Reptiles are a group (Reptilia) of tetrapod animals comprising today's turtles,

crocodilians, snakes, lizards, tuatara, and their extinct relatives. The study o
f these traditional reptile groups, historically combined with that of modern am
phibians, is called herpetology. Because crocodilians are more closely related t
o birds than to any other group of reptiles, birds are also often included as a
sub-group of reptiles by modern scientists.[1]
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 includ
e the lizard-like Hylonomus and Casineria. In addition to the living reptiles, t
here are many diverse groups that are now extinct, in some cases due to mass ext
inction events. In particular, the K Pg extinction wiped out the pterosaurs, plesi
osaurs, ornithischians, and sauropods, as well as many species of theropods (e.g
. tyrannosaurs and dromaeosaurids), crocodyliforms, and squamates (e.g. mosasaur
ids).
Modern reptiles inhabit every continent with the exception of Antarctica. Severa
l living subgroups are recognized:
Testudines (turtles, terrapins and tortoises): approximately 400 species[2]
Sphenodontia (tuatara from New Zealand): 2 species[2]
Squamata (lizards, snakes, and worm lizards): over 9,600 species[2]
Crocodilia (crocodiles, gavials, caimans, and alligators): 25 species[2]
Because some reptiles are more closely related to birds than they are to other r
eptiles (crocodiles are more closely related to birds than they are to lizards),
?many modern scientists prefer to make Reptilia a monophyletic grouping and so
also include the birds, which today contain over 10,000 species.[1][3][4][5]
Reptiles are tetrapod vertebrates, creatures that either have four limbs or, lik
e snakes, are descended from four-limbed ancestors. Unlike amphibians, reptiles
do not have an aquatic larval stage. Most reptiles are oviparous, although sever
al species of squamates are viviparous, as were some extinct aquatic clades[6]? ?t
he fetus develops within the mother, contained in a placenta rather than an eggs
hell. As amniotes, reptile eggs are surrounded by membranes for protection and t
ransport, which adapt them to reproduction on dry land. Many of the viviparous s
pecies feed their fetuses through various forms of placenta analogous to those o
f mammals, with some providing initial care for their hatchlings. Extant reptile
s 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
1 Classification
1.1 History of classification
1.1.1 Linnaeus and the 18th century
1.1.2 "Antediluvian monsters"
1.1.3 Skull openings in 20th-century classification
1.1.4 Phylogenetics and modern definition
1.2 Taxonomy
1.3 Phylogeny
1.4 The position of turtles
2 Evolutionary history
2.1 Origin of the reptiles
2.2 Rise of the reptiles
2.3 Anapsids, synapsids, diapsids and sauropsids
2.4 Permian reptiles
2.5 The Mesozoic era

2.6 Cenozoic era

3 Morphology and Physiology
3.1 Circulation
3.2 Metabolism
3.3 Respiration
3.3.1 Reptilian lungs
3.3.2 Turtles and tortoises
3.3.3 Palate
3.4 Skin
3.5 Excretion
3.6 Digestion
3.7 Nerves
3.7.1 Intelligence
3.8 Vision
3.9 Reproduction
4 Defense mechanisms
4.1 Camouflage and warning
4.2 Alternative defense in snakes
4.3 Defense in crocodilians
4.4 Shedding and regenerating tails
5 Reptiles in human culture
6 See also
7 References
8 Notes
9 Further reading
10 External links
Classification
See also: List of reptiles
History of classification
Reptiles, from Nouveau Larousse Illustr, 1897-1904: Notice the inclusion of amphi
bians (below the crocodiles).
Linnaeus and the 18th century
The reptiles were, from the outset of classification, grouped with the amphibian
s. 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
Amphibia" in his Systema Natur.[7] The terms "reptile" and "amphibian"
class "III
were largely interchangeable, "reptile" (from Latin repere, "to creep") being p
referred 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 L
innaeus.[9] Today, the two groups are still commonly treated under the same head
ing as herptiles.
"Antediluvian monsters"
An "antediluvian monster", a Mosasaurus discovered in a Maastricht limestone qua
rry, 1770 (contemporary engraving)
It was not until the beginning of the 19th century that it became clear that rep
tiles and amphibians are, in fact, quite different animals, and Pierre Andr Latre
ille erected the class Batracia (1825) for the latter, dividing the tetrapods in
to the four familiar classes of reptiles, amphibians, birds, and mammals.[10]
The British anatomist Thomas Henry Huxley made Latreille's definition popular an
d, together with Richard Owen, expanded Reptilia to include the various fossil "
antediluvian monsters", including dinosaurs and the mammal-like (synapsid) Dicyn
odon he helped describe. This was not the only possible classification scheme: I
n the Hunterian lectures delivered at the Royal College of Surgeons in 1863, Hux
ley grouped the vertebrates into mammals, sauroids, and ichthyoids (the latter c

ontaining the fishes and amphibians). He subsequently proposed the names of Saur
opsida and Ichthyopsida for the latter two groups.[11]
In 1866, Haeckel demonstrated that vertebrates could be divided based on their r
eproductive strategies, and that reptiles, birds, and mammals were united by the
amniotic egg.
The terms "Sauropsida" ("lizard faces") and "Theropsida" ("beast faces") were us
ed again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and the
ir relatives on the one hand (Sauropsida) and mammals and their extinct relative
s (Theropsida) on the other. Goodrich supported this division by the nature of t
he hearts and blood vessels in each group, and other features, such as the struc
ture of the forebrain. According to Goodrich, both lineages evolved from an earl
ier 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) includes all amniotes which are not
mammals or birds, making it a paraphyletic group.[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 r
einterpreted Sauropsida and Theropsida to exclude birds and mammals, respectivel
y. Thus his Sauropsida included Procolophonia, Eosuchia, Millerosauria, Chelonia
(turtles), Squamata (lizards and snakes), Rhynchocephalia, Crocodilia, "thecodo
nts" (paraphyletic basal Archosauria), non-avian dinosaurs, pterosaurs, ichthyos
aurs, 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 condy
le, a jaw joint formed by the quadrate and articular bones, and certain characte
ristics of the vertebrae.[14] The animals singled out by these formulations, the
amniotes other than the mammals and the birds, are still those considered repti
les today.[15]
Skull openings in 20th-century classification
Main article: Skull roof
The first reptiles had an anapsid type of skull roof, as seen in the Permian gen
us Captorhinus
The synapsid/sauropsid division supplemented another approach, one that split th
e reptiles into four subclasses based on the number and position of temporal fen
estrae, 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.[16][17] 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 rept
iles')
Euryapsida one high fenestra (above the postorbital and squamosal)
protorosa
urs (small, early lizard-like reptiles) and the marine sauropterygians and ichth
yosaurs, the latter called Parapsida in Osborn's work.
Diapsida two fenestrae
most reptiles, including lizards, snakes, crocodilian
s, dinosaurs and pterosaurs
The composition of Euryapsida was uncertain. Ichthyosaurs were, at times, consid
ered to have arisen independently of the other euryapsids, and given the older n
ame Parapsida. Parapsida was later discarded as a group for the most part (ichth
yosaurs being classified as incertae sedis or with Euryapsida). However, four (o
r three if Euryapsida is sunk into Diapsida) subclasses remained more or less un
iversal for non-specialist work throughout the 20th century. It has largely been
abandoned among recent researchers: in particular, the anapsid condition has be

en found to occur so variably among unrelated groups that it is not now consider
ed a useful distinction.[18]
Phylogenetics and modern definition
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 ph
ylogenetic taxonomy, in which all groups are defined in such a way as to be mono
phyletic; 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 therapsid
s, which were both traditionally called reptiles.[19] Birds are more closely rel
ated to crocodilians than the latter are to the rest of extant reptiles. Colin T
udge 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 t
o the formal taxon Aves. Mammalia and Aves are, in fact, subclades within the gr
and 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 Mamma
lia 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 a
nd the features it lacks: reptiles are the amniotes that lack fur or feathers. A
t best, the cladists suggest, we could say that the traditional Reptilia are 'no
n-avian, non-mammalian amniotes'.[15]
Despite the early proposals for replacing the paraphyletic Reptilia with a monop
hyletic Sauropsida which includes birds, that term was never adopted widely or,
when it was, 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 g
roup containing turtles, lizards and snakes, crocodilians, and birds, their comm
on ancestor and all its descendants. Because the actual relationship of turtles
to other reptiles was not yet well understood at this time, Gauthier's definitio
n came to be considered inadequate.[1]
A variety of other definitions were proposed by other scientists in the years fo
llowing Gauthier's paper. The first such new definition, which attempted to adhe
re to the standards of the PhyloCode, was published by Modesto and Anderson in 2
004. Modesto and Anderson reviewed the many previous definitions and proposed a
modified definition, which they intended to retain most traditional content of t
he 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 Sauro
psida, which Modesto and Anderson synonymized with Reptilia, since the latter is
more well known and more frequently used. Unlike most previous definitions of R
eptilia, however, Modesto and Anderson's definition includes birds,[1] as there
is no other way to establish a monophyletic clade that includes both lizards and
crocodiles.
Taxonomy
Classification to order level of the reptiles, after Benton, 2014.[20][21]
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
Vjushkovia, a basal archosauropmorph
Infraclass Archosauromorpha
Order Rhynchosauria
Order Protorosauria
Division Archosauriformes
Subdivision Archosauria
Infradivision Crurotarsi
Order Phytosauria
Superorder Crocodylomorpha
Order Crocodilia
Infradivision Avemetatarsalia
Infrasubdivision Ornithodira
Order Pterosauria
Superorder Dinosauria
Order Saurischia (incl. Class Aves)
Order Ornithischia
Phylogeny
The cladogram presented here illustrates the "family tree" of reptiles, and foll
ows a simplified version of the relationships found by M.S. Lee, in 2013.[22] Al
l genetic studies have supported the hypothesis that turtles are diapsids; some
have placed turtles within archosauriformes,[22][23][24][25][26][27] though a fe
w have recovered turtles as lepidosauriformes instead.[28] The cladogram below u
sed a combination of genetic (molecular) and fossil (morphological) data to obta
in its results.[22]
Amniota

Synapsida (mammals and their extinct relatives) Varanops brevirostris.jpg

Total group Reptilia
unnamed
Parareptilia

Millerettidae Milleretta BW.jpg

unnamed

Eunotosaurus
Hallucicrania

Lanthanosuchidae Lanthanosuchus NT.jpg

Procolophonia

Procolophonoidea Sclerosaurus1DB.jpg

Pareiasauromorpha Scutosaurus BW.jpg

Eureptilia

Captorhinidae Labidosaurus.jpg
Romeriida

Paleothyris
Diapsida

Araeoscelidia Spinoaequalis schultzei reconstruction.jpg

Neodiapsida

Claudiosaurus

Younginiformes Hovasaurus BW.jpg

Crown group Reptilia
Lepidosauromorpha

Kuehneosauridae
Lepidosauria

Rhynchocephalia (tuatara and their extinct relatives) Sphenodon punctatus in Wai

kanae, New Zealand.jpg

Squamata (lizards and snakes) Douthat State Park - Eastern fence lizard - 08.jpg

Archosauromorpha

Choristodera Champsosaurus BW.jpg

Prolacertiformes

Trilophosaurus Trilophosaurus BW.jpg

Rhynchosauria Paradapedon 1DB.jpg

Archosauriformes (crocodiles, birds, dinosaurs and extinct relatives) Prestosuch

us11DB.jpgGoland brun (larus fuscus) dtoure.jpg

Pantestudines

Eosauropterygia Thalassomedon BW.jpg

Placodontia Psephoderma BW.jpg

Sinosaurosphargis

Odontochelys
Testudinata

Proganochelys

Testudines (turtles) Florida Box Turtle Digon3 re-edited.jpg

The position of turtles

The placement of turtles has historically been highly variable. Classically, tur
tles were considered to be related to the primitive anapsid reptiles.[29] Molecu
lar work has usually placed turtles within the diapsids. So far three turtle gen
omes have been sequenced.[30] The results place turtles as a sister clade to the
archosaurs, the group that includes crocodiles, dinosaurs, and birds.
Evolutionary history
Main article: Evolution of reptiles
Origin of the reptiles
An early reptile Hylonomus
Mesozoic scene showing typical reptilian megafauna: dinosaurs including Europasa
urus holgeri, iguanodonts and Archaeopteryx lithographica perched on the foregro
und tree stump.
The origin of the reptiles lies about 310 320 million years ago, in the steaming s

wamps of the late Carboniferous period, when the first reptiles evolved from adv
anced reptiliomorphs.[31]
The oldest known animal that may have been an amniote is Casineria (though it ma
y have been a temnospondyl).[32][33][34] A series of footprints from the fossil
strata of Nova Scotia dated to 315 Ma show typical reptilian toes and imprints o
f scales.[35] These tracks are attributed to Hylonomus, the oldest unquestionabl
e reptile known.[36] It was a small, lizard-like animal, about 20 to 30 centimet
res (7.9 to 11.8 in) long, with numerous sharp teeth indicating an insectivorous
diet.[37] Other examples include Westlothiana (for the moment considered a rept
iliomorph rather than a true amniote)[38] and Paleothyris, both of similar build
and presumably similar habit.
Rise of the reptiles
The earliest amniotes, including stem-reptiles (those amniotes closer to modern
reptiles than to mammals), were largely overshadowed by larger stem-tetrapods su
ch as Cochleosaurus, and remained a small, inconspicuous part of the fauna until
the Carboniferous Rainforest Collapse.[39] This sudden collapse affected severa
l large groups. Primitive tetrapods were particularly devastated, while stem-rep
tiles fared better, being ecologically adapted to the drier conditions that foll
owed. Primitive tetrapods, like modern amphibians, need to return to water to la
y eggs; in contrast, amniotes, like modern reptiles
whose eggs possess a shell t
hat allows them to be laid on land were better adapted to the new conditions. Am
niotes acquired new niches at a faster rate than before the collapse and at a mu
ch faster rate than primitive tetrapods. They acquired new feeding strategies in
cluding herbivory and carnivory, previously only having been insectivores and pi
scivores.[39] 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).[40] One of the best known early stem-reptiles is
Mesosaurus, a genus from the early Permian that had returned to water, feeding o
n fish.
Anapsids, synapsids, diapsids and sauropsids
A = Anapsid, B = Synapsid, C = Diapsid
It was traditionally assumed that the first reptiles retained an anapsid skull i
nherited from their ancestors.[41] This type of skull has a skull roof with only
holes for the nostrils, eyes and a pineal eye.[29] The discoveries of synapsidlike openings (see below) in the skull roof of the skulls of several members of
Parareptilia (the clade containing most of the amniotes traditionally referred t
o as "anapsids"), including lanthanosuchoids, millerettids, bolosaurids, some ny
cteroleterids, some procolophonoids and at least some mesosaurs[42][43][44] made
it more ambiguous and it's currently uncertain whether the ancestral amniote ha
d an anapsid-like or synapsid-like skull.[44] These animals are traditionally re
ferred to as "anapsids", and form a paraphyletic basic stock from which other gr
oups evolved.[1] Very shortly after the first amniotes appeared, a lineage calle
d Synapsida split off, characterized by a temporal opening in the skull behind e
ach eye to give room for the jaw muscle to move. These are the "mammal-like amni
otes", or stem-mammals, that later gave rise to the true mammals.[45] Soon after
, another group evolved a similar trait, this time with a double opening behind
each eye, earning them the name Diapsida ("two arches").[41] The function of the
holes in these groups was to lighten the skull and give room for the jaw muscle
s to move, allowing for a more powerful bite.[29]
Turtles have been traditionally believed to be surviving parareptiles, on the ba
sis of their anapsid skull structure, which was assumed to be primitive trait.[4
6] The rationale for this classification has been disputed, with some arguing th
at turtles are diapsids that evolved anapsid skulls in order to improve their ar
mor.[31] Later morphological phylogenetic studies with this in mind placed turtl
es firmly within Diapsida.[47] All molecular studies have strongly upheld the pl
acement of turtles within diapsids, most commonly as a sister group to extant ar

chosaurs.[24][25][26][27]
Permian reptiles
With the close of the Carboniferous, the amniotes became the dominant tetrapod f
auna. While primitive, terrestrial reptiliomorphs still existed, the synapsid am
niotes 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 p
elycosaurs were replaced by the therapsids.[48]
The parareptiles, whose massive skull roofs had no postorbital holes, continued
and flourished throughout the Permian. The pareiasaurian parareptiles reached gi
ant proportions in the late Permian, eventually disappearing at the close of the
period (the turtles being possible survivors).[48]
Early in the period, the modern reptiles, or crown-group reptiles, evolved and s
plit into two main lineages: the Archosauromorpha (forebears of turtles, crocodi
les, and dinosaurs) and the Lepidosauromorpha (predecessors of modern lizards an
d tuataras). Both groups remained lizard-like and relatively small and inconspic
uous during the Permian.
The Mesozoic era
The close of the Permian saw the greatest mass extinction known (see the Permian T
riassic extinction event), a prolonged event due to the accumulation of at least
two distinct extinction pulses.[49] Most of the earlier parareptile and synapsi
d megafauna disappeared, being replaced by the true reptiles, particularly archo
sauromorphs. These were characterized by elongated hind legs and an erect pose,
the early forms looking somewhat like long-legged crocodiles. The archosaurs bec
ame the dominant group during the Triassic period, though it took 30 million yea
rs before their diversity was as great as the animals that lived in the Permian.
[49] Archosaurs developed into the well-known dinosaurs and pterosaurs, as well
as the ancestors of crocodiles. Since reptiles, first rauisuchians and then dino
saurs, dominated the Mesozoic era, the interval is popularly known as the "Age o
f Reptiles". The dinosaurs also developed smaller forms, including the feather-b
earing smaller theropods. In the Cretaceous period, these gave rise to the first
true birds.[50]
The sister group to Archosauromorpha is Lepidosauromorpha, containing lizards an
d tuataras, as well as their fossil relatives. Lepidosauromorpha contained at le
ast one major group of the Mesozoic sea reptiles: the mosasaurs, which lived dur
ing the Cretaceous period. The phylogenetic placement of other main groups of fo
ssil sea reptiles the ichthyopterygians (including ichthyosaurs) and the sauropt
erygians, which evolved in the early Triassic
is more controversial. Different a
uthors linked these groups either to lepidosauromorphs[51] or to archosauromorph
s,[52][53][54] and ichthyopterygians were also argued to be diapsids that did no
t belong to the least inclusive clade containing lepidosauromorphs and archosaur
omorphs.[55]
Cenozoic era
Varanus priscus was a giant carnivorous goanna lizard, perhaps as long as 7 metr
es and weighing up to 1,940 kilograms.[56]
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 rep
tiles, only sea turtles were left; and of the non-marine large reptiles, only th
e semi-aquatic crocodiles and broadly similar choristoderes survived the extinct
ion, with the latter becoming extinct in the Miocene.[57] Of the great host of d
inosaurs dominating the Mesozoic, only the small beaked birds survived. This dra
matic extinction pattern at the end of the Mesozoic led into the Cenozoic. Mamma
ls and birds filled the empty niches left behind by the reptilian megafauna and,
while reptile diversification slowed, bird and mammal diversification took an e

xponential turn.[40] However, reptiles were still important components of the me

gafauna, particularly in the form of giant tortoises.[58][59]
After the extinction of most archosaur and marine reptile lines by the end of th
e Cretaceous, reptile diversification continued throughout the Cenozoic. Squamat
es took a massive hit during the KT-event, only recovering ten million years aft
er it,[60] but they underwent a great radiation event once they recovered, and t
oday squamates make up the majority of living reptiles (> 95%).[2][61] Approxima
tely 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 5,70
0 living species (excluding domesticated species).[62]
Morphology and Physiology
Circulation
Thermographic image of a monitor lizard
Many squamates and turtles have a three-chambered heart consisting of two atria,
one variably partitioned ventricle, and two aortas that lead to the systemic ci
rculation. The degree of mixing of oxygenated and deoxygenated blood in the thre
e-chambered heart varies depending on the species and physiological state. Under
different conditions, deoxygenated blood can be shunted back to the body or oxy
genated blood can be shunted back to the lungs. This variation in blood flow has
been hypothesized to allow more effective thermoregulation and longer diving ti
mes for aquatic species, but has not been shown to be a fitness advantage.[63]
Crocodilians have an anatomically four-chambered heart, similar to birds, but al
so have two systemic aortas and are therefore capable of bypassing only their pu
lmonary circulation.[64] Also, some snake and lizard species (e.g., pythons and
monitor lizards) have three-chambered hearts that become functionally four-chamb
ered hearts during contraction. This is made possible by a muscular ridge that s
ubdivides the ventricle during ventricular diastole and completely divides it du
ring ventricular systole. Because of this ridge, some of these squamates are cap
able of producing ventricular pressure differentials that are equivalent to thos
e seen in mammalian and avian hearts.[65]
Metabolism
Sustained energy output (joules) of a typical reptile versus a similar size mamm
al as a function of core body temperature. The mammal has a much higher peak out
put, but can only function over a very narrow range of body temperature.
Modern reptiles exhibit some form of cold-bloodedness (i.e. some mix of poikilot
hermy, ectothermy, and bradymetabolism) so that they have limited physiological
means of keeping the body temperature constant and often rely on external source
s of heat. Due to a less stable core temperature than birds and mammals, reptili
an biochemistry requires enzymes capable of maintaining efficiency over a greate
r range of temperatures than in the case for warm-blooded animals. The optimum b
ody temperature range varies with species, but is typically below that of warm-b
looded animals; for many lizards, it falls in the 24 35 C (75 95 F) range,[66] while ext
reme heat-adapted species, like the American desert iguana Dipsosaurus dorsalis,
can have optimal physiological temperatures in the mammalian range, between 35 a
nd 40 C (95 and 104 F).[67] While the optimum temperature is often encountered when
the animal is active, the low basal metabolism makes body temperature drop rapi
dly when the animal is inactive.
As in all animals, reptilian muscle action produces heat. In large reptiles, lik
e leatherback turtles, the low surface-to-volume ratio allows this metabolically
produced heat to keep the animals warmer than their environment although they d
o not have a warm-blooded metabolism.[68] This form of homeothermy is called gig
antothermy; it has been suggested as having been common in large dinosaurs and o
ther extinct large-bodied reptiles.[69][70]
The benefit of a low resting metabolism is that it requires far less fuel to sus

tain 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.[71] A croco
dile 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.[72] 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 e
nergy output necessary for long distance chases or flying.[73] Higher energetic
capacity might have been responsible for the evolution of warm-bloodedness in bi
rds and mammals.[74] However, investigation of correlations between active capac
ity and thermophysiology show a weak relationship.[75] Most extant reptiles are
carnivores with a sit-and-wait feeding strategy, and whether reptiles are cold b
looded due to their ecology or because their metabolism is a result of their eco
logy is not clear. Energetic studies on some reptiles have shown active capaciti
es equal to or greater than similar sized warm-blooded animals.[76]
Respiration
Reptilian lungs
All reptiles breathe using lungs. Aquatic turtles have developed more permeable
skin, and some species have modified their cloaca to increase the area for gas e
xchange.[77] 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 m
usculature. This is also the same musculature that is used during locomotion. Be
cause 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 oth
er lizard species, employ buccal pumping as a complement to their normal "axial
breathing." This allows the animals to completely fill their lungs during intens
e locomotion, and thus remain aerobically active for a long time. Tegu lizards a
re 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.[78]
Crocodilians actually have a muscular diaphragm that is analogous to the mammali
an diaphragm. The difference is that the muscles for the crocodilian diaphragm p
ull 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 b
ronchia form a number of double tubular chambers within each lung. On inhalation
and exhalation air moves through the airways in the same direction, thus creati
ng a unidirectional airflow through the lungs. A similar system is found in bird
s,[79] monitor lizards[80] and iguanas.[81]
Turtles and tortoises