Testudininae

 Radiated tortoises Asterochelys radiata photographed in Madagascar by czwg.

Belongs within: Testudinidae.
Contains: Geochelone.

The Testudininae contains the land tortoises most closely related to the type genus, Testudo, with many species having been referred in the past to either that genus or Geochelone. Among the members of the group are the giant tortoises of the Seychelles islands in the genus Aldabrachelys. The genus Testudo includes species found in the Mediterranean region and western Asia, including Hermann’s tortoise T. hermanni which is commonly kept in captivity around Europe. Astrochelys includes two species native to Madagascar. Kinixys, the hinged tortoises, is a genus of African tortoises that possess a hinge in the rear part of the shell allowing the rear legs to be covered after retraction.

Synapomorphies (from Gaffney & Meylan 1988): Process of surangular interdigitates with dentary, lower jaw relatively gracile; mental glands absent; medial centrale large, separating distal end of radius from contact with medial proximal carpals.

<==Testudininae [Megalochelyina, Megalochelyini, Testudinina, Testudinini]
    |  i. s.: Protestudo Chkhikvadze 1970FB-E07
    |         Dipsochelys Bour 1982N10
    |           |--D. abrupta (Grandidier 1868) [=Geochelone abrupta]N10
    |           |--D. arnoldi Bour 1982N10
    |           |--D. daudinii (Dumeril & Bibron 1835) [=Geochelone daudinii]N10
    |           |--D. giganteusWW10
    |           `--D. hololissa (Gunther 1877) [=Geochelone hololissa]N10
    |--+--GeocheloneFB-E07
    |  `--+--+--+--Chersina Gray 1831FB-E07
    |     |  |  |    `--C. angulata (Schweigger 1812)FB-E07, JPG04
    |     |  |  `--Homopus Duméril & Bibron 1835FB-E07
    |     |  |       |--H. areolatus (Thunberg 1787)JPG04
    |     |  |       |--H. boulengeri Duerden 1906JPG04
    |     |  |       |--H. femoralis Boulenger 1888JPG04
    |     |  |       `--H. signatus (Gmelin 1789)FB-E07
    |     |  `--+--Stigmochelys Gray 1873FB-E07
    |     |     |    `--*S. pardalis (Bell 1828)FB-E07, JPG04 [=Geochelone pardalisFB-E07, Psammobates pardalisFB-E07]
    |     |     |         |--S. p. pardalisFB-E07
    |     |     |         `--‘Geochelone’ p. babcockiFB-E07
    |     |     `--Psammobates Fitzinger 1835FB-E07
    |     |          |--P. geometricus (Linnaeus 1758)JPG04
    |     |          |--P. oculifer (Kuhl 1820)FB-E07
    |     |          `--P. tentorius (Bell 1828)JPG04
    |     `--+--Astrochelys Gray 1873 [incl. Angonoka Le et al. 2006]FB-E07
    |        |    |--*A. radiata (Shaw 1802)FB-E07, JPG04 [=Geochelone radiataFB-E07, Testudo radiataUSDI77]
    |        |    `--A. yniphora (Vaillant 1885)FB-E07, JPG04 (see below for synonymy)
    |        `--+--+--AcinixysGM88
    |           |  `--Pyxis Bell 1827FB-E07
    |           |       |--P. arachnoides Bell 1827JPG04
    |           |       `--P. planicauda (Grandidier 1867)JPG04
    |           `--+--MegalochelysGM88
    |              `--Aldabrachelys Loveridge & Williams 1957FB-E07, N10
    |                   |--A. abrupta (Grandidier 1868)FB-E07
    |                   |--A. elephantinaCP03
    |                   |--A. gigantea (Schweigger 1812)ICZN13 (see below for synonymy)
    |                   |    |--A. g. giganteaH81
    |                   |    `--‘Geochelone’ g. gouffei (Rothschild 1906)H81
    |                   |--A. grandidieri (Vaillant 1885)M94, N10 (see below for synonymy)
    |                   `--A. robustaM94
    `--+--+--Malacochersus Lindholm 1929FB-E07
       |  |    `--M. tornieri (Siebenrock 1903)FB-E07
       |  `--Indotestudo Lindholm 1929FB-E07
       |       |--I. forstenii (Schlegel & Müller 1844)FB-E07, JPG04
       |       `--+--I. elongata (Blyth 1853)FB-E07, JPG04
       |          `--I. travancorica (Boulenger 1907)FB-E07, JPG04
       `--Testudo Linnaeus 1758FB-E07, L58 (see below for synonymy)
            |  i. s.: T. abingdoniiS99
            |         T. antakyensis Perälä 1996TK03
            |         T. carinata Linnaeus 1758L58
            |         T. carolina Linnaeus 1758L58
            |         T. clausaJ23
            |         T. cultrata Cope 1873 (n. d.)H96
            |         T. denticulata Gray 1866G66
            |         T. ephippiumS99
            |         T. exornata Lambe 1906 (n. d.)H96
            |         T. forsteniB28
            |         T. klettiana Cope 1875C77
            |         T. loveridgiiLW08
            |         T. lutaria Linnaeus 1758L58
            |         T. microphycesS99
            |         T. orbicularis Linnaeus 1758L58 [=Emys orbicularisK08]
            |         T. perses Perälä 2002TK03
            |         T. pusilla Linnaeus 1758L58
            |         T. scabra Linnaeus 1758L58
            |         T. tabulata Gray 1866G66
            |         T. undata Cope 1875C77
            |         T. weissingeriCP03
            |         T. werneri Perälä 2001JPG04
            |--+--T. hermanni Gmelin 1789FB-E07, JPG04 (see below for synonymy)
            |  |    |--T. h. hermanniM55
            |  |    `--T. h. robertmertensiM55
            |  `--T. horsfieldii Gray 1844FB-E07, JPG04 [=*Agrionemys horsfieldiiFB-E07]
            `--+--*T. graeca Linnaeus 1758JPG04, FB-E07, TK03 [incl. T. ibera Pallas 1814FB-E07]
               |    |--T. g. graecaTK03
               |    |--T. g. anamurensis Weissinger 1987TK03
               |    |--T. g. armeniaca Chkhikvadze & Bakradze 1991TK03
               |    |--T. g. ibera Pallas 1814TK03
               |    `--T. g. terrestris Forsskål 1775TK03
               `--+--T. kleinmanni Lortet 1883FB-E07, JPG04
                  `--T. marginata Schoepff 1792FB-E07

Aldabrachelys gigantea (Schweigger 1812)ICZN13 [=Testudo giganteaICZN13, Geochelone (Aldabrachelys) giganteaICZN13; incl. Aldabrachelys arnoldiFB-E07, Testudo dussumieri Gray 1831 (nom. rej.)ICZN13, A. dussumieriFB-E07, Dipsochelys dussumieriICZN13, Geochelone dussumieriN10, Aldabrachelys hololissaFB-E07]

Aldabrachelys grandidieri (Vaillant 1885)M94, N10 [=Geochelone grandidieriN10, Dipsochelys grandidieriN10, Testudo grandidieriM94]

Astrochelys yniphora (Vaillant 1885)FB-E07, JPG04 [=*Angonoka yniphoraFB-E07, Geochelone yniphoraFB-E07, Testudo yniphoraUSDI77]

Testudo Linnaeus 1758FB-E07, L58 [incl. Agrionemys Khozatsky & Młynarski 1966FB-E07, Chersine Merrem 1820FB-E07, Eurotestudo Lapparent de Broin et al. 2006FB-E07, Medaestia Wussow 1916FB-E07]

Testudo hermanni Gmelin 1789FB-E07, JPG04 [=Agrionemys hermanniFB-E07, *Eurotestudo hermanniFB-E07, Protestudo hermanniFB-E07; incl. Testudo boettgeriFB-E07, Eurotestudo boettgeriFB-E07, T. graeca Merrem 1820 non Linnaeus 1758FB-E07, *Chersine graecaFB-E07, *Medaestia graecaFB-E07, T. hercegovinensisFB-E07, E. hercegovinensisFB-E07]

*Type species of generic name indicated

References

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[CP03] Claude, J., E. Paradis, H. Tong & J.-C. Auffray. 2003. A geometric morphometric assessment of the effects of environment and cladogenesis on the evolution of the turtle shell. Biological Journal of the Linnean Society 79: 485–501.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4(2): i–iv, 1–370.

[FB-E07] Fritz, U., & O. R. P. Bininda-Emonds. 2007. When genes meet nomenclature: tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone. Zoology 110: 298–307.

[GM88] Gaffney, E. S., & P. A. Meylan. 1988. A phylogeny of turtles. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 157–219. Clarendon Press: Oxford.

[G66] Gray, J. E. 1866. Notes on some young specimens of tortoises (Testudo). Proceedings of the Zoological Society of London 1866: 305–306.

[H81] Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141–158.

[H96] Hutchison, J. H. 1996. Testudines. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 337–353. Cambridge University Press.

[ICZN13] ICZN. 2013. Opinion 2316: Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea; Reptilia, Testudines): usage of the specific name conserved by maintenance of a designated neotype, and suppression of Testudo dussumieri Gray, 1831 (currently Dipsochelys dussumieri). Bulletin of Zoological Nomenclature 70 (1): 61–65.

[J23] James, E. 1823. Account of an Expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and ’20, by order of the Hon. J. C. Calhoun, sec’y of war: under the command of Major Stephen H. Long. From the notes of Major Long, Mr. T. Say, and other gentlemen of the exploring party vol. 1. H. C. Carey & I. Lea: Philadelphia.

[JPG04] Joyce, W. G., J. F. Parham & J. A. Gauthier. 2004. Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology 78 (5): 989–1013.

[K08] Korsós, Z. 2008. History of the herpetological collection of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 37–93.

[LW08] Li, C., X.-C. Wu, O. Rieppel, L.-T. Wang & L.-J. Zhao. 2008. An ancestral turtle from the Late Triassic of southwestern China. Nature 456: 497–501.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M94] MacPhee, R. D. E. 1994. Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bulletin of the American Museum of Natural History 220: 1–214.

[M55] Mertens, R. 1955. Die Amphibien und Reptilien der Insel Elba. Senckenbergiana Biologica 36: 287–296.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[S99] Stanley, S. M. 1999. Earth System History. W. H. Freeman and Company: New York.

[TK03] Türkozan, O., Y. Kumlutaş, H. Arikan, Ç. Ilgaz & A. Avcı. 2003. Morphological and serological comparison of Mediterranean spur-thighed tortoises, Testudo graeca, from the Aegean region and southeastern Turkey. Zoology in the Middle East 29: 41–50.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

[WW10] White, A. W., T. H. Worthy, S. Hawkins, S. Bedford & M. Spriggs. 2010. Megafaunal meiolaniid horned turtles survived until early human settlement in Vanuatu, southwest Pacific. Proceedings of the National Academy of Sciences of the USA 107 (35): 15512–15516.

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