Turtle

Turtles are reptiles of the order Testudines (or Chelonii[3]) characterized by a special bony or
cartilaginous shell developed from their ribs and acting as a shield.[4] "Turtle" may refer to the order
Turtles
[5]
as a whole (American English) or to fresh-water and sea-dwelling testudines (British English). Temporal range:
Late Jurassic Present ,[1] 1570 Ma
The order Testudines includes both extant (living) and extinct species. The earliest known members of
Pre OS D C P T J K Pg N
this group date from 157 million years ago,[1] making turtles one of the oldest reptile groups and a
more ancient group than snakes or crocodilians. Of the 327 known species alive today, some are
highly endangered.[6][7]

Turtles are ectothermsanimals commonly called cold-bloodedmeaning that their internal

temperature varies according to the ambient environment. However, because of their high metabolic
rate, leatherback sea turtles have a body temperature that is noticeably higher than that of the
surrounding water.

Turtles are classified as amniotes, along with other reptiles, birds, and mammals. Like other amniotes,
turtles breathe air and do not lay eggs underwater
, although many species live in or around water
. Florida Box Turtle (Terrapene carolina)

Scientific classification
The study of turtles is called cheloniology, after the Greek word for turtle. It is also sometimes called
testudinology, after the Latin name for turtles. Kingdom: Animalia
Phylum: Chordata
Class: Reptilia

Contents Clade: Testudinata

Turtle, tortoise, or terrapin Order: Testudines

Batsch, 1788 [2]
Anatomy and morphology
Neck retraction
Subgroups[1]
Head
Shell Cryptodira
Skin and molting Pleurodira
Limbs
Meiolaniidae
Behavior and see text
Senses
Communication
Diversity
Intelligence
Ecology and life history 14 extant families with ca. 300
Diet species
Systematics and evolution
Phylogeny of living turtles
Classification of turtles
Fossil record
Genomics
In captivity blue: sea turtles, black: land turtles
As food, traditional medicine, and cosmetics
Conservation status
Testudinology research organizations
See also
Notes
References
Further reading
External links
Turtle, tortoise, or terrapin
Turtle, tortoise, and terrapin

Green sea turtle African spurred tortoise Red-eared slider (terrapin)

The word chelonian is popular among veterinarians, scientists, and conservationists working with these animals as a catch-all name for any member of
the superorder Chelonia, which includes all turtles living and extinct, as well as their immediate ancestors. Chelonia is based on the Greek word
chelone "tortoise", "turtle" (another relevant word is chelys "tortoise"),[8][9] also denoting armor or interlocking shields;[10] testudines, on the other
hand, is based on the Latin word testudo "tortoise".[11] "Turtle" may either refer to the order as a whole, or to particular turtles that make up a form taxon
that is not monophyletic.

The meaning of the word turtle differs from region to region. In North America, all chelonians are commonly called turtles, including terrapins and
tortoises.[12][13] In Great Britain, the wordturtle is used for sea-dwelling species, but not for tortoises.

The term tortoise usually refers to any land-dwelling, non-swimming chelonian.[13] Most land-dwelling chelonians are in the Testudinidae family, only
one of the fourteen extant turtle families.[14]

Terrapin is used to describe several species of small, edible, hard-shell turtles, typically those found in brackish waters, and is an Algonquian word for
turtle.[12][15]

Some languages do not have this distinction, as all of these are referred to by the same name. For example, in Spanish, the word tortuga is used for
turtles, tortoises, and terrapins. A sea-dwelling turtle istortuga marina, a freshwater species tortuga de ro, and a tortoise tortuga terrestre.[16]

Anatomy and morphology

The largest living chelonian is the leatherback sea turtle (Dermochelys coriacea), which reaches a shell length of 200 cm (6.6 ft) and can reach a weight
of over 900 kg (2,000 lb). Freshwater turtles are generally smaller, but with the largest species, the Asian softshell turtle Pelochelys cantorii, a few
individuals have been reported up to 200 cm (6.6 ft). This dwarfs even the better-known alligator snapping turtle, the largest chelonian in North America,
which attains a shell length of up to 80 cm (2.6 ft) and weighs as much as 113.4 kg (250 lb).[17] Giant tortoises of the genera Geochelone, Meiolania, and
others were relatively widely distributed around the world into prehistoric times, and are known to have existed in North and South America, Australia,
and Africa. They became extinct at the same time as the appearance of man, and it is assumed humans hunted them for food. The only surviving giant
[18]
tortoises are on the Seychelles and Galpagos Islands and can grow to over 130 cm (51 in) in length, and weigh about 300 kg (660 lb).

[19]
The largest ever chelonian wasArchelon ischyros, a Late Cretaceous sea turtle known to have been up to 4.6 m (15 ft) long.

The smallest turtle is the speckled padloper tortoise of South Africa. It measures no more than 8 cm (3.1 in) in length and weighs about 140 g (4.9 oz).
Two other species of small turtles are the American mud turtles and musk turtles that live in an area that ranges from Canada to South America. The shell
length of many species in this group is less than 13 cm (5.1 in) in length.

Neck retraction
Turtles are divided into two groups, according to how they retract their necks into their shells (something the ancestral Proganochelys could not do). The
Cryptodira retract their necks backwards while contracting it under their spine, whereas the
Pleurodira contract their necks to the side.

Head
Most turtles that spend most of their lives on land have their eyes looking down at objects in front of them. Some aquatic turtles, such as snapping turtles
and soft-shelled turtles, have eyes closer to the top of the head. These species of turtle can hide from predators in shallow water, where they lie entirely
submerged except for their eyes and nostrils. Near their eyes, sea turtles possess glands that produce salty tears that rid their body of excess salt taken in
from the water they drink.
Turtles have rigid beaks and use their jaws to cut and chew
Neck retraction in turtles
food. Instead of having teeth, which they appear to have lost
about 150-200 million years ago,[20] the upper and lower
jaws of the turtle are covered by horny ridges. Carnivorous
turtles usually have knife-sharp ridges for slicing through
their prey. Herbivorous turtles have serrated-edged ridges that
help them cut through tough plants. They use their tongues to
swallow food, but unlike most reptiles, they cannot stick out
their tongues to catch food.

Pleurodires retract their neck Cryptodires retract their neck

sideways backwards
Shell
The upper shell of the turtle is called the carapace. The lower
shell that encases the belly is called the plastron. The carapace and plastron are joined together on the turtle's sides by bony structures called bridges. The
inner layer of a turtle's shell is made up of about 60 bones that include portions of the backbone and the ribs, meaning the turtle cannot crawl out of its
shell. In most turtles, the outer layer of the shell is covered by horny scales called
scutes that are part of its outer skin, or epidermis. Scutes are made up of
the fibrous protein keratin that also makes up the scales of other reptiles. These scutes overlap the seams between the shell bones and add strength to the
shell. Some turtles do not have horny scutes; for example, the leatherback sea turtle and the soft-shelled turtles have shells covered with leathery skin
instead.

The rigid shell means that turtles cannot breathe as other reptiles do, by changing the volume of their chest cavities via expansion and contraction of the
ribs. Instead, they breathe in two ways. First, they employ limb pumping, sucking air into their lungs and pushing it out by moving the limbs in and out
relative to the shell. Secondly, when the abdominal muscles that cover the posterior opening of the shell contract, the pressure inside the shell and lungs
decreases, drawing air into the lungs, allowing these muscles to function in much the same way as the mammalian diaphragm. A second set of abdominal
muscles face the opposite way and when they contract they expel air under positive pressure.[21] The shape of the shell gives helpful clues about how a
turtle lives. Most tortoises have a large, dome-shaped shell that makes it difficult for predators to crush the shell between their jaws. One of the few
exceptions is the Africanpancake tortoise, which has a flat, flexible shell that allows it to hide in rock crevices. Most aquatic turtles have flat, streamlined
shells, which aid in swimming and diving. American snapping turtles and musk turtles have small, cross-shaped plastrons that give them more efficient
leg movement for walking along the bottom of ponds and streams. Another exception is the Belawan Turtle (Cirebon, West Java), which has sunken-back
soft-shell.

The color of a turtle's shell may vary. Shells are commonly colored brown, black, or olive green. In some species, shells may have red, orange, yellow, or
grey markings, often spots, lines, or irregular blotches. One of the most colorful turtles is the eastern
painted turtle, which includes a yellow plastron and a
black or olive shell with red markings around the rim.

Tortoises, being land-based, have rather heavy shells. In contrast, aquatic and soft-shelled turtles have lighter shells that help them avoid sinking in water
and swim faster with more agility. These lighter shells have large spaces called fontanelles between the shell bones. The shells of leatherback sea turtles
are extremely light because they lack scutes and contain many fontanelles.

It has been suggested by Jackson (2002) that the turtle shell can function as pH buffer. To endure through anoxic conditions, such as winter periods
trapped beneath ice or within anoxic mud at the bottom of ponds, turtles utilize two general physiological mechanisms. In the case of prolonged periods
fers and uptakes lactic acid.[22]
of anoxia, it has been shown that the turtle shell both releases carbonate buf

Skin and molting

As mentioned above, the outer layer of the shell is part of the skin; each scute (or plate) on the shell
corresponds to a single modified scale. The remainder of the skin has much smaller scales, similar to the
skin of other reptiles. Turtles do not molt their skins all at once as snakes do, but continuously in small
pieces. When turtles are kept in aquaria, small sheets of dead skin can be seen in the water (often
appearing to be a thin piece of plastic) having been sloughed off when the animals deliberately rub
themselves against a piece of wood or stone. Tortoises also shed skin, but dead skin is allowed to
accumulate into thick knobs and plates that provide protection to parts of the body outside the shell.

By counting the rings formed by the stack of smaller, older scutes on top of the larger, newer ones, it is
Tail of a snapping turtle
possible to estimate the age of a turtle, if one knows how many scutes are produced in a year.[23] This
method is not very accurate, partly because growth rate is not constant, but also because some of the
scutes eventually fall away from the shell.
Limbs
Terrestrial tortoises have short, sturdy feet. Tortoises are famous for moving slowly, in part because of their heavy, cumbersome shells, which restrict
stride length.

Amphibious turtles normally have limbs similar to those of tortoises, except that the feet are webbed and often
have long claws. These turtles swim using all four feet in a way similar to the dog paddle, with the feet on the left
and right side of the body alternately providing thrust. Large turtles tend to swim less than smaller ones, and the
very big species, such as alligator snapping turtles, hardly swim at all, preferring to walk along the bottom of the
river or lake. As well as webbed feet, turtles have very long claws, used to help them clamber onto riverbanks and
floating logs upon which they bask. Male turtles tend to have particularly long claws, and these appear to be used
to stimulate the female while mating. While most turtles have webbed feet, some, such as the pig-nosed turtle,
have true flippers, with the digits being fused into paddles and the claws being relatively small. These species
swim in the same way as sea turtles do (see below).

Sea turtles are almost entirely aquatic and have flippers instead of feet. Sea turtles fly through the water, using the
up-and-down motion of the front flippers to generate thrust; the back feet are not used for propulsion but may be
used as rudders for steering. Compared with freshwater turtles, sea turtles have very limited mobility on land, and Skeleton of snapping turtle
(Chelydra serpentina)
apart from the dash from the nest to the sea as hatchlings, male sea turtles normally never leave the sea. Females
must come back onto land to lay eggs. They move very slowly and laboriously
, dragging themselves forwards with
their flippers.

Behavior

Senses
Turtles are thought to have exceptional night vision due to the unusually large number of rod cells in their retinas. Turtles have color vision with a wealth
of cone subtypes with sensitivities ranging from the near ultraviolet (UVA) to red. Some land turtles have very poor pursuit movement abilities, which are
normally found only in predators that hunt quick-moving prey
, but carnivorous turtles are able to move their heads quickly to snap.

Communication
While typically thought of as mute, turtles make various sounds when communicating. Tortoises may be
vocal when courting and mating. Various species of both freshwater and sea turtles emit numerous types
of calls, often short and low frequency, from the time they are in the egg to when they are adults. These
[24]
vocalizations may serve to create group cohesion when migrating.

Intelligence
It has been reported that wood turtles are better than white rats at learning to navigate mazes.[25] Case
The Arrau turtle has a sizable vocal
studies exist of turtles playing.[25] They do, however, have a very low encephalization quotient (relative
repertoire.[24]
brain to body mass), and their hard shells enable them to live without fast reflexes or elaborate predator
avoidance strategies.[26] In the laboratory, turtles (Pseudemys nelsoni) can learn novel operant tasks and
[27]
have demonstrated a long-term memory of at least 7.5 months.

Ecology and life history

Although many turtles spend large amounts of their lives underwater, all turtles and tortoises breathe air and must surface at regular intervals to refill their
lungs. They can also spend much or all of their lives on dry land. Aquatic respiration in Australian freshwater turtles is currently being studied. Some
species have large cloacal cavities that are lined with many finger-like projections. These projections, called papillae, have a rich blood supply and
increase the surface area of the cloaca. The turtles can take up dissolved oxygen from the water using these papillae, in much the same way that fish use
gills to respire.[28]

Like other reptiles, turtles lay eggs that are slightly soft and leathery. The eggs of the largest species are spherical while the eggs of the rest are elongated.
Their albumen is white and contains a different protein from bird eggs, such that it will not coagulate when cooked. Turtle eggs prepared to eat consist
mainly of yolk. In some species, temperature determines whether an egg develops into a male or a female: a higher temperature causes a female, a lower
temperature causes a male. Large numbers of eggs are deposited in holes dug into mud or sand. They are
then covered and left to incubate by themselves. Depending on the species, the eggs will typically take
70120 days to hatch. When the turtles hatch, they squirm their way to the surface and head toward the
water. There are no known species in which the mother cares for her young.

Sea turtles lay their eggs on dry, sandy beaches. Immature sea turtles are not cared for by the adults.
Turtles can take many years to reach breeding age, and in many cases, breed every few years rather than
annually.

Researchers have recently discovered a turtle's organs do not gradually break down or become less Play media
efficient over time, unlike most other animals. It was found that the liver, lungs, and kidneys of a Sea turtle swimming
centenarian turtle are virtually indistinguishable from those of its immature counterpart. This has
[29]
inspired genetic researchers to begin examining the turtle genome for longevity genes.

A group of turtles is known as a bale.[30]

Diet
A turtle's diet varies greatly depending on the environment in which it lives. Adult turtles typically eat
aquatic plants; invertebrates such as insects, snails, and worms; and have been reported to occasionally
eat dead marine animals. Several small freshwater species are carnivorous, eating small fish and a wide
range of aquatic life. However, protein is essential to turtle growth and juvenile turtles are purely
carnivorous.

Sea turtles typically feed on jellyfish, sponge, and other soft-bodied organisms. Some species of sea
turtle with stronger jaws have been observed to eat shellfish while some species, such as the green sea
gely made up of algae.[31]
turtle, do not eat any meat at all and, instead, have a diet lar
A green sea turtle grazing on
seagrass
Systematics and evolution
Based on body fossils, the first proto-turtles are believed to have existed in the late
Triassic Period of the Mesozoic
era, about 220 million years ago, and their shell, which has remained a remarkably stable body plan, is thought to
have evolved from bony extensions of their backbones and broad ribs that expanded and grew together to form a
complete shell that offered protection at every stage of its evolution, even when the bony component of the shell
was not complete. This is supported by fossils of the freshwater Odontochelys semitestaceaor "half-shelled turtle
with teeth", from the late Triassic, which have been found near Guangling in southwest China. Odontochelys
displays a complete bony plastron and an incomplete carapace, similar to an early stage of turtle embryonic
development.[32] Prior to this discovery, the earliest-known fossil turtle ancestors, like Proganochelys, were
terrestrial and had a complete shell, offering no clue to the evolution of this remarkable anatomical feature. By the
late Jurassic, turtles had radiated widely, and their fossil history becomes easier to read. Life restoration of
Odontochelys semitestacea,
Their exact ancestry has been disputed. It was believed they are the only surviving branch of the ancient the oldest known turtle
evolutionary grade Anapsida, which includes groups such as procolophonids, millerettids, protorothyrids, and relative with a partial shell
pareiasaurs. All anapsid skulls lack a temporal opening while all other extant amniotes have temporal openings
(although in mammals, the hole has become the zygomatic arch). The millerettids, protorothyrids, and pareiasaurs
became extinct in the latePermian period and the procolophonoids during the Triassic.[33]

However, it was later suggested that the anapsid-like turtle skull may be due to reversion rather than to anapsid descent. More recent morphological
phylogenetic studies with this in mind placed turtles firmly within diapsids, slightly closer to Squamata than to Archosauria.[34][35] All molecular studies
have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria,[36] or, more commonly, as a sister group to extant
archosaurs,[37][38][39][40] though an analysis conducted by Lyson et al. (2012) recovered turtles as the sister group of lepidosaurs instead.[41] Reanalysis
of prior phylogenies suggests that they classified turtles as anapsids both because they assumed this classification (most of them studying what sort of
anapsid turtles are) and because they did not sample fossil and extant taxa broadly enough for constructing the cladogram. Testudines were suggested to
have diverged from other diapsids between 200 and 279 million years ago, though the debate is far from settled.[34][37][42] Even the traditional placement
of turtles outside Diapsida cannot be ruled out at this point. A combined analysis of morphological and molecular data conducted by Lee (2001) found
turtles to be anapsids (though a relationship with archosaurs couldn't be statistically rejected).[43] Similarly, a morphological study conducted by Lyson et
al. (2010) recovered them as anapsids most closely related to Eunotosaurus.[44] A molecular analysis of 248 nuclear genes from 16 vertebrate taxa
suggests that turtles are a sister group to birds and crocodiles (the Archosauria).[45] The date of separation of turtles and birds and crocodiles was
estimated to be 255 million years ago. The most recent common ancestor of living turtles, corresponding to the split between Pleurodira and Cryptodira,

[1][46]
was estimated to have occurred around 157 million years ago.[1][46] The oldest definitive crown-group turtle
(member of the modern clade Testudines) is the species Caribemys oxfordiensis from the late Jurassic period
(Oxfordian stage).[1] Through utilizing the first genomic-scale phylogenetic analysis of ultraconserved elements
(UCEs) to investigate the placement of turtles within reptiles, Crawford et al. (2012) also suggest that turtles are a
[47]
sister group to birds and crocodiles (the Archosauria).

The first genome-wide phylogenetic analysis was completed by Wang et al. (2013). Using the draft genomes of
Chelonia mydas and Pelodiscus sinensis, the team used the largest turtle data set to date in their analysis and
concluded that turtles are likely a sister group of crocodilians and birds (Archosauria).[48] This placement within
the diapsids suggests that the turtle lineage lost diapsid skull characteristics as it now possesses an anapsid-like
skull.

The earliest known fully shelled member of the turtle lineage is the late Triassic Proganochelys. This genus
already possessed many advanced turtle traits, and thus probably indicates many millions of years of preceding "Chelonia" from Ernst
Haeckel's Kunstformen der
turtle evolution; this is further supported by evidence from fossil tracks from the Early Triassic of the United
Natur, 1904
States (Wyoming and Utah) and from the Middle Triassic of Germany, indicating that proto-turtles already existed
as early as Early Triassic.[49] Proganochelys lacked the ability to pull its head into its shell, had a long neck, and
had a long, spiked tail ending in a club. While this body form is similar to that of
ankylosaurs, it resulted from convergent evolution.

Turtles are divided into twoextant suborders: the Cryptodira and the Pleurodira. The Cryptodira is the larger of the two groups and includes all the marine
turtles, the terrestrial tortoises, and many of the freshwater turtles. The Pleurodira are sometimes known as the side-necked turtles, a reference to the way
they retract their heads into their shells. This smaller group consists primarily of various freshwater turtles.

Phylogeny of living turtles

[50] and Guillon et al. (2012).[51]
The following phylogeny of living turtles is based on the work of Crawford et al. (2015)

Pleurodira
Pelusios

Pelomedusidae

Pelomedusa

Pelomedusoides

Erymnochelys

Podocnemididae Peltocephalus

Podocnemis

Cheloides: Chelidae Chelodininae

Pseudemydura

Chelodina

Elusor

Flaviemys

Rheodytes

Elseya

Emydura

Myuchelys

Hydromedusinae Hydromedusa
 ?Rhinemys

Chelus

Chelinae Mesoclemmys

Phrynops

Platemys

Acanthochelys

Cryptodira Trionychia
Carettochelyidae Carettochelys

Trionychidae
Lissemys

Cyclanorbinae
Cyclanorbis

Cycloderma

Trionychinae
Trionyx

Pelochelys
Trionychini

Chitra

Rafetus

Apalone

Pelodiscus

Palea
Pelodiscini
Dogania

Amyda

Nilssonia

Durocryptodira
Macroclemys
Chelydridae

Chelydra

Dermatemydidae Dermatemys

Chelydroidea
Claudius
Staurotypinae
Staurotypus
Kinosternidae
 Sternotherus
Kinosterninae

Americhelydia Kinosternon

Dermochelyidae Dermochelys

Natator

Chelonia
Chelonioidea

Cheloniidae
Eretmochelys

Caretta

Lepidochelys

Testudinoidea Emysternia
Platysternidae Platysternon

Emydidae Emydinae
?Actinemys

Glyptemys

Emys [incl. Emydoidea]

Clemmys

Terrapene

Deirochelys

Chrysemys

Pseudemys

Deirochelyinae

Trachemys

Malaclemys

Graptemys

Testuguria Testudinidae
Manouria

Gopherinae

Gopherus
 Gopherus

Testunidinae
Malacochersus

Testunidini Indotestudo

Testudo

Dipsochelys

Pyxidini Astrochelys

Pyxis

Chersina

Homopus

Psammobates

Chelonoidis

Geochelonini Kinixys

Geochelone

Geoemydidae
Rhinoclemmyinae Rhinoclemmys

Geoemydinae
Geoemyda

Siebenrockiella [incl. Panayanemys]

Malayemys

Orlitia

Geoclemys

Morenia

Kachuga [incl. Pangshura]

Hardella
 Batagur [incl. Callagur]

Vijayachelys

Melanochelys

Mauremys [incl. Annamemys;

Cathaiemys; Emmenia; Chinemys]

Cuora [incl. Cistoclemmys]

Leucocephalon

Sacalia

Heosemys

Notochelys

Cyclemys

Classification of turtles
Order Testudinata Klein 1760[52]

Genus Pappochelys Schoch & Sues 2015

Family ProganochelyidaeBaur 1887
Family AustralochelidaeGaffney & Kitching 1994 sensu Lee 1997
Chart of the two extant suborders,
Family Proterochersidae Nopcsa 1928
extinct groups that existed within
Clade Mesochelydia these two suborders are shown as
Family IndochelyidaeDatta, Manna, Ghosh & Das 2000 well
Family HeckerochelyidaeSukhanov 2006
Clade Perichelydia

Family ChelycarapookidaeWarren 1969

Family Sichuanchelyidae Tong et al. 2012
Family Solemydidae de Lapparent de Broin & Murelaga 1996

Clade Meiolaniformes

Family Meiolaniidae Lydekker 1887

Family OtwayemyidaeGaffney et al. 1998
Genus Trapalcochelys Sterli, de la Fuente & Cerda 2013
Genus Chubutemys Gaffney et al. 2007
Two-month-old hypomelanistic
Genus Peligrochelys Sterli & de la Fuente 2012
snapping turtle
Family Kallokibotiidae Nopcsa 1923
Clade Testudines Linnaeus, 1758

Suborder Pleurodira Cope 1864

Family ApertotemporalidaeKhne 1937

 Family PlatychelyidaeBrn 1965 sensu Gaffney, Tong & Buffetaut 2006
Family Dortokidae Lapparent de Broin & Murelaga 1996
Family Notoemyidae Fernandez & Fuente 1994
Superfamily CheloidesGray 1825 sensu Gaffney, Tong & Buffetaut 2006

Family Chelidae Gray 1825

Superfamily Pelomedusoides Cope 1868 sensu Broin 1988

Family Araripemydidae Price 1973

Family Pelomedusidae (African sideneck turtles)
Family EuraxemydidaeGaffney, Tong & Buffetaut 2006
Family Bothremydidae Baur 1891
Family Podocnemididae Cope 1868 (Madagascan big-headedand American sideneck river turtles)
Suborder Cryptodira Dumril & Bibron 1835

Infraorder Paracryptodira

Family Pleurosternidae Cope 1868

Family Compsemyidae
Family Baenidae Cope 1882
Infraorder Eucryptodira Gaffney 1975a sensu Gaffney 1984

Family Macrobaenidae Sukhanov 1964

Family Eurysternidae Dollo 1886
Family Plesiochelyidae Baur 1888
Family Xinjiangchelyidae Nesov 1990
Clade Centrocryptodira

Family OsteopygidaeZangerl 1953

Family Sinemydidae Yeh 1963
Clade Polycryptodira Gaffney 1988

Clade Pantrionychia

Family Adocidae
Superfamily Trionychoidea Gray 1870

Family Carettochelyidae (pignose turtles)

Family Trionychidae (softshell turtles)
Superfamily Testudinoidea Baur 1893

Family Haichemydidae Sukhanov & Narmandakh 2006

Family Lindholmemydidae Chkhikvadze 1970
Family Sinochelyidae Chkhikvadze 1970
Family Emydidae (pond, box, and water turtles)
Family Geoemydidae (Asian river turtles, Asian leaf turtles, Asian box turtles, and roofed turtles)
Family Testudinidae (true tortoises)
Family Geoemydidae
Clade Americhelydia

Family Chelydridae (snapping turtles)

Superfamily Kinosternoidea

Family Dermatemydidae (river turtles)

Family Kinosternidae (mud turtles)
Superfamily Chelonioidea (sea turtles)

Family Toxochelyidae Baur 1895

Family Cheloniidae (green sea turtles and relatives)
Family Thalassemydidae
Family Dermochelyidae (leatherback sea turtles)
Family Protostegidae Cope, 1872

Fossil record Sea turtle at Henry Doorly Zoo,

Omaha NE
Turtle fossils of hatchling and nestling size have been documented in the scientific literature.[53]
Paleontologists from North Carolina State University have found the fossilized remains of the world's
[54]
largest turtle in a coal mine in Colombia. The specimen named as Carbonemys cofrinii is around 60 million years old and nearly 2.4 m (8 ft) long.
On a few rare occasions, paleontologists have succeeded in unearthing large numbers of Jurassic or Cretaceous turtle skeletons accumulated in a single
area (the Nemegt Formation in Mongolia, the Turtle Graveyard in North Dakota, or the Black Mountain Turtle Layer in Wyoming). The most spectacular
find of this kind to date occurred in 2009 in Shanshan County in Xinjiang, where over a thousand ancient freshwater turtles apparently died after the last
[55][56]
water hole in an area dried out during a major drought.

Though absent from New Zealand in recent times, turtle fossils are known from the Miocene Saint Bathans Fauna, represented by a meiolaniid and
pleurodires.[57]

Genomics
[58]
Turtles possess diverse chromosome numbers (2n = 2866) and a myriad of chromosomal rearrangements have occurred during evolution.

In captivity
Turtles, particularly small terrestrial and freshwater turtles, are commonly kept as pets. Among the most
popular are Russian tortoises, spur-thighed tortoises, and red-eared sliders.[59]

In the United States, due to the ease of contracting salmonellosis through casual contact with turtles, the
U.S. Food and Drug Administration (FDA) established a regulation in 1975 to discontinue the sale of
turtles under 4 in (100 mm).[60] It is illegal in every state in the U.S. for anyone to sell any turtles under
4 inches (10 cm) long. Many stores and flea markets still sell small turtles due to a loophole in the FDA
[61][62]
regulation which allows turtles under 4 in (100 mm) to be sold for educational purposes.

Some states have other laws and regulations regarding possession of red-eared sliders as pets because A red-eared slider basking on a
they are looked upon as invasive species or pests where they are not native, but have been introduced floating platform under a sun lamp
through the pet trade. As of July 1, 2007, it is illegal in Florida to sell any wild type red-eared slider.[63]
Unusual color varieties such as albino and pastel red-eared sliders, which are derived from captive
breeding, are still allowed for sale.

As food, traditional medicine, and cosmetics

The flesh of turtles, calipash or calipee, was and still is considered a delicacy in a number of
cultures.[6] Turtle soup has been a prized dish in Anglo-American cuisine,[64] and still remains so in
some parts of Asia. Gopher tortoise stew was popular with some groups in Florida.[65]

Left: The window of a restaurant serving
guilinggao, decorated with a ("turtle")
character
Right: Turtle plastrons among other plants
and animals parts are used in traditional
Chinese medicines. (Other items in the
image are dried lingzhi, snake, luo han
guo, and ginseng)
herbs) is the guilinggao jelly;[69] these days, though, it is typically made with only herbal ingredients.
Turtles remain a part of the traditional diet on the island of Grand Cayman, so much so that when
wild stocks became depleted, a turtle farm was established specifically to raise sea turtles for their
meat. The farm also releases specimens to the wild as part of an effort to repopulate the Caribbean
Sea.[66]
Fat from turtles is also used in the Caribbean and in Mexico as a main ingredient in cosmetics,
marketed under its Spanish namecrema de tortuga.[67]
Turtle plastrons (the part of the shell that covers a tortoise from the bottom) are widely used in
traditional Chinese medicine; according to statistics, Taiwan imports hundreds of tons of plastrons
every year.[68] A popular medicinal preparation based on powdered turtle plastron (and a variety of

Conservation status
In February 2011, the Tortoise and Freshwater Turtle Specialist Group published a report about the top 25 species of turtles most likely to become extinct,
with a further 40 species at very high risk of becoming extinct. This list excludes sea turtles, however, both the leatherback and the Kemp's ridley would
make the top 25 list. The report is due to be updated in four years time allowing to follow the evolution of the list. Between 48 and 54% of all 328 of their
species considered threatened, turtles and tortoises are at a much higher risk of extinction than many other vertebrates. Of the 263 species of freshwater
and terrestrial turtles, 117 species are considered Threatened, 73 are either Endangered or Critically Endangered and 1 is Extinct. Of the 58 species
belonging to the Testudinidae family, 33 species are Threatened, 18 are either Endangered or Critically Endangered, 1 is Extinct in the wild and 7 species
are Extinct. 71% of all tortoise species are either gone or almost gone. Asian species are the most endangered, closely followed by the five endemic
species from Madagascar. Turtles face many threats, including habitat destruction, harvesting for consumption, and the pet trade. The high extinction risk
for Asian species is primarily due to the long-term unsustainable exploitation of turtles and tortoises for consumption and traditional Chinese medicine,
[70]
and to a lesser extent for the international pet trade.

Efforts have been made by Chinese entrepreneurs to satisfy increasing demand for turtle meat as gourmet food and traditional medicine with farmed
turtles, instead of wild-caught ones; according to a study published in 2007, over a thousand turtle farms operated in China.[71][72] Turtle farms in
Oklahoma and Louisiana raise turtles for export to China as well.[72]

Nonetheless, wild turtles continue to be caught and sent to market in large number (as well as to turtle
farms, to be used as breeding stock[71] ), resulting in a situation described by conservationists as "the
Asian turtle crisis".[73] In the words of the biologist George Amato, "the amount and the volume of
captured turtles... vacuumed up entire species from areas in Southeast Asia", even as biologists still did
not know how many distinct turtle species live in the region.[74] About 75% of Asia's 90 tortoise and Turtles on tree branch over a lake in
[72] New Jersey.
freshwater turtle species are estimated to have become threatened.

Harvesting wild turtles is legal in a number of states in the USA.[72] In one of these states, Florida, just a
single seafood company in Fort Lauderdale was reported in 2008 as buying about 5,000 pounds of softshell turtles a week. The harvesters (hunters) are
paid about $2 a pound; some manage to catch as many as 3040 turtles (500 pounds) on a good day. Some of the catch gets to the local restaurants, while
most of it is exported to Asia. The Florida Fish and Wildlife Conservation Commission estimated in 2008 that around 3,000 pounds of softshell turtles
were exported each week viaTampa International Airport.[75]

Nonetheless, the great majority of turtles exported from the USA are farm raised. According to one estimate by the World Chelonian Trust, about 97% of
[72][76] It has been estimated
31.8 million animals harvested in the U.S. over a three-year period (November 4, 2002 November 26, 2005) were exported.
(presumably, over the same 20022005 period) that about 47% of the US turtle exports go to People's Republic of China (predominantly to Hong Kong),
another 20% to Taiwan, and 11% to Mexico.[77][78]

TurtleSAt is a smartphone app that has been developed in Australia in honor of World Turtle Day to help in the conservation of fresh water turtles in
Australia. The app will allow the user to identify turtles with a picture guide and the location of turtles using the phones GPS to record sightings and help
find hidden turtle nesting grounds. The app has been developed because there has been a high per cent of decline of fresh water turtles in Australia due to
foxes, droughts, and urban development. The aim of the app is to reduce the number of foxes and help with geting
tar feral animal control.[79]

Testudinology research organizations

Testudinology, also known as cheloniology, is the zoological study of animals belonging to the order Testudines or Chelonii. A person who specializes in
this field is called a testudinologist. Notable testudinology research or
ganizations include:

Archie Carr Center for Sea Turtle Research, University of Florida

Chelonian Research Institute, founded byPeter Pritchard, in Oviedo, Florida
Dewees Island Turtle Team
East Coast Biologists, Inc.
Ecological Associates, Inc.
Hawaii Preparatory AcademySea Turtle Research Program
Innovation Academy for Engineering, Environmental & Marine Science
Inwater Research Group
Oceana
Padre Island National Seashore, Division of Sea Turtle Science and Recovery
Marine Turtle Research Group of China, a.k.a. the National Huidong Sea Turtle Reserve (Chinese:
)
Israel Nature and Parks Authority- Sea Turtle Rescue Center (Hebrew: )
Red Sea Turtle Project
Chelonian Research Group (not to be confused with the Chelonian Research Institute of the U.S.)
Marine Turtle Research Group

See also
Animal track
Cultural depictions of turtles
Symposium on Turtle Evolution

Notes
 1. Joyce 2007
2. "Testudines" (https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=948936)
. Integrated Taxonomic
Information System.
3. Dubois & Bour 2010
4. Hutchinson 1996
5. "Oxford English Dictionary: Turtle" (http://oxforddictionaries.com/definition/english/turtle)
.
6. Barzyk 1999
7. Vitt, Laurie J.; Caldwell, Janalee P. (25 March 2013). "Herpetology: An Introductory Biology of Amphibians and Reptiles"(https://books.
google.com/books?id=Gay9N_ry79kC&pg=P A523). Academic Press. Retrieved 11 November 2017 via Google Books.
8. (http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=xelw/nh), (http://www.perseus.tufts.edu/
hopper/text?doc=Perseus:text:1999.04.0057:entry=xe/lus). Liddell, Henry George; Scott, Robert; A GreekEnglish Lexiconat the
Perseus Project.
9. Stone 2006, p. 85
10. Brennessel 2006, p. 10
11. testudo (http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0059:entry=testudo). Charlton T. Lewis and Charles
Short. A Latin Dictionary on Perseus Project.
12. Ernst & Lovich 2009, p. 3
13. Fergus 2007
14. Iverson, Kimerling & Kiester 1999
15. Harper, Douglas. "terrapin" (http://www.etymonline.com/?term=terrapin). Online Etymology Dictionary.
16. [1] (http://lema.rae.es/drae/srv/search?val=tortuga)
, Diccionario de la Real Academia Espaola
17. National Geographic 2011.
18. Connor 2009
19. Everhart 2012
20. "Long in the tooth: Genome proves turtles evolvevery slowly"(http://news.ubc.ca/2013/05/30/long-in-the-tooth-genome-proves-turtles
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21. Druzisky, K. A. and Brainerd, E. L. (1 October 2001). "Buccal oscillation and lung ventilation in a semi-aquatic turtle, Platysternon
megacephalum" (http://www.brown.edu/Departments/EEB/brainerd_lab/pdf/Druzisky-Brainerd-2001-Zool.pdf)(PDF). Zoology. 104.
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22. Jackson 2002
23. Pet Education 2012.
24. Ferrara, C. R.; Vogt, R. C.; Sousa-Lima, R. S. (2012). "Turtle Vocalizations as the First Evidence of Posthatching Parental Care in
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"Declared Turtle Trade From the United States: Destinations". World Chelonian Trust. May 2006. Retrieved 15 November 2012. (Major
destinations: 13,625,673 animals to Hong Kong, 1,365,687 to the rest of the PRC, 6,238,300 toaiwan,
T 3,478,275 to Mexico, and
1,527,771 to Japan, 945,257 to Singapore, and 596,966 to Spain.)
"Declared Turtle Trade From the United States: Observations". World Chelonian Trust. May 2006. Retrieved 15 November 2012.
Zardoya, R.; Meyer, A. (1998). "Complete mitochondrial genome suggests diapsid af
finities of turtles". Proceedigns of the National
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doi:10.1073/pnas.95.24.14226. ISSN 0027-8424. PMC 24355 . PMID 9826682.

Further reading
Pritchard, Peter Charles Howard (1979).Encyclopedia of turtles. Neptune, NJ: T.F.H. Publications. ISBN 0-87666-918-6.
External links
Chelonian studbook Collection and display of the weights/sizes of captive turtles
Biogeography and Phylogeny of the Chelonia(taxonomy, maps)
New Scientist article (including video) on how the turtle evolved its shell

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