Reptile

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For other uses, see Reptile (disambiguation).

Reptiles
Temporal range: PennsylvanianPresent, 312

0 Ma

Pre

Pg

Clockwise from above left: Green sea turtle(Chelonia

mydas), Tuatara (Sphenodon punctatus), Nile
 crocodile (Crocodylus niloticus), and Sinai

agama (Pseudotrapelus sinaitus).

Scientific classification

Kingdom: Animalia

Phylum: Chordata

Clade: Sauropsida

Class: Reptilia
Laurenti, 1768

Extant groups

Crocodilia (crocodilians)
Lepidosauria (lepidosaurs)
Rhynchocephalia (tuatara and relatives)
Squamata (lizards & snakes)
Testudines (turtles)
Aves (birds) (cladistically included)

See text for extinct groups.

Global reptile distribution

 Reptiles are tetrapod animals in the class Reptilia, comprising
today's turtles, crocodilians, snakes, amphisbaenians, lizards, tuatara, and their extinct relatives. The
study of these traditional reptile orders, historically combined with that of modern amphibians, is
called herpetology.
Because some reptiles are more closely related to birds than they are to other reptiles (e.g.,
crocodiles are more closely related to birds than they are to lizards), the traditional groups of
"reptiles" listed above do not together constitute a monophyletic grouping (or clade). For this reason,
many modern scientists prefer to consider the birds part of Reptilia as well, thereby making Reptilia
a monophyletic class.[1][2][3][4]
The earliest known proto-reptiles originated around 312 million years ago during
the Carboniferous period, having evolved from advanced reptiliomorph tetrapods that became
increasingly adapted to life on dry land. Some early examples include the lizard-
like Hylonomus and Casineria. In addition to the living reptiles, there are many diverse groups that
are now extinct, in some cases due to mass extinction events. In particular, the Cretaceous
Paleogene extinction event wiped out the pterosaurs, plesiosaurs, ornithischians, and sauropods, as
well as many species of theropods, including troodontids, dromaeosaurids, tyrannosaurids,
and abelisaurids, along with many Crocodyliformes, and squamates (e.g. mosasaurids).
Modern non-avian reptiles inhabit every continent with the exception of Antarctica. (If birds are
classed as reptiles, then all continents are inhabited.) Several living subgroups are
recognized: Testudines (turtles and tortoises), approximately 400
species;[5]Rhynchocephalia (tuatara from New Zealand), 1 species;[5][6] Squamata (lizards, snakes,
and worm lizards), over 9,600 species;[5]Crocodilia (crocodiles, gavials, caimans, and alligators), 25
species;[5] and Aves (birds), 10,000 species.[5]
Reptiles are tetrapod vertebrates, creatures that either have four limbs or, like snakes, are
descended from four-limbed ancestors. Unlike amphibians, reptiles do not have an aquatic larval
stage. Most reptiles are oviparous, although several species of squamates are viviparous, as were
some extinct aquatic clades[7]the fetus develops within the mother, contained in a placenta rather
than an eggshell. As amniotes, reptile eggs are surrounded by membranes for protection and
transport, which adapt them to reproduction on dry land. Many of the viviparous species feed
their fetuses through various forms of placenta analogous to those of mammals, with some providing
initial care for their hatchlings. Extant reptiles range in size from a tiny gecko, Sphaerodactylus
ariasae, which can grow up to 17 mm (0.7 in) to the saltwater crocodile, Crocodylus porosus, which
may reach 6 m (19.7 ft) in length and weigh over 1,000 kg (2,200 lb).

Contents
[hide]

1Classification
o 1.1Research history
o 1.2Phylogenetics and modern definition
o 1.3Taxonomy
o 1.4Phylogeny
o 1.5The position of turtles
2Evolutionary history
o 2.1Origin of the reptiles
o 2.2Rise of the reptiles
o 2.3Anapsids, synapsids, diapsids, and sauropsids
o 2.4Permian reptiles
o 2.5Mesozoic reptiles
o 2.6Cenozoic reptiles
 3Morphology and physiology
o 3.1Circulation
o 3.2Metabolism
o 3.3Respiratory system
3.3.1Turtles and tortoises
o 3.4Skin
o 3.5Excretion
o 3.6Digestion
o 3.7Nerves
3.7.1Intelligence
o 3.8Vision
o 3.9Reproduction
4Defense mechanisms
o 4.1Camouflage and warning
o 4.2Alternative defense in snakes
o 4.3Defense in crocodilians
o 4.4Shedding and regenerating tails
5Relations with humans
o 5.1In cultures and religions
o 5.2Medicine
o 5.3Other
6See also
7Further reading
8Notes
9References
10External links

Classification[edit]
See also: List of reptiles
Research history[edit]
See also: Skull roof
Reptiles, from Nouveau Larousse Illustr, 1897-1904: Notice the inclusion of amphibians (below the
crocodiles).

In the 18th century, the reptiles were, from the outset of classification, grouped with
the amphibians. Linnaeus, working from species-poor Sweden, where the common adder and grass
snake are often found hunting in water, included all reptiles and amphibians in class "III Amphibia"
in his Systema Natur.[8] The terms "reptile" and "amphibian" were largely interchangeable, "reptile"
(from Latin repere, "to creep") being preferred by the French.[9] 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.[10] Today, the two groups are still commonly treated under the same
heading as herptiles.

An "antediluvian monster", a Mosasaurus discovered in a Maastrichtlimestone quarry, 1770 (contemporary

engraving)

It was not until the beginning of the 19th century that it became clear that reptiles and amphibians
are, in fact, quite different animals, and Pierre Andr Latreille erected the class Batracia (1825) for
the latter, dividing the tetrapods into the four familiar classes of reptiles, amphibians, birds, and
mammals.[11] 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 Sauropsidaand Ichthyopsida for the latter two groups.[12] 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.[13]
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.[14]
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.[15] The animals singled
out by these formulations, the amniotes other than the mammals and the birds, are still those
considered reptiles today.[16]

The first reptiles had an anapsidtype 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 Osbornand elaborated and
made popular by Romer's classic Vertebrate Paleontology.[17][18] Those four subclasses were:

Anapsida no fenestrae cotylosaurs and Chelonia (turtles and relatives)[note 1]

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, 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
by 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.[19]
Phylogenetics and modern definition[edit]

Phylogenetic classifications group the traditional "mammal-like reptiles", like this Varanodon, with other
synapsids, not with extant reptiles.

By the early 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
are paraphyletic, since they exclude both birds and mammals. These respectively evolved from
dinosaurs and from early therapsids, which were both traditionally called reptiles.[20] Birds are more
closely related to crocodilians than the latter are to the rest of extant reptiles. Colin Tudge wrote:
Mammals are a clade, and therefore the cladists are happy to acknowledge the traditional
taxon Mammalia; and birds, too, are a clade, universally ascribed to the formal taxon Aves.
Mammalia and Aves are, in fact, subclades within the grand clade of the Amniota. But the traditional
class Reptilia is not a clade. It is just a section of the clade Amniota: the section that is left after the
Mammalia and Aves have been hived off. It cannot be defined by synapomorphies, as is the proper
way. Instead, it is 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'.[16]

Despite the early proposals for replacing the paraphyletic Reptilia with a monophyletic Sauropsida,
which includes birds, that term was never adopted widely or, when it was, was not applied
consistently.[1] 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.[1]
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 better known and more frequently used. Unlike most
previous definitions of Reptilia, however, Modesto and Anderson's definition includes birds,[1] as they
are within the clade that includes both lizards and crocodiles.
Taxonomy[edit]
See also: List of reptiles and List of snakes
Classification to order level of the reptiles, after Benton, 2014.[21][22]
 Class Reptilia
Subclass Parareptilia
Order Pareiasauromorpha
Subclass Eureptilia
Infraclass Diapsida
Order Younginiformes
Infraclass Neodiapsida
Order Testudinata (turtles)
Infraclass Lepidosauromorpha
Infrasubclass Unnamed
Infraclass Ichthyosauria
Order Thalattosauria
Superorder Lepidosauriformes
Order Rhynchocephalia (tuatara)
Order Squamata (lizards & snakes)
Infrasubclass Sauropterygia
Order Placodontia
Order Eosauropterygia
Order Plesiosauria

Mystriosuchus, a phytosaur.

Infraclass Archosauromorpha

Order Rhynchosauria
Order Protorosauria
Order Phytosauria
Division Archosauriformes
Subdivision Archosauria
Superorder Crocodylomorpha
Order Crocodilia
Infradivision Avemetatarsalia
Infrasubdivision Ornithodira

Order Pterosauria
Superorder Dinosauria
Order Saurischia (incl. Clade Aves)
Order Ornithischia
Phylogeny[edit]
The cladogram presented here illustrates the "family tree" of reptiles, and follows a simplified version
of the relationships found by M.S. Lee, in 2013.[23] All genetic studies have supported the hypothesis
that turtles are diapsids; some have placed turtles within archosauriformes,[23][24][25][26][27][28] though a few
have recovered turtles as lepidosauriformes instead.[29] The cladogram below used a combination of
genetic (molecular) and fossil (morphological) data to obtain its results.[23]
Amn
iota Synapsida (mammals and their extinct relatives)

Total group Parare

Reptilia ptilia

Millerettidae
unnamed
Eunotosaurus
Hallucicrania

Lanthanosuchidae
Procolophonia

Procolophonoidea

Pareiasauromorpha

Eureptil
ia
Captorhinidae
Rome
riida Paleothyris
Diap
sida

Araeoscelidia
Neodia
psida
Claudiosaurus

Younginiformes
Crown group Lepidosauro
Reptilia morpha

Kuehneosauridae

Lepidosa
uria Rhynchocephalia (tuatara and their
 extinct relatives)

Squamata (lizards and snakes)

Archosauro
morpha

Choristodera

Prolacertiformes

Trilophosaurus

Rhynchosauria

Archosauriformes (crocodiles, bir

ds, dinosaurs and extinct
relatives)

Pantestu
dines Eosauropterygia

Placodontia

Sinosaurosphargis

Odontochelys
Testud
inata Proganochelys

Testudines (turtles)

The position of turtles[edit]

The placement of turtles has historically been highly variable. Classically, turtles were considered to
be related to the primitive anapsid reptiles.[30] Molecular work has usually placed turtles within the
diapsids. So far three turtle genomes have been sequenced.[31] The results place turtles as a sister
clade to the archosaurs, the group that includes crocodiles, dinosaurs, and birds.[32]
Evolutionary history[edit]
Main article: Evolution of reptiles
Origin of the reptiles[edit]

An early reptile Hylonomus

Mesozoic scene showing typical reptilian megafauna: dinosaursincluding Europasaurus

holgeri, iguanodonts and Archaeopteryx lithographica perched on the foreground tree stump.

The origin of the reptiles lies about 310320 million years ago, in the steaming swamps of the
late Carboniferous period, when the first reptiles evolved from advanced reptiliomorphs.[3]
The oldest known animal that may have been an amniote is Casineria (though it may have been
a temnospondyl).[33][34][35] A series of footprints from the fossil strata of Nova Scotia dated
to 315 Ma show typical reptilian toes and imprints of scales.[36] These tracks are attributed
to Hylonomus, the oldest unquestionable reptile known.[37] It was a small, lizard-like animal, about 20
to 30 centimetres (7.9 to 11.8 in) long, with numerous sharp teeth indicating an insectivorous
diet.[38] Other examples include Westlothiana (for the moment considered a reptiliomorph rather than
a true amniote)[39] and Paleothyris, both of similar build and presumably similar habit.
Rise of the reptiles[edit]
The earliest amniotes, including stem-reptiles (those amniotes closer to modern reptiles than to
mammals), were largely overshadowed by larger stem-tetrapods, such as Cochleosaurus, and
remained a small, inconspicuous part of the fauna until the Carboniferous Rainforest
Collapse.[40] This sudden collapse affected several large groups. Primitive tetrapods were particularly
devastated, while stem-reptiles fared better, being ecologically adapted to the drier conditions that
followed. Primitive tetrapods, like modern amphibians, need to return to water to lay eggs; in
contrast, amniotes, like modern reptiles whose eggs possess a shell that allows them to be laid on
land were better adapted to the new conditions. Amniotes acquired new niches at a faster rate
than before the collapse and at a much faster rate than primitive tetrapods. They acquired new
feeding strategies including herbivory and carnivory, previously only having been insectivores and
piscivores.[40] From this point forward, reptiles dominated communities and had a greater diversity
than primitive tetrapods, setting the stage for the Mesozoic (known as the Age of Reptiles).[41] One of
the best known early stem-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
ancestors.[42] This type of skull has a skull roofwith only holes for the nostrils, eyes and a pineal
eye.[30] 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 the amniotes traditionally referred to as
"anapsids"), including lanthanosuchoids, millerettids, bolosaurids, some nycteroleterids,
some procolophonoids and at least some mesosaurs[43][44][45] made it more ambiguous and it's
currently uncertain whether the ancestral amniote had an anapsid-like or synapsid-like skull.[45] These
animals are traditionally referred to as "anapsids", and form a paraphyletic basic stock from which
other groups evolved.[1] Very shortly after the first amniotes appeared, a lineage
called Synapsida split off; this group was 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 amniotes", or stem-
mammals, that later gave rise to the true mammals.[46] Soon after, another group evolved a similar
trait, this time with a double opening behind each eye, earning them the name Diapsida ("two
arches").[42] 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.[30]
Turtles have been traditionally believed to be surviving parareptiles, on the basis of their anapsid
skull structure, which was assumed to be primitive trait.[47] The rationale for this classification has
been disputed, with some arguing that turtles are diapsids that evolved anapsid skulls in order to
improve their armor.[3] Later morphological phylogenetic studies with this in mind placed turtles firmly
within Diapsida.[48] All molecular studies have strongly upheld the placement of turtles within diapsids,
most commonly as a sister group to extant archosaurs.[25][26][27][28]
Permian reptiles[edit]
With the close of the Carboniferous, the amniotes became the dominant tetrapod fauna. While
primitive, terrestrial reptiliomorphs still existed, the synapsid amniotes 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.[49]
The parareptiles, whose massive skull roofs had no postorbital holes, continued and flourished
throughout the Permian. The pareiasaurian parareptiles reached giant proportions in the late
Permian, eventually disappearing at the close of the period (the turtles being possible survivors).[49]
Early in the period, the modern reptiles, or crown-group reptiles, evolved and split into two main
lineages: the Archosauromorpha (forebears of turtles, crocodiles, and dinosaurs) and
the Lepidosauromorpha (predecessors of modern lizards and tuataras). Both groups remained
lizard-like and relatively small and inconspicuous during the Permian.
Mesozoic reptiles[edit]
The close of the Permian saw the greatest mass extinction known (see the PermianTriassic
extinction event), an event prolonged by to the combination two or more distinct extinction
pulses.[50] Most of the earlier parareptile and synapsid megafauna disappeared, being replaced by
the true reptiles, particularly archosauromorphs. These 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.[50] Archosaurs
developed into the well-known dinosaursand pterosaurs, as well as the ancestors of crocodiles.
Since reptiles, first rauisuchians and then dinosaurs, dominated the Mesozoic era, the interval is
popularly known as the "Age of Reptiles". The dinosaurs also developed smaller forms, including the
feather-bearing smaller theropods. In the Cretaceous period, these gave rise to the first true birds.[51]
The sister group to Archosauromorpha is Lepidosauromorpha, containing lizards and tuataras, as
well as their fossil relatives. Lepidosauromorpha contained at least one major group of the Mesozoic
sea reptiles: the mosasaurs, which lived during the Cretaceous period. The phylogenetic placement
of other main groups of fossil sea reptiles the ichthyopterygians(including ichthyosaurs) and
the sauropterygians, which evolved in the early Triassic is more controversial. Different authors
linked these groups either to lepidosauromorphs[52] or to archosauromorphs,[53][54][55] and
ichthyopterygians were also argued to be diapsids that did not belong to the least inclusive clade
containing lepidosauromorphs and archosauromorphs.[56]
Cenozoic reptiles[edit]

Varanus priscus was a giant carnivorous goanna lizard, perhaps as long as 7 metres and weighing up to 1,940
kilograms.[57]

The close of the Cretaceous period saw the demise of the Mesozoic era reptilian megafauna (see
the CretaceousPaleogene 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 choristoderes survived the extinction, with the latter becoming extinct in the Miocene.[58] Of the
great host of dinosaurs dominating the Mesozoic, only the small beaked birds survived. This
dramatic extinction pattern at the end of the Mesozoic led into the Cenozoic. 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.[41] However, reptiles were still
important components of the megafauna, particularly in the form of giant tortoises.[59][60]
After the extinction of most archosaur and marine reptile lines by the end of the Cretaceous, reptile
diversification continued throughout the Cenozoic. Squamates took a massive hit during the KT-
event, only recovering ten million years after it,[61] but they underwent a great radiation event once
they recovered, and today squamates make up the majority of living reptiles (>
95%).[5][62] Approximately 10,000 extant species of traditional reptiles are known, with birds adding
about 10,000 more, almost twice the number of mammals, represented by about 5,700 living species
(excluding domesticated species).[63]

Morphology and physiology[edit]

Circulation[edit]

Thermographic image of monitor lizards

All squamates and turtles 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.[64]

A juvenile Iguana heart bisected through the ventricle, bisecting the left and right atrium.
For example, Iguana hearts, like the majority of the squamates hearts, are composed of three
chambers with two aorta and one ventricle, cardiac involuntary muscles.[65] The main structures of
the heart are the sinus venosus, the pacemaker, the left atrium, the right atruim, the atrioventriular
valve, the cavum venosum, cavum arteriosum, the cavum pulmonale, the muscular ridge, the
ventricular ridge, pulmanary veins, and paired aortic arches.[66]
Some squamate 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.[67]
Crocodilians have an anatomically four-chambered heart, similar to birds, but also have two
systemic aortas and are therefore capable of bypassing their pulmonary circulation.[68]
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.

Modern non-avian reptiles exhibit some form of cold-bloodedness (i.e. some mix
of poikilothermy, ectothermy, and bradymetabolism) so that they have limited physiological means of
keeping the body temperature constant and often rely on external sources of heat. Due to a less
stable core temperature than birds and mammals, reptilian biochemistry requires enzymes capable
of maintaining efficiency over a greater range of temperatures than in the case for warm-
blooded animals. The optimum body temperature range varies with species, but is typically below
that of warm-blooded animals; for many lizards, it falls in the 2435 C (7595 F) range,[69] 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).[70] While the optimum temperature is often encountered when the animal is active, the low
basal metabolism makes body temperature drop rapidly when the animal is inactive.
As in all animals, reptilian muscle action produces heat. In large reptiles, like leatherback turtles, the
low surface-to-volume ratio allows this metabolically produced heat to keep the animals warmer than
their environment even though they do not have a warm-bloodedmetabolism.[71] This form of
homeothermy is called gigantothermy; it has been suggested as having been common in
large dinosaurs and other extinct large-bodied reptiles.[72][73]
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.[74] 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.[75] 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.[76]Higher energetic capacity might have been
responsible for the evolution of warm-bloodedness in birds and mammals.[77] However, investigation
of correlations between active capacity and thermophysiology show a weak relationship.[78] Most
extant reptiles are carnivores with a sit-and-wait feeding strategy; whether reptiles are cold blooded
due to their ecology is not clear. Energetic studies on some reptiles have shown active capacities
equal to or greater than similar sized warm-blooded animals.[79]
Respiratory system[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.[80] 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.[81]
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
airways bronchia form a number of double tubular chambers within each lung. On inhalation and
exhalation air moves through the airways in the same direction, thus creating a unidirectional airflow
through the lungs. A similar system is found in birds,[82] monitor lizards[83] and iguanas.[84]
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.[citation needed]
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 those turtles breathe. The varied 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).[85]
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.[86] 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.[86]
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.[87] 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[citation needed]. The scales found in turtles and crocodiles are of dermal, rather than
epidermal, origin and are properly termed scutes[citation needed]. 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.
Shedding. Reptiles shed their skin through a process called ecdysis which occurs continuously
throughout their lifetime. In particular, younger reptiles tend to shed once every 56 weeks while
adults shed 3-4 times a year.[88] O Younger reptiles shed more because of their rapid growth rate.
Once full size, the frequency of shedding drastically decreases. The process of ecdysis involves
forming a new layer of skin under the old one. Proteolytic enzymes and lymphatic fluid is secreted
between the old and new layers of skin. Consequently, this lifts the old skin from the new one
allowing for shedding to occur.[89] Snakes will shed from the head to the tail while lizards shed in a
patchy pattern.[89] Dysecdysis, a common skin disease in snakes and lizards will occur when
ecdysis, or shedding, fails.[90] There are numerous reasons why shedding fails and can be related to
inadequate humidity and temperature, nutritional deficiencies, dehydration and traumatic
injuries.[89] Nutritional deficiencies decreases proteolytic enzymes while dehydration reduces
lymphatic fluids to separate the skin layers. Traumatic injuries on the other hand, forms scars that
will not allow new scales to form and disrupt the process of ecdysis.[90]
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]

A colubrid snake, Dolichophis jugularis, eating a legless lizard, Pseudopus apodus. Most reptiles are
carnivorous, and many primarily eat other reptiles.

Gastroliths 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.[91] 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.[75]
While modern reptiles are predominantly carnivorous, during the early history of reptiles several
groups produced some herbivorous megafauna: in the Paleozoic, the pareiasaurs; and in
the Mesozoic several lines of dinosaurs.[41] 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.[92]
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.[92] 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.[93][94] Salt water
crocodiles also use gastroliths as ballast, stabilizing them in the water or helping them to dive.[95] A
dual function as both stabilizing ballast and digestion aid has been suggested for gastroliths found
in plesiosaurs.[96]
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.[97] Due to their short cochlea, reptiles use electrical
tuning to expand their range of audible frequencies.
Intelligence[edit]
See also: Animal cognition
Reptiles are generally considered less intelligent than mammals and birds.[30] 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,[98] though larger reptiles can show more complex brain development.
Larger lizards, like the monitors, are known to exhibit complex behavior, including
cooperation.[99] Crocodiles have relatively larger brains and show a fairly complex social structure.
The Komodo dragon is even known to engage in play,[100] as are turtles, which are also considered to
be social creatures[citation needed] and sometimes switch between monogamy and promiscuity in their
sexual behavior.[citation needed] One study found that wood turtles were better than white rats at learning to
navigate mazes.[101]
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]
Crocodilian egg diagram
1. eggshell, 2. yolk sac, 3. yolk (nutrients), 4. vessels, 5. amnion, 6. chorion, 7. air space, 8. allantois, 9.
albumin (egg white), 10. amniotic sac, 11. crocodile embryo, 12. amniotic fluid

Most reptiles reproduce sexually, for example this Trachylepis maculilabrisskink

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, only one of which is
typically used in each session. Tuatara, however, lack copulatory organs, and so the male and
female simply press their cloacas together as the male discharges sperm.[102]
Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion, chorion,
and allantois are present during embryoniclife. The eggshell (1) protects the crocodile embryo (11)
and keeps it from drying out, but it is flexible to allow gas exchange. The chorion (6) aids in gas
exchange between the inside and outside of the egg. It allows carbon dioxide to exit the egg and
oxygen gas to enter the egg. The albumin (9) further protects the embryo and serves as a reservoir
for water and protein. The allantois (8) is a sac that collects the metabolic waste produced by the
embryo. The amniotic sac (10) contains amniotic fluid (12) which protects and cushions the embryo.
The amnion (5) aids in osmoregulation and serves as a saltwater reservoir. The yolk sac (2)
surrounding the yolk (3) contains protein and fat rich nutrients that are absorbed by the embryo via
vessels (4) that allow the embryo to grow and metabolize. The air space (7) provides the embryo
with oxygen while it is hatching. This ensures that the embryo will not suffocate while it is hatching.
There are no larval stages of development. Viviparity and ovoviviparity have evolved in many extinct
clades of reptiles and in squamates. In the latter group, many species, including all boas and most
vipers, utilize this mode of reproduction. The degree of viviparity varies; some species simply retain
the eggs until just before hatching, others provide maternal nourishment to supplement the yolk, and
yet others lack any yolk and provide all nutrients via a structure similar to the mammalian placenta.
The earliest documented case of viviparity in reptiles is the Early Permianmesosaurs,[103] 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.[7]
Asexual reproduction has been identified in squamates in six families of lizards and one snake. In
some species of squamates, a population of females is able to produce a unisexual diploid clone of
the mother. This form of asexual reproduction, called parthenogenesis, occurs in several species
of gecko, and is particularly widespread in the teiids (especially Aspidocelis) and lacertids (Lacerta).
In captivity, Komodo dragons (Varanidae) have reproduced by parthenogenesis.
Parthenogenetic species are suspected to occur among chameleons, agamids, xantusiids,
and typhlopids.
Some reptiles exhibit temperature-dependent sex determination (TDSD), in which the incubation
temperature determines whether a particular egg hatches as male or female. TDSD is most common
in turtles and crocodiles, but also occurs in lizards and tuatara.[104] To date, there has been no
confirmation of whether TDSD occurs in snakes.[105]

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.[106] 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 and warning[edit]

A camouflaged Phelsuma deubiaon 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 color vision. Thus the skins of many
reptiles have cryptic coloration of plain or mottled gray, green, and brown to allow them to blend into
the background of their natural environment.[107] Aided by the reptiles' capacity for remaining
motionless for long periods, the camouflage of many snakes is so effective that people or domestic
animals are most typically bitten because they accidentally step on them.[108]
When camouflage fails to protect them, blue-tongued skinks will try to ward off attackers by
displaying their blue tongues, and the frill-necked lizard will display its brightly colored frill. These
same displays are used in territorial disputes and during courtship.[109] If danger arises so suddenly
that flight is useless, crocodiles, turtles, some lizards, and some snakes hiss loudly when confronted
by an enemy. Rattlesnakesrapidly vibrate the tip of the tail, which is composed of a series of nested,
hollow beads to ward of approaching danger.
In contrast to the normal drab coloration of most reptiles, the lizards of the
genus Heloderma (the Gila monster and the beaded lizard) and many of the coral snakes have high-
contrast warning coloration, warning potential predators they are venomous.[110] A number of non-
venomous North American snake species have colorful markings similar to those of the coral snake,
an oft cited example of Batesian mimicry.[111][112]
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 American hognose snakes 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.[113] The main weapon in all crocodiles is the bite, which can
generate very high bite force. Many species also possess canine-like teeth. These are used primarily
for seizing prey, but are also used in fighting and display.[114]
Shedding and regenerating tails[edit]
Main article: Autotomy
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 detached tails of leopard geckos can wiggle for up to 20 minutes.[115] In
many species the tails are of a separate and dramatically more intense color than the rest of the
body so as to encourage potential predators to strike for the tail first. In the shingleback skink and
some species of geckos, the tail is short and broad and resembles the head, so that the predators
may attack it rather than the more vulnerable front part.[116]
Reptiles that are capable of shedding their tails can partially regenerate them over a period of
weeks. The new section will however contain cartilage rather than bone, and will never grow to the
same length as the original tail. It is often also distinctly discolored compared to the rest of the body
and may lack some of the external sculpting features seen in the original tail.[117]

Relations with humans[edit]

Main article: Reptiles in culture
In cultures and religions[edit]
Main article: Reptiles in culture

The 1897 painting of fighting "Laelaps" (now Dryptosaurus) by Charles R. Knight

Dinosaurs have been widely depicted in culture since the English palaeontologist Richard
Owen coined the name dinosaur in 1842. As soon as 1854, the Crystal Palace Dinosaurs were on
display to the public in south London.[118][119] One dinosaur appeared in literature even earlier,
as Charles Dickens placed a Megalosaurus in the first chapter of his novel Bleak House in
1852.[120] The dinosaurs featured in books, films, television programs, artwork, and other media have
been used for both education and entertainment. The depictions range from the realistic, as in the
television documentaries of the 1990s and first decade of the 21st century, or the fantastic, as in
the monster movies of the 1950s and 1960s.[119][121][122]
The snake or serpent has played a powerful symbolic role in different cultures. In Egyptian history,
the Nile cobra adorned the crown of the pharaoh. It was worshipped as one of the gods and was
also used for sinister purposes: murder of an adversary and ritual suicide (Cleopatra). In Greek
mythology snakes are associated with deadly antagonists, as a chthonic symbol, roughly translated
as earthbound. The nine-headed Lernaean Hydra that Hercules defeated and the
three Gorgon sisters are children of Gaia, the earth. Medusa was one of the three Gorgon sisters
who Perseus defeated. Medusa is described as a hideous mortal, with snakes instead of hair and
the power to turn men to stone with her gaze. After killing her, Perseus gave her head to Athena who
fixed it to her shield called the Aegis. The Titans are depicted in art with their legs replaced by
bodies of snakes for the same reason: They are children of Gaia, so they are bound to the
earth.[123] In Hinduism, snakes are worshipped as gods, with many women pouring milk on snake
pits. The cobra is seen on the neck of Shiva, while Vishnu is depicted often as sleeping on a seven-
headed snake or within the coils of a serpent. There are temples in India solely for cobras
sometimes called Nagraj (King of Snakes), and it is believed that snakes are symbols of fertility. In
the annual Hindu festival of Nag Panchami, snakes are venerated and prayed to.[124] In religious
terms, the snake and jaguar are arguably the most important animals in ancient Mesoamerica. "In
states of ecstasy, lords dance a serpent dance; great descending snakes adorn and support
buildings from Chichen Itza to Tenochtitlan, and the Nahuatl word coatl meaning serpent or twin,
forms part of primary deities such as Mixcoatl, Quetzalcoatl, and Coatlicue."[125] In Christianity and
Judaism, a serpent appears in Genesis to tempt Adam and Eve with the forbidden fruit from the Tree
of Knowledge of Good and Evil.[126]
The turtle has a prominent position as a symbol of steadfastness and tranquility in religion,
mythology, and folklore from around the world.[127] A tortoise's longevity is suggested by its long
lifespan and its shell, which was thought to protect it from any foe.[128] In the cosmological myths of
several cultures a World Turtle carries the world upon its back or supports the heavens.[129]
Medicine[edit]

The Rod of Asclepiussymbolizes medicine

Deaths from snakebites are uncommon in many parts of the world, but are still counted in tens of
thousands per year in India.[130] Snakebite can be treated with antivenom made from the venom of
the snake. To produce antivenom, a mixture of the venoms of different species of snake is injected
into the body of a horse in ever-increasing dosages until the horse is immunized. Blood is then
extracted; the serum is separated, purified and freeze-dried.[131] The cytotoxic effect of snake venom
is being researched as a potential treatment for cancers.[132]
Geckos have also been used as medicine, especially in China.[133]
Other[edit]
Further information: Crocodilia Interactions with humans
Crocodiles are protected in many parts of the world, and are farmed commercially. Their hides are
tanned and used to make leather goods such as shoes and handbags; crocodile meat is also
considered a delicacy.[134] The most commonly farmed species are the saltwater and Nile crocodiles.
Farming has resulted in an increase in the saltwater crocodile population in Australia, as eggs are
usually harvested from the wild, so landowners have an incentive to conserve their habitat. Crocodile
leather is made into wallets, briefcases, purses, handbags, belts, hats, and shoes. Crocodile oil has
been used for various purposes.[135]
In the Western world, some snakes (especially docile species such as the ball python and corn
snake) are kept as pets.[136]

See also[edit]
Evolution of reptiles
Herpetophobia
List of reptiles
Lists of reptiles by region
List of threatened reptiles and amphibians of the United States
Ophidiophobia
 Reptile Database

Further reading[edit]
Colbert, Edwin H. (1969). Evolution of the Vertebrates (2nd ed.). New York: John Wiley and Sons
Inc. ISBN 978-0-471-16466-1.
Landberg, Tobias; Mailhot, Jeffrey; Brainerd, Elizabeth (2003). "Lung ventilation during treadmill
locomotion in a terrestrial turtle, Terrapene carolina". Journal of Experimental Biology. 206 (19): 3391
3404. PMID 12939371. doi:10.1242/jeb.00553.
Laurin, Michel and Gauthier, Jacques A.: Diapsida. Lizards, Sphenodon, crocodylians, birds, and their
extinct relatives, Version 22 June 2000; part of The Tree of Life Web Project
Pianka, Eric; Vitt, Laurie (2003). Lizards Windows to the Evolution of Diversity. University of California
Press. pp. 116118. ISBN 978-0-520-23401-7.
Pough, Harvey; Janis, Christine; Heiser, John (2005). Vertebrate Life. Pearson Prentice Hall. ISBN 978-0-
13-145310-4.

Notes[edit]
1. Jump up^ This taxonomy does not reflect modern molecular evidence, which places turtles within
Diapsida.

References[edit]
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