Therapsids
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Therapsida is a clade comprising a major group of
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 ...
n
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 that includes
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 ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, resulting in a more "standing"
quadrupedal Quadrupedalism is a form of Animal locomotion, locomotion in which animals have four legs that are used to weight-bearing, bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four l ...
posture, as opposed to the lower sprawling posture of many
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and
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. Therapsids evolved from earlier synapsids commonly called "
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", specifically within the
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 ...
, more than 279.5 million years ago. They replaced the pelycosaurs as the dominant large land animals in the
Guadalupian The Guadalupian is the second and middle Series (stratigraphy), series/Epoch (geology), epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico an ...
through to the Early Triassic. In the aftermath of the
Permian–Triassic extinction event The Permian–Triassic extinction event (also known as the P–T extinction event, the Late Permian extinction event, the Latest Permian extinction event, the End-Permian extinction event, and colloquially as the Great Dying,) was an extinction ...
, therapsids declined in relative importance to the rapidly diversifying
archosaurian Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistic sense of the t ...
sauropsid 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 dino ...
s (
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
ns,
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, etc.) during the Middle Triassic. The therapsids include the
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s, the group that gave rise to mammals (
Mammaliaformes Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as ...
) in the Late Triassic around 225 million years ago, the only therapsid clade that survived beyond the end of the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
. The only other group of therapsids to have survived into the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
, the
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
s, became extinct towards the end of the period. The last surviving group of non-mammaliaform cynodonts were the
Tritylodontidae Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, with several mammalian traits including erect limbs, endothermy, and some d ...
, which became extinct during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
.


Characteristics


Jaw and teeth

Therapsids'
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 were larger than those of the pelycosaurs. The jaws of some therapsids were more complex and powerful, and the
teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
were differentiated into frontal
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s for nipping, great lateral canines for puncturing and tearing, and
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
for shearing and chopping food.


Posture

Therapsid legs were positioned more vertically beneath their bodies than were the sprawling legs of
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and pelycosaurs. Also compared to these groups, the feet were more symmetrical, with the first and last toes short and the middle toes long, an indication that the foot's
axis An axis (: axes) may refer to: Mathematics *A specific line (often a directed line) that plays an important role in some contexts. In particular: ** Coordinate axis of a coordinate system *** ''x''-axis, ''y''-axis, ''z''-axis, common names ...
was placed parallel to that of the animal, not sprawling out sideways. This orientation would have given a more
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 ...
-like gait than the
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
-like gait of the pelycosaurs.


Physiology

The physiology of therapsids is poorly understood. Most Permian therapsids had a pineal foramen, indicating that they had a
parietal eye A parietal eye (third eye, pineal eye) is a part of the epithalamus in some vertebrates. The eye is at the top of the head; is photoreceptive; and is associated with the pineal gland, which regulates circadian rhythmicity and hormone production ...
like many modern reptiles and amphibians. The parietal eye serves an important role in thermoregulation and the
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural oscillation that repeats roughly every 24 hours. Circadian rhythms can refer to any process that originates within an organism (i.e., Endogeny (biology), endogenous) and responds to the env ...
of ectotherms, but is absent in modern mammals, which are
endothermic An endothermic process is a chemical or physical process that absorbs heat from its surroundings. In terms of thermodynamics, it is a thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, ...
. Near the end of the Permian, dicynodonts,
therocephalia Therocephalia is an extinct clade of therapsids (mammals and their close extinct relatives) from the Permian and Triassic periods. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their te ...
ns and cynodonts show parallel trends towards loss of the pineal foramen, and the foramen is completely absent in probainognathian cynodonts. Evidence from oxygen isotopes, which are correlated with body temperature, suggests that most Permian therapsids were ectotherms and that endothermy evolved convergently in dicynodonts and cynodonts near the end of the Permian. In contrast, evidence from histology suggests that endothermy is shared across Therapsida, whereas estimates of blood flow rate and lifespan in the mammaliaform ''
Morganucodon ''Morganucodon'' ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represent ...
'' suggest that even early mammaliaforms had reptile-like metabolic rates. Evidence for respiratory turbinates, which have been hypothesized to be indicative of endothermy, was reported in the therocephalian '' Glanosuchus'', but subsequent study showed that the apparent attachment sites for turbinates may simply be the result of distortion of the skull.


Integument

The evolution of integument in therapsids is poorly known, and there are few fossils that provide direct evidence for the presence or absence of fur. The most basal synapsids with unambiguous direct evidence of fur are docodonts, which are mammaliaforms very closely related to crown-group mammals. Two "mummified" juvenile specimens of the dicynodont '' Lystrosaurus murrayi'' preserve skin impressions; the skin is hairless, leathery, and dimpled, somewhat comparable to elephant skin. Fossilized facial skin from the dinocephalian ''
Estemmenosuchus ''Estemmenosuchus'' (meaning "crowned crocodile" in Greek language, Greek) is an extinct genus of large, early omnivore, omnivorous therapsid. It is believed and interpreted to have lived during the middle part of the Middle Permian around 267 m ...
'' has been described as showing that the skin was glandular and lacked both scales and hair.
Coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
s containing what appear to be hairs have been found from the
Late Permian 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 ...
. Though the source of these hairs is not known with certainty, they may suggest that hair was present in at least some Permian therapsids. The closure of the pineal foramen in probainognathian cynodonts may indicate a mutation in the regulatory gene Msx2, which is involved in both the closure of the skull roof and the maintenance of hair follicles in mice. This suggests that hair may have first evolved in probainognathians, though it does not entirely rule out an earlier origin of fur. Whiskers probably evolved in probainognathian cynodonts. Some studies had inferred an earlier origin for whiskers based on the presence of foramina on the snout of therocephalians and early cynodonts, but the arrangement of foramina in these taxa actually closely resembles lizards, which would make the presence of mammal-like whiskers unlikely.


Evolutionary history

Therapsids
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
from a group of pelycosaurs called sphenacodonts. Therapsids became the dominant land animals in the Middle
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 ...
, displacing the pelycosaurs. Therapsida consists of four major
clades In biology, a clade (), also known as a 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 to taxonomy ...
: the
dinocephalia Dinocephalians (terrible heads) are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. ...
ns, the herbivorous
anomodont Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores. Anomodonts were very diverse during the Middle Pe ...
s, the carnivorous
biarmosuchia Biarmosuchia is an extinct clade of non-mammalian synapsids from the Permian. Biarmosuchians are the most basal group of the therapsids. They were moderately-sized, lightly built carnivores, intermediate in form between basal sphenacodont " pel ...
ns, and the mostly carnivorous
theriodont The theriodonts (clade Theriodontia) are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts. Naming In 1876, ...
s. After a brief burst of evolutionary diversity, the dinocephalians died out in the later Middle Permian (
Guadalupian The Guadalupian is the second and middle Series (stratigraphy), series/Epoch (geology), epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico an ...
) but the anomodont
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
s as well as the theriodont
gorgonopsia Gorgonopsia (from the Greek Gorgon, a mythological beast, and 'aspect') is an extinct clade of Saber-toothed predator, sabre-toothed therapsids from the Middle Permian, Middle to the Upper Permian, roughly between 270 and 252 million years ago. ...
ns and therocephalians flourished, being joined at the very end of the Permian by the first of the
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s. Like all land animals, the therapsids were seriously affected by the
Permian–Triassic extinction event The Permian–Triassic extinction event (also known as the P–T extinction event, the Late Permian extinction event, the Latest Permian extinction event, the End-Permian extinction event, and colloquially as the Great Dying,) was an extinction ...
, with the very successful gorgonopsians and the biarmosuchians dying out altogether and the remaining groups—dicynodonts, therocephalians and cynodonts—reduced to a handful of species each by the earliest
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
. Surviving dicynodonts were represented by two families of disaster taxa ( Lystrosauridae and Myosauridae), the scarcely known '' Kombuisia'', and a single group of large stocky
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s, the Kannemeyeriiformes, which were the only dicynodont lineage to thrive during the Triassic. They and the medium-sized cynodonts (including both carnivorous and herbivorous forms) flourished worldwide throughout the Early and Middle Triassic. They disappear from the fossil record across much of
Pangea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia (continent), Siberia during the Carboniferous period ...
at the end of the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
(Late Triassic), although they continued for some time longer in the wet equatorial band and the south. Some exceptions were the still further derived
eucynodonts Eucynodontia ("true dog teeth") is a clade of cynodont therapsids including mammals and most non-mammalian cynodonts. The oldest eucynodonts are known from the Early Triassic and possibly Late Permian. Eucynodontia includes two major subgroups, ...
. At least three groups of them survived. They all appeared in the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
period. The extremely mammal-like family,
Tritylodontidae Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, with several mammalian traits including erect limbs, endothermy, and some d ...
, survived into the Early
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. Another extremely mammal-like family,
Tritheledontidae Tritheledontidae, the tritheledontids or ictidosaurs, is an extinct family of small to medium-sized (about 10 to 20 cm long) cynodonts. They were highly mammal-like, specialized cynodonts, although they still retained a few reptile-like ...
, are unknown later than the Early
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
.
Mammaliaformes Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as ...
was the third group, including ''Morganucodon'' and similar animals. Some taxonomists refer to these animals as "mammals", though most limit the term to the mammalian
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
. The non-eucynodont cynodonts survived the Permian–Triassic extinction; ''
Thrinaxodon ''Thrinaxodon'' is an extinct genus of cynodonts which lived in what are now South Africa and Antarctica during the Late Permian - Early Triassic. ''Thrinaxodon'' lived just before, during, and right after the Permian–Triassic mass extinction ...
'', ''
Galesaurus ''Galesaurus'' (from the Greek roots for 'weasel' and 'lizard') is an extinct genus of carnivorous cynodont therapsid that lived between the Induan and the Olenekian stages of the Early Triassic in what is now South Africa. It was incorrectly cl ...
'' and ''
Platycraniellus ''Platycraniellus'' is an extinct genus of carnivorous cynodonts from the Early Triassic. It is known from the ''Lystrosaurus'' Assemblage Zone of the Normandien Formation in South Africa. ''P. elegans'' is the only species in this genus based ...
'' are known from the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
. By the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
, however, only the eucynodonts remained. The therocephalians, relatives of the cynodonts, managed to survive the Permian–Triassic extinction and continued to diversify through the Early Triassic period. Approaching the end of the period, however, the therocephalians were in decline to eventual extinction, likely outcompeted by the rapidly diversifying Saurian lineage of
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The earliest traditionally identified diapsids, the araeosc ...
s, equipped with sophisticated respiratory systems better suited to the very hot, dry and oxygen-poor world of the End-Triassic. Dicynodonts were among the most successful groups of therapsids during the Late Permian; they survived through to near the end of the Triassic. Mammals are the only living therapsids. The mammalian crown group, which evolved in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
period, radiated from a group of mammaliaforms that included the docodonts. The mammaliaforms themselves evolved from
probainognathia Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and most b ...
ns, a lineage of the eucynodont suborder.


Classification

Six major groups of therapsids are generally recognized: Biarmosuchia, Dinocephalia, Anomodontia, Gorgonopsia, Therocephalia and Cynodontia. A clade uniting therocephalians and cynodonts, called
Eutheriodontia Eutheriodontia is a clade of therapsids which appear during the Middle Permian and which includes therocephalians and cynodonts, this latter group including mammals and related forms. With the dicynodonts, they form one of two lineages of ther ...
, is well supported, but relationships among the other four clades are controversial. The most widely accepted hypothesis of therapsid relationships, the Hopson and Barghausen paradigm, was first proposed in 1986. Under this hypothesis, biarmosuchians are the earliest-diverging major therapsid group, with the other five groups forming the Eutherapsida, and within Eutherapsida, gorgonopsians are the sister taxon of eutheriodonts, together forming the
Theriodontia The theriodonts (clade Theriodontia) are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts. Naming In 1876, R ...
. Hopson and Barghausen did not initially come to a conclusion about how dinocephalians, anomodonts and theriodonts were related to each other, but subsequent studies suggested that anomodonts and theriodonts should be classified together as the Neotherapsida. However, there remains debate over these relationships; in particular, some studies have suggested that anomodonts, not gorgonopsians, are the sister taxon of Eutheriodontia, other studies have found dinocephalians and anomodonts to form a clade, and both the phylogenetic position and monophyly of Biarmosuchia remain controversial. In addition to the six major groups, there are several other lineages and species of uncertain classification. ''
Raranimus ''Raranimus'' is an extinct genus of therapsids of the Middle Permian. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the Qingtoushan Formation, outcropping in the Qilian Mountains of Gansu, China. Th ...
'' from the early Middle Permian of China is likely to be the earliest-diverging known therapsid. ''
Tetraceratops ''Tetraceratops insignis'' ("four-horned face emblem") is an extinct synapsid from the Early Permian that was formerly considered the earliest known representative of Therapsida, a group that includes mammals and their close extinct relatives. It ...
'' from the Early Permian of the United States has been hypothesized to be an even earlier-diverging therapsid, but more recent study has suggested it is more likely to be a non-therapsid sphenacodontian.


Biarmosuchia

Biarmosuchia is the most recently recognized therapsid clade, first recognized as a distinct lineage by Hopson and Barghausen in 1986 and formally named by Sigogneau-Russell in 1989. Most biarmosuchians were previously classified as gorgonopsians. Biarmosuchia includes the distinctive Burnetiamorpha, but support for the monophyly of Biarmosuchia is relatively low. Many biarmosuchians are known for extensive cranial ornamentation.


Dinocephalia

Dinocephalia comprises two distinctive groups, the
Anteosauria Anteosaurs are a group of large, primitive carnivore, carnivorous dinocephalian therapsids with large canines and incisors and short limbs, that are known from the Middle Permian of South Africa, Russia, China, and Brazil. Some grew very large, w ...
and Tapinocephalia. Historically, carnivorous dinocephalians, including both anteosaurs and titanosuchids, were called titanosuchians and classified as members of Theriodontia, while the herbivorous Tapinocephalidae were classified as members of Anomodontia.


Anomodontia

Anomodontia includes the dicynodonts, a clade of tusked, beaked herbivores, and the most diverse and long-lived clade of non-cynodont therapsids. Other members of Anomodontia include ''
Suminia ''Suminia'' is an extinct genus of basal anomodont that lived during the Tatarian age of the late Permian, spanning approximately from 268–252 Ma.Rybczynski N. 2000. Cranial anatomy and phylogenetic position of Suminia getmanovi, a basal anomod ...
,'' which is thought to have been a climbing form.


Gorgonopsia

Gorgonopsia is an abundant but morphologically homogeneous group of
saber-toothed predator A saber-tooth (alternatively spelled sabre-tooth) is any member of various extinct groups of predatory therapsids, predominantly carnivoran mammals, that are characterized by long, curved saber-shaped canine teeth which protruded from the mouth ...
s.


Therocephalia

It has been suggested that Therocephalia might not be monophyletic, with some species more closely related to cynodonts than others. However, most studies regard Therocephalia as monophyletic.


Cynodontia

Cynodonts are the most diverse and longest-lived of the therapsid groups, as Cynodontia includes mammals. Cynodonts are the only major therapsid clade to lack a Middle Permian fossil record, with the earliest-known cynodont being '' Charassognathus'' from the
Wuchiapingian In the geologic timescale, the Wuchiapingian or Wujiapingian (from in the Liangshan area of Hanzhong, Shaanxi Province ) is an age or stage of the Permian. It is also the lower or earlier of two subdivisions of the Lopingian Epoch or S ...
age of the Late Permian. Non-mammalian cynodonts include both carnivorous and herbivorous forms.


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 ...
*
Timeline of the evolutionary history of life The timeline of the evolutionary history of life represents the current scientific theory outlining the major events during the development of life on planet Earth. Dates in this article are consensus estimates based on scientific evidence, main ...
*
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


Further reading

* Benton, M. J. (2004). ''Vertebrate Palaeontology'', 3rd ed., Blackwell Science. * Carroll, R. L. (1988). ''Vertebrate Paleontology & Evolution''. W. H. Freeman & Company, New York. * Kemp, T. S. (2005). ''The origin and evolution of mammals''.
Oxford University Press Oxford University Press (OUP) is the publishing house of the University of Oxford. It is the largest university press in the world. Its first book was printed in Oxford in 1478, with the Press officially granted the legal right to print books ...
. * Romer, A. S. (1966). ''Vertebrate Paleontology''. University of Chicago Press, 1933; 3rd ed. * Bennett, A. F., & Ruben, J. A. (1986).
The metabolic and thermoregulatory status of therapsids
." In ''The ecology and biology of mammal-like reptiles''. Smithsonian Institution Press, Washington, DC, 207-218. * * Ross, R.P., Ross, C.A. (2023). ''Permian Period, geochronology''. Encyclopedia Britannica. https://www.britannica.com/science/Permian-Period * Padian, Kevin (2013-09). "A Review of "Forerunners of Mammals: Radiation, Histology, Biology"". ''Journal of Vertebrate Paleontology''. 33 (5): 1250–1251.


External links

*
Therapsida: Mammals and extinct relatives
''Tree of Life'' *

''Palaeos'' {{Taxonbar, from=Q322241 Cisuralian first appearances Extant Permian first appearances Taxa named by Robert Broom