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For other uses, see Tortoise (disambiguation).
A. gigantea Aldabra Giant Tortoise.jpg
Aldabra giant tortoise
(Aldabrachelys gigantea)
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Cryptodira
Superfamily: Testudinoidea
Family: Testudinidae
Batsch, 1788
Type species
Testudo graeca
Linnaeus, 1758

Tortoises (/ˈtɔːr.təs.z/) are a family, Testudinidae, of land-dwelling reptiles in the order Testudines. Tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The carapace is fused to both the vertebrae and ribcage, and tortoises are unique among vertebrates in that the pectoral and pelvic girdles are inside the ribcage rather than outside. Tortoises can vary in size from a few centimeters to two meters. They are usually diurnal animals with tendencies to be crepuscular depending on the ambient temperatures. They are generally reclusive animals.

Use of the terms turtle, tortoise, and terrapin

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.[1] These terms are common names and do not reflect precise biological or taxonomic distinctions.[2]

Tile with two rabbits, two snakes, and a tortoise, illustration for Zakariya al-Qazwini's book, Iran, 19th century

The 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.[1] General American usage agrees; turtle is often a general term (although some restrict it to aquatic turtles); 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).[3][4][5][6] In America, for example, the members of the genus Terrapene dwell on land, yet are referred to as box turtles rather than tortoises.[2]

British 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.[6] In Britain, terrapin is used to refer to a larger group of semiaquatic turtles than the restricted meaning in America.[4][7]

Australian usage is different from both American and British usage.[6] Land tortoises are not native to Australia, yet traditionally freshwater turtles have been called "tortoises" in Australia.[8] 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.[8]



Adult male leopard tortoise, South Africa
Young African sulcata tortoise

Female tortoises dig nesting burrows in which they lay from one to 30 eggs.[9] 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.[10] 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 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.

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.[11]

Male tortoise
Adult male tortoise

Tortoises generally have one of the longest lifespans of any animal, and some individuals are known to have lived longer than 150 years.[12] Because of this, they symbolize longevity in some cultures, such as China. 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 Captain 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.[13] The record for the longest-lived vertebrate is exceeded only by one other, a koi named Hanako whose death on July 17, 1977, ended a 226-year lifespan.[14]

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 (sometimes spelled with two ds) 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.[15]

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.[16] Harriet died on June 23, 2006, just shy of her 176th birthday.

Timothy, a 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.

Jonathan, a Seychelles giant tortoise living on the island of St Helena, may be as old as 182[17][18] or 178 years.[19]

Sexual dimorphism

Many species of tortoises are sexually dimorphic, though the differences between males and females vary from species to species. 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.

In most tortoise species, the female tends to be larger than the male. 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.


The brain of a tortoise is extremely small. The tortoises, from Central and South America, do not have an area in the brain called hippo-campus, 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, an area that human's use for actions such as decision making.[20] In the 17th century, Francesco Redi performed an experiment involving removing the brain of a land tortoise, which then proceeded to live six months.[citation needed] Freshwater tortoises, when subject to the same experiment, continued similarly, but did not live so long. Redi also cut the head off a tortoise entirely, and it lived for 23 days.[21][22][23]

General information

Giant tortoises move very slowly on dry land, at only 0.17 mph (0.27 km/h).[24] The fastest recorded tortoise speed is 5 mph (8.0 km/h).[25]

The tortoise starts digging the ground to form its hybernaculum at the first signs of autumn. It digs with its fore-feet in a very slow motion, and prefers swampy grounds where it could bury itself in mud. It starts losing its appetite for food as the temperature drops until it stops eating. During hibernation it stops breathing as well. When the weather warms up suddenly it stops its digging, and starts it again as soon as the temperature drops. It wakes up from hibernation in the spring, but it does not start eating immediately. Gradually it gains its appetite and energy as the temperature warms up. During hot summer days, tortoises eat voraciously, and spend many hours sleeping. They start sleeping in late afternoon until late the next morning. Although tortoises love warm weather, they avoid hot sun, hiding under green leaves or between vegetation.[26]


Baby tortoise feeding on lettuce
Tortoise feeding on a cactus
Tortoise feeding on a cactus

Most land-based tortoises are herbivores, feeding on grasses, weeds, leafy greens, flowers, and some fruits, although some omnivorous species are in this family. 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. As different tortoise species vary greatly in their nutritional requirements, it is essential to thoroughly research the dietary needs of an individual turtles.


This species list largely follows van Dijk et al. (2014)[27] and Rhodin et al. (2015).[28]

Skeleton of a tortoise
Fossil of the extinct Ergilemys insolitus
Achilemys cassouleti, the most primitive testudine

Family Testudinidae Batsch 1788[29]

In religion

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

In Hinduism, Kurma (Sanskrit: 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.



Cultural depictions


See also


  1. ^ a b Simoons, Frederick J. (1991). Food in China: A Cultural and Historical Inquiry. CRC Press. ISBN 084938804X. p. 358.
  2. ^ a b Burton, Maurice and Burton, Robert (2002). International Wildlife Encyclopedia. Marshall Cavendish. ISBN 0761472665. p. 2796 .
  3. ^ Orenstein, Ronald Isaac (2001). Turtles, Tortoises and Terrapins: Survivors in Armor. Firefly Books. ISBN 1770851194
  4. ^ a b "Turtle". Retrieved 2012-09-16. 
  5. ^ What is the difference between turtles, terrapins, and tortoises?, North Carolina Aquariums (July 1997).
  6. ^ a b c Dawkins, Richard (2009). The Greatest Show on Earth: The Evidence for Evolution. Free Press. ISBN 1416594795. p. 174.
  7. ^ Endangered Wildlife and Plants of the World, Vol. 1. Marshall Cavenish. (2001). ISBN 0761471952. p. 1476.
  8. ^ a b Romanowski, Nick (2010). Wetland Habitats: A Practical Guide to Restoration and Management. CSIRO Publishing. ISBN 9780643096462. p. 134.
  9. ^ Highfield, Andy. "Tortoise Trust Egg F.A.Q". Retrieved 2009-04-07. 
  10. ^ Highfield, Andy. "Tortoise egg incubation". Retrieved 2009-04-07. 
  11. ^ Veterinary Services Department, Drs. Foster & Smith, Inc. "Shells: Anatomy and Diseases of Turtle and Tortoise Shells". Retrieved 2013-10-22. 
  12. ^ Moon, J. C.; McCoy, E. D.; Mushinsky, H. R.; Karl, S. A. (2006). "Multiple Paternity and Breeding System in the Gopher Tortoise, Gopherus polyphemus". Journal of Heredity. 97 (2): 150–157. doi:10.1093/jhered/esj017. PMID 16489146. 
  13. ^ "Tortoise Believed to Have Been Owned by Darwin Dies at 176". Associated Press via FOXNews. 2006-06-26. Archived from the original on July 1, 2006. 
  14. ^ Barton, Laura (2007-04-12). "Will You Still Feed Me...". London: The Guardian. Retrieved 2013-01-08. 
  15. ^ "World | South Asia | 'Clive of India's' tortoise dies". BBC News. 2006-03-23. Retrieved 2009-04-07. 
  16. ^ Thomson, S., Irwin, S. and Irwin, T. (1995). "Harriet, the Galapagos tortoise: disclosing one and a half centuries of history". Intermontanus. 4 (5): 33–35. 
  17. ^ Meet Jonathan, St Helena's 182-year-old giant tortoise BBC, March 13, 2014
  18. ^ Jonathan the 176-year-old tortoise revealed as world's oldest animal in Boer War photo Daily Mail, December 5, 2008
  19. ^ Boer War memento puts years on Jonathan the tortoise. The Times, December 4, 2008
  20. ^ "Tortoises Show Off Smarts by Mastering Touch-Screen Tech". Live Science. Retrieved 2016-12-01. 
  21. ^ Cuvier, Georges; Smith, Charles Hamilton; Pidgeon, Edward (1831). The animal kingdom arranged in conformity with its organization. 9. Printed for G. B. Whittaker. pp. 54–. 
  22. ^ The London Literary Gazette and Journal of Belles Lettres, Arts, Sciences, Etc. H. Colburn. 1831. pp. 277–. 
  23. ^ Broderip, William John (1852). Leaves from the Note Book of a Naturalist. E. Littell & Company. p. 83 (left column). In the beginning of November he opened the skull of a land-tortoise, removed every particle of brain, and cleaned the cavity out… instead of dying or remaining motionless, it groped its way about…. 
  24. ^ Grolier Encyclopedia (2003). The Great Book of Knowledge, The Speed of Animals, pp. 278.
  25. ^ Grolier Encyclopedia (2007). The Great Book of Knowledge, The Speed of Animals, p. 297.
  26. ^ The Natural History of Selborne by Gilbert White, pages on tortoise.
  27. ^ Turtle Taxonomy Working Group: van Dijk, P.P., Iverson, J.B., Rhodin, A.G.J., Shaffer, H.B., and Bour, R. (2014). Turtles of the World, 7th edition: annotated checklist of taxonomy, synonymy, distribution with maps, and conservation status. In: Rhodin, A.G.J., Pritchard, P.C.H., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Iverson, J.B., and 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 5(7):000.329–479 doi:10.3854/crm.5.000.checklist.v7.2014.
  28. ^ 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". Chelonian Research Monographs. 5(8):000e.1–66.
  29. ^ 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.
  30. ^ Loveridge, Arthur; Williams, Ernest E. (1957). "Revision of the African tortoises and turtles of the suborder Cryptodira". Bulletin of the Museum of Comparative Zoology. 115 (6): 163–557. 
  31. ^ Gray, John Edward (1873). "Notes on the genera of turtles (Oiacopodes), and especially on their skeletons and skulls". Proceedings of the Zoological Society of London: 395–411. 
  32. ^ 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.
  33. ^ a b c d Fitzinger, Leopold J. (1835). "Entwurf einer systematischen Anordnung der Schildkröten nach den Grundsätzen der natürlichen Methode". Annalen des Wiener Museums der Naturgeschichte. 1: 105–128. 
  34. ^ Testudinidae, The Reptile Database
  35. ^ Austin, J. J.; Nicholas Arnold, E. (2001). "Ancient mitochondrial DNA and morphology elucidate an extinct island radiation of Indian Ocean giant tortoises (Cylindraspis)". Proceedings of the Royal Society B: Biological Sciences. 268 (1485): 2515. doi:10.1098/rspb.2001.1825. 
  36. ^ The Recently Extinct Plants and Animals Database Extinct Reptiles: Geochelone burchardi
  37. ^ The Recently Extinct Plants and Animals Database Extinct Reptiles: Geochelone vulcanica
  38. ^ Rafinesque, Constantine Samuel (1832). "Description of two new genera of soft shell turtles of North America". Atlantic Journal and Friend of Knowledge. 1: 64–65. 
  39. ^ Duméril, André Marie Constant and Bibron, Gab riel. 1834. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Premier. Paris: Roret, 439 pp.
  40. ^ Lindholm, Wassili A. (1929). "Revidiertes Verzeichnis der Gattungen der rezenten Schildkröten nebst Notizen zur Nomenklatur einiger Arten". Zoologischer Anzeiger. 81: 275–295. 
  41. ^ Gray, John Edward (1834). "Characters of several new species of freshwater tortoises (Emys) from India and China". Proceedings of the Zoological Society of London. 2: 53–54. 
  42. ^ Falconer, H.; Cautley, P.T. (1837). "On additional fossil species of the order Quadrumana from the Siwalik Hills". Journal of the Asiatic Society of Bengal. 6: 354–360. 
  43. ^ Bell, T. (1827). "XVI. On two new Genera of Land Tortoises" (PDF). Transactions of the Linnean Society of London. 15 (2): 392. doi:10.1111/j.1095-8339.1826.tb00122.x. 

Further reading

  • Chambers, Paul (2004). A Sheltered Life: The Unexpected History of the Giant Tortoise. London: John Murray. ISBN 0-7195-6528-6. 
  • Ernst, C. H.; Barbour, R. W. (1989). Turtles of the World. Washington, DC: Smithsonian Institution Press. 
  • Gerlach, Justin (2004). Giant Tortoises of the Indian Ocean. Frankfurt: Chimiara. 
  • Antoinette C. van der Kuyl; Donato L. Ph. Ballasina; John T. Dekker; Jolanda Maas; Ronald E. Willemsen; Jaap Goudsmit (February 2002). "Phylogenetic Relationships among the Species of the Genus Testudo (Testudines: Testudinidae) Inferred from Mitochondrial 12S rRNA Gene Sequences". Molecular Phylogenetics and Evolution. 22 (2): 174–183. doi:10.1006/mpev.2001.1052. ISSN 1055-7903. PMID 11820839. 

External links