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Taxonomy of the family Testudinidae
Bibliography of the genus Chelonoidis
Biodiversity of the family Testudinidae








Genus
Chelonoidis

South American Tortoises

Südamerikanische Landschildkröten

Sydamerikanske Landskildpadder*

1835 Chelonoidis Fitzinger (type species: Testudo boiei Wagler 1833)
1870 Gopher Gray (type species: Testudo chilensis Gray 1870; syn. Fritz & Havas 2006)
1873 Elaphantopus Gray [not Elephantopus Agassiz 1846 (Siphonophora)] (type species: Testudo planiceps Gray 1854 [nomen dubium, not Testudo planiceps Schoepff 1801]; syn. Fritz & Havas 2006)
1929 Pampatestudo Lindholm (type species: Testudo chilensis Gray 1870; syn. Fritz & Havas 2006)

Contents: 13 species, of which 10 (76.9%) are endemic. In addition to this, 4 species are considered extinct (abingdonii, duncanensis, niger, phantasticus), all of which were endemic.
Remarks: Previously included in Testudo (e.g., Wermuth & Mertens 1977) and, more recently, Geochelone (e.g., King & Burke 1989). All Galapagos populations of Chelonoidis were previously regarded as subspecies or synonyms of a single species, C. niger (in older literature referred to as elephantopus). These populations are now regarded as a group of separate species, although some have been disputed. The nomenclature and taxonomy used herein follows Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk (2017). Olson & David (2014) pointed out that the grammatical gender of Chelonoidis is masculine.
Distribution: South America, Galapagos Islands, Panama, West Indies.
Reported from: Argentina, Bolivia, Brazil (Acre, Alagoas, Amapa, Amazonas, Bahia, Ceara, Distrito Federal, Espirito Santo, Goias, Maranhao, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Para [incl. Marajo Island], Paraiba, Pernambuco, Piaui, Rio de Janeiro, Rio Grande do Norte, Rondonia, Roraima, Sergipe, Tocantins), Colombia, Ecuador (incl. Galapagos Islands [Espanola, Isabela, Pinzon, San Cristóbal, Santa Cruz, Santiago]), French Guiana, Guyana, Panama, Paraguay, Peru, Saint Kitts & Nevis (Saint Kitts), Saint Lucia, Saint Vincent & the Grenadines (Saint Vincent), Surinam, Trinidad & Tobago (Trinidad), Venezuela.


Chelonoidis abingdonii (extinct)

Pinta Giant Tortoise, Abingdon Giant Tortoise, Floreana Giant Tortoise

Pinta-Riesenschildkröte

Pinta-elefantskildpadde

1877 Testudo abingdonii Günther
Testudo elephantopus abingdonii Mertens & Wermuth 1955
Geochelone abingdonii Pritchard [lapsus] 1967
Geochelone elephantopus abingdoni Pritchard 1967
Chelonoidis abingdonii Bour 1980
Chelonoidis elephantopus abingdoni Obst 1985
Geochelone nigra abingdoni Stubbs 1989
Chelonoidis nigra abingdonii David 1994

Remarks: See remarks under genus. This species became extinct on 24 June, 2012 (Dijk, Iverson, Shaffer, Bour & Rhodin 2012). However, wild hybrid populations with abingdoni ancestry were recently discovered on the remote Wolf volcano on Isabela, presumably descendants of tortoises translocated from Pinta by mariners (Miller, Quinzin, Poulakakis, Gibbs, Beheregaray, Garrick, Russello, Ciofi, Edwards, Hunter, Tapia, Rueda, Carrion, Valdivieso & Caccone 2017).
Distribution: Galapagos Islands (Pinta). Extinct.


Chelonoidis becki

Northern Isabela Giant Tortoise*, Volcán Wolf Giant Tortoise

Nördliche Isabela-Riesenschildkröte*, Wolf-Riesenschildkröte

Nordlig Isabela-elefantskildpadde*

1901 Testudo becki Rothschild
Geochelone becki Pritchard 1967
Geochelone elephantopus becki Pritchard 1967
Chelonoidis becki Bour 1980
Chelonoidis elephantopus becki Obst 1985
Geochelone nigra becki Stubbs 1989
Chelonoidis nigra becki Artner 2003

Remarks: See remarks under genus.
Distribution: Galapagos Islands (Isabela).


Chelonoidis carbonarius

South American Red-footed Tortoise

Köhlerschildkröte

Rødfodet Skovskildpadde

1824 Testudo carbonaria Spix
Geochelone carbonaria Williams 1960
Chelonoidis carbonaria Obst 1978
1831 Testudo hercules truncata Gray in Griffith & Pidgeon (Nowak-Kemp & Fritz 2010)
1833 Testudo boiei Wagler (Fritz & Havas 2006)
Geochelone boiei Fitzinger 1835
Chelonoidis boiei Fitzinger 1843

Remarks: Wild populations have been confirmed for Nicaragua, but whether they are native or introduced populations is still unclear (Salazar-Saavedra, Loza, Fernández, Martínez-Fonseca, Dwyer & Sunyer 2015), although some have regarded them as introduced (e.g., Köhler 2001; Sunyer & Köhler 2010; Sunyer 2014). Presence of natural populations in Trinidad highly questionable, but it may have been introduced (Murphy 1997). West Indian populations are presumably all introduced.
Introduced to: Anguilla (Anguilla Island), Antigua & Barbuda (incl. Antigua, Barbuda), Barbados, Brazil (Rio de Janeiro), British Virgin Islands, Colombia (Providencia), Dominica, Grenada, Guadeloupe (incl. Desirade, Iles de la Petite Terre [incl. Terre-de-Bas], Iles des Saintes, Marie-Galante), Martinique, Montserrat, Nicaragua (Islas del Maiz: Isla Grande del Maiz), Saba, Saint Barthelemy, Saint Kitts & Nevis (Saint Kitts), Saint Lucia, Saint Martin, Saint Vincent & the Grenadines (incl. Union Island), Sint Eustatius, Trinidad & Tobago (Trinidad), Venezuela (Los Testigos, Margarita), U.S. Virgin Islands (incl. St. Croix), USA (Florida).
Distribution: Argentina, Bolivia, Brazil (Alagoas, Amapa, Amazonas, Bahia, Ceara, Distrito Federal, Espirito Santo, Goias, Maranhao, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Para [incl. Marajo Island], Paraiba, Pernambuco, Piaui, Rio Grande do Norte, Rondonia, Roraima, Sergipe, Tocantins), Colombia, French Guiana, Guyana, Panama, Paraguay, Surinam, Venezuela.


Chelonoidis chathamensis

San Cristóbal Giant Tortoise, Chatham Island Giant Tortoise

San Cristobal-Riesenschildkröte*

San Cristobal-elefantskildpadde*

1907 Testudo chathamensis Van Denburgh
Testudo elephantopus chathamensis Mertens & Wermuth 1955
Geochelone chathamensis Pritchard 1967
Geochelone elephantopus chathamensis Pritchard 1967
Chelonoidis chathamensis Bour 1980
Chelonoidis elephantopus chathamensis Obst 1985
Geochelone nigra chathamensis Stubbs 1989
Chelonoidis nigra chathamensis David 1994

Remarks: See remarks under genus. Regarded as extinct by Fritz & Havas (2006), but as Endangered by Dijk, Iverson, Shaffer, Bour & Rhodin (2012).
Introduced to: Galapagos Islands (Rabida).
Distribution: Galapagos Islands (San Cristóbal).


Chelonoidis chilensis

Argentine Tortoise, Monte Tortoise

Patagonische Landschildkröte, Argentinische Landschildkröte

Argentinsk Landskildpadde*

1870 Testudo (Gopher) chilensis Gray
Geochelone chilensis Loveridge & Williams 1957
Chelonoidis chilensis Bour 1980
1870 Testudo argentina Sclater [substitute name for Testudo chilensis Gray 1870] (Fritz & Havas 2006)
1973 Geochelone donosobarrosi Freiberg (Fritz & Havas 2006)
Geochelone chilensis donosobarrosi Pritchard 1979
Chelonoidis chilensis donosobarrosi Obst 1985
Chelonoidis donosobarrosi Cei 1986
1973 Geochelone petersi Freiberg (Fritz & Havas 2006)
Chelonoidis chilensis petersi Obst 1985
Chelonoidis petersi Cei 1986

Other common names:
petersi: Chaco Tortoise
Remarks: Some authors recognize petersi as a separate species (e.g., Cei 1986; Bickham, Iverson, Parham, Philippen, Rhodin, Shaffer, Spinks & van Dijk 2007). Others recognize donosobarrosi as a separate species (e.g., Cei 1986). Fritz & Havas (2006) recognized only chilensis as a valid species for the time being, pending further studies.
Distribution: Argentina, Bolivia, Paraguay.


Chelonoidis darwini

Santiago Giant Tortoise, James Island Giant Tortoise

Santiago-Riesenschildkröte

Santiago-elefantskildpadde*

1907 Testudo darwini Van Denburgh
Geochelone darwini Pritchard 1967
Geochelone elephantopus darwini Pritchard 1967
Chelonoidis darwini Bour 1980
Chelonoidis elephantopus darwini Obst 1985
Geochelone nigra darwini Crumly in King & Burke 1989
Chelonoidis nigra darwinii David 1994
Chelonoidis elephantopus darwinii Rogner 1996

Remarks: See remarks under genus.
Distribution: Galapagos Islands (Santiago).


Chelonoidis denticulatus

South American Yellow-footed Tortoise, South American Forest Tortoise, South American Tortoise, Jaboty Tortoise

Waldschildkröte

Gulfodet Skovskildpadde

1766 Testudo denticulata Linnaeus
Chersine denticulata Merrem 1820
Geochelone denticulata Fitzinger 1835
Chelonoidis denticulata Fróes 1957
1782 Testudo tabulata Walbaum [nomen illegitimum] (Fritz & Havas 2006)
1792 Testudo tessellata Schneider (Fritz & Havas 2006)
Chersine tessellata Merrem 1820
1812 Testudo terrestris americana Schweigger (Fritz & Havas 2006)
1812 Testudo terrestris brasiliensis Schweigger (Fritz & Havas 2006)
1812 Testudo terrestris surinamensis Schweigger (Fritz & Havas 2006)
1812 Testudo terrestris var. caymanensis Schweigger (Fritz & Havas 2006)
1824 Testudo cagado Spix (Fritz & Havas 2006)
1824 Testudo hercules Spix (Fritz & Havas 2006)
1824 Testudo sculpta Spix (Fritz & Havas 2006)

Remarks: Records from eastern Brazil (Bahia, Distrito Federal, Espirito Santo, Goias, Minas Gerais, Rio de Janeiro, Rio Grande do Norte, Sergipe) may have been introduced (Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk 2017) and are regarded as such herein.
Introduced to: Brazil (Bahia, Distrito Federal, Espirito Santo, Goias, Minas Gerais, Rio de Janeiro, Rio Grande do Norte, Sergipe), Guadeloupe.
Distribution: Bolivia, Brazil (Acre, Amapa, Amazonas, Maranhao, Mato Grosso, Mato Grosso do Sul, Para [incl. Marajo Island], Rondonia, Roraima, Tocantins), Colombia, Ecuador, French Guiana, Guyana, Peru, Surinam, Trinidad & Tobago (Trinidad), Venezuela.


Chelonoidis donfaustoi

Eastern Santa Cruz Giant Tortoise

Östliche Santa-Cruz-Riesenschildkröte

Østlig Santa Cruz-elefantskildpadde*

2015 Chelonoidis donfaustoi Poulakakis, Edwards & Caccone in Poulakakis, Edwards, Chiari, Garrick, Russello, Benavides, Watkins-Colwell, Glaberman, Tapia, Gibbs, Cayot & Caccone

Remarks: Previously regarded as a population of porteri.
Distribution: Galapagos Islands (Santa Cruz).



Chelonoidis duncanensis (extinct)

Pinzón Giant Tortoise, Duncan Island Giant Tortoise

Pinzon-Riesenschildkröte

Pinzon-elefantskildpadde*

1875 Testudo ephippium Günther (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Testudo elephantopus ephippium Mertens & Wermuth 1955
Geochelone elephantopus ephippium Pritchard 1967
Geochelone ephippium Pritchard 1967
Chelonoidis ephippium Bour 1980
Chelonoidis elephantopus ephippium Obst 1985
Geochelone nigra ephippium Stubbs 1989
Chelonoidis nigra ephippium David 1994
1996 Geochelone nigra duncanensis Garman in Pritchard
Chelonoidis nigra duncanensis Artner 2003
Chelonoidis duncanensis Joseph-Ouni 2004

Remarks: See remarks under genus.
Distribution: Galapagos Islands (Pinzon).


Chelonoidis guntheri

Sierra Negra Giant Tortoise

Sierra-Negra-Riesenschildkröte*

Sierra Negra-elefantskildpadde*

1889 Testudo güntheri Baur [nomen dubium] (Nowak-Kemp & Fritz 2010)
Geochelone elephantopus guentheri Pritchard 1967
Geochelone guntheri Pritchard 1967
Chelonoidis guentheri Bour 1980
Chelonoidis elephantopus guentheri Obst 1985
Geochelone nigra guntheri Crumly in King & Burke 1989
Chelonoidis nigra guentheri David 1994

Remarks: See remarks under genus. Previously regarded as an invalid name (e.g., Dijk, Iverson, Rhodin, Shaffer & Bour 2014). Considered a valid species herein, following Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk (2017).
Distribution: Galapagos Islands (Isabela).


Chelonoidis hoodensis

Española Giant Tortoise, Hood Island Giant Tortoise

Española-Riesenschildkröte

Española-elefantskildpadde*

1907 Testudo hoodensis Van Denburgh
Testudo elephantopus hoodensis Mertens & Wermuth 1955
Geochelone elephantopus hoodensis Pritchard 1967
Geochelone hoodensis Pritchard 1967
Chelonoidis hoodensis Bour 1980
Chelonoidis elephantopus hoodensis Obst 1985
Geochelone nigra hoodensis Stubbs 1989
Chelonoidis nigra hoodensis David 1994

Remarks: See remarks under genus.
Distribution: Galapagos Islands (Espanola).


Chelonoidis microphyes

Volcán Darwin Giant Tortoise, Darwin Volcano Giant Tortoise, Tagus Cove Giant Tortoise

Volcán-Darwin-Riesenschildkröte*

Volcán Darwin-elefantskildpadde*

1875 Testudo microphyes Günther [nomen dubium] (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Geochelone elephantopus microphyes Pritchard 1967
Geochelone microphyes Pritchard 1967
Chelonoidis microphyes Bour 1980
Chelonoidis elephantopus microphyes Obst 1985
Geochelone nigra microphyes Crumly in King & Burke 1989
Chelonoidis nigra microphyes David 1994
1917 Testudo macrophyes Garman (Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk 2017)

Remarks: See remarks under genus. Previously regarded as a nomen dubium (e.g., Dijk, Iverson, Rhodin, Shaffer & Bour 2014). Considered a valid species herein, following Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk (2017).
Distribution: Galapagos Islands (Isabela).



Chelonoidis niger (extinct)

Black Giant Tortoise, Floreana Giant Tortoise, Charles Island Giant Tortoise, (Galapagos Tortoise)

Schwarze Riesenschildkröte*

Sort Elefantskildpadde*

1824 Testudo nigra Quoy & Gaimard [nomen dubium] (Olson 2015)
Chelonoidis nigra Bour 1985
Geochelone nigra Pritchard 1986
1824 Testudo californiana Quoy & Gaymard [nomen dubium] (Olson 2015)
Testudo californica Gray 1831 [error or unjustified emendation] (Olson 2015)
1827 Testudo elephantopus Harlan [nomen dubium] (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Geochelone elephantopus Pritchard 1967
Chelonoidis elephantopus Bour 1980
1889 Testudo galapagoensis Baur (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Testudo elephantopus galapagoensis Mertens & Wermuth 1955
Geochelone elephantopus galapagoensis Pritchard 1967
Chelonoidis galapagoensis Bour 1980
Chelonoidis elephantopus galapagoensis Obst 1985
Geochelone nigra galapagoensis Crumly in King & Burke 1989
Chelonoidis nigra galapagoensis David 1994

Remarks: Previously included all Galapagos populations of the genus (see remarks under genus). Pritchard (1996) revalidated nigra as the name to be used for this species, instead of elephantopus, used until then. Olson (2015) concluded that the taxa nigra and californiana are doubtful names (nomina dubia), thus currently unavailable names, based on the unknown origin of the type (same for both taxa). Consequently, the first available name becomes galapagoensis, however, Olson (2015) did not formally revalidate this name for the species, presumably awaiting molecular evidence that might determine the exact origin of nigra and californiana. The type of nigra, supposedly from California, arrived in Hawaii on the ship the Eagle. The Eagle could have stopped over in Galapagos, but there is no record of this (Olson 2015). Subsequently, Olson & Humphrey (2017) presented circumstantial evidence that elephantopus came from Floreana, thus suggesting that elephantopus should take precedence for the Floreana Giant Tortoise. Some authors have returned to using elephantopus rather than niger (e.g., Miller, Quinzin, Poulakakis, Gibbs, Beheregaray, Garrick, Russello, Ciofi, Edwards, Hunter, Tapia, Rueda, Carrion, Valdivieso & Caccone 2017). The name nigra is herein retained for the species, following Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk (2017). The species became extinct c. 1850 (Dijk, Iverson, Shaffer, Bour & Rhodin 2012). However, wild hybrid populations with niger ancestry were recently discovered on the remote Wolf volcano on Isabela, presumably descendants of tortoises translocated from Floreana by mariners. A captive breeding program has been initiated, which will, over the next decades, return niger tortoises to Floreana (Miller, Quinzin, Poulakakis, Gibbs, Beheregaray, Garrick, Russello, Ciofi, Edwards, Hunter, Tapia, Rueda, Carrion, Valdivieso & Caccone 2017).
Distribution: Galapagos Islands (Floreana). Extinct.


Chelonoidis phantasticus (extinct)

Fernandina Giant Tortoise, Narborough Island Giant Tortoise

Fernandina-Riesenschildkröte

Fernandina-elefantskildpadde*

1907 Testudo phantastica Van Denburgh
Testudo elephantopus phantastica Mertens & Wermuth 1955
Geochelone elephantopus phantastica Pritchard 1967
Geochelone phantastica Pritchard 1967
Chelonoidis phantastica Bour 1980
Chelonoidis elephantopus phantastica Obst 1985
Geochelone nigra phantastica Stubbs 1989
Chelonoidis nigra phantastica David 1994

Remarks: See remarks under genus. This species became extinct c. 1960 (Dijk, Iverson, Shaffer, Bour & Rhodin 2012).
Distribution: Galapagos Islands (Fernandina). Extinct.


Chelonoidis porteri

Western Santa Cruz Giant Tortoise, Indefatigable Island Giant Tortoise, La Caseta Giant Tortoise

Westliche Santa-Cruz-Riesenschildkröte

Vestlig Santa Cruz-elefantskildpadde*

1835 Testudo nigrita Dumeril & Bibron [nomen dubium] (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Testudo elephantopus nigrita Mertens & Wermuth 1955
Geochelone nigrita Pritchard 1967
Chelonoidis nigrita Bour 1980
Geochelone elephantopus nigrita Groombridge 1982
Chelonoidis elephantopus nigrita Obst 1985
Geochelone nigra nigrita Crumly in King & Burke 1989
Chelonoidis nigra nigrita David 1994
1903 Testudo porteri Rothschild
Geochelone elephantopus porteri Pritchard 1967
Geochelone porteri Fritts in Bowman, Beron & Leviton 1983
Geochelone nigra porteri Stubbs 1989
Chelonoidis nigra perteri [lapsus] Müller & Schmidt 1995

Remarks: See remarks under genus. Now restricted to La Reserva in western Santa Cruz. Previously included the eastern population now known as donfaustoi.
Distribution: Galapagos Islands (Santa Cruz).


Chelonoidis vandenburghi

Volcán Alcedo Giant Tortoise, Alcedo Volcano Giant Tortoise

Volcán-Alcedo-Riesenschildkröte*

Volcán Alcedo-elefantskildpadde*

1930 Testudo vandenburghi De Sola (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Geochelone elephantopus vandenburgi [lapsus] Pritchard 1967
Geochelone vandenburghi Pritchard 1967
Chelonoidis vandenburghi Bour 1980
Chelonoidis elephantopus vandenburghi Obst 1985
Geochelone nigra vandenburghi Crumly in King & Burke 1989
Chelonoidis nigra vandenburghi David 1994

Remarks: See remarks under genus. Previously regarded as a synonym of vicina (e.g., Dijk, Iverson, Rhodin, Shaffer & Bour 2014). Considered a valid species herein, following Rhodin, Iverson, Bour, Fritz, Georges, Shaffer & Dijk (2017).
Distribution: Galapagos Islands (Isabela).


Chelonoidis vicina

Southern Isabela Giant Tortoise, Cerro Azul Giant Tortoise, Iguana Cove Giant Tortoise, (Albemarle Island Giant Tortoise)

Südliche Isabela-Riesenschildkröte*, Wolf-Riesenschildkröte

Sydlig Isabela-elefantskildpadde*

1875 Testudo vicina Günther
Geochelone vicina Pritchard 1967
Geochelone nigra vicina Stubbs 1989
Chelonoidis elephantopus vicina Rogner 1996
Chelonoidis nigra vicina Artner 2003
Chelonoidis vicina Joseph-Ouni 2004

Remarks: See remarks under genus. Previously included guentheri, microphyes, and vandenburghi.
Distribution: Galapagos Islands (Isabela).