Sauropsida
Encyclopedia
Sauropsida is a group of amniotes that includes all existing reptiles and birds and their fossil ancestors, including the dinosaur
s, the immediate ancestors of birds. Sauropsida is distinguished from Theropsida ("beast faces"), more commonly called Synapsid
a, which includes mammals and their fossil ancestors.
and Archaeopteryx
, that were starting to become known at the time.
In the Hunterian lectures delivered at the Royal College of Surgeons
in 1863, Huxley grouped the vertebrate classes
informally into mammals, sauroids, and ichthyoids (the latter containing the anamniotes
), based on the gaps in physiological traits and lack of transitional fossil
s that seem to exist between the three groups. He subsequently proposed the names of Sauropsida and Ichthyopsida for the two latter.
It has been noted that species on the mammalian line (synapsids) like Dicynodon
, described as a reptile by Richard Owen
as early as 1845, were beginning to become known at the time Huxley gave his systematic lecture. Thus, under the original definition, Sauropsida contained not only the the groups usually associated with it today, but also several mammal-line species.
synapsids from South Africa
had become well known, allowing palaeontologists to trace synapsid evolution in much greater detail. The term Sauropsida was taken up by E.S. Goodrich
in 1916 much like Huxley's, to include lizards, birds and their relatives. He distinguished them from mammal
s and their extinct relatives, which he included in the sister group Theropsida (now usually replaced with the name Synapsida). Goodrich's classification thus differs somewhat from Huxley's, in which the non-mammalian synapsids (or at least the dicynodontians) fell under the sauropsids. 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, the Protosauria ("first lizards"), which included some Paleozoic
amphibian
s as well as early reptile
s predating the sauropsid/synapsid split (and thus not true sauropsids).
, Eosuchia
, Millerosauria
, Chelonia
(turtles), Squamata
(lizards and snakes), Rhynchocephalia
, Crocodilia
, "thecodont
s" (paraphyletic
basal
Archosaur
ia), non-avian dinosaur
s, pterosaur
s, ichthyosaur
s, and sauropyterygians
.
This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer
's classic Vertebrate Paleontology
) into four subclasses according to the positioning of temporal fenestrae, openings in the sides of the skull behind the eyes. Since the advent of phylogenetic nomenclature
, the term Reptilia has fallen out of favor with many taxonomists, who have used Sauropsida in its place to include a monophyletic group containing the traditional reptiles and the birds.
rather than a clade for as long as evolution
has been recognised. Reclassifying reptiles has been among the key aims of phylogenetic nomenclature
.
The term Sauropsida had from the mid 20th century been used to denote all species not on the synapsid side after the synapsid/sauropsid split, a branch-based clade
. This group encompasses all now-living reptiles as well as birds, and as such is comparable to Goodrich's classification, the difference being that better resolution of the early amniote tree has split up most of the Goodrich's "Protosauria". Some later cladistic work has used Sauropsida more restrictively, to signify the crown group
, i.e. all descendants of the last common ancestor of extant reptiles and birds. A number of phylogenetic stem, node and crown definitions have been published, anchored in a variety of fossil and extant organisms, thus there is currently no consensus of the actual definition (and thus content) of Sauropsida as a phylogenetic unit.
Some taxonomists, such as Benton (2004), have co-opted the term to fit into traditional rank-based classifications, making Sauropsida and Synapsida class-level taxa to replace the traditional Class Reptilia, while Modesto and Anderson (2004), using the PhyloCode
standard, have suggested replacing the name Sauropsida with their redefinition of Reptilia, arguing that the latter is by far better known and should have priority.
.
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
s, the immediate ancestors of birds. Sauropsida is distinguished from Theropsida ("beast faces"), more commonly called Synapsid
Synapsid
Synapsids are a group of animals that includes mammals and everything more closely related to mammals than to other living amniotes. They are easily separated from other amniotes by having an opening low in the skull roof behind each eye, leaving a bony arch beneath each, accounting for their name...
a, which includes mammals and their fossil ancestors.
Huxley and the fossil gaps
The term Sauropsida ("lizard faces") has a long history, and hails back to Thomas Henry Huxley, and his opinion that birds had risen from the dinosaurs. He based this chiefly on the fossils of HesperornisHesperornis
Hesperornis is a genus of flightless aquatic birds that spanned the first half of the Campanian age of the Late Cretaceous period . One of the lesser-known discoveries of the paleontologist O. C. Marsh in the late 19th century Bone Wars, it was an important early find in the history of avian...
and Archaeopteryx
Archaeopteryx
Archaeopteryx , sometimes referred to by its German name Urvogel , is a genus of theropod dinosaur that is closely related to birds. The name derives from the Ancient Greek meaning "ancient", and , meaning "feather" or "wing"...
, that were starting to become known at the time.
In the Hunterian lectures delivered at the Royal College of Surgeons
Royal College of Surgeons of England
The Royal College of Surgeons of England is an independent professional body and registered charity committed to promoting and advancing the highest standards of surgical care for patients, regulating surgery, including dentistry, in England and Wales...
in 1863, Huxley grouped the vertebrate classes
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
informally into mammals, sauroids, and ichthyoids (the latter containing the anamniotes
Anamniotes
The anamniotes are an informal group of vertebrates that lack the amnion during fetal development. These animals are not able to have embryos that develop on land, thus they lay their eggs in water exclusively...
), based on the gaps in physiological traits and lack of transitional fossil
Transitional fossil
A transitional fossil is any fossilized remains of a lifeform that exhibits characteristics of two distinct taxonomic groups. A transitional fossil is the fossil of an organism near the branching point where major individual lineages diverge...
s that seem to exist between the three groups. He subsequently proposed the names of Sauropsida and Ichthyopsida for the two latter.
It has been noted that species on the mammalian line (synapsids) like Dicynodon
Dicynodon
Dicynodon is a type of dicynodont therapsid that flourished during the Permian period between 251 and 299 million years ago. Like all dicynodonts, it was herbivorous. This animal was toothless, except for prominent tusks, hence the name...
, described as a reptile by Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...
as early as 1845, were beginning to become known at the time Huxley gave his systematic lecture. Thus, under the original definition, Sauropsida contained not only the the groups usually associated with it today, but also several mammal-line species.
Sauropsids redefined
By the early 20th century, the fossils of PermianPermian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...
synapsids from South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...
had become well known, allowing palaeontologists to trace synapsid evolution in much greater detail. The term Sauropsida was taken up by E.S. Goodrich
Edwin Stephen Goodrich
Edwin Stephen Goodrich , was an English zoologist, specialising in comparative anatomy, embryology, paleontology, and evolution. He held the Linacre Chair of Zoology in the University of Oxford from 1921 to 1946...
in 1916 much like Huxley's, to include lizards, birds and their relatives. He distinguished them from mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s and their extinct relatives, which he included in the sister group Theropsida (now usually replaced with the name Synapsida). Goodrich's classification thus differs somewhat from Huxley's, in which the non-mammalian synapsids (or at least the dicynodontians) fell under the sauropsids. 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, the Protosauria ("first lizards"), which included some Paleozoic
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...
s as well as early reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...
s predating the sauropsid/synapsid split (and thus not true sauropsids).
Detailing the reptile family tree
In 1956, D.M.S. Watson observed that sauropsids and synapsids diverged very early in the reptilian evolutionary history, and so he divided Goodrich's Protosauria between the two groups. He also reinterpreted the Sauropsida and Theropsida to exclude birds and mammals respectively, making them paraphyletic, unlike Goodrich's definition. Thus his Sauropsida included ProcolophoniaProcolophonia
The Procolophonia are a suborder 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 but are now considered a clade of Parareptilia...
, Eosuchia
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....
, Millerosauria
Millerettid
The millerettids are an extinct group of anapsids that lived in South Africa during the Upper Permian. They were small insectivores and probably resembled modern lizards in appearance and lifestyle.-External links:*...
, Chelonia
Turtle
Turtles are reptiles of the order Testudines , characterised by a special bony or cartilaginous shell developed from their ribs that acts as a shield...
(turtles), Squamata
Squamata
Squamata, or the scaled reptiles, is the largest recent order of reptiles, including lizards and snakes. Members of the order are distinguished by their skins, which bear horny scales or shields. They also possess movable quadrate bones, making it possible to move the upper jaw relative to the...
(lizards and snakes), Rhynchocephalia
Sphenodontia
Sphenodontia is an order of lizard-like reptiles that includes only one living genus, the tuatara , and only two living species...
, Crocodilia
Crocodilia
Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period . They are the closest living relatives of birds, as the two groups are the only known survivors of the Archosauria...
, "thecodont
Thecodont
Thecodont , now considered an obsolete term, was formerly used to describe a diverse range of early archosaurs that first appeared in the Latest Permian and flourished until the end of the Triassic period...
s" (paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...
basal
Basal (phylogenetics)
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...
Archosaur
Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of modern birds and crocodilians. This group also includes all extinct non-avian dinosaurs, many extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most...
ia), non-avian dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
s, pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...
s, ichthyosaur
Ichthyosaur
Ichthyosaurs were giant marine reptiles that resembled fish and dolphins...
s, and sauropyterygians
Sauropterygia
Sauropterygia were a group of very successful aquatic reptiles that flourished during the Mesozoic before they became extinct at the end of the era. They were united by a radical adaptation of their shoulder, designed to support powerful flipper strokes...
.
This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer
Alfred Romer
Alfred Sherwood Romer was an American paleontologist and comparative anatomist and a specialist in vertebrate evolution.-Biography:...
's classic Vertebrate Paleontology
Vertebrate Paleontology (Romer)
Vertebrate Paleontology is an advanced textbook on vertebrate paleontology by Alfred Sherwood Romer, published by the University of Chicago Press. It went through three editions and for many years constituted a very authoritative work and the definitive coverage of the subject. A condensed...
) into four subclasses according to the positioning of temporal fenestrae, openings in the sides of the skull behind the eyes. Since the advent of phylogenetic nomenclature
Phylogenetic nomenclature
Phylogenetic nomenclature or phylogenetic taxonomy is an alternative to rank-based nomenclature, applying definitions from cladistics . Its two defining features are the use of phylogenetic definitions of biological taxon names, and the lack of obligatory ranks...
, the term Reptilia has fallen out of favor with many taxonomists, who have used Sauropsida in its place to include a monophyletic group containing the traditional reptiles and the birds.
Cladistics and the Sauropsida
The class Reptilia has been known to be an evolutionary gradeEvolutionary grade
In alpha taxonomy, a grade refers to a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit.-Definition:...
rather than a clade for as long as evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
has been recognised. Reclassifying reptiles has been among the key aims of phylogenetic nomenclature
Phylogenetic nomenclature
Phylogenetic nomenclature or phylogenetic taxonomy is an alternative to rank-based nomenclature, applying definitions from cladistics . Its two defining features are the use of phylogenetic definitions of biological taxon names, and the lack of obligatory ranks...
.
The term Sauropsida had from the mid 20th century been used to denote all species not on the synapsid side after the synapsid/sauropsid split, a branch-based clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. This group encompasses all now-living reptiles as well as birds, and as such is comparable to Goodrich's classification, the difference being that better resolution of the early amniote tree has split up most of the Goodrich's "Protosauria". Some later cladistic work has used Sauropsida more restrictively, to signify the crown group
Crown group
A crown group is a group consisting of living representatives, their ancestors back to the most recent common ancestor of that group, and all of that ancestor's descendants. The name was given by Willi Hennig, the formulator of phylogenetic systematics, as a way of classifying living organisms...
, i.e. all descendants of the last common ancestor of extant reptiles and birds. A number of phylogenetic stem, node and crown definitions have been published, anchored in a variety of fossil and extant organisms, thus there is currently no consensus of the actual definition (and thus content) of Sauropsida as a phylogenetic unit.
Some taxonomists, such as Benton (2004), have co-opted the term to fit into traditional rank-based classifications, making Sauropsida and Synapsida class-level taxa to replace the traditional Class Reptilia, while Modesto and Anderson (2004), using the PhyloCode
PhyloCode
The International Code of Phylogenetic Nomenclature, known as the PhyloCode for short, is a developing draft for a formal set of rules governing phylogenetic nomenclature...
standard, have suggested replacing the name Sauropsida with their redefinition of Reptilia, arguing that the latter is by far better known and should have priority.
Phylogeny
The cladogram presented here illustrates the "family tree" of sauropsids, and follows a simplified version of the relationships found by Laurin and Gauthier (1996), presented as part of the Tree of Life Web ProjectTree of Life Web Project
The Tree of Life Web Project is an ongoing Internet project providing information about the diversity and phylogeny of life on Earth. This collaborative peer reviewed project began in 1995, and is written by biologists from around the world....
.