Osteology in the Cuora galbinifrons complex suggests conspecifity of C. bourreti and C. galbinifrons, with notes on shell osteology and phalangeal formulae within the Geoemydidae

Thirty-six complete skeletons, three shells, and x-rays of the extremities of 32 additional turtles of the Cuora galbinifrons complex have been compared with 38 other geoemydid species from 19 genera. Cuora bourreti differs from C. galbinifrons and C. picturata by a lost phalanx in the fourth finger and fourth toe. Individuals with a shell shape intermediate between C. bourreti and C. galbinifrons, as found on Hainan Island (China), have either the reduced phalangeal formula of C. bourreti (manus: 2-3-3-2-2, pes: 2-3-3-2-1) or the complete number of phalanges (manus: 2-3-3-3-2, pes: 2-3-3-3-1). Only in C. flavomarginata did we also register a lost phalanx in the fourth digit of manus and pes; in the pes of C. mouhotii the same character state may occur. In C. flavomarginata the fifth digit of the pes is also lacking. Some other terrestrial and semiterrestrial geoemydids (Cuora mccordi, Heosemys spinosa, and in part C. mouhotii and Leucocephalon yuwonoi) display a similar pattern of phalangeal reduction, resulting in the loss of the fifth digit of the pes. Likewise, in tortoises (Testudinidae), a further group of terrestrial chelonians, and the terrestrial turtle genus Terrapene (Emydidae) the loss of phalanges or complete digits is known to occur. Malayemys subtrijuga, Morenia petersi, Pangshura smithii and Siebenrockiella crassicollis differ from all other studied geoemydid taxa by an additional phalanx in the fifth digit of the manus (2-3-3-3-3); one P. smithii has on one body side three phalanges in the fifth digit of the pes (2-3-3-3-3). These are highly aquatic turtles with extensive toe webbing. Probably, longer digits (and thus a higher phalangeal number) are a favorable prerequisite for swimming while phalangeal loss seems to be the consequence of walking. Cuora bourreti and C. picturata have consistently in the bony carapace a very rare character state regarding the articulation of the rib tips with the peripheral plates. In both species the rib tips are intercalated between two peripheral plates in the bridge region. Intercalated rib tips like those in C. bourreti and C. picturata were found only in C. m. mouhotii, but not in the southern subspecies C. mouhotii obsti . I nC. galbinifrons and all other geoemydid taxa studied the rib tips articulate on the bridge within the underlying peripheral and not between two peripherals. In turtles morphologically intermediate between C. bourreti and C. galbinifrons both characters states are found. This suggests that such individuals are hybrids or intergrades. Three known- locality specimens from Hainan Island display both extremes and an intermediate character state. This, together with external morphology and the occurrence of both phalangeal formulae in approximately the same frequency on that island, argues for genetic introgression of C. bourreti on the Hainan population of C. galbinifrons. We conclude that our findings qualify C. galbinifrons and C. bourreti under the Biological Species Concept as conspecific.