Reptiles Temporal range: Pennsylvanian - Recent 312–0 Ma PreЄ Є O S D C P T J K Pg N
Clockwise from above left: Green turtle (Chelonia mydas), Tuatara (Sphenodon punctatus), Nile Crocodile (Crocodylus niloticus), and Sinai Agama (Pseudotrapelus sinaitus).
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
clade:	Amniota
Class:	Reptilia Laurenti, 1768
Subgroups
Anapsida (=Parareptilia?) Testudines (traditional) Eureptilia Saurischia (only living subgroup Aves) Crocodilia Sphenodontia Squamata Testudines (molecular) For extinct orders, see text

Reptiles (Reptilia) are members of a class of air-breathing, ectothermic (cold-blooded) vertebrates^[1] which are characterized by skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors. Modern reptiles inhabit every continent with the exception of Antarctica. Reptiles originated around 320-310 million years ago during the Carboniferous period, having evolved from advanced reptile-like amphibians that became increasingly adapted to life on dry land. Four living orders are typically recognized:

• Crocodilia (crocodiles, gavials, caimans, and alligators): 23 species
• Sphenodontia (tuataras from New Zealand): 2 species
• Squamata (lizards, snakes, and worm lizards): approximately 9,150 species^[2]
• Testudines (turtles, terrapins and tortoises): over 300 species

Aves (birds), which are not cold-blooded or scaly, are not included in this list. Crocodiles are more closely related to birds than they are to lizards, however, so some writers who prefer a more unified (monophyletic) grouping also include the birds: over 10,000 species.

Unlike amphibians, reptiles do not have an aquatic larval stage. As a rule, reptiles are oviparous (egg-laying), although certain species of squamates retain the eggs until hatching and a few are viviparous. 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 may reach 6 m (19.7 ft) in length and weigh over 1,000 kg (2,200 lb).

The study of reptiles and amphibians is called herpetology.

Classification[link]

History of classification[link]

Reptiles, from Nouveau Larousse Illustré, 1897-1904. Notice the inclusion of amphibians (below the crocodiles).

Linnaeus and the 18th century[link]

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.^[3] The terms "reptile" and "amphibian" were largely interchangeable, "reptile" (from Latin repere, "to creep") being preferred by the French.^[4] 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.^[5] Today, it is still common to treat the two groups under the same heading as herptiles.

"Antediluvian monsters"[link]

An "antediluvian monster", a Mosasaurus discovered in a Maastricht limestone quarry, 1770 (contemporary engraving)

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.^[6]

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.^[7]

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") which included some Paleozoic amphibians as well as early reptiles.^[8]

The traditional class Reptilia (green field) is a paraphyletic group comprising all non-avian and non-mammalian amniotes.^{[note 1]}

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.^[9]

In the late 19th century, a number of definitions of Reptilia were offered. The traits listed by Lydekker in 1896, for example, include a single occipital condyle, a jaw joint formed by the quadrate and articular bones, and certain characteristics of the vertebrae.^[10] The animals singled out by these formulations, the amniotes other than the mammals and the birds, are still those considered reptiles today.

Skull openings in 20th-century classification[link]

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 and made popular by Romer's classic Vertebrate Paleontology.^[11][12] Those four subclasses were:

• Anapsida – no fenestrae - cotylosaurs and Chelonia (turtles and relatives)^{[note 2]}
• 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[link]

By the 21st century, vertebrate paleontologists were beginning to adopt 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 animals like dinosaurs and early therapsids that were traditionally called reptiles.^[13] 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'.^[14]

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.^[15] 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. Because the actual relationship of turtles to other reptiles was not yet well understood at this time, Gauthier's definition came to be considered inadequate.^[15] A variety of other definitions were proposed by other scientists in the years following Gauthier's paper. The first such new definition, which attempted to adhere to the standards of the PhyloCode, was published by Modesto and Anderson in 2004. Modesto and Anderson 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 the more common definition of Sauropsida, which Modesto and Anderson synonymized with Reptilia, since the latter is more well known and more frequently used. Unlike most previous definitions of Reptilia, however, Modesto and Anderson's definition includes birds.^[15]

Taxonomy[link]

Classification to order level, after Benton, 2005.^{[16][note 2]}

• 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 Ornithischia
                • Order Saurischia
                  • Class Aves

Phylogeny[link]

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,^[17] with information on the relationships of the most primitive reptiles after Muller and Reisz (2006).^{[18][note 2]}

1. ^ Some workers in phylogenetic nomenclature employ the term "Reptilia" in a more restrictive sense, see Sauropsida
2. ^ ^{a b c} The above taxonomy and phylogeny do not reflect modern molecular evidence that places turtles within Diapsida.

Evolutionary history[link]

Origin of the reptiles[link]

An early reptile Hylonomus

Mesozoic scene showing typical reptilian megafauna: dinosaurs including Europasaurus holgeri, iguanodonts and Archaeopteryx lithographica perched on the foreground tree stump.

Megalania was a giant, carnivorous goanna that might have grown to as long as 7 metres, and weighed up to 1,940 kilograms (Molnar, 2004).

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.^[19] The oldest known animal that may have been an amniote, i.e. a primitive reptile rather than an advanced amphibian is Casineria.^[20][21] A series of footprints from the fossil strata of Nova Scotia, dated to 315 million years show typical reptilian toes and imprints of scales.^[22] The tracks are attributed to Hylonomus, the oldest unquestionable reptile known.^[23] It was a small, lizard-like animal, about 20 to 30 cm (8–12 in) long, with numerous sharp teeth indicating an insectivorous diet.^[24] Other examples include Westlothiana (for the moment considered a reptiliomorph amphibian rather than a true amniote)^[25] and Paleothyris, both of similar build and presumably similar habit.

Rise of the reptiles[link]

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.^[26] 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 possess a shell that allows them to 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.^[26] 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).^[27] 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[link]

A = Anapsid, B = Synapsid, C = Diapsid

The first reptiles retained an anapsid skull inherited from their amphibian ancestors.^[28] This type of skull has a skull roof with only holes for the nostrils, eyes and a pineal eye.^[29] These animals are generally referred to as anapsids, and form a paraphyletic basic stock from which other groups evolved.^[15] Very shortly after the reptiles appeared, a lineage called Synapsida split off, characterized by a temporal opening in the skull behind each eye to give room for the jaw muscle to move. These are the "mammal-like reptiles" that later gave rise to the true mammals.^[30] Soon after, another group evolved a similar trait, this time with a double opening behind each eye, earning them the name Diapsida ("two arches").^[28] The function of the holes in these groups was to lighten the skull and give room for the jaw muscles to move, allowing for a more powerful bite.^[29] Diapsids and the anapsid reptiles back to the last common ancestor of diapsids and synapsids are classed as the "true reptiles", Sauropsida.^[8]

Turtles have been traditionally believed to be surviving reptilian anapsids, on the basis of their skull structure.^[31] The rationale for this classification has been disputed, with some arguing that turtles are diapsids that reverted to this primitive state in order to improve their armor (see Parareptilia).^[19] Later morphological phylogenetic studies with this in mind placed turtles firmly within Diapsida.^[32] All molecular studies have strongly upheld the placement of turtles within diapsids, most commonly as a sister group to extant archosaurs.^{[33][34][35][36]}

Permian reptiles[link]

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 pelycosaurs were replaced by the therapsids.^[37]

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).^[37]

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"[link]

The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic extinction event), a prolonged event due to the accumulation of at least two distinct extinction pulses.^[38] 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.^[38] Archosaurs developed 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.^[37]

The lepidosauromorph diapsids may have been ancestral to the sea reptiles.^[39] 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 mammals being the only survivors of the line by the late Cretaceous.

Cenozoic era[link]

The close of the Cretaceous period saw the demise of the Mesozoic era reptilian megafauna (see the Cretaceous–Paleogene extinction event). Of the large marine reptiles, only sea turtles were left, and of the non-marine large reptiles, only the semi-aquatic crocodiles and broadly similar Champsosaurus survived the extinction, with the latter becoming extinct in the Miocene.^[40] Of the great host of dinosaurs dominating the Mesozoic, only the small feathered birds survived. This dramatic extinction pattern at the end of the “Age of Reptiles” led into the “Age of Mammals”. Mammals and birds filled the empty niches left behind by the reptilian megafauna, and while reptile diversification slowed, bird and mammal diversification took an exponential turn.^[27]

After the extinction of most archosaur and marine reptile lines end of the Cretaceous, 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 living reptiles (> 95%), most species being lizards.^[2] There are approximately 9,500 extant species of reptiles,^[2] compared with 5,500 species of mammals.

Systems[link]

Circulation[link]

Thermographic image of a monitor lizard

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.^[41]

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.^[42] 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.^[43]

Metabolism[link]

Sustained energy output (Joules) of a typical reptile versus a similar size mammal as a function of core body temperature. The mammal has a much higher peak output but can only function over a very narrow range of body temperatures.

All 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 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 to 35 °C (75 to 95 °F) range,^[44] 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).^[45] 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 although they do not have a warm-blooded metabolism.^[46] This form of homeothermy is called gigantothermy; it has been suggested as having been common in large dinosaurs and other extinct large-bodied reptiles.^[47][48]

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.^[49] 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.^[50] 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.^[51] Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals.^[52] However, investigation of correlations between active capacity and thermophysiology show a weak relationship.^[53] Most extant reptiles are carnivores with a sit-and-wait feeding strategy, and whether reptiles are cold blooded due to their ecology or because 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.^[54]

Respiration[link]

Reptilian lungs[link]

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.^[55] 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.^[56] 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[link]

Red-eared slider taking a gulp of air

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.^[57] 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.^[57]

Palate[link]

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[link]

Skin of a sand lizard, showing squamate reptiles iconic scales

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.^[58] 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[link]

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[link]

Gastrolithes from a plesiosaur

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[link]

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.^[59] 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.^[29] The size of their brain relative to their body is much less than that of mammals, the encephalization quotient being about one tenth of that of mammals,^[60] though larger reptiles show more complex brain development. Larger lizards like the monitors are known to exhibit complex behavior, including cooperation.^[61] Crocodiles have relatively larger brains and show a fairly complex social structure. The Komodo dragon is even known to engage in play.^[62]

Vision[link]

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[link]

Most reptiles reproduce sexually such as this Trachylepis maculilabris skink

Reptiles have amniotic eggs with hard or leathery shells, requiring internal fertilization.

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.^[63]

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.^[64] To date, there has been no confirmation of whether TDSD occurs in snakes.^[65]

Defense mechanisms[link]

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.^[66] 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.

Camouflage[link]

A camouflaged Phelsuma deubia on a palm frond

Reptiles tend to avoid confrontation through camouflage. Two major groups of reptile predators are birds and other reptiles, both of whom has well developed colour vision. Thus many reptiles uses a variety of grays, greens, and browns, allowing these animals can blend remarkably well into the background of their natural environment.^[67]

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.

Alternative defense in snakes[link]

Camouflage will not always fool a predator. When caught out, snake 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 American corn snake or European grass snake, play dead when in danger.

Defense in crocodiles[link]

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.

A White-headed dwarf gecko with shed tail

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.^[68]

Shedding and regenerating tails[link]

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[link]

• List of reptiles
• Lists of reptiles by region
• Herpetophobia
• Ophidiophobia

References[link]

Further reading[link]

External links[link]

Find more about Reptile on Wikipedia's sister projects:
	Definitions and translations from Wiktionary
	Images and media from Commons
	Learning resources from Wikiversity
	News stories from Wikinews
	Quotations from Wikiquote
	Source texts from Wikisource
	Textbooks from Wikibooks

• Data related to Reptilia at Wikispecies
• Reptilia at Wikibooks
• Reptile Phylogeny
• Reptile images
• Sri Lanka Wild Life Information Database

Trent Reznor
Background information
Birth name	Michael Trent Reznor
Born	(1965-05-17) May 17, 1965 (age 47) Mercer, Pennsylvania, U.S.
Genres	Industrial rock, industrial metal, post-industrial, dark ambient, alternative rock, experimental rock
Occupations	Musician, songwriter, sound designer, record producer, film score composer
Instruments	Vocals, guitar, piano, synthesizer, mellotron, keyboards, bass guitar, saxophone, cello, double bass, drums, tuba, sousaphone, harmonium, marimba, pan flute, harpsichord, vibraphone, swarmatron
Years active	1982–present
Labels	The Null Corporation, Nothing, Interscope, Universal, TVT
Associated acts	Option 30, The Innocent, Exotic Birds, Nine Inch Nails, How to Destroy Angels, Tapeworm, Atticus Ross, Marilyn Manson, David Fincher
Website	www.nin.com

Michael Trent Reznor (born May 17, 1965) is an American singer–songwriter, composer, and record producer. As both a vocalist and a multi-instrumentalist, Reznor has led the industrial rock project Nine Inch Nails beginning in 1988, and as of 2010 he and his wife, Mariqueen Maandig, are also members of the post-industrial^[1][2] band How to Destroy Angels alongside fellow composer Atticus Ross,^[1][3] with whom Reznor has scored two films. Reznor was previously associated with the bands Option 30, Exotic Birds, and Tapeworm, among others. Reznor left Interscope Records in 2007, and is now an independent recording artist.

Reznor began creating music early in his life, and cites his Western Pennsylvania childhood as an early influence. After being involved with a number of synthesizer-based bands in the mid-80s, Reznor gained employment at Right Track Studios in Cleveland, Ohio and began creating his own music during the studio's closing hours under the moniker Nine Inch Nails. Reznor's first release as Nine Inch Nails, Pretty Hate Machine (1989), was a commercial and critical success, and he has since released seven major studio releases. Outside of Reznor's chief project Nine Inch Nails, he has contributed to many other artists' albums, including Marilyn Manson and Saul Williams. In 1997, Reznor appeared in Time magazine's list of the year's most influential people, and in Spin magazine he was described as "the most vital artist in music."^[4]

Reznor, in collaboration with Atticus Ross, composed the score for the soundtrack to The Social Network, a 2010 theatrical film about the founding of Facebook. The duo won the 2010 Golden Globe Award for Best Original Score and the Academy Award for Best Original Score for their collaboration. Reznor and Ross also composed the score to the US film adaptation of the novel The Girl with the Dragon Tattoo, which was released in late 2011. Both soundtrack albums were released by The Null Corporation, Reznor's own independent record label.

Early life[link]

Reznor was born in Mercer, Pennsylvania, halfway between Pittsburgh and Erie, the son of Nancy Lou (née Clark) and Michael Reznor.^[5] After his parents divorced, he lived with his maternal grandparents, while his sister Tera lived with their mother.^[6] Reznor is a direct descendant of George Reznor, who founded the Reznor Company, a heating and air conditioning company, in 1888. The family sold the business in the 1960s.^[5][7]

Reznor began playing the piano at the age of five and showed an early aptitude for music. In a 1995 interview, his grandfather, Bill Clark, remarked, "Music was his life, from the time he was a wee boy. He was so gifted."^[8] His former piano teacher Rita Beglin said "Reznor always reminded me of Harry Connick, Jr." when he played.^[8]

Reznor has acknowledged that his sheltered life in Pennsylvania left him feeling isolated from the outside world. In a 1994 interview with Rolling Stone, he references his choices in the music industry.

However, Reznor later said, "I don't want to give the impression it was a miserable childhood."^[10]

At Mercer Area Junior/Senior High School, Reznor learned to play the tenor saxophone and tuba. He was a member of both the jazz and marching band. Former Mercer High School band director Dr. Hendley Hoge remembered Reznor as "very upbeat and friendly."^[8] Reznor also became involved in theater while in high school. He was voted "Best in Drama" by classmates for his roles as Judas in Jesus Christ Superstar and Professor Harold Hill in The Music Man. Reznor graduated from this high school in 1983 and enrolled at Allegheny College, where he studied computer engineering.^[11]

Music career[link]

Early musical projects[link]

While he was enrolled in Allegheny College, Reznor joined local band Option 30, playing three shows a week with them. After only a year of college, Reznor dropped out and moved to Cleveland, Ohio to pursue a career in music.^[8] In 1985, he joined The Innocent as a keyboardist. They released one album, Livin' in the Street, but Reznor quit after just three months. In 1986, Reznor joined local band Exotic Birds, during this time, Reznor also contributed on keyboards to the band Slam Bamboo. He also appeared with them as the fictional band "The Problems" in the 1987 film Light of Day.^[12]

Reznor got a job at Cleveland's Right Track Studio as an assistant engineer and janitor.^[13] Studio owner Bart Koster later commented that Reznor was "so focused in everything he [did]. When that guy waxed the floor, it looked great."^[8] Reznor asked Koster for permission to record demos of his own songs for free during unused studio time. Koster agreed, remarking that it cost him "just a little wear on [his] tape heads".^[8] While assembling these, the earliest Nine Inch Nails recordings, Reznor was unable to find a band that could articulate his songs as he wanted. Instead, inspired by Prince, he played all the instruments himself, except drums.^[14] This role remains Reznor's on most of the band's studio recordings, though he has occasionally involved other musicians and assistants. Several labels responded favorably to the demo material, and Reznor signed with TVT Records.^[13] Nine selections from the Right Track demos were unofficially released in 1988 as Purest Feeling; many of these songs had appeared in revised form on his 1989 debut studio album under the stage name Nine Inch Nails.

Formation of Nine Inch Nails[link]

Reznor performing at the Lollapalooza festival, 1991.

Most of Reznor's work as a musician has been as founding and primary member of Nine Inch Nails. Nine Inch Nails' debut album, Pretty Hate Machine was released in 1989. It was a moderate commercial success, and was certified Gold in 1992.^[15] Amid pressure from Reznor's record label to produce a follow-up to Pretty Hate Machine, Reznor secretly began recording under various pseudonyms to avoid record company interference, resulting in the Broken EP, released in 1992.^[16] In the summer of 1991 Nine Inch Nails was included in the Lollapalooza package tour. They later won a Grammy Award in 1993 for the song "Wish" in the Best Heavy Metal Performance category.^[17]

Nine Inch Nails' second full-length album, The Downward Spiral, entered the Billboard 200 chart in 1994 at number two,^[18] and remains the highest-selling Nine Inch Nails release in the United States.^[15] To record the album, Reznor rented and moved into the 10050 Cielo Drive mansion, site of the 1969 Manson Family murders.^[19] Reznor built a studio space in the house, which he renamed Le Pig, after the word that was scrawled on the front door in Sharon Tate's blood by her murderers.^[19] Reznor told Entertainment Weekly that despite the notoriety attached to the house, he chose to record there because, "I looked at a lot of places, and this just happened to be the one I liked most".^[19]

Nine Inch Nails toured extensively over the next few years, including a performance at Woodstock '94, although he admitted to the audience that he did not like to play large venues.^[20] Around this time, Reznor's studio perfectionism,^[21] struggles with addiction, and bouts of writer's block prolonged the production of a follow-up.^[22]

Collaboration with other artists[link]

One of Reznor's earliest collaborations was a Ministry side project in 1990 under the name of 1000 Homo DJs. Reznor sang vocals on a cover of Black Sabbath's Supernaut. Due to legal issues with Reznor's label, Reznor's vocals had to be distorted enough to make his voice indistinguishable. The band also recorded additional versions with Al Jourgensen doing vocals.^[23] While there is still debate as to which version is Reznor and which is Jourgensen, it has been definitively stated that Reznor's vocals were used in the TVT Record's Black Box Box set.^[24] Reznor sang backing vocals on Tori Amos' 1994 album Under the Pink on a track called "Past The Mission." Reznor produced Marilyn Manson's first album Portrait of an American Family (1994), several tracks on Marilyn Manson's albums Smells Like Children (1995), and Antichrist Superstar (1996), as well as the film soundtracks for Oliver Stone's Natural Born Killers (1994) and David Lynch's Lost Highway (1997). Reznor is credited for "Driver Down" and "Videodrones; Questions" on the soundtrack for Lost Highway, while another track, "The Perfect Drug", is credited to Nine Inch Nails instead. In 1998 Reznor produced a remix of Notorious B.I.G.'s song "Victory" which also featured Busta Rhymes.^[25] He also unofficially worked with Garrett Ahlstrom on "Big Black Bomb" a remix on "Burn" he stated on the spiral, "I never worked on the song but I still consider it an official mix"