorsal fin rays ，Cirrhinus lu and C. inornatus new species SYSTEMATICS AND BIOLOGY OF CIRRHINUS 175 always always or usually have 9， and C. rubirostris new species 10. Scales Scales in the main longitud 泊al (l ateralline) scale row ，useful in distinguishing species ， r組 ge from 32 to 60. Transverse scale rows ， also useful ，r，叩 ge 仕om 611/4 (C. lu and C. C. soi) to 13/118 (C. microlepis). Vertebral Vertebral counts range from 32 to 41 ， with abdominal vertebrae about twice 部 numerous as as caudal. In In most species the number of scales in the lateralline scale row is 血e same as or very close close to the number of vertebrae. Th us C. lobatus normally has 32 vertebrae and 32-33 scales scales in the lateralline scale row ， C. lu 39 -4 1 vertebrae 佃 d 39 -4 0 scales. Th ere 紅 etwo main exceptions ， C. fulungee and C. microlepis ， in which the scales 紅 'e more numerous 白血也 e vertebrae. Cirrhinus fulungee has 3ι37 vertebrae and 42-52 scales in the lateral line line row ， C. microlepis 40 vertebrae 阻 d 53 -6 0 scales. No Cirrhinus have lateral line scales scales fewer 白佃 vertebrae. Perhaps Perhaps the most characteristic color feature of the genus is 出e pinkish ，rosy ，orangish or or sometimes blood red color of 白e pectoral ，pelvic ，anal ，and lower p紅 t of the caudal fin observed observed to some extent in all or ne 紅 ly all species. Such coloration is not unique ， but is more 仕'equent in Cirrhinus than in any other labeoin genus. Several of 血e species have other other distinctive color features. 百le dorsal fin of all species of Cirrhinus has more or less s紅ong 加te 町'adial pigmentation ， composed entirely of mel 組 ophores. In the Indian and Burmese species 血is inte 町adial pigmentation pigmentation tends to occupy ne 紅 ly all of the interradial surface ， while in Southeast Asian species species it tends to be res 凶cted to the middle or middle and distal po 凶 ons. In some species ，especially the smaller Southeast Asian ones ， the dorsal fin has a black margin. Wi 出 a single peculiar exception ， the other fins lack 釦 y dis 出lctive pigmentation features due due to melanophores. 百le one exception is the caudal fin of the holo 勿pe of C. caudiguttatus which which has a number of small but regularly arr 祖 ged dusky spots. No additional material definitely definitely referable to 白is species has come to light ，and its status as a valid species is doub 出1. Several Several Ci"hinus species have humeral marks but others do no t. In most species with a humeral mark it is black ， but C. molitorella ザpically has a bluish green humeral mark. Most species have body whitish or silvery (rosy to bluish in C. microlepis) ， without any distinctive distinctive marks. Longitudinal stripes occur in 伽田species ， C. lineatus ， C. fulungee ，and C. C. ariza. In some Cyprinidae with longitudinal stripes (e.g. La bωbarbus) 血es 凶pes go down the center of the scale rows. In such species 白es 住ipe on the lateral line scale row runs runs right down the lateral line. In Cirrhinus ，however ， the stripes 紅 e aligned with the obliquely obliquely overlapping zone of the scale rows ， so that the lateralline has s凶pes immediately above above and below it ，rather 由加 as 凶pe s回 ddling it. Some large sp 田 imens of C. ariza are are u凶que for the genus in having a broad midlateral s凶pe. La beo yunnanensis Chaudh 町 y，1911 ，佃 d L. decorus Peters ，1880 ， both described 貨'om Yunnan ，were 出 signed to 出e genus Cirrhinus by Banarescu ， 1972. Th ese species probably probably do not belong in Cirrhinus. 百le systematic status of both has been considered in in the recent monograph on Yunnanese Cyprinidae by Chu and Chen ， 1989. Th ese authors place place L. yunnanensis in La beo Cuvier ，1817 ， and L. decorus in Sinilabeo Rendahl ， 1932. Th e only Yunnanese species of Cirrhinus 由ey recog 凶ze is C. molitorella. 176 176 TYSON R. ROBERTS

Mouthparts Mouthparts of Cirrhinus are similar in most respects to those of another tropical Asian labeoin labeoin genus ， Tylognathus Heckel ， 1843. But Varicorhinus diplos ωmus Heckel ， 1838 (type (type species of Tylognathus) and other species provisionally referred to Tylognathus by Reid ， 1985: 287 have a well developed ethmoid furrow (absent in Cirrhinus).

Key to Species of Cirrhinus

Dorsal Dorsal fin branched rays usually 8 (9 in some C. ariza) …...・ H ・....・ H ・..… H ・H ・...・ H ・H ・H ・....2

Dorsal Dorsal fin branched rays 9 or more...... ・H ・....・ H ・-…....・ H ・-…....・ H ・.....・ H ・-…....・ H ・....・ H ・...・ ...7

2 Scales in lateral series 32-35 …...・ H ・..… H ・H ・...... ・ H ・..…… H ・H ・...・ H ・.....・ H ・H ・H ・.....・ H ・..…...・ H ・..3

Scales Scales in lateral series 42-52 …H ・H ・...・ H ・-…....・ H ・...... ・ H ・..… H ・H ・...……....・ H ・....・ H ・.c. fulungee

3 Transverse scale rows 6/1/4 (Mekong).... ・H ・....・ H ・-…...・ H ・...・ H ・...・ H ・H ・H ・.....c. ornat 伊innis

Transverse Transverse scale rows more than 6/ 1/ 4・…...・ H ・....・ H ・....・ H ・-…....・ H ・-…....・ H ・-…...・ H ・-… H ・H ・.4

4 Body with longitudinal stripes ・...... ・ H ・....・ H ・....・ H ・.....・ H ・-…....・ H ・...・ H ・-…....・ H ・....・ H ・....・ H ・.....5

Body without stripes... …H ・H ・...・ H ・H ・.....・ H ・..… H ・H ・...・ H ・..… H ・H ・...・ H ・.....・ H ・....・ H ・...・ H ・.....・ H ・...6 5 barbels Rostral present ， well developed; maxillary barbels absent; to 30 cm

(India ，Bangladesh).. …....・ H ・...... C. ariza Rostral Rostral barbels absent; minute maxill 訂 y barbels present; to 12 cm

(Mekong ，Chao Phraya) …H ・H ・-…....・ H ・...・ H ・...... ・ H ・-… H ・H ・-… H ・H ・H ・H ・...・ H ・-… H ・H ・.c. lineatus 6 Head large and broad; snout weakly or not projecting; total vertebrae 33 or more; to 20 cm (Mekong ，Chao Phraya)…••••••••••••••••••••••••••••.•••••••••.••••••••••••.•• ...... C. siamensis Head smaller and n紅 rower; snout often strongly projecting; total vertebrae usually 32;

to to 12 cm.... ・H ・....・ H ・...・ H ・....・ H ・-…....・ H ・...... 7

7 Caudal peduncle with mor 巴 or less bold spot (Chao Phraya) ・・ H ・H ・..c. caudimaculatus

Caudal Caudal peduncle without spot (Mekong) ・・ H ・H ・....・ H ・....・ H ・H ・H ・H ・H ・...・ H ・...・ H ・-… ...c. lobatus

8 Dorsal fin branched rays 11 or more.... …・・・ H ・H ・....・ H ・...... ・ H ・...・ H ・...... ・ H ・.....・ H ・-… H ・H ・-・ 9

Dorsal Dorsal fin branched rays 9 or 10... ・H ・...... ・ H ・....・ H ・...・ H ・H ・....・ H ・...・ H ・.....・ H ・..…...・ H ・.....・ H ・.11

9 Rostral barbels absent; lateral line scales 53 -6 0 (Mekong ，Chao Phraya) …H ・H ・...... c. ...c. microlepis

Rostral barbels barbels Rostral present; lateralline scales 39 -4 6.. ・H ・...... ・ H ・...・ H ・...... ・ H ・..… H ・H ・-… H ・H ・...10 10 10 Body with reticulated pattem due to concentrations of melanophores aligned with overlapping overlapping zone of scales; to 40 cm (China ，Indo-China ，Th ailand)..... C. molitorella

Body without reticulate pattem...... ・ H ・....・ H ・....…・・・ H ・H ・-…....・ H ・...・ H ・-……....・ H ・.....・ H ・-… 11

11 11 Body elongate ，predorsal profile nearly s凶 ight; pectoral ，pelvic ， anal and caudal fins colorless ，dusky ， or pinkish; to 1m (native to Indian subcontinent including Myanmar ，

but but widely introduced into China ，Indo-China ，Thailand)... ・H ・...・ H ・.....・ H ・..c. cirrhosus SYSTEMATICS AND BIOLOGY OF CIRRHINUS 177

Body relatively 凶 ncate ，predorsal profile steep; pectoral ，pelvic ， and anal fins ，組 d lower lower caudal fin lobe usually bright red; to 20 cm (l ower Mekong ，Chao Phraya) … …C. jullieni

12 12 Branched dorsal fin rays usually 10; snout strongly tuberculate..... ・H ・....c. rubirostris Branched Branched dorsal fin rays usually 9; snout without tubercles or weakly tuberculate.

…...・ H ・.13

13 13 Rostral barbels present ，maxillary barbels absen t....・ H ・-…....・ H ・..… H ・H ・....・ H ・.....・ H ・.c. lu

Rostral barbels barbels Rostral absent ，maxillary barbels presen t....…....・ H ・-…...・ H ・....・ H ・....c. inomatus

Cirrhinus Cirrhinus cirrhosus (Bloch ， 1795) Fig.2 Fig.2

Cyprinus Cyprinus cirrhosus Bloch ， 1795: 52 ， p l. 411 (reproduced here as Fig. 2) (type locality rivers rivers and lakes of the coast of Malabar). Cyprinus Cyprinus mrigala Hamilton ， 1822: 279 ，386 ， p l. 6 ，fig. 79 (type locality “ ponds and fresh waters waters of the Gangetic provinces"). Cirrhina Cirrhina rubripinnis Valenciennes in Cuvier & Valenciennes ， 1842: 288 ， p l. 479 (type locality locality "etangs de Calcutta"). Dangila Dangila leschenaulti Valenciennes in Cuvier & Valenciennes ， 1842: 235 ， p l. 471 (type locality locality “、 ea 卸ux douces de Pondi 比cher 可y" 勺) Cαi かrr 油hi 仇na pl ωumbe αValenciennes in Cuv 吋ie ぽr & Valenciennes ， 1842: 289 (type locality Irr awaddy). Cirrhina Cirrhina blochii Valenciennes in Cuvier & Valenciennes ， 1842: 290 (unw 紅 Tanted substite name for Cyprinus cirrhosus Bloch). Cirrhina Cirrhina mrigala ，Valenciennes in Cuvier & Valenciennes ， 1842: 294; Day ， 1877: 547 ， p l. 129 ，fig. 4). Mrigala Mrigala Bleeker ， 1859: 427. Mrigala Mrigala buchanani Bleeker ， 1860: 226 (replacement name for Cyprinus mrigala). Cirrhina Cirrhina leschenaultii ，Gunther ， 1868: 36. ?Cirrhina ?Cirrhina macrops Steindachner ， 1870: 636 (type locality Madras). Cirrhina Cirrhina cirrhosa ，Day ， 1877: 547 ， p l. 131 fig. 3. Cirrhinus Cirrhinus horai Lakshm 組組， 1966: 59 (type locality Godavari Ri ver at Rajahmundry ， An dhra Pradesh). Cirrhinus Cirrhinus chaudhryi Srivastava ， 1968: 30 (type locality “ ponds near Naua Dumari ， 11 miles miles south of Gor 此 hpur" ， Uttar Pradesh ， northem India).

Type material examined.-MNHN A3362 ， 219 mm ，Irrawaddy ，Reynaud (holotype C. plumbea); plumbea); MNHN 3854 ， 3: 160-255 mm ， India (syntypes C. rubripinnis); MNHN 3852 ， 184 184 mm ， Pondichery (holotype C. leschenaulti). Non-type Non-type material exarnined. -C AS (SU) 41125 ， 163 mm ，Bisrampur ，Central Pr ov. ，India ， 13 13 Dec. 1040 ，A. W. He 町'e; BMNH 1889.2. 1. 29 4- 96 ，3: 74.2-101 mm ，Madras ， F. Day; USNM 44790 ， 164 mm ，Bhamo ， upper Irr awaddy ，1885-89 ， L. Fea. 178 178 TYSON R. ROBERTS

Di8gnωis. 針。 bably the largest sp 田 ies of Cirrhinus ，叫 aining 90 cm or 1m (D ay ， 1877: 1877: 548; Jhingran & Kh an ， 1979: 33); rostral barbels present ， length about h必f eye diameter; diameter; maxillary barbels smaller 血叩 ros 住al barbels or absent; rostral cap without f1IIl briae or papillae ，its oral margin entire; upper 1培 absent; lower lip absent 町 weakly developed ，represented by a tr 佃 sverse series of small papillae near m 紅 g泊 of lower horny jaw sheath; lateralline scales 39 -4 6 ，transverse scale rows 7/116 ，circumpeduncular scales 20; 20; branched dorsal fm rays 12-15; ve 由 brae 26+13=39. Tubercles Tubercles on rostral cap weakly developed ，represented by small tubercle scars 泊 most larger specirnens examined.

Discussion. Discussion. Recent authors have recognized 由ree 1紅 ge species of Cirrhinus in In dia (J AY 成 AM ， 1981; T此，WAR& 加 NG 貼 N ， 1991). According to 伽 se au 白'O rs ，白 e sp 田 ies most c'O mm 'O nly used 泊 aquacul 旬飽佃d most widely dis 位ibuted is C. ，!，- rigala; C. cirrhosus ， also also cultured ，occ 町 s in rivers of peninsular In dia flowing into 白e Bay of Bengal ，合 om 血e Godavari Godavari to the Cauvery; while C. macrops ， not used in aquacul 加re ，is comm 'O n in the fishery fishery catches of the Godavari Ri ver at Rajamundry and also 'O ccurs at Madras (TALWAR & JH 別 GRAN ， 1991: 172). Cirrhinus Cirrhinus cirrhosus supposedly differs from C. mrigala in having maxillary b紅 bels well well developed instead of small or absent 仰町G 臥 N & KHAN ， 1979: 1) ， and 15-16 ins 伽 d of of only 12-13 branched dorsal fm rays (T 此，WAR & JH 町 G 貼 N ， 1991: 170). Since its placement placement by Gunther ， 1868: 36 (wi 出 question m 釘 k) 叩 d 也en by Day ， 1877: 547 (wi 出'O ut question question m 釘 'k) as se 凶or s戸lO nym of C. leschenaultii ， C. cirrhosus almost invariably has been been regarded as av a1i d species with two p厄rs of b紅 bels. 百le pr 'O blem is 白at Bloch's plate plate 'O f the holotype of C. cirrhosus shows only two b紅 bels ，and 由ey are both defmitely ros 位al barbels (Fig. 2). 百le unavoidable conclusion is 由at C. cirrhosus is 血e most se 凶'O r synonym for the species usually known as C. mrigala. 百le name C. cirrhosus should not be suppressed in favor of C. mrigala because C. cirrhosus has been recognized by m 組 y authors authors as av a1i d species ， and is ，泊 addition ， type species of the genus. 百le holotype of C. leschenaulti has a pair of rostral and ap 耐 of maxillary barbels: length length 'O f rostral barbels about one-half eye diameter; m 拡 ill 紅 y barbels much thinner and n'O t qui 飽 s'O long ， length about 215 eye diameter; total gill rakers on first gill arch at least 73 ， perhaps a few missing at anterior end of lower arch ， so total might be ne 紅 'er 80; branched branched dorsal fin rays 14; vertebrae 26+13=39. 1 have comp 紅 ed this specimen direc t1 y with with specimens 'O f comparable size identified 踊 C. mrigala (i. e. ，lacking maxill 紅 ybarbels) and fmd them ex 回 mely simil 低 Cirrhinus Cirrhinus horai Lakshmanan ， 1966 was based on presumably wild stocks of spaw 凶ng fishes fishes caught 泊恥Go daveri Ri ver at R 桝 lffi Us句， An dhra Pra desh. Tι ，WAR&J 剛 GRAN (1991) (1991) placed it as a synonym of C. macrops Steindachner ，1870 ，a species generally not recognized recognized since its original description. 1 have not examined type material of either C. horai horai or C. macrops. Morphological differences between these two n'O minal taxa and 血e more readily available cultivated examples of C. cirrhosus might be due to in 位制pecific vanation. vanation.

stribution. Di stribution. Cirrhinus cirrhosus has been so widely transported 泊 connection with aquacul 制民社lat its na 佃raldis 住ibution no longer can be determined. Day ，who regarded SYSTEMAT ICS AND B IO LOGY OF C IRRHI NUS 17 9

同

Figut 巴 1. V0 1ll erOpal alal orga n (Ci rrh il//I 可 m birosl ri s，265 1ll1ll )

へ'J切 llUH Il :J ?U 冗υj 'J"( ，') ;おも、、山下、、:\~ Ir ~、、L A 号、、、石、、，\~~l( 、 ~\t 吋，:>-，>~，，，，，.，.、ふ~.、

Figur e 2. Ci rrhi l/ lI s cirrl /O s lI s. Bl och ，1 795 ， p l. 4 11 (p hologra ph co url esy Ca rl J. Ferr a ri s). No te pr ese n ce of onl y a singl e pa ir ofb arbe ls( ros lr al) 180 TYSON R. R OilERTS

Figure 3. Cirrhin11 s ari ~a . Buchanan , 1807, pi. 3 1.

Figure 4. Cirrhin11s ariza, 164 111 111 , Patna mark et (Ganges basin).

Fi gure 5. Cirrhintts inom a t11 s. holotype, I I 5 111111 , M anda lay. SYSTEMATICS AND BIOLOGY OF CIRRI-liNUS 181

Figure 6. Cirrhin11s )111/ieni. Above, Phnom Penh (Mekong basin); be low, Sukhothai (Chao Phraya basin ).

Figure 7. Cirrhin11s lin ea/lis. 80.4 111111. Mekong nea r That Phanom. 182 182 T YSON R. R OsE RT S

F igur e 8. C irrh illl ls loba fll s，70 mm grav id femal e，M ekong below Khon e Fall s

Fig ur e 9. Ci rrhillll s 111 ，1 45 mm ，Inl 巴 Lak e

Fi glll 巴 10. Ci rrhillll s mi crol ep is. Up p巴r，ju ve nil e from Gr e at L ake ，Ca m bod ia; lower ，v io let or or pu rpli sh hu ed adu lt ，m id dle M ekong SYSTEMA TICS AND BIOLOGY OF CIRRHINUS 183

C. C. cirrhosus 佃 d C. mrigala as distinct species ， gave the distribution of C. mrigala as “Cutch ，S 泊d，Pu吋 ab ，Bengal ，組 d Burma" 釦 d “rivers and t創立 s in Bengal ，Deccan ，N.W. Pr ovinces ，Pu niab ，Sind ，Cutch ， and Burma" and that of C. cirrhosus as “Godavery ，Ki stna [=Kr ishna] and Cauvery rivers ， and generally in south India" (Day ， 1877: 547-548). JHINGRAN & KHAN (1979) indicate 伽 t C. cirrhosus (called by them C. mrigal α)， has been been widely introduced in southem India ， which is certainly 佐ue. The point is ，however ， 出at the naturally occurring distribution is not known and is now unknowable. If there were were formerly two distinct species ， one more southerly with two well developed pairs of barbels barbels and one more northerly with (usually) only a single pair of barbels ， both have by now been repeatedly 叩 d massively introduced into each other' s natural ranges. Th ere has almost almost certainly been widespread introgressive hybridization of introduced and any na 知rally occurring occurring stocks due to artificial production of seed for aquaculture by fisheries stations throughout throughout In dia.

Cirrhinus Cirrhinus ariza (Buchanan ， 1807) new combination Figs.3 -4

Cyprinus Cyprinus ariza Buchanan ， 1807: 344 ， p l. 31 (reproduced as Fig. 3) (type locality “a small clear clear stre 創 n called the Vedawati" ， a tributary of 白e Tungabahdra R.， Kri shna basin; no type type specimens). Cyprinus Cyprinus ariza H 創凶lton ， 1822: 286 (“ rivers of Bengal as well as those of the peninsula of of India"). Cyprinus Cyprinus reba Ham i1 ton ， 1822: 280 ，386 (type locality Bengal; Behar; no type specimens). ?Gobio limnophilus McClelland ， 1838: 279 ， p l. 55 ，fig. 3 (type locality “ ponds in Bengal 勺 location of type material unknown). ?Gobio ?Gobio isurus McClelland ， 1838: 277 (type locality upper Assam; location of Type material u此 nown). Cyprinus Cyprinus bangon Buchanan ，in 民1c Clelland ，1838 ， p l. 107 ，fig. 2 (figure only; no type specimens). specimens). Cirrhina Cirrhina dussumieri Valenciennes in Cuvier & Valenciennes ， 1842: 291 ， p l. 480 (type locality locality eaux douces de Mysore). Chondrostoma Chondrostoma gangeticum Valenciennes in Cuvier & Valenciennes ， 1844: 399 (type locality Gange). Gange). Cirrhina Cirrhina reba Day ， 1877; 549 ， p l. 130 ，fig. 3. La beo ariza Day ， 1877; 544 ， p l. 132 ，fig. 5 (“ Wynaad and Bowany river at the foot of the Neilghe 町Yh i1l s in Madras ，叫a Is 叩o Cauvery rive 釘r" 勺) ?μ be ωO αU似バri切za α Ta 剖lw 紅 &J拍hi 均n昭喝1gran 創佃札n仏1，1ω99 引1: 刈19 兜8.

Type material examined.-MNHN 3153 ， 73.0 mm ，Gange ，Reynaud (holotype C. gangeticum). gangeticum). Non-type Non-type material exarnined. -C AS 24237 ，5: 63.7-103 mm ， Indus R. 325 mi N ofKarachi and and 5 mi N of Su は ur ， 1-11 Nov. 1968 ， E.S. Herald; CAS 29653 ， 85.8 mm ，Ar abian Sea off off Karachi ，Pakistan ，22 Oct. 1973 ， F.B. Steiner; CAS(SU) 52929 ， 6: 64.5-75.9 mm ， Nepal ，Biratnagar ，27-30 Nov. 1955 ，A. C. Taft; CAS 50369 ， 126 mm ，Nepal ，Terai ， 184 184 TYSON R. ROBERTS

Kalaiya Kalaiya market ， 12 km E of Birganj ，1 May 1975 ，T. R. Roberts; CAS 91612 ， 5: 86.5-185 mm ， Patna market ，April-May ，1996 ，T .R. Roberts; CAS(SU) 41128 ， 117 mm ，Tozpur market ，Assam ，no date ，Zoological Survey of In dia; CAS(SU) 34568 ， 2: 74. 0-- 83.0 mm ， Mahanadi ，Siliguri ， 10 April 1937 ，A. W. Herre; CAS 79176 ， 128 mm ， Hirakud Reservoir 組 d Samb a1 pur market ， Orissa state ，Mahanadi basin ，22-24 Feb. 1985 ，T. R. Roberts; CAS 61823 ， 85.1 mm ，India ， Orissa State ，Mahanadi basin ， Sonepur market ， Feb. 1985 ，T. R. Roberts; Roberts; CAS(SU) 34569 ， 2: 13 6-- 146 mm ，Deol a1 i， Nasik Dis t.， Bombay ，1935-36 ，A. G. L. Fraser; Fraser; BMNH 1938.2.22.8 6-- 87 ， 2: 131-180 mm ， Godaveri R. headwaters ，Deolali ， Nasik Dis t.， Bombay Na t. Hi st. Soc.; CAS 62067 ，2: 102-106 ， and CAS 62093 ，6: 88.5-217 mm ， Tungabahdra R. and Reservoir at Hospet and Kampli ，28 Jan.-3 Feb. 1985 ，T. R. Roberts; CAS(SU) 34567 ， 3: 81. 5-90.8 mm ， near Kodur ，Cuddapah ， South India ， 10 April 1937 ， A. W. Herre; CAS 62032 ， 9: 49. 4-- 89 .4 mm ， Cauvery basin NW and WNW of Mysore ， Karnataka Karnataka State ，5-8 Jan. 1985 ，T. R. Roberts; CAS(SU) 34566 ， 124 mm ，Pu lta ，India ， April April 1937 ，A.W. He 町 e.

Diagnosis. Diagnosis. A moderately large species ，att 沿凶ng 30 cm (TALWAR & KHAN 1991: 174); 174); body with thin s凶 pes mostly above lateral line; larger fish sometimes with a broad midlater a1 s回pe (not observed in any other Cirrhinus); later a1 sc a1 e series 32-35; transverse sc a1 e rows 7-8/ 1/ 5--6; dors a1 fin branched rays 8-9; vertebrae 22-24+ 11 ・12=34 (4) ，35 (3). Color Color in life variable ，overall dull dirty white or greyish ，silvery or yellow; thin stripes v紅 y from subdued to bold. Comments. Cyprinus ariza Buchanan ， 1807 is one of the earliest described freshwater fish fish species of India ，and is included on the classic work on Gangetic fishes by Hamilton (1822). (1822). McClelland ， 1838: 357 noted its similarity to his new species Gobio isur 附組d G. G. limnophilus ， both subsequently recog 凶zed as junior synonyms of C. reba. Jerdon ， 1849: 1849: 308 noted the similarity between C. ariza and C. bangon and suggested they might be conspecific. Cyprinus bangon has been placed as a synonym of C. reba by most authors ， and 1 concur. Buchanan's drawing of C. ariza shows the exposed m 紅 gins ofthe sc a1 es in all rows above the lateral line and two rows below it with very distinctive diamond-shaped diamond-shaped dark outlines. Such a color feature ，a1 though not observed exactly as illustrated illustrated in 叩 y species of Indian freshwater fish known to me ，is closely approached by some specimens of the species commonly identified as C. reba ， in which the longitudinal stripes stripes between the sc a1 e rows tend to zig-zag. Th e original description of Cyprinus ariza by Francis Buchanan (1 762 ー1829) ，who formally formally changed his name to Francis Hamilton in 1818 ，was based on a fish with exception a1 1y bold longitudin a1 位 ipes. In many individu a1 s，especi a1 1y those coming from muddy water or having been dead for some time ， the stripes are much more subdued. Hamilton Hamilton (1822) reported C. ariza from Beng a1 and southem India but did not compare his his Cyprinus reba and C. ariza directly. His description of C. reba includes several characters characters not treated in that of C. ariza ， such as variation in coloration. Two substantive differences differences he does mention 紅 e 11 instead of 12 dorsal fin rays [8 instead of 9 branched rays] rays] and “ minute" rostral present barbels instead of absen t. These differences represent individu a1 variation. In In the fish collection of the Southem Research Station of the Zoologic a1 Survey of

India India in Madras 1 examined two lots identified ad C. reba 血at may repr 回l ent a distinct SYSTEMATICS AND BIOLOGY OF CIRRHINUS 185

species species characterized by fewer transverse scale rows ，i.e. only 6/114. The specimens consequently consequently have fewer longitudinal stripes and thus look quite distinctive ， but no additional differences differences were noted to distinguish them from C. reba. The lots are ZSIISRS F. 4542 ， 2: 2: 95-110 mm ， Maruthar Damn ，Tirunvelv 巴li district ， southern India ，2 April 95 ，M.B. Ragunathan; Ragunathan; and ZSIISRS F. 4255 ， 131 mm ， Tampraparni R. near Palayam Kotai ，8 March 1995 ， M.S. Ravichandra. Distribution. Distribution. Indus plain and adjoining hilly areas (Pakistan); Ganges-Brahmaputra basin basin (India ，Nepal ， and Bangladesh); Mahanadi ，Kr ishna ，Cauvery ， and some smaller basins basins in southern India; Karnapouli and adjacent smaller basins in Chittagong Hill Tracts (Bangladesh). (Bangladesh).

Cirrhinus Cirrhinus caudimaculatus Fowler ， 1934 new combination

Tylognathus Tylognathus caudimaculatus Fowler ， 1934: 133 ，figs. 89-90 (type locality Chieng Mai). Crossocheilus Crossocheilus caudiguttatus Fowler ， 1934: 137 ，fig. 103 (type locality Chieng Mai ， north Siam [probably=Menam Ping]). Cirrhinus Cirrhinus (Henicorhynchus) caudiguttatus ，Bohlke ， 1984: 72. Henicorhynchus Henicorhynchus caudimaculatus ，Rainboth ， 1996: 111.

Type material examined.-ANSP 58332 ， 42.5 mm ， Chieng Mai ，1 Jan. 1933 ，R.M. de Schauensee Schauensee (holotype T. caudimaculatus); ANSP 58333 ， 26.6 mm ，collected with holotype (paratype (paratype T. caudimaculatus); ANSP 58452 ，6 1. 0 mm ，Chieng Mai (no further 10cality=Menam 10cality=Menam Ping) ，5 Feb. 1933 ，R.M. de Schauensee (holotype C. caudiguttatus); ANSP 59089-91 ，3: 28.5-35.0 mm ，same data as ANSP 58452 (paratypes C. caudiguttatus). Additional Additional materia l.- The following lots ，all from the Chao Phraya basin ， are identified as C. C. caudimaculatus: USNM 117769 ， 27: 45.0-78.5 mm ， upper Nan River at Ban Kh wang ， northern northern Siam ， 31 March 1936 ， H.G. Deignan; USNM 119493 ， 2: 59. 6- 60 .5 mm ，Chao Phraya Phraya at Bangsai ，central Thailand ， 27 Nov. 1923 ，H.M. Smith (paratypes H. lobatus); USNM 119491 ， 68.0 mm ，Chao Phraya at Paknampo ， 19 Nov. 円1923 ，H

H. H. lob αω tu 附の); S USNM 119494 ， 73.5 mm ， Pasak R. at Dha Luang ，central Thailand ，20 Aug. 1923 ，H.M. Smith (paratype H. lobatus); USNM 108093 ，3: 70.2-73.3 mm ，Menam Sak above above Dha Luang ， 10 Dec. 1923 ，H.M. Smith; USNM 107880 ，6: 70. 6- 86.5 mm ，Menam Nan at Nan ， 29 March 1936 ， H.G. Deignan; USNM 119514 ， 57.3 mm ，Menam Kon ， tributary tributary of Menam Nan ， 21 April 1936 ， H.G. Deignan; USNM 108092 ，4: 60.6-73.2 mm ， Phetchabun ，Chao Phraya above Bangkok ，4 Jan. 1925 ，H.M. Smith; USNM 108094 ， 45.7 mm ，Bung Borapet ， 19 Nov. 1923 ，H.M. Smith; USNM 109783 ， 7: 72.1-90.7 mm ，Nan Ri ver ， 23 Aprill930 ，Luang Praserth Aksorn; CAS 91781 ， 55.2 mm ，Menam Wang 79 km by road N of Lampang and 6 km E of highway 1035 ， 28 Feb. 1991 ，T. R. Roberts.

Comments. Cirrhinus caudimaculatus appe 紅 s to be a valid species known only from the the Chao Phraya basin. is It very similar to the Mekong species C. lobatus ，仕om which it it consistently differs in having a dark round spot on the caudal peduncle (a feature not present present in any C. lobatus observed by me from the Mekong basin of Thailand ， Laos or Cambodia). Cambodia). Like C. lobatus ，it is a small ，small-headed species ， often (but not always) 186 TYSON R. ROBER 百

with with a strongly project 泊g snout not observed in other species. Cirrhinus siamensis from the the Chao Phraya basin occasionally has a peduncul 紅 mark like C. caudimaculatus ， but its head is much bigger and the mark is (always?) fain t. Coloration in life not recorded. 1 have not found any additional specimens 血at agree with the holotype of C. caudiguttatus caudiguttatus as described and figured by Fowler. Wh en 1 examined it in November 1992 ， the the caudal fm of 血e holotype still showed the distinctive pattem of small round spots (not known in 姐 y other specimen of Cirrhinus) described and figured by Fowler. Fowler did not not specifically mention these spots in 血ep 紅 atypes ，which 紅 'e much smaller specimens than than the holotype ，組d 由e spots were not evident in 由ep 紅 'atypes when 1 examined them at 血e same time as 白e holotype. Except Except for the caudal fin coloration of the holotype ， the type specimens of C. caudiguttatus 紅 e similar to much larger type specimens of C. caudimaculatus ， also described from Chieng Mai by Fowler. 百le holotype of C. caudiguttatus has ve 巾 brae 21+11=32. This This is the first time 白紙 C. caudimaculatus and C. caudiguttatus have been identified 出 being being the same species. As both were described in 血e same paper by Fowler ，1934 ，1 hereby hereby select C. caudimaculatus 創出e senior 0 均配 tive synynom. Cirrhinus Cirrhinus caudimaculatus is said to be extremely common in central Thailand (RA 別 BO 百 ζ1996: 111) ， but 1 have collected few spec 耐lens in the Chao Phraya since 1 began collecting there in 1970. 百le species app 釘 'ently is greatly reduced and may be ne 紅 ing extinction.

Distribution. Distribution. Cirrhinus caudimaculatus is known only 企om the Chao Phraya basin.

Ci"hinus fulungee (Sykes ， 1841)

Chondrostoma fulungee ，Sykes ， 1841: 358 (type loc a1i ty Du凶 un). Gymnostomus fulungee ，Gunther ， 1868: 76. Cirrhina Cirrhina fulungee ，Day ， 1877: 549 ， p l. 132 ，fig. 1.

Type material examined.-None. Apparently there is no extant type material of C. fulungee. In In November 1992 Oliver Crimmen and 1 searched in the Sykes fish collection deposited in 白e BMNH (including his dried spec 加 ens) for type material but did not find 飢 y. Non- 旬pe material examined. -C AS(S U) 41123 ， 5: 66.9-95.7 mm ，Poona Dis t.， Bombay Pr es. ，3 April 1937 ，Zo ological Survey of In dia; CAS(SU) 41124 ， 2: 88. 0- 91. 8 mm ， Mugao ，Dharwar ，Bombay Pre s. ， 21 July 1937 ，Zo ological Survey of In dia; CAS 61967 ， 2: 2: 102-107 mm ， Bedti or Gangavali Ri ver ， about 15 km E of Yellapur ，N Kann ara Dis t.， Karn ataka State ，25 Jan. 1985 ， T .R. Roberts; CAS(S U) 34563 ， 2: 63.5-80.2 mm ，Kodur ， Cuddapah ， South In dia ，April1937 ，A. W. He 町 e; BMNH 1889.2. 1. 362 ， 88.3 mm ，Malab 紅， F. F. Day. Diagnosis. Diagnosis. A small or moderatedly large species of (to 30 cm according to TALWAR &JH 町 GRAN ， 1991: 171); ros 回 1 barbels present; body with 也in s凶pes; scales 泊 lateral series series 42-52; 位叩sverse scale rows 10- 111117; dorsal fm branched rays 8; vertebrae 23-25+12-13=36 (5) ，37 (6). 百le two specimens from 由e Gangav a1i (CAS 61967 ， 102 and 107 mm) 紅 'e markedly SYSTEMATICS AND BIOLOGY OF CIRRHINUS 187

tuberculate. tuberculate. The entire rostral cap ， snout between nos 凶Is ， and lacrimal 釘 ea is covered with with large tubercles. A l1 other specimens of C. fulungee examined are non-tuberculate. Th e 80.2-mm specimen from Kodur ，CAS(SU) 34563 ，appe 紅 s to be a ripening female. 百lI s is remarkably small size for sexual maturity for a species supposedly attaining 30 cm.

Distribution. Distribution. Cirrhinus fulungee occ 町 s in “Poona and Deccan" according to Day ， 1877: 1877: 549.

Ci"hinus inornatus new species 日g.5

Holotype.-εAS 91772 ， 115 mm ，Myanmar ，Mandalay market ， 13-25 April 1993 ，T .R. Roberts. Roberts.

P 紅 atypes. -C AS 91776 ， 8: 76.8-129 mm ，same data as holotype; CAS 88903 ， 106 mm ，

Pagan Pagan market ， 13 April1996 ，C.J. Fe 町 aris 佃 d D. Cata 凶a; CAS 91774 ， 9: 63.3-86.8 mm ， Pagan Pagan market (Irr awaddy R ふ8 Nov. 1996 ，C.J. Ferr 組 s; CAS 91775 ， 122 mm ， Myitkyina market market (Irr awaddy b槌 in) ，20 ー21 Apri11996 ，C.J. Ferraris; CAS 91773 ， 6: 94.ι116 mm ， Taungoo market (Sittang R よ 7 April 1996 ，C.J. Ferraris and D. Catania.

Diagnosis. Diagnosis. Total gil1 rakers on first gill arch 35; vomero ・palatine org 佃 with only 5 pairs pairs of fimbriate lame l1 a; lame l1 ae comparativ e1 y broad ， each with up to 20 short funbriae; branched branched dorsal fin rays usually 9. Lateral scales 35; transverse scale series 7/ 1/ 5; circumpeduncular circumpeduncular scales 20; vertebrae 22 ・24+12-13=35 (4) ，36 (1)， or 37 (2). Body du l1 white white or silvery overall; fins dusky ，∞ lorless; humeral mark present ，otherwise no dis 出 ctive markings. markings.

Comments. In出 e Irr awaddy Cirrhinus inornatus co ・occ 町 s with a sp 即 ies of Bangana ， possibly possibly B. devdevi Hora ， which it superflcially resembles. Th e Bangana differs in having a more prominent snout ，a vertically elongate peduncul 釘 m 紅 k， and 11 instead of only 9- 10 10 branched dorsal fin rays.

Distribution. Distribution. Cirrhinus inornatus is known only from 白e Irr awaddy and Sittang basins basins in Myanm 低

Ci"hinus jullieni Sauvage ， 1878 Fig.6

Cirrhina Cirrhina jullieni Sauvage ， 1878: 237 (type locality Stung-Strang [=S 伽 ngTreng ，Mekong basin ， Cambodia]). nec nec Cirrhina jullieni ，Sauvage ， 1881: 174 ， p l. 6 ，fig. 2 (=Cirrhinus siamensis; based on syntype syntype of Morara siamensis). Rohita Rohita sima Sauvage 1881: 177 (nec Rohita sima Sauvage 1878). nec nec Osteochilus spilopleura Fowler ， 1935 (=c. moli ωrella). Cirrhina Cirrhina chinensis ，Fang ， 1942: 168. 188 188 TYSON R. ROBERl 百

nec nec Cirrhinus jullieni ，Smith ， 1945: 162 (=c. siamensis). Cirrhinus Cirrhinus jullieni Rainbo 血， 1996: 107.

Type material exar 凶ned. 一MNl王 N 8586 ， 109 mm ，Fl. MιKong ，Cochinchine ，1874 ，Jullien (l ectotype C. jullieni ， designated by Banarescu ， 1972); MNHN B. 2960 ， 5: 88.0 ー95.5 mm ， Cochinchine ，Jullien (p 翻 lectotypes C. jullieni); M Nl王 N 3999 ， 90.2 mm ，Cochinchine ， Jullien Jullien (p 訂 'alectotypeι jullieni). Non-type Non-type material examined.-MEKONG BASIN: MNHN 1983.19 ， 145 mm ， Tonle Sap ， 7J 加 .1963 ， F. d'Aubenton; MNHN 1983.129 ， 89.5 mm ， Tonle Sap Km 9，29 Jan. 1961 ， F. F. d' Aubenton; MNHN 1983.198 ， 84.5 mm ， Tonle Sap km 9，2 Nov. 1961 ， F. d' Aubenton; CAS 68202 ， 4: 83.ι110 mm; CAS 91610 ， 13: 94.7-104 mm ，Phnom Penh market ， 18 Jan.-20 Jan.-20 Feb. 1994 ，K. E. Witte. CHAO PHRAYA BASIN: CAS 79168 ， 12: 80.2-101 mm ， 組 d MNHN 199 0- 621 ， 4: 78. 0- 109 mm ， Sukhothai market (Menam Yom) 23 Febru ぽ Y 1989 ，T. R. Robe 氏s; CAS 91748 ， 104 mm ，Sawankhalok market (Menam Yom) ，36 km N of of Sukhothai ，2 Feb. 1989 ，T. R. Roberts. Diagnosis. Diagnosis. A medium-sized species ，attaining at least 20 cm; predorsal profile relatively ste 叩， ros 回 1 barbels well developed ， length more than one-half eye diameter; maxillary barbels barbels absent; branched dorsal fin rays 14- 16; scales in lateral series 36; transverse scale rows 8/ 1/ 6; vertebrae 22 ・24+ 11=34-35. Body overall silvery or bluish silvery ， with or without faint humeral mark; no peduncular spot; spot; p田 toral ，pelvic ，組 d anal fins and lower one 血ird or two- 白irds of lower caudal fin lobe lobe often blood red. 1n other Cirrhinus with red caudal fins ， usually the entire fin is red. Cirrhinus Cirrhinus jullieni is somewhat similar to the much larger species C. molitorella ，from which it differs markedly in coloration. Also ，it is deeper bodied ，especially anteriorly ，and has has a larger ，more falcate anal fin ， the distal end of which extends posteriorly to below posteriormost posteriormost scales on caudal fin. Discussion. Discussion. This species was known for a long time only from the type specimens from the Lower Mekong basin. As pointed out by Fang ， 1942: 168 ，由 e redescription 叩 d figure figure of C. jullieni by Sauvage ， 1881 is not based on 白e type specimens of 血at species ， but but rather on specimens of a different species. The species subsequently w 部 identified as Henicorhynchus Henicorhynchus [=Cirrhinusl siamensis by Banarescu ， 1983: 16. Banarescu's identification is is co 町民t; the description and figure published by Sauvage ， 1881 is actually 白紙 of a syntype syntype of C. siamensis (originally described as Morara siamensis ，a species not otherwise fig 町 'ed by Sauvage). Specimens recently collected from the Chao Phraya basin were c<;>mp 紅 ed by me directly directly with types and other material of C. jullieni from the Lower Mekong in Cambodia and they 紅 'e very similar.

Di stribution. Cirrhinus jullieni occ 町 s in the Chao Phraya basin in central Thailand and the lower Mekong basin in Cambodia and in southemmost Laos below Kh one Falls. 1t 1t seems to be absent in the Mekong above the waterfalls. Pr evious records 仕om Thailand (e.g. (e.g. SM 町 H ， 1945; SODSUK ， 1988) app ぽ 'ently 紅 'e all based on C. siamensis. SYSTEMATICS AND BIOLOGY OF CIRRHINUS 189

Cirrhinus Cirrhinus lineatus Smith ， 1945 Fig.7 Fig.7

Cirrhinus Cirrhinus lineatus Smith ， 1945: 163 ，fig. 25 (type loc a1 ity Larn Ton Lang ， a tributary of the the Menarn Sak ，central τbailand [Chao Phraya basin]). Henicorhynchus Henicorhynchus cη 'P topogon ，Rainbo 出， 1996: 111 (Chao Phraya and middle Mekong; misidentification). misidentification). Type materi a1 examined.-USNM 107960 ， 118 mm ，Lam Tong Lang ， 19 July 1925 ，H.M. Smith Smith (holotype C. lineatus); USNM 119484 ， 109 mm ，Me Fang ，tributary of Me Kok ， Mekong basin ， 12 July 1936 ， H.G. Deignan (paratype C. lineatus). Non-type Non-type material examined.- CAS 79169 ， 5: 64.9-95.2 mm ，mouth of Huay Ngao where where it flows into Mekong R. 1 km S of Ban Chaem Pong (about 30 km S of Chiang Kh ong) ， 12 May 1990 ，T. R. Roberts; CAS 91764 ， 7: 50.1-63.1 mm ，Menarn Kok at Th a Ton and up to 5 km downstrearn ， 15 May 1990 ，T .R. Roberts; CAS 91766 ， 9: 54.5-106 mm ，Mekong mainstrearn from Pak Ing to Jom Paeng (about 4- 5 km downstrearn from Pak Ing) ，16- 18 Jan. 1989 ，T. R. Roberts; BMNH 1980.12.17.121-122 ，2: 48.2-55.7 ， Payao Lake ，N Th ailand ， Oct. 1978 ，J. Karnasuta; CAS 91763 ，4: 41.4- 50.5 mm ，Se Kh one ne 訂 Stung Stung Treng ，6 Feb. 1994 ， T.R. Roberts; CAS 91765 ， 101 mm ，0 Changni ，sm a1 1 strearn on road from Ann Long Mea to Ban Lung ，Ratanakiri prov. ，Carnbodia ， Feb. 1994 ，T. R. Roberts Roberts Di agnosis. A small species of Cirrhinus; longitudinal stripes on body; no rostral barbels; barbels; small max i1l ary barbels; branched dors a1 -fin rays 8; sc a1 es in lateral series 34- 35; 仕ansverse scales rows 6/115; vertebrae 20 ー22+10 ー13=32 (5) ， 33 (6). The s住ipes v町 individu a1 1y from faint to very bold. When bold ， they may assume a zig-zag course ， as in in the South Asian species C. ariza ， which has very similar stripes. Comment. The holotype of Henicorhynchus cη 'ptopogon Fowler ，1935 ， is a Lo boche i/ os.

stribution. Di stribution. CirrJ 旨inus lineatus is known from the Chao Phraya and Mekong basins (from (from southem Yunnan to the Mekong delta).

Cirrhinus Cirrhinus lobatus (Smith ， 1945) Fig.8 Fig.8

Henicorhynchus Henicorhynchus lobatus Smith ， 1945: 257 ，fig. 49 (type locality Me Kok ne 紅 Chiengrai [Mekong basin]). Cirrhinus Cirrhinus lobatus Roberts and Baird ， 1996: 247 (migrations at Kh one F a1 1s; sexuality). Type material exarnined.-USNM 119490 ， 98.0 mm ，Me Kok at Chieng R む， 2 March 1934 ，H 119498 119498 (印 pa 紅ra 瓜type H. lobatus); USNM 119492 ，3: 69.2-101 mm ，Bung Borapet ，20 Nov. 1923 ，H.M. Smith (paratypes H. lobatus); USNM 119493 ，2: 60.5-6 1. 0 mm ，Menarn Chao Phraya Phraya at Bangsai ，Centr a1 Thailand ， 27 Nov. 1923 ，H.M. Smith (paratypes H. lobatus); 190 190 TYSON R. ROBERTS

USNM 119494 ， 73.5 mm ， Pasak R. at Dha Luang ，Central Th ailand ，20 Aug. 1923 ，H.M. Smi 血 (p 紅 atypes H. loba ω s); USNM 119491 ， 68.0 mm ，Menam Chao Phraya ，Paknampo ， Cen 住a1 Siam ， 19 Nov. 1923 ，H.M. Smi 血 (p 紅 atypes H. lobat 凶). Non-type Non-type materi a1 examined. -C AS 91769 ，8: 56.9-102 mm ，Menam Kok at Th a Ton and up to 5 km downstream ， 15 May 1990 ，T. R. Roberts; CAS 91768 ，Menam Chi at Maha Chana Chai ， 13 March 1991 ，T. R. Roberts; CAS 91770 ， 25: 41. 7-85.6 mm ，and CAS 91771 ， 24: 43.3-88.1 mm ， Yasothon market [=Menam Chi] ， 13 March 1991 ，T. R. Roberts; CAS 91767 ， 4: 43. 4- 50.5 mm ，Se Kh one at Stung Treng ，6 Feb. 1994 ，T. R. Roberts.

Diagnosis. Diagnosis. A small species ，1紅 'gest recorded specimen just ov ぽ 100 mm in s刷 d制 length; length; snout often but not a1 ways strongly projecting; head relatively small ，especi a1 1y comp 紅 'ed to 出e closely related and somewhat 1紅 ger species C. siamensis; branched dors a1 fin fin rays 8; scales in later a1 series 32-33; vertebrae usu a1 1y 32; differs 食om all other species of of Cirrhinus so f:紅部 known in being a protogynous hermaphrodite (see below).

Comments. In Jun e- July 19931 observed reproductive migrations of Cirrhinus 組 d o白er sm a1 1 cypri 凶d species in 血.e Mekong mainstream just below Le e Pee Waterf a1 1s 加 southem Laos. It was immediately apparent 白紙 there were two ex 位emely abundant sm a1 1 species species of Cirrhinus. One of them was identified as C. siamensis and the other on1 y 出 C. C. sp. by ROBERTS ， 1993. Cirrhinus sp. was subsequen t1 y identified as Cirrhinus lobatus in in ROBERTS & BAIRD ， 1995. Th e type materi a1 of H. lobatus comprises 白ree species. Only 白eωpotypic paratype from the Me Kok (U SNM 108091) is conspecific wi 白白e holotype. 百le r邸 t of the p紅 atypes ，a1 1 from the Chao Phraya basin ，a1 1 examined and reidentified by me ，釘 e C. caudimaculatus caudimaculatus and C. siamensis.

Biological Biological observations. Sexu a1 1y active Cyprinidae 紅 'e among the easiest of fishes to to sex accurately. Gr avid fem a1 es ， with grea t1 y distended abdomens ，readily discharge 句 gs with slight finger pressure. Th em a1 es discharge 凶 1勾 white 凶 lt equally readily. In live live or very f詑sh fish it is often possible to sex accurately a1 1 sexu a1 1y active individu a1 s， even even in very sm a1 1 species ， without resort 泊g to microscopic examination. Mi gration of reproductively active C. lobatus was observed 泊 Jun ←Ju1 y 1993 at B 組 H 組 g Kh one ，a fishing vil1 age some 4 km below Lee PI 田 Waterfalls (the Spirit Trap) on 白e mainstream Mekong Ri ver in southem Laos (near the border with Cambodia). Ne 紅 Iy a1 1 individu a1 fish caught at 也is time were in spaw 凶ng condition. Tw o samples were sexed ， one of 83 individu a1 s taken on 22 June 1993 and the other of 302 individu a1 s taken on 1 July 1993. 百lese were caught in wide-mouthed basket traps or kah (s 閃 ROBERTS ， 1993) 叩 d were ex 創世田d wi 由泊 an hour remov a1 of 血e traps from the water. Results Results were reported in ROBERTS & BAIRD (1996). 百le sm a1 1est individu a1 s in both S鉱 nples ，45-70 mm Sl ， were nearly a1 1 gravid fem a1 es. A few fish within 血is size range could could not be sexed; some of these appe 紅 ed to be malnourished or otherwise 泊 poor condition. condition. In the 22 June sample a1 1 ripe m a1 es were over 80 mm SL ，and 泊由e1 July sample sample a1 1 were over 70 mm SL. In bo 由 S佃 lples the few fish over 90 mm SL were a1 1 ripe ripe m a1 es. In both samples ripe fem a1 es outnumbered ripe m a1 es by more 出an3to 1. 官 le data suggest 血at C. lobatus is a protogynous (“ fem a1 e frrst") hermaphrodite. Distribution. Distribution. Cirrhinus lobatus is known on1 y from the Mekong basin. SYS1 芭MA 百CS AND BIOLOGY OF CIRRHINUS 191

C 加 'hinus lu new name Fig.9

Cirrhina Cirrhina latia ，Ann andale ， 1918: 46 (1 叫e Lake). Crossocheilus Crossocheilus horai B 叩紅白cu ， 1986: 153 ，fig. 11 (type l'O cality Inle Lak e).

Type materi a1 examined.-USNM 191451 ， 121 mm ， Inle Lake ， Feb. 1958 ， C.J. Sh 'O ntz (h 'O l'O type C. horai); USNM 288034 ， 3: 101-112 mm ，same c'O llecti 'O n data 槌 USNM 191451 191451 (p 釘 'atypes C. horai).

N 'O materi n-type a1 examined. -C AS 81548; 24: 95.9-155 mm ，Inle Lake ，25 Feb.-7 March 1994 ， T.R. R 'O berts.

Dia 伊 osis. Cirrhinus lu is 'O ne 'O f the m 'O st distinctive members 'O f the genus Cirrhinus ， characterized characterized by an excepti 'O na1 1y terete b'O dy f'O rm; largest kn 'O wn specimen 155 mm; stral r'O stral barbels present ，m 凱 ill 紅 y barbels absent; d'O rs a1 fin branched rays 9; sc a1 es in later a1 series 39 -4 1; transverse sc a1 er 'O ws 6/1/4; circumpeduncular sc a1 es 1ι18; vertebrae 25-27+13-15=38(1) ，39(5) ，40(7) ，41 (1).

Discussion. Discussion. In his 'O rigi na1 descripti 'O n，Ban 紅白cureferred 出is sp 即 ies t 'O Crossocheilus. Crossocheilus Crossocheilus and Cirrhinus 釘 e (at least superfici a1 1y) very similar mo 中h'O l'O gic 叫ly ，制d may be cl 'O sely related t 'O each 'O ther. Evidently the m 'O st imp 'O rtant difference between them inv 'O lves the r'O s回 1 cap. In Crossocheilus (and in sever a1 cl 'O sely related labe 'O in gene ra1泊 cluding Epalzeorhynchos ，Garra ， and Paracrossocheilus) 血， e rostr a1 cap is enl 紅 ged and m 'O re 'O r less deeply furr 'O wed by a series 'O f papill 'O se funbriae which project from its 'O ra1 m 釘 gin (see ROBERTS ，1982 ，fig. 8; BANARESCU ，1986 ，figs. 1-2). In Cirrhinus 血e r'O s位a1 cap is n'O tn 'O tably enlarged; it either has n 'O papill 'O se fimbriae at a1 1，'O r has weakly devel 'O ped papill 'O se fimbriae that d 'O n'O tf 'O rm furr 'O ws. Th e difference is very 紺政ing ，and is is related t 'O feeding behavi 'O r. Crossocheilus are a1 ga1 br 'O wsers ，scrap 泊g a1 gae fr 'O m hard

subs 回 te (e.g. r'O cks ，l'O gs) by me 佃 s 'O f the r'O str a1 cap. Cirrhinus ，'O n the 'O ther hand ， feed mainly mainly by me 姐 s 'O f sucti 'O n，and d 'O n'O t use 血 er 'O str a1 cap t 'O scrape f'O'O d 企om 血 e substrate. substrate. Th er 'O str a1 cap 'O f C. lu d'O es have a papill 'O se m 紅 gin ， but the cap is much sh 'O rter and less less 'O verhang 泊gthan 泊 Crossocheilus ，and 血 e papillae are n'O t arranged 'O n well devel 'O ped fimbriae fimbriae which pr 'O ject 合eely 企om its 'O ra1 margin. Th e extent and devel 'O pment 'O f 血 e r'O s仕組 papillaein 由is species 'O ccurs n'O rm a1 1y in s'O me 'O ther species assigned t 'O Cirrhinus ， such such as C. fulungee and C. ariza. In additi 'O n，individu a1 specimens 'O f still 'O ther Cir 時 inus species ，including C. lobatus ， have c'O mparably devel 'O ped r'O s位a1 papillae.

Banarescu Banarescu suggested 由at 白e species might be 血e same species 鎚 sp 民泊lens identified 部 Crossochilus latius fr 'O m 'O ther parts 'O f Myanmar ，but 血 is very d'O ub 伯11. In P紅白ul 紅， he listed Crossocheilus latius 'O f Vincigue 町a 企om Meetan [Tenasserim pr 'O vince] ，Myanmar 部 p'O ssibly 出e same species. 1 have examined Vinciguerra's specimens (MSNG 17349 ， 2: 2: 108-114 mm) and find 白紙 they ar 芭 very typic a1 Crossocheilus ， with a large ，br 'O ad r'O strum and well devel 'O ped rostr a1 fimbriae and papillae.

Nomenclatural note. Transfer 'O f Crossocheilus horai Banarescu ， 1986 t'O血 e genus Cirrhinus Cirrhinus makes it a ju 凶'O rh 'O m 'O nym 'O f Cirrhinus horai L 紘 shmanan ， 1966. Because it has has n 'O existing replacement name ，1 have c'O ined the new replacement name C. lu. 192 TYSON R. ROBERTS

Etymology.τ 'h e new scientific name C. lu is from the Intha vemacular n創即時a lu ， reported reported by Ann and a1 e， 1918: 38) ， and still in use for 出is species at the time of my visit to to ln1 e in 1994.

Di stribution. 百lis sp 田 ia1i zed sp 民 ies app 紅 'ently occ 町 s only 泊 ln1 e Lake ，a wonde 曲 11 place place jus t1 y famous for its beautiful and varied endemic fishes (Ann an da1 e， 1918).

Ci"hinus microlepis Sauvage 1878 Fig.l0

Cirrhina Cirrhina microlepis Sauvage ， 1878: 236 (type loc a1 ity Me-Kong at Tma- Kr e). Cirrhina Cirrhina aurata Sauvage ， 1878: 236 (type loc a1 ity Pn om-Penh). La beo (μ beo) aurovittatus Sauvage ， 1878: La beo pruol Tir 却し 1885; Kottelat ， 1986: 14 (designation of lectotype; placed as synonym of of C. microlepis). Type materi a1 examined. 一MNI 王N 3849 ， 118 mm ，Pn om-Penh ，1874 ，Ju l1i en (syntype C. aurata). aurata).

Non 勾 pe materi a1 examined. 一MNI 町 1987-127 ，2: 193-195 mm ，Cochinchine ，1921 ， de Kr empf; USNM 104935 ， 2: 122-146 mm ，Chao Phraya at Nontaburi ， 16 Dec. 1926 ，H.M. Smith. Smith.

Diagnosis. Diagnosis. A large sp 田 ies wi 血 very small sc a1 es ，no barbels ， and distinctive coloration. Juveniles Juveniles silvery wi 由 red caud a1 fin ，larger fish with head and body violaceous ，rosy ， or bluish bluish and caud a1 fm dusky. Sc a1 es in later a1 series 5ι60 ，transverse sc a1 e rows 13/118; dors a1 fin rays iv 11-12-!-; vertebrae 27+13 =4 0(3). 2 Distribution. Distribution. Cirrhinus microlepis occ 町 s in the Chao Phraya 組 d Mekong basins of Th ailand ，Laos ，Cambodia ，and Vietnam. A record 企'O m Chantabun in southeast Th ailand (Smi 出， 1945: 164) is doubtfu l.

Ci"hinus mo 前'o rella (Valenciennes in Cuv. & Val. ， 1844) Fig.11-12

Le uciscus molitorella V a1 enciennes in Cuvier & V a1 enciennes ， 1844: 359 (type loc a1 ity China; China; based on pre 吋.o usly unpublished painting made 泊 Canωifi for Dussumier deposited in in Biblioth 句ue Cen 紅白， MNI 王N ，Ms 5037 ， reproduced here as Fig. 11). Leuciscus Leuciscus chevanilla V a1 enciennes in Cuvier & Valenciennes ， 1844: 358 (type loc a1 ity not given ， but based on a Chinese painting ，a copy of which is 泊出e Biblioth 句ue Centr a1 e， MNHN ，Ms 515 ，document XVII C 44). Cirrhina Cirrhina chinensis Gun 血er ， 1868: 36 (type loc a1i ty China). La beo (Diplocheilichthys) gamieri Sauvage ， 1884: 210 ， pl. 7 ，fig. 4 (type loc a1i ty environs de de Hanoi); B 佃紅白cu ， 1972: 254 ，fig. 3 (design 副 on and figure of lectotype). La beo jordani Oshim a， 1919: 204 ， p l. 49 fig. 3 (type loc a1 ity hatchery at Shori ， Formosa; stock stock in 住'O duced from South China). SYSTEMATICS AND BIOLOGY OF CIRRHINUS 193

Cirrhina Cirrhina melanostigma Fowler & Bean ， 1922: 4 ，fig. 1 (type locality Koroton ， Formosa). La beo collaris Nichols & Pope ， 1927: 362 ，fig. 28 (type locality Nodoa ， Hainan). La beo pingi Wu ， 1931: 20 ，fig. 3 (type locality presumably Ming Ri ver ， Foochow). Osteochilus Osteochilus prosemion Fowler ， 1934: 116 ，figs. 6ι7 (type locality Chiengmai [Chao Phraya Phraya basin]); Fowler ， 1937: 183 (Kemarat ， Mekong). Osteochilus Osteochilus spilopleura Fowler ， 1935: 115 ，figs. 52-3 (type locality Srisawat ，Me k1 0ng basin). basin). Osteochilus Osteochilus macrosemion Fowler ， 1935: 116 (type locality Srisawat ，Me k1 0ng basin). La beo stigmapleura Fowler ， 1937: 202 ，figs. 167-70 (type locality Kemrat=Mekong R. in Thailand); Thailand); Bohlke ， 1984: 91 (placed as synonym of Cirrhinus prosemion). Cirrhinus Cirrhinus mrigala prosemion ，Banarescu ， 1983: 14. Cirrhinus Cirrhinus macrosemion ，Sodsuk ， 1988: 31. Type material examined.-BMNH 197 1.1 2.3 0.2 ， 143 mm ，China ，J. R. Reeves (holotype C. C. chinensis; designated neotype C. molitorella below); MNHN 188 4- 81 ，208 mm ， Hanoi (Tonkin) ，Harmand (l ectotype L. garnieri); USNM 84168 ， 3: 13 0- 139 mm ，Koroton ， Formosa ， H. Sauter (holotype and p紅 'atypes C. melanostigma); AMNH 8399 ， 197 mm ， China ，Kwangtung ， Hainan Tao ，Nodoa ，1922-23 ， C. Pope (holotype L. collaris); ANSP 59095 ，9 1. 8 mm ，Chieng Mai ， 15-23 Dec. 1932 ，R. M. deSchauensee (holotype O. prosemion); prosemion); ANSP 5909 6- 97 ， 2: 85.3-89.9 mm ，same da ぬ as ANSP 59095 ，paratypes O. prosemion; prosemion; ANSP 60808 ， 159 mm ，Srisawat ， July 1934 ，R. M. deSchauensee (holotype O. spilopleura); spilopleura); ANSP 60809 ， 138 mm ，Srisawat ， July 1934 ，R. M. deSchauensee (holotype O. O. macrosemion); ANSP 68169 ， 76.0 mm ，Kemrat ，1936 ，R. M. deSchauensee (holotype L. L. stigmapleura); ANSP 68170 ー78 ，9: 36.0 ー73.8 mm ，same data 部 ANSP 68169 (para 旬pes L. L. stigmapleura). Non-type Non-type material examined. -C HINA: AMNH 37020 ，2: 90.5-130 mm ，Foochow ， Fukien Pr ov. ，March-Sep t. 1926 ， C. Pope; MNHN B. 2643 ，5: 12 6- 140 mm ， Kouang-Si; MEKONG BASIN: CAS 79171 ， 145 mm ，Mekong R. ne 紅 Ban Ha Bia ，3 0-4 0 krn NW of Chiang Kh an on highway 2186 ， 12 March 1990 ，T. R. Roberts; CAS 79172 ， 2: 136 ー147 mm ， Nakom Phanom market ，4- 7 March 1991 ，T .R. Roberts; CAS 79173 ， 2: 107-113 mm ， rapids rapids in mainstream of Mekong R. about 12 km S of That Phanom ，6 April 1991 ，T. R. Roberts; Roberts; CAS 79170 ， 2: 64 .4ー 76.6 mm ，mouth of Huay Ngao where it flows into Mekong R. R. about 30 krn SE of Chiang Kh ang ， 12 May 1990 ，T. R. Roberts; MEKLONG BASIN: CAS 79174 ， 10: 76.5-144 mm ，Huay Kh a Kh aeng at Greung Gr ai ，Huay Kh a Kh aeng Wildlife Wildlife Sanctuary ，Me k1 0ng basin ， 21 Jan. 1991 ，T. R. Roberts; CAS 79175 ， 2: 82.8 ー93.7 mm ，Huay Sangkalia 7 krn N of Sang k1 aburi on road to Chedi Sam Ong ， 11 Feb. 1989 ， T. R. Roberts; TAPI BAS 別: CAS 81656 ， 2: 188-194 mm ， Fish landing at S end of Chiao Lan Reservoir ，28 Feb. 1989 ，T. R. Roberts. Di agnosis. A moderately large and distinctively colored species of Cirrhinus ，attaining at at least 40 cm standard leng 由; rostral barbels relatively well developed ， length half or more of eye diameter; branched dorsal fin rays 11-15; lateralline scales 38 -4 0; transverse scale scale rows 8/1/5; vertebrae 23-25+11-13=34-38. Cirrhinus Cirrhinus molitorella differs from all other species of the genus in having more or less discrete discrete marks on each scale on the upper ，middle ， and sometimes lower p紅 ts of the body. 194 TYSON R. ROBERTS

Th e marks 紅 'e f'O rmed by melan 'O ph 'O res c'O ncen 回 ted near the middle 'O f the anteri 'O r scale shield shield where the scale is partially 'O verlapped by the free dis ta1 marg 泊'O f the scale in 合ont 'O f i t. Such marks ，visible in m 'O st 合'eshly caught and recently preserved specimens ， are s'O me 也nes 泊.t ensely devel 'O ped ，giv 泊 g the fish a markedly reticulated appe 紅組問. Cirrhinus molitorella molitorella 'O ften has a very intense humeral mark which is vivid bluish-green in life. In 'O ther Cirrhinus the humeral marks 紅 'e black 'O r bluish-blac k.

Commeots. Pぽ t 'O f the syn 'O nymy 'O f C. molitorella given ab 'O ve was w 'O rked 'O ut by Banarescu ， 1972: 253-54. Use 'O f the juni 'O r syn 'O nym C. chinensis by Banarescu and s'O me 'O ther auth 'O rs f'O r this species presumably is based 'O n the n'O ti 'O n 白紙白e se 凶'O r syn 'O nym C. C. molitorella is an 'O men nudum (see Banarescu ， 1972: 251-52). Valenciennes's 'O riginal acc 'O unt ，alth 'O ugh based 'O n an unpublished figure ，definitely includes a descripti 'O n S 'O that the the name C. molitorella is available. Th e 'O riginal i1l us 位ati 'O n 'O f C. molitorella (Fig. 11) is 'O ne 'O f 24 fish paintings made f'O r J.J. Dussumier in Cant 'O n in 1820. It is clearly 'O fa Cirrhinus ，and is an excellent likeness likeness 'O f the species called C. chinensis 'O r C. molitorella by vari 'O us auth 'O rs. In 'O rder t 'O rem 'O ve 紐 yp 'O ssible d'O ubt as t'O血 .e identity 'O f 白e species represented by the well- kn 'O wn and available name C. molitorella ，血 eh 'O l'Oザ pe 'O f C. chinensis 但MNH 197 1.1 2.30.2 ， 143 143 mm) is hereby designated ne 'O type 'O f C. molitorella. Wild and cultivated st 'O cks 'O f C. molitorella di 能 rc 'O nsiderably 泊 appearance and behavi 'O r. Wild st 'O cks 紅 e str 'O ngly migrat 'O ry ， while the cultivated st 'O cks pr 'O bably have l 'O st the migrat 'O ry behavi 'O r. Cultivated st 'O cks (and p 'O ssibly als 'O wild fish) from China typically typically have discrete marks 'O n scales very b'O ld ， as 'O pp 'O sed t 'O fish 企om wild st 'O cks in S 'O utheast Asia. On 'O ne 'O ccasi 'O n 泊 Bangk 'O k1 'O bserved feeding behavi 'O r 'O fa number 'O f wild wild fish 企om Th ailand and cultivated fish fr 'O m China in a1 紅 'ge cement h'O lding tank with very very clear water ab 'O ut 1. 5m deep. 百le cultivated fish were readily distinguished by their darker darker c'O l'O rati 'O n. Wh en fed' with pellets ， the wild fish dashed ab 'O ut ， catching individual pellets 泊 midwater ， but were n'O t interested in the pellets after 出ey reached the b'O tt 'O m. Th e cultivated fish did n'O t chase the pellets in midwater ，and '0凶 y fed 'O n them after 白ey settled settled 'O n 血 .e b'O tt 'O m 'O f the tank.

Distribu 組00. Cirrhinus molitorella is widely dis 凶buted in s'O uthern China (where it is is extensively cultured and has been widely transp 'O rted). It 'O cc 町 s na 加rally in much 'O f Th ailand and In d'O china ，including the Mekl 'O ng and middle and l'O wer Mek 'O ng basins. It app 紅 ently has n'O t been c 'O llected in the Cha 'O Phraya basin. In tr 'O duced t 'O Hainan ，H 'O ng K 'O ng ，Taiwan ，Singap 'O re ，In d 'O nesia. Aquacultural st 'O ck rep 'O rtedly has been in 位oduced recently recently 企'O m Chin a-in to- the Mek 'O ng basin in La 'O s. Sp 巴:cimens rep 'O rted here 企om 由e Tapi basin (peninsular Th ailand) p 'O ssibly represent 白e furthest s'O uthward range 'O f any naturally 'O ccurring Cirrhinus. 百 e Tapi is als 'O the s'O uthernm 'O st kn 'O wn l 'O cality f'O r the 1紅 ge bagrid catfish Mystus microphthalmus (se 凶'O r syn 'O nym 'O f M. wyckoides). B 'O th species might 'O we their presence in the Tapi t 'O intr 'O ducti 'O n， but n 'O rec 'O rds 'O f 白is have c'O me t 'O my attenti 'O n and l'O cal 百凶 fisheries fficers 'O fficers interviewed by me believe 白紙 b'O th species 'O ccur na 旬 rally in the Tapi. SYSτ 'E MATICS AND BIOLOGY OF CIRRHINUS 195

Cirrhinus Cirrhinus ornatipinn お new species Fig. Fig. 13

Holotype. -C AS 91756 ， 69.7 mm ，roadside ditch on highway 24 at km 150 marker ， 179 km by road E of Nakom Ratchasima ，27 May 1991 ，T. R. Roberts.

Paratypes.-CAS 91762 ， 6: 51. 5-6 7.3 mm ，collected with holotype; CAS 91757 ， 27: 37.3 0 60.1 60.1 mm ，i凶 gation canal about 20 km S of Phibun M 阻 gs 油 an ， 16 Sep t. 1990 ，T. R. Roberts; Roberts; CAS 91760 ， 9: 55.7-90.5 mm ，roadsidecnals 5-30 km S of Phibun Mangsahan ， 16 16 Sep t. 1990 ，T. R. Roberts; CAS 91759 ， 15: 58.6 ー73.2 mm ，Nakhon Phanom market ，2 6-- 31 31 May 1990 ，T. R. Roberts; CAS 91758 ， 4: 83.7-93.9 mm ，Nakhon Phanom market ，4 ー7 March 1991 ，T. R. Roberts; ANSP 172554 ， 2: 57.1 -61. 7 mm ，and USNM 331212 ， 2: 63.8- 76 .5 mm ， Phibun Mangsahan market ， 15 Sep t. 1990 ，T. R. Roberts.

Non-type Non-type material examined.-CAS 91761 ， 26: 49. 仏-6 7.3 mm ， Phibun Mangsahan market ， 15 15 Sep t. 1990 ，T. R. Roberts. Di agnosis. Perhaps the smallest species of Cirrhinus (l argest known specimen 90.5 mm standard length); readily distinguished by its small head ， short blunt snout ， very deep caudal caudal peduncle ，組 d usually having the pectoral ，pelvic ，anal ， but not the caudal fin red ， reddish-orange ， or at least rose-tinted; the pelvic and anal fins often tipped with white; barbels barbels absent; dorsal fin branched rays 8; scales in lateral series 34- 35; tr 組 sverse scale rows 6/ 1/ 4; vertebrae 21-22+11-13=33(14) ，34(10) ， or 35(2). In all other species of Cirrhinus Cirrhinus the pectoral fins are colorless ，dusky ， or at most pale pink or rose-tinted.

Etymology. 百le specific n創 ne “omatipinnis" (Latin) refers to the colorful fins. Di stribution. Cirrhinus ornatipinnis is known only from the Middle Mekong basin of of northeastem Thailand (Isan). Unlike other Cirrhinus familiar to me ，it seems to be res 住icted to m 釘 ginal or tempor 訂 y habitats ，including roadside canals ，where 1 observed it it on numerous occasions. 1 have never seen it in flowing water habitats of the Mekong and its 凶butaries where C. siamensis ， C. lineatus ， and C. lobatus usually occ 町. So far as as known it does not occur sympa 凶 cally with other species of the genus.

Cirrhinus Cirrhinus rubirosT お new species Fig. Fig. 14

Holotype. 一一 MNHN 1992.1043 ， 140 mm ， Tenasserim Ri ver ， 12 March 1992 ，T. R. Roberts. Paratypes.-MNHN 1992.1044. 2: 11ι118 mm ，collected with holotype; ZRCS 22933 ， 138 138 mm ， Tuler Kloh ， Tenasserim basin ，March 1992 ，T. R. Roberts; CAS 91753 ， 110 mm ， backwater backwater at huge rocky ledge in Tenasserim Ri ver midway between Htee-tah 佃 d Baowashung ， 12 March 1992 ，T. R. Roberts; CAS 91754 ， 106 mm ， Tenasserim Ri ver upstream 仕om Htee-tah ，8-9 March 1992 ， T .R. Roberts; CAS 91755 ， 172 mm ， Tenasserim Ri ver between Htee-tah and Baowashung ，March 1992 ，T. R. Roberts. Diagnosis. Diagnosis. Cirrhinus rubirostris differs from all other Cirrhinus in having both sexes with with well developed rostral tubercles ，and rostral tubercles and snout pinkish or rosy red. 196 196 TYSON R. ROBER.τ 百

Ros 位al barbels absent ，m 砿 ill ぽ yb 紅 hels presen t. Dorsal fm wi 血 10 branched rays. Scales in in lateral series 37 ，predorsal scales 12-13 ，位佃 sverse scale rows 7/116 ，circumpeduncular scales scales 21. Vertebrae 22+13=35 (3). Palate Palate anteriorly with numerous delicate folds ， followed by palatall 創 nellar organ with 6 pairs of tuberculate lamellae (Fig. 1). Roof of branchial chamber wi 血 about 100 組 teroven 佐必 ly directed tuberculate digitiform papill 鵠. Gi ll rakers on frrst gill 紅 .ch 43. Head and body iridescent gree 凶sh and bluish dorsally ，yellowish laterally ， and silvery white white ventrally. Humeral mark bluish black. Al l fins reddish or pale rose except anal fin; anal anal fin very pale rose or whitish. Dorsal half of iris red ，rest white.

Etymology. Etymology. Th e sp 田 ies name rub 註os 凶 s，a La出 adjective ，refers to the red color of 出e ros 位al tubercles and of the snout generally; it also refers to the Karen n釘ne for this species ， niya gwoh nadee ，“ 'red-nosed fish".

Distribution. Distribution. Cirrhinus rubirostris is known only from 血e Tenasserim Ri ver basin of of southeastem Burma.

Cirrhinus Cirrhinus siamensis (Sauvage 1878) Fi g.1S

Morara siamensis Bleeker ， 1865a: 35 (nomen nudum); Bleeker ， 1865b: 175 (nomen nudum); nudum); Sauvage ， 1881: 164 ，187 ，pl. 6 ，fig. 2 (type locality Bangkok; figure erroneously labelled labelled Cirrhina jullienz). Aspidoparia Aspidoparia siamensis ，Smi 也， 1945: 124 (doubt expressed 回 to identity of genus and of species). species). ?η lognathus siamensis 何回 Morara siamensis Bl 即:k er) de Beaufort ， 1927: 9 (type locality

Payao sw 創 np [=Mekong basin] and Chao Phraya at Lopburi). Tylognathus Tylognathus brunneus Fowler ， 1934: 131 ，figs. 87-88 (type locality No 凶 1 Si 倒的. ?Tylognathus ?Tylognathus en 伽 ema Fowler ， 1934: 134 ，figs. 101-102 (type locality Bangkok). Cirrhinus Cirrhinus marg 初ipinnis Fowler ， 1937: 173 ，figs. 108-109 (type locality Phitsanulok=Chao Phraya Phraya basin). Cirrhinus Cirrhinus sauvagei Fang ， 1942: 168 (type locality Me-Kong). Crossocheilus Crossocheilus reba ，Smi 血， 1945: 270 ー71 (Chao Phraya). ?Crossocheilus ?Crossocheilus thai Fowler ， 1944: 49 (type locality Bangkok; aquarium spec 泊施n).

Type material exax 凶ned. 一 M Nl王 N 1839 ， 4: 77.5-92 .3 mm ，Bangkok ，1862 ， Bocourt (syntypes (syntypes M. siamensis); MNl削 8598 ， 93.0 mm ，and MNHN B. 2961 ， 7: 76.ι96.8 mm ， Cochinchine ，Jullien (syntypes C. sauvage l); ANSP 58369 ， 72.3 mm ， North Siam (holotype T. T. brunneus); ANSP 59092 ， 47.7 mm ， Silom canal ，Bangkok (holotype T. en 伽 ema); ANSP 68069 ， 115 mm ，Phitsanulok ，百 lailand (holotype C. marginipinnis); ANSP 71336 ， 78.7 78.7 mm ，Bangkok ， A. Viehoever (h olotype C. thai). Non-type Non-type material exaxruned.-MEKONG BASIN: MNHN 9546 ， 8: 79.8-98.5 mm ， Cochinchine ，1874 ，Jullien; CAS 91751 ， 109 mm ，Pak Mun ，May 1991 ，T. R. Roberts; CAS 91750 ， 3: 79.8-83.7 mm ， Yasothon market [Menam Chi] ， 13 March 1991 ，T. R. Roberts; CHAO PHRA YA BASIN: MNl町 A. 5080 ， 4: 84. 0- 94.8 mm ，Bangkok 1882 ，H 紅 mand; SYS T EMAT ICS AN D BI O LOGY OF C IRR /-I INUS 197

Fi gur e 1 1. Cirrhi l/lI s 川 o!i lorell α. P 山 nl ll1 g0 11 引 Ik C0 111111i ss io n巴db y D uss u l11 ier in Ca nt o n

A rchi ves MN トIN 司 D OC U111 enl 5037 (ph OlO co pyri g hl Bib l. Ce nlr ale，MNHN ，1 994)

Fig 山 巴 12. Cirrhi l/ lI s 1/ /{)li lO rell" トIu ay K hは Kh ιleng at Gr ucng Gra i

F ig lll 巴 13. C irrhi l/ II .¥ ο川 /((Iipi l/I/ i.¥. 69.7 111111 ho lolype，Thai land (Me kon g ba sin )

F igure 14. Cirrhi l/ II .¥ m iJ im .¥lr is， ho lo lype，1 40 111111 ‘T enassc ri l11 R ive l 19 8 T YSON R. R OsER TS

Fig Ul 巴 15. Ci rrhill l/ ss iolll ell sis ，1 35 111111 ，Na ko rn Sawa n (C h ao Phray a)

礼 R ¥

l 〆 ~ ~ -~ ~ ~ ~

Figllr e 16 “Ev 巴r-c hangin g leaders hip str al巴gy " to find path of leas t res istan ce by sc hoo lin g fis h l11 igr atin g lI pstl 巴al11 pa st waterfa ll s a nd rap id s.S impliti ed dia gram ba s ed on obs巴rv巴d or Sll Sp巴ctecl behav iol

of of Ci rrhill l/ s loba ll.l s at K hone Fa ll s on th e 111 巴ko ng l11 ain slr ea l11 in sOll th e rn Lao s. AJTOW S rep rese nt Khon e fall ltlin e. A rr ows pointin g down str 巴al11 repr es 巴nt hi gh ancl oth erw is巴 iI11 passab le barri ers to

up str ea l11 fi sh 日10V 巴111 巴nt s. A rro ws pointin g c1 0wn str ea m or in bot h c1 irection s incli ca te pla c巴s fi sh ca n pa ss lI pstr ea l11 ，S OI11 巴11111 巴s a icl ecl by cO llnt erclln 巴nt s SYSτ 百MA' 百 CS AND BIOLOGY OF CIRRHINUS 199

KUMF 656 ， 6: 61 -6 7 mm ， Sikuk Ri ver at Ban D 佃 g，26 Nov. 1923 ，H.M. Smi 白 (orig. id. Crossocheilus Crossocheilus reba by Smith); CAS 79177 ， 3: 107-118 mm ，Uttaradit market (Nan R.， Chao Phraya basin) ，9J 佃 U 紅 y 1989 ，T. R. Roberts; CAS 91752 ， 120 mm ，Nakom Sawan market ，5 -6 Feb. 1989 ， T.R. Roberts; CAS 79178 ， 15: 78.2-116 mm ，Nakom Sawan market ，4 March 1990 ，T. R. Roberts; BANGPAKONG BASIN: CAS 79179 ， 74.5 mm ， Menam Bangpakong ne 紅Pr achinburi ，6 Dec. 1990 ，T. R. Roberts; CAS 91749 ， 4: 57. 4- 65.2 65.2 mm ，Menam Bangpakong at B 佃Kh ao Cha-chan ， 19 km S of Sa Kaeo on highway 317 ， 23 March 1989 ，T. R. Roberts. Di agnosis. Cirrhinus siamensis is a medium sized species (to 20 cm) ，usu a1 1y with a much larger and broader head than sma1 1er species such 部 C. caudimaculatus ， C. lineatus ， C. C. lobatus and C. soi. Freshly collected specimens of these species can be identified readily ， but difficulties arise with older preserved specimens. Further study is needed to discover discover better diagnostic characters. Most radiographed specimens of C. siamensis have 33-34 vertebrae ， while those of C. lobatus have only 32.

Discussion. Discussion. Confusion in 由e identification of 也is very common speci 邸 with C. jullieni jullieni (a relatively rare species) was caused by publication of the origin a1 figure of Morara siamensis mislabelled as C. jullieni (Sauvage 1881 ， pl. 6 ，fig. 2) ， while at 白esame 血ne no fi 思.rr e was published of the 住ue C. jullieni ， a relatively species. rare For many years 血.e type 叩配imensof “ 'Morara" siamensis were 凶 spl 邸 ed ， which further compounded 出e problem. 百le type spec 泊lens of T. brunneus 佃 d of C. marginipinnis 紅 'e a1 1 typic a1 1紅 ge- headed headed C. siamensis. 百le C. en 初tema holotype has a relatively sm a1 1 head ， not unlike C. C. lobatus ， but without a projecting or pointed snout 部 in many C. lobatus. Specimens of of C. siamensis 姐 d C. lobatus of 血is size 釘 e often difficult to distinguish. Sm a1 1 Cirrhinus ，known as pla soi in 百凶， were the dominant fishes in the massive fish fish migrations 由at formerly occurred 泊 cen 位叫百lailand. An evocative description of the migrations migrations is given by Smith ， 1945: 270 ー71. Samples identified by Smi 也 as C. reba have been been examined by me 佃 d reidentified as C. siamensis. Cirrhinus caudimaculatus may a1 so have been 佃 important (perhaps 白e most 加 port 姐 t) component of these migrations. Distribution. Distribution. Chao Phraya and Mekong basins of Th ailand ，Laos ，Cambodia ， and Vietnam. Vietnam.

BIOLO Gl CAL OBSERVATIONS ON CIRRHINUS LOBATUS

Cirrhinus Cirrhinus lobatus is endemic to 出e Mekong basin ， where it may be the single most abundant abundant fish species. It is certainly an ecologic a1 keystone species. Specimens have been collected collected in numerous 凶butaries as sm a1 1 as 2-3 m wide in widely separated loc a1 ities in τ'h ailand ，Laos ，and Cambodia. In terms of absolute numbers ，it undoubtedly is 血e most abundant abundant fish sp 民 ies in 白e major migrations 血at occur in 血e mainstream of the Mekong Ri ver below Kh one Fa1 1s every December-February and May-July ， where there is an important 紅白an a1 fisheries. It probably is 出es 泊.g le most important forage or prey species species for the many piscivorous fish sp 田 ies present 出ere ， and may a1 so be heavily preyed 200 TYSON R. ROBERTS

up 'O n by the local p'O p叫柑'O n 'O f 血 e Irr awaddy d'O lphin ， Orcaella brevirostris. 百1e cl 'O sely related related species C. caudimaculatus ， endemic t 'O the Cha 'O Phraya ， might f'O rme r1 y have been 血 es 泊 gle m 'O st abundant fish s戸 cies in 出 at basin ， but 出 is hyp 'O thesis lacks adequ a:旬 d'O cumentati 'O n. Cirrhinus Cirrhinus lobatus is 'O ne 'O f the lead species in the massive migrati 'O ns 'O f cypri 凶d fishes fishes m 'O ving up the Mek 'O ng mainstream 泊也e Kh'O ne Falls area. M 'O st 'O f the falls represent represent an unp 槌 sable physical 'O bstacle t'O曲 e migrat 'O rs ，but 出 ey can be av 'O ided by g'O ing up 'O ne 'O r tw 'O 'O f the large “h 'O'O" 'O r channels (m 'O st importan t1 yH 'O'O Sah 'O ng) and smallerpa 出ways ，血 us by-passing the impassable falls and m 'O re d出icult rapids. 百1e 紅 ea 泊 v'O lved has 'On1 y recen t1 y been mapped ， and the c'O mplexity 'O f the migrat 'O ry pa 血 ways d'O cumented (ROBERTS & BAIRD ， 1996). Ini tially 1 hyp 'O thesized 白紙由e migrating fishes w 'O uld “加旬matically" find 'O r remember the pa 伽明白紙w 'Ou1 d take them p制御falls and and rapids ， but 'O bservati 'O ns 'O f large numbers 'O f fishes repeatedly g'O泊 .g up dead ends and being being st 'O pped by 也 em 'O st impassable falls ca 凶 ed this hyp 'O thesis t 'O be rej 配飽d. Subsequent 'O bservati 'O ns suggested that 血 e migrating fish find their way up 姐 eam largely by 凶 al and err 'O r. Th e fish eviden t1 y are empl 'O ying a 抑制跡“出e ever-chan 仰 g leadership s回.t egy f'O r fmding the pa:血 way 'O f least resistance" ， which c組'O nly be success fu1 when very 1紅 'ge numbers numbers 'O f individuals are migrating. As the migrating fish m 'O ve ups 位e創 n ，and leading fish 釘 e bl 'O cked 'O r f ai1 t 'O fmd a way 'O nw 紅白， s'O me fishωrn back and find 'O ther rou 飽s. 羽田 the leadersbip 'O f the migrat 'O rs is c'O nstan t1 y changing ，until s'O me leaders 紅 e again successful successful in ge 凶ng past 'O bsta c1 es and take large numbers 'O ff 'Ol1'O wers with them (Figure 16). 官邸 s回 .tegy bears a str 'O ng resemblance t'O， and may be derived 企'O m ，血e ever- changing changing leadership s凶飽gy 'O f sch 'O'O ling fish inv 'O lved in predat 'O r av 'O idance. 百1e c'O njuncti 'O n 'O f ex 回 me abundance ， key ec 'O l 'O gical r'O le ，and migrat ぽ y behavi 'O r f'O und 泊 C. lobatus is ass 'O ciated with 戸田'O ci 'O us sequential hermaphr 'O ditism. Data previ 'O usly published indicate 也a: tall individu a1 s bec 'O me sexu a1 1y ripe fem a1 es at a sm a1 1 size size (and presumed c'O rresp 'O ndingly y'O ung age) ， with 'O nly a relatively few chan ，回 ng int 'O sexually sexually ripe m a1 es at much 1紅 ger sizes 侭.o BERTS & BAIRD ， 1996: 247-248 ，figs. 22-23). Such a sexu a1 s回旬gyh 部 n'O t been identified 泊 any 'O ther species 'O f Ci"hinus ，'O r indeed in 佃 y 'O ther Mek 'O ng species 'O f Cyprinid 鵠. It may a1 s 'O characterize the cl 'O sely related sp 田 ies C. caudimaculatus 血 the Cha 'O Phray a， but n 'O da 旬 is available f'O r 白紙 sp 郎 ies. A p 'O ssible explanati 'O nf 'O r the adap ぬ.ti 'O n 'O f sequenti a1 hermaphr 'O ditism in C. lobatus is 白 e size “size a1 1'O cati 'O nm 'O del" (Gms 乱闘， 1969). In 'O ne versi 'O n 'O f 也 is m 'O del ，'O rigin a1 1y based 'O n fish species ， the advantage t 'O 1紅 'ge ma1 esis 血a: t they d'O minate the aggressive c'O mpetiti 'O n f'O r fem a1 es. In C. lobatus ， the adv 阻 .tage in having sm a1 1 females and large m a1 es may inv 'O lve male (and fem a1 e) migrat 'O ry s回.t egy f'O r mating m 'O re than m a1 e aggressive c'O mpetiti 'O nf 'O r mates. 百1at is ， m a1 es may still be c'O mpeting f'O r fem a1 es ， but d 'O S 'O primarily primarily by being m 'O re “ capable migrat 'O rs". In曲 ec 'O ntext 'O f 出 e “ever-changing leadership" leadership" hyp 'O thesis 'O ffered here ， being be 恥 r migra 旬， rs may m 悶 n，泊 1伺.8 t in part ， being being better b'O血槌 leaders and as f'Ol1'O wers. A1 s 'O， m a1 es may migrate 'O ver l'O nger distances 血組 fe ma1 es ， and p釘 ticipate in m 'O re spawning 加 uts. Pr eliminary 'O bservati 'O ns suggest 也 at it is usu a1 1y 'O r a1 ways the sm a1 1er C. lobatus 白紙 紅 e 白 e migrat 'O ry leaders. Th e fem a1 es 血 us expend c 'O nsiderably m 'O re energy 也組曲e m a1 es fmding pa:血 ways past physi 'O graphic a1 barriers such as 出 e rapids and waterf a1 1s. It may a1 s 'O be 白紙 fem a1 es migrate relatively sh 'O rter distances bef 'O re spaw 凶ng ，出鉱山.ey SYSTEM A:百 CS AND BIOLOGY OF CIRRHINUS 201

出en spawn totally ， and drop out of the migration ， while the 1紅 'ger males may spawn with one one group of females ， then continue on upstream to spawn with other groups of females. Whil e the spaw 凶ng migrations of C. lobatus take place over hundreds or perhaps thousands of of kilometers ，it may be 血at the distances migrated by individuals 紅 'e much shorter ， with the the smallest (and most numerous) females migrating the shortest distances ， and with “new" migrators migrators (male as well as female) re 伊d紅 ly recruited along the route as 曲.e migr ョ， tion moves upstream. Such a migratory scen 組 o fits well with the observation of pr' 田∞ious sequential sequential hermaphroditism and 由 e hypothesized size adv 組 tage model for C. lobatus. An other implication is 白紙 the so-called long-distance migration of some riverine fish species species differ fundamentally from the 加 e long distance migrations of some birds and m 倍加 e fishes.

ACKNOWLEDG 島ffiNTS

My sincere thanks to 出 e followings curators and staff for their help: Roland Billard ， Jean Jean Claude Hureau ，Jeanine Able ， Marie Louise Bauchot ， Martine Desoutter ，組dMonique Ducreux ，島町HN; William N. Eschmeyer ，Tomio Iwamoto ，Dave Catania ， and Carl J. Fe 町紅is ，CAS; Oliver Crimmen ，Gordon Howes ， and Nigel Merritt ，B 島町H;H 佃 Nijssen and and Isaac Isbrucker ，ZMA; Eugenia Bohlke and Bill Smi 由-V aniz ，ANSP; Rema Davi ， Zo ological Survey of In dia ， Madras. Most of the specimen examination and writing was done in 血e Dep 紅 tment of Ichthyology ，California Academy of Sciences 加 d 泊出 e Laboratory ofIchthyology ，Museum National National d' Hi stoire Naturelle ，Paris. Many sp 田 imens reported upon were collected during fieldwork fieldwork supported by the Smithsonian In stitution; Smithsonian Tropical Research Ins titute: stitute In stitute for Biological Exploration; and 血e Committee for Research and Exploration of the the National Geographic S∞ iety (Gr ant 5141-93).

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