YASUKAWA IT AI.. — PIIYl OCl'NY OF GEOEMYDIM-; TURTLES 



121 



-Siebenrockiolla crassicollis 



- Orlilui borneonsis 



Sticalia spp. 



— Mauremys japonica 

 M.niromys leprosa species group 

 M.Hircmys mulica 

 Mauremys annamensis 



I Notochelys plalynota 



I Cyclemys spp 

 I Inosomys spinosa 



Hoosemys grandis 



— Molanochelys Iriiuga 



Rhinoclemmys areolata 

 Rhinoclemmys pulcherrlma 

 nhinoclommys funerea 



Rhinoclemmys punclulaha 



Cuora amboinensis 



Cuora zhoui 



Cuora Irilasciala species group 

 Cuora mccordi 



' inmoclommys annulal<\ 

 — Rhinoclemmys rubid. i 



■ lyidea mouholii 



- 1 isloclemmys galbinilrons 



Cistoclemmys tlavomargnata 



■ oemyda silvatica 



, Geoemyda spengleri 



Geoemyda japonica 



Fk;. 4. IMiylogram by ncii;hhi)i-ji)iiiiiig ihciIukI oI the sublainily Cicocniyciiiiac and its relatives. 



PvxkIch IkkI \\\o aiiloapiMiUM jihic cluirac- 

 Il'i slaIc^: 2^ (I), NlioiiL' sciraiioii o\ posic- 

 rioi cai apace mai yin; ami ''1 (1), posiei ioi 

 licaJ skill siilnliv uleJ iiilo small scales. 

 MinmpInK ol ( icocniycia (Stem S) was 

 sii]ipoiicd In ri\e eliaiactor stales: 10 ( 1 ). 

 \ eiil I nmeJial iiieeliiiL' ol maxillae: (I), 

 enioplasium sepaialed liom i'lilo liiimei al 

 scam; S pieseiiee ol ilie juvai ^|iKulia 

 ti)jugal eoiiiaei; P (1), \eiv weak ma\illa 

 ptciy};oii.l eoiuaei; aiul IS (1) iMoeessus 

 trochlearis iiticmii loiiiieJ uiili piooiie aiul 

 qiKuliaie. I he loimei lUD eharaetcr stales 

 \\eie iiiikiiie m the Ikitaiiur idac. in addi 

 iioii, ilie (/. v/j(V/t,'/('r/- (/. japonici/ ekule 

 (Niein 1) \\as suppoiieJ In ihroc syiiapo 

 moiplue eliaiaeiei stales: ^ (2), t'oianieii 

 nei\i \uliam somelimes located on piei> 

 goid; 15 (2), lorauicn pracpalatmum open- 

 ing within picnuivilla; and 23 (1). strong 

 scrraliiMi ai pivsicrior carapace margin, 

 riie til St uas unique uithiu the Balagur- 

 kl.ie. 



1 he pluloi'iam, ohtamed liy the neighbor- 

 joining; (NI) method using an absolute 

 distance in.iiiiv is shown in l ie. 4. This 



iMuKiyram is laiL'el> icleiilica! in topology 

 wiih ilie siiici consensus nee from tlic 

 lxiisimon\ anahsis (I-ig. 3), and thus is 

 iniei ineied as suiiporiiiig the validity of the 

 killer. One ol liie ditTercnccs between the 

 siricl consensus nee and the NJ phylogram 

 la\ in ihe lelaiionships within the Rliino- 

 clvinnivs \. In ihe latter tree, R. pulchcr- 

 nimi uas uinieJ noi with the R. fiincrea-R. 

 ininctularui group but with A', an'oluia. 

 \noiher (.litterence was in the relationships 

 anion;' A', uriniilaia. R. rtihida, and the 

 ( 'iMock'fnniys- Pyxidea-Gcoemyda group. 

 In the N.I phylogram. R. anniilala and R. 

 rtihida were united, collectively constituting 

 the first outgroup to the Cisloclemmys- 

 Pv\t(lcit CictH'tnydii group. 



Discussion 



Monophyly of the Hatasuridav 



Based on the analyses of morphological 

 and mithochondrial l)N.\ characters for 

 major lestudinc laxa including one haiagur- 

 inc. one gcocmydinc. three cmydid. and 

 one icsiudinid genera. Shaffer et al. (I')***) 



