Pelycosaurs
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Pelycosaur ( ) is an older term for basal or primitive Late
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s, excluding the
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
, but this is now thought to be incorrect, and seen as outdated. Because it excludes the advanced synapsid group
Therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
a, the term is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
and contrary to modern formal naming practice. Thus the name ''pelycosaurs'', similar to the term ''mammal-like reptiles'', had fallen 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 used where needed, instead.


Etymology

The term ''pelycosaur'' has been fairly well abandoned by paleontologists because it no longer matches the features that distinguish a clade. The modern word was created from Greek meaning 'wooden bowl' or 'axe' and meaning 'lizard'. The intended etymology presents a problem, since ''pelycosaur'' as created basically means 'basin lizard', but there is a ambiguity between several original Greek words in that is 'wooden bowl' and is 'basin', whereas is 'ax' (like 'double-edged ax'). Pelycosauria is a
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
taxon because it excludes the
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s. 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 fos ...
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 cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
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 insec ...
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
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur o ...
, having differentiated teeth and a developing hard palate. The pelycosaurs appeared during the Late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
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 ...
, 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 Ar ...
. They were succeeded by the
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s.


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 ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodont ...
'', '' Sphenacodon'', ''
Edaphosaurus ''Edaphosaurus'' (, meaning "pavement lizard" for dense clusters of 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 to ...
'', and '' Ophiacodon''. Pelycosaur fossils have been found mainly in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
, although some small, late-surviving forms are known from
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-ei ...
and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
. Unlike
lepidosauria The Lepidosauria (, from Greek meaning ''scaled lizards'') is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes snakes, lizards, and amphisbaenians. Squamata contains over 9,000 species, m ...
n reptiles, pelycosaurs 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 w ...
. Fossil evidence from some
varanopids Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an '' Archaeothyris''-like synapsid in the Late Carbonife ...
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 amp ...
s, presumably overlain by horny
scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "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 birds. The term is also used to describe the anterior po ...
s. The belly was covered in rectangular scutes, looking like those present in
crocodile Crocodiles (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 even more loosely to include all extant me ...
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 Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s, sengis,
moonrat The moonrat (''Echinosorex gymnura'') is a southeast Asian species of mammal in the family Erinaceidae (hedgehogs and gymnures). It is the only species in the genus ''Echinosorex''. The moonrat is a fairly small, primarily carnivorous animal whi ...
s, 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 93 species in 18 genera. Opossums originated in South America and entered No ...
s and other
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s, and as regular dermal armour with underlying bone in the
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, alo ...
. At least two pelycosaur
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
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 ma ...
, consisting of elongated vertebral spines: the edaphosaurids and the
sphenacodontid Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family (biology), family of small to large, advanced, carnivore, carnivorous, Late Pennsylvanian to Guadalupian, middle Permian pelycosaurs. The most recent one, ''Dimetrodon ang ...
s. In life, this would 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 temperatur ...
device or as a mating display.


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 A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
), 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. In traditional classification, the order Pelycosauria is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
in that the
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s (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 approach, in which taxon names are defined by a '' type'', which ...
. 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 relat ...
, Therapsida is a separated order from Pelycosauria, and
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
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 * ** 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 la ...
** Family
Caseidae Caseidae are an extinct family of 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 south-central part of the United States (Texas, Okla ...
** Family
Varanopidae Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an '' Archaeothyris''-like synapsid in the Late Carbonife ...
** Family
Ophiacodontidae Ophiacodontidae is an extinct family of early eupelycosaurs from the Carboniferous and Permian. '' Archaeothyris'', and ''Clepsydrops'' were among the earliest ophiacodontids, appearing in the Late Carboniferous. Ophiacodontids are among the mos ...
** Family
Edaphosauridae Edaphosauridae is a family of mostly large (up to 3 meters 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 a ...
** Family
Sphenacodontidae Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family of small to large, advanced, carnivorous, Late Pennsylvanian to middle Permian pelycosaurs. The most recent one, ''Dimetrodon angelensis'', is from the late Kungurian or ...
* Order
Therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
a


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 sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked l ...
*
List of pelycosaurs This list of pelycosaurs is an attempt to create a comprehensive listing of all genera that have ever been included in the synapsida excluding therapsida and purely vernacular terms. The list includes all commonly accepted genera, but also genera t ...
*
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 animals with vertebrae or a notochord. It also tries to connect, by us ...


Notes


References

*


External links

* * {{Taxonbar, from=Q319520 Synapsids Paraphyletic groups