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Chelonoidis

{{Short description|Genus of tortoises}}

{{Automatic taxobox

| fossil_range = {{geological range|21|0}}{{cite web|url=http://www.fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_name=Chelonoidis|title=Fossilworks:Chelonoidis}}

| image = Geochelone denticulata - couple Suriname.jpg

| image_caption = A pair of yellow-footed tortoises

| taxon = Chelonoidis

| authority = Fitzinger, 1835

| subdivision_ranks = Species

| subdivision = * C. carbonarius

}}

Chelonoidis is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835.{{cite encyclopedia | author1=Turtle Taxonomy Working Group | author-link1 = Turtle Taxonomy Working Group | last2 = Rhodin | first2 = A.G.J. | last3 = Iverson | first3 = J.B. | last4 = Bour | first4 = R. | last5 = Fritz | first5 = U. | last6 = Georges | first6 = A. | last7 = Shaffer | first7 = H.B. | last8 = van Dijk | first8 = P.P. | editor-last1 = Rhodin | editor-first1 = A.G.J. | editor-last2 = Iverson | editor-first2 = J.B. | editor-last3 = van Dijk | editor-first3 = P.P. | editor-last4 = Saumure | editor-first4 = R.A. | editor-last5 = Buhlmann | editor-first5 = K.A. | editor-last6 = Pritchard | editor-first6 = P.C.H. | editor-last7 = Mittermeier | editor-first7 = R.A. | encyclopedia = Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs | title = Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status | url = http://images.turtleconservancy.org/documents/2017/crm-7-checklist-atlas-v8-2017.pdf | access-date = 14 October 2017 | edition = 8th | year = 2017 | series = 7 | isbn = 978-1-5323-5026-9 | doi = 10.3854/crm.7.checklist.atlas.v8.2017 | pages = 1–292}}

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.{{Cite journal | last = Cione | first = A. L. | author2 = Tonni, E. P. | author3 = Soibelzon, L. | title = The Broken Zig-Zag: Late Cenozoic large mammal and tortoise extinction in South America | journal = Rev. Mus. Argentino Cienc. Nat. |series=Nueva Serie | volume = 5 | issue = 1 | pages = 1–19 | year = 2003 | issn = 1514-5158 | doi = 10.22179/REVMACN.5.26 | doi-access = free }}

Taxonomy

They were formerly assigned to Geochelone, but a 2006 genetic analysis indicated that they were actually most closely related to hingeback tortoises.{{cite journal|last=Le|first=M.|author2=Raxworthy, C. J.|author3=McCord, W. P.|author4=Mertz, L.|date=2006-05-05|title=A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes|url=http://www.iucn-tftsg.org/wp-content/uploads/file/Articles/Le_etal_2006.pdf|url-status=dead|journal=Molecular Phylogenetics and Evolution|volume=40|issue=2|pages=517–531|doi=10.1016/j.ympev.2006.03.003|pmid=16678445|archive-url=https://web.archive.org/web/20120316211953/http://www.iucn-tftsg.org/wp-content/uploads/file/Articles/Le_etal_2006.pdf|archive-date=2012-03-16}} However, a more recent genetic analysis of mtDNA has found that they are actually most closely related to the lineage containing Centrochelys and Geochelone. Their ancestors apparently floated across the Atlantic from Africa to South America in the Oligocene. 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. 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.{{cite journal|last1=Kehlmaier|first1=Christian|last2=Barlow|first2=Axel|last3=Hastings|first3=Alexander K.|last4=Vamberger|first4=Melita|last5=Paijmans|first5=Johanna L. A.|last6=Steadman|first6=David W.|last7=Albury|first7=Nancy A.|last8=Franz|first8=Richard|last9=Hofreiter|first9=Michael|last10=Fritz|first10=Uwe|date=11 January 2017|title=Tropical ancient DNA reveals relationships of the extinct Bahamian giant tortoise Chelonoidis alburyorum|journal=Proceedings of the Royal Society B|volume=284|issue=1846|pages=20162235|doi=10.1098/rspb.2016.2235|pmc=5247498|pmid=28077774}}

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.{{Cite journal|last1=Kehlmaier|first1=Christian|last2=Albury|first2=Nancy A.|last3=Steadman|first3=David W.|last4=Graciá|first4=Eva|last5=Franz|first5=Richard|last6=Fritz|first6=Uwe|date=2021-02-09|title=Ancient mitogenomics elucidates diversity of extinct West Indian tortoises|journal=Scientific Reports|language=en|volume=11|issue=1|pages=3224|doi=10.1038/s41598-021-82299-w|pmid=33564028|pmc=7873039|bibcode=2021NatSR..11.3224K|issn=2045-2322|doi-access=free}} This was followed by the Turtle Taxonomy Working Group and the Reptile Database in 2021.{{Cite web |title=Chelonoidis niger |url=https://reptile-database.reptarium.cz/species.php?genus=Chelonoidis&species=niger |access-date=2022-03-24 |website=The Reptile Database}}

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.

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.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. [http://www.iucn-tftsg.org/wp-content/uploads/file/Accounts/crm_5_000e_fossil_checklist_v1_2015.pdf Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians.] {{Webarchive|url=https://web.archive.org/web/20190718101604/http://www.iucn-tftsg.org/wp-content/uploads/file/Accounts/crm_5_000e_fossil_checklist_v1_2015.pdf|date=2019-07-18}} Chelonian Research Monographs 5(8) {{doi|10.3854/crm.5.000e.fossil.checklist.v1.2015}}

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.{{cite journal| title=A new species of extinct Late Quaternary giant tortoise from Hispaniola| year=2017| last1=Turvey| first1=Samuel T.| last2=Almonte| first2=Juan| last3=Hansford| first3=James| last4=Scofield| first4=R. Paul| last5=Brocca| first5=Jorge L.| last6=Chapman| first6=Sandra D.| journal=Zootaxa| volume=4277| issue=1| pages=1–16| pmid=30308657| doi=10.11646/zootaxa.4277.1.1| url=https://www.researchgate.net/publication/317604287 }}

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.

''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.{{cite journal |last1= Olson|first1=Storrs L |last2= David|first2=Normand |date=1 January 2014 |title=The gender of the tortoise genus Chelonoidis Fitzinger, 1835 (Testudines: Testudinidae)|journal=Proceedings of the Biological Society of Washington |volume=126 |issue=4 |pages=393–394 |doi=10.2988/0006-324X-126.4.393 |s2cid=83706022 }}

= Extant and recently extinct species =

Listed alphabetically:

Relationships of the genus, after Kehlmaier et al, 2017 and 2021.{{clade|{{clade

|label1=Chelonoidis

|1={{clade

|1={{clade

|1=Chelonoidis carbonarius (red-footed tortoise)

|2=Chelonoidis denticulatus (yellow-footed tortoise)

}}

|2={{clade

|label1=

|1={{clade

|1={{extinct}}Chelonoidis alburyorum (Bahamian giant tortoise)

}}

|label2=

|2={{clade

|1=Chelonoidis chilensis (Chaco tortoise)

|2=Chelonoidis niger (Galápagos tortoise)

}}

}}

}}

}}|style=font-size:100%;line-height:80%}}

= Fossil species =

== Late Quaternary fossil species ==

Listed alphabetically:

  • {{extinct}} C. lutzae – Lutz’s giant tortoise (Late Pleistocene of Argentina)
  • {{extinct}} C. marcanoi (Quaternary of the Dominican Republic) (nomen dubium as holotype fossil does not allow for distinguishing between species){{Cite web|title=Fossilworks: Chelonoidis marcanoi|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=356324|access-date=17 December 2021|website=fossilworks.org}}{{Cite journal|last1=Albury|first1=Nancy Ann|last2=Franz|first2=Richard (Professor)|last3=Rímoli|first3=Renato O.|last4=Lehman|first4=Phillip|last5=Rosenberger|first5=Alfred L.|date=2018-08-09|title=Fossil land tortoises (Testudines, Testudinidae) from the Dominican Republic, West Indies, with a description of a new species. |journal=American Museum Novitates|issue=3904|pages=1–28 |doi=10.1206/3904.1 |hdl=2246/6903|s2cid=92186485 |url=http://digitallibrary.amnh.org/handle/2246/6903|language=en-US}}
  • {{extinct}} C. pucara (Late Pleistocene of Buenos Aires province, Argentina){{cite journal |last1=Agnolín |first1=F.L. |last2=Chimento|first2=N.R.|year=2025|title=A giant among giants: a new land tortoise from the Pleistocene of the Argentine pampas

|journal=Rivista Italiana di Paleontologia e Stratigrafia |volume=131|issue=2|url=https://riviste.unimi.it/index.php/RIPS/article/view/27229|doi=10.54103/2039-4942/27229|doi-access=free}}

== Other fossil species ==

Listed alphabetically:{{Cite web|title=Mindat.org|url=https://www.mindat.org/taxon-5220265.html|access-date=2021-03-29|website=www.mindat.org}}

  • {{extinct}} C. australis (Miocene to Pliocene of Argentina){{Cite web|title=Fossilworks: Chelonoidis australis|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=105268|access-date=17 December 2021|website=fossilworks.org}}
  • {{extinct}} C. gallardoi (Miocene of Argentina, Late Pleistocene of Ecuador){{Cite web|title=Fossilworks: Chelonoidis gallardoi|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=105267|access-date=17 December 2021|website=fossilworks.org}}
  • † C. gringorum (Miocene of Argentina){{Cite web|title=Fossilworks: Chelonoidis gringorum|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=356116|access-date=17 December 2021|website=fossilworks.org}}
  • † C. hesternus (Miocene of Colombia){{Cite web|title=Fossilworks: Chelonoidis hesterna|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=132796|access-date=17 December 2021|website=fossilworks.org}}

References

{{Reflist}}

{{Testudinidae}}

{{Taxonbar|from=Q2080003}}

Category:Turtle genera

Category:Taxa named by Leopold Fitzinger