Size The largest
living species of turtle (and fourth-largest
reptile) is the
leatherback turtle, which can reach over in length and weigh over . The largest known turtle was
Archelon ischyros, a
Late Cretaceous sea turtle up to long, wide between the tips of the front flippers, and estimated to have weighed over . The smallest living turtle is
Chersobius signatus of South Africa, measuring no more than in length and weighing .
Shell of a tortoise skeleton|alt=Photograph of one half of a tortoise skeleton, cut in half vertically showing the vertebrae following curving along the carapace The shell of a turtle is unique among
vertebrates and serves to protect the animal and provide shelter from the elements. It is primarily made of 50–60 bones and consists of two parts: the domed, dorsal (back)
carapace and the flatter, ventral (belly)
plastron. They are connected by lateral (side) extensions of the plastron. The shoulder girdle in turtles is made up of two bones, the scapula and the
coracoid. Both the shoulder and pelvic girdles of turtles are located within the shell and hence are effectively within the rib cage. The trunk ribs grow over the shoulder girdle during development. Turtle scutes are usually structured like
mosaic tiles, but some species, like the
hawksbill sea turtle, have overlapping scutes on the carapace. The shapes of turtle shells vary with the adaptations of the individual species, and
sometimes with sex. Land-dwelling turtles are more dome-shaped, which appears to make them more resistant to being crushed by large animals. Aquatic turtles have flatter, smoother shells that allow them to cut through the water. Sea turtles in particular have streamlined shells that reduce
drag and increase stability in the open ocean. Some turtle species have pointy or spiked shells that provide extra
protection from predators and
camouflage against the leafy ground. The lumps of a tortoise shell can tilt its body when it gets flipped over, allowing it to flip back. In male tortoises, the tip of the plastron is thickened and used for butting and ramming during combat. Shells vary in flexibility. Some species, such as
box turtles, lack the lateral extensions and instead have the carapace bones fully fused or
ankylosed together. Several species have hinges on their shells, usually on the plastron, which allow them to expand and contract.
Softshell turtles have rubbery edges, due to the loss of bones. The leatherback turtle has hardly any bones in its shell, but has thick
connective tissue and an outer layer of leathery skin.
Head and neck The turtle's skull is unique among living
amniotes (which includes reptiles, birds and mammals); it is solid and rigid with no openings for muscle attachment (
temporal fenestrae). Muscles instead attach to recesses in the back of the skull. Turtle skulls vary in shape, from the long and narrow skulls of softshells to the broad and flattened skull of the
mata mata. Some turtle species have developed large and thick heads, allowing for greater muscle mass and stronger bites. In several species, some individuals can develop larger heads and thicker muscles than others, a phenomenon known as 'megacephaly'. This is likely due to differences in diet. Turtles that are carnivorous or
durophagous (eating hard-shelled animals) have the most powerful bites. For example, the durophagous
Mesoclemmys nasuta has a bite force of . Species that are
insectivorous,
piscivorous (fish-eating), or
omnivorous have lower bite forces. Living turtles lack teeth but have beaks made of
keratin sheaths along the edges of the jaws. The necks of turtles are highly flexible, possibly to compensate for their rigid shells. Some species, like sea turtles, have short necks while others, such as
snake-necked turtles, have long ones. Despite this, all turtle species have eight
neck vertebrae, a consistency not found in other reptiles but similar to mammals. Some snake-necked turtles have both long necks and large heads, limiting their ability to lift them when not in water. Some turtles have folded structures in the
larynx or
glottis that vibrate to produce sound. Other species have
elastin-rich
vocal cords.
Limbs and locomotion Due to their heavy shells, turtles are slow-moving on land. A
desert tortoise moves at only . By contrast, sea turtles can swim at . Compared to other reptiles, turtles tend to have reduced tails, but these vary in both length and thickness among species and between sexes.
Snapping turtles and the
big-headed turtle have longer tails; the latter uses it for balance while climbing. The
cloaca is found underneath and at the base, and the tail itself houses the reproductive organs. Hence, males have longer tails to contain the penis. In sea turtles, the tail is longer and more
prehensile in males, who use it to grasp mates. Several turtle species have spines on their tails.
Senses has an exceptional seven types of color-detecting cells in its eyes. Sea turtles orient themselves on land by night, using visual features detected in dim light. They can use their eyes in clear surface water, muddy coasts, the darkness of the deep ocean, and also above water. Unlike in terrestrial turtles, the
cornea (the curved surface that lets light into the eye) does not help to focus light on the retina, so focusing underwater is handled entirely by the lens, behind the cornea. The cone cells contain oil droplets placed to shift perception toward the red part of the spectrum, improving color discrimination. Visual acuity, studied in hatchlings, is highest in a horizontal band with retinal cells packed about twice as densely as elsewhere. This gives the best vision along the visual horizon. Sea turtles do not appear to use
polarized light for orientation as many other animals do. The deep-diving leatherback turtle lacks specific adaptations to low light, such as large eyes, large lenses, or a reflective
tapetum. It may rely on seeing the
bioluminescence of prey when hunting in deep water. Turtles have no ear openings; the
eardrum is covered with scales and encircled by a bony
otic capsule, which is absent in other reptiles. Their hearing thresholds are high in comparison to other reptiles, reaching up to 500
Hz in air, but underwater they are more attuned to lower frequencies. The
loggerhead sea turtle has been shown experimentally to respond to low sounds, with maximal sensitivity between 100 and 400 Hz. Turtles have
olfactory (smell) and
vomeronasal receptors along the nasal cavity, the latter of which are used to detect chemical signals. Experiments on green sea turtles showed they could learn to respond to a selection of different odorant chemicals such as
triethylamine and
cinnamaldehyde, which were detected by olfaction in the nose. Such signals could be used in navigation.
Breathing nose-breathing at river surface|alt=photo of a river turtle with only its nose above water The rigid shell of turtles is not capable of expanding and making room for the lungs, as in other amniotes, so they have had to evolve special adaptations for respiration. The lungs of turtles are attached directly to the carapace above while below, connective tissue attaches them to the organs. They have multiple lateral (side) and medial (middle) chambers (the numbers of which vary between species) and one terminal (end) chamber. The lungs are ventilated using specific groups of abdominal muscles attached to the organs that pull and push on them. During exhalation, the contraction of the
transversus abdominis muscle propels the organs into the lungs and expels air. Conversely, during inhalation, the relaxing and flattening of the
oblique abdominis muscle pulls the transversus back down, allowing air back into the lungs. Some species can
respire through the cloaca, which contains large sacs that are lined with many finger-like projections that take up dissolved
oxygen from the water.
Circulation emerging from period of
brumation, in which it buried itself in mud. Turtles have multiple circulatory and physiological adaptations to enable them to go long periods without breathing. Turtles are capable of enduring periods of anaerobic respiration longer than many other vertebrates. This process breaks down sugars incompletely to
lactic acid, rather than all the way to
carbon dioxide and water as in
aerobic (oxygen-based) respiration.
Osmoregulation In sea turtles, the bladder is one unit and in most freshwater turtles, it is double-lobed. Sea turtle bladders are connected to two small accessory bladders, located at the sides to the neck of the urinary bladder and above the
pubis. Arid-living tortoises have bladders that serve as reserves of water, storing up to 20% of their body weight in fluids. The fluids are normally low in
solutes, but higher during droughts when the reptile gains
potassium salts from its plant diet. The bladder stores these salts until the tortoise finds fresh drinking water. To regulate the amount of salt in their bodies, sea turtles and the
brackish-living
diamondback terrapin secrete excess salt in a thick sticky substance from their
tear glands. Because of this, sea turtles may appear to be "crying" when on land.
Thermoregulation s, regulate their temperature by basking in the sun.|alt=cooter turtles basking in sunshine near their pond Turtles, like other reptiles, have a limited ability to
regulate their body temperature. This ability varies between species, and with body size. Small pond turtles regulate their temperature by crawling out of the water and basking in the sun, while small terrestrial turtles move between sunny and shady places to adjust their temperature. Large species, both terrestrial and marine, have sufficient mass to give them substantial
thermal inertia, meaning that they heat up or cool down over many hours. The
Aldabra giant tortoise weighs up to some and is able to allow its temperature to rise to some on a hot day, and to fall naturally to around by night. Some giant tortoises seek out shade to avoid overheating on sunny days. On
Grand Terre Island, food is scarce inland, shade is scarce near the coast, and the tortoises compete for space under the few trees on hot days. Large males may push smaller females out of the shade, and some then overheat and die. == Behavior==