Cynodont
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The cynodonts () (
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 term, ...
Cynodontia) are a clade of
eutheriodont 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 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 that first appeared in the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
(approximately 260 mya), and extensively diversified after the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
. Cynodonts had a wide variety of lifestyles, including carnivory and herbivory.
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 or ...
s are cynodonts, as are their extinct ancestors and close relatives, having evolved from advanced probainognathian cynodonts during the Late Triassic. All other cynodont lines went extinct, with the last known non-mammalian cynodont group, 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, bearing several mammalian traits like erect limbs, endothermy and detail ...
, having its youngest records in the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
.


Description

Early cynodonts have many of the skeletal characteristics of
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 or ...
s. The teeth were fully differentiated and the braincase bulged at the back of the head. Outside of some
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. ...
mammals (notably the
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
ns), all cynodonts probably laid eggs. The
temporal fenestrae The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
were much larger than those of their ancestors, and the widening of the
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
in a more mammal-like skull would have allowed for more robust jaw musculature. They also have the
secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...
that other primitive therapsids lacked, except the therocephalians, who were the closest relatives of cynodonts. (However, the secondary palate of cynodonts primarily comprises the maxillae and palatines as in mammals, whereas the secondary palate of the therocephalians primarily comprises the maxillae and the vomer.) The
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
was the largest bone in their lower jaw. The cynodonts probably had some form of
warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The onl ...
metabolism. This has led to many reconstructions of cynodonts as having
fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
. Being endothermic they may have needed it for
thermoregulation 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 ...
, but fossil evidence of their fur (or lack thereof) has been elusive. Modern mammals have
Harderian gland The Harderian gland is a gland found within the eye's orbit that occurs in tetrapods (reptiles, amphibians, birds and mammals) that possess a nictitating membrane. The gland can be compound tubular or compound tubuloalveolar, and the fluid it s ...
s secreting lipids to coat their fur, but the telltale imprint of this structure is only found from the primitive mammal '' Morganucodon'' and onwards. Nonetheless, recent studies on
Permian The Permian ( ) is a geologic period and 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 251.9 Mya. It is the last period of the Paleoz ...
synapsid
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 is de ...
s show that more basal therapsids may have had fur, and at any rate fur was already present in Mammaliaformes such as '' Castorocauda'' and ''
Megaconus ''Megaconus'' is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, ''Megaconus mammaliaformis'' was first described in the journal ''Nature'' in 2013. ''Meg ...
''. Early cynodonts had numerous small
foramina In anatomy and osteology, a foramen (;Entry "foramen"
in
on their snout bones, similar to reptiles. This suggests that they had immobile, non-muscular lips like those of lizards, and lacked muscular cheeks. As a muscular, mobile face is necessary to perform whisking movements and to avoid damage to
whisker Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser ...
s, it is unlikely that early cynodonts had whiskers. In prozostrodontian cynodonts, the group that includes mammals, the foramina are replaced by a single large infraorbital foramen, which indicates that the face had become muscular and that whiskers would have been present. Derived cynodonts developed
epipubic Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them). They first o ...
bones. These served to strengthen the torso and support abdominal and hindlimb musculature, aiding them in the development of an erect gait, but at the expense of prolonged pregnancy, forcing these animals to give birth to highly
altricial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
young as in modern marsupials and
monotremes Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials ( Metatheria). Monotremes are typified by structural differences in their br ...
. Only
placentals Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
, and perhaps ''
Megazostrodon ''Megazostrodon'' is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old.
'' and ''
Erythrotherium ''Erythrotherium'' (meaning "red beast") is an extinct genus of basal mammaliaforms from the Late Triassic to Lower Jurassic. It is related to ''Morganucodon''. Only one species is recorded, ''Erythrotherium parringtoni'', from Red Beds, Stormber ...
'', would lose these. A specimen of ''
Kayentatherium ''Kayentatherium'' is an extinct genus of tritylodontid cynodonts that lived during the Early Jurassic. It is one of two tritylodonts from the Kayenta Formation of northern Arizona, United States. ''Kayentatherium'' means "Kayenta Beast", and is ...
'' does indeed demonstrate that at least tritylodontids already had a fundamentally marsupial-like reproductive style, but produced much higher litters at around 38 perinates or possibly eggs. Cynodonts are the only known synapsid lineage to have produced aerial locomotors, with gliding and flying being known in haramiyidans and various mammal groups. The largest known non-mammalian cynodont is '' Scalenodontoides'', a
traversodontid Traversodontidae is an extinct family of herbivorous cynodonts. Traversodonts were primarily Gondwanan, with many species known from Africa and South America. Recently, traversodonts have also been found from Europe and eastern North America. Tra ...
, which has been estimated to have a maximum skull length of approximately based on a fragmentary specimen.


Evolutionary history

The closest relatives of cynodonts are
therocephalia Therocephalia is an extinct suborder of eutheriodont therapsids (mammals and their close relatives) from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their ...
ns, with which they form the clade
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 therap ...
. The earliest cynodonts are known early
Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...
(early
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 Series. The ...
) aged sediments of the ''Tropidostoma'' Assemblage Zone, in the
Karoo Supergroup The Karoo Supergroup is the most widespread stratigraphic unit in Africa south of the Kalahari Desert. The supergroup consists of a sequence of units, mostly of nonmarine origin, deposited between the Late Carboniferous and Early Jurassic, a peri ...
of South Africa, belonging to the basal family Charassognathidae. Fossils of Permian cynodonts are relatively rare outside of South Africa, with the most widely distributed genus being ''
Procynosuchus ''Procynosuchus'' (Greek: "Before dog crocodile") is an extinct genus of cynodonts from the Late Permian. It is considered to be one of the earliest and most basal cynodonts. It was 60 cm (2 ft) long. Remains of ''Procynosuchus'' have ...
,'' which is known from South Africa, Germany, Tanzania, Zambia, and possibly Russia. Cynodonts expanded rapidly in diversity after the Permian-Triassic extinction event. Post-Early Triassic cynodonts were dominated by members of the advanced clade
Eucynodontia 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, ...
, which has two main subdivisions, the predominantly herbivorous
Cynognathia Cynognathia ("dog jaw") is one of two major clades of cynodonts, the other being Probainognathia. Cynognathians included the large carnivorous genus '' Cynognathus'' and the herbivorous traversodontids. Cynognathians can be identified by several ...
and the predominantly carnivorous Probainognathia. During the Early and Middle Triassic, cynodont diversity was dominated by members of Cynognathia, and members of Probainognathia would not become prominent until the Late Triassic (early
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
). Almost all Middle Triassic cynodonts are known from Gondwana, with only one genus ('' Nanogomphodon'') having been found in the Northern Hemisphere. Among the most dominant groups of Middle and Late Triassic cynodonts is the herbivorous
Traversodontidae Traversodontidae is an extinct family of herbivorous cynodonts. Traversodonts were primarily Gondwanan, with many species known from Africa and South America. Recently, traversodonts have also been found from Europe and eastern North America. Tra ...
, predominantly in Gondwana, which reached a peak diversity in the Late Triassic. Mammaliaformes originated from probainognathian cynodonts during the Late Triassic. Early Mammaliaformes were small bodied insectivores. Only two groups of non-mammaliaform cynodonts existed beyond the end of the Triassic, both belonging to Probainognathia. The first is the insectivorous
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 ana ...
, which briefly lasted into the Early Jurassic. The second is the herbivorous
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, bearing several mammalian traits like erect limbs, endothermy and detail ...
, which first appeared in the latest Triassic, which were abundant and diverse during the Jurassic, predominantly in the Northern Hemisphere, persisted into the Early Cretaceous in Asia. The geologically youngest known tritylodontid is '' Fossiomanus'' from the 120 million years old
Jiufotang Formation The Jiufotang Formation ( Chinese: 九佛堂组, pinyin: ''jiǔfótáng zǔ'') is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see ...
of China. During their
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
, the number of cynodont
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
bones reduced. This move towards a single bone for the mandible paved the way for other bones in the jaw, the
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 oth ...
and angular, to migrate to the cranium, where they function as parts of the mammalian hearing system. Cynodonts also developed a
secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...
in the roof of the mouth. This caused air flow from the nostrils to travel to a position in the back of the mouth instead of directly through it, allowing cynodonts to chew and breathe at the same time. This characteristic is present in all mammals.


Taxonomy

Richard Owen named Cynodontia in 1861, which he assigned to Anomodontia as a family.
Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a British- South African doctor and palaeontologist. He qualified as a medical practitioner in 1895 and received his DSc in 1905 from the University of Glasgow. From 1903 to 1910, he ...
(1913) reranked Cynodontia as an infraorder, since retained by others, including Colbert and Kitching (1977), Carroll (1988), Gauthier ''et al.'' (1989), and Rubidge and Cristian Sidor (2001). Olson (1966) assigned Cynodontia to
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, Ric ...
, Colbert and Kitching (1977) to Theriodontia, and Rubridge and Sidor (2001) to
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 therap ...
.
William King Gregory William King Gregory (May 19, 1876 – December 29, 1970) was an American zoologist, renowned as a primatologist, paleontologist, and functional and comparative anatomist. He was an expert on mammalian dentition, and a leading contributor t ...
(1910), Broom (1913), Carroll (1988), Gauthier ''et al.'' (1989), Hopson and Kitching (2001) and Botha ''et al.'' (2007) all considered Cynodontia as belonging to Therapsida. Botha ''et al.'' (2007) seems to have followed Owen (1861), but without specifying taxonomic rank.


Phylogeny

Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
from Ruta, Botha-Brink, Mitchell and Benton (2013) showing one hypothesis of cynodont relationships:


Distribution

Cynodonts have been found in South America, India, Africa, Antarctica, Asia, Europe and North America.


See also

*
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
*
Prehistoric mammal This is an incomplete list of prehistoric mammals. It does not include extant mammals or recently extinct mammals. For extinct primate species, see: list of fossil primates.Mikko's Phylogeny Archiv Mammaliaformes ' *Genus †'' Adelobasileus ...
*
Tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
* Triassic-Jurassic extinction event


References


Further reading

* *Davis, Dwight (1961). "Origin of the Mammalian Feeding Mechanism". Am. Zoologist, 1:229–234.


External links


Palaeos cynodonts



Bennett and Ruben 1986. The Metabolic and Thermoregulatory Status of Therapsids

BBC cynodonts
{{Taxonbar, from=Q131801 Lopingian first appearances Extant Permian first appearances Fossil taxa described in 1861 Taxa named by Richard Owen