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Checklist of Chelonians of the World

Compiled by UWE FRITZ and PETER HAVAŠ
at the request of the CITES Nomenclature Committee and the German Agency for Nature Conservation

Här är listan över landsköldpaddor.

Aldabrachelys Loveridge & Williams, 1957

Comment: Frazier (2006) reviewed the confusing history of the usage of the generic names Aldabrachelys Loveridge & Williams, 1957 and Dipsochelys Bour, 1982. Using molecular genetic data, Austin et al. (2003) and Palkovacs et al. (2003) provided evidence that all non-Madagascan taxa within Aldabrachelys represent only a single species.

Aldabrachelys gigantea (Schweigger, 1812)

Distribution: Aldabra, Granitic Seychelles.

Astrochelys Gray, 1873

Astrochelys radiata (Shaw, 1802)

Distribution: Southwestern and southern Madagascar (southwestern Antandroy Territory along the coast between Amboasary and Morambe); introduced on Mauritius and Réunion.

Astrochelys yniphora (Vaillant, 1885)

Distribution: Northwestern Madagascar (vicinity of Baly Bay).

Chelonoidis Fitzinger, 1835

Chelonoidis carbonaria (Spix, 1824)

Distribution: Southermost central America (southeastern Panama), part of northern half of South America (west of the Andes in Chocó of Colombia and east of the Andes (Colombia, Venezuela, and the Guyanas to eastern Brazil, south to Rio de Janeiro, and west to eastern Bolivia, Paraguay, and northern Argentina), Trinidad and some islands of the Lesser Antilles.

Chelonoidis chilensis (Gray, 1870)

Distribution: South-central South America (southwestern Bolivia, western Paraguay, and northwestern Argentina southward to about 40°S in northern Patagonia).

Comment: Taxonomy of the Chelonoidis chilensis complex is still in flux. Freiberg (1973) described the large-sized Geochelone donosobarrosi from the southernmost part of the range of Ch. chilensis (Gray, 1870) and the small-sized G. petersi from within the northernmost part of the range of Ch. chilensis. While Buskirk (1993) failed to differentiate Ch. chilensis and Ch. donosobarrosi (Freiberg, 1973) morphologically, Cei (1986) treated all three taxa as valid species. Cabrera (1998) and Richard (1999) recognized only two species. Cabrera (1998) used the name G. petersi Freiberg, 1973 for northern populations and G. chilensis for southern populations (with G. donosobarrosi Freiberg, 1973 as junior synonym). Richard (1999) used the name G. chilensis for northern populations (with G. petersi as junior synonym) and G. donosobarrosi for southern populations. Ernst et al. (2000) suggested that the Chelonian description of G. petersi could have been based on small-sized males of Ch. chilensis. Considering the well-known phenotypic plasticity of other testudinids (Fritz et al. 2005a, 2006b, 2007), we feel it appropriate to recognize only a monotypic Ch. chilensis until a phylogeographic investigation is undertaken to determine geographic variation.

Chelonoidis denticulata (Linnaeus, 1766)

Distribution: Northern half of South America east of the Andes, Trinidad Island.

Chelonoidis nigra (Quoy & Gaimard, 1824)

Distribution: Galápagos Islands (Ecuador).

Chelonoidis nigra abingdonii (Günther, 1877)

Distribution: Formerly Abingdon Island (Pinta), Galápagos Islands; extinct in the wild, one captive survivor (Pritchard 1996).

Comment: Many references for the name ephippium refer to tortoises from Duncan Island.

Chelonoidis nigra becki (Rothschild, 1901)

Distribution: Northern and western slopes of Volcán Wolf, Albemarle Island (Isabela), Galápagos Islands (Pritchard 1996).

Chelonoidis nigra darwini (Van Denburgh, 1907)

Distribution: James Island (San Salvador or Santiago), Galápagos Islands; tortoises on Jervis Island (Rábida) are thought to be introduced (Pritchard 1996).

Chelonoidis nigra duncanensis Garman, 1996

Distribution: Duncan Island (Pinzón), Galápagos Islands (Pritchard 1996).

Comment: The Duncan Island subspecies is often referred to under the name Geochelone or Chelonoidis nigra ephippium (Günther, 1875).

Chelonoidis nigra hoodensis (Van Denburgh, 1907)

Distribution: Hood Island (Española), Galápagos Islands (Pritchard 1996).

Chelonoidis nigra phantastica (Van Denburgh, 1907)

Distribution: Narborough Island (Fernandina), Galápagos Islands (Pritchard 1996).

Chelonoidis nigra porteri (Rothschild, 1903)

Distribution: Southern Indefatigable Island (Santa Cruz), Galápagos Island (Pritchard 1996).

Comment: For this subspecies the name nigrita is often used. Russello et al. (2005) demonstrated that a cryptic second taxon exists on Indefatigable.

Chelonoidis nigra vicina (Günther, 1875)

Distribution: Southern and central Albemarle Island (Isabela), Galápagos Islands, from Iguana Cove, Cerro Azul, to Sierra Negra, Volcan Alcedo, and to Tagus Cove, Volcan Darwin (Pritchard 1996).

Comment: Often giant tortoises from Volcán Alcedo (central Albemarle) and Volcán Darwin (northern Albemarle) are treated as distinct subspecies (vandenburghi and microphyes, respectively). For tortoises from southwestern Albemarle the name elephantopus is often used.

Chersina Gray, 1831

Chersina angulata (Schweigger, 1812)

Distribution: Coastal regions of the Cape Provinces, South Africa; Namibian records are questionable.

Geochelone Fitzinger, 1835

Geochelone elegans (Schoepff, 1794)

Distribution: Pakistan, peninsular India from Orissa in the east and Sind and Kutch in the west southward to the tip, Sri Lanka.

Geochelone platynota (Blyth, 1863)

Distribution: Central Myanmar.

Geochelone sulcata (Miller, 1779)

Distribution: From Ethiopia and Sudan westward through the dry regions of Chad, Niger, and Mali to southern Mauritania and Senegal.

Gopherus Rafinesque, 1832

Gopherus agassizii (Cooper, 1863)

Distribution: Southwestern U.S.A. (southern Nevada, extreme southwestern Utah, southwestern California, western Arizona.) and adjacent Mexico (northern Baja California, cape region of Baja California Sur, western Sonora including Tiburón Island in the Gulf of California, northwestern Sinaloa).

Gopherus berlandieri (Agassiz, 1857)

Distribution: Texas, U.S.A. and north-eastern Mexico (Tamaulipas, northeastern San Luis Potosí, northern and eastern Nuevo León, and Coahuila).

Gopherus flavomarginatus Legler, 1959

Distribution: North-central Mexico (Bolson de Mapimi of southeastern Chihuahua, southwestern Coahuila, and northeastern Durango).

Gopherus polyphemus (Daudin, 1801)

Distribution: Southeastern U.S.A. (southwestern South Carolina south along the Atlantic Coastal Plain through Florida, and west along the Gulf Coastal Plain to southeastern Louisiana).

Homopus Duméril & Bibron, 1835

Homopus areolatus (Thunberg, 1787)

Distribution: Northern, Western, and Eastern Cape Provinces of South Africa.

Homopus boulengeri Duerden, 1906

Distribution: Karoo Plain of South Africa.

Homopus femoralis Boulenger, 1888

Distribution: Western part of Western Cape, Eastern Cape, southwest Free State Provinces, and possibly extreme southwest North-West Province, South Africa.

Homopus signatus (Gmelin, 1789)

Distribution: Western and southwestern South Africa; southern Namibia.

Homopus signatus signatus (Gmelin, 1789)

Distribution: Western South Africa and southern Namibia.

Homopus signatus cafer (Daudin, 1801)

Distribution: Southwestern South Africa.

Indotestudo Lindholm, 1929

Indotestudo elongata (Blyth, 1853)

Distribution: From Nepal, Bangladesh, and northeastern India (Jalpaiguri, West Bengal, and Singhbhum in Bihar) southward through Myanmar, Laos, Thailand, Cambodia, and from Guangxi, China to Penang, Malaysia.

Indotestudo forstenii (Schlegel & Müller, 1844)

Distribution: Sulawesi and Halmahera, Indonesia.

Indotestudo travancorica (Boulenger, 1907)

Distribution: Southwestern India (states of Karnataka, Kerala and Tamil Nadu).

Kinixys Bell, 1827

Kinixys belliana (Gray, 1831)

Distribution: From Somalia southward to Swaziland and Kwazulu-Natal and westward into the Democratic Republic of Congo and across West Africa to Senegal; Madagascar.

Kinixys belliana belliana (Gray, 1831)

Distribution: Eastern Africa from Somalia and Ethiopia to the northeastern Democratic Republic of Congo southward to Uganda, southern Malawi, Zimbabwe, and Zululand; northwestern Madagascar (vicinity of Bay of Ambaro, Ambanja, Nosy Faly, likely to be introduced there; Bour 1985).

Kinixys belliana nogueyi (Lataste, 1886)

Distribution: West Africa from Senegal eastward to Cameroon and the Central African Republic.

Kinixys erosa (Schweigger, 1812)

Distribution: West Africa (from Gambia eastward to Uganda, and southward to Cabinda and northern Angola).

Kinixys homeana Bell, 1827

Distribution: West Africa (from Liberia eastward to eastern Democratic Republic of Congo).

Kinixys lobatsiana (Power, 1927)

Distribution: Southeastern Botswana and northern Republic of South Africa.

Kinixys natalensis Hewitt, 1935

Distribution: Southern Mozambique, Swaziland and eastern Republic of South Africa.

Kinixys spekii Gray, 1863

Distribution: Angola eastward to southern Kenya and northeastern Republic of South Africa.

Malacochersus Lindholm, 1929

Malacochersus tornieri (Siebenrock, 1903)

Distribution: Patchy, highly localized distribution in Kenya, Tanzania and extreme northeastern Zambia as reviewed by Chansa & Wagner (2006).

Manouria Gray, 1852

Manouria emys (Schlegel & Müller, 1844)

Distribution: Northeastern India, Myanmar, and from Thailand through Malaysia to Sumatra andBorneo.

Manouria emys emys (Schlegel & Müller, 1844)

Distribution: Southern Thailand (provinces of Ranong and Nakorn Sri Thammarat) through Malaysia to Sumatra and Borneo; this subspecies ranges as far north as the Khao Luang Massif of Nakhon Sri Thammarat Province, being separated from Manouria emys phayrei (Blyth, 1853) by a tectonic slide fault area, the Surat gap (Wirot 1979, Ernst et al. 2000, P.P. van Dijk, pers. comm.).

Manouria emys phayrei (Blyth, 1853)

Distribution: Northeastern India (Assam, Meghalaya, and Nagaland), Myanmar, and from Tak Province (northern Thailand) as far south as Ranong and western Surat Thani Province, and may extend into northern Phang-nga Province, Thailand (Wirot 1979, Ernst et al. 2000, P.P. van Dijk, pers. comm.).

Manouria impressa (Günther, 1882)

Distribution: From the Karenni Hills in Myanmar to Malaysia and Vietnam.

Psammobates Fitzinger, 1835

Psammobates geometricus (Linnaeus, 1758)

Distribution: Southwestern Western Cape Province, South Africa.

Psammobates oculifer (Kuhl, 1820)

Distribution: Southern Africa from Northern Province, North-West, Mpumalanga and the western Free State northwestward through the Kalahari Desert of Botswana and Namibia almost to Angola.

Psammobates tentorius (Bell, 1828)

Distribution: From the Great Namaqualand of Namibia southeastward to Eastern Cape, South Africa.

Psammobates tentorius tentorius (Bell, 1828)

Distribution: Southern and eastern Karoo from Grahamstown to Matjiesfontein, Republic of South Africa.

Psammobates tentorius trimeni (Boulenger, 1886)

Distribution: Extreme western Cape Provinces of the Republic of South Africa from Lambert's Bay north to beyond the Orange River in Great Namaqualand in Namibia.

Psammobates tentorius verroxii (Smith, 1839)

Distribution: From the Northern Cape Province of the Republic of South Africa northwestward to the Great Namaqualand of Namibia; perhaps southern Free State.

Pyxis Bell, 1827

Pyxis arachnoides Bell, 1827

Distribution: Southwestern and southern coast of Madagascar.

Pyxis arachnoides arachnoides Bell, 1827

Distribution: Southern part of southwestern coast of Madagascar in the vicinity of the Onilahy River.

Pyxis arachnoides brygooi (Vuillemin & Domergue, 1972)

Distribution: Northern part of southwestern coast of Madagascar, southwest of the Mangoky River, between Morombe, Lake Ihotry, and Fanemotra Bay.

Pyxis arachnoides oblonga Gray, 1869

Distribution: Southern coast of Madagascar from La Linta to Lake Anony.

Pyxis planicauda (Grandidier, 1867)

Distribution: Southwestern coast of Madagascar, between the Monrondava and Tsiribihina Rivers.

Stigmochelys Gray, 1873

Stigmochelys pardalis (Bell, 1828)

Distribution: Sub-Saharan Africa, from southern Sudan and Ethiopia southward through eastern Africa to South Africa and westward to southern Angola and Namibia.

Comment: Until recently, Stigmochelys pardalis (Bell, 1828) was included by most authors in the genus Geochelone Fitzinger, 1835 (e.g. Loveridge & Williams 1957, Pritchard 1979, Ernst et al. 2000). According to mtDNA and nDNA data, S. pardalis forms a well-supported clade with the much smaller Psammobates spp. (Le et al. 2006) and not with other African and Malagasy tortoises of Geochelone sensu lato, as suggested by morphology (Gerlach 2001). We do not follow the suggestion of Le et al. (2006) to place S. pardalis into the genus Psammobates Fitzinger, 1835 however, acknowledging the most likely sister group relation of the large-sized S. pardalis to the dwarfed Psammobates spp. and the considerable morphological differences.

Pronounced sequence divergences of samples of Stigmochelys pardalis pardalis and S. p. Babcocki studied by Le et al. (2006) support distinction of both subspecies.

Stigmochelys pardalis pardalis (Bell, 1828)

Distribution: In Namibia possibly as far north as Rehoboth, more certainly south of a line from Kolmanskop (near Luederitz Bay) to Keetmanshoop south to the Cape (Loveridge & Williams 1957).

Stigmochelys pardalis babcocki (Loveridge, 1935)

Distribution: Sudan and Ethiopia south to Natal, west through Cape Province to Namibia, where it intergrades with Stigmochelys pardalis pardalis, but is dominant north of 27°S and in southern Angola; absent from northern Angola (Loveridge & Williams 1957).

Testudo Linnaeus, 1758

Testudo graeca Linnaeus, 1758

Distribution: Southern Spain and Balearic Islands; northern Africa (from Morocco to Libya), southeastern Balkans, Caucasus and Turkey eastward to Iran and through Syria, Lebanon and Jordan southward to Israel; introduced to some northern Mediterranean islands.

Comment: A forthcoming publication (Fritz et al. 2007) provides evidence that many taxa within the Testudo graeca complex are invalid and that all taxa are conspecific. For the time being, and to avoid unnecessary taxonomic upheaval, we treat all currently recognized taxa within the T. graeca complex as subspecies. See also comment under Testudo hermanni Gmelin, 1789 below.

Testudo graeca graeca Linnaeus, 1758

Distribution: Northeastern Morocco (eastern part of Mediterranean coast and adjacent inland regions),

Algeria (Pieh & Perälä 2004); introduced on several islands in the northern Mediterranean.

Testudo graeca anamurensis Weissinger, 1987

Distribution: Levant without central and southern coastal part (Perälä 2002a).

Testudo graeca armeniaca Chkhikvadze & Bakradze, 1991

Distribution: Araxes Valley and adjacent regions in Turkey, Armenia, Azerbaijan, and Iran (Pieh et al. 2002).

Testudo graeca buxtoni Boulenger, 1921

Distribution: Southwestern corner of Caspian Sea, Iran (Perälä 2004).

Testudo graeca cyrenaica Pieh & Perälä, 2002

Distribution: Cyrenaica Peninsula, Libya (Pieh & Perälä 2002).

Testudo graeca floweri Bodenheimer, 1935

Distribution: Coastal plains of Gaza, Israel and Lebanon (Perälä 2002a, Perälä & Shacham 2004).

Testudo graeca ibera Pallas, 1814

Distribution: Southeastern Balkan Peninsula, Asia Minor, part of Caucasus Region (Buskirk et al. 2001).

Testudo graeca lamberti Pieh & Perälä, 2004

Distribution: Northern Morocco (western part of Mediterranean coast; Pieh & Perälä 2004).

Testudo graeca marokkensis Pieh & Perälä, 2004

Distribution: Northwestern Morocco (Atlantic coast and adjacent inland region; Pieh & Perälä 2004).

Testudo graeca nabeulensis (Highfield, 1990)

Distribution: Tunisia, adjacent Libya (Pieh & Perälä 2002).

Testudo graeca nikolskii Chkhikvadze & Tuniyev, 1986

Distribution: Northern Black Sea coast of Russia and Georgia (Buskirk et al. 2001).

Testudo graeca pallasi Chkhikvadze & Bakradze, 2002

Distribution: Caspian Sea coastal area in Dagestan (Russia) and adjacent Azerbaijan (Danilov et al. 2004).

Testudo graeca perses Perälä, 2002

Distribution: Zagros Mountains (Iran) and adjacent mountain chains (Perälä 2002a).

Testudo graeca soussensis Pieh, 2001

Distribution: Southwestern Morocco, region of Marrakech and region southwest of High Atlas Mountains (Pieh & Perälä 2004).

Testudo graeca terrestris Forsskål, 1775

Distribution: Northern Mesopotamia (Bour & Perälä 2004, Perälä & Bour 2004).

Testudo graeca zarudnyi Nikolsky, 1896

Distribution: Eastern Iran (Perälä 2002a).

Testudo hermanni Gmelin, 1789

Distribution: Most of the southern and central Balkan Peninsula; Apennine Peninsula, Corsica, Sardinia, Sicily, relict populations on the Spanish and French Mediterranean coast; introduced on Balearic Islands (Cheylan 2001).

Comment: The nomenclatural history of Testudo hermanni Gmelin, 1789 is overshadowed by name confusions, leading to a considerable mixing-up of data for different taxa in many publications.

Before Flower (1925, 1926) put forward that the name Testudo hermanni Gmelin, 1789 has to beapplied to the "Greek tortoise", this species was known for decades under the name Testudo graeca Linnaeus, 1758, while the name Testudo ibera Pallas, 1814 was used for spur-thighed tortoises that are now known as Testudo graeca. This leads until today to confusion of data for both involved species. After the pioneering work of Wermuth (1952), demonstrating the existence of two subspecies within Testudo hermanni, the western subspecies was recognized for more than three decades as Testudo hermanni robertmertensi Wermuth, 1952 and the eastern subspecies as Testudo hermanni hermanni Gmelin, 1789. Bour (1987) rediscovered the type specimen of Testudo hermanni Gmelin, 1789, a representative of the western subspecies, and restricted its type locality to Collobrières, Département Var, France. This resulted in a reversal of the meaning of the subspecies name Testudo hermanni hermanni, which was no longer valid for the eastern, but for the western subspecies, and the resurrection of the name Testudo hermanni boettgeri Mojsisovic, 1889 for the eastern subspecies. This situation contributed to much upheaval because several authors did not notice the change in the application of the nominotypical subspecies.

The elevation of both subspecies to full species level and distinction of several western Balkan populations as third distinct species (Testudo hercegovinensis Werner, 1899), as suggested by Perälä (2002b, 2004) and Bour (2004a, B) , was not confirmed by a rangewide phylogeography (Fritz et al. 2006b).

Testudo hermanni hermanni Gmelin, 1789

Distribution: Apennine Peninsula, Corsica, Sardinia, Sicily, relict populations on the Spanish and French Mediterranean coast; introduced on Balearic Islands (Chelyan 2001).

Testudo hermanni boettgeri Mojsisovics, 1889

Distribution: Most of the southern and central Balkan Peninsula (Cheylan 2001).

Testudo horsfieldii Gray, 1844

Distribution: Kazakhstan, Turkmenistan, Uzbekistan, Kyrgyzstan and from Iran, Afghanistan, andPakistan to western Xinjiang, China.

Testudo horsfieldii horsfieldii Gray, 1844

Distribution: From Iran, Afghanistan, and Pakistan to western Xinjiang, China.

Testudo horsfieldii kazachstanica (Chkhikvadze, 1988)

Distribution: Kazakhstan and Turkmenistan.

Testudo horsfieldii rustamovi (Chkhikvadze, Amiranashvili & Ataev, 1990)

Distribution: Kopet-Dag region of southwestern Turkmenistan and western Kazakhstan.

Testudo kleinmanni Lortet, 1883

Distribution: Mediterranean coast of Libya and Egypt; Negev Desert (Israel).

Comment: Baha el Din (2006) synonymized Testudo werneri Perälä, 2001 with T. kleinmanni Lortet, 1883. This is in line with a completely lacking mtDNA differentiation of T. kleinmanni from western Libya compared to tortoises from the Negev Desert (Široký & Fritz in press).

Testudo marginata Schoepff, 1792

Distribution: Greece and adjacent Albania; introduced on Sardinia and perhaps in mainland Italy (Bringsøe et al. 2001).

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