20 



During this period (1970-1979), life history, ecology, and behavior of Phoxinus phoxinus 

 attracted many ichthyologists' attention. Wootton & Mills (1979) studied the breeding be- 

 havior of P. phoxinus in a small lake in Europe. Stott & Buckley (1979) observed P. pho- 

 xinus avoiding their home range when water was deoxygenated but back home when 

 oxygen was restored. Wootton et al. (1980) studied the relationships between body weight 

 and daily food consumption. Settles & Hoyt (1978) studied the reproductive biology of 

 P. erythro gast er from Kentucky. 



Some papers describing hybrids were published in these years, such as those by Legendre 

 (1970b) and Stasiak (1972) (Phoxinus neogaeus x P. eos), and Greenfield et al. (1973) 

 (P. erythrogaster x Notropis cornutus). 



1980-1994 



Joswiak (1980) studied the karyotype of five known nearctic Phoxinus species and P. pho- 

 xinus. He demonstrated 2N=50 for all studied Phoxinus species, and his karyotype data 

 supported the lumping of Phoxinus and Chrosomus. Based on the Nei distance among the 

 Phoxinus species, Joswiak (1980) proposed a phylogenetic tree for Phoxinus. The tree 

 shows that P. eos and erythrogaster were derived from P. oreas-like ancestor. Joswiak con- 

 sidered his study an evidence supporting the Bering bridge connection for the migration 

 of Phoxinus from Eurasia to North America. 



Coad (1984) described some P. phoxinus specimens with deformed vertebral columns, and 

 Banbura (1985) reported a case of P. phoxinus without pectoral fins. 

 Leuciscinae is a most problematic subfamily in cyprinids to which Phoxinus is generally 

 assigned. A few serial papers on the anatomy and relationships of Leuciscinae were pu- 

 blished by Howes (1978, 1984, 1985, 1991), though the monophyly of the subfamily is 

 still an open question. Having studied the anatomy and phylogeny of some Chinese carps 

 (e.g., Ctenopharyngodon and Hypophthalmichthys), Howes (1984) proposed that at least 

 four monophyletic groups could be recognized within Leuciscinae: aspinine, abramine, al- 

 burine (raised to subfamily level in Howes 1991), and phoxine with two genera, Phoxi- 

 nus and Couesius (Howes 1985, 1991). Howes (1985) reviewed the synonyms of Phoxinus 

 and proposed a new synonym list for the genus. This synonym list is a modification of 

 those in Berg (1949) and Banarescu (1964). According to Howes (1985, 1991), Phoxinus 

 and Couesius share the comb-like tuberculation on the breast scales in breeding males 

 (Howes 1985) and form a sistergroup 1 . Howes (1985) is an important contribution to the 

 study of Phoxinus because this was the first one to review the synonyms of Phoxinus 

 systematically. 



Kim & Kang (1986) studied the osteology of Moroco keumgang from Korea. Based on 

 the comparison between Moroco and Phoxinus, the authors claimed Moroco as the si- 

 stergroup of Phoxinus. They proposed the following synapomorphies for the two genera: 

 the presence of the anterior myodome, the lower part of the interorbital septum extended 



') However, as discussed below and Chen & Arratia (1996), the breast tuberculation of the two ge- 

 nera shows a great difference. 



