'''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 [[turtle]]s, tortoises have a [[turtle shell|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''' ({{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 [[turtle shell|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 tortoise|Galápagos giant tortoise]], growing to more than {{convert|1.2|m|ft}} in length, whereas others like the [[Chersobius signatus|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|pages=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 [[Diurnality|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 tortoise]]s 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.
Tortoises can vary in size with some species, such as the [[Galápagos tortoise|Galápagos giant tortoise]], growing to more than {{convert|1.2|m|ft}} in length, whereas others like the [[Chersobius signatus|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 [[Diurnality|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 tortoise]]s 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==
==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>[[File:SmithTestudoSkeleton.jpg|thumb|Skeleton of a tortoise|210x210px]]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.<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|url-status=dead|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 turtle]]s rather than tortoises.<ref name="WildlifeEncy" />
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>[[File:SmithTestudoSkeleton.jpg|thumb|Skeleton of a tortoise|210x210px]]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.<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 turtle]]s rather than tortoises.<ref name="WildlifeEncy" />
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.<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>
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" />
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" />
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 (eggs)|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|url-status=dead}}</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 [[Turtle shell#Plastron|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 [[herbivory|herbivorous]] species commonly will consume [[worm]]s or insect [[larva]]e 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>
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 (eggs)|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 [[Turtle shell#Plastron|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 [[herbivory|herbivorous]] species commonly will consume [[worm]]s or insect [[larva]]e 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]]|url-status=dead|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>
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 [[Tonga]]n 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>
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 [[Tonga]]n 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 Zoological Gardens|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=http://news.bbc.co.uk/1/hi/world/south_asia/4837988.stm|title='Clive of India's' tortoise dies|work=BBC News|date=2006-03-23|access-date=2009-04-07}}</ref>
The [[Alipore Zoological Gardens|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 (turtle)|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 [[HMS Beagle|''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.
[[Harriet (turtle)|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 [[HMS Beagle|''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 (tortoise)|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=http://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>
[[Timothy (tortoise)|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 (tortoise)|Jonathan]], a [[Dipsochelys hololissa|Seychelles giant tortoise]] living on the island of [[St Helena]], may be as old as {{Years ago|1832}} years.<ref>[https://www.bbc.com/news/magazine-26543021 Meet Jonathan, St Helena's 182-year-old giant tortoise] BBC, March 13, 2014</ref>
[[Jonathan (tortoise)|Jonathan]], a [[Dipsochelys hololissa|Seychelles giant tortoise]] living on the island of [[St Helena]], may be as old as {{Years ago|1832}} years.<ref>[https://www.bbc.com/news/magazine-26543021 Meet Jonathan, St Helena's 182-year-old giant tortoise] BBC, March 13, 2014</ref>
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===Brain===
===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>
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>
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.<ref>{{cite book|last1=Cuvier|first1=Georges|last2=Smith|first2=Charles Hamilton|last3=Pidgeon|first3=Edward|title=The animal kingdom arranged in conformity with its organization|url=https://books.google.com/books?id=_n1IAAAAMAAJ&pg=PA54|year=1831|publisher=Printed for G. B. Whittaker|pages=54–|volume=9}}</ref><ref>{{cite book|title=The London Literary Gazette and Journal of Belles Lettres, Arts, Sciences, Etc|url=https://books.google.com/books?id=HYdHAAAAYAAJ&pg=PA277|year=1831|publisher=H. Colburn|pages=277–}}</ref><ref>{{cite book|last=Broderip|first=William John|title=Leaves from the Note Book of a Naturalist|url=https://archive.org/details/leavesfromnotebo00brodrich|year=1852|publisher=E. Littell & Company|page=[https://archive.org/details/leavesfromnotebo00brodrich/page/83 83] (left column)|quote=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....}}</ref>
==Distribution==
==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.
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 tortoise]]s. 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=978-0226143972}}</ref>
Many large islands are or were characterized by species of [[giant tortoise]]s. 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 tortoise]]s could be found on nearly every major island group, including the [[Bahamas]], the [[Greater Antilles]] (including [[Cuba]] and [[Hispaniola]]), the [[Lesser Antilles]], the [[Canary Island]]s, [[Malta]], the [[Seychelles]], the [[Mascarene Islands]] (including [[Mauritius]] and [[Réunion|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 [[Meiolaniidae|meiolaniid]] stem turtles are thought to have filled the same niche. [[Giant tortoise]]s 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]]
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 tortoise]]s could be found on nearly every major island group, including the [[Bahamas]], the [[Greater Antilles]] (including [[Cuba]] and [[Hispaniola]]), the [[Lesser Antilles]], the [[Canary Island]]s, [[Malta]], the [[Seychelles]], the [[Mascarene Islands]] (including [[Mauritius]] and [[Réunion|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 [[Meiolaniidae|meiolaniid]] stem turtles are thought to have filled the same niche. [[Giant tortoise]]s 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]]
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==Taxonomy==
==Taxonomy==
This species list largely follows [[Turtle Taxonomy Working Group]] (2021)<ref>{{Cite book |last=Rhodin |first=Anders G.J. |url=http://dx.doi.org/10.3854/crm.8.checklist.atlas.v9.2021 |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=978-0965354097 |doi-access=free }}</ref>
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>[[August Batsch|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>
[[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>[[August Batsch|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 }}</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 }}</ref>
* '''''[[Pyxis (turtle)|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| url-status = dead}}</ref>
* '''''[[Pyxis (turtle)|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>
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|pages=3224|doi=10.1038/s41598-021-82299-w|pmid=33564028|pmc=7873039|bibcode=2021NatSR..11.3224K|issn=2045-2322|doi-access=free}}</ref>
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
{{clade
|label1=Testudinidae
|label1=Testudinidae
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[[File:Awatoceanofmilk01.JPG|upright|thumb|Bas-relief from [[Angkor Wat]], Cambodia, shows [[Samudra manthan]]-Vishnu in the centre, his turtle [[Avatar]] [[Kurma]] below, [[asura]]s and [[deva (Hinduism)|devas]] to left and right]][[File:Tile al-Qazwini Louvre MAO1194.jpg|thumb|Tile with two [[rabbits]], two [[snake]]s, and a tortoise, illustration for [[Zakariya al-Qazwini]]'s book [[Aja'ib al-Makhluqat|''ʿAjāʾib al-makhlūqāt'']], [[Iran]], 19th century.]]
[[File:Awatoceanofmilk01.JPG|upright|thumb|Bas-relief from [[Angkor Wat]], Cambodia, shows [[Samudra manthan]]-Vishnu in the centre, his turtle [[Avatar]] [[Kurma]] below, [[asura]]s and [[deva (Hinduism)|devas]] to left and right]][[File:Tile al-Qazwini Louvre MAO1194.jpg|thumb|Tile with two [[rabbits]], two [[snake]]s, and a tortoise, illustration for [[Zakariya al-Qazwini]]'s book [[Aja'ib al-Makhluqat|''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 |pages=eabq2574 |doi=10.1126/sciadv.abq2574 |pmid=36630487 |pmc=9833658 |bibcode=2023SciA....9.2574K |issn=2375-2548}}</ref>
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==
==In human culture==
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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 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 space===
===In space: the 1968 trip to the Moon===
In September, 1968, two [[Russian tortoise]]s became the first animals to fly to and circle the Moon. Their [[Zond 5]] mission brought them back to Earth safely.
In September, 1968, two [[Russian tortoise]]s became the first animals to fly to and circle the Moon. Their [[Zond 5]] mission brought them back to Earth safely.
===In the United Kingdom===
In 2015, the University of Lincoln held a ribbon-cutting ceremony to open the new Joseph Banks Laboratories. Instead of using scissors, the honor went to Charles Darwin, a tortoise from the School of Life Sciences.
The "ribbon" was made from dandelion and rocket leaves, which the tortoise chewed through to mark the opening.
[[File:Charles Darwin Ribbon-cutting Ceremony.jpg|thumb|Featured photo of Charles Darwin's ribbon-cutting ceremony, 2015.]]
===As pets===
===As pets===
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==Further reading==
==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=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=9780874744149}}
* {{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 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}}
* {{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}}
Tortoises (Template:IPAc-enTemplate:Respell) 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.[1]
Tortoises can vary in size with some species, such as the Galápagos giant tortoise, growing to more than Template:Convert in length, whereas others like the speckled Cape tortoise have shells that measure only Template:Convert long.[2] Several lineages of tortoises have independently evolved very large body sizes in excess of Template:Convert, 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.
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.[3] These terms are common names and do not reflect precise biological or taxonomic distinctions.[4]
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.[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 the diamondback terrapin (Malaclemys terrapin).[5][6][7][8] In America, for example, the members of the genus Terrapene dwell on land, yet are referred to as box turtles rather than tortoises.[4]
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.[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 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.[11] 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.[12]
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.[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 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.[15]
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.[16]
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.[17] 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.[18]
Many species of tortoises are sexually 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.
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.[22]
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.[23] 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.[24]
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.[26] 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.[27]
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.[48]
Script error: No such module "Labelled list hatnote".In Hinduism, Kurma (Template:Langx) was the second Avatar of Vishnu. Like the MatsyaAvatara, 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.Script error: No such module "Unsubst".
In Ancient Greek mythology, Hermes crafts the first lyre from a tortoise.[52]
In space: the 1968 trip to the Moon
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.
In the United Kingdom
In 2015, the University of Lincoln held a ribbon-cutting ceremony to open the new Joseph Banks Laboratories. Instead of using scissors, the honor went to Charles Darwin, a tortoise from the School of Life Sciences.
The "ribbon" was made from dandelion and rocket leaves, which the tortoise chewed through to mark the opening.
↑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.
↑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.
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↑Duméril, André Marie Constant and Bibron, Gab riel. 1834. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Premier. Paris: Roret, 439 pp.
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