Tortoise

{{Short description|Family of land turtles}} {{About|the reptile|other uses}} {{pp-move-indef}} {{Automatic taxobox | name = Testudinidae | fossil_range = {{Fossil range|Eocene|Recent}} | image = A. gigantea Aldabra Giant Tortoise.jpg | image_caption = Aldabra giant tortoise (''Aldabrachelys gigantea'') | taxon = Testudinidae

| authority = Batsch, 1788 | type_genus = ''Testudo'' | type_genus_authority = Linnaeus, 1758 | subdivision_ranks = Subgroups | subdivision = }}

'''Tortoises''' ({{IPAc-en|ˈ|t|ɔːr|t|ə|s|.|ᵻ|z}} {{respell|TOR|təs|iz}}) are reptiles of the family '''Testudinidae''' of the order Testudines (Latin for "tortoise"). Like other testudines, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like other members of the suborder Cryptodira, they retract their necks and heads directly backward into the shell to protect them.<ref>{{Cite web |last=Harris |first=Samantha |date= |title=How Strong Is A Tortoise Shell? |url=https://www.tortoiseknowledge.com/how-strong-is-a-tortoise-shell/ |access-date=23 January 2023 |website=Tortoise Knowledge |language=en-US}}</ref>

Tortoises can vary in size with some species, such as the Galápagos giant tortoise, growing to more than {{convert|1.2|m|ft}} in length, whereas others like the speckled Cape tortoise have shells that measure only {{convert|6.8|cm|in}} long.<ref>{{Cite book|title=encyclopedia of LIFE|publisher=Miles Kelly|year=2017|isbn=978-1-78617-327-0|page=211}}</ref> Several lineages of tortoises have independently evolved very large body sizes in excess of {{convert|100|kg|lbs}}, including the Galapagos giant tortoise and the Aldabra giant tortoise. They are usually diurnal animals with tendencies to be crepuscular depending on the ambient temperatures. They are generally reclusive animals. Tortoises are the longest-living land animals in the world, although the longest-living species of tortoise is a matter of debate. Galápagos tortoises are noted to live over 150 years, but an Aldabra giant tortoise named Adwaita may have lived an estimated 255 years. In general, most tortoise species can live 80–150 years. Tortoises are typically slow-moving.

==Terminology== Differences exist in usage of the common terms turtle, tortoise, and terrapin, depending on the variety of English being used; usage is inconsistent and contradictory.<ref name="Simoons">Simoons, Frederick J. (1991). ''Food in China: A Cultural and Historical Inquiry''. CRC Press. {{ISBN|084938804X}}. p. 358.</ref> These terms are common names and do not reflect precise biological or taxonomic distinctions.<ref name="WildlifeEncy">Burton, Maurice and Burton, Robert (2002). ''International Wildlife Encyclopedia''. Marshall Cavendish. {{ISBN|0761472665}}. p. 2796.</ref>thumb|class=skin-invert-image|Skeleton of a tortoise|210x210pxThe American Society of Ichthyologists and Herpetologists uses "turtle" to describe all species of the order Testudines, regardless of whether they are land-dwelling or sea-dwelling, and uses "tortoise" as a more specific term for slow-moving terrestrial species.<ref name="Simoons" /> General American usage agrees; turtle is often a general term; tortoise is used only in reference to terrestrial turtles or, more narrowly, only those members of Testudinidae, the family of modern land tortoises; and terrapin may refer to turtles that are small and live in fresh and brackish water, in particular the diamondback terrapin (''Malaclemys terrapin'').<ref>Orenstein, Ronald Isaac (2001). ''Turtles, Tortoises and Terrapins: Survivors in Armor''. Firefly Books. {{ISBN|1770851194}}</ref><ref name="SanDiegoZoo">{{cite web|url=http://www.sandiegozoo.org/animalbytes/t-turtle.html|title=Turtle|publisher=Sandiegozoo.org|access-date=2012-09-16|archive-url=https://web.archive.org/web/20101206125855/http://www.sandiegozoo.org/animalbytes/t-turtle.html|archive-date=2010-12-06}}</ref><ref name="NCA">[http://www.ncaquariums.com/ask-the-aquarium/what-is-the-difference-between-turtles-terrapins-and-tortoises What is the difference between turtles, terrapins, and tortoises?] {{webarchive|url=https://web.archive.org/web/20150505004238/http://www.ncaquariums.com/ask-the-aquarium/what-is-the-difference-between-turtles-terrapins-and-tortoises|date=2015-05-05}}, North Carolina Aquariums (July 1997).</ref><ref name="Dawkins">Dawkins, Richard (2009). ''The Greatest Show on Earth: The Evidence for Evolution''. Free Press. {{ISBN|1416594795}}. p. 174.</ref> In America, for example, the members of the genus ''Terrapene'' dwell on land, yet are referred to as box turtles rather than tortoises.<ref name="WildlifeEncy" />

British and International English usage, by contrast, tends not to use "turtle" as a generic term for all members of the order, and also applies the term "tortoises" broadly to all land-dwelling members of the order Testudines, regardless of whether they are actually members of the family Testudinidae.<ref name="Dawkins" /> In Britain, terrapin is used to refer to a larger group of semiaquatic turtles than the restricted meaning in America.<ref name="SanDiegoZoo" /><ref name="Endangered">''Endangered Wildlife and Plants of the World'', Vol. 1. Marshall Cavenish. (2001). {{ISBN|0761471952}}. p. 1476.</ref>

Australian usage is different from both American and British usage.<ref name="Dawkins" /> Land tortoises are not native to Australia, and traditionally freshwater turtles have been called "tortoises" in Australia.<ref name="Romanowski">Romanowski, Nick (2010). ''Wetland Habitats: A Practical Guide to Restoration and Management''. CSIRO Publishing. {{ISBN|9780643096462}}. p. 134.</ref> Some Australian experts disapprove of this usage—believing that the term tortoises is "better confined to purely terrestrial animals with very different habits and needs, none of which are found in this country"—and promote the use of the term "freshwater turtle" to describe Australia's primarily aquatic members of the order Testudines because it avoids misleading use of the word "tortoise" and also is a useful distinction from marine turtles.<ref name="Romanowski" />

==Biology==

Most species of tortoises lay small clutch sizes, seldom exceeding 20 eggs, and many species have clutch sizes of only 1–2 eggs. Incubation is characteristically long in most species, the average incubation period are between 100 and 160.0 days. Egg-laying typically occurs at night, after which the mother tortoise covers her clutch with sand, soil, and organic material. The eggs are left unattended, and depending on the species, take from 60 to 120 days to incubate.<ref name="Incubation">{{cite web|author=Highfield, Andy|url=http://www.tortoisetrust.org/articles/eggfaq.html|title=Tortoise egg incubation|publisher=Tortoisetrust.org|access-date=2009-04-07|archive-url=https://web.archive.org/web/20130905231033/http://www.tortoisetrust.org/articles/eggfaq.html|archive-date=2013-09-05}}</ref> The size of the egg depends on the size of the mother and can be estimated by examining the width of the cloacal opening between the carapace and plastron. The plastron of a female tortoise often has a noticeable V-shaped notch below the tail which facilitates passing the eggs. Upon completion of the incubation period, a fully formed hatchling uses an egg tooth to break out of its shell. It digs to the surface of the nest and begins a life of survival on its own. They are hatched with an embryonic egg sac which serves as a source of nutrition for the first three to seven days until they have the strength and mobility to find food. Juvenile tortoises often require a different balance of nutrients than adults, so they may eat foods which a more mature tortoise would not. For example, the young of a strictly herbivorous species commonly will consume worms or insect larvae for additional protein.<ref>{{Cite web |title=Feeding your baby: 6–12 months |url=https://www.unicef.org/parenting/food-nutrition/feeding-your-baby-6-12-months |access-date=2022-07-13 |website=www.unicef.org |language=en}}</ref>

The number of concentric rings on the carapace, much like the cross-section of a tree, can sometimes give a clue to how old the animal is, but, since the growth depends highly on the accessibility of food and water, a tortoise that has access to plenty of forage (or is regularly fed by its owner) with no seasonal variation will have no noticeable rings. Moreover, some tortoises grow more than one ring per season, and in some others, due to wear, some rings are no longer visible.<ref>{{cite web|title=Shells: Anatomy and Diseases of Turtle and Tortoise Shells|url=http://www.peteducation.com/article.cfm?c=17+1797&aid=2700|website=PetEducation.com|publisher=Drs. Foster & Smith|archive-url=https://web.archive.org/web/20131023060541/http://www.peteducation.com/article.cfm?c=17+1797&aid=2700|archive-date=2013-10-23|access-date=2013-10-22}}</ref>

Tortoises generally have one of the longest lifespans of any animal, and some individuals are known to have lived longer than 150 years.<ref>{{cite journal|last1=Moon|first1=J. C.|last2=McCoy|first2=E. D.|last3=Mushinsky|first3=H. R.|last4=Karl|first4=S. A.|title=Multiple Paternity and Breeding System in the Gopher Tortoise, ''Gopherus polyphemus''|doi=10.1093/jhered/esj017|journal=Journal of Heredity|volume=97|issue=2|pages=150–157|year=2006|pmid=16489146|url=https://www.researchgate.net/publication/7288354|doi-access=free|citeseerx=10.1.1.572.6767}}</ref> Because of this, they symbolize longevity in some cultures, such as Chinese culture. The oldest tortoise ever recorded, and one of the oldest individual animals ever recorded, was Tu'i Malila, which was presented to the Tongan royal family by the British explorer James Cook shortly after its birth in 1777. Tu'i Malila remained in the care of the Tongan royal family until its death by natural causes on May 19, 1965, at the age of 188.<ref>{{cite news|url=https://www.foxnews.com/story/tortoise-believed-to-have-been-owned-by-darwin-dies-at-176|title=Tortoise Believed to Have Been Owned by Darwin Dies at 176|work=Fox News|agency=Associated Press|date=2006-06-26|url-status=live|archive-url=https://web.archive.org/web/20060701141233/http://www.foxnews.com/story/0,2933,200831,00.html|archive-date=July 1, 2006}}</ref>

The Alipore Zoo in India was the home to Adwaita, which zoo officials claimed was the oldest living animal until its death on March 23, 2006. Adwaita (also spelled Addwaita) was an Aldabra giant tortoise brought to India by Lord Wellesley, who handed it over to the Alipur Zoological Gardens in 1875 when the zoo was set up. West Bengal officials said records showed Adwaita was at least 150 years old, but other evidence pointed to 250. Adwaita was said to be the pet of Robert Clive.<ref>{{cite news|url=https://news.bbc.co.uk/2/hi/south_asia/4837988.stm|title='Clive of India's' tortoise dies|work=BBC News|date=2006-03-23|access-date=2009-04-07}}</ref>

Harriet was a resident at the Australia Zoo in Queensland from 1987 to her death in 2006; she was believed to have been brought to England by Charles Darwin aboard the ''Beagle'' and then on to Australia by John Clements Wickham.<ref name="SII1">{{cite journal|last1=Thomson|first1=S.|last2=Irwin|first2=S.|last3=Irwin|first3=T.|year=1995|title=Harriet, the Galapagos tortoise: disclosing one and a half centuries of history|journal=Intermontanus|volume=4|issue=5|pages=33–35}}</ref> Harriet died on June 23, 2006, just shy of her 176th birthday.

Timothy, a female spur-thighed tortoise, lived to be about 165 years old. For 38 years, she was carried as a mascot aboard various ships in Britain's Royal Navy. Then in 1892, at age 53, she retired to the grounds of Powderham Castle in Devon. Up to the time of her death in 2004, she was believed to be the United Kingdom's oldest resident.<ref>{{cite news|url=https://news.bbc.co.uk/2/hi/uk_news/england/devon/3607053.stm|title=Timmy the tortoise dies aged 160|date=2004-04-07|access-date=2019-05-30|language=en-GB}}</ref>

Jonathan, a Seychelles giant tortoise living on the island of St Helena, may be as old as {{Years ago|1832}} years.<ref>{{cite web|url=https://www.bbc.com/news/magazine-26543021|title= Meet Jonathan, St Helena's 182-year-old giant tortoise|publisher= BBC|date= March 13, 2014}}</ref>

DNA analysis of the genomes of the long-lived tortoises, Lonesome George, the iconic last member of ''Chelonoidis abingdonii'', and the Aldabra giant tortoise ''Aldabrachelys gigantea'' led to the detection of lineage-specific variants affecting DNA repair genes that might contribute to their long lifespan.<ref name="Quesada2019">{{cite journal |vauthors=Quesada V, Freitas-Rodríguez S, Miller J, Pérez-Silva JG, Jiang ZF, Tapia W, Santiago-Fernández O, Campos-Iglesias D, Kuderna LF, Quinzin M, Álvarez MG, Carrero D, Beheregaray LB, Gibbs JP, Chiari Y, Glaberman S, Ciofi C, Araujo-Voces M, Mayoral P, Arango JR, Tamargo-Gómez I, Roiz-Valle D, Pascual-Torner M, Evans BR, Edwards DL, Garrick RC, Russello MA, Poulakakis N, Gaughran SJ, Rueda DO, Bretones G, Marquès-Bonet T, White KP, Caccone A, López-Otín C |title=Giant tortoise genomes provide insights into longevity and age-related disease |journal=Nat Ecol Evol |volume=3 |issue=1 |pages=87–95 |date=January 2019 |pmid=30510174 |pmc=6314442 |doi=10.1038/s41559-018-0733-x }}</ref>

===Dimorphism=== Many species of tortoises are sexually dimorphic, though the differences between males and females vary from species to species.<ref>{{cite web|url=http://www.tortoisesupply.com/SexingYourTortoises|title=Sexing Your Tortoise|website=Tortoise Supply|access-date=13 February 2017}}</ref> In some species, males have a longer, more protruding neck plate than their female counterparts, while in others, the claws are longer on the females.

The male plastron is curved inwards to aid reproduction. The easiest way to determine the sex of a tortoise is to look at the tail. The females, as a general rule, have smaller tails, dropped down, whereas the males have much longer tails which are usually pulled up and to the side of the rear shell.

===Brain=== The brain of a tortoise is extremely small. Red-footed tortoises, from Central and South America, do not have an area in the brain called the hippocampus, which relates to emotion, learning, memory and spatial navigation. Studies have shown that red-footed tortoises may rely on an area of the brain called the medial cortex for emotional actions, an area that humans use for actions such as decision making.<ref>{{cite news|url=http://www.livescience.com/47155-tortoise-touchscreen-learning.html|title=Tortoises Show Off Smarts by Mastering Touch-Screen Tech|newspaper=Live Science|access-date=2016-12-01}}</ref>

==Distribution== Tortoises are found from southern North America to southern South America, around the Mediterranean basin, across Eurasia to Southeast Asia, in sub-Saharan Africa, Madagascar, and some Pacific islands. They are absent from Australasia. They live in diverse habitats, including deserts, arid grasslands, and scrub to wet evergreen forests, and from sea level to mountains. Most species, however, occupy semiarid habitats.

Many large islands are or were characterized by species of giant tortoises. Part of the reason for this is that tortoises are good at oceanic dispersal. Despite being unable to swim, tortoises are able to survive long periods adrift at sea because they can survive months without food or fresh water. Tortoises have been known to survive oceanic dispersals of more than 740 km.<ref>{{cite journal|last1=Gerlach|first1=Justin|last2=Muir|first2=Catharine|last3=Richmond|first3=Matthew D.|date=28 December 2006|title=The first substantiated case of trans-oceanic tortoise dispersal|journal=Journal of Natural History|volume=40|issue=41–43|pages=2403–2408|doi=10.1080/00222930601058290|bibcode=2006JNatH..40.2403G |s2cid=86037101|url=https://zenodo.org/record/5230669}}</ref> Once on islands tortoises faced few predators or competitors and could grow to large sizes and become the dominant large herbivores on many islands due to their low metabolic rate and reduced need for fresh water compared to mammals.<ref>{{cite book |last1=Goodman |first1=Steven M.|last2=Jungers|first2=William L. |title=Extinct Madagascar: picturing the island's past |date=2014 |location=Chicago |isbn=978-0-226-14397-2}}</ref> Today there are only two living species of giant tortoises, the Aldabra giant tortoise on Aldabra Atoll and the dozen subspecies of Galapagos giant tortoise found on the Galapagos Islands. However, until recently giant tortoises could be found on nearly every major island group, including the Bahamas, the Greater Antilles (including Cuba and Hispaniola), the Lesser Antilles, the Canary Islands, Malta, the Seychelles, the Mascarene Islands (including Mauritius and Reunion), and Madagascar. Most of these tortoises were wiped out by human arrival. Many of these giant tortoises are not closely related (belonging to different genera such as ''Megalochelys'', ''Chelonoidis'', ''Centrochelys'', ''Aldabrachelys'', ''Cylindraspis'', and ''Hesperotestudo''), but are thought to have independently evolved large body size through convergent evolution. Giant tortoises are notably absent from Australasia and many south Pacific islands, but the distantly related meiolaniid stem turtles are thought to have filled the same niche. Giant tortoises are also known from the Oligocene-Pliocene of mainland North America, South America, Europe, Asia, and Africa, but are all now extinct, which is also attributed to human activity.<ref>{{cite journal |last1=Cione |first1=Alberto |last2=Tonni |first2=Eduardo |last3=Soibelzon |first3=Leooldo |title=The Broken Zig-Zag: Late Cenozoic large mammal and tortoise extinction in South America |journal=Revista del Museo Argentino de Ciencias Naturales |series=Nueva Serie |year=2003 |volume=5 |issue=1 |pages=1–19|doi=10.22179/REVMACN.5.26 |doi-access=free }}</ref>[[File:Gopher Tortoise snacking on Opuntia ( Nopales ) cactus at Smyrna Dunes Park - Flickr - Andrea Westmoreland.jpg|alt=Tortoise feeding on a cactus|thumb|Gopher tortoise feeding on cactus]]

==Diet== Tortoises are generally considered to be strict herbivores, feeding on grasses, weeds, leafy greens, flowers, and some fruits. However, hunting and eating of birds has been observed on occasion.<ref>{{Cite journal|last1=Zora|first1=Anna|last2=Gerlach|first2=Justin|date=2021-08-23|title=Giant tortoises hunt and consume birds|journal=Current Biology|language=English|volume=31|issue=16|pages=R989–R990|doi=10.1016/j.cub.2021.06.088|issn=0960-9822|pmid=34428417|doi-access=free|bibcode=2021CBio...31.R989Z }}</ref> Pet tortoises typically require diets based on wild grasses, weeds, leafy greens and certain flowers. Certain species consume worms or insects and carrion in their normal habitats. Too much protein is detrimental in herbivorous species, and has been associated with shell deformities and other medical problems. Different tortoise species vary greatly in their nutritional requirements.

== Behavior == Communication in tortoises is different from many other reptiles. Because they are restricted by their shell and short limbs, visual communication is not a strong form of communication in tortoises. Tortoises use olfactory cues to determine the sex of other tortoises so that they can find a potential mate. Tactile communication is important in tortoises during combat and courtship. In both combat and courtship, tortoises use ramming to communicate with other individuals.<ref>{{Cite journal|last=Auffenberg|first=Walter|date=1977-02-01|title=Display Behavior in Tortoises|journal=American Zoologist|volume=17|issue=1|pages=241–250|doi=10.1093/icb/17.1.241|issn=0003-1569|doi-access=free}}</ref>

==Taxonomy== This species list largely follows Turtle Taxonomy Working Group (2021)<ref>{{Cite book |last=Rhodin |first=Anders G.J. |title=Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (9th Ed.). |date=2021-11-15 |publisher=Chelonian Research Foundation and Turtle Conservancy |isbn=978-0-9910368-3-7 |series=Chelonian Research Monographs|volume=8 |doi=10.3854/crm.8.checklist.atlas.v9.2021 |s2cid=244279960 }}</ref> and the Turtle Extinctions Working Group (2015).<ref>{{cite book |collaboration=Turtle Extinctions Working Group |author1=Rhodin, A.G.J. |author2=Thomson, S. |author3=Georgalis, G. |author4=Karl, H.-V. |author5=Danilov, I.G. |author6=Takahashi, A. |author7=de la Fuente, M.S. |author8=Bourque, J.R. |author9=Delfino M. |author10=Bour, R. |author11=Iverson, J.B. |author12=Shaffer, H.B. |author13=van Dijk, P.P. |title=Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians |issue=8 |pages=000e.1–66 |volume=5 |doi=10.3854/crm.5.000e.fossil.checklist.v1.2015 |year=2015 |series=Chelonian Research Monographs |isbn=978-0-9653540-9-7 |doi-access=free }}</ref> [[File:TortoiseSkeleton.jpg|thumb|A skeleton of Aldabra giant tortoise found in Cousin Island (Seychelles).]][[File:Radiated Tortoise (Astrochelys radiata) (10292092024).jpg|thumb|Radiated tortoise (''Astrochelys radiata'')]]'''Family Testudinidae''' Batsch 1788<ref name=bat88>Batsch, A.J.G.C. (1788). ''Versuch einer Anleitung zur Kenntniss und Geschichte der Thiere und Mineralien. Erster Theil. Allgemeine Geschichte der Natur; besondre der Säugthiere, Vögel, Amphibien und Fische.'' Jena: Akademischen Buchandlung, 528 pp.</ref> * {{Extinct}}'''''Alatochelon'''''<ref>{{cite journal|author1=Adán Pérez-García |author2=Evangelos Vlachos |author3=Xabier Murelaga |year=2020 |title=A large testudinid with African affinities in the post-Messinian (lower Pliocene) record of south-eastern Spain |journal=Palaeontology |volume=63 |issue= 3|pages= 497–512|doi=10.1111/pala.12468 |s2cid=214232312 |hdl=11336/146829 |hdl-access=free }}</ref> ** {{Extinct}} ''Alatochelon myrteum'' * '''''Aldabrachelys''''' Loveridge and Williams 1957:166<ref name=lov&will>{{cite journal|author=Loveridge, Arthur |author2= Williams, Ernest E.|year=1957|title= Revision of the African tortoises and turtles of the suborder Cryptodira|journal= Bulletin of the Museum of Comparative Zoology |volume=115|issue=6|pages=163–557|url=http://biostor.org/reference/871}}</ref> ** ''Aldabrachelys gigantea'' Aldabra giant tortoise. *** ''A. g. gigantea'' Aldabra tortoise. *** ''A. g. arnoldi'' Arnold's giant tortoise. *** ''A. g. daudinii'' Daudin's giant tortoise. *** ''A. g. hololissa'' Domed Seychelles giant tortoise. ** †''Aldabrachelys abrupta'' Late Holocene, extinct ''circa'' 1200 AD ** †''Aldabrachelys grandidieri'' Late Holocene, extinct ''circa'' 884 AD * [[File:Chelonoidis carbonarius 386013561.jpg|thumb|Red-footed tortoise (''Chelonoidis carbonarius'')]][[File:Chelonoidis porteri 114750622.jpg|thumb|Galápagos giant tortoise (''Chelonoidis niger'')]]'''''Astrochelys''''' Gray, 1873:4<ref name=gray1873>{{cite journal|author=Gray, John Edward|title= Notes on the genera of turtles (''Oiacopodes''), and especially on their skeletons and skulls|url=http://biostor.org/reference/59893|journal= Proceedings of the Zoological Society of London|year= 1873|volume= 1873|pages=395–411}}</ref> ** ''Astrochelys radiata'', radiated tortoise ** ''Astrochelys yniphora'', angonoka tortoise, (Madagascan) plowshare tortoise * '''''Centrochelys''''' Gray 1872:5<ref name=gray1872>Gray, John Edward. (1872). "Appendix to the Catalogue of Shield Reptiles in the Collection of the British Museum. Part I. Testudinata (Tortoises)". London: British Museum, 28 pp.</ref> ** {{Extinct}} ''Centrochelys atlantica'' ** {{Extinct}} ''Centrochelys burchardi'' Tenerife giant tortoise ** {{Extinct}} ''Centrochelys marocana'' ** {{Extinct}} ''Centrochelys robusta'' Maltese giant tortoise ** ''Centrochelys sulcata'', African spurred tortoise, sulcata tortoise ** {{Extinct}} ''Centrochelys vulcanica'' Gran Canaria giant tortoise * '''''Chelonoidis''''' Fitzinger 1835:112<ref name=fits1835/> ** {{Extinct}} ''Chelonoidis alburyorum'' Abaco tortoise, Late Pleistocene, extinct ''c.'' 1400 CE ** ''Chelonoidis carbonarius'', red-footed tortoise ** ''Chelonoidis chilensis'', Chaco tortoise, Argentine tortoise or southern wood tortoise ** {{Extinct}} ''Chelonoidis cubensis'' Cuban giant tortoise ** ''Chelonoidis denticulatus'' Brazilian giant tortoise, yellow-footed tortoise **{{extinct}} ''C. dominicensis'' Dominican giant tortoise<ref>{{Cite web|title=Fossilworks: Chelonoidis dominicensis|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=374950|access-date=17 December 2021|website=fossilworks.org}}</ref> **{{Extinct}} ''Chelonoidis lutzae'' Lutz's giant tortoise, Late Pleistocene ** {{Extinct}} ''Chelonoidis monensis'' Mona tortoise ** [[File:Angulate tortoise (Chersina angulata) 1.jpg|thumb|Angulate tortoise (''Chersina angulata'')]]''Chelonoidis niger'' Galapagos giant tortoise<ref>[http://reptile-database.reptarium.cz/advanced_search?taxon=Testudinidae&submit=Search Testudinidae], The Reptile Database</ref> ** {{Extinct}} ''Chelonoidis sellovii'' Southern Cone giant tortoise, Pleistocene ** {{Extinct}} ''Chelonoidis sombrerensis'' Sombrero giant tortoise, Late Pleistocene * [[File:Homopus signatus - the worlds smallest tortoise.jpg|thumb|Speckled padloper (''Chersobius signatus'')]]'''''Chersina''''' Gray 1830:5 ** ''Chersina angulata'', angulated tortoise, South African bowsprit tortoise * {{Extinct}} '''''Cheirogaster''''' Bergounioux 1935:78 ** †''Cheirogaster gymnesica'' Late Pliocene to Early Pleistocene ** †''Cheirogaster schafferi'' Pliocene to Early Pleistocene * [[File:Ergilemys insolitus.jpg|thumb|†''Ergilemys insolitus'' fossil]]'''''Chersobius''''' Fitzinger, 1835 ** ''Chersobius boulengeri'', Karoo padloper, Karoo dwarf tortoise, Boulenger's Cape tortoise ** ''Chersobius signatus'', speckled padloper tortoise ** ''Chersobius solus'', Nama padloper, Berger's Cape tortoise * †'''''Cylindraspis''''' Fitzinger 1835:112<ref name=fits1835>{{cite journal|author=Fitzinger, Leopold J. |year=1835|title= Entwurf einer systematischen Anordnung der Schildkröten nach den Grundsätzen der natürlichen Methode|journal=Annalen des Wiener Museums der Naturgeschichte|volume= 1|pages=105–128}}</ref> (all species extinct) following Austin and Arnold, 2001:<ref name=aa01>{{Cite journal | last1 = Austin | first1 = J. J. | last2 = Nicholas Arnold | first2 = E. | doi = 10.1098/rspb.2001.1825 | title = Ancient mitochondrial DNA and morphology elucidate an extinct island radiation of Indian Ocean giant tortoises (Cylindraspis) | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 268 | issue = 1485 | pages = 2515–23 | year = 2001 | pmid = 11749704| pmc = 1088909}}</ref> ** †''Cylindraspis indica'', synonym ''Cylindraspis borbonica'', Reunion giant tortoise ** †''Cylindraspis inepta'', saddle-backed Mauritius giant tortoise or Mauritius giant domed tortoise ** †''Cylindraspis peltastes'', domed Rodrigues giant tortoise ** †''Cylindraspis triserrata'', domed Mauritius giant tortoise or Mauritius giant flat-shelled tortoise ** †''Cylindraspis vosmaeri'', saddle-backed Rodrigues giant tortoise * [[File:Indian star tortoise (2016) 7365.jpg|thumb|Indian star tortoise (''Geochelone elegans'')]]{{Extinct}} '''''Ergilemys''''' Ckhikvadze, 1984<ref>{{cite web |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=341551&is_real_user=1 |title=Ergilemys |author=|website=Paleobiology Database |publisher=Paleobio DB |access-date=March 23, 2022 }}</ref> ** ''Ergilemys bruneti'' ** ''Ergilemys insolitus'' ** ''Ergilemys saikanensis'' * [[File:Gopher Tortoise at Smyrna Dunes Park - Flickr - Andrea Westmoreland (1).jpg|thumb|Gopher tortoise (''Gopherus polyphemus'')]]'''''Geochelone''''' Fitzinger 1835:112<ref name=fits1835 /> ** ''Geochelone elegans'', Indian star tortoise ** ''Geochelone platynota'', Burmese star tortoise * '''''Gopherus''''' Rafinesque 1832:64<ref name=raf32>{{cite journal|author=Rafinesque, Constantine Samuel|year=1832|title= Description of two new genera of soft shell turtles of North America|journal= Atlantic Journal and Friend of Knowledge |volume=1|pages=64–65}}</ref> ** ''Gopherus agassizii'', Mojave desert tortoise, Agassiz's desert tortoise ** ''Gopherus berlandieri'', Texas tortoise, Berlandier's tortoise ** ''Gopherus flavomarginatus'', Bolson tortoise ** ''Gopherus morafkai'', Sonoran desert tortoise, Morafka's desert tortoise ** ''Gopherus polyphemus'', gopher tortoise ** ''Gopherus evgoodei'', Sinaloan desert tortoise, Goode's thornscrub tortoise * {{Extinct}} '''''Hadrianus''''' ** ''Hadrianus corsoni'' (syn. ''H. octonarius'') ** ''Hadrianus robustus'' ** ''Hadrianus schucherti'' ** ''Hadrianus utahensis'' * {{Extinct}} '''''Hesperotestudo''''' ** ''Hesperotestudo alleni'' ** ''Hesperotestudo angusticeps'' ** ''Hesperotestudo brontops'' ** ''Hesperotestudo equicomes'' ** ''Hesperotestudo impensa'' ** ''Hesperotestudo incisa'' ** ''Hesperotestudo johnstoni'' ** ''Hesperotestudo kalganensis'' ** ''Hesperotestudo niobrarensis'' ** ''Hesperotestudo orthopygia'' ** ''Hesperotestudo osborniana'' ** ''Hesperotestudo percrassa'' ** ''Hesperotestudo riggsi'' ** ''Hesperotestudo tumidus'' ** ''Hesperotestudo turgida'' ** ''Hesperotestudo wilsoni'' * [[File:Homopus areolatus 166592733 (cropped).jpg|thumb|Parrot-beaked padloper (''Homopus areolatus'')]]'''''Homopus''''' Duméril and Bibron 1834:357<ref name=dum&bib34>Duméril, André Marie Constant and Bibron, Gab riel. 1834. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Premier. Paris: Roret, 439 pp.</ref> ** ''Homopus areolatus'', common padloper, parrot-beaked tortoise, beaked Cape tortoise ** ''Homopus femoralis'', greater padloper, greater dwarf tortoise * [[File:Indotestudo elongata 88503209.jpg|thumb|Elongated tortoise (''Indotestudo elongata'')]]'''''Indotestudo''''' Lindholm, 1929 ** ''Indotestudo elongata'', elongated tortoise, yellow-headed tortoise ** ''Indotestudo forstenii'', Forsten's tortoise, East Indian tortoise ** ''Indotestudo travancorica'', Travancore tortoise * [[File:Kinixys lobatsiana 15532559.jpg|thumb|Lobatse hinge-back tortoise (''Kinixys lobatsiana'')]]'''''Kinixys''''' ** ''Kinixys belliana'', Bell's hinge-back tortoise ** ''Kinixys erosa'', forest hinge-back tortoise, serrated hinge-back tortoise ** ''Kinixys homeana'', Home's hinge-back tortoise ** ''Kinixys lobatsiana'', Lobatse hinge-back tortoise ** ''Kinixys natalensis'', Natal hinge-back tortoise ** ''Kinixys spekii'', Speke's hinge-back tortoise * '''''Malacochersus''''' Lindholm 1929:285<ref name=lind29>{{cite journal|author=Lindholm, Wassili A.|year= 1929|title=Revidiertes Verzeichnis der Gattungen der rezenten Schildkröten nebst Notizen zur Nomenklatur einiger Arten|journal= Zoologischer Anzeiger |volume=81|pages=275–295}}</ref> ** ''Malacochersus tornieri'', pancake tortoise * [[File:Manouria impressa.jpg|thumb|Impressed tortoise (''Manouria impressa'')]]'''''Manouria''''' Gray 1854:133<ref name=gray34>{{cite journal|author=Gray, John Edward|title= Characters of several new species of freshwater tortoises (Emys) from India and China|journal= Proceedings of the Zoological Society of London|year=1834|volume= 2|pages=53–54}}</ref> ** ''Manouria emys'', Asian giant tortoise, brown tortoise (mountain tortoise) ** ''Manouria impressa'', impressed tortoise * {{Extinct}} '''''Megalochelys''''' Falconer, H. and Cautley, P.T. 1837.<ref name=fal&caut37>{{cite journal|author=Falconer, H. |author2=Cautley, P.T. |year=1837|title= On additional fossil species of the order Quadrumana from the Siwalik Hills|journal= Journal of the Asiatic Society of Bengal |volume=6|pages=354–360}}</ref> ** {{Extinct}} ''Megalochelys atlas'', Atlas tortoise, Extinct – Pliocene to Pleistocene ** {{Extinct}} ''Megalochelys cautleyi'', Cautley's giant tortoise * [[File:Western 'Namaqualand' tent tortoise.jpg|thumb|Tent tortoise (''Psammobates tentorius'')]][[File:Titanochelon perpiniana full.jpg|thumb|†''Titanochelon perpiniana'' ]]'''''Psammobates''''' Fitzinger 1835:113<ref name=fits1835 /> ** ''Psammobates geometricus'', geometric tortoise ** ''Psammobates oculifer'', serrated tent tortoise, Kalahari tent tortoise ** ''Psammobates tentorius'', African tent tortoise * '''''Pyxis''''' Bell 1827:395<ref name=bell1827>{{Cite journal| doi = 10.1111/j.1095-8339.1826.tb00122.x| url = http://www.iucn-tftsg.org/wp-content/uploads/file/Articles/Bell_1827.pdf| title = XVI. On two new Genera of Land Tortoises| journal = Transactions of the Linnean Society of London| volume = 15| issue = 2| pages = 392–401| year = 1827| last1 = Bell| first1 = T.| access-date = 2015-08-28| archive-url = https://web.archive.org/web/20180905175555/http://www.iucn-tftsg.org/wp-content/uploads/file/Articles/Bell_1827.pdf| archive-date = 2018-09-05}}</ref> ** ''Pyxis arachnoides'', (Madagascan) spider tortoise ** ''Pyxis planicauda'', flat-backed spider tortoise, (Madagascan) flat-tailed tortoise, flat-tailed spider tortoise * '''''Stigmochelys''''' Gray, 1873 ** ''Stigmochelys pardalis'', leopard tortoise * {{Extinct}} '''''Stylemys''''' ** ''Stylemys botti'' ** ''Stylemys calaverensis'' ** ''Stylemys canetotiana'' ** ''Stylemys capax'' ** ''Stylemys conspecta'' ** ''Stylemys copei'' ** ''Stylemys emiliae'' ** ''Stylemys frizaciana'' ** ''Stylemys karakolensis'' ** ''Stylemys nebrascensis'' (syn. ''S. amphithorax'') ** ''Stylemys neglectus'' ** ''Stylemys oregonensis'' ** ''Stylemys pygmea'' ** ''Stylemys uintensis'' ** ''Stylemys undabuna'' *{{Extinct}} '''''Titanochelon''''' ** {{extinct}}''Titanochelon gymnesica'' (Bate, 1914) Balearic Islands, Pliocene ** {{extinct}} ''Titanochelon bolivari'' (Hernandez-Pacheco, 1917) (type) Iberian Peninsula, Miocene ** {{extinct}} ''Titanochelon bacharidisi'' (Vlachos et al., 2014) Greece, Bulgaria, Late Miocene ** {{extinct}} ''Titanochelon perpiniana'' (Deperet 1885) France, Pliocene ** {{extinct}}''Titanochelon schafferi'' (Szalai, 1931) Samos, Greece, Miocene ** {{extinct}}''Titanochelon vitodurana'' (Biedermann 1862) Switzerland, Early Miocene ** {{extinct}}''Titanochelon kayadibiensis'' Karl, Staesche & Safi, 2021, Anatolia, Miocene ** {{extinct}}''Titanochelon eurysternum'' (Gervais, 1848–1852) France, Miocene[[File:Tunisian tortoise (Testudo graeca nabeulensis) male Cap Bon.jpg|thumb|Greek tortoise (''Testudo graeca'')]] ** {{extinct}}''Titanochelon ginsburgi'' (de Broin, 1977 ) France, Miocene ** {{extinct}}''Titanochelon leberonensis'' (Depéret, 1890) France, Miocene * '''''Testudo''''' ** ''Testudo graeca'', Greek tortoise, spur-thighed tortoise, Moorish tortoise ** ''Testudo hermanni'', Hermann's tortoise ** ''Testudo horsfieldii'', Russian tortoise ** ''Testudo kleinmanni'', Egyptian tortoise, including Negev tortoise ** ''Testudo marginata'', marginated tortoise ** {{extinct}}''Testudo sahakyanae'' (Vlachos & Vasilyan, 2026 ) Armenia, Pliocene<ref>{{Cite journal|last1=Vlachos |first1=E. |last2=Vasilyan |first2=D. |title=A new species of a bony spur-thighed tortoise from the Early Pliocene (MN15) of Jradzor site, Armenia, and the origin of the ''Testudo graeca'' (Testudines, Testudinidae) complex |year=2026 |journal=Swiss Journal of Palaeontology |volume=145 |pages=553–578 |doi=10.3897/sjp.145.182210 |doi-access=free }}</ref>

==Phylogeny== A molecular phylogeny of tortoises, following Le et al. (2006: 525):<ref name=Le>{{cite journal |url=https://www.researchgate.net/publication/7103480 |vauthors=Le M, Raxworthy CJ, McCord WP, Mertz L |title=A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes |journal=Molecular Phylogenetics and Evolution |volume=40 |issue=2 |pages=517–31 |date=August 2006 |pmid=16678445 |doi=10.1016/j.ympev.2006.03.003}}</ref>

{{clade |label1=Testudinidae |1={{clade |1={{clade |1=''Ergilemys'' |2=''Manouria'' |3=''Gopherus'' }}

|2={{clade |1={{clade |1=''Indotestudo'' |2=''Testudo'' |3=''Malacochersus'' }}

|2={{clade |1={{clade |1=''Centrochelys sulcata'' |label2=''Geochelone'' |2={{clade |1=''Geochelone platynota'' |2=''Geochelone elegans'' }} }}

|2={{clade |1={{clade |1={{clade |1=''Chersina'' |2=''Homopus'' }} |2={{clade |1=''Stigmochelys'' |2=''Psammobates'' }} }} |2={{clade |1=''Aldabrachelys'' |2=''Pyxis'' |3=''Astrochelys radiata'' |4=''Astrochelys yniphora'' }} }}

|3={{clade |1=''Kinixys'' |2=''Chelonoidis'' }} }} }} }} }}

A separate phylogeny via mtDNA analysis was found by Kehlmaier et al. (2021):<ref>{{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|page=3224|doi=10.1038/s41598-021-82299-w|pmid=33564028|pmc=7873039|bibcode=2021NatSR..11.3224K|issn=2045-2322|doi-access=free}}</ref> {{clade |label1=Testudinidae |1={{clade |1={{clade |1=''Manouria'' |2={{clade |1=''Gopherus'' |2={{clade |1={{clade |1=''Testudo'' |2={{clade |1=''Indotestudo'' |2={{clade |1=''Agrionemys'' |2=''Malacochersus'' }} }} }} |3={{clade |1={{extinct}}''Cylindraspis'' |2={{clade |1={{clade |1={{clade |1={{clade |1={{clade |1={{clade |1=''Chersina'' |2=''Chersobius'' }} |2=''Homopus'' }} |2={{clade |1=''Psammobates'' |2=''Stigmochelys'' }} }} |2={{clade |1={{clade |1=''Aldabrachelys'' |2={{clade |1=''Pyxis'' |2=''Astrochelys'' }} }} |2={{clade |1=''Kinixys'' |2={{clade |1={{clade |1=''Centrochelys'' |2=''Geochelone'' }} |2=''Chelonoidis'' }} }} }} }} }} }} }} }} }} }} }} }} [[File:Awatoceanofmilk01.JPG|upright|thumb|Bas-relief from Angkor Wat, Cambodia, shows Samudra manthan-Vishnu in the centre, his turtle Avatar Kurma below, asuras and devas to left and right]][[File:Tile al-Qazwini Louvre MAO1194.jpg|thumb|Tile with two rabbits, two snakes, and a tortoise, illustration for Zakariya al-Qazwini's book ''ʿAjāʾib al-makhlūqāt'', Iran, 19th century.]] In 2023 Kehlmaier again recovered a very similar phylogeny to the 2021 one, which further reaffirmed the evolutionary distinctiveness of the extinct ''Cylindraspis'', but swapped the position of ''Gopherus'' and ''Manouria'', making ''Gopherus'' the most basal genus.<ref>{{Cite journal |last1=Kehlmaier |first1=Christian |last2=Graciá |first2=Eva |last3=Ali |first3=Jason R. |last4=Campbell |first4=Patrick D. |last5=Chapman |first5=Sandra D. |last6=Deepak |first6=V. |last7=Ihlow |first7=Flora |last8=Jalil |first8=Nour-Eddine |last9=Pierre-Huyet |first9=Laure |last10=Samonds |first10=Karen E. |last11=Vences |first11=Miguel |last12=Fritz |first12=Uwe |date=2023-01-13 |title=Ancient DNA elucidates the lost world of western Indian Ocean giant tortoises and reveals a new extinct species from Madagascar |journal=Science Advances |language=en |volume=9 |issue=2 |article-number=eabq2574 |doi=10.1126/sciadv.abq2574 |pmid=36630487 |pmc=9833658 |bibcode=2023SciA....9.2574K |issn=2375-2548}}</ref>

==In human culture== {{Further information|Cultural depictions of turtles}} {{See also|World Turtle}}In Hinduism, Kurma ({{langx|sa|कुर्म}}) was the second Avatar of Vishnu. Like the Matsya Avatara, Kurma also belongs to the Satya Yuga. Vishnu took the form of a half-man, half-tortoise, the lower half being a tortoise. He is normally shown as having four arms. He sat on the bottom of the ocean after the Great Flood. A mountain was placed on his back by the other gods so they could churn the sea and find the ancient treasures of the Vedic peoples.{{Citation needed|date=November 2024}}

In Judaism, tortoises are seen as unclean animals.<ref>[https://www.stepbible.org/?q=version=ESV|reference=Lev.11&options=NVHUG Leviticus 11]</ref> Early Christians also viewed tortoises as unclean.<ref>{{Cite journal|last=Thomas|first=Richard|title=TORTOISES AND THE EXOTIC ANIMAL TRADE IN BRITAIN FROM MEDIEVAL TO 'MODERN'|url=http://www.britishcheloniagroup.org.uk/sites/default/files/u8/v8n1thomas_wm.pdf|journal=Testudo|volume=8|via=British Chelonia Group site}}</ref>

In Chinese and Japanese folk religion, tortoises are considered oracular animals. Tortoise shells were used by ancient Chinese as oracle bones to make predictions.<ref>{{Cite book |last=Raven |first=James |title=The Oxford Illustrated History of the Book |publisher=Oxford University Press |year=2020 |isbn=978-0-19-870298-6 |edition=Illustrated}}</ref>

In Ancient Greek mythology, Hermes crafts the first lyre from a tortoise.<ref>{{cite journal |last1=Shelmerdine |first1=Susan C. |title=Hermes and the Tortoise: A Prelude to Cult |journal=Greek, Roman, and Byzantine Studies |date=1984-09-11 |volume=25 |issue=3 |pages=201–208 |url=https://grbs.library.duke.edu/index.php/grbs/article/view/5481/5295 |access-date=11 November 2024}}</ref>

In September, 1968, two Russian tortoises became the first animals to fly to and circle the Moon. Their Zond 5 mission brought them back to Earth safely.

==Gallery== <gallery perrow="5"> File:Tortoise-Hatchling.jpg|Baby ''Testudo marginata'' emerges from its egg File:Baby tortoise.jpg|Baby tortoise, less than a day old File:Tortoise closeup.jpg|Young, 20-year-old Tanzanian leopard tortoise feeding on grass File:Aldabra.giant.tortoise.arp.jpg|Aldabra giant tortoise, ''Geochelone gigantea'' File:Leopards tortoise.jpg|22-year-old leopard tortoise File:Geochelone sulcata -Oakland Zoo -feeding-8a.jpg|African spurred tortoise from the Oakland Zoo File:Tortoise mating.jpg|Pair of African spurred tortoises mate in a zoo File:TurtleRideIfrog.jpg|Boy rides a tortoise at a zoo File:Small tortoises mating.webm|Video of tortoises mating File:Young Hermann's Tortoise.jpg|Young ''Testudo hermanni'' </gallery>

==References== {{Reflist|30em}}

==Further reading== * {{cite book|title=A Sheltered Life: The Unexpected History of the Giant Tortoise|last=Chambers|first=Paul|year=2004|publisher=John Murray|location=London|isbn=978-0-7195-6528-1}} * {{cite book|title=Turtles of the World|url=https://archive.org/details/turtlesofworld00erns|url-access=registration|last1=Ernst|first1=C. H.|last2=Barbour|first2=R. W.|year=1989|publisher=Smithsonian Institution Press|location=Washington, DC|isbn=978-0-87474-414-9}} * {{cite book|title=Giant Tortoises of the Indian Ocean|last=Gerlach|first=Justin|year=2004|publisher=Chimiara|location=Frankfurt}} * {{cite journal|date=February 2002|title=Phylogenetic Relationships among the Species of the Genus Testudo (Testudines: Testudinidae) Inferred from Mitochondrial 12S rRNA Gene Sequences|journal=Molecular Phylogenetics and Evolution|volume=22|issue=2|pages=174–183|doi=10.1006/mpev.2001.1052|pmid=11820839|issn=1055-7903|author1=Antoinette C. van der Kuyl|author2=Donato L. Ph. Ballasina|author3=John T. Dekker|author4=Jolanda Maas|author5=Ronald E. Willemsen|author6=Jaap Goudsmit}}

==External links== {{Commons|Testudinidae}} {{Wikispecies|Testudinidae}} * [http://reptile-database.reptarium.cz/advanced_search?taxon=Testudinidae&submit=Search Family Testudinidae (Tortoises)], The Reptiles Database * {{usurped|1=[https://web.archive.org/web/20020111174914/http://www.chelonia.org/ Chelonia]}}: Conservation and care of turtles. * [https://web.archive.org/web/20181104023757/http://tortoisestream.com/ Live Tortoise Stream ]: Live Tortoise Stream

Category:Tortoises