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Order Testudines
Suborder Cryptodira
Superfamily Testudinoidea
Family Geoemydidae (including Geoemydinae and Batagurinae)
(Asian River Turtles, Leaf and Roofed Turtles, Asian Box Turtles)
Keeled Box Turtle (Pyxidea mouhotii)
© Wayne van Devender
Appearance and morphology: Turtles with oval to oblong and moderately domed or flattened carapaces; the plastron is large and occasionally hinged. The jaw closure mechanism articulates on a trochlear surface of the otic capsule and is enclosed in a synovial capsule. An epipterygoid is present in the skull; the internal carotid canal lies in the pterygoid, and the parietal but not the postorbital touches the squamosal. The facial nerve lacks a hyomandibular branch. The plastron lacks a mesoplastron, and the plastral buttresses usually articulate firmly with the costals of the carapace; the carapace has 11 pairs of sutured peripherals around its margin and a nuchal without costiform processes. The neck withdraws vertically, and this mechanism is reflected in a anteroventrally oriented articular surface of the first thoracic vertebra; other vertebral traits are the exclusion of the 10th thoracic vertebra from the sacral complex and procoelous caudal vertebra. The pelvic girdle flexibly articulates with the plastron, and the ilium lacks a thelial process (after Zug et al. 2001).
Size: carapax from 13 cm (Geoemyda spengleri, Heosemys silvatica) to 80 cm (Orlitia borneensis).
Distribution: Southern Europe to Japan in Asia, East Asia, Central America and northern South America.
Habitat: mostly aquatic or semiaquatic. Although most species live in freshwater habitats, and a few even in mountain streams (e.g. Cyclemys dentata, Cuora trifasciata) or estuaries (Batagur baska, Callagur borneoensis). Callagur borneoensis is a batagurine that nests on sea beaches along with nesting sea turtles. Geoemyda spengleri , Heosemys silvatica, and Pyxidea mouhotii are terrestrial. However, even within a single genus habitat preferences can vary markedly. For example, Rhinoclemmys has totally terrestrial species (R. annulata) and highly aquatic species (R. annulata)
Reproduction: Most species produce fewer than 10 eggs per clutch, although several appear to ahve multiple clutches during a single season. Geoclemys hamiltoni has 18-30 eggs, Batagur baska on average 20 eggs, Callagur borneoensis 15-25 eggs per clutch. Incubation periods are unknown for most species but appears to be commonly around 3 to 5 months.
Food: Most species are herbivorous (Kachuga smithi), or omnivorous, but some are carnivorous (e.g. Malayemys, Mauremys).
Taxonomic notes: Gaffney & Meylan (1988) state that the Bataguridae would be paraphyletic without the inclusion of the Testudinidae. The generic subdivision into Batagurinae and Geoemydinae follows OBST (2003). Melanochelys, Heosemys are sometimes considered as subgenera of Geoemyda. Annamemys has been synonymized with Mauremys. Cistoclemmys is treated here as a synonym of Cuora. Pyxidea has been synonymized with Cuora by SPINKS e al. (2004) who also included Chinemys in Mauremys and Hieremys in Heosemys. Another change suggested by SPINKS et al. (2004) is the revalidation of Pangshura for several species of Kachuga. Note that according to these authors the subfamilial classification of the Geoemydidae needs to be revised significantly. See the figure below for generic relationships based on DNA sequence data.
List of Genera (after Gaffney and Meylan 1988 and modified after Obst 2003):
Subfamily Batagurinae
Callagur and Kachuga have been synonymized with Batagur (see note below).
Subfamily Geoemydinae
The genus Pyxidea has been synonymized with Cuora. Barth et al. (2004), baserd on DNA sequence data, suggested to lump all species of Mauremys, Chinemys, and Ocadia in one expanded genus, Mauremys, or split Mauremys into 4 genera: Ocadia Gray 1870 is the oldest available name for the clade containing 'M .' japonica, all Chinemys species, and O.sinensis. For the clade containing 'M.' annamensis and 'M .'mutica, Cathaiemys Lindholm 1931 is available, and Emmenia Gray,1870 for 'M .'caspica and 'M .'rivulata. Mauremys Gray,1869 would have to be restricted to M.leprosa.
Hieremys has recently been synonymized with Heosemys (DIESMOS et al. 2005). Panyaenemys was erected as a new clade and subgenus including Siebenrockiella leytensis (formerly Heosemys leytensis.) but not Siebenrockiella crassicollis..
Phylogenetic relationships among Geoemydid genera (after SPINKS et al. 2004)
This phylogenetic is based on analyses of the protein-coding cyt b mtDNA, 12S ribosomal (rDNA) mtDNA, and a 1 kb intronfrom the R35 neural transmitter gene. For details see SPINKS et al. 2004. Note that Le et al. (2007) and Praschag et al. (2007) found a different topology for the Geoemydidae in which Kachuga was paraphyletic and thus suggested to combine all species of Kachuga and Callagur in the genus Batagur.
References:
BARTH, D.; D. BERNHARD; G. FRITZSCH & U. FRITZ (2004)
The freshwater turtle genus Mauremys (Testudines, Geoemydidae) &emdash; a textbook example of an east&endash;west disjunction or a taxonomic misconcept?
Zoologica Scripta 33 (3): 213&endash;221
Diesmos, Arvin; Parham, James F.; Stuart, Bryan L.; Brown, Rafe M. (2005)
The phylogenetic position of the recently rediscovered Philippine forest turtle (Bataguridae: Heosemys leytensis).
Proceedings of the California Academy of Sciences 56(3):31-41.
Ernst,C.H. & Barbour,R.W. (1989)
Turtles of the World
Smithsonian Institution Press, Washington D.C. - London
ISBN 0-87474-414-8
Fritz,U. & Wischuf,T. 1997
Zur Systematik west-asiatisch-südosteuropäischer Bachschildkröten (Gattung Mauremys) (Reptilia: Testudines: Bataguridae).
Zool. Abh. Staatl. Mus. Tierk. Dresden 49 (13): 223-260
Fritz,U., Gaulke,M. & Lehr,E. 1997
Revision der südostasiatischen Dornschildkröten-Gattung Cyclemys Bell 1834, mit Beschreibung einer neuen Art.
Salamandra 33 (3): 183-212
Gaffney-E-S; Meylan-P-A (1988)
A phylogeny of turtles.
SYSTEMATICS ASSOCIATION SPECIAL VOLUME (No. 35A) 1988: 157-219
Le, Minh; William P. McCord and John B. Iverson (2007)
On the paraphyly of the genus Kachuga (Testudines: Geoemydidae).
Molecular Phylogenetics and Evolution 45: 398-404
Obst, F.J. (2003)
Sumpfschildkröten.
Draco 4 (13): 4-18
Praschag, P.; Hundsdörfer, A.K. & Fritz, U. (2007)
Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variation (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura).
Zoologica Scripta 36 (5): 429–442
Schaefer, I. (2005)
Zacken-Erdschildkröten - Die Gattung Geoemyda.
Natur und Tier Verlag, Münster, 143 pp.
Schilde, M. (2004)
Asiatische Sumpfschildkröten - Die Familie Geoemydidae in Südostasien, China und Japan.
Natur und Tier Verlag (Münster), 192 pp.
Spinks, Phillip Q.; Shaffer, H. Bradley; Iverson, John B; McCord, William P. (2004)
Phylogenetic hypotheses for the turtle family Geoemydidae.
Molecular Phylogenetics and Evolution 32 (1):164-182
Spinks, Phillip Q.; Shaffer, H. Bradley 2007
Conservation phylogenetics of the Asian box turtles (Geoemydidae, Cuora): mitochondrial introgression, numts, and inferences from multiple nuclear loci.
Conservation Genetics 8(3):641-657
Wermuth,H. & Mertens,R. (1996)
Schildkröten, Krokodile, Brückenechsen
(reprint of the 1961 edition with an updated appendix by F.J. Obst), 506 pp.
Gustav Fischer Verlag, Jena
Zug,G.R.; Vitt, L.J. & Caldwell, J.P. (2001)
Herpetology, 2nd ed.
Academic Press San Diego, London, [...]XIV + 630 pp.
Online information:
Emydidae in King & Burke: CROCODILIAN, TUATARA, AND TURTLE SPECIES OF THE WORLD.
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