Reptiles Temporal range: Pennsylvanian – Holocene 312–0Ma 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
Extant orders
Anapsida (=Parareptilia?) Testudines (traditional) Eureptilia Crocodilia Sphenodontia Squamata Testudines (molecular) For extinct orders, see text

Traditionally, reptiles are members of the class Reptilia comprising the amniotes that are neither birds nor mammals.^[1] (The amniotes are the vertebrates with eggs featuring an amnion, a double membrane that permits the embryo to breathe effectively on land.) Living reptiles, in that sense, can be distinguished from other tetrapods in that they are cold-blooded and bear scutes or scales.

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. Many groups are extinct, including dinosaurs, pterosaurs, and aquatic groups such as the ichthyosaurs. Modern reptiles inhabit every continent with the exception of Antarctica. Several living subgroups are recognized:

• Testudines (turtles, terrapins and tortoises): approximately 330 species^[2]
• Sphenodontia (tuataras from New Zealand): 2 species^[2]
• Squamata (lizards, snakes, and worm lizards): over 9,400 species^[2]
• Crocodilia (crocodiles, gavials, caimans, and alligators): 25 species^[2]

Although they have scutes on their feet and lay eggs, birds have historically been excluded from the reptiles. They therefore do not appear on the list above. However, as some reptiles are more closely related to birds than they are to other reptiles — crocodiles are more closely related to birds than they are to lizards — cladistic writers who prefer a more unified (monophyletic) grouping usually also include the birds, which include over 10,000 species.^[3][4][5] (See Sauropsida.)

Reptiles are tetrapod vertebrates, either having four limbs or, like snakes, being descended from four-limbed ancestors. Unlike amphibians, reptiles do not have an aquatic larval stage. Most reptiles are oviparous (egg-laying), although several species of squamates and of extinct aquatic clades are viviparous^[6] — the fetus develops within the mother, contained in a placenta rather than an eggshell. As amniotes, reptile eggs are surrounded by membranes for protection and transport that adapt them to reproduction on dry land. Many of the viviparous species feed their fetuses through various forms of placenta analogous to those of mammals, with some providing initial care for their hatchlings. Extant reptiles range in size from a tiny gecko, Sphaerodactylus ariasae, which can grow up to 17 mm (0.7 in) to the saltwater crocodile, Crocodylus porosus, which may reach 6 m (19.7 ft) in length and weigh over 1,000 kg (2,200 lb).

Traditionally, the reptiles are one of the three classes of amniotes, the others being the mammals and the birds. By the middle of the 20th century, it had become common to split the reptiles into four groups based on their skull openings. However, as reptiles in the traditional sense are the ancestors of both birds and mammals, many taxonomists have more recently adopted a cladistic approach, preferring to group animals based exclusively on shared ancestry rather than on shared features; this orientation has led some to abandon reptiles as a systematic unit or redefine the reptiles in a manner that includes birds.

The historically combined study of reptiles and amphibians is called herpetology.

Classification [edit]

History of classification [edit]

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

Linnaeus and the 18th century [edit]

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.^[7] The terms "reptile" and "amphibian" were largely interchangeable, "reptile" (from Latin repere, "to creep") being preferred by the French.^[8] 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.^[9] Today, the two groups are still commonly treated under the same heading as herptiles.

"Antediluvian monsters" [edit]

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.^[10]

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.^[11]

In 1866, Haeckel demonstrated that vertebrates could be divided based on their reproductive strategies, and that reptiles, birds, and mammals were united by the amniotic egg.

The terms "Sauropsida" ("lizard faces") and "Theropsida" ("beast faces") were used again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and their relatives on the one hand (Sauropsida) and mammals and their extinct relatives (Theropsida) on the other. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, Protosauria ("first lizards") in which he included some animals today considered reptile-like amphibians, as well as early reptiles.^[12]

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, 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.^[13]

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.^[14] The animals singled out by these formulations, the amniotes other than the mammals and the birds, are still those considered reptiles today.^[1]

Skull openings in 20th-century classification [edit]

The first reptiles had an anapsid type of skull roof, as seen in the Permian genus Captorhinus

The synapsid/sauropsid division supplemented another approach, one that split 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.^[15][16] 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. It has largely been abandoned among recent researchers; in particular, the anapsid condition has been found to occur so variably among unrelated groups that it is not now considered a useful distinction.^[17]

Phylogenetics and modern definition [edit]

Phylogentic classifications group mammal-like reptiles like this Varanodon with other synapsids, not with extant reptiles.

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.^[18] 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'.^[1]

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.^[19] 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.^[19] 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^[19], as there is no other way to establish a monophyletic clade that includes both lizards and crocodiles.

Taxonomy [edit]

Classification to order level, after Benton, 2005.^{[20][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)
      • 

        

        Vjushkovia, a basal archosauropmorph
        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 [edit]

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,^[21] with information on the relationships of the most primitive reptiles after Muller and Reisz (2006)^[22] and the placement of Testudines following Lyson and Gilbert (2009).^[23]

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

Evolutionary history [edit]

Origin of the reptiles [edit]

An early reptile Hylonomus

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

The origin of the reptiles lies about 310–320 million years ago, in the steaming swamps of the late Carboniferous period, when the first reptiles evolved from advanced reptiliomorph labyrinthodonts.^[24] The oldest known animal that may have been an amniote, i.e. a primitive reptile rather than an advanced amphibian is Casineria (though it may have been a temnospondyl amphibian).^[25][26][27] A series of footprints from the fossil strata of Nova Scotia dated to 315 Ma show typical reptilian toes and imprints of scales.^[28] The tracks are attributed to Hylonomus, the oldest unquestionable reptile known.^[29] It was a small, lizard-like animal, about 20 to 30 centimetres (7.9 to 12 in) long, with numerous sharp teeth indicating an insectivorous diet.^[30] Other examples include Westlothiana (for the moment considered a reptiliomorph amphibian rather than a true amniote)^[31] and Paleothyris, both of similar build and presumably similar habit.

Rise of the reptiles [edit]

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.^[32] 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.^[32] 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).^[33] 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 [edit]

A = Anapsid, B = Synapsid, C = Diapsid

It was traditionally assumed that the first reptiles retained an anapsid skull inherited from their amphibian ancestors.^[34] This type of skull has a skull roof with only holes for the nostrils, eyes and a pineal eye.^[35] The discoveries of synapsid-like openings (see below) in the skull roof of the skulls of several members of Parareptilia (the clade containing most of reptiles traditionally referred to as anapsids), including lanthanosuchoids, millerettids, bolosaurids, some nycteroleterids, some procolophonoids and at least some mesosaurs^[36][37][38] made it more ambiguous and it's currently uncertain whether the ancestral reptile had an anapsid-like or synapsid-like skull.^[38] These animals are traditionally referred to as anapsids, and form a paraphyletic basic stock from which other groups evolved.^[19] 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.^[39] Soon after, another group evolved a similar trait, this time with a double opening behind each eye, earning them the name Diapsida ("two arches").^[34] 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.^[35] Diapsids and the anapsid reptiles back to the last common ancestor of diapsids and synapsids are classed as the "true reptiles", Sauropsida.^[12]

Turtles have been traditionally believed to be surviving reptilian anapsids, on the basis of their skull structure.^[40] 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).^[24] Later morphological phylogenetic studies with this in mind placed turtles firmly within Diapsida.^[41] All molecular studies have strongly upheld the placement of turtles within diapsids, most commonly as a sister group to extant archosaurs.^{[42][43][44][45]}

Permian reptiles [edit]

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 became drier, resulting in a change of fauna: The pelycosaurs were replaced by the therapsids.^[46]

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).^[46]

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" [edit]

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.^[47] 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.^[47] 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.^[46]

The sister group to Archosauromorpha is Lepidosauromorpha, containing squamates and rhynchocephalians, as well as their fossil relatives. Lepidosauromorpha contained at least one major group of the Mesozoic sea reptiles: the mosasaurs, which emerged during the Cretaceous period. The phylogenetic placement of other main groups of fossil sea reptiles – the sauropterygians and the ichthyosaurs, which evolved in the early Triassic and in the Middle Triassic respectively – is more controversial. Different authors linked these groups either to lepidosauromorphs^[48] or to archosauromorphs,^[49][50][51] and ichthyosaurs were also argued to be diapsids that did not belong to the least inclusive clade containing lepidosauromorphs and archosauromorphs.^[52]

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.

The megalania was a giant carnivorous goanna, perhaps as long as 7 metres and weighing up to 1,940 kilograms.^[53] Early humans in Australia may have encountered this reptile.

Cenozoic era [edit]

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.^[54] 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.^[33]

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] Approximately 9,800 extant species of reptiles are known, compared with about 5,700 species of mammals alive today (excluding domesticated species).^[55]

Systems [edit]

Circulation [edit]

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.^[56]

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.^[57] 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.^[58]

Metabolism [edit]

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 temperature.

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°–35°C (75°–95°F) range,^[59] 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).^[60] 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.^[61] This form of homeothermy is called gigantothermy; it has been suggested as having been common in large dinosaurs and other extinct large-bodied reptiles.^[62][63]

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.^[64] 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.^[65] 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.^[66] Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals.^[67] However, investigation of correlations between active capacity and thermophysiology show a weak relationship.^[68] 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.^[69]

Respiration [edit]

Reptilian lungs [edit]

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.^[70] 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.^[71]

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." The lung portions of the bronchia form a number of double tubular chambers within each lung. On inhalation, air moves into the upper chambers, and on exhalation exits through the lower chambers, thus creating an unidirectional airflow through the lungs. A similar, though highly expanded system is found in birds.^[72]

Turtles and tortoises [edit]

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.^[73] 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.^[73]

Palate [edit]

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 [edit]

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.^[74] 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 [edit]

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 [edit]

Watersnake Malpolon monspessulanus eating a lizard. Most reptiles are carnivorous, and many primarily eat other reptiles.

Gastrolithes from a plesiosaur

Most reptiles are insectivorous or 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.^[75] 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.^[65]

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.^[33] 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.^[76]

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.^[76] Fossil gastroliths have been found associated with both ornithopods and sauropods, though whether they actually functioned as a gastric mill in the latter is disputed.^[77][78] Salt water crocodiles also use gastroliths as ballast, stabilizing them in the water or helping them to dive.^[79] A dual function as both stabilizing ballast and digestion aid has been suggested for gastrolithes found in plesiosaurs.^[80]

Nerves [edit]

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.^[81] 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.^[35] 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,^[82] though larger reptiles can show more complex brain development. Larger lizards, like the monitors, are known to exhibit complex behavior, including cooperation.^[83] Crocodiles have relatively larger brains and show a fairly complex social structure. The Komodo dragon is even known to engage in play.^[84]

Vision [edit]

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 [edit]

Most reptiles reproduce sexually such as this Trachylepis maculilabris skink

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

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 discharges sperm.^[85]

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 in many extinct clades of reptiles and in squamates. In the latter group, many species, including all boas and most vipers, utilize this mode of reproduction. The degree of viviparity varies: some species simply retain the eggs until just before hatching, others provide maternal nourishment to supplement the yolk, and yet others lack any yolk and provide all nutrients via a structure similar to the mammalian placenta. The earliest documented case of viviparity in reptiles is the Early Permian mesosaurs,^[86] although some individuals or taxa in that clade may also have been oviparous because a putative isolated egg has also been found. Several groups of Mesozoic marine reptiles also exhibited viviparity, such as mosasaurs, ichthyosaurs, and Sauropterygia, a group that include pachypleurosaurs and Plesiosauria.^[6]

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.^[87] To date, there has been no confirmation of whether TDSD occurs in snakes.^[88]

Defense mechanisms [edit]

Many small reptiles, such as snakes and lizards that 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.^[89] 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 [edit]

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 which have well developed colour vision. Thus the skins of many reptiles are plain or mottled gray, green, and brown to allow them to blend into the background of their natural environment.^[90]

If the danger arises so suddenly that flight is useless, 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 [edit]

Camouflage will not always fool a predator. When caught out, snake species will adopt different defensive tactics and 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 large and threatening. Failure of this strategy 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 crocodilians [edit]

When a crocodilian is concerned about its safety, it will gape to expose the teeth and yellow tongue. If this doesn't work, the crocodilian gets a little more agitated and typically begins to make hissing sounds. After this, the crocodilian 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 the opponent onto land or galloping after it.^[91]

Shedding and regenerating tails [edit]

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 than the rest of the body so as to encourage potential predators to strike for the tail first.

See also [edit]

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

References [edit]

Further reading [edit]

External links [edit]

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