Reptiles, as commonly defined, are a group of
tetrapod
A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s with an
ectotherm
An ectotherm (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Dav ...
ic metabolism and
amniotic development. Living traditional reptiles comprise four
orders
Order, ORDER or Orders may refer to:
* A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* H ...
:
Testudines
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
,
Crocodilia
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
,
Squamata
Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...
, and
Rhynchocephalia
Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a speciose g ...
. About 12,000 living species of reptiles are listed in the
Reptile Database
The Reptile Database is a scientific database that collects taxonomic information on all living reptile species (i.e. no fossil species such as dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared ...
. The study of the traditional reptile orders, customarily in combination with the study of modern
amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, is called
herpetology
Herpetology (from Ancient Greek ἑρπετόν ''herpetón'', meaning "reptile" or "creeping animal") is a branch of zoology concerned with the study of amphibians (including frogs, salamanders, and caecilians (Gymnophiona)) and reptiles (in ...
.
Reptiles have been subject to several conflicting
taxonomic
280px, Generalized scheme of taxonomy
Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...
definitions.
In
Linnaean taxonomy
Linnaean taxonomy can mean either of two related concepts:
# The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus th ...
, reptiles are gathered together under the
class
Class, Classes, or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used d ...
Reptilia ( ), which corresponds to common usage. Modern
cladistic taxonomy
Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
regards that group as
paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, since
genetic and
paleontological
Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
evidence has determined that
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s (class Aves), as members of
Dinosauria, are more closely related to living
crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s than to other reptiles, and are thus nested among reptiles from an evolutionary perspective. Many cladistic systems therefore redefine Reptilia as a
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
(
monophyletic
In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria:
# the grouping contains its own most recent co ...
group) including birds, though the precise definition of this clade varies between authors.
Others prioritize the clade
Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
, which typically refers to all
amniote
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s more closely related to modern reptiles than to
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s.
The earliest known proto-reptiles originated from the
Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period, having evolved from advanced
reptiliomorph
Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defi ...
tetrapods which became increasingly adapted to life on dry land. The earliest known
eureptile
Sauropsida (Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern reptiles and birds (which, as theropod dinosaur ...
("true reptile") was ''
Hylonomus
''Hylonomus'' (; ''hylo-'' "forest" + ''nomos'' "dweller") is an extinct genus of reptile that lived during the Bashkirian stage of the Late Carboniferous. It is the earliest known crown group amniote and the oldest known unquestionable reptile, ...
'', a small and superficially lizard-like animal which lived in
Nova Scotia
Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
during the
Bashkirian
The Bashkirian is in the International Commission on Stratigraphy geologic timescale the lowest stage (stratigraphy), stage or oldest age (geology), age of the Pennsylvanian (geology), Pennsylvanian. The Bashkirian age lasted from to Mega annu ...
age of the
Late Carboniferous
Late or LATE may refer to:
Everyday usage
* Tardy, or late, not being on time
* Late (or the late) may refer to a person who is dead
Music
* Late (The 77s album), ''Late'' (The 77s album), 2000
* Late (Alvin Batiste album), 1993
* Late!, a pseudo ...
, around .
[ Genetic and fossil data argues that the two largest lineages of reptiles, ]Archosauromorpha
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) than to lepidosaurs (such as tuataras, lizards, ...
(crocodilians, birds, and kin) and Lepidosauromorpha
Lepidosauromorpha (in PhyloCode known as Pan-Lepidosauria) is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains tw ...
(lizards, and kin), diverged during the Permian
The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
period. In addition to the living reptiles, there are many diverse groups that are now extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
, in some cases due to mass extinction events. In particular, the Cretaceous–Paleogene extinction event
The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
wiped out the pterosaur
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s, plesiosaurs
The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia.
Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
, and all non-avian dinosaurs
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
alongside many species of crocodyliforms and squamates
Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles; most members of which are commonly known as lizards, with the group also including snakes. With over 11,991 species, it is also the second-largest order ...
(e.g., mosasaur
Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s). Modern non-bird reptiles inhabit all the continents except Antarctica.
Reptiles are tetrapod vertebrates
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
, creatures that either have four limbs or, like snakes, are descended from four-limbed ancestors. Unlike amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, reptiles do not have an aquatic larval stage. Most reptiles are oviparous
Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
, although several species of squamates are viviparous
In animals, viviparity is development of the embryo inside the body of the mother, with the maternal circulation providing for the metabolic needs of the embryo's development, until the mother gives birth to a fully or partially developed juve ...
, as were some extinct aquatic clades – the fetus develops within the mother, using a (non-mammalian) placenta rather than contained in an eggshell
An eggshell is the outer covering of a hard-shelled egg (biology), egg and of some forms of eggs with soft outer coats.
Worm eggs
Nematode eggs present a two layered structure: an external vitellin layer made of chitin that confers mechanical ...
. 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 fetus
A fetus or foetus (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic development, embryonic stage, the fetal stage of development takes place. Pren ...
es through various forms of placenta analogous to those of mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, with some providing initial care for their hatchlings. Extant
Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to:
* Extant hereditary titles
* Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English
* Exta ...
reptiles range in size from a tiny gecko, ''Sphaerodactylus ariasae
''Sphaerodactylus ariasae'', commonly called the Jaragua sphaero or the Jaragua dwarf gecko, is the smallest species of lizard in the family Sphaerodactylidae.
Description
''Sphaerodactylus ariasae'' is the world's smallest known reptile. The ...
'', which can grow up to to the saltwater crocodile
The saltwater crocodile (''Crocodylus porosus'') is a crocodilian native to saltwater habitats, brackish wetlands and freshwater rivers from India's east coast across Southeast Asia and the Sundaland to northern Australia and Micronesia. It ha ...
, ''Crocodylus porosus'', which can reach over in length and weigh over .
Classification
Research history
In the 13th century, the category of ''reptile'' was recognized in Europe as consisting of a miscellany of egg-laying creatures, including "snakes, various fantastic monsters, lizards, assorted amphibians, and worms", as recorded by Beauvais
Beauvais ( , ; ) is a town and Communes of France, commune in northern France, and prefecture of the Oise Departments of France, département, in the Hauts-de-France Regions of France, region, north of Paris.
The Communes of France, commune o ...
in his ''Mirror of Nature''.
In the 18th century, the reptiles were, from the outset of classification, grouped with the amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s. Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, working from species-poor Sweden
Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...
, where the common adder and grass snake
The grass snake (''Natrix natrix''), sometimes called the ringed snake or water snake, is a Eurasian semi-aquatic non- venomous colubrid snake. It is often found near water and feeds almost exclusively on amphibians.
Subspecies
Many subspecie ...
are often found hunting in water, included all reptiles and amphibians in class
Class, Classes, or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used d ...
in his '' Systema Naturæ''.
The terms ''reptile'' and ''amphibian'' were largely interchangeable, ''reptile'' (from Latin ''repere'', 'to creep') being preferred by the French. J.N. Laurenti was the first to formally use the term ''Reptilia'' for an expanded selection of reptiles and amphibians basically similar to that of Linnaeus. Today, the two groups are still commonly treated under the single heading herpetology
Herpetology (from Ancient Greek ἑρπετόν ''herpetón'', meaning "reptile" or "creeping animal") is a branch of zoology concerned with the study of amphibians (including frogs, salamanders, and caecilians (Gymnophiona)) and reptiles (in ...
.
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 P.A. Latreille erected the class ''Batracia'' (1825) for the latter, dividing the tetrapod
A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s into the four familiar classes of reptiles, amphibians, birds, and mammals. The British anatomist T.H. Huxley made Latreille's definition popular and, together with Richard Owen
Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
, expanded Reptilia to include the various fossil "antediluvian
The antediluvian (alternatively pre-diluvian or pre-flood) period is the time period chronicled in the Bible between the fall of man and the Genesis flood narrative in biblical cosmology. The term was coined by Thomas Browne (1605–1682). The n ...
monsters", including dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s and the mammal-like (synapsid
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
) ''Dicynodon
''Dicynodon'' (from Ancient Greek '' δίς'' "two" and '' κυνόδους'' "canine teeth", often translated to "two canine-teeth" or "two dog-teeth") is a genus of dicynodont therapsid that lived in southern and eastern Africa during the Lat ...
'' he helped describe. This was not the only possible classification scheme: In the Hunterian lectures delivered at the Royal College of Surgeons
The Royal College of Surgeons is an ancient college (a form of corporation) established in England to regulate the activity of surgeons. Derivative organisations survive in many present and former members of the Commonwealth. These organisations ...
in 1863, Huxley grouped the vertebrates into mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, sauroids, and ichthyoids (the latter containing the fishes and amphibians). He subsequently proposed the names of Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
and Ichthyopsida
The anamniotes are an informal group of craniates comprising all fish and amphibians, which lay their eggs in aquatic environments. They are distinguished from the amniotes (reptiles, birds and mammals), which can reproduce on dry land either b ...
for the latter two groups. In 1866, Haeckel
Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
demonstrated that vertebrates could be divided based on their reproductive strategies, and that reptiles, birds, and mammals were united by the amniotic egg
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the C ...
.
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 mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s 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.
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
Procolophonia is an extinct suborder (clade) of herbivorous reptiles that lived from the Middle Permian till the end of the Triassic period. They were originally included as a suborder of the Cotylosauria (later renamed Captorhinida Carroll ...
, Eosuchia
Eosuchians are an extinct order of diapsid reptiles. Depending on which taxa are included the order may have ranged from the late Carboniferous to the Eocene but the consensus is that eosuchians are confined to the Permian and Triassic.
Eosuchi ...
, Millerosauria, Chelonia (turtles), Squamata
Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...
(lizards and snakes), Rhynchocephalia
Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a speciose g ...
, Crocodilia
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
, " thecodonts" (paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
basal Archosaur
Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
ia), non- avian dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s, pterosaur
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s, ichthyosaur
Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides.
Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s, and sauropterygia
Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns.
In the late 19th century, a number of definitions of Reptilia were offered. The biological traits listed by Lydekker in 1896, for example, include a single occipital condyle
The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra.
The condyles are oval or reniform (kidney-shaped) in shape ...
, a jaw joint formed by the quadrate and articular
The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals.
Anatomy
In most vertebrates, the articular bone is connected to two o ...
bones, and certain characteristics of the vertebrae
Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
. The animals singled out by these formulations, the amniote
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s other than the mammals and the birds, are still those considered reptiles today.
The synapsid/sauropsid division supplemented another approach, one that split the reptiles into four subclasses based on the number and position of temporal fenestra
Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
e, openings in the sides of the skull behind the eyes. This classification was initiated by Henry Fairfield Osborn
Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was professor of anatomy at Columbia University, president of the American Museum of Natural History for 25 y ...
and elaborated and made popular by Romer
Romer, Römer, Roemer, or similar may refer to:
* Romer (surname), includes a list of people with the name
* Romer (tool), a cartographic device also known as a reference card
* Rømer scale, a disused temperature scale named after Ole Rømer
* ...
's classic ''Vertebrate Paleontology
Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct vertebrates (animals with vertebrae and their descendants). It also t ...
''. Those four subclasses were:
* Anapsid
An anapsid is an amniote whose skull lacks one or more skull openings (fenestra, or fossae) near the temples. Traditionally, the Anapsida are considered the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Dia ...
a – no fenestrae – cotylosaurs and chelonia (turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s and relatives)
* Synapsida
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rep ...
– one low fenestra – pelycosaur
Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
s and therapsids (the 'mammal-like reptiles
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
')
* Euryapsida __NOTOC__
Euryapsida is a polyphyletic (unnatural, as the various members are not closely related) group of sauropsids that are distinguished by a single temporal fenestra, an opening behind the orbit, under which the post-orbital and squamosal bo ...
– one high fenestra (above the postorbital and squamosal) – protorosaurs (small, early lizard-like reptiles) and the marine sauropterygia
Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns and ichthyosaurs, the latter called Parapsida in Osborn's work.
* Diapsida – two fenestrae – most reptiles, including lizards, snakes, crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s, dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s and pterosaur
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s.
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 merged 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.
Phylogenetics and modern definition
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 clade, monophyletic; that is, groups which include all descendants of a particular ancestor. The reptiles as historically defined are paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, since they exclude both birds and mammals. These respectively evolved from dinosaurs and from early therapsids, both of which were traditionally called "reptiles". Birds are more closely related to crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s than the latter are to the rest of extant reptiles. Colin Tudge wrote:
Mammals are a clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, and therefore the Phylogenetic nomenclature, cladists are happy to acknowledge the traditional taxon Mammalia; and birds, too, are a clade, universally ascribed to the formal taxon bird, 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 synapomorphy, 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'.
Despite the early proposals for replacing the paraphyletic Reptilia with a monophyletic Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
, which includes birds, that term was never adopted widely or, when it was, was not applied consistently.
When Sauropsida was used, it often had the same content or even the same definition as Reptilia. In 1988, Jacques Gauthier proposed a cladistics, 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. While Gauthier's definition was close to the modern consensus, nonetheless, it became considered inadequate because the actual relationship of turtles to other reptiles was not yet well understood at this time.[ Major revisions since have included the reassignment of synapsids as non-reptiles, and classification of turtles as diapsids.][ Gauthier 1994 and Laurin and Reisz 1995's definition of Sauropsida defined the scope of the group as distinct and broader than that of Reptilia, encompassing Mesosauridae as well as Reptilia ''sensu stricto''.]
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, as they are within the clade that includes both lizards and crocodiles.][
]
Taxonomy
General classification of extinct and living reptiles, focusing on major groups.
* Reptilia/Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
**Parareptilia
** Eureptilia
***Captorhinidae
***Diapsida
****Araeoscelidia
****Neodiapsida
*****Drepanosauromorpha (placement uncertain)
*****Younginiformes (paraphyletic)
*****Ichthyosauromorpha (placement uncertain)
*****Thalattosauria (placement uncertain)
*****Sauria
******Lepidosauromorpha
Lepidosauromorpha (in PhyloCode known as Pan-Lepidosauria) is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains tw ...
*******Lepidosauriformes
********Rhynchocephalia
Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a speciose g ...
(tuatara)
********Squamata
Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...
(lizards and snakes)
******Choristodera (placement uncertain)
******Sauropterygia (placement uncertain)
******Pantestudines (turtles and kin, placement uncertain)
******Archosauromorpha
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) than to lepidosaurs (such as tuataras, lizards, ...
*******Protorosauria (paraphyletic)
*******Rhynchosauria
*******Allokotosauria
*******Archosauriformes
********Phytosauria
********Archosauria
*********Pseudosuchia
**********Crocodilia
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
(crocodilians)
*********Avemetatarsalia/Ornithodira
**********Pterosauria
**********Dinosauria
***********Ornithischia
***********Saurischia (including birds (Bird, Aves))
Phylogeny
The cladistics#Cladograms, cladogram presented here illustrates the "family tree" of reptiles, and follows a simplified version of the relationships found by M.S. Lee, in 2013. All Molecular phylogenetics, genetic studies have supported the hypothesis that turtles are diapsids; some have placed turtles within Archosauromorpha,[ though a few have recovered turtles as Lepidosauromorpha instead. The cladogram below used a combination of genetic (molecular) and fossil (morphological) data to obtain its results.][
]
The position of turtles
The placement of turtles has historically been highly variable. Classically, turtles were considered to be related to the primitive anapsid reptiles. Molecular work has usually placed turtles within the diapsids. As of 2013, three turtle genomes have been sequenced. The results place turtles as a sister clade to the archosaurs, the group that includes crocodilians, non-avian dinosaurs, and birds. However, in their comparative analysis of the timing of organogenesis, Werneburg and Sánchez-Villagra (2009) found support for the hypothesis that turtles belong to a separate clade within Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
, outside the saurian clade altogether.
Evolutionary history
Origin of the reptiles
The origin of the reptiles lies about 310–320 million years ago, in the steaming swamps of the late Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period, when the first reptiles evolved from advanced Reptiliomorpha, reptiliomorphs.[
]
The oldest known animal that may have been an amniote
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
is ''Casineria'' (though it may have been a Temnospondyli, temnospondyl). A series of footprints from the fossil strata of Nova Scotia
Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
dated to show typical reptilian toes and imprints of scales. These tracks are attributed to ''Hylonomus
''Hylonomus'' (; ''hylo-'' "forest" + ''nomos'' "dweller") is an extinct genus of reptile that lived during the Bashkirian stage of the Late Carboniferous. It is the earliest known crown group amniote and the oldest known unquestionable reptile, ...
'', the oldest unquestionable reptile known.
It was a small, lizard-like animal, about long, with numerous sharp teeth indicating an insectivorous diet. Other examples include ''Westlothiana'' (for the moment considered a reptiliomorph
Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defi ...
rather than a true amniote
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
) and ''Paleothyris'', both of similar build and presumably similar habit.
However, microsaurs have been at times considered true reptiles, so an earlier origin is possible.
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, such as ''Cochleosaurus'', and remained a small, inconspicuous part of the fauna until the Carboniferous Rainforest Collapse. 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. 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). One of the best known early stem-reptiles is ''Mesosaurus'', a genus from the Cisuralian, Early Permian that had returned to water, feeding on fish.
A 2021 examination of reptile diversity in the Carboniferous and the Permian suggests a much higher degree of diversity than previously thought, comparable or even exceeding that of synapsids. Thus, the "First Age of Reptiles" was proposed.
Anapsids, synapsids, diapsids, and sauropsids
It was traditionally assumed that the first reptiles retained an anapsid skull inherited from their ancestors. This type of skull has a skull roof with only holes for the nostrils, eyes and a pineal eye. The discoveries of synapsid
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
-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 Lanthanosuchoidea, lanthanosuchoids, Millerettidae, millerettids, Bolosauridae, bolosaurids, some Nycteroleteridae, nycteroleterids, some Procolophonoidea, procolophonoids and at least some mesosaurs made it more ambiguous and it is currently uncertain whether the ancestral amniote had an anapsid-like or synapsid-like skull.[ These animals are traditionally referred to as "anapsids", and form a ]paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
basic stock from which other groups evolved.[ Very shortly after the first amniotes appeared, a lineage called ]Synapsida
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rep ...
split off; this group was characterized by a temporal opening in the skull behind each eye giving room for the jaw muscle to move. These are the "mammal-like amniotes", or stem-mammals, that later gave rise to the true mammals. Soon after, another group evolved a similar trait, this time with a double opening behind each eye, earning them the name Diapsida ("two arches").[ 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.]
Turtles have been traditionally believed to be surviving parareptiles, on the basis of their anapsid skull structure, which was assumed to be primitive trait. The rationale for this classification has been disputed, with some arguing that turtles are diapsids that evolved anapsid skulls, improving their armor. Later morphological phylogenetics, phylogenetic studies with this in mind placed turtles firmly within Diapsida. All Molecular phylogenetics, molecular studies have strongly upheld the placement of turtles within diapsids, most commonly as a sister group to extant archosaurs.
Permian reptiles
With the close of the Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
, the amniotes became the dominant tetrapod fauna. While primitive, terrestrial reptiliomorpha, 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.[Edwin Harris Colbert, Colbert, E.H. & Morales, M. (2001): ''Evolution of the Vertebrates, Colbert's Evolution of the Vertebrates: A History of the Backboned Animals Through Time''. 4th edition. John Wiley & Sons, Inc, New York. .]
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).
Early in the period, the modern reptiles, or Sauria, crown-group reptiles, evolved and split into two main lineages: the Archosauromorpha
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) than to lepidosaurs (such as tuataras, lizards, ...
(forebears of turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s, crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s, and dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s) and the Lepidosauromorpha
Lepidosauromorpha (in PhyloCode known as Pan-Lepidosauria) is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains tw ...
(predecessors of modern lizards, snakes, and tuataras). Both groups remained lizard-like and relatively small and inconspicuous during the Permian.
Mesozoic reptiles
The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic extinction event), an event prolonged by the combination of two or more distinct extinction pulses. Most of the earlier parareptile and synapsid megafauna disappeared, being replaced by the true reptiles, particularly archosauromorpha, 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. Archosaurs developed into the well-known dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s and pterosaur
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s, as well as the ancestors of crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s. 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 theropoda, theropods. In the Cretaceous period, these gave rise to the first true birds.
The sister group to Archosauromorpha is Lepidosauromorpha
Lepidosauromorpha (in PhyloCode known as Pan-Lepidosauria) is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains tw ...
, 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 ichthyosaur
Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides.
Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s) and the sauropterygia
Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns, which evolved in the early Triassic – is more controversial. Different authors linked these groups either to lepidosauromorphs or to archosauromorphs, and ichthyopterygians were also argued to be diapsids that did not belong to the least inclusive clade containing lepidosauromorphs and archosauromorphs.
Cenozoic reptiles
The close of the Cretaceous period saw the demise of the Mesozoic era reptilian megafauna (see the Cretaceous–Paleogene extinction event
The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
, also known as K-T extinction event). Of the large marine reptiles, only sea turtles were left; and of the non-marine large reptiles, only the semi-aquatic crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s and broadly similar Choristodera, choristoderes survived the extinction, with last members of the latter, the lizard-like ''Lazarussuchus'', becoming extinct in the Miocene. 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.[ However, reptiles were still important components of the megafauna, particularly in the form of large and giant tortoises.]
After the extinction of most archosaur and marine reptile lines by the end of the Cretaceous, reptile diversification continued throughout the Cenozoic. Squamata, Squamates took a massive hit during the K–Pg event, only recovering ten million years after it, but they underwent a great radiation event once they recovered, and today squamates make up the majority of living reptiles (> 95%). 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 List of domesticated animals, domesticated species).
Morphology and physiology
Circulation
All Lepidosauria, lepidosaurs and turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s have a three-chambered heart consisting of two atrium (heart), atria, one variably partitioned ventricle (heart), 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 (biology), fitness advantage.
For example, iguana hearts, like the majority of the Squamata, squamate hearts, are composed of three chambers with two aorta and one ventricle, cardiac involuntary muscles. The main structures of the heart are the sinus venosus, the pacemaker, the Atrium (heart), left atrium, the Atrium (heart), right atrium, the Heart valve, atrioventricular valve, the cavum venosum, cavum arteriosum, the cavum pulmonale, the muscular ridge, the ventricular ridge, pulmonary veins, and paired aortic arches.
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 cardiac cycle, ventricular diastole and completely divides it during systole (medicine), ventricular systole. Because of this ridge, some of these squamata, squamates are capable of producing ventricular pressure differentials that are equivalent to those seen in mammalian and avian hearts.
Crocodilia
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns have an anatomically four-chambered heart, similar to bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, but also have two systemic aortas and are therefore capable of bypassing their pulmonary circulation. In turtles, the ventricle is not perfectly divided, so a mix of aerated and nonaerated blood can occur.
Metabolism
Modern non-avian reptiles exhibit some form of Ectotherm, cold-bloodedness (i.e. some mix of poikilothermy, ectotherm
An ectotherm (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Dav ...
y, 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 bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s and mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, 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 range, while extreme heat-adapted species, like the American desert iguana ''Dipsosaurus dorsalis'', can have optimal physiological temperatures in the mammalian range, between . 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-blooded metabolism. This form of homeothermy is called gigantothermy; it has been suggested as having been common in large dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s and other extinct large-bodied reptiles.
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. 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. 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. Higher energetic capacity might have been responsible for the evolution of warm-bloodedness in birds and mammals. However, investigation of correlations between active capacity and Thermoregulation, thermophysiology show a weak relationship. 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.
Respiratory system
All reptiles breathe using lungs. Aquatic turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s have developed more permeable skin, and some species have modified their cloaca to increase the area for gas exchange. Even with these adaptations, breathing is never fully accomplished without lungs. Lung ventilation is accomplished differently in each main reptile group. In squamata, 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 Carrier's constraint, 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. Tupinambis, Tegu lizards are known to possess a proto-thoracic diaphragm, 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.
Crocodilia
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns 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 "Liver, hepatic piston". The Bronchus, airways 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, monitor lizards and iguanas.
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 Skink, 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.
Turtles and tortoises
How turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s 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 Breathing, breathe. The varied results indicate that turtles 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 (''Indian flapshell turtle, 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 thoracic diaphragm, diaphragm), turtles are capable of pushing their viscera up and down, resulting in effective respiration, since many of these muscles have attachment points in conjunction with their forelimbs (indeed, many of the muscles expand into the limb pockets during contraction).
Breathing during locomotion has been studied in three species, and they show different patterns. Adult female green sea turtles do not breathe as they crutch along their nesting beaches. They hold their breath during terrestrial locomotion and breathe in bouts as they rest. North American box turtles breathe continuously during locomotion, and the ventilation cycle is not coordinated with the limb movements. This is because they use their abdominal muscles to breathe during locomotion. The last species to have been studied is the red-eared slider, which also breathes during locomotion, but takes smaller breaths during locomotion than during small pauses between locomotor bouts, indicating that there may be mechanical interference between the limb movements and the breathing apparatus. Box turtles have also been observed to breathe while completely sealed up inside their shells.[
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Sound production
Compared with frogs, birds, and mammals, reptiles are less vocal. Sound production is usually limited to wikt:hiss, hissing, which is produced merely by forcing air though a partly closed glottis and is not considered to be a true vocalization. The ability to vocalize exists in crocodilians, some lizards and turtles; and typically involves vibrating fold-like structures in the larynx or glottis. Some geckos and turtles possess true vocal cords, which have elastin-rich connective tissue.
Hearing in snakes
Hearing in humans relies on 3 parts of the ear; the outer ear that directs sound waves into the ear canal, the middle ear that transmits incoming sound waves to the inner ear, and the inner ear that helps in hearing and keeping their balance. Unlike humans and other mammals, snakes do not possess an outer ear, a middle ear, and a Tympanum (anatomy), tympanum but have an inner ear structure with cochleas directly connected to their jawbone. They are able to feel the vibrations generated from the sound waves in their jaw as they move on the ground. This is done by the use of mechanoreceptors, sensory nerves that run along the body of snakes directing the vibrations along the spinal nerves to the brain. Snakes have a sensitive auditory perception and can tell which direction sound being made is coming from so that they can sense the presence of prey or predator but it is still unclear how sensitive snakes are to sound waves traveling through the air.
Skin
Reptilian skin is covered in a horny Epidermis (zoology), 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 dermis, dermal layer that produces leather in mammals.
Exposed parts of reptiles are protected by reptile scales, scales or scutes, sometimes with a bony base (osteoderms), forming armour (zoology), armor. In lepidosaurs, such as lizards and snakes, the whole skin is covered in overlapping epidermis (zoology), 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 lepidosaurs. The scales found in turtles and crocodiles are of dermis, 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.
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 five to six weeks while adults shed three to four times a year. 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. Proteolysis, Proteolytic enzymes and Lymph, lymphatic fluid is secreted between the old and new layers of skin. Consequently, this lifts the old skin from the new one allowing shedding to occur. Snakes will shed from the head to the tail while lizards shed in a "patchy pattern". Dysecdysis, a common skin disease in snakes and lizards, will occur when ecdysis, or shedding, fails. There are numerous reasons why shedding fails and can be related to inadequate humidity and temperature, nutritional deficiencies, dehydration and traumatic injuries. Nutritional deficiencies decrease proteolytic enzymes while dehydration reduces lymphatic fluids to separate the skin layers. Traumatic injuries on the other hand, form scars that will not allow new scales to form and disrupt the process of ecdysis.
Excretion
Excretion is performed mainly by two small kidneys. In diapsids, uric acid is the main nitrogenous waste product; turtles, like mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, excrete mainly urea. Unlike the mammalian kidney, kidneys of mammals and birds, Kidney (vertebrates)#Reptile kidney, 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 (anatomy), colon to aid in the reabsorption of water. Some are also able to take up water stored in the Urinary bladder, bladder. Excess salts are also excreted by nasal and lingual salt glands in some reptiles.
In all reptiles, the urinogenital ducts and the rectum both empty into an organ called a cloaca. In some reptiles, a midventral wall in the cloaca may open into a urinary bladder, but not all. It is present in all turtles and tortoises as well as most lizards, but is lacking in the monitor lizard, the legless lizards. It is absent in the snakes, alligators, and crocodiles.
Many turtles and lizards have proportionally very large bladders. Charles Darwin noted that the Galapagos tortoise had a bladder which could store up to 20% of its body weight. Such adaptations are the result of environments such as remote islands and deserts where water is very scarce. Other desert-dwelling reptiles have large bladders that can store a long-term reservoir of water for up to several months and aid in osmoregulation.
Turtles have two or more accessory urinary bladders, located lateral to the neck of the urinary bladder and dorsal to the pubis, occupying a significant portion of their body cavity. Their bladder is also usually bilobed with a left and right section. The right section is located under the liver, which prevents large stones from remaining in that side while the left section is more likely to have Bladder stone (animal), calculi.
Digestion
Most reptiles are insectivorous or carnivorous and have simple and comparatively short digestive tracts due to meat being fairly simple to break down and digest. Digestion is slower than in mammals, reflecting their lower resting metabolism and their inability to divide and masticate their food. Their poikilotherm metabolism has very low energy requirements, allowing large reptiles like crocodiles and large constrictors to live from a single large meal for months, digesting it slowly.[
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 ]dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s.[ Today, ]turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s are the only predominantly herbivorous reptile group, but several lines of Agamidae, agamas and Iguanidae, iguanas have evolved to live wholly or partly on plants.
Herbivorous reptiles face the same problems of mastication as herbivorous mammals but, lacking the complex teeth of mammals, many species swallow rocks and pebbles (so called gastroliths) to aid in digestion: The rocks are washed around in the stomach, helping to grind up plant matter.[ Fossil gastroliths have been found associated with both ornithopods and sauropods, though whether they actually functioned as a gastric mill in the latter is disputed. Salt water crocodiles also use gastroliths as Sailing ballast, ballast, stabilizing them in the water or helping them to dive. A dual function as both stabilizing ballast and digestion aid has been suggested for gastroliths found in plesiosaurs.
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Nerves
The reptilian nervous system contains the same basic part of the amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
brain, but the reptile cerebrum and cerebellum are slightly larger. Most typical sense organs are well developed with certain exceptions, most notably the snake's lack of external ears (middle and inner ears are present). There are twelve pairs of cranial nerves. Due to their short cochlea, reptiles use electrical tuning to expand their range of audible frequencies.
Vision
Most reptiles are Diurnality, 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.
Reptiles usually have excellent vision, allowing them to detect shapes and motions at long distances. They often have poor vision in low-light conditions. Birds, crocodiles and turtles have three types of Photoreceptor cell, photoreceptor: rod cell, rods, single Cone cell, cones and double cones, which gives them sharp color vision and enables them to see ultraviolet wavelengths. The lepidosaurs appear to have lost the duplex retina and only have a single class of receptor that is cone-like or rod-like depending on whether the species is diurnal or nocturnal. In many burrowing species, such as blind snakes, vision is reduced.
Many Lepidosauria, lepidosaurs have a photosensory organ on the top of their heads called the parietal eye, which are also called third eye, pineal eye or pineal gland. This "eye" does not work the same way as a normal eye does as it has only a rudimentary retina and lens and thus, cannot form images. It is, however, sensitive to changes in light and dark and can detect movement.[
Some snakes have extra sets of visual organs (in the loosest sense of the word) in the form of Loreal pit, pits sensitive to infrared radiation (heat). Such heat-sensitive pits are particularly well developed in the pit vipers, but are also found in boidae, boas and pythonidae, pythons. These pits allow the snakes to sense the body heat of birds and mammals, enabling pit vipers to hunt rodents in the dark.
Most reptiles, as well as birds, possess a nictitating membrane, a translucent third eyelid which is drawn over the eye from the inner corner. In crocodilians, it protects its eyeball surface while allowing a degree of vision underwater. However, many squamates, geckos and snakes in particular, lack eyelids, which are replaced by a transparent scale. This is called the brille, spectacle, or eyecap. The brille is usually not visible, except for when the snake molts, and it protects the eyes from dust and dirt.
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Reproduction
Reptiles generally sexual reproduction, 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 hemipenis, 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.
Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion (5), chorion (6), and allantois (8) are present during embryonic life. 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 squamates and many extinct clades of reptiles. Among squamates, many species, including all boas and most vipers, use 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
The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
mesosaurs, 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 mosasaur
Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s, ichthyosaur
Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides.
Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s, and Sauropterygia, a group that includes pachypleurosaurs and Plesiosauria.[
Asexual reproduction has been identified in squamata, 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 teiidae, teiids (especially ''Aspidocelis'') and lacertidae, lacertids (''Lacerta (genus), Lacerta''). In captivity, Komodo dragons (Varanidae) have reproduced by parthenogenesis.
Parthenogenetic species are suspected to occur among chameleons, agamidae, agamids, night lizard, xantusiids, and typhlopidae, 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. To date, there has been no confirmation of whether TDSD occurs in snakes.
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Longevity
Giant tortoises are among the longest-lived vertebrate animals (over 100 years by some estimates) and have been used as a model for studying longevity.[Quesada V, Freitas-Rodríguez S, Miller J, Pérez-Silva JG, Jiang ZF, Tapia W, Santiago-Fernández O, Campos-Iglesias D, Kuderna LFK, Quinzin M, Álvarez MG, Carrero D, Beheregaray LB, Gibbs JP, Chiari Y, Glaberman S, Ciofi C, Araujo-Voces M, Mayoral P, Arango JR, Tamargo-Gómez I, Roiz-Valle D, Pascual-Torner M, Evans BR, Edwards DL, Garrick RC, Russello MA, Poulakakis N, Gaughran SJ, Rueda DO, Bretones G, Marquès-Bonet T, White KP, Caccone A, López-Otín C. Giant tortoise genomes provide insights into longevity and age-related disease. Nat Ecol Evol. 2019 Jan;3(1):87-95. doi: 10.1038/s41559-018-0733-x. Epub 2018 Dec 3. PMID 30510174; PMCID: PMC6314442] DNA analysis of the genomes of Lonesome George, the iconic last member of ''Pinta Island tortoise, Chelonoidis abingdonii'', and the Aldabra giant tortoise ''Aldabrachelys gigantea'' led to the detection of lineage-specific variants affecting DNA repair genes that might contribute to our understanding of increased lifespan.[
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Cognition
Reptiles are generally considered less intelligent than mammals and birds. The Brain-to-body mass ratio, 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, though larger reptiles can show more complex brain development. Larger lizards, like the monitor lizard, monitors, are known to exhibit complex behavior, including cooperation and cognitive abilities allowing them to optimize their foraging and Territory (animal), territoriality over time. Crocodiles have relatively larger brains and show a fairly complex social structure. The Komodo dragon is even known to engage in play, as are turtles, which are also considered to be social creatures, and sometimes switch between monogamy and promiscuity in their sexual behavior. One study found that wood turtles were better than white rats at learning to navigate mazes. Another study found that giant tortoises are capable of learning through operant conditioning, visual discrimination and retained learned behaviors with long-term memory. Sea turtles have been regarded as having simple brains, but their flippers are used for a variety of foraging tasks (holding, bracing, corralling) in common with marine mammals.
There is evidence that reptiles are sentient and able to feel emotions including anxiety and pleasure.
Defense mechanisms
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, Conflict avoidance, avoidance is the most common form of defense in reptiles. At the first sign of danger, most snakes and lizards crawl away into the undergrowth, and turtles and crocodiles will plunge into water and sink out of sight.
Camouflage and warning
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 Crypsis, cryptic coloration of plain or mottled gray, green, and brown to allow them to blend into the background of their natural environment. 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.
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. If danger arises so suddenly that flight is useless, 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 to ward off 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. 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.
Alternative defense in snakes
Camouflage does not always fool a predator. When caught out, snake species adopt different defensive tactics and use a complicated set of behaviors when attacked. Some species, like cobras or hognose snakes, 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, which use snake venom, 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
The grass snake (''Natrix natrix''), sometimes called the ringed snake or water snake, is a Eurasian semi-aquatic non- venomous colubrid snake. It is often found near water and feeds almost exclusively on amphibians.
Subspecies
Many subspecie ...
, thanatosis, play dead when in danger; some, including the grass snake, exude a foul-smelling liquid to deter attackers.
Defense in crocodilians
When a crocodilian
Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
is concerned about its safety, it will gape to expose the teeth and tongue. If this does not 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.
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. The main weapon in all crocodiles is the bite, which can generate very high bite force. Many species also possess Canine tooth, canine-like teeth. These are used primarily for seizing prey, but are also used in fighting and display.
Shedding and regenerating tails
Geckos, skinks, and some 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 thrash, 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. The tail grows back in most species, but some, like crested geckos, lose their tails for the rest of their lives. 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 Tiliqua rugosa, 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.
Reptiles that are capable of shedding their tails can partially regeneration (biology), 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.
Relations with humans
In cultures and religions
Dinosaurs have been widely depicted in culture since the English palaeontologist Richard Owen
Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
coined the name ''dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
'' in 1842. As soon as 1854, the Crystal Palace Dinosaurs were on display to the public in south London. One dinosaur appeared in literature even earlier, as Charles Dickens placed a ''Megalosaurus'' in the first chapter of his novel ''Bleak House'' in 1852.
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 documentary film, documentaries of the 1990s and first decade of the 21st century, to the fantastic, as in the monster movies of the 1950s and 1960s.
The snake or serpent has played a powerful Serpent (symbolism), symbolic role in different cultures. In History of Egypt, Egyptian history, the Nile cobra adorned the crown of the pharaoh. It was snake worship, worshipped as one of the gods and was also used for sinister purposes: murder of an adversary and ritual suicide (Cleopatra VII of Egypt, 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 (mythology), 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 Titan (mythology), 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. In Hinduism, snake worship, 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. 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." 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.
The turtle has a prominent position as a symbol of steadfastness and tranquility in religion, mythology, and folklore from around the world. A tortoise's longevity is suggested by its long lifespan and its shell, which was thought to protect it from any foe. In the cosmological myths of several cultures a ''World Turtle'' carries the world upon its back or supports the heavens.
Medicine
Deaths from snakebites are uncommon in many parts of the world, but are still counted in tens of thousands per year in India. 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. The cytotoxic effect of snake venom is being researched as a potential treatment for cancers.
Gila monsters produce compounds that reduce plasma glucose; one of these substances is now used in the anti-diabetes drug exenatide (Byetta), a glucagon-like peptide-1 (GLP-1) receptor agonist like Semaglutide, semiglutide (Ozempic). Another toxin from Gila monster saliva has been studied for use as an anti-Alzheimer's drug.
Gecko, Geckos have also been used as folk medicine, especially in China, without any evidence that they have any active compounds. Turtles have been used in Chinese traditional medicine for thousands of years, with every part of the turtle believed to have medical benefits (again, without scientific evidence). Growing demand for turtle meat has placed pressure on vulnerable wild populations of turtles.
Commercial farming
Crocodiles are protected in many parts of the world, and are crocodile farm, farmed commercially. Their hides are tanned and used to make leather goods such as shoes and handbags; crocodile meat is also considered a delicacy. 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.
Snakes are also farmed, primarily in East Asia, East and Southeast Asia, and their production has become more intensive in the last decade. Snake farming has been troubling for conservation in the past as it can lead to overexploitation of wild snakes and their natural prey to supply the farms. However, farming snakes can limit the hunting of wild snakes, while reducing the slaughter of higher-order vertebrates like cows. The energy efficiency of snakes is higher than expected for carnivores, due to their ectothermy and low metabolism. Waste protein from the poultry and pig industries is used as feed in snake farms. Snake farms produce meat, Snakeskin, snake skin, and antivenom.
Turtle farming is another known but controversial practice. Turtles have been farmed for a variety of reasons, ranging from food to traditional medicine, the pet trade, and scientific conservation. Demand for turtle meat and medicinal products is one of the main threats to turtle conservation in Asia. Though commercial breeding would seem to insulate wild populations, it can stoke the demand for them and increase wild captures.[ Even the potentially appealing concept of raising turtles at a farm to release into the wild is questioned by some veterinarians who have had some experience with farm operations. They caution that this may introduce into the wild populations infectious diseases that occur on the farm, but have not (yet) been occurring in the wild.
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Reptiles in captivity
A herpetarium is a zoo, zoological exhibition space for reptiles and amphibians.
In the Western world, some snakes (especially relatively docile species such as the Python regius, ball python and corn snake) are sometimes kept as pets. Numerous species of lizard are kept as pets, including Pogona, bearded dragons, iguanas, anoles, and geckos (such as the popular leopard gecko and the crested gecko).
Turtles and tortoises are increasingly popular pets, but keeping them can be challenging due to their particular requirements, such as temperature control, the need for UV light sources, and a varied diet. The long lifespans of turtles and especially tortoises mean they can potentially outlive their owners. Good hygiene and significant maintenance is necessary when keeping reptiles, due to the risks of ''Salmonella'' and other pathogens. Regular hand-washing after handling is an important measure to prevent infection.
See also
* Amphibian and reptile tunnel
* List of reptiles
* Lists of reptiles by region
* Reptile Database
The Reptile Database is a scientific database that collects taxonomic information on all living reptile species (i.e. no fossil species such as dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared ...
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Notes
References
Further reading
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* Duellman, William E., Berg, Barbara (1962),
Type Specimens of Amphibians and Reptiles in the Museum of Natural History, the University of Kansas
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External links
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* — an online full text copy of a 22 volume 13,000 page summary of the state of reptile research.
{{Portal bar, Reptiles
Reptiles,
Extant Pennsylvanian first appearances
Paraphyletic groups
Articles containing video clips