22 



strated that this species "appears to prefer cold, clear water running over a sand and stone 

 substrate and was often together with Nemacheilus, Misgurnus, and young salmonids." 

 Reproductive biology of Phoxinus phoxinus was studied by Mills (1987). Mills (1987) 

 reported the reproductive size for P. phoxinus at 50 mm length. The behavior of P. pho- 

 xinus was described by some European ichthyologists. Partridge (1980) and Magurran & 

 Girling (1986) studied the schooling of the species. Kennedy (1981) observed the homing 

 tendency of the species. Legkiy & Popoya (1984) demonstrated that one of the important 

 factors for the downstream migration of P. phoxinus was a change in their photoreaction 

 (negative response). Pfeiffer et al. (1985) indicated P. phoxinus bear alarm cells through 

 the year. Bioenergetic biology of the species was studied by Cui & Wootton (1988a, b, 

 1989). Distribution and ecology of P. cumberlandensis was studied by O'Bara (1990, 



1991) . 



Wheeler (1991) discussed the status of the type specimens of P. phoxinus. The specimens 

 were in very poor condition and Wheeler was not able to identify whether the specimens 

 were the type of the species (see the discussion in the species account of P. phoxinus), 

 thus a problem was raised if the specimen is the "real" type of P. phoxinus. 



Summary 



Since the description of Cyprinus phoxinus was published (Linnaeus 1758), almost 240 

 years have passed. In the past more than 200 years, ichthyologists have paid much atten- 

 tion to the genus Phoxinus, publishing hundreds of studies related to these fishes. The 

 main contributions during these years can be summarized as follows: 



1. Numerous records of the geographic distribution of the genus; 



2. Accumulation of data on life history, ecology and behavior of some species, such as P. 

 phoxinus and erythro gaster\ 



3. Some knowledge of osteology and external morphology of some species; and 



4. Synonyms of some species. 



PHYLOGENETIC RELATIONSHIPS OF PHOXINUS AND THE RELATED GENERA 



Monophyly of the Hemitremian clade and its position in the family Cyprinidae 



According to Chen et al. (1984) and Cavender & Coburn (1992), Cyprinidae comprises 

 two major lineages, the subfamilies Cyprininae and Leuciscinae (Cavender & Coburn 



1992) (Fig.l). Phoxinins, defined by the disconnection between temporal and preopercu- 

 lomandibular sensory canals, is one of the eight clades in Leuciscinae. Consisting of Shi- 

 ner, Chub, and Western clades (Fig.l), phoxinins include almost all North American 

 minnows (excluding Notemigonus crysoleucas which belongs to the Eurasian leuciscins) 

 and a few Eurasian genera (Cavender & Coburn 1992; Coburn & Cavender 1992). 



The term "phoxinins" used in this publication has different content from that of "phoxi- 

 nins" of Howes (1985). "Phoxinins" of Howes contains two genera, Phoxinus and Coue- 

 sius (Howes 1985, 1991). When I refer to "phoxinins" of Howes, I use "phoxinins of 

 Howes". The term "phoxinins" used herein refers to that of Cavender & Coburn (1992) 

 and Coburn & Cavender (1992). 



