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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Cladus: Archelosauria
Division: Pan-Testudines
Division: Testudinata
Ordo: Testudines
Subordo: Cryptodira
Superfamilia: Testudinoidea

Familia: Geoemydidae
Subfamiliae: Geoemydinae - Rhinoclemmydinae


Geoemydidae Theobald, 1868


Vernacular names
Deutsch: Altwelt-Sumpfschildkröten
English: Geoemydid turtles
日本語: イシガメ科
한국어: 남생이과
македонски: Слатководни желки
ไทย: วงศ์เต่านา
Türkçe: Eski Dünya bataklık kaplumbağasıgiller

The Geoemydidae (formerly known as Bataguridae) are one of the largest and most diverse families[4] in the order Testudines (turtles), with about 70 species. The family includes the Eurasian pond and river turtles and Neotropical wood turtles. Members of this family are commonly called Leaf turtle.

Geoemydidae are turtles of various sizes (from about 10 to 80 cm (4 to 30 in) in length) with often a high degree of sexual dimorphism. They usually have webbed toes, and their pelvic girdles articulate with their plastrons flexibly. Their necks are drawn back vertically. Their carapaces have 24 marginal scutes. The plastron is composed of 12 scutes and has no mesoplastron; the pectoral and abdominal scutes contact the marginal scutes.

Some other features include a single articulation between the fifth and sixth cervical vertebrae, the lack of a hyomandibular branch of the facial nerve, and an epipterygoid bone in the skull.

Geoemydidae live in tropics and subtropics of Asia, Europe and North Africa; the only genus in Central and South America is Rhinoclemmys. Their habitats include freshwater ecosystems, coastal marine areas, and tropical forests. Most are herbivorous, but some are omnivorous or carnivorous species. In mating, the males are usually much more active than females. A relatively small number of eggs per clutch is common, produced several times a year. Some species have a temperature-dependent sex determination system, while others possess different sex chromosomes; one known species (Siebenrockiella crassicolis) exhibits XX/XY sex determination, while another species (Pangshura smithii) exhibits ZZ/ZW sex determination.[5]

About 70% of the extant species have been reported to be in endangered or vulnerable condition.
Systematics and evolution

Traditional systematics placed the geoemydids in the family Emydidae as the subfamily Batagurinae. In the 1980s, the subfamily was elevated to the family status and renamed to Geoemydidae according to the ICZN rules.

Most fossil and molecular data support their close relationship to the family Testudinidae.

The intrafamilial taxonomy is not well established yet, due to the large number and diversity of species. The family is usually divided into two subfamilies and 19 genera.[4] Several species are known to give viable hybrids, which makes the systematics even more complicated.[6]
Subfamilies and genera

The following genera are classified under Geoemydidae.[4]

Family Geoemydidae
Genus †Banhxeochelys
Genus †Duboisemys
Subfamily Geoemydinae
Genus Batagur (six species, including part of Kachuga)
Genus Cuora, Asian box turtle (10 species) (including Cistoclemmys)
Genus Cyclemys (seven species)
Genus Geoclemys (monotypic genus)
Genus Geoemyda (two species)
Genus Hardella (monotypic genus)
Genus Heosemys (formerly in Geoemyda)
Genus Leucocephalon (formerly in Geoemyda and Heosemys)
Genus Malayemys (three species)
Genus Mauremys, pond turtles (including Annamemys, Cathaiemys and Emmenia)
Genus Melanochelys (two species)
Genus Morenia (two species)
Genus Notochelys (monotypic genus)
Genus Orlitia (monotypic genus)
Genus Pangshura (four species) (formerly in Kachuga)
Genus Sacalia, "eyed" turtles
Genus Siebenrockiella (two species, one subgenus Panyaenemys, formerly under Heosemys)
Genus Vijayachelys, cane turtle (formerly in Geoemyda and Heosemys)
Subfamily Rhinoclemmydinae
Genus Rhinoclemmys, Neotropical wood turtles


As of the early 2013, six species of the family Geoemydidae are on the CITES Appendix I, and 30 more are on the treaty's Appendix II. A joint China-US proposal for a March 2013 CITES participants' conference seeks to add 15 more Geoemydidae species to the convention's Appendix II.[7] [8]

Theobald, William, Jr. 1868. Catalogue of Reptiles in the Museum of the Asiatic Society of Bengal. Journal of the Asiatic Society, Extra Number, 88 pp.
Gray, John Edward. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London 1869:165–225.
Gray, John Edward. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum. Part I. Testudinata (Tortoises). London: British Museum, 120 pp.
Turtle Taxonomy Working Group [van Dijk , P.P., Iverson, J.B., Shaffer, H.B., Bour, R., and Rhodin, A.G.J.]. 2012. Turtles of the World, 2012 update: annotated checklist of taxonomy, synonymy, distribution, and conservation status. Chelonian Research Monographs No. 5, pp. 000.243–000.328, doi:10.3854/crm.5.000.checklist.v5.2012, "Turtles of the World, 2012 update: annotated checklist of taxonomy, synonymy, distribution, and conservation status" (PDF). Archived from the original (PDF) on 2013-06-16. Retrieved 2014-04-19.
Badenhorst, Daleen; Stanyon, Roscoe; Engstrom, Tag; Valenzuela, Nicole (2013-04-01). "A ZZ/ZW microchromosome system in the spiny softshell turtle, Apalone spinifera, reveals an intriguing sex chromosome conservation in Trionychidae". Chromosome Research. 21 (2): 137–147. doi:10.1007/s10577-013-9343-2. ISSN 1573-6849. PMID 23512312. S2CID 14434440.
Buskirk, James R.; Parham, James F. & Feldman, Chris R. (2005): On the hybridisation between two distantly related Asian turtles (Testudines: Sacalia × Mauremys). Salamandra 41: 21-26. PDF fulltext[permanent dead link]
"convention On International Trade in Endangered Species of Wild Fauna and Flora: Sixteenth meeting of the Conference of the parties, Bangkok (Thailand), 3-14 March 2013. Consideration of Proposals for Amendment of Appendices I and II (CoP16 Prop. 32)" (PDF). Archived from the original (PDF) on 2013-02-22. Retrieved 2013-03-04.
Dinny McMahon, China Backs Tortoise in Race to Protect Endangered Species, 2013-03-04

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