Cynodont

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Cynodonts
Temporal range: Late Permian-Recent, 260–0 Ma
Belesodon magnificus.JPG
Fossil of Chiniquodon in the Staatliches Museum für Naturkunde Stuttgart
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Therapsida
Clade: Eutheriodontia
Clade: Cynodontia
Owen, 1861
Clades

The cynodonts (lit.'dog-teeth') (clade Cynodontia) are a clade of therapsids that first appeared in the Late Permian (approximately 260 Ma), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety of lifestyles, including carnivory and herbivory. Mammals (including humans) 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 having its last records in the Early Cretaceous.

Taxonomy[edit]

Richard Owen named Cynodontia in 1861, which he assigned to Anomodontia as a family.[1] Robert Broom (1913) reranked Cynodonia as an infraorder, since retained by others, including Colbert and Kitching (1977), Carroll (1988), Gauthier et al. (1989), and Rubidge and Cristian Sidor (2001).[2] Olson (1966) assigned Cynodontia to Theriodontia, Colbert and Kitching (1977) to Theriodontia, and Rubridge and Sidor (2001) to Eutheriodontia. William King Gregory (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.[3][4]


Sauropsida (reptiles)

Synapsida

Caseasauria

†"Pelycosaurs"

Therapsida

Biarmosuchia

Dinocephalia

Anomodontia

Gorgonopsia

Therocephalia

Cynodontia

Evolutionary history[edit]

Thrinaxodon from the Early Triassic of South Africa

Together with the extinct gorgonopsians and the therocephalians, the cynodonts themselves are part of a group of therapsids called theriodonts.

The earliest cynodonts are known early Lopingian (early Wuchiapingian) aged sediments of the Tropidostoma Assemblage Zone, in the Karoo Supergroup 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, which is known from South Africa, Germany, Tanzania, Zambia, and possibly Russia.[5]

Cynodonts expanded rapidly in diversity after the Permian-Triassic extinction event. Post-Early Triassic cynodonts were dominated by members of the advanced clade Eucynodontia, which has two main subdivisions, the predominantly herbivorous Cynognathia 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).[6] 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, predominantly in Gondwana, which reached a peak diversity in the Late Triassic. Mammaliaformes originated from probainognathian cynodonts during the Late Triassic. Only two groups of non-mammaliaform cynodonts existed beyond the end of the Triassic, both belonging to Probainognathia. The first is the insectivorous Tritheledontidae, which briefly lasted into the Early Jurassic. The second is the herbivorous Tritylodontidae, which first appeared in the latest Triassic, which were abundant and diverse during the Jurassic, predominantly in the Northern Hemisphere, and would persist into the Early Cretaceous.[7]

During their evolution, the number of cynodont jaw bones reduced. This move towards a single bone for the mandible paved the way for other bones in the jaw, the articular and angular, to migrate to the cranium, where they function as parts of the mammalian hearing system.

Cynodonts also developed a secondary palate 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.

Characteristics[edit]

Bienotherium yunnanense

Early cynodonts have many of the skeletal characteristics of mammals. The teeth were fully differentiated and the braincase bulged at the back of the head. Outside of some crown-group mammals (notably the therians), all cynodonts probably laid eggs. The temporal fenestrae were much larger than those of their ancestors, and the widening of the zygomatic arch in a more mammal-like skull would have allowed for more robust jaw musculature. They also have the secondary palate 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 was the largest bone in their lower jaw.

The cynodonts probably had some form of warm-blooded metabolism. This has led to many reconstructions of cynodonts as having fur. Being endothermic they may have needed it for thermoregulation, but fossil evidence of their fur (or lack thereof) has been elusive. Modern mammals have Harderian glands secreting lipids to coat their fur, but the telltale imprint of this structure is only found from the primitive mammal Morganucodon and onwards.[8] Nonetheless, recent studies on Permian synapsid coprolites show that more basal therapsids may have had fur,[9] and at any rate fur was already present in Mammaliaformes such as Castorocauda and Megaconus.

Early cynodonts had numerous small foramina on their snout bones, similar to reptiles. This suggests that they had immobile, non-muscular lips like those of lizards, and lacked muscular cheeks.[10] As a muscular, mobile face is necessary to perform whisking movements and to avoid damage to whiskers, it is unlikely that early cynodonts had whiskers.[11][12] 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 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 young as in modern marsupials and monotremes. Only placentals, and perhaps Megazostrodon and Erythrotherium, would lose these.[13][14] A specimen of Kayentatherium 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.[15]

Cynodonts are the only known synapsid lineage to have produced aerial locomotors, with gliding and flying being known in haramiyidans[16] and various mammal groups.

Phylogeny[edit]

Below is a cladogram from Ruta, Botha-Brink, Mitchell and Benton (2013) showing one hypothesis of cynodont relationships:[6]

Cynodontia 

Charassognathus

Dvinia

Procynosuchus

 Epicynodontia 

Cynosaurus

Galesaurus

Progalesaurus

Nanictosaurus

Thrinaxodon

Platycraniellus

 Eucynodontia 

Cynognathia

Probainognathia

→ †Cynognathia 

Cynognathus

 †Gomphodontia 

Diademodon

 †Trirachodontidae 

Beishanodon

Sinognathus

Trirachodon

Cricodon

Langbergia

 †Traversodontidae 

Andescynodon

Pascualgnathus

Scalenodon

Luangwa

Traversodon

"Scalenodon" attridgei

Mandagomphodon

Nanogomphodon

Arctotraversodon

Boreogomphodon

 †Massetognathinae 

Massetognathus

Dadadon

Santacruzodon

 †Gomphodontosuchinae 

Menadon

Gomphodontosuchus

Protuberum

Exaeretodon

Scalenodontoides

→ Probainognathia 

Lumkuia

Ecteninion

Aleodon

Chiniquodon

Probainognathus

Trucidocynodon

Therioherpeton

 †Tritheledontidae 

Riograndia

Chaliminia

Elliotherium

Diarthrognathus

Pachygenelus

 †Brasilodontidae 

Brasilitherium

Brasilodon

 †Tritylodontidae 

Oligokyphus

Kayentatherium

Tritylodon

Beinotherium

 Mammaliaformes 

Mammalia

Sinoconodon

Morganucodon

Distribution[edit]

Cynodonts have been found in South America, India, Africa, Antarctica,[17] Asia,[18] Europe[19] and North America.[20]

See also[edit]

References[edit]

  1. ^ Classification of R. Owen 1861.
  2. ^ Classification of B. S. Rubidge and C. A. Sidor 2001
  3. ^ R. Broom. 1913. A revision of the reptiles of the Karroo. Annals of the South African Museum 7(6):361–366
  4. ^ S. H. Haughton and A. S. Brink. 1954. A bibliographical list of Reptilia from the Karroo Beds of South Africa. Palaeontologia Africana 2:1–187
  5. ^ Huttenlocker, Adam K.; Sidor, Christian A. (1 December 2020). "A Basal Nonmammaliaform Cynodont from the Permian of Zambia and the Origins of Mammalian Endocranial and Postcranial Anatomy". Journal of Vertebrate Paleontology. 40 (5): e1827413. doi:10.1080/02724634.2020.1827413. ISSN 0272-4634.
  6. ^ a b Ruta, M.; Botha-Brink, J.; Mitchell, S. A.; Benton, M. J. (2013). "The radiation of cynodonts and the ground plan of mammalian morphological diversity". Proceedings of the Royal Society B: Biological Sciences. 280 (1769): 20131865. doi:10.1098/rspb.2013.1865. PMC 3768321. PMID 23986112.
  7. ^ Abdala, Fernando; Gaetano, Leandro C. (2018), Tanner, Lawrence H. (ed.), "The Late Triassic Record of Cynodonts: Time of Innovations in the Mammalian Lineage", The Late Triassic World, Cham: Springer International Publishing, 46, pp. 407–445, doi:10.1007/978-3-319-68009-5_11, ISBN 978-3-319-68008-8, retrieved 24 May 2021
  8. ^ Ruben, J.A.; Jones, T.D. (2000). "Selective Factors Associated with the Origin of Fur and Feathers". American Zoologist. 40 (4): 585–596. doi:10.1093/icb/40.4.585.
  9. ^ Bajdek, Piotr (2015). "Microbiota and food residues including possible evidence of pre-mammalian hair in Upper Permian coprolites from Russia". Lethaia. 49 (4): 455–477. doi:10.1111/let.12156.
  10. ^ Estes, Richard (1961). "Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus". Bulletin Museum of Comparative Zoology. 125: 165–180.
  11. ^ Benoit, J.; Manger, P. R.; Rubidge, B. S. (9 May 2016). "Palaeoneurological clues to the evolution of defining mammalian soft tissue traits". Scientific Reports. 6 (1): 25604. doi:10.1038/srep25604. ISSN 2045-2322. PMC 4860582.
  12. ^ Benoit, Julien; Ruf, Irina; Miyamae, Juri A.; Fernandez, Vincent; Rodrigues, Pablo Gusmão; Rubidge, Bruce S. (2020). "The Evolution of the Maxillary Canal in Probainognathia (Cynodontia, Synapsida): Reassessment of the Homology of the Infraorbital Foramen in Mammalian Ancestors". Journal of Mammalian Evolution. 27 (3): 329–348. doi:10.1007/s10914-019-09467-8. eISSN 1573-7055. ISSN 1064-7554.
  13. ^ Michael L. Power, Jay Schulkin. The Evolution of the Human Placenta. pp. 68–.
  14. ^ Jason A. Lillegraven, Zofia Kielan-Jaworowska, William A. Clemens, Mesozoic Mammals: The First Two-Thirds of Mammalian History, University of California Press, 17 December 1979 – 321
  15. ^ Hoffman, Eva A.; Rowe, Timothy B. (2018). "Jurassic stem-mammal perinates and the origin of mammalian reproduction and growth". Nature. 561 (7721): 104–108. Bibcode:2018Natur.561..104H. doi:10.1038/s41586-018-0441-3. PMID 30158701. S2CID 205570021.
  16. ^ Meng, Qing-Jin; Grossnickle, David M.; Di, Liu; Zhang, Yu-Guang; Neander, April I.; Ji, Qiang; Luo, Zhe-Xi (2017). "New gliding mammaliaforms from the Jurassic". Nature. 548 (7667): 291–296. Bibcode:2017Natur.548..291M. doi:10.1038/nature23476. PMID 28792929. S2CID 205259206.
  17. ^ Abdala, Fernando; Ribeiro, Ana Maria (2010). "Distribution and diversity patterns of Triassic cynodonts (Therapsida, Cynodontia) in Gondwana". Palaeogeography, Palaeoclimatology, Palaeoecology. 286 (3–4): 202–217. Bibcode:2010PPP...286..202A. doi:10.1016/j.palaeo.2010.01.011.
  18. ^ Lucas, Spencer G. (2001). Chinese Fossil Vertebrates. New York City, Chichester, West Sussex: Columbia University Press. p. 133. ISBN 0-231-08482-X. Retrieved 30 November 2019.
  19. ^ Fraser, Nicholas C.; Sues, Hans-Dieter (1997). In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods. Cambridge University Press.
  20. ^ Sues, Hans-Dieter; Olsen, Paul E.; Carter, Joseph G. (1999). "A Late Triassic Traversodont Cynodont From the Newark Supergroup of North Carolina" (PDF). Journal of Vertebrate Paleontology. 19 (2): 351–354. doi:10.1080/02724634.1999.10011146.[permanent dead link]

Further reading[edit]

  • Hopson, J.A.; Kitching, J.W. (2001). "A probainognathian cynodont from South Africa and the phylogeny of non-mammalian cynodonts". Bull. Mus. Comp. Zool. 156: 5–35.
  • Davis, Dwight (1961). "Origin of the Mammalian Feeding Mechanism". Am. Zoologist, 1:229–234.

External links[edit]