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Current Herpetol. 20(2) 2001 



marginal in Mauremys japonica, the M. 

 leprosa species-group, Sacalia, and some- 

 times in Notochelys, M. annamensis, M. 

 mutica, the Cuora trifasciata species- 

 group, Cu. zhoui, and Melanochelys. 

 Thus, it is obviously inadequate to use 

 these characters to unite Sacalia and 

 Notochelys. In the present analyses, the 

 Notochelys-Cyclemys clade is supported by 

 three synapomorphic character states, and 

 we thus consider this relationship to be far 

 more likely than the Notochelys-Sacalia 

 monophyly. 



Except for the position of Notochelys 

 (see above), our results largely support 

 Hirayama's (1984) hypothesis in that they 

 also suggest: polyphyly of Cuora (sensu 

 lato) and validity of Cistoclemmys; close 

 relationships among Cistocleminys, Pyxidea 

 and Geoemyda; and polyphyly of Rhino- 

 clemmys. Sites et al. (1984), on the basis 

 of the analysis of allozymic variation 

 among 22 batagurid species, suggested that 

 Cuora and Cistocleminys are most closely 

 related to each other. This has been one of 

 the major reasons to regard Cistoclemmys 

 as a junior synonym of Cuora (e.g., Ernst 

 and Barbour, 1989; McCord and Iverson, 

 1991; Iverson, 1992). However, Sites et al. 

 (1984) examined very small numbers of 

 individuals for most species. The number 

 of loci examined in that study was also 

 very small. Our results showed that Cisto- 

 clemmys is related more closely to Pyxidea 

 and Geoemyda than to Cuora, although 

 Cistoclemmys, Pyxidea, and Geoemyda are 

 well differentiated from each other (Fig. 4). 

 Based on the external comparisons of 

 subspecies of Cistoclemmys galbinifrons 

 and Pyxidea, Fritz and Obst (1997) con- 

 cluded that Ci. g. serrata is a distinct 

 species which narrows the gap between 

 Cuora (sensu lato) and Pyxidea. They 

 doubted the validity of both Cistoclemmys 

 and Pyxidea. However, the number of 

 specimens of Ci. g. serrata examined by 

 them was very small. Moreover, they did 

 not examine internal characters, in which 



we did find a number of remarkable differ- 

 ences among Cistoclemmys, Cuora, and 

 Pyxidea. We thus consider the three genera 

 to be valid. 



The results of this study confirmed the 

 polyphyly of Rhinoclemmys as proposed 

 by Hirayama (1984), because they divided 

 Rhinoclemmys into two groups belonging 

 to different clades. Of these, Rhinoclem- 

 mys A consisted of two aquatic species, 

 punctularia and funerea, and two terres- 

 trial species, areolata and pulcherrima. 

 Rhinoclemmys B, on the other hand, 

 consisted of two terrestrial species with 

 unnotched, hooked beaks, anmilata and 

 rubida, that were not united into a mono- 

 phyletic group by the parsimonious analy- 

 sis, but did constitute a single unit in the 

 NJ phylogram. We thus discuss the 

 relationship of these two Rhinoclemmys B 

 species below. 



Based exclusively on the coloration and 

 the relative depth of the shell, Ernst (1978) 

 demonstrated that Rhinoclemmys (as Cal- 

 lopsis) can be divided into three groups, 

 the pulcherrima-rubida group, punctularia 

 group (as consisting of R. punctularia, R. 

 annulata, R. diademata, R. funerea, R. 

 melanosterna, and R. nasuta), and R. 

 areolata. This grouping substantially 

 contradicts our results, especially in the 

 positions of R. annulata, R. pulcherrima, 

 and R. rubida. Considering the remarkable 

 intersubspecific variation in those characters 

 in both R. pulcherrima and R. rubida 

 (Pritchard and Trebbau, 1984; Ernst and 

 Barbour, 1989), the grouping proposed by 

 Ernst (1978) is dubious. 



Sites et al. (1981), in the assessment of 

 allozymic variation among R. areolata, R. 

 funerea, R. pulcherrima, R. punctularia, 

 and R. rubida, suggested close affinity 

 among R. areolata, R. punctularia, and R. 

 funerea, and distinct differentiation among 

 the areolata-punctularia-funerea assem- 

 blage, R. pulcherrima, and R. rubida. Our 

 results are in accord with those of Sites et 

 al. (1981) in showing close relationship 



