Chelonoidis Temporal range: 21–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N ^[1]

A pair of yellow-footed tortoises
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira
Superfamily:	Testudinoidea
Family:	Testudinidae
Genus:	Chelonoidis Fitzinger, 1835^[2]
Species
C. carbonarius C. chilensis C. denticulatus C. niger

Chelonoidis is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835.^[2] They are found in South America and the Galápagos Islands, and formerly had a wide distribution in the West Indies.

The multiple subspecies of the Galápagos tortoise (C. niger) are among the largest extant terrestrial chelonians. Giant members of the genus, such as Lutz's giant tortoise (C. lutzae) were also present in mainland South America and the West Indies during the Pleistocene, and the latter into the Holocene.^[3]

Taxonomy

They were formerly assigned to Geochelone, but a 2006 genetic analysis indicated that they were actually most closely related to hingeback tortoises.^[4] However, a more recent genetic analysis of mtDNA has found that they are actually most closely related to the lineage containing Centrochelys and Geochelone.^[5] Their ancestors apparently floated across the Atlantic from Africa to South America in the Oligocene.^[4] This crossing was made possible by their ability to float with their heads up and to survive up to six months without food or water.^[4] Based on mtDNA analysis, the extant Chelonoidis members can be divided into two lineages, with one containing the red-footed tortoise (C. carbonarius) and yellow-footed tortoise (C. denticulatus), and the other containing the Chaco tortoise (C. chilensis) and the Galapagos tortoises (C. niger). The now-extinct West Indian radiation is thought to group with the Chaco and Galapagos tortoises but is significantly basal to both, and was a rather evolutionary distinct lineage, having diverged well before any of the modern species in the genus did and only 7 mya after the divergence of Chelonoidis from African tortoises.^[6]

A 2021 study found that the extent of divergence among the species in the Galápagos and Bahamian Chelonoidis radiations may have been overestimated, and supported subsuming many of the species in both complexes to being subspecies of two parent species; C. alburyorum for the Bahamas and Turks and Caicos Islands, and C. niger for the Galápagos.^[5] This was followed by the Turtle Taxonomy Working Group and the Reptile Database in 2021.^[2]^[7]

The names of several species names in the genus have often been misspelled, beginning in the 1980s when Chelonoidis was elevated to genus and mistakenly treated as feminine, an error recognized and fixed in 2017.^[2]

Distribution

Presently, Chelonoidis are distributed throughout most of South America, as well as most of the Galápagos; the genus extended north into Central America during the Pleistocene and most of the West Indies up to the late Holocene. C. carbonarius and C. denticulatus are presently found on some of the Lesser Antillean islands, but the provenance of these individuals is unknown, and they could have been introduced by Amerindians during pre-Columbian times.

In the West Indies

In prehistoric times, a large insular radiation of giant Chelonoidis existed on many West Indian islands, including the Bahamas, Greater Antilles and Lesser Antilles. While some species such as those on Curaçao, Anguilla, and Barbados are thought to have gone extinct during the Late Pleistocene, other species such as those on the Greater Antilles and the Bahamas are thought to have been driven to extinction after the first humans arrived on the islands, from 7000 BC onwards. The Bahamian tortoise (C. alburyorum) was one of the last-surviving of these species, persisting up to 1170 AD on the Abacos, up to 1200 AD on Grand Turk, and up to 1400 AD on the Middle Caicos, just under a century prior to European colonization of the islands.^[6]^[8]

A 2017 study found that some of these species such as those from Hispaniola were specialists adapted to dry, open habitats such as Hispaniolan dry forests and had a major role in shaping them; following a decrease in the extent of these ecosystems after the end of the Pleistocene, these tortoises were restricted to refugia habitats up until their extinction.^[9]

A 2021 study identified two distinct genetic lineages within the Bahaman tortoise, C. alburyorum, but also sometimes found the remains of members of both lineages on the same island, even though the lineages would have only separated with geographic isolation. This indicates that the early inhabitants of the West Indies were successfully transporting the large tortoises across islands, presumably for the purpose of consumption, and thus causing the remains geographically-isolated lineages to co-occur on the same islands.^[5]

Chelonoidis species

Note that the genus name Chelonoidis is masculine under the rules of the ICZN, and adjectival species names must agree in gender; the species names below are displayed in keeping with this rule, and may differ from how they commonly appear in publications.^[10]

Extant and recently extinct species

Listed alphabetically:

C. carbonarius – red-footed tortoise^[2]
C. chilensis – Chaco tortoise^[2]
C. denticulatus – yellow-footed tortoise^[2]
C. niger – Galápagos tortoise – with the following subspecies:^[2]^[7]
- † C. n. abingdonii – Pinta Island tortoise (extinct as of 2012, but could be bred back from hybrids and/or persist as lone individuals)^[2]^[8]
- C. n. becki – Volcán Wolf giant tortoise^[2]
- C. n. chathamensis – San Cristobal giant tortoise^[2]
- C. n. darwini – Santiago Island giant tortoise^[2]
- C. n. donfaustoi – eastern Santa Cruz tortoise^[2]^[11]^[12]
- C. n. duncanensis – Pinzon Island giant tortoise^[2]
- C. n. guntheri – Sierra Negra giant tortoise^[2]
- C. n. hoodensis – Hood Island giant tortoise^[2]
- C. n. microphyes – Volcán Darwin giant tortoise^[2]
- † C. n. niger – Floreana giant tortoise (driven to extinction in the mid-19th century, but could be bred back from hybrids)^[2]
- C. n. phantasticus – Fernandina Island tortoise^[2]^[8] (rediscovered in February 2019)^[13]
- C. n. porteri – western Santa Cruz tortoise^[2]
- †C. n. 'Santa Fe Island lineage' – Santa Fe Island tortoise (undescribed, driven to extinction in the mid-19th century)^[14]
- C. n. vandenburghi – Volcán Alcedo giant tortoise^[2]
- C. n. vicina – Cerro Azul giant tortoise^[2]
† C. alburyorum – Bahamian tortoise (Late Holocene of The Bahamas and Turks & Caicos Islands)^[6]^[5]^[8]
† C. cubensis – Cuban giant tortoise (Early Holocene of Cuba)^[8]
† C. gersoni (Pleistocene-Holocene, Hispaniola)^[15]
† C. marcanoi^[16] (=C. dominicensis^[17]^[15]) (Pleistocene-Holocene, Hispaniola)
† C. monensis – Mona tortoise (Late Holocene of Isla de Mona, Puerto Rico)^[8]

Relationships of the genus, after Kehlmaier et al, 2017 and 2021.^[5]

Chelonoidis

	Chelonoidis carbonarius (red-footed tortoise)

	Chelonoidis denticulatus (yellow-footed tortoise)

	†Chelonoidis alburyorum (Bahamian giant tortoise)

	Chelonoidis chilensis (Chaco tortoise)

	Chelonoidis niger (Galápagos tortoise)

Fossil species

Late Quaternary fossil species

Listed alphabetically:

† C. lutzae – Lutz’s giant tortoise (Late Pleistocene of Argentina)^[8]
† C. marcanoi (Quaternary of the Dominican Republic) (nomen dubium as holotype fossil does not allow for distinguishing between species)^[9]^[18]^[17]
† C. sombrerensis – Sombrero giant tortoise (Late Pleistocene of Sombrero, Anguilla)^[8]

Other fossil species

Listed alphabetically:^[19]

† C. australis (Miocene to Pliocene of Argentina)^[20]
† C. gallardoi (Miocene of Argentina, Late Pleistocene of Ecuador)^[21]
† C. gringorum (Miocene of Argentina)^[22]
† C. hesternus (Miocene of Colombia)^[23]^[10]^[5]

References

^ "Fossilworks:Chelonoidis".
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v Turtle Taxonomy Working Group; Rhodin, A.G.J.; Iverson, J.B.; Bour, R.; Fritz, U.; Georges, A.; Shaffer, H.B.; van Dijk, P.P. (2017). "Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status" (PDF). In Rhodin, A.G.J.; Iverson, J.B.; van Dijk, P.P.; Saumure, R.A.; Buhlmann, K.A.; Pritchard, P.C.H.; Mittermeier, R.A. (eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs. 7 (8th ed.). pp. 1–292. doi:10.3854/crm.7.checklist.atlas.v8.2017. ISBN 978-1-5323-5026-9. Retrieved 14 October 2017.
^ Cione, A. L.; Tonni, E. P.; Soibelzon, L. (2003). "The Broken Zig-Zag: Late Cenozoic large mammal and tortoise extinction in South America". Rev. Mus. Argentino Cienc. Nat. Nueva Serie. 5 (1): 1–19. doi:10.22179/REVMACN.5.26. ISSN 1514-5158.
^ ^a ^b ^c Le, M.; Raxworthy, C. J.; McCord, W. P.; Mertz, L. (2006-05-05). "A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes" (PDF). Molecular Phylogenetics and Evolution. 40 (2): 517–531. doi:10.1016/j.ympev.2006.03.003. PMID 16678445. Archived from the original (PDF) on 2012-03-16.
^ ^a ^b ^c ^d ^e ^f Kehlmaier, Christian; Albury, Nancy A.; Steadman, David W.; Graciá, Eva; Franz, Richard; Fritz, Uwe (2021-02-09). "Ancient mitogenomics elucidates diversity of extinct West Indian tortoises". Scientific Reports. 11 (1): 3224. Bibcode:2021NatSR..11.3224K. doi:10.1038/s41598-021-82299-w. ISSN 2045-2322. PMC 7873039. PMID 33564028.
^ ^a ^b ^c Kehlmaier, Christian; Barlow, Axel; Hastings, Alexander K.; Vamberger, Melita; Paijmans, Johanna L. A.; Steadman, David W.; Albury, Nancy A.; Franz, Richard; Hofreiter, Michael; Fritz, Uwe (11 January 2017). "Tropical ancient DNA reveals relationships of the extinct Bahamian giant tortoise Chelonoidis alburyorum". Proceedings of the Royal Society B. 284 (1846): 20162235. doi:10.1098/rspb.2016.2235. PMC 5247498. PMID 28077774.
^ ^a ^b "Chelonoidis niger". The Reptile Database. Retrieved 2022-03-24.
^ ^a ^b ^c ^d ^e ^f ^g ^h Turtle Extinctions Working Group (Rhodin, A.G.J., Thomson, S., Georgalis, G., Karl, H.-V., Danilov, I.G., Takahashi, A., de la Fuente, M.S., Bourque, J.R., Delfino, M., Bour, R., Iverson, J.B., Shaffer, H.B., and van Dijk, P.P.). 2015. Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians. Archived 2019-07-18 at the Wayback Machine Chelonian Research Monographs 5(8) doi:10.3854/crm.5.000e.fossil.checklist.v1.2015
^ ^a ^b Turvey, Samuel T.; Almonte, Juan; Hansford, James; Scofield, R. Paul; Brocca, Jorge L.; Chapman, Sandra D. (2017). "A new species of extinct Late Quaternary giant tortoise from Hispaniola". Zootaxa. 4277 (1): 1–16. doi:10.11646/zootaxa.4277.1.1. PMID 30308657.
^ ^a ^b Olson, Storrs L; David, Normand (1 January 2014). "The gender of the tortoise genus Chelonoidis Fitzinger, 1835 (Testudines: Testudinidae)". Proceedings of the Biological Society of Washington. 126 (4): 393–394. doi:10.2988/0006-324X-126.4.393. S2CID 83706022.
^ Poulakakis, Nikos; Edwards, Danielle L.; Chiari, Ylenia; Garrick, Ryan C.; Russello, Michael A.; Benavides, Edgar; Watkins-Colwell, Gregory J.; Glaberman, Scott; Tapia, Washington; Gibbs, James P.; Cayot, Linda J.; Caccone, Adalgisa (2015). "Description of a New Galapagos Giant Tortoise Species (Chelonoidis; Testudines: Testudinidae) from Cerro Fatal on Santa Cruz Island". PLOS ONE. 10 (10): e0138779. Bibcode:2015PLoSO..1038779P. doi:10.1371/journal.pone.0138779. PMC 4619298. PMID 26488886.
^ Marris, Emma (21 October 2015). "Genetics probe identifies new Galapagos tortoise species". Nature News. doi:10.1038/nature.2015.18611. S2CID 182351587.
^ Van Denburgh, J. (1914). "The gigantic land tortoises of the Galapagos archipelago". Proceedings of the California Academy of Sciences. Series 4. 2 (1): 203–374. Retrieved 2012-01-11.
^ "Taxonomy browser (Chelonoidis Santa Fe island lineage)". www.ncbi.nlm.nih.gov. Retrieved 2021-03-29.
^ ^a ^b Viñola-López, Lázaro W; Almonte, Juan N (2022-07-22). "Revision of the fossil land tortoises (Testudines: Testudinidae) from Hispaniola with the description of a new species". Novitates Caribaea (20): 11–29. doi:10.33800/nc.vi20.302. ISSN 2079-0139.
^ Turvey, Samuel T.; Almonte, Juan; Hansford, James; Scofield, R. Paul; Brocca, Jorge L.; Chapman, Sandra D. (2017-06-15). "A new species of extinct Late Quaternary giant tortoise from Hispaniola". Zootaxa. 4277 (1): 1. doi:10.11646/zootaxa.4277.1.1. ISSN 1175-5334. PMID 30308657.
^ ^a ^b Albury, Nancy Ann; Franz, Richard (Professor); Rímoli, Renato O.; Lehman, Phillip; Rosenberger, Alfred L. (2018-08-09). "Fossil land tortoises (Testudines, Testudinidae) from the Dominican Republic, West Indies, with a description of a new species". American Museum Novitates (3904): 1–28. doi:10.1206/3904.1. hdl:2246/6903. S2CID 92186485.
^ "Fossilworks: Chelonoidis marcanoi". fossilworks.org. Retrieved 17 December 2021.
^ "Mindat.org". www.mindat.org. Retrieved 2021-03-29.
^ "Fossilworks: Chelonoidis australis". fossilworks.org. Retrieved 17 December 2021.
^ "Fossilworks: Chelonoidis gallardoi". fossilworks.org. Retrieved 17 December 2021.
^ "Fossilworks: Chelonoidis gringorum". fossilworks.org. Retrieved 17 December 2021.
^ "Fossilworks: Chelonoidis hesterna". fossilworks.org. Retrieved 17 December 2021.

Tortoise family

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Subclass: Anapsida
Order: Testudines
Suborder: Cryptodira
Superfamily: Testudinoidea
Family: Testudinidae

Genera

Species of the tortoise family

Agrionemys

Russian tortoise

Chaco tortoise
Red-footed tortoise
Yellow-footed tortoise
†Chelonoidis alburyorum
†Chelonoidis cubensis
†Chelonoidis lutzae
†Chelonoidis monensis
†Chelonoidis sellovii
†Chelonoidis sombrerensis

Galápagos tortoise

Chelonoidis complex

†Cheirogaster

†Cheirogaster bacharidisi
†Cheirogaster gymnesica
†Cheirogaster schafferi

Chersina

Angulate tortoise

†Hadrianus

†Hadrianus corsoni
†Hadrianus majusculus
†Hadrianus octonaria
†Hadrianus robustus
†Hadrianus schucherti
†Hadrianus utahensis
†Hadrianus vialovi

†Hesperotestudo

†Hesperotestudo percrassa

Pancake tortoise

Leopard tortoise

†Stylemys

†Stylemys botti
†Stylemys calaverensis
†Stylemys canetotiana
†Stylemys capax
†Stylemys conspecta
†Stylemys copei
†Stylemys emiliae
†Stylemys frizaciana
†Stylemys karakolensis
†Stylemys nebrascensis
†Stylemys neglectus
†Stylemys oregonensis
†Stylemys pygmea
†Stylemys uintensis
†Stylemys undabuna

Testudo

Phylogenetic arrangement of turtles based on turtles of the world 2017 update: Annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status. Key: †=extinct.

Testudines

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Diapsida
Order: Testudines

Genus

Chelonioidea
(Sea turtles)

Cheloniidae	†Allopleuron Caretta †Carolinochelys Chelonia †Eochelone Eretmochelys †Gigantatypus †Glarichelys †Itilochelys Lepidochelys †Mexichelys †Miocaretta Natator †Pacifichelys †Syllomus †Tasbacka
Dermochelyidae	†Arabemys †Corsochelys †Cosmochelys Dermochelys †Eosphargis †Mesodermochelys †Psephophorus
	†Euclastes †Peritresius †Procolpochelys †Protosphargis †Puppigerus

Kinosternoidea

Dermatemydidae	Dermatemys
Kinosternidae	Claudius †Hoplochelys Kinosternon Staurotypus Sternotherus

Testudinoidea

Emydidae	†Acherontemys Chrysemys Clemmys Deirochelys Emys Actinemys Emydoidea Glyptemys Graptemys Malaclemys Pseudemys Terrapene Trachemys †Wilburemys
Geoemydidae	Batagur †Banhxeochelys Cuora Cyclemys Geoclemys Geoemyda Hardella Heosemys Leucocephalon Malayemys Mauremys Melanochelys Morenia Notochelys Orlitia Pangshura Rhinoclemmys Sacalia Siebenrockiella Vijayachelys
Platysternidae	Platysternon
Testudinidae	Aldabrachelys Astrochelys Centrochelys Chelonoidis Chersina Cylindraspis †Cymatholcus †Floridemys Geochelone Gopherus †Hadrianus †Hesperotestudo Homopus Indotestudo Kinixys Malacochersus Manouria †Megalochelys †Oligopherus Psammobates Pyxis †Solitudo Stigmochelys †Stylemys Testudo

Trionychia

Carettochelyidae	†Allaeochelys †Anosteira Carettochelys
Trionychidae	Amyda Apalone †Axestemys Chitra Cyclanorbis Cycloderma †Drazinderetes Dogania †Gilmoremys †Hutchemys †Khunnuchelys Lissemys Nilssonia Palea †Palaeoamyda Pelochelys Pelodiscus Rafetus Trionyx
	†Basilochelys †Sinaspideretes

Chelydridae	Chelydra †Chelydrops †Chelydropsis †Emarginachelys †Macrocephalochelys Macrochelys †Planiplastron †Protochelydra
†Nanhsiungchelyidae	†Anomalochelys †Basilemys †Jiangxichelys
†Protostegidae	†Alienochelys †Archelon †Atlantochelys †Bouliachelys †Calcarichelys †Cratochelone †Desmatochelys †Iserosaurus †Notochelone †Ocepechelon †Pneumatoarthrus †Protostega †Rhinochelys †Santanachelys †Terlinguachelys
	†Adocus †Argillochelys †Bashuchelys †Ctenochelys †Prionochelys †Toxochelys

Pleurodira

†Araripemydidae	†Araripemys
†Bothremydidae	†Araiochelys †Arenila †Azabbaremys †Bothremys †Cearachelys †Chedighaii †Chupacabrachelys †Eotaphrosphys †Foxemys †Galianemys †Ilatardia †Inaechelys †Itapecuruemys †Jainemys †Kinkonychelys †Kurmademys †Labrostochelys †Nigeremys †Phosphatochelys †Polysternon †Puentemys †Rosasia †Rhothonemys †Sankuchemys †Taphrosphys †Ummulisani †Zolhafah
Chelidae	Acanthochelys Chelodina Chelus Elseya Elusor Emydura Hydromedusa †Lomalatachelys Mesoclemmys Myuchelys Phrynops Platemys †Prochelidella Pseudemydura Rheodytes Rhinemys †Yaminuechelys
Pelomedusidae	Pelomedusa Pelusios
Podocnemididae	†Albertwoodemys †Bauruemys †Brontochelys †Caninemys †Carbonemys †Cerrejonemys †Cordichelys Erymnochelys †Lapparentemys †Latentemys Peltocephalus Podocnemis †Stupendemys
†Sahonachelyidae	†Sahonachelys †Sokatra

	†Caribemys †Caririemys †Tacuarembemys

Phylogenetic arrangement of turtles based on Turtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status. † = extinct.

See also List of Testudines families