HP-A New Future

This is just world filler chapter , ignore this chapter 1234556677888877

Turtles are reptiles of the order Testudines characterized by a special bony or cartilaginous shell developed from their ribs and acting as a shield.[3] "Turtle" may refer to the order as a whole (American English) or to fresh-water and sea-dwelling testudines (British English).[4] The order Testudines includes both extant (living) and extinct species. The earliest known members of this group date from the Middle Jurassic,[1] making turtles one of the oldest reptile groups and a more ancient group than snakes or crocodilians. Of the 356 known species[2] alive today, some are highly endangered.[2]

Turtles are ectotherms—animals commonly called cold-blooded—meaning 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.

Differences exist in usage of the common terms turtle, tortoise, and terrapin, depending on the variety of English being used.[5] These terms are common names and do not reflect precise biological or taxonomic distinctions.[6]

Turtle may either refer to the order as a whole, or to particular turtles that make up a form taxon that is not monophyletic, or may be limited to only aquatic species. Tortoise usually refers to any land-dwelling, non-swimming chelonian.[7] Terrapin is used to describe several species of small, edible, hard-shell turtles, typically those found in brackish waters.

In North America, all chelonians are commonly called turtles. Tortoise is used only in reference to fully terrestrial turtles or, more narrowly, only those members of Testudinidae, the family of modern land tortoises.[8][7] Terrapin may refer to small semi-aquatic turtles that live in fresh and brackish water, in particular the diamondback terrapin (Malaclemys terrapin).[9][10][11][12] Although the members of the genus Terrapene dwell mostly on land, they are referred to as box turtles rather than tortoises.[6] The American Society of Ichthyologists and Herpetologists uses "turtle" to describe all species of the order Testudines, regardless of whether they are land-dwelling or sea-dwelling, and uses "tortoise" as a more specific term for slow-moving terrestrial species.[5]

In the United Kingdom, the word turtle is used for water-dwelling species, including ones known in the US as terrapins, but not for terrestrial species, which are known only as tortoises.

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 for turtles, χελώνη chelone; Greek χέλυς chelys "tortoise" is also used in the formation of scientific names of chelonians.[13] Testudines, on the other hand, is based on the Latin word for tortoise, testudo.[14] Terrapin comes from an Algonquian word for turtle.[8][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 is tortuga marina, a freshwater species tortuga de río, and a tortoise tortuga terrestre.[16]

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 tortoises are on the Seychelles and Galápagos Islands and can grow to over 130 cm (51 in) in length, and weigh about 300 kg (660 lb).[18]

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

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.

Turtles are divided into two groups, according to how they retract their necks into their shells (something the ancestral Proganochelys could not do). The mechanism of neck retraction differs phylogenetically: the suborder Pleurodira retracts laterally to the side, anterior to shoulder girdles, while the suborder Cryptodira retracts straight back, between shoulder girdles.[20] These motions are largely due to the morphology and arrangement of cervical vertebrae. Of all recent turtles, the cervical column consists of nine joints and eight vertebrae, which are individually independent.[21] Since these vertebrae are not fused and are rounded, the neck is more flexible, being able to bend in the backwards and sideways directions.[20] The primary function and evolutionary implication of neck retraction is thought to be for feeding rather than protection.[22] Neck retraction and reciprocal extension allows the turtle to reach out further to capture prey while swimming. Neck expansion creates suction when the head is thrust forward and the oropharynx is expanded, and this morphology suggests the retraction function is for feeding purposes as the suction helps catch prey.[22] The protection the shell provides the head when it is retracted is therefore not the main function of retraction, thus is an exaptation.[23] As for the difference between the two methods of retraction, both Pleurodirans and Cryptodirans use the quick extension of the neck as a method of predation, so the difference in retraction mechanism is not due to a difference in ecological niche.[24]

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 semitestacea or "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.[52] 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.

Their exact ancestry has been disputed. It was believed they are the only surviving branch of the ancient evolutionary grade 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.[53]

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.[54][55] All molecular studies have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria,[56] or, more commonly, as a sister group to extant archosaurs,[57][58][59][60] though an analysis conducted by Lyson et al. (2012) recovered turtles as the sister group of lepidosaurs instead.[61] 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.[54][57][62] 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).[63] Similarly, a morphological study conducted by Lyson et al.. (2010) recovered them as anapsids most closely related to Eunotosaurus.[64] A molecular analysis of 248 nuclear genes from 16 vertebrate taxa suggests that turtles are a sister group to birds and crocodiles (the Archosauria).[65] 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, was estimated to have occurred around 157 million years ago.[66][67] The oldest definitive crown-group turtle (member of the modern clade Testudines) is the species Caribemys oxfordiensis from the late Jurassic period (Oxfordian stage).[66] 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 sister group to birds and crocodiles (the Archosauria).[68]

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).[69] 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 turtle evolution; this is further supported by evidence from fossil tracks from the Early Triassic of the United States (Wyoming and Utah) and from the Middle Triassic of Germany, indicating that proto-turtles already existed as early as the Early Triassic.[70] 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 two extant suborders: Cryptodira and 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. Until 3,000 years ago, the family Meiolaniidae was also extant, but this family is outside the Testudines crown group, belonging to Perichelydia.

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 family Testudinidae, 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, and to a lesser extent for the international pet trade.[92]

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.[93][94] Turtle farms in Oklahoma and Louisiana raise turtles for export to China.[94]

Turtles on tree branch over a lake in New Jersey.

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[93]), resulting in a situation described by conservationists as "the Asian turtle crisis".[95] 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.[96] About 75% of Asia's 90 tortoise and freshwater turtle species are estimated to have become threatened.[94]

Harvesting wild turtles is legal in a number of states in the US.[94] 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 30–40 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 via Tampa International Airport.[97]

Nonetheless, the great majority of turtles exported from the US are farm raised. According to one estimate by the World Chelonian Trust, about 97% of 31.8 million animals harvested in the U.S. over a three-year period (November 4, 2002 – November 26, 2005) were exported.[94][98] It has been estimated (presumably, over the same 2002–2005 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.[99][100]

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 targeting feral animal control.[101]

Queensland's shark culling program, which has killed roughly 50,000 sharks since 1962, has also killed thousands of turtles as bycatch.[102][103] Over 5,000 marine turtles have been killed in Queensland's "shark control" program (which uses shark nets and drum lines).[103] The program has also killed 719 loggerhead turtles and 33 hawksbill turtles (hawksbill turtles are critically endangered).[103] New South Wales has a "shark control" program which has killed many turtles: its program uses shark nets,[104][105] in which more than 5,000 marine turtles have been caught.[106]

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. Immersion periods vary between 60 seconds and 1 hour depending on the species.[47] 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.[48]

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 70–120 days to hatch.[citation needed] 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 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.[49]

A group of turtles is known as a bale.[50]

Turtles are known for displaying a wide variety of mating behaviors, however, they are not known for forming pair-bonds or for being part of a social group.[39] Once fertilization has occurred and an offspring has been produced, neither parent will provide care for the offspring once it's hatched.[39] Females generally outnumber males in various turtle species (such as green turtles), and as a result, most males will engage in multiple copulation with multiple partners throughout their lifespan.[40] However, due to the sexual dimorphism present in most turtle species, males must develop different courting strategies or use alternate methods to gain access to a potential mate.[41] Most terrestrial species have males that are larger than females, and fighting between males often determines a hierarchical order in which the higher up the order an individual is, the better the chance is of the individual getting access to a potential mate.[41] For most semi-aquatic species and bottom-walking aquatic species, combat occurs less often. Males belonging to semi-aquatic and bottom-walking species instead often use their larger size advantage to forcibly mate with a female.[41] In fully aquatic species, males are often smaller than females and therefore they cannot use the same strategy as their semi-aquatic relatives, which relies on overpowering the females with strength. Males in this category resort to using courtship displays in an attempt to gain mating access to a female.[41]

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 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 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. 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 of anoxia, it has been shown that the turtle shell both releases carbonate buffers and uptakes lactic acid.[26]

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 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.

Reptiles are tetrapod animals in the class Reptilia, comprising today's turtles, crocodilians, snakes, amphisbaenians, lizards, tuatara, and their extinct relatives. The study of these traditional reptile orders, historically combined with that of modern amphibians, is called herpetology.

Because some reptiles are more closely related to birds than they are to other reptiles (e.g., crocodiles are more closely related to birds than they are to lizards), the traditional groups of "reptiles" listed above do not together constitute a monophyletic grouping or clade (consisting of all descendants of a common ancestor). For this reason, many modern scientists prefer to consider the birds part of Reptilia as well, thereby making Reptilia a monophyletic class, including all living diapsids.[1][2][3][4] The term reptiles is sometimes used as shorthand for 'non-avian Reptilia'.[5][6]

The earliest known proto-reptiles originated around 312 million years ago during the Carboniferous period, having evolved from advanced reptiliomorph tetrapods that became increasingly adapted to life on dry land. Some early examples include the lizard-like Hylonomus and Casineria. In addition to the living reptiles, there are many diverse groups that are now extinct, in some cases due to mass extinction events. In particular, the Cretaceous–Paleogene extinction event wiped out the pterosaurs, plesiosaurs, ornithischians, and sauropods, alongside many species of theropods, crocodyliforms, and squamates (e.g., mosasaurs).

Modern non-avian reptiles inhabit all the continents except Antarctica, although some birds are found on the periphery of Antarctica. Several living subgroups are recognized: Testudines (turtles and tortoises), 350 species;[7] Rhynchocephalia (tuatara from New Zealand), 1 species;[7][8] Squamata (lizards, snakes, and worm lizards), over 10,200 species;[7] and Crocodilia (crocodiles, gharials, caimans, and alligators), 24 species.[7]

Reptiles are tetrapod vertebrates, creatures that either have four limbs or, like snakes, are descended from four-limbed ancestors. Unlike amphibians, reptiles do not have an aquatic larval stage. Most reptiles are oviparous, although several species of squamates are viviparous, as were some extinct aquatic clades[9] – the fetus develops within the mother, contained in a placenta rather than an eggshell. As amniotes, reptile eggs are surrounded by membranes for protection and transport, which adapt them to reproduction on dry land. Many of the viviparous species feed their fetuses through various forms of placenta analogous to those of mammals, with some providing initial care for their hatchlings. Extant reptiles range in size from a tiny gecko, Sphaerodactylus ariasae, which can grow up to 17 mm (0.7 in) to the saltwater crocodile, Crocodylus porosus, which can reach 6 m (19.7 ft) in le In the 13th century the category of reptile was recognized in Europe as consisting of a miscellany of egg-laying creatures, including "snakes, various fantastic monsters, lizards, assorted amphibians, and worms", as recorded by Vincent of Beauvais in his Mirror of Nature.[10] In the 18th century, the reptiles were, from the outset of classification, grouped with the amphibians. Linnaeus, working from species-poor Sweden, where the common adder and grass snake are often found hunting in water, included all reptiles and amphibians in class "III – Amphibia" in his Systema Naturæ.[11] The terms reptile and amphibian were largely interchangeable, reptile (from Latin repere, 'to creep') being preferred by the French.[12] Josephus Nicolaus Laurenti was the first to formally use the term Reptilia for an expanded selection of reptiles and amphibians basically similar to that of Linnaeus.[13] Today, the two groups are still commonly treated under the single heading herpetology.ngth and weigh over 1,000 kg (2,200 lb).

It was not until the beginning of the 19th century that it became clear that reptiles and amphibians are, in fact, quite different animals, and Pierre André Latreille erected the class Batracia (1825) for the latter, dividing the tetrapods into the four familiar classes of reptiles, amphibians, birds, and mammals.[14] The British anatomist Thomas Henry Huxley made Latreille's definition popular and, together with Richard Owen, expanded Reptilia to include the various fossil "antediluvian monsters", including dinosaurs and the mammal-like (synapsid) Dicynodon he helped describe. This was not the only possible classification scheme: In the Hunterian lectures delivered at the Royal College of Surgeons in 1863, Huxley grouped the vertebrates into mammals, sauroids, and ichthyoids (the latter containing the fishes and amphibians). He subsequently proposed the names of Sauropsida and Ichthyopsida for the latter two groups.[15] In 1866, Haeckel demonstrated that vertebrates could be divided based on their reproductive strategies, and that reptiles, birds, and mammals were united by the amniotic egg.

The terms Sauropsida ('lizard faces') and Theropsida ('beast faces') were used again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and their relatives on the one hand (Sauropsida) and mammals and their extinct relatives (Theropsida) on the other. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features, such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, Protosauria ("first lizards") in which he included some animals today considered reptile-like amphibians, as well as early reptiles.[16]

In 1956, D.M.S. Watson observed that the first two groups diverged very early in reptilian history, so he divided Goodrich's Protosauria between them. He also reinterpreted Sauropsida and Theropsida to exclude birds and mammals, respectively. Thus his Sauropsida included Procolophonia, Eosuchia, Millerosauria, Chelonia (turtles), Squamata (lizards and snakes), Rhynchocephalia, Crocodilia, "thecodonts" (paraphyletic basal Archosauria), non-avian dinosaurs, pterosaurs, ichthyosaurs, and sauropterygians.[17]

Modern non-avian reptiles exhibit some form of cold-bloodedness (i.e. some mix of poikilothermy, ectothermy, and bradymetabolism) so that they have limited physiological means of keeping the body temperature constant and often rely on external sources of heat. Due to a less stable core temperature than birds and mammals, reptilian biochemistry requires enzymes capable of maintaining efficiency over a greater range of temperatures than in the case for warm-blooded animals. The optimum body temperature range varies with species, but is typically below that of warm-blooded animals; for many lizards, it falls in the 24°–35 °C (75°–95 °F) range,[74] while extreme heat-adapted species, like the American desert iguana Dipsosaurus dorsalis, can have optimal physiological temperatures in the mammalian range, between 35° and 40 °C (95° and 104 °F).[75] While the optimum temperature is often encountered when the animal is active, the low basal metabolism makes body temperature drop rapidly when the animal is inactive.

As in all animals, reptilian muscle action produces heat. In large reptiles, like leatherback turtles, the low surface-to-volume ratio allows this metabolically produced heat to keep the animals warmer than their environment even though they do not have a warm-blooded metabolism.[76] This form of homeothermy is called gigantothermy; it has been suggested as having been common in large dinosaurs and other extinct large-bodied reptiles.[77][78]

The benefit of a low resting metabolism is that it requires far less fuel to sustain bodily functions. By using temperature variations in their surroundings, or by remaining cold when they do not need to move, reptiles can save considerable amounts of energy compared to endothermic animals of the same size.[79] A crocodile needs from a tenth to a fifth of the food necessary for a lion of the same weight and can live half a year without eating.[80] Lower food requirements and adaptive metabolisms allow reptiles to dominate the animal life in regions where net calorie availability is too low to sustain large-bodied mammals and birds.

It is generally assumed that reptiles are unable to produce the sustained high energy output necessary for long distance chases or flying.[81] Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals.[82] However, investigation of correlations between active capacity and thermophysiology show a weak relationship.[83] Most extant reptiles are carnivores with a sit-and-wait feeding strategy; whether reptiles are cold blooded due to their ecology is not clear. Energetic studies on some reptiles have shown active capacities equal to or greater than similar sized warm-blooded animals.[84]

Reptiles generally reproduce sexually, though some are capable of asexual reproduction. All reproductive activity occurs through the cloaca, the single exit/entrance at the base of the tail where waste is also eliminated. Most reptiles have copulatory organs, which are usually retracted or inverted and stored inside the body. In turtles and crocodilians, the male has a single median penis, while squamates, including snakes and lizards, possess a pair of hemipenes, only one of which is typically used in each session. Tuatara, however, lack copulatory organs, and so the male and female simply press their cloacas together as the male discharges sperm.[123]

Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion, chorion, and allantois are present during embryonic life. The eggshell (1) protects the crocodile embryo (11) and keeps it from drying out, but it is flexible to allow gas exchange. The chorion (6) aids in gas exchange between the inside and outside of the egg. It allows carbon dioxide to exit the egg and oxygen gas to enter the egg. The albumin (9) further protects the embryo and serves as a reservoir for water and protein. The allantois (8) is a sac that collects the metabolic waste produced by the embryo. The amniotic sac (10) contains amniotic fluid (12) which protects and cushions the embryo. The amnion (5) aids in osmoregulation and serves as a saltwater reservoir. The yolk sac (2) surrounding the yolk (3) contains protein and fat rich nutrients that are absorbed by the embryo via vessels (4) that allow the embryo to grow and metabolize. The air space (7) provides the embryo with oxygen while it is hatching. This ensures that the embryo will not suffocate while it is hatching. There are no larval stages of development. Viviparity and ovoviviparity have evolved in many extinct clades of reptiles and in squamates. In the latter group, many species, including all boas and most vipers, utilize this mode of reproduction. The degree of viviparity varies; some species simply retain the eggs until just before hatching, others provide maternal nourishment to supplement the yolk, and yet others lack any yolk and provide all nutrients via a structure similar to the mammalian placenta. The earliest documented case of viviparity in reptiles is the Early Permian mesosaurs,[124] although some individuals or taxa in that clade may also have been oviparous because a putative isolated egg has also been found. Several groups of Mesozoic marine reptiles also exhibited viviparity, such as mosasaurs, ichthyosaurs, and Sauropterygia, a group that include pachypleurosaurs and Plesiosauria.[9]

Asexual reproduction has been identified in squamates in six families of lizards and one snake. In some species of squamates, a population of females is able to produce a unisexual diploid clone of the mother. This form of asexual reproduction, called parthenogenesis, occurs in several species of gecko, and is particularly widespread in the teiids (especially Aspidocelis) and lacertids (Lacerta). In captivity, Komodo dragons (Varanidae) have reproduced by parthenogenesis.

Parthenogenetic species are suspected to occur among chameleons, agamids, xantusiids, and typhlopids.

Some reptiles exhibit temperature-dependent sex determination (TDSD), in which the incubation temperature determines whether a particular egg hatches as male or female. TDSD is most common in turtles and crocodiles, but also occurs in lizards and tuatara.[125] To date, there has been no confirmation of whether TDSD occurs in snakes.[126]

Dinosaurs have been widely depicted in culture since the English palaeontologist Richard Owen coined the name dinosaur in 1842. As soon as 1854, the Crystal Palace Dinosaurs were on display to the public in south London.[143][144] One dinosaur appeared in literature even earlier, as Charles Dickens placed a Megalosaurus in the first chapter of his novel Bleak House in 1852.[145] The dinosaurs featured in books, films, television programs, artwork, and other media have been used for both education and entertainment. The depictions range from the realistic, as in the television documentaries of the 1990s and first decade of the 21st century, or the fantastic, as in the monster movies of the 1950s and 1960s.[144][146][147]

The snake or serpent has played a powerful symbolic role in different cultures. In Egyptian history, the Nile cobra adorned the crown of the pharaoh. It was worshipped as one of the gods and was also used for sinister purposes: murder of an adversary and ritual suicide (Cleopatra). In Greek mythology snakes are associated with deadly antagonists, as a chthonic symbol, roughly translated as earthbound. The nine-headed Lernaean Hydra that Hercules defeated and the three Gorgon sisters are children of Gaia, the earth. Medusa was one of the three Gorgon sisters who Perseus defeated. Medusa is described as a hideous mortal, with snakes instead of hair and the power to turn men to stone with her gaze. After killing her, Perseus gave her head to Athena who fixed it to her shield called the Aegis. The Titans are depicted in art with their legs replaced by bodies of snakes for the same reason: They are children of Gaia, so they are bound to the earth.[148] In Hinduism, snakes are worshipped as gods, with many women pouring milk on snake pits. The cobra is seen on the neck of Shiva, while Vishnu is depicted often as sleeping on a seven-headed snake or within the coils of a serpent. There are temples in India solely for cobras sometimes called Nagraj (King of Snakes), and it is believed that snakes are symbols of fertility. In the annual Hindu festival of Nag Panchami, snakes are venerated and prayed to.[149] In religious terms, the snake and jaguar are arguably the most important animals in ancient Mesoamerica. "In states of ecstasy, lords dance a serpent dance; great descending snakes adorn and support buildings from Chichen Itza to Tenochtitlan, and the Nahuatl word coatl meaning serpent or twin, forms part of primary deities such as Mixcoatl, Quetzalcoatl, and Coatlicue."[150] In Christianity and Judaism, a serpent appears in Genesis to tempt Adam and Eve with the forbidden fruit from the Tree of Knowledge of Good and Evil.[151]

The turtle has a prominent position as a symbol of steadfastness and tranquility in religion, mythology, and folklore from around the world.[152] A tortoise's longevity is suggested by its long lifespan and its shell, which was thought to protect it from any foe.[153] In the cosmological myths of several cultures a World Turtle carries the world upon its back or supports the heavens.[154]

Eagles are large, powerfully built birds of prey, with heavy heads and beaks. Even the smallest eagles, such as the booted eagle (Aquila pennata), which is comparable in size to a common buzzard (Buteo buteo) or red-tailed hawk (B. jamaicensis), have relatively longer and more evenly broad wings, and more direct, faster flight – despite the reduced size of aerodynamic feathers. Most eagles are larger than any other raptors apart from some vultures. The smallest species of eagle is the South Nicobar serpent eagle (Spilornis klossi), at 450 g (1 lb) and 40 cm (16 in). The largest species are discussed below. Like all birds of prey, eagles have very large, hooked beaks for ripping flesh from their prey, strong, muscular legs, and powerful talons. The beak is typically heavier than that of most other birds of prey. Eagles' eyes are extremely powerful. It is estimated that the martial eagle, whose eye is more than twice as long as a human eye, has a visual acuity 3.0 to 3.6 times that of humans. This acuity enables eagles to spot potential prey from a very long distance.[2] This keen eyesight is primarily attributed to their extremely large pupils which ensure minimal diffraction (scattering) of the incoming light. The female of all known species of eagles is larger than the male.[3][4]

Eagles normally build their nests, called eyries, in tall trees or on high cliffs. Many species lay two eggs, but the older, larger chick frequently kills its younger sibling once it has hatched. The dominant chick tends to be a female, as they are bigger than the male. The parents take no action to stop the killing.[5][6]

Due to the size and power of many eagle species, they are ranked at the top of the food chain as apex predators in the avian world. The type of prey varies by genus. The Haliaeetus and Ichthyophaga eagles prefer to capture fish, though the species in the former often capture various animals, especially other water birds, and are powerful kleptoparasites of other birds. The snake and serpent eagles of the genera Circaetus, Terathopius, and Spilornis predominantly prey on the great diversity of snakes found in the tropics of Africa and Asia. The eagles of the genus Aquila are often the top birds of prey in open habitats, taking almost any medium-sized vertebrate they can catch. Where Aquila eagles are absent, other eagles, such as the buteonine black-chested buzzard-eagle of South America, may assume the position of top raptorial predator in open areas. Many other eagles, including the species-rich genus Spizaetus, live predominantly in woodlands and forest. These eagles often target various arboreal or ground-dwelling mammals and birds, which are often unsuspectingly ambushed in such dense, knotty environments. Hunting techniques differ among the species and genera, with some individual eagles having engaged in quite varied techniques based their environment and prey at any given time. Most eagles grab prey without landing and take flight with it, so the prey can be carried to a perch and torn apart.[7]

The bald eagle is noted for having flown with the heaviest load verified to be carried by any flying bird, since one eagle flew with a 6.8 kg (15 lb) mule deer fawn.[8] However, a few eagles may target prey considerably heavier than themselves; such prey is too heavy to fly with, thus it is either eaten at the site of the kill or taken in pieces back to a perch or nest. Golden and crowned eagles have killed ungulates weighing up to 30 kg (66 lb) and a martial eagle even killed a 37 kg (82 lb) duiker, 7–8 times heavier than the preying eagle.[7][9] Authors on birds David Allen Sibley, Pete Dunne, and Clay Sutton described the behavioral difference between hunting eagles and other birds of prey thus (in this case the bald and golden eagles as compared to other North American raptors):[10]

They have at least one singular characteristic. It has been observed that most birds of prey look back over their shoulders before striking prey (or shortly thereafter); predation is after all a two-edged sword. All hawks seem to have this habit, from the smallest kestrel to the largest Ferruginous – but not the Eagles.

Among the eagles are some of the largest birds of prey: only the condors and some of the Old World vultures are markedly larger. It is regularly debated which should be considered the largest species of eagle. They could be measured variously in total length, body mass, or wingspan. Different lifestyle needs among various eagles result in variable measurements from species to species. For example, many forest-dwelling eagles, including the very large harpy eagle, have relatively short wingspans, a feature necessary for being able to maneuver in quick, short bursts through densely forested habitats.[7] Eagles in the genus Aquila, though found almost strictly in open country, are superlative soarers, and have relatively long wings for their size.[7]

These lists of the top five eagles are based on weight, length, and wingspan, respectively. Unless otherwise noted by reference, the figures listed are the median reported for each measurement in the guide Raptors of the World[11] in which only measurements that could be personally verified by the authors were listed.[7]

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They could be measured variously in total length, body mass, or wingspan. Different lifestyle needs among various eagles result in variable measurements from species to species. For example, many forest-dwelling eagles, including the very large harpy eagle, have relatively short wingspans, a feature necessary for being able to maneuver in quick, short bursts through densely forested habitats Turtles are divided into two groups, according to how they retract their necks into their shells (something the ancestral Proganochelys could not do). The mechanism of neck retraction differs phylogenetically: the suborder Pleurodira retracts laterally to the side, anterior to shoulder girdles, while the suborder Cryptodira retracts straight back, between shoulder girdles.[20] These motions are largely due to the morphology and arrangement of cervical vertebrae. Of all recent turtles, the cervical column consists of nine joints and eight vertebrae, which are individually independent.[21] Since these vertebrae are not fused and are rounded, the neck is more flexible, being able to bend in the backwards and sideways directions.[20] The primary function and evolutionary implication of neck retraction is thought to be for feeding rather than protection.[22]

Turtles are reptiles of the order Testudines characterized by a special bony or cartilaginous shell developed from their ribs and acting as a shield.[3] "Turtle" may refer to the order as a whole (American English) or to fresh-water and sea-dwelling testudines (British English).[4] The order Testudines includes both extant (living) and extinct species. The earliest known members of this group date from the Middle Jurassic,[1] making turtles one of the oldest reptile groups and a more ancient group than snakes or crocodilians. Of the 356 known species[2] alive today, some are highly endangered.[2]

Turtles are ectotherms—animals commonly called cold-blooded—meaning 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.

Differences exist in usage of the common terms turtle, tortoise, and terrapin, depending on the variety of English being used.[5] These terms are common names and do not reflect precise biological or taxonomic distinctions.[6]

Turtle may either refer to the order as a whole, or to particular turtles that make up a form taxon that is not monophyletic, or may be limited to only aquatic species. Tortoise usually refers to any land-dwelling, non-swimming chelonian.[7] Terrapin is used to describe several species of small, edible, hard-shell turtles, typically those found in brackish waters.

In North America, all chelonians are commonly called turtles. Tortoise is used only in reference to fully terrestrial turtles or, more narrowly, only those members of Testudinidae, the family of modern land tortoises.[8][7] Terrapin may refer to small semi-aquatic turtles that live in fresh and brackish water, in particular the diamondback terrapin (Malaclemys terrapin).[9][10][11][12] Although the members of the genus Terrapene dwell mostly on land, they are referred to as box turtles rather than tortoises.[6] The American Society of Ichthyologists and Herpetologists uses "turtle" to describe all species of the order Testudines, regardless of whether they are land-dwelling or sea-dwelling, and uses "tortoise" as a more specific term for slow-moving terrestrial species.[5]

In the United Kingdom, the word turtle is used for water-dwelling species, including ones known in the US as terrapins, but not for terrestrial species, which are known only as tortoises.

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 for turtles, χελώνη chelone; Greek χέλυς chelys "tortoise" is also used in the formation of scientific names of chelonians.[13] Testudines, on the other hand, is based on the Latin word for tortoise, testudo.[14] Terrapin comes from an Algonquian word for turtle.[8][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 is tortuga marina, a freshwater species tortuga de río, and a tortoise tortuga terrestre.[16]

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 tortoises are on the Seychelles and Galápagos Islands and can grow to over 130 cm (51 in) in length, and weigh about 300 kg (660 lb).[18]

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

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.

Turtles are divided into two groups, according to how they retract their necks into their shells (something the ancestral Proganochelys could not do). The mechanism of neck retraction differs phylogenetically: the suborder Pleurodira retracts laterally to the side, anterior to shoulder girdles, while the suborder Cryptodira retracts straight back, between shoulder girdles.[20] These motions are largely due to the morphology and arrangement of cervical vertebrae. Of all recent turtles, the cervical column consists of nine joints and eight vertebrae, which are individually independent.[21] Since these vertebrae are not fused and are rounded, the neck is more flexible, being able to bend in the backwards and sideways directions.[20] The primary function and evolutionary implication of neck retraction is thought to be for feeding rather than protection.[22] Neck retraction and reciprocal extension allows the turtle to reach out further to capture prey while swimming. Neck expansion creates suction when the head is thrust forward and the oropharynx is expanded, and this morphology suggests the retraction function is for feeding purposes as the suction helps catch prey.[22] The protection the shell provides the head when it is retracted is therefore not the main function of retraction, thus is an exaptation.[23] As for the difference between the two methods of retraction, both Pleurodirans and Cryptodirans use the quick extension of the neck as a method of predation, so the difference in retraction mechanism is not due to a difference in ecological niche.[24]

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 semitestacea or "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.[52] 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.

Their exact ancestry has been disputed. It was believed they are the only surviving branch of the ancient evolutionary grade 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.[53]

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.[54][55] All molecular studies have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria,[56] or, more commonly, as a sister group to extant archosaurs,[57][58][59][60] though an analysis conducted by Lyson et al. (2012) recovered turtles as the sister group of lepidosaurs instead.[61] 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.[54][57][62] 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).[63] Similarly, a morphological study conducted by Lyson et al.. (2010) recovered them as anapsids most closely related to Eunotosaurus.[64] A molecular analysis of 248 nuclear genes from 16 vertebrate taxa suggests that turtles are a sister group to birds and crocodiles (the Archosauria).[65] 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, was estimated to have occurred around 157 million years ago.[66][67] The oldest definitive crown-group turtle (member of the modern clade Testudines) is the species Caribemys oxfordiensis from the late Jurassic period (Oxfordian stage).[66] 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 sister group to birds and crocodiles (the Archosauria).[68]

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).[69] 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 turtle evolution; this is further supported by evidence from fossil tracks from the Early Triassic of the United States (Wyoming and Utah) and from the Middle Triassic of Germany, indicating that proto-turtles already existed as early as the Early Triassic.[70] 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 two extant suborders: Cryptodira and 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. Until 3,000 years ago, the family Meiolaniidae was also extant, but this family is outside the Testudines crown group, belonging to Perichelydia.

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 family Testudinidae, 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, and to a lesser extent for the international pet trade.[92]

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.[93][94] Turtle farms in Oklahoma and Louisiana raise turtles for export to China.[94]

Turtles on tree branch over a lake in New Jersey.

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[93]), resulting in a situation described by conservationists as "the Asian turtle crisis".[95] 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.[96] About 75% of Asia's 90 tortoise and freshwater turtle species are estimated to have become threatened.[94]

Harvesting wild turtles is legal in a number of states in the US.[94] 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 30–40 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 via Tampa International Airport.[97]

Nonetheless, the great majority of turtles exported from the US are farm raised. According to one estimate by the World Chelonian Trust, about 97% of 31.8 million animals harvested in the U.S. over a three-year period (November 4, 2002 – November 26, 2005) were exported.[94][98] It has been estimated (presumably, over the same 2002–2005 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.[99][100]

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 targeting feral animal control.[101]

Queensland's shark culling program, which has killed roughly 50,000 sharks since 1962, has also killed thousands of turtles as bycatch.[102][103] Over 5,000 marine turtles have been killed in Queensland's "shark control" program (which uses shark nets and drum lines).[103] The program has also killed 719 loggerhead turtles and 33 hawksbill turtles (hawksbill turtles are critically endangered).[103] New South Wales has a "shark control" program which has killed many turtles: its program uses shark nets,[104][105] in which more than 5,000 marine turtles have been caught.[106]

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. Immersion periods vary between 60 seconds and 1 hour depending on the species.[47] 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.[48]

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 70–120 days to hatch.[citation needed] 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 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.[49]

A group of turtles is known as a bale.[50]

Turtles are known for displaying a wide variety of mating behaviors, however, they are not known for forming pair-bonds or for being part of a social group.[39] Once fertilization has occurred and an offspring has been produced, neither parent will provide care for the offspring once it's hatched.[39] Females generally outnumber males in various turtle species (such as green turtles), and as a result, most males will engage in multiple copulation with multiple partners throughout their lifespan.[40] However, due to the sexual dimorphism present in most turtle species, males must develop different courting strategies or use alternate methods to gain access to a potential mate.[41] Most terrestrial species have males that are larger than females, and fighting between males often determines a hierarchical order in which the higher up the order an individual is, the better the chance is of the individual getting access to a potential mate.[41] For most semi-aquatic species and bottom-walking aquatic species, combat occurs less often. Males belonging to semi-aquatic and bottom-walking species instead often use their larger size advantage to forcibly mate with a female.[41] In fully aquatic species, males are often smaller than females and therefore they cannot use the same strategy as their semi-aquatic relatives, which relies on overpowering the females with strength. Males in this category resort to using courtship displays in an attempt to gain mating access to a female.[41]

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 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 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. 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 of anoxia, it has been shown that the turtle shell both releases carbonate buffers and uptakes lactic acid.[26]

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 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.

Reptiles are tetrapod animals in the class Reptilia, comprising today's turtles, crocodilians, snakes, amphisbaenians, lizards, tuatara, and their extinct relatives. The study of these traditional reptile orders, historically combined with that of modern amphibians, is called herpetology.

Because some reptiles are more closely related to birds than they are to other reptiles (e.g., crocodiles are more closely related to birds than they are to lizards), the traditional groups of "reptiles" listed above do not together constitute a monophyletic grouping or clade (consisting of all descendants of a common ancestor). For this reason, many modern scientists prefer to consider the birds part of Reptilia as well, thereby making Reptilia a monophyletic class, including all living diapsids.[1][2][3][4] The term reptiles is sometimes used as shorthand for 'non-avian Reptilia'.[5][6]

The earliest known proto-reptiles originated around 312 million years ago during the Carboniferous period, having evolved from advanced reptiliomorph tetrapods that became increasingly adapted to life on dry land. Some early examples include the lizard-like Hylonomus and Casineria. In addition to the living reptiles, there are many diverse groups that are now extinct, in some cases due to mass extinction events. In particular, the Cretaceous–Paleogene extinction event wiped out the pterosaurs, plesiosaurs, ornithischians, and sauropods, alongside many species of theropods, crocodyliforms, and squamates (e.g., mosasaurs).

Modern non-avian reptiles inhabit all the continents except Antarctica, although some birds are found on the periphery of Antarctica. Several living subgroups are recognized: Testudines (turtles and tortoises), 350 species;[7] Rhynchocephalia (tuatara from New Zealand), 1 species;[7][8] Squamata (lizards, snakes, and worm lizards), over 10,200 species;[7] and Crocodilia (crocodiles, gharials, caimans, and alligators), 24 species.[7]

Reptiles are tetrapod vertebrates, creatures that either have four limbs or, like snakes, are descended from four-limbed ancestors. Unlike amphibians, reptiles do not have an aquatic larval stage. Most reptiles are oviparous, although several species of squamates are viviparous, as were some extinct aquatic clades[9] – the fetus develops within the mother, contained in a placenta rather than an eggshell. As amniotes, reptile eggs are surrounded by membranes for protection and transport, which adapt them to reproduction on dry land. Many of the viviparous species feed their fetuses through various forms of placenta analogous to those of mammals, with some providing initial care for their hatchlings. Extant reptiles range in size from a tiny gecko, Sphaerodactylus ariasae, which can grow up to 17 mm (0.7 in) to the saltwater crocodile, Crocodylus porosus, which can reach 6 m (19.7 ft) in le In the 13th century the category of reptile was recognized in Europe as consisting of a miscellany of egg-laying creatures, including "snakes, various fantastic monsters, lizards, assorted amphibians, and worms", as recorded by Vincent of Beauvais in his Mirror of Nature.[10] In the 18th century, the reptiles were, from the outset of classification, grouped with the amphibians. Linnaeus, working from species-poor Sweden, where the common adder and grass snake are often found hunting in water, included all reptiles and amphibians in class "III – Amphibia" in his Systema Naturæ.[11] The terms reptile and amphibian were largely interchangeable, reptile (from Latin repere, 'to creep') being preferred by the French.[12] Josephus Nicolaus Laurenti was the first to formally use the term Reptilia for an expanded selection of reptiles and amphibians basically similar to that of Linnaeus.[13] Today, the two groups are still commonly treated under the single heading herpetology.ngth and weigh over 1,000 kg (2,200 lb).

It was not until the beginning of the 19th century that it became clear that reptiles and amphibians are, in fact, quite different animals, and Pierre André Latreille erected the class Batracia (1825) for the latter, dividing the tetrapods into the four familiar classes of reptiles, amphibians, birds, and mammals.[14] The British anatomist Thomas Henry Huxley made Latreille's definition popular and, together with Richard Owen, expanded Reptilia to include the various fossil "antediluvian monsters", including dinosaurs and the mammal-like (synapsid) Dicynodon he helped describe. This was not the only possible classification scheme: In the Hunterian lectures delivered at the Royal College of Surgeons in 1863, Huxley grouped the vertebrates into mammals, sauroids, and ichthyoids (the latter containing the fishes and amphibians). He subsequently proposed the names of Sauropsida and Ichthyopsida for the latter two groups.[15] In 1866, Haeckel demonstrated that vertebrates could be divided based on their reproductive strategies, and that reptiles, birds, and mammals were united by the amniotic egg.

The terms Sauropsida ('lizard faces') and Theropsida ('beast faces') were used again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and their relatives on the one hand (Sauropsida) and mammals and their extinct relatives (Theropsida) on the other. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features, such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, Protosauria ("first lizards") in which he included some animals today considered reptile-like amphibians, as well as early reptiles.[16]

In 1956, D.M.S. Watson observed that the first two groups diverged very early in reptilian history, so he divided Goodrich's Protosauria between them. He also reinterpreted Sauropsida and Theropsida to exclude birds and mammals, respectively. Thus his Sauropsida included Procolophonia, Eosuchia, Millerosauria, Chelonia (turtles), Squamata (lizards and snakes), Rhynchocephalia, Crocodilia, "thecodonts" (paraphyletic basal Archosauria), non-avian dinosaurs, pterosaurs, ichthyosaurs, and sauropterygians.[17]

Modern non-avian reptiles exhibit some form of cold-bloodedness (i.e. some mix of poikilothermy, ectothermy, and bradymetabolism) so that they have limited physiological means of keeping the body temperature constant and often rely on external sources of heat. Due to a less stable core temperature than birds and mammals, reptilian biochemistry requires enzymes capable of maintaining efficiency over a greater range of temperatures than in the case for warm-blooded animals. The optimum body temperature range varies with species, but is typically below that of warm-blooded animals; for many lizards, it falls in the 24°–35 °C (75°–95 °F) range,[74] while extreme heat-adapted species, like the American desert iguana Dipsosaurus dorsalis, can have optimal physiological temperatures in the mammalian range, between 35° and 40 °C (95° and 104 °F).[75] While the optimum temperature is often encountered when the animal is active, the low basal metabolism makes body temperature drop rapidly when the animal is inactive.

As in all animals, reptilian muscle action produces heat. In large reptiles, like leatherback turtles, the low surface-to-volume ratio allows this metabolically produced heat to keep the animals warmer than their environment even though they do not have a warm-blooded metabolism.[76] This form of homeothermy is called gigantothermy; it has been suggested as having been common in large dinosaurs and other extinct large-bodied reptiles.[77][78]

The benefit of a low resting metabolism is that it requires far less fuel to sustain bodily functions. By using temperature variations in their surroundings, or by remaining cold when they do not need to move, reptiles can save considerable amounts of energy compared to endothermic animals of the same size.[79] A crocodile needs from a tenth to a fifth of the food necessary for a lion of the same weight and can live half a year without eating.[80] Lower food requirements and adaptive metabolisms allow reptiles to dominate the animal life in regions where net calorie availability is too low to sustain large-bodied mammals and birds.

It is generally assumed that reptiles are unable to produce the sustained high energy output necessary for long distance chases or flying.[81] Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals.[82] However, investigation of correlations between active capacity and thermophysiology show a weak relationship.[83] Most extant reptiles are carnivores with a sit-and-wait feeding strategy; whether reptiles are cold blooded due to their ecology is not clear. Energetic studies on some reptiles have shown active capacities equal to or greater than similar sized warm-blooded animals.[84]

Reptiles generally reproduce sexually, though some are capable of asexual reproduction. All reproductive activity occurs through the cloaca, the single exit/entrance at the base of the tail where waste is also eliminated. Most reptiles have copulatory organs, which are usually retracted or inverted and stored inside the body. In turtles and crocodilians, the male has a single median penis, while squamates, including snakes and lizards, possess a pair of hemipenes, only one of which is typically used in each session. Tuatara, however, lack copulatory organs, and so the male and female simply press their cloacas together as the male discharges sperm.[123]

Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion, chorion, and allantois are present during embryonic life. The eggshell (1) protects the crocodile embryo (11) and keeps it from drying out, but it is flexible to allow gas exchange. The chorion (6) aids in gas exchange between the inside and outside of the egg. It allows carbon dioxide to exit the egg and oxygen gas to enter the egg. The albumin (9) further protects the embryo and serves as a reservoir for water and protein. The allantois (8) is a sac that collects the metabolic waste produced by the embryo. The amniotic sac (10) contains amniotic fluid (12) which protects and cushions the embryo. The amnion (5) aids in osmoregulation and serves as a saltwater reservoir. The yolk sac (2) surrounding the yolk (3) contains protein and fat rich nutrients that are absorbed by the embryo via vessels (4) that allow the embryo to grow and metabolize. The air space (7) provides the embryo with oxygen while it is hatching. This ensures that the embryo will not suffocate while it is hatching. There are no larval stages of development. Viviparity and ovoviviparity have evolved in many extinct clades of reptiles and in squamates. In the latter group, many species, including all boas and most vipers, utilize this mode of reproduction. The degree of viviparity varies; some species simply retain the eggs until just before hatching, others provide maternal nourishment to supplement the yolk, and yet others lack any yolk and provide all nutrients via a structure similar to the mammalian placenta. The earliest documented case of viviparity in reptiles is the Early Permian mesosaurs,[124] although some individuals or taxa in that clade may also have been oviparous because a putative isolated egg has also been found. Several groups of Mesozoic marine reptiles also exhibited viviparity, such as mosasaurs, ichthyosaurs, and Sauropterygia, a group that include pachypleurosaurs and Plesiosauria.[9]

Asexual reproduction has been identified in squamates in six families of lizards and one snake. In some species of squamates, a population of females is able to produce a unisexual diploid clone of the mother. This form of asexual reproduction, called parthenogenesis, occurs in several species of gecko, and is particularly widespread in the teiids (especially Aspidocelis) and lacertids (Lacerta). In captivity, Komodo dragons (Varanidae) have reproduced by parthenogenesis.

Parthenogenetic species are suspected to occur among chameleons, agamids, xantusiids, and typhlopids.

Some reptiles exhibit temperature-dependent sex determination (TDSD), in which the incubation temperature determines whether a particular egg hatches as male or female. TDSD is most common in turtles and crocodiles, but also occurs in lizards and tuatara.[125] To date, there has been no confirmation of whether TDSD occurs in snakes.[126]

Dinosaurs have been widely depicted in culture since the English palaeontologist Richard Owen coined the name dinosaur in 1842. As soon as 1854, the Crystal Palace Dinosaurs were on display to the public in south London.[143][144] One dinosaur appeared in literature even earlier, as Charles Dickens placed a Megalosaurus in the first chapter of his novel Bleak House in 1852.[145] The dinosaurs featured in books, films, television programs, artwork, and other media have been used for both education and entertainment. The depictions range from the realistic, as in the television documentaries of the 1990s and first decade of the 21st century, or the fantastic, as in the monster movies of the 1950s and 1960s.[144][146][147]

The snake or serpent has played a powerful symbolic role in different cultures. In Egyptian history, the Nile cobra adorned the crown of the pharaoh. It was worshipped as one of the gods and was also used for sinister purposes: murder of an adversary and ritual suicide (Cleopatra). In Greek mythology snakes are associated with deadly antagonists, as a chthonic symbol, roughly translated as earthbound. The nine-headed Lernaean Hydra that Hercules defeated and the three Gorgon sisters are children of Gaia, the earth. Medusa was one of the three Gorgon sisters who Perseus defeated. Medusa is described as a hideous mortal, with snakes instead of hair and the power to turn men to stone with her gaze. After killing her, Perseus gave her head to Athena who fixed it to her shield called the Aegis. The Titans are depicted in art with their legs replaced by bodies of snakes for the same reason: They are children of Gaia, so they are bound to the earth.[148] In Hinduism, snakes are worshipped as gods, with many women pouring milk on snake pits. The cobra is seen on the neck of Shiva, while Vishnu is depicted often as sleeping on a seven-headed snake or within the coils of a serpent. There are temples in India solely for cobras sometimes called Nagraj (King of Snakes), and it is believed that snakes are symbols of fertility. In the annual Hindu festival of Nag Panchami, snakes are venerated and prayed to.[149] In religious terms, the snake and jaguar are arguably the most important animals in ancient Mesoamerica. "In states of ecstasy, lords dance a serpent dance; great descending snakes adorn and support buildings from Chichen Itza to Tenochtitlan, and the Nahuatl word coatl meaning serpent or twin, forms part of primary deities such as Mixcoatl, Quetzalcoatl, and Coatlicue."[150] In Christianity and Judaism, a serpent appears in Genesis to tempt Adam and Eve with the forbidden fruit from the Tree of Knowledge of Good and Evil.[151]

The turtle has a prominent position as a symbol of steadfastness and tranquility in religion, mythology, and folklore from around the world.[152] A tortoise's longevity is suggested by its long lifespan and its shell, which was thought to protect it from any foe.[153] In the cosmological myths of several cultures a World Turtle carries the world upon its back or supports the heavens.[154]

Eagles are large, powerfully built birds of prey, with heavy heads and beaks. Even the smallest eagles, such as the booted eagle (Aquila pennata), which is comparable in size to a common buzzard (Buteo buteo) or red-tailed hawk (B. jamaicensis), have relatively longer and more evenly broad wings, and more direct, faster flight – despite the reduced size of aerodynamic feathers. Most eagles are larger than any other raptors apart from some vultures. The smallest species of eagle is the South Nicobar serpent eagle (Spilornis klossi), at 450 g (1 lb) and 40 cm (16 in). The largest species are discussed below. Like all birds of prey, eagles have very large, hooked beaks for ripping flesh from their prey, strong, muscular legs, and powerful talons. The beak is typically heavier than that of most other birds of prey. Eagles' eyes are extremely powerful. It is estimated that the martial eagle, whose eye is more than twice as long as a human eye, has a visual acuity 3.0 to 3.6 times that of humans. This acuity enables eagles to spot potential prey from a very long distance.[2] This keen eyesight is primarily attributed to their extremely large pupils which ensure minimal diffraction (scattering) of the incoming light. The female of all known species of eagles is larger than the male.[3][4]

Eagles normally build their nests, called eyries, in tall trees or on high cliffs. Many species lay two eggs, but the older, larger chick frequently kills its younger sibling once it has hatched. The dominant chick tends to be a female, as they are bigger than the male. The parents take no action to stop the killing.[5][6]

Due to the size and power of many eagle species, they are ranked at the top of the food chain as apex predators in the avian world. The type of prey varies by genus. The Haliaeetus and Ichthyophaga eagles prefer to capture fish, though the species in the former often capture various animals, especially other water birds, and are powerful kleptoparasites of other birds. The snake and serpent eagles of the genera Circaetus, Terathopius, and Spilornis predominantly prey on the great diversity of snakes found in the tropics of Africa and Asia. The eagles of the genus Aquila are often the top birds of prey in open habitats, taking almost any medium-sized vertebrate they can catch. Where Aquila eagles are absent, other eagles, such as the buteonine black-chested buzzard-eagle of South America, may assume the position of top raptorial predator in open areas. Many other eagles, including the species-rich genus Spizaetus, live predominantly in woodlands and forest. These eagles often target various arboreal or ground-dwelling mammals and birds, which are often unsuspectingly ambushed in such dense, knotty environments. Hunting techniques differ among the species and genera, with some individual eagles having engaged in quite varied techniques based their environment and prey at any given time. Most eagles grab prey without landing and take flight with it, so the prey can be carried to a perch and torn apart.[7]

The bald eagle is noted for having flown with the heaviest load verified to be carried by any flying bird, since one eagle flew with a 6.8 kg (15 lb) mule deer fawn.[8] However, a few eagles may target prey considerably heavier than themselves; such prey is too heavy to fly with, thus it is either eaten at the site of the kill or taken in pieces back to a perch or nest. Golden and crowned eagles have killed ungulates weighing up to 30 kg (66 lb) and a martial eagle even killed a 37 kg (82 lb) duiker, 7–8 times heavier than the preying eagle.[7][9] Authors on birds David Allen Sibley, Pete Dunne, and Clay Sutton described the behavioral difference between hunting eagles and other birds of prey thus (in this case the bald and golden eagles as compared to other North American raptors):[10]

They have at least one singular characteristic. It has been observed that most birds of prey look back over their shoulders before striking prey (or shortly thereafter); predation is after all a two-edged sword. All hawks seem to have this habit, from the smallest kestrel to the largest Ferruginous – but not the Eagles.

Among the eagles are some of the largest birds of prey: only the condors and some of the Old World vultures are markedly larger. It is regularly debated which should be considered the largest species of eagle. They could be measured variously in total length, body mass, or wingspan. Different lifestyle needs among various eagles result in variable measurements from species to species. For example, many forest-dwelling eagles, including the very large harpy eagle, have relatively short wingspans, a feature necessary for being able to maneuver in quick, short bursts through densely forested habitats.[7] Eagles in the genus Aquila, though found almost strictly in open country, are superlative soarers, and have relatively long wings for their size.[7]

These lists of the top five eagles are based on weight, length, and wingspan, respectively. Unless otherwise noted by reference, the figures listed are the median reported for each measurement in the guide Raptors of the World[11] in which only measurements that could be personally verified by the authors were listed.[7]

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They could be measured variously in total length, body mass, or wingspan. Different lifestyle needs among various eagles result in variable measurements from species to species. For example, many forest-dwelling eagles, including the very large harpy eagle, have relatively short wingspans, a feature necessary for being able to maneuver in quick, short bursts through densely forested habitats Turtles are divided into two groups, according to how they retract their necks into their shells (something the ancestral Proganochelys could not do). The mechanism of neck retraction differs phylogenetically: the suborder Pleurodira retracts laterally to the side, anterior to shoulder girdles, while the suborder Cryptodira retracts straight back, between shoulder girdles.[20] These motions are largely due to the morphology and arrangement of cervical vertebrae. Of all recent turtles, the cervical column consists of nine joints and eight vertebrae, which are individually independent.[21] Since these vertebrae are not fused and are rounded, the neck is more flexible, being able to bend in the backwards and sideways directions.[20] The primary function and evolutionary implication of neck retraction is thought to be for feeding rather than protection.[22]