Japanese Journal of Ichthyology 魚 類 学 雑 誌 Vol.l.25,No.11978 25巻1号1978年 Comparative Morphology and Interspecific Relationships of the Cyprinid Genus Puntius Yasuhiko Taki,Akari Katsuyama and Tousei Urushido (Received November 22,1977) Abstract Interspecific relationships were investigated in 23 species of the cyprinid genus Puntius on the basis of the features of infraorbital bones ,pharyngeal bones and their teeth, lateral folds and barbels on the snout ,and color pattern.Six phyletic groups were recognized in these fishes.In general,species belonging to the same group have overlapped or about equal geographic distributions.Between Southeast Asian and Indo-Ceylonese groups a more or less distinct gap is found in the degree of development of lateral folds and barbels . The results of this study indicate that Bleeker's(1863)division of Puntius into the subgenera Puntius,Barbodes and Capoeta is not acceptable from a phylogenetic viewpoint . catalogue number(in the case of catalogued Introduction specimen)are given in order. The genus Puntius Hamilton is the largest Southeast Asian species.P.altus(Gunther): in number of species of all cyprinid genera 25;41.5-105.0 mm;Thailand,Vietnam; in Asian tropics.Fishes of this genus occur IBRP 3112,3218,3283,6092,6485.P. throughout the region from Pakistan to schwanenfeldi(Bleeker):17;35.0•`163.0 southern China,inhabiting various types of mm;Thailand,Vietnam,Indonesia;IBRP fresh waters.The status of Puntius is ob- 3187,3197, 3282,6705.P.gonionotus scure;the delimitation and nomenclatural (Bleeker):14;70.0-182.5 mm;Laos,Thai- validity of the genus have remained unsettled land,Vietnam,Indonesia;IBRP 3180,3195, (Hora and Mukerji,1934;Smith,1945;Hoede- 3234,3289.P.orphoides(Valenciennes): man,1958;Myers,1960), largely owing to 15; 66.0-122.0 mm; Laos,Thailand,Viet- the scantiness in the knowledge of its inter- nam;IBRP 3079,3182,3285,4376,6223 . and infrageneric relationships. P.tetrazona(Bleeker):24;23.5-47.0 mm; The primary aim of this study is to in- Indonesia;IBRP 3184,6874.P. partipen- vestigate relationships among species of Puntius tazona Fowler:24;21.0-35.5 mm;Thai- on morphological bases,in order to provide land;IBRP 3068,3166,3261. P. pentazona data necessary for the assessment of evolu- (Boulenger): 11; 21.5-51.0 mm; Iodonesia; tionary lines within the genus.To this end, IBRP 3265. P. rhomboocellatus Koumans: internal and external characters were ex- 2; 27.5, 32.0 mm; Indonesia;IBRP 3190. amined on a total of 23 species. P.everetti(Boulenger):18;16.0-83.0 mm; 5 of 18,Indonesia;IBRP 6726,6754;13 Materials and methods of 18,aquarium-rised non-catalogued speci- This study was based primarily on speci- mens.P.lateristriga(Valenciennes):15; mens collected from Southeast Asia and de- 11.0-66.0 mm;Indonesia;IBRP 6720,6768. posited in the Institute for Breeding Research, P.fasciatus (Bleeker):18;20.0-88.5 mm; Tokyo University of Agriculture(abbrevia- 10 of 18,Indonesia;IBRP 3266,6618;8 tion for catalogue number,IBRP).It also of 18,aquarium-raised non-catalogued speci- dealt with specimens obtained from aquarium mens.P.binotatus(Valenciennes): 12; 49.0 fish suppliers in Tokyo and those raised in 82.5 mm;Vietnam,Malaysia;IBRP 6231,•` the laboratory.In the following list of 6736,6875.P.oligolepis(Bleeker):4;24.0 materials,number of specimens,their range •`32.0 mm;1 of 4,Indonesia;IBRP 6827;3 for standard length,collecting locality and of 4, non-catalogued specimens from an ―1― 魚類学雑誌 Japan.J.Ichthyol.25(1),1978 aquarium fish dealer.P.stigmatosomus anterior.Terms pertaining to pharyngeal Smith:8;37.5-68.0 mm;Laos,Vietnam; bones and their teeth used in this study are IBRP 3094,6231,6736.P.leiacanthus shown in Fig.2. (Bleeker).26;35.5,•`83.0 mm;Laos, Thai- Comparative observations land,Vietnam;IBRP 3038,3211,3235, 4772,5209,6353. Infraorbital bones.In all species under Indian and Ceylonese species.All of the study the infraorbital series consists of a following materials are non-catalogued speci- lacrimal,three infraorbitals,and a dermo- mens obtained from aquarium fish dealers. sphenotic (Fig.1).The antorbital is missing. P.chola(Hamilton):2;50.0,55.0 mm.P. The third and fourth infraorbitals are fused arulius(Jerdon):3;53.0,-64.5 mm.P. together into a large bone bordering the bot- tom of the orbit.This fused element is filamentosus(Valenciennes): 2;34.5,85.0 mm.P.titteya Deraniyagala:3;16.0•`27.0 termed infraorbital 3+4.In P.conchonius, mm.P.conchonius(Hamilton):9;41.0, P.cumingi,P.ticto and P.nigrofasciatus,the •`62.5 mm.P.cumingi(Gunther):2;37.5, infraorbital 3+4 is much broader than other 44.0 mm.P.ticto (Hamilton):8;30.5•`49.0 infraorbital elements,with the outer margin mm.P.nigrofasciatus(Gunther):12;25.0 greatly elevated(Fig.1A).Whereas in the •` 43.0 mm. remaining species the bone is only as broad Infraorbital bones were observed on alizarin- as the other infraorbitals(Fig.1B). stained transparent specimens.Identification Pharyngeal bones and their teeth.All of of the bones was based on Nelson(1969). the 23 species examined have three rows of Pharyngeal bones were removed from the pharyngeal teeth.In general,the first(outer) pharynx, stained with alizarin red,and then row has two slender teeth,the second(mid- cleaned.For cyprinid pharyngeal bones and dle)row three teeth slightly thicker than the their teeth, Chu's(1935)terminology has first-row teeth,and the main(third or inner) been employed by many authors.He used row four or five teeth,of which the third the terms'anterior'and'posterior'(e.g., and fourth teeth are large and broad.The anterior edentulous process, posterior angle) tooth formula for each species is shown in to indicate the positions of regions of the Table 1. bones.The pharyngeal bones of cyprinids, The number of teeth in the main row is however,are in reality more upright than stable in the 2,3,5-5,3,2 species.While in recumbent,and there are some cases where two of the 2,3,4-4,3,2 species,P.tetrazona this indication does not apply(e.g.,the and P. lateristriga,a fifth tooth is added to anterior angle is sometimes posterior to the the main row in some individuals.Of the posterior angle in actual position).We there- total of 23 specimens of P.tetrazona,two fore consider it proper to use'dorsal'in had 2,3,5-4,3,2 teeth,one 2,3,4-5,3,2,and place of posterior and'ventral'instead of one 2,3,5-5,3,2;of the 15 specimens of P. Fig.1.Two different types of infraorbital bones in Puntius.A,P.comingi,37.5 mm SL;B,P. orphoides,66.0 mm SL.Scales indicate 1 mm. Abbreviations;D,dermosphenotic;L,lacrimal; 2•`5,infraorbitals 2-5. ―2― Taki,Katsuyama and Urushido:Morphology and Relationships of Puntius ―3― 魚類学雑誌 Japan.J.Ichthyol.25(1),1978 lateristriga,one was 2,3,4-5,3,2,and another species.In the combination of the structural one 2,3,5-4,3,2.Variations in the number features of the pharyngeal bones and teeth, of teeth in the minor rows appear arbitrary six different types are recognized in the 23 and are observed in both the 2,3,4-4,3,2 species examined in this study(Fig.2). and 2,3,5-5,3,2 species but more frequently Characters of each type are briefly described in the former.The deviant formulae ob- as follows.Type A:pharyngeal bones very served are:1,3,4,-4,3,2 (P.tetrazona,1 broad and stout;dorsal limb broad, its tip specimen;P.fasciatus,1),2,3,4-4,3,1(P. truncate;ventral limb short;upper angle lateristriga,1),2,4,4-4,3,2(P.tetrazona,1; hardly recognizable;teeth congregated,thick, P.lateristriga,1),1,3,5-5,3,1(P.altus,1), swollen at apex.Type B:pharyngeal bones 2, 5,6-7,3,2(P.everetti,1),and 3,3,4-4,3,2 narrow; tip of dorsal limb pointed,some- (P.partipentazona,1). times slightly recurved in-and upward;up- In contrast to the inconstancy in tooth per angle conspicuous;teeth coarsely set, formula,the structure of the pharyngeal slender.Type C: pharyngeal bones stout, bones and their teeth are uninform within a broad;tip of dorsal limb truncate;upper Fig.2.Fronto-dorsal aspects of left pharyngeal and its teeth in Puntius to show six different types of their structure.A,P.gonionotus, 115.0mm SL;B.P.tetrazona,36.5 mm SL;C.P.fasciatus, 88.5 mm SL;D,P.binotatus,56.0 mm SL;E,P.filamentosus,34.5 mm SL;E',P.conchonius, 50.5 mm SL.Scales indicate 2 mm.Abbreviations:DL,dorsal limb;UA,upper angle;VL, ventral limb;I-III,first,second and main rows of teeth;5,fifth tooth in the main row. ―4― Taki,Katsuyama and Urushido:Morphology and Relationships of Puntius angle more or less conspicuous;teeth ag- gregated,broad.Type D.pharyngeal bones rather broad;tip of dorsal limb blunt,greatly recurved in- and forward;no upper angle; teeth aggregated,moderate in size.Type E:pharyngeal bones broad;tip of dorsal limb pointed;upper angle unrecognizable; teeth congregated,moderate in size.Type E':pharyngeal bones and their teeth resemble those of the type E in shape and structure, but teeth more coarsely set(this type was Fig.3.Semidiagrammatic illustration to show represented by E'because of its similarity to the location and naming of black body the type E).Characters of the pharyngeal markings in Puntius.Abbreviations:AB, bones and their teeth of each species are anal-basal marking; CD,caudal marking; indicated in Table 1 by type.