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Current Herpetol. 20(2) 2001 



subfamilies to separate families (Batagur- 

 idae, Emydidae, and Testudinidae, respec- 

 tively) of the superfamily Testudinoidea, 

 making the Bataguridae the largest and 

 most divergent family in the order 

 Testudines. They also divided this family 

 into two subfamilies, Batagurinae and 

 Geoemydinae (Gaffney and Meylan, 1988). 

 The former currently contains 22 species of 

 12 genera (David, 1994; McCord and 

 Iverson, 1994) distributed over most of 

 temperate to tropical Asia, except for the 

 western part (Iverson, 1992). The latter 

 consists of 45 species of 12 genera (David, 

 1994; McCord et al., 1995, 2000; Fritz et 

 al., 1997; Fritz and Wischuf, 1997; 

 McCord, 1997), occurring in northern 

 Africa, southern Europe, most parts of 

 temperate to tropical Asia, central America, 

 and northern South America (Iverson, 

 1992). 



The first attempt to clarify the phylogeny 

 of the Bataguridae was made by Hirayama 

 (1984) (as the Batagurinae: see above), 

 utilizing cladistic analyses of 37 species 

 representing all recognized genera mainly 

 on the basis of morphological characters. 

 Two major groups were recognized, of 

 which one, consisting of Batagur, Callagur, 

 Chineinys, Geocleinys, Hardella, Hier- 

 emys, Kachuga, Malayemys, Morenia, 

 Ocadia, Orlitia, and Siebenrockiella, was 

 characterized by relatively broad triturating 

 surfaces of jaws (=Batagurinae sensu 

 Gaffney and Meylan, 1988), whereas the 

 other, accommodating Cistocleinmys, 

 Cuora, Cyclemys, Geoemyda, Heosemys, 

 Mauremys, Melanochelys, Notochelys, 

 Pyxidea, Rhinoclemmys and Sacalia, was 

 characterized by relatively narrow tritu- 

 rating surfaces of jaws (=Geoemyd- 

 inae). Hirayama (1984), while defining the 

 broad triturating surfaces as a synapo- 

 morph of the Batagurinae, regarded the 

 narrow triturating surfaces in the Geoemy- 

 dinae as a primitive condition of the Bat- 

 aguridae. He thus considered Geoemydinae 

 as paraphyletic, with the Rhinoclemmys 



annuIata-R. rubida-Geoemyda-Cistoclem- 

 mys-Pyxidea clade being monophyletic 

 with the family Testudinidae rather than 

 with the other geoemydine genera. In this 

 hypothesis, the Batagurinae is the sister 

 group of the Geoemydinae plus the Testu- 

 dinidae, and the Bataguridae is also para- 

 phyletic. 



With respect to batagurine monophyly 

 and phylogeny within this subfamily, 

 Gaffney and Meylan (1988) approved 

 Hirayama's (1984) view because most char- 

 acters Hirayama employed show a low 

 degree of homoplasy in their cladogram. 

 They also shared the view of possible 

 non-monophyly of the Bataguridae with 

 Hirayama, but doubted the geoemydine 

 phylogeny hypothesized by Hirayama 

 because of a very high degree of 

 homoplasy postulated therein. Since then, 

 no comprehensive systematic studies have 

 been carried out for the Bataguridae, 

 leaving both the geoemydine phylogeny 

 and the batagurid monophyly problematic. 



Hirayama's (1984) paper was essentially 

 an abstract of a much larger work of 

 character analysis (not yet pubHshed), and 

 thus a number of important issues, such as 

 detailed descriptions of character states, 

 were omitted (Gaffney and Meylan, 1988). 

 In addition, a number of taxonomic 

 changes have been made for geoemydine 

 turtles since Hirayama's (1984) analysis 

 (David, 1994; Obst and Reimann, 1994; 

 McCord et al., 1995; Yasukawa et al., 

 1996; Fritz and Wischuf, 1997; Fritz et 

 al., 1997; Iverson and McCord, 1997a; 

 McCord, 1997; Lehr et al., 1998), though a 

 few recent authors suspect some of these 

 changes to have been misled by artificial 

 hybrids and thus are untenable (van Dijk, 

 2000; Shi and Parham, 2001; Das, personal 

 communication). These conditions have 

 made it strongly desirable to conduct 

 additional analyses for the geoemydine 

 phylogeny with detailed methodological 

 descriptions and considerations of those 

 taxonomic changes. We therefore analyzed 



