Pelycosaurian
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Pelycosaur ( ) is an older term for basal or primitive Late
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
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 ...
s, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an
order 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 ...
, but this is now thought to be incorrect and outdated. Because it excludes the advanced synapsid group Therapsida, the term is
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 ...
and contrary to modern formal naming practice. Thus the name ''pelycosaurs'', similar to the term ''mammal-like reptiles'', fell out of favor among scientists by the 21st century, and is only used informally, if at all, in the modern scientific literature. The terms stem mammals, protomammals, and basal or primitive synapsids are instead used where needed.


Etymology

The modern word was created from Greek meaning 'basin' and meaning 'lizard'. The term ''pelycosaur'' has been fairly well abandoned by paleontologists because it no longer matches the features that distinguish a clade. Pelycosauria is 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 ...
taxon because it excludes the therapsids. For that reason, the term is sometimes avoided by proponents of a strict cladistic approach.
Eupelycosauria Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308million years ago during the Early Pennsylvanian epoch, with the fo ...
is used to designate the clade that includes most pelycosaurs, along with the Therapsida and Mammalia. In contrast to "pelycosaurs", Eupelycosauria is a proper
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.
Caseasauria Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insect ...
is a pelycosaur side-branch, or clade, that did not leave any descendants.


Evolutionary history

The pelycosaurs appear to have been a group of synapsids that have direct ancestral links with the
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, having differentiated teeth and a developing hard palate. The pelycosaurs appeared during the
Late 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), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
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 ...
and reached their apex in the early part of the Permian, remaining the dominant land animals for some 40 million years. A few continued into the
Capitanian In the geologic timescale, the Capitanian is an age or stage of the Permian. It is also the uppermost or latest of three subdivisions of the Guadalupian Epoch or Series. The Capitanian lasted between and million years ago. It was preceded by t ...
, but they experienced a sharp decline in diversity in the late
Kungurian In the geologic timescale, the Kungurian is an age or stage of the Permian. It is the latest or upper of four subdivisions of the Cisuralian Epoch or Series. The Kungurian lasted between and million years ago (Ma). It was preceded by the Arti ...
. They were succeeded by the therapsids.


Description

Some species were quite large, growing to a length of or more, although most species were much smaller. Well-known pelycosaurs include the
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
'', ''
Sphenacodon ''Sphenacodon'' (meaning "wedge point tooth") is an extinct genus of synapsid that lived from about 300 to about 280 million years ago (Ma) during the Late Carboniferous and Early Permian periods. Like the closely related ''Dimetrodon'', ''Sphena ...
'', ''
Edaphosaurus ''Edaphosaurus'' (, meaning "pavement lizard" for dense clusters of its teeth) is a genus of extinct edaphosaurid synapsids that lived in what is now North America and Europe around 303.4 to 272.5 million years ago, during the Late Carboniferous ...
'', and ''
Ophiacodon ''Ophiacodon'' (meaning "snake tooth") is an extinct genus of synapsid belonging to the family Ophiacodontidae that lived from the Late Carboniferous to the Early Permian in North America and Europe. The genus was named along with its type specie ...
''. Pelycosaur fossils have been found mainly in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, although some small, late-surviving forms are known from
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
and
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. Unlike
lepidosauria The Lepidosauria (, from Greek meaning ''scaled lizards'') is a Order (biology), superorder or Class (biology), subclass of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata also includes Lizard, lizards and Snake, snakes. Sq ...
n reptiles, pelycosaurs might have lacked reptilian epidermal
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...
. Fossil evidence from some
varanopids Varanopidae is an extinct family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus ''Varanus'') and may have filled a similar niche. T ...
shows that parts of the skin were covered in rows of
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amph ...
s, presumably overlain by horny
scute A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "Scutum (shield), shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of Bird anatomy#Scales, birds. The ter ...
s. The belly was covered in rectangular scutes, looking like those present in
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s. Parts of the skin ''not'' covered in scutes might have had naked, glandular skin like that found in some mammals. Dermal scutes are also found in a diverse number of extant mammals with conservative body types, such as in the tails of some
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s, sengis, moonrats, the
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 126 species in 18 genera. Opossums originated in South America and entered North A ...
s, and other
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s, and as regular dermal armour with underlying bone in the
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
. At least two pelycosaur
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 ...
s independently evolved a tall
sail A sail is a tensile structure, which is made from fabric or other membrane materials, that uses wind power to propel sailing craft, including sailing ships, sailboats, windsurfers, ice boats, and even sail-powered land vehicles. Sails may b ...
, consisting of elongated vertebral spines: the
edaphosaurid Edaphosauridae is a family of mostly large (up to or more) Late Carboniferous to Early Permian synapsids. Edaphosaur fossils are so far known only from North America and Europe. Characteristics They were the earliest known herbivorous amniotes ...
s and the
sphenacodontid Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family of sphenacodontoid synapsids. Small to large, advanced, carnivorous, Late Pennsylvanian to middle Permian "pelycosaurs". The most recent one, ''Dimetrodon angelensis'', is ...
s. In life, this may have been covered by skin, and likely functioned as a
thermoregulatory Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
device or as a
mating display A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), ...
.


Taxonomy

In phylogenetic nomenclature, "Pelycosauria" is not used formally, since it does not constitute a group of ''all'' organisms descended from some common ancestor (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 ...
), because the group specifically excludes the therapsids which are descended from pelycosaurs. Instead, it represents a paraphyletic "grade" of basal synapsids leading up to the clade Therapsida. In 1940, the group was reviewed in detail, and every species known at the time described, with many illustrated, in an important monograph by
Alfred Sherwood Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
and
Llewellyn Price Llewellyn Ivor Price (October 9, 1905 – June 9, 1980) was one of the first Brazilian paleontologists. His work contributed not only to the development of Brazilian but also to global paleontology. He collected ''Staurikosaurus'' in 1936, th ...
. In traditional classification, the order Pelycosauria is
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 ...
in that the therapsids (the "higher" synapsids) have emerged from them. That means Pelycosauria is a grouping of animals that does not contain all descendants of its common ancestor, as is often required by
phylogenetic nomenclature Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional method, by which taxon names are defined by a '' type'', which c ...
. In
evolutionary taxonomy Evolutionary taxonomy, evolutionary systematics or Darwinian classification is a branch of biological classification that seeks to classify organisms using a combination of phylogenetic relationship (shared descent), progenitor-descendant relati ...
, Therapsida is a separated order from Pelycosauria, 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 (having evolved from therapsids) are separated from both as their own class. This use has not been recommended by a majority of systematists since the 1990s, but several paleontologists nevertheless continue using this word. The following classification was presented by Benton in 2004. * Order Pelycosauria * ** Suborder
Caseasauria Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insect ...
*** Family
Eothyrididae Eothyrididae is an extinct family of very primitive, insectivorous synapsids. Only three genera are known, '' Eothyris'', '' Vaughnictis'' and '' Oedaleops'', all from the early Permian of North America. Their main distinguishing feature is the l ...
*** Family
Caseidae Caseidae are an Extinction, extinct Family (biology), family of Basal (phylogenetics), basal synapsids that lived from the Late Carboniferous to Middle Permian between about 300 and 265 million years ago. Fossils of these animals come from the so ...
** Suborder
Eupelycosauria Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308million years ago during the Early Pennsylvanian epoch, with the fo ...
*** Family
Varanopidae Varanopidae is an extinct family (biology), family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus ''Varanus'') and may have filled ...
*** Family
Ophiacodontidae Ophiacodontidae is an extinct family of early synapsids from the Carboniferous and Permian. '' Archaeothyris'', and '' Clepsydrops'' were among the earliest ophiacodontids, appearing in the Late Carboniferous. Ophiacodontids are among the most b ...
*** Family
Edaphosauridae Edaphosauridae is a family of mostly large (up to or more) Late Carboniferous to Early Permian synapsids. Edaphosaur fossils are so far known only from North America and Europe. Characteristics They were the earliest known herbivorous amniotes ...
*** Infraorder
Sphenacodontia Sphenacodontia is a stem-based taxon, stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes ''Haptodus baylei'', ''Haptodus garnettensis'' and ''Sphenacodon ferox'', but not ''Edaphos ...
**** Family
Sphenacodontidae Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family (biology), family of sphenacodontoidea, sphenacodontoid synapsids. Small to large, advanced, carnivore, carnivorous, Late Pennsylvanian to Guadalupian, middle Permian "pelyc ...
* Order Therapsida


See also

*
Evolution of mammals The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian (geology), Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synaps ...
*
List of pelycosaurs This list of pelycosaurs is an attempt to create a comprehensive listing of all Genus, genera that have ever been included in the synapsida excluding therapsida and purely vernacular terms. The list includes all commonly accepted genera, but also g ...
*
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 ...


Notes


References

*


External links

* * {{Taxonbar, from=Q319520 Synapsida Paraphyletic groups Taxa named by Edward Drinker Cope