82 



Current Herpetol. 20(2) 2001 



examined, and this clearly indicates an 

 early divergence of the ancestral form of 

 R. zhengi from the common ancestor of 

 the brown frogs from both Europe and 

 Asia, including R. saiiteri from Taiwan 

 (Chou and Lin, 1997; Tanaka-Ueno et al., 

 1998b). 



We have already rejected Fei et al.'s 

 (1990) and Dubois' (1992) idea of regarding 

 R. saiiteri from Taiwan as a member of a 

 distinct genus or subgenus Pseudorana, and 

 suggested that the species be included in the 

 subgenus Rana of the genus Rana (Dubois, 

 1992) in our previous studies of the Taiwan- 

 ese, Japanese, and Russian brown frogs 

 (Tanaka-Ueno et al., 1998a, b). The present 

 study again demonstrated a monophyly of 

 the brown frogs including R. saiiteri, and 

 parallel divergences of this species in the 

 course of brown frog evolution. 



Rana saiiteri is surely divergent from 

 other brown frogs in having unique larval 

 characteristics of complicate dentition on a 

 large mouth and abdominal sucker in 

 adaptation to a torrential life (Chou and 

 Lin, 1997), but these characteristics can be 

 viewed as resuUs of convergent evolution 

 (Tanaka-Ueno et al., 1998a). 



Zhao (1999) first gave only a brief 

 diagnosis of R. zhengi without providing 

 pictures, and later redescribed it in longer 

 form and showing a color photograph of 

 two specimens that are not clearly desig- 

 nated as type series (Zhao, 2000a). In the 

 latter paper, Zhao (2000a) also compared 

 this species with R. sangzhiensis and R. 

 johnsi under an implicit presumption of 

 their close affinities to R. zhengi. These 

 latter two species have been assigned to 

 members of Pseudorana (Fei et al., 1990; 

 Dubois, 1992). 



Zhao (2000a) also described tadpoles of 

 R. zhengi with rather complicate dentition 

 (mostly 1:4-4/4-4:1), but never mentioned 

 the presence or absence of an abdominal 

 sucker. Rana johnsi, which is now 

 regarded as a distinct species (e.g., Inger et 

 al., 1999), was first described as a subspe- 



cies of R. sauteri (see Smith, 1921) and is 

 superficially similar to the latter species in 

 adult morphology. However, larvae of this 

 species lack an abdominal sucker or large 

 mouth parts, although dentition is compli- 

 cate (1:5-5/4-4:1; Matsui's observation of 

 the type series in the Natural History 

 Museum, London). Most probably, this 

 species belongs to a lineage distant from 

 brown frogs of the R. temporaria group. 



In the same journal (Sichuan Journal of 

 Zoology) as he described R. zhengi (Zhao, 

 1999, 2000a), Zhao (2000b) referred to 

 convergent properties of R. sauteri with 

 torrent-dweUing Ainolops species in the 

 structure of larval ventral sucker without 

 mentioning R. zhengi. He should have 

 described this type of modification in R. 

 zhengi if it was ever present in this species. 



Thus, both morphologically and geneti- 

 cally, R. zhengi is considered to be remote 

 from R. sauteri. In order to clarify the 

 exact phylogenetic position of R. zhengi in 

 the genus Rana, it is necessary to examine 

 biochemically representatives of many sub- 

 genera proposed by Dubois (1992), as well 

 as morphologically related species such as 

 R. johnsi and R. sangzhiensis. 



Acknowledgments 



We thank Prof. E.-M. Zhao for provid- 

 ing literature. We are grateful to C. 

 McCarthy and B. T. Clarke (BMNH) for 

 allowing MM to examine specimens under 

 their care. This research was partly sup- 

 ported by Grants-in Aid from the Ministry 

 of Education, Science, Sports and Culture 

 of Japan (Nos. 07454234 and 10041166) to 

 MM, and by the Research Fellowships of 

 the Japan Society for the Promotion of 

 Science for Young Scientists (No. 5397) to 

 TT-U. 



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