Turtles, Tortoises, and Terrapins are reptiles of the Order Testudines (all living turtles belong to the crown group Chelonia), most of whose body is shielded by a special bony or cartilagenous shell developed from their ribs. The Order Testudines includes both extant (living) and extinct species, the earliest known turtles being from around 215 million years ago, making turtles one of the oldest reptile groups, and a much more ancient group than lizards and snakes. About 300 species are alive today; some are highly endangered. Turtles are ectothermic.
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Turtles vary widely in size, although marine turtles tend to
be relatively larger animals than their land and freshwater
relatives.
The largest chelonian is a marine turtle, the great leatherback
sea turtle, which reaches a shell length of 200 cm (80 inches)
and can reach a weight of over 900 kg (2,000 lb, or 1 short
ton). Freshwater turtles are generally smaller, but with the
largest species, the Asian softshell turtle Pelochelys cantorii,
a few individuals have been reported to measure up to 200 cm or
80 in (Das, 1991). 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 (31˝ in) and a weight of
about 60 kg (170 lb).
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 that
humans hunted them for food. The only surviving giant tortoises
are on the Seychelles and Galápagos Islands and can grow to over
130 cm (50 in) in length, and weigh about 300 kg (670 lb).
The largest ever chelonian was Archelon 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 in) in length and
weighs about 140 g (5 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 in) in
length.
Turtles are broken down into two groups, according to how they
evolved a solution to the problem of withdrawing their neck into
their shell (something the ancestral Proganochelys could not
do): the Cryptodira, which can draw their neck in while folding
it under their spine; and the Pleurodira, which fold their neck
to the side.
Most turtles that spend most of their life 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 turtles can
hide from predators in shallow water where they lie entirely
submerged except for their eyes and nostrils. Sea turtles
possess glands near their eyes that produce salty tears that rid
their body of excess salt taken in from the water they drink.
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 (UV A) to Red.
Some land turtles have very poor pursuit movement abilities,
which are normally reserved for predators that hunt quick moving
prey. Carnivorous turtles can move their heads quickly to snap
though.
Turtles have a rigid beak. Turtles use their jaws to cut and
chew food. Instead of teeth, 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. Turtles use their tongues to swallow
food, but they can't, unlike most reptiles, stick out their
tongues to catch food.
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 includes 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 a fibrous protein
called 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 shape of the shell gives helpful clues to how the 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 African pancake 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.
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 and these
markings are 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 landbased, 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 shell of a leatherback
turtle is extremely light because they lack scutes and contain
many fontanelles.
As mentioned above, the outer layer of the shell is part of the
skin, each scute (or plate) on the shell corresponding to a
single modified scale. The remainder of the skin is composed of
skin with much smaller scales, similar to the skin of other
reptiles. Turtles and terrapins do not moult their skins all in
one go, as snakes do, but continuously, in small pieces. When
kept in aquaria, small sheets of dead skin can be seen in the
water (often appearing to be a thin piece of plastic) when it
has been sloughed off, often when the animal deliberately rubs
itself against a piece of wood or stone. Tortoises also shed
skin, but a lot of dead skin is allowed to accumulate into thick
knobs and plates that provide protection to parts of the body
outside the shell.
The scutes on the shell are never molted, and, as they
accumulate over time, the shell becomes thicker. By counting the
rings formed by the stack of smaller, older scutes on top of the
larger, newer ones, it is possible to estimate the age of a
turtle, if you know how many scutes are produced in a year. 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.
Terrestrial tortoises have short, sturdy feet. Tortoises are
famous for moving slowly, in part because of their heavy,
cumbersome shell but also because of the relatively inefficient
sprawling gait that they have, with the legs being bent, as with
lizards rather than being straight and directly under the body,
as is the case with mammals.
The 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
simply walk along the bottom of the river or lake. As well as
webbed feet, turtles also have very long claws, used to help
them clamber onto riverbanks and floating logs, upon which they
like to 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, a few turtles, such
as the pig-nose turtles, 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 (see below).
Sea turtles are almost entirely aquatic and instead of feet they
have flippers. 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 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. The back flippers are used to dig
the burrow and then fill it back with sand once the eggs have
been deposited.
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Even though many spend large amounts of their lives underwater,
all turtles and tortoises are air-breathing reptiles, and must
surface at regular intervals to refill their lungs with fresh
air. They can also spend a lot 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
serve to 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.
Turtles lay eggs, like other reptiles, which 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 than do 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.
When the turtles hatch they squirm their way to the surface and
make for the water. There are no known species wherein the
mother cares for the young.
Sea turtles lay their eggs on dry sandy beaches, and are highly
endangered largely as a result of beach development and over
hunting. Immature sea turtles are not raised by either parent.
Turtles can take many years to reach breeding age. Often turtles
only breed every few years or more.
Researchers have recently discovered a turtle’s organs do not
gradually break down or become less efficient over time, unlike
most other animals. It was found that the liver, lungs and
kidneys of a centenarian turtle are virtually indistinguishable
from those of its immature counterpart. This has inspired
genetic researchers to begin examining the turtle genome for
longevity genes.
The first turtles are believed to have existed in the early
Triassic Period of the Mesozoic era, about 200 million years
ago. Their exact ancestry is disputed. It was believed that they
are the only surviving branch of the ancient clade Anapsida,
which includes groups such as procolophonoids, millerettids,
protorothyrids and 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 late Permian period, and the procolophonoids
during the Triassic.
However, it was recently suggested that the anapsid-like turtle
skull may be due to reversion rather than to anapsid descent.
More recent phylogenetic studies with this in mind placed
turtles firmly within diapsids, slightly closer to Squamata than
to Archosauria. All molecular studies have strongly upheld this
new phylogeny, though some place turtles closer to Archosauria.
Re-analysis 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. As of 2003,
the consensus is that Testudines diverged from other diapsids
between 285 and 270 million years ago. Future analyses may show
the turtles to be relatives of the placodonts.
A new phylogenetic analysis agrees with prior analyses nesting
turtles with pareiasaurs within the much larger clade,
Lepidosauromorpha. The closest pareiasaur to turtles appears to
be a rarely-studied form, Stephanospondylus. Indeed turtles are
related to other reptiles without temporal openings.
The earliest known modern turtle is proganochelys, though this
species already had many advanced turtle traits, and thus
probably had many millions of years of preceding "turtle"
evolution and species in its ancestry. It did lack the ability
to pull its head into its shell (and it had a long neck), and
had a long, spiked tail ending in a club, implying an ancestry
occupying a similar niche to the ankylosaurs (though,
presumably, only parallel evolution).
If you own more than one turtle, it is important that they both
receive eat equal amounts of food. Water turtles must be in
water to swallow. They will not eat if they are too cool.
Turtles are generally messy when they eat, so keep their water
clean and free of debris, as small decomposing particles can get
trapped under rocks and logs and contaminate the turtle's
environment. Another good practice is to feed turtles live fish
at least once a week, to make sure they exercise. Turtles like
to hunt, and they dive in after small fish.
Aquatic and terrestrial turtles have similar feeding habits,
though aquatic turtles eat meat even when they are older,
whereas terrestrial turtles have a tendency to eat more
vegetation. Older turtles eat less meat because they require
less protein; therefore, special care must be taken not to
overfeed turtles with protein-rich food. As for vegetation,
turtles like strawberries, raspberries, tomatoes, and some dark
leafy green vegetables, including lettuce, cabbage, alfalfa, and
clover. Turtles also eat animal or high-protein foods such as
rehydrated dried cat, dog, or trout pellets, quality prepared
food such as 'Reptomin', earthworms, snails, crickets,
grasshoppers, slugs, and molluscs. Variety and calcium are key
to a good diet.
Turtles should be fed as much as they will eat within a 5 minute
period. In most cases, feeding 3 times per week, preferably in
the morning, is quite adequate. Daily feeding is hardly ever
required with aquatic turtles, though it may be appropriate with
juveniles. An overfed aquatic turtle will become fatty, obese,
and lethargic, and you will very quickly experience serious tank
hygiene problems - and an almost certain outcome of that will be
a dramatic increase in the incidence of infectious disease.
Have a water dish in the turtle's terrarium or pen, since it
will be a clean source of water for it to drink from, as its
swimming water becomes contaminated with urine and feces.
The water area, which should be about 1/3 to 1/2 of their space,
must be deep enough for the turtles to submerge themselves
completely and to be able to swim freely. A land basking area is
also required. Easy access to the land area must be possible; a
sloping ramp is usually the best approach.
For most North American turtles, the kinds most often kept as
pets, water temperature should be maintained around 82-85
degrees Fahrenheit -- or in the range 23 C (73 F) to 29 C (84
F). Excessively high temperatures (over 32 C; 90 F) or prolonged
periods at too low a temperature (less than 20 C; 68 F) can be
dangerous. The turtle needs to be able to bask, in a crawl-out
space with temperature of about 90 F. Turtles need plenty of UV
light, which result in the production of Vitamin D3, which in
turn promotes the proper utilization of calcium in the turtle
and prevents bone deformities and soft shell problems. UV rays
will not pass through glass.
Plants or leaves are important, as these provide shelter, and
are particularly important for baby turtles which, in the wild,
are considered a prey item. It is natural for them to want to
hide among plants, such as Elodea and Vallisneria. In the
absence of plant cover, baby turtles will become stressed, and
may become ill.
If a turtle seems lethargic and has cloudy skin patches, or
milky eyes, it may be because of poor nutrition or disease. A
veterinarian who knows about reptiles can determine whether the
animal's diet is to blame, or if there are other factors
involved.

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