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).