Cyprininae 



CD 



tincins 

 rasborins 

 acheilognathins 



gobionins 

 xenocyprins 



cultrins 



Shiner Clade -c 



c 



Western Clade \ 



Chub Clade 5 



leuciscins 



Fig.l: Phylogenetic hypothesis of Cyprinidae, showing the position of phoxinins. The synapomor- 

 phies supporting each nodes are as following. Node A: Absence of uncinate process on epibranchi- 

 als I and II; subtemporal fossa deep and well-developed; interorbital septum formed by orbitosphenoid 

 and a dorsal component of the parasphenoid. Node B: Rib head and parapophysis of fifth rib modi- 

 fied for greater mobility; origin of dorsal fin behind pelvic insertion; absence of anterior maxillary 

 barbel and accompanying foramen in maxilla. Node C: Crest of neural complex (supraneural) divi- 

 ded dorsally (forked); anterior (free) supraneural in contact with neural complex; three unbranched 

 dorsal fin rays. Node D: Scale without basal radii. Node E: Oval scales with modified circuli in po- 

 sterior field and apical radii only. Node F: Pterotic elongated, its anterior end reaching in front of 

 the anterior opening of the trigeminal-facial chamber. Node G: Anterior fork of the pelvic bone re- 

 duced; 48 diploid chromosomes with longest pair submetacentric. Node H: Supraorbital canal dis- 

 connected from the infraorbital canal; pharyngeal teeth in one or two rows. Node I: Phoxinin scale 

 type; basal radii of scale absence; scale with numerous apical radii; infraorbital and preoperculo- 

 mandibular canals not joined; opercular canal absent; preethmoid projecting anteroventrally when 

 viewed anteriorly. Node J: Postorbital cranium elongated; anterior and posterior angles of pharynge- 

 al arch rounded; rib of vertebra 4 with an anterior process (from Chen et al. 1984, Cavender & Coburn 

 1992, Coburn & Cavender 1992). 



The Chub clade within the phoxinins (Fig.l) contains the Exoglossin clade, Tribolodon, 

 and some other "genera (Fig. 2), including Phoxinus, defined as Hemitremian clade herein 

 (see below for detail). The Exoglossin clade is well diagnosed by synapomorphies (Co- 

 burn & Cavender 1992) such as the shortened mandibular canal terminating posterior to 

 the mental foramen. However, some problems are present in the relationships of other ge- 

 nera in the Chub clade [Tribolodon + Hemitremians] as discussed by Coburn & Caven- 

 der (1992). First, only two Eurasian genera of phoxinins (Tribolodon and Rhynchocypris) 

 were studied, obviously, "the examination of additional Eurasian phoxinin genera is clear- 

 ly required" (Coburn & Cavender 1992:329). Second, all the characters supporting the 

 nodes below the Exoglossin clade in Coburn & Cavender (1992: Fig. 13) are either re- 



