Tortoises (/ˈtɔːrtəs.ɪz/TOR-təs-iz) arereptiles of the familyTestudinidae of the orderTestudines (Latin for "tortoise"). Like otherturtles, tortoises have ashell to protect frompredation and other threats. The shell in tortoises is generally hard, and like other members of the suborderCryptodira, they retract their necks and heads directly backward into the shell to protect them.[1]
Tortoises can vary in size with some species, such as theGalápagos giant tortoise, growing to more than 1.2 metres (3.9 ft) in length, whereas others like theSpeckled cape tortoise have shells that measure only 6.8 centimetres (2.7 in) long.[2] Several lineages of tortoises have independently evolved very large body sizes in excess of 100 kilograms (220 lb), including the Galapagos giant tortoise and theAldabra giant tortoise. They are usuallydiurnal animals with tendencies to becrepuscular depending on the ambient temperatures. They are generallyreclusive 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 namedAdwaita may have lived an estimated 255 years. In general, most tortoise species can live 80–150 years.
Tortoises are placid and slow-moving, with an average walking speed of 0.2–0.5 km/h.[citation needed]
Differences exist in usage of the common termsturtle, tortoise, andterrapin, depending on the variety of English being used; usage is inconsistent and contradictory.[3] These terms are common names and do not reflect precise biological or taxonomic distinctions.[4]
Skeleton of a tortoise
TheAmerican 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.[3] 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 thediamondback terrapin (Malaclemys terrapin).[5][6][7][8] In America, for example, the members of the genusTerrapene dwell on land, yet are referred to asbox turtles rather than tortoises.[4]
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.[8] In Britain, terrapin is used to refer to a larger group of semiaquatic turtles than the restricted meaning in America.[6][9]
Australian usage is different from both American and British usage.[8] Land tortoises are not native to Australia, and traditionally freshwater turtles have been called "tortoises" in Australia.[10] 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.[10]
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 herclutch with sand, soil, and organic material. The eggs are left unattended, and depending on the species, take from 60 to 120 days to incubate.[11] The size of the egg depends on the size of the mother and can be estimated by examining the width of thecloacal opening between thecarapace andplastron. 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 formedhatchling uses anegg 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 strictlyherbivorous species commonly will consumeworms or insectlarvae for additional protein.[12]
The number of concentric rings on the carapace, much like the cross-section of atree, 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 offorage (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.[13]
Tortoises generally have one of the longest lifespans of any animal, and some individuals are known to have lived longer than 150 years.[14] Because of this, they symbolize longevity in some cultures, such asChinese culture. The oldest tortoise ever recorded, and one of the oldest individual animals ever recorded, wasTu'i Malila, which was presented to theTongan royal family by the British explorerJames 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.[15]
TheAlipore Zoo in India was the home toAdwaita, which zoo officials claimed was the oldest living animal until its death on March 23, 2006. Adwaita (also spelled Addwaita) was anAldabra 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 ofRobert Clive.[16]
Harriet was a resident at theAustralia Zoo in Queensland from 1987 to her death in 2006; she was believed to have been brought to England byCharles Darwin aboard theBeagle and then on to Australia byJohn Clements Wickham.[17] Harriet died on June 23, 2006, just shy of her 176th birthday.
Timothy, a femalespur-thighed tortoise, lived to be about 165 years old. For 38 years, she was carried as amascot aboard various ships in Britain'sRoyal Navy. Then in 1892, at age 53, she retired to the grounds ofPowderham Castle inDevon. Up to the time of her death in 2004, she was believed to be the United Kingdom's oldest resident.[18]
Many species of tortoises aresexually dimorphic, though the differences between males and females vary from species to species.[21] 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.
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 thehippocampus, 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.[22]
In the 17th century,Francesco Redi performed an experiment that involved removing the brain of a land tortoise, which then proceeded to live six months. Freshwater tortoises, when subjected 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.[23][24][25]
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 fromAustralasia. 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 ofgiant tortoises. Part of the reason for this is that tortoises are good atoceanic 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.[26] Once on islands tortoises faced few predators or competitors and could grow to large sizes and become the dominant largeherbivores on many islands due to their low metabolic rate and reduced need for fresh water compared to mammals.[27]
Tortoises are generally considered to be strictherbivores, feeding on grasses, weeds, leafy greens, flowers, and some fruits. However, hunting and eating of birds has been observed on occasion.[29] Pet tortoises typically require diets based on wild grasses, weeds, leafy greens and certain flowers. Certain species consumeworms orinsects 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.
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 useolfactory 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.[30]
In 2023 Kehlmaier again recovered a very similar phylogeny to the 2021 one, which further reaffirmed the evolutionary distinctiveness of the extinctCylindraspis, but swapped the position ofGopherus andManouria, makingGopherus the most basal genus.[51]
InHinduism,Kurma (Sanskrit:कुर्म) was the secondAvatar ofVishnu. Like theMatsyaAvatara, Kurma also belongs to theSatya 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 theocean after theGreat Flood. Amountain was placed on his back by the othergods so they could churn the sea and find the ancient treasures of theVedic peoples.[citation needed]
In September, 1968, twoRussian tortoises became the first animals to fly to and circle the Moon. TheirZond 5 mission brought them back to Earth safely.
^Thomson, S.; Irwin, S.; Irwin, T. (1995). "Harriet, the Galapagos tortoise: disclosing one and a half centuries of history".Intermontanus.4 (5):33–35.
^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....
^Goodman, Steven M.; Jungers, William L. (2014).Extinct Madagascar : picturing the island's past. Chicago.ISBN978-0226143972.{{cite book}}: CS1 maint: location missing publisher (link)
^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.; van Dijk, P.P.; et al. (Turtle Extinctions Working Group) (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. Vol. 5. pp. 000e.1–66.doi:10.3854/crm.5.000e.fossil.checklist.v1.2015.ISBN978-0965354097.
^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.
^Adán Pérez-García; Evangelos Vlachos; Xabier Murelaga (2020). "A large testudinid with African affinities in the post-Messinian (lower Pliocene) record of south-eastern Spain".Palaeontology.63 (3):497–512.doi:10.1111/pala.12468.S2CID214232312.
^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.
^abcdFitzinger, Leopold J. (1835). "Entwurf einer systematischen Anordnung der Schildkröten nach den Grundsätzen der natürlichen Methode".Annalen des Wiener Museums der Naturgeschichte.1:105–128.
^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.
^Duméril, André Marie Constant and Bibron, Gab riel. 1834. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Premier. Paris: Roret, 439 pp.
^Lindholm, Wassili A. (1929). "Revidiertes Verzeichnis der Gattungen der rezenten Schildkröten nebst Notizen zur Nomenklatur einiger Arten".Zoologischer Anzeiger.81:275–295.
^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.
^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.
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.ISSN1055-7903.PMID11820839.
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