33 



ted breast scale was illustrated in Howes (1985: Fig. 2) for Cone sins, but not discussed in 

 his text. Our study also shows the serrated breast scales present in Couesius (Chen & Ar- 

 ratia 1996). - TS 10: Breast scales not deeply embedded [0]. or deeply embedded [1] in 

 breeding males. - TS 11: Apical margin of breast scales in breeding male smooth [0]. or 

 serrated [1]. 



11. Tuberculation on breast scales (Fig.llA-C). Based on our observation and published 

 data, all cyprinids. except Phoxinus. lack tubercles on the apical margin of breast scales 

 in breeding males (Chen & Arratia 1996). In Phoxinus. each breast scale bears a series of 

 sharp tipped tubercles (generally two to six tubercles) on its apical margin (Fig.llC). Ma- 

 les of Tribolodon hakonensis also have a tuberculated margin in the breast scales, but the 

 arrangement of the breast scales is different from that in Phoxinus (Cavender & Coburn 

 1992). I interpret the tuberculation in Tribolodon homoplastic with that of Phoxinus be- 

 cause of the different arrangement of breast scales of the two genera, and the lack of the 

 anteriorly located anal pterygiophores in Tribolodon (Coburn & Cavender 1992). 



In the outgroups, breeding males do not bear tubercles on the breast scales. In the ingroup, 

 only three genera, Margariscus, Semotilus. and Couesius, bear one or two small tubercles 

 at the middle of each scale, but no tubercles are present at the apical margin (Chen & 

 Arratia 1996). The presence of centrally placed tubercles on the breast scales is interpre- 

 ted as apomorphic. the absence of the tubercles on this area of scales as plesiomorphic. 

 Chen & Arratia (1996) summarize the history of the study on the tuberculation in Phoxi- 

 nus. They compare the tuberculation on breast scales in Phoxinus and Margariscus-Cli- 

 nostomus-Couesius-group of Hubbs & Brown (1929). According to Chen & Arratia (1996), 

 two differences in breast tuberculation between Phoxinus and the Margariscus-Clinosto- 

 mus-Couesius-group (defined as Margariscus-group hereafter) are present. (1) The breast 

 scales are deeply embedded, and only the apical margin of the scale can be observed in 

 Phoxinus; whereas the breast scales are loose, not deeply embedded, and most of a scale 

 can be observed in the Margariscus-group. (2) In Phoxinus. tubercles are large, tall, sharp, 

 and present at the apical margin of the breast scales, and each scale bears up to six tu- 

 bercles (even more in some species of the genus); whereas in Margariscus-gvoup, the tu- 

 bercles are small, short, and centrally placed on the scale, and each scale bears only two 

 to three tubercles. 



TS 12: The breast scale in breeding males bearing no tubercles on the apical margin [0], 

 or bearing a series of sharp-tipped tubercles at the apical margin [1]. - TS 13: Each breast 

 scale of breeding males bearing no tubercle at its center [0], or centrally bearing one to 

 three [1] small tubercles. 



12. Tuberculation on scales on caudal peduncle (Fig.llD-F). In breeding males of most 

 members in the ingroup and in the outgroups, each scale on the ventrolateral part of the 

 caudal peduncle bears one or two tubercles on its middle portion, but no tubercles are pre- 

 sent on its apical margin (Fig.llD-E). In Phoxinus. however, each scale in this region 

 bears a few. up to 12, well-developed and sharp-tipped tubercles at its apical margin 

 (Fig. 1 IF). - TS 14: In breeding males, scales on the ventrolateral part of caudal pedun- 

 cle bearing no tubercles on the apical margin [0]; or bearing two or more tubercles on the 

 apical margin [1]. 



