turtle

turtle, a reptile of the order Chelonia, with strong, beaked, toothless jaws and, usually, an armorlike shell. The shell normally consists of bony plates overlaid with horny shields. The upper portion, or carapace, covers the turtle's back and sides, and the lower portion, or plastron, covers the belly; the two parts are joined at the sides. Exceptions are the rare plateless turtles of New Guinea and the marine leatherback turtle, which is encased in a thick, ossified skin resembling a carapace. When startled, most turtles withdraw their heads straight back into their shells, the neck folding into an S-shaped curve. However, in the side-necked turtles of the Southern Hemisphere, the head moves sideways and tucks next to the shoulder.

Turtles are found throughout most of the temperate and tropical world and in the open ocean; of the 270 known species, 42% are rare or threatened with extinction. Many turtles and their eggs are valued as food. Edible species include several marine turtles, the green turtle (traditional ingredient of turtle soup), the diamondback terrapin, and the soft-shelled turtles. Catching females when they lay eggs on land has contributed to a serious decline in many species, since it can take 10 to 30 years for some turtles to reach sexual maturity.

Different types of turtle are variously adapted to living on land, in freshwater, or in the ocean, but all turtles breathe by means of lungs, and all lay eggs on land. The land-living species, especially those of the family Testudinidae, are commonly called tortoises. The name terrapin is generally applied to large freshwater or brackish water species, especially those used for food. Turtle species are either herbivorous or carnivorous but rarely both. They range in length from a few inches to over 6 ft (2 m), most being between 5 in. and 15 in. (13-38 cm) long. Many specimens have survived more than 50 years in captivity; one giant tortoise is known to have lived for 176 years, and another is believed to have lived about 250 years.

Turtles existed 200 million years ago, at the time of the earliest dinosaurs; these early land-dwelling turtles could not retract their necks. By 120 million years ago some turtles had adapted to an aquatic life, although a 220-million-year-old ancestor of turtles that had only a bony breastplate may have been aquatic. Many of the living families of turtles existed in the Cretaceous period and have undergone very little change since then. On the basis of morphological (body structure) evidence, turtles were thought to be the oldest surviving group of reptiles. However, molecular studies comparing genes in different reptile groups indicate that turtles, along with crocodiles, are the most modern of reptiles.

Types of Turtles

Turtles are classified in 12 families. The Northern Hemisphere's largest family is that of common freshwater turtles (Emydidae), which includes about a third of all turtle species and is abundant in S and E Asia, E North America, and Central America. Members of this group have webbed feet; many spend most of the time in freshwater ponds or marshes; some live in brackish estuaries. They include such well-known North American turtles as the pond turtles (including the spotted, wood, and Muhlenberg's turtles), the painted turtle, the sliders, the diamondback terrapin, and the Blanding's turtle. The box turtle, which is primarily terrestrial, belongs to this family. Land tortoises (Testudinidae) form the second largest family. Tortoises have high-domed shells, move on club-shaped feet, are vegetarian, and live in warm regions throughout the world. The musk turtles and mud turtles (family Kinosternidae) are common small turtles of the E United States, and are found only in the Americas. The soft-shelled turtles (family Trionychidae) are flat-bodied, carnivorous freshwater turtles of the Northern Hemisphere, with a leathery covering instead of horny shields on their shells. The snapping turtle family (Chelydridae) is a North American group that includes the common snapper and the alligator snapper.

Marine turtles are classified in two families. The family Chelonidae includes five sea turtle species of tropical and subtropical distribution: the green turtle, the loggerhead, the hawksbill (or tortoiseshell turtle), the Kemp's ridley, and the olive ridley. The family Dermochelidae includes only one species, the leatherback, or leatherneck, largest and heaviest of all turtles, weighing as much as 1100 lbs (500 kg). Marine turtles lack toes, and their legs are oarlike, allowing speeds of nearly 20 mph (32 kph) in the water. With the exception of the loggerhead, all are endangered, either by pollution with plastic debris, which some turtles eat by mistake, or by commercial fishing, especially shrimp trawling. Commercial trade in all endangered sea turtles is banned; however, many wild turtles are skinned for leather and tortoiseshell ornaments, or taken for food.

Classification

Turtles are classified in the phylum Chordata, subphylum Vertebrata, class Reptilia, order Chelonia.

Species (Streptopelia turtur, family Columbidae) of migratory European pigeon that winters in northern Africa. It is about 11 in. (28 cm) long and has a reddish brown body, blue-gray head, and white-tipped tail. A ground feeder, it eats prodigious amounts of small seeds. The name is applied to other temperate and tropical Old World Streptopelia species of slim, fast-flying game birds. The ringed turtledove, or ringdove, has feral populations in California and Florida, U.S.; the laughing and spotted doves have also been introduced outside their native habitats.

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Any of approximately 300 species (order Testudines) of reptiles that have a bony shell overlaid with horny shields and are found in most parts of the world. Turtles have existed for 200 million years, making them the oldest of all surviving reptiles. Most species are aquatic or semiaquatic; some are terrestrial. Turtles eat plants, animals, or both. They are toothless, have a horny beak, and range from less than 4 in. (10 cm) to more than 5 ft (1.5 m) long. They have sturdy, sprawling limbs with short feet or paddlelike flippers (marine turtles). Some species bend the neck sideways, but most pull the head and neck backward into the shell. Almost half of the known turtle species are rare, threatened, or endangered. Seealso box turtle; painted turtle; snapping turtle; softshell turtle; terrapin; tortoise.

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Any of more than 30 species (family Trionychidae) of swift-moving carnivorous turtles found in North America, Africa, and Asia. Thriving in fresh waters that have soft sandy or muddy bottoms, they often lie buried in mud or sand and occasionally emerge to bask in the sun. They have a pancakelike leathery shell, webbed feet, a long neck, and an elongated snout. Aggressive when captured, they can deliver rapid, vicious bites. North American species have a grayish or brown shell, 14–18 in. (35–45 cm) long. Two Old World species grow to 24 in. (60 cm) or more.

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Either of two species (family Chelydridae) of edible, omnivorous, freshwater turtles found in North and Central America. They are tan to black and have a rough upper shell, a small cross-shaped lower shell, a long tail, and a large head with hooked jaws. Known for their fierceness, they lunge at aggressors and prey and bite them with their powerful jaws. The common snapping turtle (Chelydra serpentina) has a shell 8–12 in. (20–30 cm) long and weighs 10–35 lbs (4.5–16 kg). The alligator snapping turtle (Macroclemys temmincki), the largest freshwater turtle in the U.S., has a shell 16–28 in. (40–70 cm) long and weighs 40–155 lbs (18–70 kg). It lies quietly on the bottom of slow moving bodies of water, luring fishes by means of a wormlike appendage on the floor of its open mouth.

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Species (Chrysemys picta, family Emydidae) of brightly marked North American turtle found from southern Canada to northern Mexico. It has a smooth shell, 4–7 in. (10–18 cm) long, with red and yellow markings on its relatively flat, black or greenish brown upper shell. It usually lives in quiet, shallow bodies of fresh water, especially those with thickly planted mud bottoms, feeding on plants, small animals, and some carrion. It often basks in large groups on logs and other objects. In many areas it hibernates.

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Any of several species of terrestrial turtles found in the U.S. and Mexico (genus Terrapene) or the subtropical and tropical regions of Asia (genus Cuora). Box turtles have a high, rounded upper shell (carapace) that grows to a maximum length of about 7 in. (18 cm). The lower shell (plastron) is hinged across the centre and can be drawn very tightly against the carapace to form a protective “box” that completely encloses the turtle's soft parts. Box turtles feed on earthworms, insects, mushrooms, and berries and are often kept as pets.

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(born circa 1752, near Fort Wayne, Ind.—died July 14, 1812, Fort Wayne, Ind., U.S.) American Indian leader. Chief of the Miami tribe, he led raids on settlements in the Northwest Territory in the early 1790s. Defeated by Gen. Anthony Wayne at the Battle of Fallen Timbers (1794), he was obliged to sign the Treaty of Greenville (1795), which ceded to the U.S. much of Ohio and parts of Illinois, Indiana, and Michigan. He then advocated peace and prevented the Miami from joining the Shawnee confederacy of Tecumseh.

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Turtles 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 cartilaginous shell developed from their ribs. The Order Testudines includes both extant (living) and extinct species. The earliest known turtles date from 215 million years ago, making turtles one of the oldest reptile groups and a more ancient group than lizards and snakes. About 300 species are alive today, and some are highly endangered.

Turtles cannot breathe in water, but they can hold their breath for various periods of time.

Like other reptiles, turtles are "cold-blooded" (or poikilothermic — "of varying temperature). Like other amniotes (reptiles, dinosaurs, birds, and mammals), they breathe air and do not lay eggs underwater, although many species live in or around water. The largest turtles are aquatic.

Anatomy and morphology

The largest chelonian is 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 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 (31½ 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.

Neck folding

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 contracting it under their spine; and the Pleurodira, which contract their neck to the side.

Head

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, but carnivorous turtles are able to move their heads quickly to snap.

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 cannot, unlike most reptiles, stick out their tongues to catch food.

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 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 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 shell of a leatherback turtle is extremely light because they lack scutes and contain many fontanelles.

Skin and molting

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 molt 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) having been sloughed off 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.

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.

Limbs

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, 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 (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 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. .

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 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 "papilla", 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.

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 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 head toward the water. There are no known species in which the mother cares for the 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 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.

Taxonomy

Turtles are divided into three suborders, one of which, the Paracryptodira, is extinct. The two extant suborders are 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 withdraw their heads into their shells. This smaller group consists primarily of various freshwater turtles.

Evolutionary history

The first proto-turtles are believed to have existed in the early Triassic Period of the Mesozoic era, about 200 million years ago, and their shell, which has remained a remarkably stable body plan, is thought to have evolved from rows of protective armour plates that gradually fused to one another and eventually to the reptile's ribs and vertebrae as well, creating a complete shell that offered protection at every stage of its evolution. By the late Jurassic, turtles had radiated widely, and their fossil history becomes easier to read.

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 procolophonids, 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. 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. As of 2003, the consensus is that Testudines diverged from other diapsids between 200 and 279 million years ago.

The earliest known fully-shelled turtle is the late-Triassic 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 only through parallel evolution).

Turtle, tortoise, or terrapin?

As pets

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 (or terrapin).

Due to the ease of contracting salmonella through casual contact with turtles, the FDA established a regulation in 1975 to discontinue the sale of turtles under 4 inches. It is illegal in every state in the U.S. for anyone to sell any turtles under 4 inches long. Many stores and flea markets still sell small turtles due to a loophole in the FDA regulation which allows turtles under 4 inches to be sold for "educational" purposes.

Some states have other laws and regulations regarding possession of Red-eared Sliders (abbreviated as "RES") as pets because they are looked upon as invasive species or pests where they are not native but have been introduced through the pet trade. As of July 1, 2007 it is illegal in Florida to sell any wild type RES. Unusual color varieties such as albino and pastel RES, which are derived from captive breeding, are still allowed for sale.

As food

The flesh of turtles was, or still is, considered a delicacy in a number of cultures. Turtle soup has been a prized dish in Anglo-American cuisine, and still remains so in some parts of the Far East.

Guilinggao jelly was a Chinese medicine preparation containing powdered shell of a certain turtle species; these days, though, it is typically made with only herbal ingredients.

See also

Further reading

• Iskandar, DT (2000). Turtles and Crocodiles of Insular Southeast Asia and New Guinea. ITB, Bandung.
• Pritchard, Pether C H (1979). Encyclopedia of Turtles. T.F.H. Publications.

References

External links

• UC Berkeley Museum of Paleontology
• Chelonian studbook Collection and display of the weights/sizes of captive turtles
• John M. Legler & Arthur Georges, Biogeography and Phylogeny of the Chelonia (taxonomy, maps)
• The word 'turtle' in many different languages