21 



and contacting the parasphenoid. and a relativer) straight parasphenoid (Kim & Kang 

 1986). 



Chen et al. (1984) published a hypothesis of phylogenetic relationships among the major 

 groups of Cyprinidae. They proposed ten subfamilies, among which Phoxinus belongs to 

 the subfamily Leuciscinae. The content of the subfamily Leuciscinae of Chen et al. (1984) 

 is similar to that of Gosline (1978). However. Howes (1991) complained that the analy- 

 sis of Chen et al. (1984) was not a cladistic one. although the authors claimed so. Chen 

 et al. (1984) did not provide synapomorphies to support some sistergroup relationships 

 and some subfamilies are recognized by plesiomorphies only. 



Cavender & Coburn (1987) studied the evolutionär) relationships among eastern North 

 American cyprinids. They stated that Phoxinus, Semotilus, Couesius, Hemitremia, and Cli- 

 nostomus share a unique type of anal fin suspension, and the anterior placement of the 

 anal fin pterygiophores. In a later study of the phylogenetic relationships of cyprinids, Ca- 

 vender & Coburn (1992) recognized two subfamilies, i.e.. Cyprininae and Leuciscinae. in 

 the family Cyprinidae. Phoxinus was placed in the phoxinin tribe of Leuciscinae. 

 Chen (1986a. b. 1987a. b) studied the osteology and phylogeny of Chinese leuciscines. 

 He proposed the Chinese leuciscines a monophyletic group and included Phoxinus in the 

 subfamily Leuciscinae. 



During the study of fish fauna of Huang He (Yellow River) in China. Chen (1988a) de- 

 scribed a new Phoxinus species. P. tchangi from the Huang He tributary. The olfactory 

 organ of Phoxinus was studied by Chen (1988b). Chen (1988b) considered the olfactory 

 organ of Phoxinus a primitive condition in Cyprinidae because the types of the olfactory 

 organ in Phoxinus were commonly found in Cyprinidae. 



The sixth North American Phoxinus species. P. tennesseensis, was described by Starnes & 

 Jenkins (1988) from the Tennessee River drainage, USA. Starnes & Jenkins (1988) also 

 proposed P. cumberlandensis the sister species of [P. oreas + tennesseensis]. and P. eos 

 the sister species of P. erythrogaster. 



Banarescu (1989) analyzed the distribution patterns of European freshwater fishes. Two 

 patterns were proposed: vicariant and dispersal. In this paper. Banarescu considered P. 

 phoxinus the vicariant sister of P. neogaeus though no evidence was discussed (Banares- 

 cu 1989). 



Bugutskaya (1987. 1988a. b. 1989. 1990. 1991) published several papers on the osteolo- 

 gy and classification of Leuciscinae based mainly on the Eurasian genera. She recognized 

 eight tribes in the subfamily: Leuciscini. Alburnini. Phoxini. Abramini, Aspinini, 

 Elopichthyini. Pseudoaspini. and Hypophthalmichthvini. 



The external morphology and biology of P. phoxinus from River Skawa of Poland were 

 studied by Heese (1981. 1984). He stated the maximum age of P. phoxinus to be more 

 than six years. Sexual dimorphism of the species was shown in the shape of pectoral and 

 ventral fins, the coloration, and the tuberculation (Heese 1981. 1984). Heese's result was 

 similar to that of Frost (1943). 



Based on field work in Mongolia in 1984, Travers (1989) described the Mongolian fish 

 fauna. P. phoxinus was caught from "Arctic and Pacific basins."* Travers (1989) demon- 



