Reptiles are animals in the (Linnaean) class Reptilia. They are characterized by breathing air, laying shelled eggs (except for some vipers and constrictor snakes that give live birth), and having skin covered in scales and/or scutes. Reptiles are classically viewed as having a "cold-blooded" metabolism. They are tetrapods (either having four limbs or being descended from four-limbed ancestors). Modern reptiles inhabit every continent with the exception of Antarctica, and four living orders are currently recognized:

Crocodilia (crocodiles, gavials, caimans, and alligators): 23 species

Sphenodontia (tuataras from New Zealand): 2 species

Squamata (lizards, snakes, and worm lizards): approximately 7,900 species

Testudines (turtles, terrapins and tortoises): approximately 300 species

Unlike amphibians, reptiles do not have an aquatic larval stage. As a rule, reptiles are oviparous (egg-laying), although certain species of squamates are capable of giving live birth. This is achieved by either ovoviviparity (egg retention) or viviparity (birth of offspring without the development of calcified eggs). 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 1.7 cm (0.6 in) to the saltwater crocodile, ''Crocodylus porosus'', which may reach 6 m in length and weigh over 1,000 kg.

The study of reptiles and amphibians is called herpetology.

Classification

History of classification

Linnaeus and 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. The terms "reptile" and "amphibian" were largely interchangeable, "reptile" (from Latin ''repere'', "to creep") being preferred by the French. 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. Today, it is still common to treat the two groups under the same heading as herptiles.

"Antediluvian monsters"

Not until the beginning of the 19th century did it become 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.

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 two.

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") which included some Paleozoic amphibians as well as early reptiles.

In 1956 D.M.S. Watson observed that the first two groups diverged very early in reptilian history, and 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.

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. By the end of the 19th century, the class Reptilia had come to include all the amniotes except birds and mammals. Thus reptiles were defined as the set of animals that includes the extant crocodiles, alligators, tuatara, lizards, snakes, amphisbaenians, and turtles, as well as fossil groups like dinosaurs, synapsids and the primitive pareiasaurs. This is still the common definition of the term.

Skull openings in 20th century classification

The synapsid/sauropsid division supplemented, but was never as popular during the 20th century as a Linneaean approach splitting the reptiles into four subclasses based on the number and position of ''temporal fenestrae'', openings in the sides of the skull behind the eyes. This classification was initiated by Henry Fairfield Osborn and elaborated an made popular by Romer's classic ''Vertebrate Paleontology''. Those four subclasses were: Anapsida – no fenestrae - cotylosaurs and Chelonia (turtles and relatives)

Synapsida – one low fenestra - pelycosaurs and therapsids (the 'mammal-like reptiles')

Euryapsida – one high fenestra (above the postorbital and squamosal) - protorosaurs (small, early lizard-like reptiles) and the marine sauropterygians and ichthyosaurs, the latter called Parapsida in Osborn's work.

Diapsida – two fenestrae - most reptiles, including lizards, snakes, crocodilians, dinosaurs and pterosaurs

The composition of Euryapsida was uncertain. Ichthyosaurs were at times considered to have arisen independently of the other euryapsids, and given the older name Parapsida. Parapsida was later discarded as a group for the most part (ichthyosaurs being classified as ''incertae sedis'' or with Euryapsida). However, the scheme with four (or three if Euryapsida is sunk into Diapsida) subclasses remained more or less universal for non-specialist work throughout the 20th century, and has only been challenged with the rising popularity of phylogenetic nomenclature.

Phylogenetics and modern definition

By the 21st century, most vertebrate paleontologists had adopted phylogenetic taxonomy, in which all groups are defined in such a way as to be monophyletic; that is, groups include all descendants of a particular ancestor. The reptiles as historically defined would be paraphyletic, since they exclude both birds and mammals, although these also evolved from the original reptiles. Colin Tudge wrote:

Mammals are a clade, and therefore the cladists are happy to acknowledge the traditional taxon Mammalia; and birds, too, are a clade, universally ascribed to the formal taxon Aves. Mammalia and Aves are, in fact, subclades within the grand clade of the Amniota. But the traditional class Reptilia is not a clade. It is just a section of the clade Amniota: the section that is left after the Mammalia and Aves have been hived off. It cannot be defined by synapomorphies, as is the proper way. It is instead defined by a combination of the features it has and the features it lacks: reptiles are the amniotes that lack fur or feathers. At best, the cladists suggest, we could say that the traditional Reptilia are 'non-avian, non-mammalian amniotes'.

Despite the early proposals for replacing the paraphyletic Reptilia for a monophyletic Sauropsida, that term was never adopted widely or, when it was, applied consistently. When Sauropsida was used, it often had the same content or even the same definition as Reptilia. In 1988 Jacques Gauthier proposed a cladistic definition of Reptilia as a monophyletic node-based crown group containing turtles, lizards and snakes, crocodilians, and birds, their common ancestor and all its descendants. The proposed suffered under the actual relationship of turtles to other reptiles not being fully understood. A variety of other definitions were proposed by other scientists in the years following Gauthier's paper. The first which attempted to adhere to the standards of the PhyloCode was published by Modesto and Anderson in 2004. They reviewed the many previous definitions, and proposed a modified definition which they intended to retain most traditional content of the group while keeping it stable and monophyletic. They defined Reptilia as all amniotes closer to ''Lacerta agilis'' and ''Crocodylus niloticus'' than to ''Homo sapiens''. This stem-based definition is equivalent to that of Sauropsida, which Modesto and Anderson synonymized with Reptilia, since the latter is more well known and more frequently used, despite their definition including birds.

Taxonomy

Classification to order level, after Benton, 2005.

Series Amniota

* Class Synapsida

** Order Pelycosauria*

** Order Therapsida

*** Class Mammalia

* Class Sauropsida

** Subclass Anapsida

*** Order Testudines (turtles)

*** A series of unassigned anapsid families, corresponding to Captorhinida, Mesosauria and Procolophonomorpha

** Subclass Diapsida

*** Order Araeoscelidia

*** Order Younginiformes

*** Infraclass Ichthyosauria

*** Infraclass Lepidosauromorpha

**** Superorder Sauropterygia

***** Order Placodontia

***** Order Nothosauroidea

***** Order Plesiosauria

**** Superorder Lepidosauria

***** Order Sphenodontia (tuatara)

***** Order Squamata (lizards & snakes)

*** Infraclass Archosauromorpha

**** Order Prolacertiformes

**** Division Archosauria

***** Subdivision Crurotarsi

****** Superorder Crocodylomorpha

******* Order Crocodilia

******* Order Phytosauria

******* Order Rauisuchia

******* Order Rhynchosauria

***** Subdivision Avemetatarsalia

****** Infradivision Ornithodira

******* Order Pterosauria

******* Superorder Dinosauria

******** Order Saurischia

********* Class Aves

******** Order Ornithischia

Phylogeny

The cladogram presented here illustrates the "family tree" of reptiles, and follows a simplified version of the relationships found by Laurin and Gauthier (1996), presented as part of the Tree of Life Web Project.

|3=?Testudines (turtles, tortoises, and terrapins) }} }} }} }} |label2=Romeriida |2= }} }} }} }} }} }} }} }}

Evolutionary history

Origin of the reptiles

The origin of the reptiles lies about 320–310 million years ago, in the steaming swamps of the late Carboniferous period, when the first reptiles evolved from advanced reptiliomorph labyrinthodonts. The oldest known animal that may have been an amniote, i.e. a primitive reptile rather than an advanced amphibian is ''Casineria''. A series of footprints from the fossil strata of Nova Scotia, dated to 315 million years show typical reptilian toes and imprints of scales. The tracks are attributed to ''Hylonomus'', the oldest unquestionable reptile known. It was a small, lizard-like animal, about 20 to 30 cm (8–12 in) long, with numerous sharp teeth indicating an insectivorous diet. Other examples include ''Westlothiana'' (for the moment considered a reptiliomorph amphibian rather than a true amniote) and ''Paleothyris'', both of similar build and presumably similar habit.

Rise of the reptiles

Earliest reptiles were largely overshadowed by bigger labyrinthodont amphibians such as ''Cochleosaurus'', and remained a small, inconspicuous part of the fauna until the Carboniferous Rainforest Collapse. This sudden collapse affected several large groups, amphibians were particularly devastated, while reptiles fared better, being ecologically adapted to the drier conditions that followed. Amphibians must return to water to lay eggs; in contrast, reptiles - whose eggs have a hard shell and can therefore be laid on land - were better adapted to the new conditions. Reptiles acquired new niches at a faster rate than before the collapse and at a much faster rate than amphibians. They acquired new feeding strategies including herbivory and carnivory, previously only having been insectivores and piscivores. From this point forward, reptiles dominated communities and had a greater diversity than amphibians, setting the stage for the Mesozoic (known as the Age of Reptiles). One of the best known early reptiles is ''Mesosaurus'', a genus from the early Permian that had returned to water, feeding on fish.

Anapsids, synapsids, diapsids and sauropsids

The first reptiles were anapsids, having a solid skull with holes for only nose, eyes, spinal cord, etc. Very soon after the first reptiles appeared, they split into two branches. One branch, Synapsida (including both "mammal-like reptiles" and modern, extant mammals such as humans), had one opening in the skull roof behind each eye; the other branch, Diapsida, possessed a hole in their skulls behind each eye, along with a second hole located higher on the skull. The function of the holes in both groups was to lighten the skull and give room for the jaw muscles to move, allowing for a more powerful bite. The rationale for this classification was disputed, with some arguing that turtles are diapsids that reverted to this primitive state in order to improve their armor (see Parareptilia). All molecular studies have strongly upheld the placement of turtles within diapsids, most commonly as a sister group to extant archosaurs.

Permian reptiles

With the close of the Carboniferous, reptiles became the dominant tetrapod fauna. While the terrestrial reptiliomorph labyrinthodonts still existed, the synapsids evolved the first truly terrestrial megafauna (giant animals) in the form of pelycosaurs such as ''Edaphosaurus'' and the carnivorous ''Dimetrodon''. In the mid-Permian period the climate turned dryer, resulting in a change of fauna: The primitive pelycosaurs were replaced by the more advanced therapsids.

The anapsid reptiles, whose massive skull roofs had no postorbital holes, continued and flourished throughout the Permian. The pareiasaurs reached giant proportions in the late Permian, eventually disappearing at the close of the period (the turtles being possible survivors).

Early in the period, the diapsid reptiles split into two main lineages, the archosaurs (forefathers of crocodiles and dinosaurs) and the lepidosaurs (predecessors of modern snakes, lizards, and tuataras). Both groups remained lizard-like and relatively small and inconspicuous during the Permian.

The Mesozoic era, the "Age of Reptiles"

The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic extinction event), a prolonged event due to the accumalation of at lest two distinct extinction pulses. Most of the earlier anapsid/synapsid megafauna disappeared, being replaced by the archosauromorph diapsids. The archosaurs were characterized by elongated hind legs and an erect pose, the early forms looking somewhat like long-legged crocodiles. The archosaurs became the dominant group during the Triassic period, though it took 30 million years before their diversity was as great as the animals that lived in the Permian. Archosaurs developing into the well-known dinosaurs and pterosaurs, as well as crocodiles and phytosaurs. Some of the dinosaurs developed into the largest land animals ever to have lived, making the Mesozoic era popularly known as the "Age of Reptiles". The dinosaurs also developed smaller forms, including the feather-bearing smaller theropods. In the mid-Jurassic period, these gave rise to the first birds.

The lepidosauromorph diapsids may have been ancestral to the sea reptiles. These reptiles developed into the sauropterygians in the early Triassic and the ichthyosaurs during the Middle Triassic. The mosasaurs also evolved in the Mesozoic era, emerging during the Cretaceous period.

The therapsids came under increasing pressure from the dinosaurs in the early Mesozoic and developed into increasingly smaller and more nocturnal forms, the first mammals being the only survivors of the line by the late Jurassic.

Demise of the dinosaurs

The close of the Cretaceous period saw the demise of the Mesozoic era reptilian megafauna (see the Cretaceous–Tertiary extinction event). Of the large marine reptiles, only sea turtles are left, and, of the dinosaurs, only the small feathered theropods survived in the form of birds. The end of the “Age of Reptiles” led into the “Age of Mammals”. Mammals and birds filled the empty niches left behind by reptiles and while reptile diversification slowed bird and mammal diversification took an exponential turn.

Reptile diversification continued throughout the Cenozoic, with squamates undergoing a greater radiation than they did in the Mesozoic. Today squamates make up the majority of extant reptiles today (over 90%). There are approximately 8,700 extant species of reptiles, compared with 5,400 species of mammals.

Systems

Circulation

Most reptiles have a three-chambered heart consisting of two atria, one variably partitioned ventricle, and two aortas that lead to the systemic circulation. The degree of mixing of oxygenated and deoxygenated blood in the three-chambered heart varies depending on the species and physiological state. Under different conditions, deoxygenated blood can be shunted back to the body or oxygenated blood can be shunted back to the lungs. This variation in blood flow has been hypothesized to allow more effective thermoregulation and longer diving times for aquatic species, but has not been shown to be a fitness advantage.

There are some exceptions to the general physiology. For instance, crocodilians have an anatomically four-chambered heart, but also have two systemic aortas and are therefore capable of bypassing only their pulmonary circulation. Also, some snake and lizard species (e.g., pythons and monitor lizards) have three-chambered hearts that become functionally four-chambered hearts during contraction. This is made possible by a muscular ridge that subdivides the ventricle during ventricular diastole and completely divides it during ventricular systole. Because of this ridge, some of these squamates are capable of producing ventricular pressure differentials that are equivalent to those seen in mammalian and avian hearts.

Metabolism

All reptiles exhibit some form of cold-bloodedness (i.e. some mix of poikilothermy, ectothermy, and bradymetabolism). This means that most reptiles 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 warm-blooded animals. The optimum body temperature range varies with species, but is typically below that of warm-blooded animals, in the 24°–35°C range for many lizards, 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. 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.

Like 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, despite not having a warm-blooded metabolism. This form of homeothermy is called gigantothermy, and has been suggested as having been common in large dinosaurs and other extinct large-bodied reptiles.

The benefits 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 endotherm animals of the same size. A crocodile need from a fifth to a tenth of the food necessary for a lion of the same weight, and can live half a year without eating. Lower food requirements and adaptive metabolisms allow reptiles to dominate the animal life in regions where net calorie production 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. Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals. However, investigation of correlations between active capacity and thermophysiology show a weak relationship. Most extant reptiles are carnivores with a sit-and-wait feeding strategy, and whether reptiles are cold blooded due to their ecology, or if their metabolism is a result of their ecology is not clear. Energetic studies on some reptiles have shown active capacities equal to, or greater than similar sized warm-blooded animals.

Respiration

Reptilian lungs

All reptiles breathe using lungs. Aquatic turtles have developed more permeable skin, and some species have modified their cloaca to increase the area for gas exchange. Even with these adaptations, breathing is never fully accomplished without lungs. Lung ventilation is accomplished differently in each main reptile group. In squamates, the lungs are ventilated almost exclusively by the axial musculature. This is also the same musculature that is used during locomotion. Because of this constraint, most squamates are forced to hold their breath during intense runs. Some, however, have found a way around it. Varanids, and a few other lizard species, employ buccal pumping as a complement to their normal "axial breathing." This allows the animals to completely fill their lungs during intense locomotion, and thus remain aerobically active for a long time. Tegu lizards are known to possess a proto-diaphragm, which separates the pulmonary cavity from the visceral cavity. While not actually capable of movement, it does allow for greater lung inflation, by taking the weight of the viscera off the lungs. Crocodilians actually have a muscular diaphragm that is analogous to the mammalian diaphragm. The difference is that the muscles for the crocodilian diaphragm pull the pubis (part of the pelvis, which is movable in crocodilians) back, which brings the liver down, thus freeing space for the lungs to expand. This type of diaphragmatic setup has been referred to as the "hepatic piston."

Turtles and tortoises

How turtles and tortoises breathe has been the subject of much study. To date, only a few species have been studied thoroughly enough to get an idea of how turtles do it. The results indicate that turtles and tortoises have found a variety of solutions to this problem.

The difficulty is that most turtle shells are rigid and do not allow for the type of expansion and contraction that other amniotes use to ventilate their lungs. Some turtles such as the Indian flapshell (''Lissemys punctata'') have a sheet of muscle that envelops the lungs. When it contracts, the turtle can exhale. When at rest, the turtle can retract the limbs into the body cavity and force air out of the lungs. When the turtle protracts its limbs, the pressure inside the lungs is reduced, and the turtle can suck air in. Turtle lungs are attached to the inside of the top of the shell (carapace), with the bottom of the lungs attached (via connective tissue) to the rest of the viscera. By using a series of special muscles (roughly equivalent to a diaphragm), turtles are capable of pushing their viscera up and down, resulting in effective respiration, since many of these muscles have attachment points in conjunction with their forelimbs (indeed, many of the muscles expand into the limb pockets during contraction).

Breathing during locomotion has been studied in three species, and they show different patterns. Adult female green sea turtles do not breathe as they crutch along their nesting beaches. They hold their breath during terrestrial locomotion and breathe in bouts as they rest. North American box turtles breathe continuously during locomotion, and the ventilation cycle is not coordinated with the limb movements. This is because they use their abdominal muscles to breathe during locomotion. The last species to have been studied is the red-eared slider, which also breathes during locomotion, but takes smaller breaths during locomotion than during small pauses between locomotor bouts, indicating that there may be mechanical interference between the limb movements and the breathing apparatus. Box turtles have also been observed to breathe while completely sealed up inside their shells.

Palate

Most reptiles lack a secondary palate, meaning that they must hold their breath while swallowing. Crocodilians have evolved a bony secondary palate that allows them to continue breathing while remaining submerged (and protect their brains against damage by struggling prey). Skinks (family Scincidae) also have evolved a bony secondary palate, to varying degrees. Snakes took a different approach and extended their trachea instead. Their tracheal extension sticks out like a fleshy straw, and allows these animals to swallow large prey without suffering from asphyxiation.

Skin

Reptilian skin is covered in a horny epidermis, making it watertight and enabling reptiles to live on dry land, in contrast to amphibians. Compared to mammalian skin, that of reptiles is rather thin and lacks the thick dermal layer that produces leather in mammals. Exposed parts of reptiles are protected by scales or scutes, sometimes with a bony base, forming armor. In lepidosaurians such as lizards and snakes, the whole skin is covered in overlapping epidermal scales. Such scales were once thought to be typical of the class Reptilia as a whole, but are now known to occur only in lepidosaurians. The scales found in turtles and crocodiles are of dermal, rather than epidermal, origin and are properly termed scutes. In turtles, the body is hidden inside a hard shell composed of fused scutes.

Lacking a thick dermis, reptilian leather is not as strong as mammalian leather. It is used in leather-wares for decorative purposes for shoes, belts and handbags, particularly crocodile skin. Due to reptiles lacking feathers or fur, reptiles are used as pets by people with allergies.

Excretion

Excretion is performed mainly by two small kidneys. In diapsids, uric acid is the main nitrogenous waste product; turtles, like mammals, excrete mainly urea. Unlike the kidneys of mammals and birds, reptile kidneys are unable to produce liquid urine more concentrated than their body fluid. This is because they lack a specialized structure called a loop of Henle, which is present in the nephrons of birds and mammals,. Because of this, many reptiles use the colon to aid in the reabsorption of water. Some are also able to take up water stored in the bladder. Excess salts are also excreted by nasal and lingual salt glands in some reptiles.

Digestion

Most reptiles are carnivorous and have rather simple and comparatively short digestive tracts, meat being fairly simple to break down and digest. Digestion is slower than in mammals, reflecting their lower resting metabolism and their inability to divide and masticate their food. Their poikilotherm metabolism has very low energy requirements, allowing large reptiles like crocodiles and the large constrictors to live from a single large meal for months, digesting it slowly.

While modern reptiles are predominately carnivorous, during the early history of reptiles several groups produced some herbivorous megafauna: in the Paleozoic the pareiasaurs and the synapsid dicynodonts, and in the Mesozoic several lines of dinosaurs. Today the turtles are the only predominantly herbivorous reptile group, but several lines of agamas and iguanas have evolved to live wholly or partly on plants.

Herbivorous reptiles face the same problems of mastication as herbivorous mammals but, lacking the complex teeth of mammals, many species swallow rocks and pebbles (so called gastroliths) to aid in digestion: The rocks are washed around in the stomach, helping to grind up plant matter. Fossil gastroliths have been found associated with sauropods. Sea turtles, crocodiles, and marine iguanas also use gastroliths as ballast, helping them to dive.

Nerves

The reptilian nervous system contains the same basic part of the amphibian brain, but the reptile cerebrum and cerebellum are slightly larger. Most typical sense organs are well developed with certain exceptions, most notably the snake's lack of external ears (middle and inner ears are present). There are twelve pairs of cranial nerves. Due to their short cochlea, reptiles use electrical tuning to expand their range of audible frequencies.

Reptiles are generally considered less intelligent than mammals and birds. Though larger reptiles show more complex brain development. Larger lizards like the monitors are known to exhibit complex behavior, including cooperation. Crocodiles have relatively larger brains and show a fairly complex social structure. The Komodo dragon is even known to engage in play.

Vision

Most reptiles are diurnal animals. The vision is typically adapted to daylight conditions, with color vision and more advanced visual depth perception than in amphibians and most mammals. In some species, such as blind snakes, vision is reduced. Some snakes have extra sets of visual organs (in the loosest sense of the word) in the form of pits sensitive to infrared radiation (heat). Such heat-sensitive pits are particularly well developed in the pit vipers, but are also found in boas and pythons. These pits allow the snakes to sense the body heat of birds and mammals, enabling pit vipers to hunt rodents in the dark.

Reproduction

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. Tuataras, however, lack copulatory organs, and so the male and female simply press their cloacas together as the male excretes sperm.

Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion, chorion, and allantois are present during embryonic life. There are no larval stages of development. Viviparity and ovoviviparity have evolved only in squamates, and 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.

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 tuataras. To date, there has been no confirmation of whether TDSD occurs in snakes.

Defense mechanisms

Many small reptiles such as snakes and lizards which live on the ground or in the water are vulnerable to being preyed on by all kinds of carnivorous animals. Thus avoidance is the most common form of defense in reptiles. At the first sign of danger, most snakes and lizards crawl away into the undergrowth, and turtles and crocodiles will plunge into water and sink out of sight. Reptiles may also avoid confrontation through camouflage. Using a variety of grays, greens, and browns, these animals can blend remarkably well into the background of their natural environment.

If the danger arises so suddenly that flight may be harmful, then crocodiles, turtles, some lizards, and some snakes hiss loudly when confronted by an enemy. Rattlesnakes rapidly vibrate the tip of the tail, which is composed of a series of nested, hollow beads.

If all this does not deter an enemy, different species will adopt different defensive tactics.

Snakes use a complicated set of behaviors when attacked. Some will first elevate their head and spread out the skin of their neck in an effort to look bigger and more threatening. Failure of this may lead to other measures practiced particularly by cobras, vipers, and closely related species, who use venom to attack. The venom is modified saliva, delivered through fangs from a venom gland. Some non-venomous snakes, such as the corn snake, play dead when in danger.

When a crocodile is concerned about its safety, it will gape to expose the teeth and yellow tongue. If this doesn't work, the crocodile gets a little more agitated and typically begins to make hissing sounds. After this, the crocodile will start to change its posture dramatically to make itself look more intimidating. The body is inflated to increase apparent size. If absolutely necessary it may decide to attack an enemy. Some species try to bite immediately. Some will use their heads as sledgehammers and literally smash an opponent, some will rush or swim toward the threat from a distance, even chasing them onto land or galloping after them.

Geckos, skinks, and other lizards that are captured by the tail will shed part of the tail structure through a process called autotomy and thus be able to flee. The detached tail will continue to wiggle, creating a deceptive sense of continued struggle and distracting the predator's attention from the fleeing prey animal. The animal can partially regenerate its tail over a period of weeks. The new section will contain cartilage rather than bone, and the skin may be distinctly discolored compared to the rest of the body. The tails are often a separate and dramatically more vivid color then the rest of the body, as to attract potential predators to strike for the tail first.

See also

List of reptiles

Lists of reptiles by region

Herpetophobia

Ophidiophobia

References

Further reading

External links

Sri Lanka Wild Life Information Database

The Reptile Database

Reptile Phylogeny

World Reptile Amphibian Information Center

Reptile images

Category:Articles with inconsistent citation formats Category:Obsolete taxonomic groups

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Coordinates	34°03′″N118°15′″N
Name	Johnny Cage
Series	''Mortal Kombat'' series
Firstgame	''Mortal Kombat'' (1992)
Voiceactor	Jeff Bennett (''The Journey Begins'')Jeff Pilson (''MK2011'')
Liveactor	Daniel Pesina (''MK'', ''MKII'')Chris Alexander (''MKT'')Linden Ashby (first film)Chris Conrad (second film)Jeff Durbin (''Live Tour'')Matt Mullins (''Rebirth'') and (''Legacy'')
Inuniverse	}}

Johnny Cage, whose real name is John Carlton, is a video game character from the ''Mortal Kombat'' fighting game series.

In video games

Johnny Cage introduced as a playable character in the first ''Mortal Kombat'' game. He enters into the titular tournament to prove he does not use special effects in the films he stars in. In the sequel ''Mortal Kombat II'', Cage goes to another tournament in the realm of Outworld after allying with the warriors who wish to protect Earth led by the thunder god Raiden.

He does not reappear in the series until ''Mortal Kombat Trilogy'', where he was killed by Shao Kahn's forces invading Earth. He is shortly revived to help his former comrades defeat Shao Kahn. In the next game, ''Mortal Kombat 4'', Cage is allowed to continue his life after seeking Raiden's help to revive and joins friends once again to defeat the forces from the former god Shinnok who plans to initiate a war between the realms.

By ''Mortal Kombat: Deadly Alliance'', Cage once again joins Raiden's group, in order to stop the menace from the alliance of sorcerers Quan Chi and Shang Tsung. However, Cage is killed alongside most of his comrades and he is resurrected by the Dragon King Onaga to use him as a slave. He is freed from his mind control by the ninja Ermac and Liu Kang's ghost. He returns in ''Mortal Kombat: Armageddon'' as a playable character, participating in the new tournament Armageddon that will decide mankind's fate. In the ensuing battle, Cage is decapitated by an unknown opponent.

He has also appeared in the spin off game ''Mortal Kombat: Shaolin Monks'' that reprises the events from ''Mortal Kombat II''. Besides being a supporting character across the game, he is also playable in the fighting mode.

He reappears in the 2011 ''Mortal Kombat'' game. As the game retells the events from the first tournaments, Cage joins Raiden's forces to save Earth. He and Sonya Blade become the only Earthrealm warriors to survive the course of the game's plot.

Character design

The character was modeled after Jean-Claude Van Damme. Cage's appearance in the first game was loosely based on Van Damme's portrayal of Dux Ryu Ninjutsu founder Frank Dux in the 1988 movie ''Bloodsport'', from his outfit to his "Split Punch". Designed to be a Hollywood star, Cage is meant to be comic relief character in contrast to more serious characters like Liu Kang and Raiden. Johnny Cage's real name came from Midway game artist John Carlton, who worked on the ''NBA Jam'' series.

In other media

Johnny Cage is one of the main characters in the ''Mortal Kombat'' comic book series. His portrayal in the comics is faithful to the games, keeping both his desire to prove his moves are real and his personality traits. He appears during the whole ''Blood & Thunder'' series, most of the time alongside Sonya, with whom he shares a relationship in the comics. His most serious moment is during issue 4 when he, under Raiden's advice, decides to not seek the power of the Tao Te Zhan, since he felt it would be like faking his moves in his films. During the second series, ''Battlewave'', he is seen resuming his acting career when Sonya asks him to help with an investigation after Jax was brutally attacked. He initially refuses, but later reconsiders what's important and decides to follow with Jax into the Outworld. In the meantime, he sends his musclebound bodyguard, Bo, to protect Liu Kang.

In the first ''Mortal Kombat'' movie, Johnny Cage was played by Linden Ashby, whose portrayal was faithful to the games in keeping with Cage's personality and his comic relief role. During the film, he fights with both Scorpion and Goro in Mortal Kombat, and manages to defeat both.

Johnny Cage reappears in ''Mortal Kombat: Annihilation''. He was played by Chris Conrad. In the opening scene, Shao Kahn takes Sonya Blade hostage, and Cage attempts to save her. He is killed when Kahn snaps his neck.

Matt Mullins appears as Johnny Cage in the internet video, ''Mortal Kombat: Rebirth''. He was an action movie star, whose career was dying. To allow his talents to still be used, he became an undercover agent for Jackson Briggs. However, Alan Zane / Baraka killed him shortly thereafter in battle. Mullins reprised his role in ''Mortal Kombat: Legacy'', in which he plays a career failing Johnny Cage, a former "ass-kicking teen heartthrob from ''Power Rangers''."

Reception

UGO.com ranked Johnny Cage fourth on their "Top 11 Mortal Kombat Characters" list, praising his role as the comic relief of the series with the added statement that "there's nothing better than kicking somebody's ass with a fighter that's supposed to be a joke". GameDaily put him 23rd in their feature "Top 25 Gaming Hunks", stating "If you can't marry Brad Pitt, you can always settle for Mortal Kombat's Johnny Cage". 1UP.com ranked him nineth in their "They Is Risen" feature, which covered the ten most notorious video game resurrections. The publication noted that one of the most remarkable part of his character was that he was one of the few ones who died during the storyline. Cage's Split Punch was in GamesRadar's article "Gaming’s most devastating genital attacks" with its author commenting that "It’s hardly Cage’s most impressive or most damaging move, but it’s inarguably his best." In another article, the split-punch was listed as his signature move with notes that while it is performed in a "decidedly un-Street Fighter way", it was the first "silly" moment in the franchise. Johnny Cage's split-punch on Goro in the ''Mortal Kombat'' film was rated by UGO.com as one the top 11 Game Movie Moments. He was also ninth in Game Revolution's "Top 10 Old School Mortal Kombat Characters" with praise on his special moves that made him stand out. IGN listed him as a character they would like to see as downloadable content for ''Mortal Kombat vs. DC Universe'', stating Johnny "remains a favorite among MK heroes, even though he's spent a good portion of the series' life in a coffin".

References

Category:Fictional American people Category:Fictional actors Category:Fictional characters based on real people Category:Fictional characters introduced in 1992 Category:Film characters Category:Fictional Jeet Kune Do practitioners Category:Fictional karateka Category:Male video game characters Category:Mortal Kombat characters

ar:جوني كايج ca:Johnny Cage es:Johnny Cage fr:Johnny Cage it:Johnny Cage pt:Johnny Cage ru:Джонни Кейдж simple:Johnny Cage fi:Johnny Cage tr:Johnny Cage ur:جانی کیج (مارٹل کامبیٹ)