Reptiles Database

Order Testudines
Suborder Cryptodira
Superfamily Testudinoidea

Family Testudinidae (Tortoises)


Chelonoidis carbonaria.
© Peter Uetz

Appearance: They have high domed shells and stout, heavily scaled limbs (Malacochersus is an exception with a dorsoventrally compressed shell). The head and limbs can be fully withdrawn into the shell. The genera Testudo and Pyxis have hinged plastrons while members of the genus Kinyxis have, uniquely among tortoises, a hinged carapace that allows it to be lowered over the hindquarters.

Size: 10 - 125 cm carapax length. Gigantism has evolved independently in several island populations, with those of the Galapagos and Aldabra islands being the most famous.

Distribution: Testudinids are land turtles that occur mainly in Africa in Asia although a few species are known from the Americas and Europe.

Food: predominantly herbivorous or omnivorous

Habitat: Tortoises are mainly arid-adapted.

Behaviour: Juvenile turtles tend to be cryptic, because their shell doesn't protect them from predators yet.

Reproduction: About 12 eggs in Testudo graeca and Testudo (= Agrionemys) horsfieldii. the latter deposits its eggs in July and the young hatch only 9 months later (April). In contrast, the young of Testudo kleinmanni hatch already after 20 days.

Relationships: Sometimes the Batagurinae have been included as subfamily in the Testudinidae, although they are also dconsidered as separate family, Bataguridae, or as subfamily of the Emydidae.

Taxonomic notes: Geochelone is considered as a subgenus of Testudo by Wermuth & Mertens (1977), but as a valid genus by Ernst & Barbour (1989). The species list given bellow follows mainly Ernst & Barbour (1989).

Geochelone, Asterochelys, Chelonoidis, Megalochelys, and Testudo are used synonymously to some extent. Check species list of Geochelone and Testudo. The giant tortoises of Galapagos (Geochelone elephantopus ssp.) have been subdivided in a series of separate species. The Indian Ocean giant tortoises (genus Cylindraspis) seem to be extinct now (Arnold 1979, Bour 1980). Cylindraspis contains at least 5 species: borbonica (Réunion), triserrata, inepta (both Mauritius), vosmaeri, peltastes (both Rodrigues) (AUSTIN et al. 2002).

Subgenera of Testudo according to Wermuth & Mertens (1977): Acinixys, Agrionemys, Asterochelys, Chelonoidis, Chersina, Geochelone, Indotestudo, Manouria, Megalochelys, Psammobates, Pseudotestudo, Testudo.

Previously, some authors included the Platysterninae in the Testudinidae. However, they are now considered as a subfamily of the family Chelydridae.

List of genera:

Zug et al. (2001) do not include Dipsochelys and Furculachelys in their list of genera for this family. The former is then considered as a synonym of Testudo (see also Kuyl et al. 2002, for a phylogenetic analysis of Testudo sp.).

Phylogeny: Le et al. (2006) presented a comprehensive molecular analysis and suggested several changes to testudinid taxonomy. For example, they erected the new genus Angonoka for Geochelone yniphora. Their phylogeny looks as follows:

Fritz & Bininda-Emonds (2007) published a revised phylogeny based on additional sequence data and synonymized Angonoka with Testudo. They also erected the new genus Stigmochelys for Geochelone pardalis.

References:

Alderton, David (1997)
Turtles & Tortoises of the World
Facts on File, New York, 191 pp.

Arnold, E.N. 1980
Indian Ocean giant tortoises: their systematics and island adaptations.
PHILOSOPHICAL TRANSACTIONS OF THE ROYAL SOCIETY OF LONDON B BIOLOGICAL SCIENCES 286 (1011): 127-145

Bonin, FranckDevaux, Bernard Dupré, Alain (1996)
Toutes les Tortues du Monde [Turtles and Tortoises of the World]
Delachaux et niestlé, Lausanne, Switzerland, 254 pp.

Bour,R. 1980
Systematique des tortues terrestres des iles Mascareignes: genre Cylindraspis Fitzinger, 1835 (Reptilia, Chelonii).
BULLETIN DU MUSEUM NATIONAL D'HISTOIRE NATURELLE SECTION A ZOOLOGIE BIOLOGIE ET ECOLOGIE ANIMALES 2 (3): 895-904

Crumly, Charles R. (1994)
Phylogenetic systematics of North American tortoises (genus Gopherus): evidence for their classification, in: Biology of North American Tortoises.
U.S.D.I. National Biological Survey, Washington, D.C.,

Ernst,C.H. & Barbour,R.W. (1989)
Turtles of the World
Smithsonian Institution Press, Washington D.C. - London

Fritz, Uwe and Olaf R.P. Bininda-Emonds (2007)
When genes meet nomenclature: Tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone.
Zoology 110 (4): 298-307

Gerlach, Justin Canning, Laura (1998)
Taxonomy of Indian Ocean giant tortoises (Dipsochelys).
Chelonian Conservation and Biology 3 (1): 3-19

Gerlach, J. (2004)
Giant tortoises of the Indian Ocean. The genus Dipsochelys inhabiting the Seychelles Islands and the extinct giants of Madagascar and the Mascarenes.
Edition Chimaira, Frankfurt, 208 pp.

Highfield,A.C. & Martin,J. (1990)
A revision of the Testudines of North Africa, Asia and Europe.
J. Chelon. Herpetol. 1 (1): 1-12

Kuyl, Antoinette C. van der; Donato L. Ph. Ballasina, John T. Dekker, Jolanda Maas, Ronald E. Willemsen and Jaap Goudsmit (2002)
Phylogenetic Relationships among the Species of the Genus Testudo (Testudines: Testudinidae) Inferred from Mitochondrial 12S rRNA Gene Sequences.
Molecular Phylogenetics and Evolution 22: 174-183

Lamb, Trip & Lydeard, Charles (1994)
A molecular phylogeny of the gopher tortoises, with comments on familial relationships within the Testudinoidea.
Molecular Phylogenetics and Evolution 3 (4): 283-291

Le, Minh; Christopher J. Raxworthy, William P. McCord and Lisa Mertz (2006)
A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes.
Molecular Phylogenetics and Evolution 450 (2): 517-531

Palkovacs, Eric P.; Justin Gerlach and Adalgisa Caccone (2002)
The evolutionary origin of Indian Ocean tortoises (Dipsochelys).
Molecular Phylogenetics and Evolution 24 (2002) 216&endash;227

Parham, James F.; J. Robert Macey; Theodore J. Papenfuss; Chris R. Feldman; Oguz Türkozan; Rosa Polymeni and Jeffrey Boore (2005)
The phylogeny of Mediterranean tortoises and their close relatives based on complete mitochondrial genome sequences from museum specimens.
Molecular Phylogenetics and Evolution 38 (1): 50-64 [2006]

Rogner, M. (2005)
Griechische Landschildkröten (Testudo hermanni hermanni, T. h. boettgeri, T. h. hercegovinensis).
Natur und Tier Verlag, Münster, 167 pp.

Pritchard, Peter C. H. (1997)
The Galápagos tortoises: An introduction to their nomenclature and survival status.
Reptile & Amphibian Magazine 1997 (Jan./Feb.): 64-70

Walls, Jerry G. (1996)
Kinixys: the forest tortoises.
Reptile Hobbyist 1 (3): 76-81

Wermuth,H. & Mertens,R. (1977)
Liste der rezenten Amphibien und Reptilien:
Testudines, Crocodylia, Rhynchocephalia
Das Tierreich, Lfg. 100, XXVII + 174 pp.
Walter de Gruyter, Berlin-New York

Zug,G.R.; Vitt, L.J. & Caldwell, J.P. (2001)
Herpetology, 2nd ed.
Academic Press San Diego, London, [...]XIV + 630 pp.

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