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Tortoises () are
reptiles Reptiles, as most commonly defined, are the animals in the class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or o ...

reptiles
of the family Testudinidae of the order Testudines (Latin: tortoise). They are particularly distinguished from other
turtles Turtles are an Order (biology), order of reptiles known as Testudines, characterized by a turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira, side-necked turtles and Cryptodi ...

turtles
(which includes the order Chelonia) by being exclusively land-dwelling, while many (though not all) other turtle species are at least partly
aquatic Aquatic means relating to water Water (chemical formula H2O) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known li ...
. Like other turtles, tortoises have a
shell Shell may refer to: Architecture and design * Shell (structure)A shell is a type of structural element which is characterized by its geometry, being a three-dimensional solid whose thickness is very small when compared with other dimensions, and ...
to protect from
predation Predation is a biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms, including humans, and their physical env ...

predation
and other threats. The shell in tortoises is generally hard, and like other members of the suborder
Cryptodira The Cryptodira ('' el, hidden neck'') are a suborder of Testudines that includes most living tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the Latin name for tortoise). They are particula ...
, they retract their necks and heads directly backwards into the shell to protect them. Tortoises can vary in size with some species, such as the
Galápagos giant tortoise
Galápagos giant tortoise
, growing to more than in length, whereas others like the Speckled cape tortoise have shells that measure only long. Several lineages of tortoises have independently evolved very large body sizes in excess of 100 kg, including the
Galapagos giant tortoise
Galapagos giant tortoise
and the
Aldabra giant tortoise The Aldabra giant tortoise (''Aldabrachelys gigantea''), from the islands of the Aldabra Atoll in the Seychelles, is one of the largest tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the L ...
. They are usually
diurnal Diurnal ("daily Daily or The Daily may refer to: Journalism * Daily newspaper A newspaper is a Periodical literature, periodical publication containing written News, information about current events and is often typed in black ink with a ...
animals with tendencies to be
crepuscular A crepuscular animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respirat ...

crepuscular
depending on the ambient temperatures. They are generally
reclusive A recluse is a person who lives in voluntary seclusion from the public and society. The word is from the Latin ''recludere'', which means "shut up" or "sequester". Historically, the word referred to a hermit's total isolation from the world. E ...
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 The Galápagos tortoise complex or Galápagos giant tortoise complex is a species complex of 15 (13 Extant taxon, extant and 2 extinct) very large tortoise species in the genus ''Chelonoidis'' (which also contains three other species from mainla ...

Galápagos tortoise
s are noted to live over 150 years, but an
Aldabra giant tortoise The Aldabra giant tortoise (''Aldabrachelys gigantea''), from the islands of the Aldabra Atoll in the Seychelles, is one of the largest tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the L ...
named
Adwaita Adwaita (meaning "one and only" in Sanskrit Sanskrit (, attributively , ''saṃskṛta-'', nominalization, nominally , ''saṃskṛtam'') is a classical language of South Asia belonging to the Indo-Aryan languages, Indo-Aryan branch of the ...

Adwaita
may have lived an estimated 255 years. In general, most tortoise species can live 80–150 years. Tortoises are placid and very slow moving, with an average walking speed of 0.2–0.5 km/h.


Terminology

Differences exist in usage of the common terms
turtle Turtles are an Order (biology), order of reptiles known as Testudines, characterized by a turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira, side-necked turtles and Cryptodi ...

turtle
, tortoise, and
terrapin Terrapins are one of several small species of turtle Turtles are reptiles of the Order (biology), order Chelonia or Testudines . They are characterized by a special bone, bony or cartilage, cartilaginous animal shell, shell developed fro ...
, depending on the variety of English being used; usage is inconsistent and contradictory.Simoons, Frederick J. (1991). ''Food in China: A Cultural and Historical Inquiry''. CRC Press. . p. 358. These terms are common names and do not reflect precise biological or taxonomic distinctions.Burton, Maurice and Burton, Robert (2002). ''International Wildlife Encyclopedia''. Marshall Cavendish. . p. 2796. The
American Society of Ichthyologists and Herpetologists The American Society of Ichthyologists and Herpetologists (ASIH) is an international learned society A learned society (; also known as a learned academy, scholarly society, or academic association) is an organization that exists to promote an ...
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. 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 The diamondback terrapin or simply terrapin (''Malaclemys terrapin'') is a species of turtle native to the Brackish water, brackish coastal tidal marshes of the eastern and southern United States, and in Bermuda. It belongs to the monotypic genus ...
(''Malaclemys terrapin'').What is the difference between turtles, terrapins, and tortoises?
, North Carolina Aquariums (July 1997).
Dawkins, Richard (2009). '' The Greatest Show on Earth: The Evidence for Evolution''. Free Press. . p. 174. In America, for example, the members of the genus '' Terrapene'' dwell on land, yet are referred to as
box turtle Box turtles are North American turtle Turtles are reptiles of the Order (biology), order Chelonia or Testudines . They are characterized by a special bone, bony or cartilage, cartilaginous animal shell, shell developed from their ribs tha ...

box turtle
s rather than tortoises. 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. In Britain, terrapin is used to refer to a larger group of semiaquatic turtles than the restricted meaning in America.''Endangered Wildlife and Plants of the World'', Vol. 1. Marshall Cavenish. (2001). . p. 1476. Australian usage is different from both American and British usage. Land tortoises are not native to Australia, and traditionally freshwater turtles have been called "tortoises" in Australia.Romanowski, Nick (2010). ''Wetland Habitats: A Practical Guide to Restoration and Management''. CSIRO Publishing. . p. 134. 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.


Biology


Life cycle

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 Single dry-clutch friction disc. The splined hub is attached to the disc with springs to damp chatter. A clutch is a mechanical device that engages and disengages power transmission Transmission may refer to: Science and technology * Power ...
with sand, soil, and organic material. The eggs are left unattended, and depending on the species, take from 60 to 120 days to incubate. The size of the egg depends on the size of the mother and can be estimated by examining the width of the
cloacal In animal anatomy, a cloaca (plural cloacae or ) is the posterior Body orifice, orifice that serves as the only opening for the Gut (anatomy), digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, ...
opening between the
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
and
plastron The turtle shell is a shield for the ventral and dorsal parts of turtles (the Order (biology), order Testudines), completely enclosing all the vital organs of the turtle and in some cases even the head. It is constructed of modified bony elements s ...

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 In oviparous Oviparous animals are female animals that lay their egg Diagram of a chicken egg in its 9th day. Membranes: allantois, chorion, amnion, and vitellus/ yolk. An egg is the organic vessel containing the zygote in which an emb ...

hatchling
uses an
egg tooth An egg tooth is a temporary, sharp projection present on the bill Bill(s) may refer to: Common meanings * Banknote A banknote (often known as a bill (in the US and Canada), paper money, or simply a note) is a type of negotiable instrument ...
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 A herbivore is an animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular r ...
species commonly will consume
worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limb Limb can refer to: *Limb (anatomy), an appendage of a human or animal *Limb Music, a record label *Limb (album), an ...

worm
s or insect
larva A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are organisms that form the Animalia. With few exceptions, animals , , are , can , and grow from a hollow sphere of , the , during . Ove ...
e for additional protein. The number of concentric rings on the carapace, much like the cross-section of a
tree In botany, a tree is a perennial plant with an elongated Plant stem, stem, or trunk (botany), trunk, supporting branches and leaves in most species. In some usages, the definition of a tree may be narrower, including only wood plants with se ...

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 Forage is a plant material (mainly plant leaves and stems) eaten by grazing In agriculture Agriculture is the science, art and practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentism, se ...

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. Tortoises generally have one of the longest lifespans of any animal, and some individuals are known to have lived longer than 150 years. Because of this, they symbolize longevity in some cultures, such as
Chinese culture Chinese culture () is one of the world's oldest cultures, originating thousands of years ago. The culture prevails across a large geographical region in East Asia East Asia is the eastern region of Asia Asia () is Earth's large ...
. The oldest tortoise ever recorded, and one of the oldest individual animals ever recorded, was
Tu'i Malila Tu'i Malila (1777 – 16 May 1966) was a tortoise that Captain James Cook was traditionally said to have given to the royal family of Tonga Tonga (; Tongan language, Tongan: ), officially named the Kingdom of Tonga (Tongan language, Tongan ...
, which was presented to the
Tonga Tonga (, ), officially named the Kingdom of Tonga ( to, Puleʻanga Fakatuʻi ʻo Tonga), is a Polynesia Polynesia (, ; from grc, πολύς "many" and grc, νῆσος "island") ( to, Faka-Polinisia; mi, Porinihia; haw, Polenekia; ...

Tonga
n royal family by the British explorer
James Cook Captain Captain is a title for the commander of a military unit, the commander of a ship, aeroplane, spacecraft, or other vessel, or the commander of a port, fire department or police department, election precinct, etc. The captain is a milit ...

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. The record for the longest-lived vertebrate is exceeded only by one other, a
koi or more specifically , are colored varieties of the Amur carp (''Cyprinus rubrofuscus'') that are kept for decorative purposes in outdoor koi ponds or water gardens. Koi is an informal name for the colored variants of ''C. rubrofuscus'' kept ...

koi
named Hanako, whose death on July 17, 1977, ended a 226-year lifespan. The Alipore Zoo in
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...

India
was the home to
Adwaita Adwaita (meaning "one and only" in Sanskrit Sanskrit (, attributively , ''saṃskṛta-'', nominalization, nominally , ''saṃskṛtam'') is a classical language of South Asia belonging to the Indo-Aryan languages, Indo-Aryan branch of the ...

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 The Aldabra giant tortoise (''Aldabrachelys gigantea''), from the islands of the Aldabra Atoll in the Seychelles, is one of the largest tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the L ...
brought to
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...

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 Major-General Major general (abbreviated MG, maj. gen. and similar) is a military rank used in many countries. It is derived from the older rank of sergeant major general. The disappearance of the "sergeant" in the title explains the apparen ...
. Harriet was a resident at the
Australia Zoo Australia Zoo is a zoo located in the Australian state of Queensland on the Sunshine Coast, Queensland, Sunshine Coast near Beerwah, Queensland, Beerwah/Glass House Mountains National Park, Glass House Mountains. It is a member of the Zoo and Aq ...
in Queensland from 1987 to her death in 2006; she was believed to have been brought to England by
Charles Darwin Charles Robert Darwin (; ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist and biologist, best known for his contributions to the science of evolution. His proposition that all sp ...

Charles Darwin
aboard the ''Beagle'' and then on to Australia by
John Clements Wickham John Clements Wickham (21 November 17986 January 1864) was a Scotland, Scottish explorer, naval officer, magistrate and administrator. He was Executive officer, first officer on during its Second voyage of HMS Beagle, second survey mission, 1831& ...

John Clements Wickham
. Harriet died on June 23, 2006, just shy of her 176th birthday. Timothy, a female
spur-thighed tortoise The Greek tortoise (''Testudo graeca''), also known commonly as the spur-thighed tortoise, is a species of tortoise in the family In human society, family (from la, familia) is a group of people related either by consanguinity (by recog ...
, lived to be about 165 years old. For 38 years, she was carried as a
mascot A mascot is any human, animal, or object thought to bring luck Luck is the phenomenon and belief that defines the experience of notably positive, negative, or improbable events. The naturalistic interpretation is that positive and negative ...

mascot
aboard various ships in Britain's
Royal Navy The Royal Navy (RN) is the United Kingdom's naval warfare Naval warfare is combat Combat ( French for ''fight'') is a purposeful violent conflict meant to physically harm or kill the opposition. Combat may be armed (using weapon A ...
. Then in 1892, at age 53, she retired to the grounds of
Powderham Castle File:PowderhamCastle FromUnderGatehouse.jpg, 200px, Powderham Castle, west front, viewed from under the Victorian gatehouse. The leftmost tower dates from 1390–1450 as does the main high central block, which originally housed a full-height grea ...
in
Devon Devon (, archaically known as Devonshire) is a county A county is a geographical region of a country used for administrative or other purposesChambers Dictionary The ''Chambers Dictionary'' (''TCD'') was first published by William Ch ...

Devon
. Up to the time of her death in 2004, she was believed to be the United Kingdom's oldest resident. Jonathan, a living on the island of
St Helena Saint Helena () is a British possession in the South Atlantic Ocean. It is a remote volcanic tropical island west of the coast of south-western Africa, and east of Rio de Janeiro Rio de Janeiro (; ;), or simply Rio, is the List of larges ...
, may be as old as years or years.


Sexual dimorphism

Many species of tortoises are
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same species exhibit different characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most animals and some plants. Differences may in ...
, 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. 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 The hippocampus (via Latin from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its popul ...

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. In the 17th century,
Francesco Redi Francesco Redi (18 February 1626 – 1 March 1697) was an Italian Italian may refer to: * Anything of, from, or related to the country and nation of Italy ** Italians, an ethnic group or simply a citizen of the Italian Republic ** Italian langua ...

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.


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 Australasia is a region In geography Geography (from Ancient Greek, Greek: , ''geographia'', literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and phenomena of the Earth ...

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 Giant tortoises are any of several species of various large land tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the Latin name for tortoise). They are particularly distinguished from other ...
s. Part of the reason for this is that tortoises are very good at
oceanic dispersal Oceanic dispersal is a type of biological dispersal Biological dispersal refers to both the movement of individuals (animals, plants, fungi, bacteria, etc.) from their birth site to their breeding site ('natal dispersal'), as well as the movem ...
. 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. Once on islands tortoises faced few predators or competitors and could grow to very large sizes and become the dominant large
herbivores A herbivore is an animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All ...
on many islands due to their low metabolic rate and reduced need for fresh water compared to mammals. Today there are only two living species of giant tortoises, the
Aldabra giant tortoise The Aldabra giant tortoise (''Aldabrachelys gigantea''), from the islands of the Aldabra Atoll in the Seychelles, is one of the largest tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the L ...
on
Aldabra Atoll Aldabra is the world's second-largest coral atoll. It is situated in the Aldabra Group of islands in the Indian Ocean that are part of the Outer Islands (Seychelles), Outer Islands of the Seychelles, with a distance of 1,120 km (700 m ...
and the several species/subspecies of Galapagos giant tortoise found on the Galapagos Islands. However, until very recently
giant tortoise Giant tortoises are any of several species of various large land tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the Latin name for tortoise). They are particularly distinguished from other ...
s could be found on nearly every major island group, including the
Bahamas The Bahamas (), known officially as The Commonwealth of The Bahamas, is a sovereign country within the Lucayan Archipelago of the West Indies in the Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the a ...

Bahamas
, the
Greater Antilles The Greater Antilles ( es, Grandes Antillas or Antillas Mayores; french: Grandes Antilles ht, Gwo Zantiy jam, Grieta hAntiliiz) is a grouping of the larger islands in the Caribbean Sea, including Cuba, Hispaniola, Puerto Rico, Jamaica, and the ...

Greater Antilles
(including
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is a country comprising the island of Cuba, as well as Isla de la Juventud Isla de la Juventud (; en, Isle of Youth) is the second-largest Cuban islan ...

Cuba
and
Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and the region's se ...
), the
Lesser Antilles The Lesser Antilles ( es, link=no, Antillas Menores; french: link=no, Petites Antilles; pap, Antias Menor; nl, Kleine Antillen) are a group of islands in the Caribbean Sea The Caribbean Sea ( es, Mar Caribe; french: Mer des Caraïbes; ht, ...
, the
Canary Island The Canary Islands (; es, Islas Canarias, ), also known informally as ''the Canaries'', is a Spanish archipelago in the Atlantic Ocean, in a region known as Macaronesia. At their closest point to the African mainland, they are west of Morocco ...
s,
Malta Malta ( , , ), officially known as the Republic of Malta ( mt, Repubblika ta' Malta ) and formerly Melita, is a Southern European island country consisting of an archipelago in the Mediterranean Sea. It lies south of Italy, east of Tunisi ...

Malta
, the
Seychelles Seychelles (; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic island country An island country or an island nation is a country A country ...

Seychelles
, the
Mascarene Islands The Mascarene Islands (, ) or Mascarenes or Mascarenhas Archipelago is a group of islands in the Indian Ocean east of Madagascar consisting of the islands belonging to the Republic of Mauritius as well as the French department of Réunion. Their ...
(including
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label=Mauritian Creole Mauritian Creole or Morisien or formerly Morisyen ( mfe, kreol morisien, links=no ) is a French-based creole language spoken in Mauritius Mauritius ( ; french: ...

Mauritius
and ), and
Madagascar Madagascar (; mg, Madagasikara), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), and previously known as the Malagasy Republic The Malagasy Republic ( mg, Repoblika Mal ...

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 ''Megalochelys'' is an extinct genus of cryptodiran tortoises that lived from the Miocene to Pleistocene, across Asia and possibly Eastern Europe. They are noted for their giant size, which is among the largest of any known testudine, with a maxim ...
'', ''
Chelonoidis ''Chelonoidis'' is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835. They are found in South America and the Galápagos Islands, and formerly had a wide distribution in the West Indies. The members of this genus on ...
'', ''
Centrochelys ''Centrochelys'' is a genus (biology), genus of tortoise. It contains one extant species and several extinct species: * ''Centrochelys atlantica'' * ''Centrochelys burchardi'' * ''Centrochelys marocana'' * ''Centrochelys robusta'' * ''Centrochely ...
'', ''
Aldabrachelys ''Aldabrachelys'' is the recognised genus for the Seychelles and Madagascan radiations of giant tortoises, including the Aldabra giant tortoise (''Aldabrachelys gigantea''). Naming This name is problematic in that the type specimen is actually re ...
'', ''
Cylindraspis ''Cylindraspis'' is a genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer ...

Cylindraspis
'', and ''
Hesperotestudo ''Hesperotestudo'' ("Western turtle") is an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system tha ...
''), but are thought to have independently evolved large body size through
convergent evolution Convergent evolution is the independent evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; eit ...
. Giant tortoises are notably absent from
Australasia Australasia is a region In geography Geography (from Ancient Greek, Greek: , ''geographia'', literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and phenomena of the Earth ...

Australasia
and many south
Pacific The Pacific Ocean is the largest and deepest of Earth Earth is the third planet from the Sun and the only astronomical object known to harbour and support life. 29.2% of Earth's surface is land consisting of continents and islands. ...

Pacific
islands, but the distantly related
meiolaniid Meiolaniidae is an extinct family of large, possibly herbivorous stem-turtles with heavily armored heads and tails known from South America and most of Oceania. Though once believed to be cryptodires, they are not closely related to any living spe ...
turtles are thought to have filled the same niche.
Giant tortoise Giant tortoises are any of several species of various large land tortoise Tortoises () are reptile species of the family Testudinidae of the order Testudines (from the Latin name for tortoise). They are particularly distinguished from other ...
s are also known from the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define ...
-
Pliocene The Pliocene ( ; also Pleiocene) epoch (geology), Epoch is the epoch in the geologic timescale that extends from 5.333 million to 2.58North America North America is a continent A continent is any of several large landmasses. Generally identified by convention (norm), convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continen ...

North America
,
South America South America is a continent A continent is any of several large landmasses. Generally identified by convention (norm), convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continent ...

South America
,
Europe Europe is a continent A continent is any of several large landmass A landmass, or land mass, is a large region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a field of scienc ...

Europe
,
Asia Asia () is Earth's largest and most populous continent, located primarily in the Eastern Hemisphere, Eastern and Northern Hemisphere, Northern Hemisphere of the Earth, Hemispheres. It shares the continental landmass of Eurasia with the cont ...

Asia
, and
Africa Africa is the world's second-largest and second-most populous continent A continent is any of several large landmass A landmass, or land mass, is a large region In geography Geography (from Greek: , ''geographia'', ...

Africa
, but are all now extinct, which is also attributed to human activity.


Diet

Tortoises are generally considered to be strict
herbivores A herbivore is an animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All ...
, feeding on grasses, weeds, leafy greens, flowers, and some fruits. However, hunting and eating of birds has been observed on occasion. Pet tortoises typically require diets based on wild grasses, weeds, leafy greens and certain flowers. Certain species consume
worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limb Limb can refer to: *Limb (anatomy), an appendage of a human or animal *Limb Music, a record label *Limb (album), an ...

worm
s or
insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to be in ...

insect
s 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 Visual communication is the use of visual elements to convey ideas and information"7 Paramount Components of Visual Communication". ''Infographic Design Team - Infographics Design - Data Visualization''. Retrieved 2021-01-27. https://www.infographi ...
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 very important in tortoises during combat and courtship. In both combat and courtship, tortoises use ramming to communicate with other individuals.


Taxonomy

This species list largely follows
Turtle Taxonomy Working GroupThe Turtle Taxonomy Working Group (TTWG) is an informal working group of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group (TFTSG). It is composed of a number of leading turtle Turtles are reptiles of the Order (biology), order Che ...
(2019) and the Turtle Extinctions Working Group (2015). Family Testudinidae Batsch 1788August 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. * ''Agrionemys'' Khosatzky & Młynarski, 1966 ** ''Russian tortoise, Agrionemys horsfieldii'', Russian tortoise * ''Alatochelon'' ** ''Alatochelon myrteum'' * ''
Aldabrachelys ''Aldabrachelys'' is the recognised genus for the Seychelles and Madagascan radiations of giant tortoises, including the Aldabra giant tortoise (''Aldabrachelys gigantea''). Naming This name is problematic in that the type specimen is actually re ...
'' Loveridge and Williams 1957:166 ** ''Aldabra giant tortoise, Aldabrachelys gigantea'' Aldabran giant tortoise. ** Extinction, †''Aldabrachelys abrupta'' Late Holocene, extinct ''circa'' 1200 AD ** Extinction, †''Aldabrachelys grandidieri'' Late Holocene, extinct ''circa'' 884 AD * ''Astrochelys'' John Edward Gray, Gray, 1873:4 ** ''radiated tortoise, Astrochelys radiata'', radiated tortoise ** ''angonoka, Astrochelys yniphora'', angonoka tortoise, (Madagascan) plowshare tortoise * ''
Centrochelys ''Centrochelys'' is a genus (biology), genus of tortoise. It contains one extant species and several extinct species: * ''Centrochelys atlantica'' * ''Centrochelys burchardi'' * ''Centrochelys marocana'' * ''Centrochelys robusta'' * ''Centrochely ...
'' Gray 1872:5Gray, 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. ** ''Centrochelys atlantica'' ** ''Centrochelys burchardi'' ** ''Centrochelys marocana'' ** ''Centrochelys robusta'' ** ''African spurred tortoise, Centrochelys sulcata'', African spurred tortoise, sulcata tortoise ** ''Centrochelys vulcanica'' * ''
Chelonoidis ''Chelonoidis'' is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835. They are found in South America and the Galápagos Islands, and formerly had a wide distribution in the West Indies. The members of this genus on ...
'' Leopold Fitzinger, Fitzinger 1835:112 ** ''Chelonoidis alburyorum'' Abaco tortoise, Late Pleistocene, extinct ''c.'' 1400 CE ** ''red-footed tortoise, Chelonoidis carbonarius'', red-footed tortoise ** ''Argentine tortoise, Chelonoidis chilensis'', Chaco tortoise, Argentine tortoise or southern wood tortoise ** ''Chelonoidis cubensis'' Cuban giant tortoise ** ''Yellow-footed tortoise, Chelonoidis denticulatus'' Brazilian giant tortoise, yellow-footed tortoise * ''Chelonoidis dominicensis, C. dominicensis'' Dominican giant tortoise ** ''Chelonoidis lutzae'' Lutz's giant tortoise, Late Pleistocene ** ''Chelonoidis monensis'' Mona tortoise ** ''Chelonoidis niger'' complex: *** ''Chelonoidis abingdonii'', Pinta Island giant tortoise, Abingdon Island giant tortoise (extinct) *** ''Chelonoidis becki'', Wolf Volcano giant tortoise, Cape Berkeley giant tortoise *** ''Chelonoidis chathamensis'', San Cristobal giant tortoise, Chatham Island giant tortoise *** ''Chelonoidis darwini'', San Salvador giant tortoise, James Island giant tortoise *** ''Chelonoidis duncanensis'', Pinzon giant tortoise, Duncan Island giant tortoise *** ''Chelonoidis hoodensis'', Española giant tortoise, Hood Island giant tortoise *** ''Chelonoidis niger'', Floreana giant tortoise, Charles Island giant tortoise (extinct) *** ''Chelonoidis phantasticus'', Fernandina giant tortoise, Narborough Island giant tortoise *** ''Chelonoidis porteri'', Santa Cruz giant tortoise, Indefatigable Island giant tortoise *** ''Chelonoidis vicina'', Isabela Island giant tortoise, Albemarle Island giant tortoise ** ''Chelonoidis sellovii'' Southern Cone giant tortoise, Pleistocene ** ''Chelonoidis sombrerensis'' Sombrero giant tortoise, Late Pleistocene * ''Chersina'' Gray 1830:5 ** ''Angulate tortoise, Chersina angulata'', angulated tortoise, South African bowsprit tortoise * ''Cheirogaster'' Bergounioux 1935:78 ** †''Cheirogaster gymnesica'' Late Pliocene to Early Pleistocene ** †''Cheirogaster schafferi'' Pliocene to Early Pleistocene * ''Chersobius'' Fitzinger, 1835 ** ''Chersobius boulengeri'', Karoo padloper, Karoo dwarf tortoise, Boulenger's Cape tortoise ** ''Chersobius signatus'', speckled padloper tortoise ** ''Chersobius solus'', Nama padloper, Berger's Cape tortoise * †''
Cylindraspis ''Cylindraspis'' is a genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer ...

Cylindraspis
'' Fitzinger 1835:112 (all species extinct) following Austin and Arnold, 2001: ** ''Reunion giant tortoise, Cylindraspis indica'', synonym ''Cylindraspis borbonica'', Reunion giant tortoise ** ''saddle-backed Mauritius giant tortoise, Cylindraspis inepta'', saddle-backed Mauritius giant tortoise or Mauritius giant domed tortoise ** ''domed Rodrigues giant tortoise, Cylindraspis peltastes'', domed Rodrigues giant tortoise ** ''domed Mauritius giant tortoise, Cylindraspis triserrata'', domed Mauritius giant tortoise or Mauritius giant flat-shelled tortoise ** ''saddle-backed Rodrigues giant tortoise, Cylindraspis vosmaeri'', saddle-backed Rodrigues giant tortoise * ''Geochelone'' Fitzinger 1835:112 ** ''Geochelone burchardi'' Tenerife giant tortoise ** ''Geochelone vulcanica'' Gran Canaria giant tortoise ** ''Indian star tortoise, Geochelone elegans'', Indian star tortoise ** ''Burmese star tortoise, Geochelone platynota'', Burmese star tortoise ** ''Geochelone robusta'' Malta giant tortoise * ''Gopherus'' Constantine Rafinesque, Rafinesque 1832:64 ** ''Desert tortoise, Gopherus agassizii'', Mojave desert tortoise, Agassiz's desert tortoise ** ''Texas tortoise, Gopherus berlandieri'', Texas tortoise, Berlandier's tortoise ** ''Bolson tortoise, Gopherus flavomarginatus'', Bolson tortoise ** ''Sonoran Desert tortoise, Gopherus morafkai'', Sonoran desert tortoise, Morafka's desert tortoise ** ''Gopherus polyphemus'', gopher tortoise * ''Hadrianus (genus), Hadrianus'' ** ''Hadrianus corsoni'' (syn. ''H. octonarius'') ** ''Hadrianus robustus'' ** ''Hadrianus schucherti'' ** ''Hadrianus utahensis'' * ''
Hesperotestudo ''Hesperotestudo'' ("Western turtle") is an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system tha ...
'' ** ''Hesperotestudo alleni'' ** ''Hesperotestudo angusticeps'' ** ''Hesperotestudo brontops'' ** ''Hesperotestudo equicomes'' ** ''Hesperotestudo impensa'' ** ''Hesperotestudo incisa'' ** ''Hesperotestudo johnstoni'' ** ''Hesperotestudo kalganensis'' ** ''Hesperotestudo niobrarensis'' ** ''Hesperotestudo orthopygia'' ** ''Hesperotestudo osborniana'' ** ''Hesperotestudo percrassa'' ** ''Hesperotestudo riggsi'' ** ''Hesperotestudo tumidus'' ** ''Hesperotestudo turgida'' ** ''Hesperotestudo wilsoni'' * ''Homopus'' Duméril and Bibron 1834:357Duméril, André Marie Constant and Bibron, Gab riel. 1834. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Premier. Paris: Roret, 439 pp. ** ''Homopus areolatus'', common padloper, parrot-beaked tortoise, beaked Cape tortoise ** ''Homopus femoralis'', greater padloper, greater dwarf tortoise * ''Indotestudo'' Wassili Adolfovitch Lindholm, Lindholm, 1929 ** ''Indotestudo elongata'', elongated tortoise, yellow-headed tortoise ** ''Indotestudo forstenii'', Forsten's tortoise, East Indian tortoise ** ''Indotestudo travancorica'', Travancore tortoise * ''Kinixys'' ** ''Kinixys belliana'', Bell's hinge-back tortoise ** ''Kinixys erosa'', forest hinge-back tortoise, serrated hinge-back tortoise ** ''Kinixys homeana'', Home's hinge-back tortoise ** ''Kinixys lobatsiana'', Lobatse hinge-back tortoise ** ''Kinixys natalensis'', Natal hinge-back tortoise ** ''Kinixys spekii'', Speke's hinge-back tortoise * ''Malacochersus'' Lindholm 1929:285 ** ''pancake tortoise, Malacochersus tornieri'', pancake tortoise * ''Manouria'' Gray 1854:133 ** ''Manouria emys'', Asian giant tortoise, brown tortoise (mountain tortoise) ** ''Impressed tortoise, Manouria impressa'', impressed tortoise * ''
Megalochelys ''Megalochelys'' is an extinct genus of cryptodiran tortoises that lived from the Miocene to Pleistocene, across Asia and possibly Eastern Europe. They are noted for their giant size, which is among the largest of any known testudine, with a maxim ...
'' Falconer, H. and Cautley, P.T. 1837. ** ''Megalochelys atlas'', Atlas tortoise, Extinct – Pliocene to Pleistocene ** ''Megalochelys cautleyi'', Cautley's giant tortoise * ''Psammobates'' Fitzinger 1835:113 ** ''geometric tortoise, Psammobates geometricus'', geometric tortoise ** ''Psammobates oculifer'', serrated tent tortoise, Kalahari tent tortoise ** ''Psammobates tentorius'', African tent tortoise * ''Pyxis (genus), Pyxis'' Bell 1827:395 ** ''spider tortoise, Pyxis arachnoides'', (Madagascan) spider tortoise ** ''flat-backed spider tortoise, Pyxis planicauda'', flat-backed spider tortoise, (Madagascan) flat-tailed tortoise, flat-tailed spider tortoise * ''Stigmochelys'' Gray, 1873 ** ''Leopard tortoise, Stigmochelys pardalis'', leopard tortoise * ''Stylemys'' ** ''Stylemys botti'' ** ''Stylemys calaverensis'' ** ''Stylemys canetotiana'' ** ''Stylemys capax'' ** ''Stylemys conspecta'' ** ''Stylemys copei'' ** ''Stylemys emiliae'' ** ''Stylemys frizaciana'' ** ''Stylemys karakolensis'' ** ''Stylemys nebrascensis'' (syn. ''Stylemys amphithorax, S. amphithorax'') ** ''Stylemys neglectus'' ** ''Stylemys oregonensis'' ** ''Stylemys pygmea'' ** ''Stylemys uintensis'' ** ''Stylemys undabuna'' * ''Testudo (genus), Testudo'' ** ''Spur-thighed tortoise, Testudo graeca'', Greek tortoise, spur-thighed tortoise, Moorish tortoise ** ''Hermann's tortoise, Testudo hermanni'', Hermann's tortoise ** ''Kleinmann's tortoise, Testudo kleinmanni'', Egyptian tortoise, including Negev tortoise ** ''Marginated tortoise, Testudo marginata'', marginated tortoise


Phylogeny

A molecular phylogeny of tortoises, following Le et al. (2006: 525): A separate phylogeny via mtDNA analysis was found by Kehlmaier ''et al.'' (2021):


In human culture


In religion

In Hinduism, Kurma ( sa, कुर्म) was the second Avatar of Vishnu. Like the Matsya Avatara, Kurma also belongs to the Satya Yuga. Vishnu took the form of a half-man, half-tortoise, the lower half being a tortoise. He is normally shown as having four arms. He sat on the bottom of the ocean after the Deluge (mythology), Great Flood. A mountain was placed on his back by the other deity, gods so they could churn the sea and find the ancient treasures of the Vedic period, Vedic peoples. In Judaism and early Christianity tortoises were seen as unclean animals. Tortoise shells were used by ancient Chinese as oracle bones to make predictions. The tortoise is a symbol of the Ancient Greek god, Hermes.


As pets

See Turtle#As_pets, turtle


As food

See turtle soup


In fiction

Teenage Mutant Ninja Turtles is an entertainment franchise. The four anthropomorphic turtle brothers trained in ninjitsu are shown to fight evil in New York City.


Gallery

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


See also

* Cultural depictions of turtles * Jackson ratio * Tortoise Protection Group


References


Further reading

* * * *


External links


Family Testudinidae (Tortoises)
The Reptile Database
Chelonia
Conservation and care of turtles.
Live Tortoise Stream
Live Tortoise Stream {{Authority control Tortoises, Testudinidae, Taxa named by August Batsch