Cricodon

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Cricodon
Temporal range: 251–242  Ma
Partial Cricodon metabolus skeleton reconstruction.png
The partial skeletal reconstruction of Cricodon metabolus with a tool as reference for size.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: Cynodontia
Clade: Neogomphodontia
Genus: Cricodon
A. W. Crompton, 1955
Type species
C. metabolus

Cricodon is an extinct genus of trirachodontid cynodonts that lived during the Early Triassic and Middle Triassic periods of Africa. [1] [2] A. W. Crompton named Cricodon based on the ring-like arrangement of the cuspules on the crown of a typical postcanine tooth. [3] The epithet of the type species, C. metabolus, indicates the change in structure of certain postcanines resulting from replacement. [3]

Contents

Discovery

Cricodon was first discovered in the Tanzanian Manda Beds of South Africa. [3] [1] Broili & Schröder (1936) [4] were the first to describe Cricodon, yet were not able to provide a name for the taxon, which at the time was only known from 5 teeth. Extensive and in-depth descriptions of fossils from the Manda Beds were provided by A. W. Crompton (1955). [3] Crompton provided the name Cricodon as more fossil discoveries were found and a more complete view of the skeleton could be created. [3]

Fossil evidence of Cricodon has also been discovered in the Karoo Beds, specifically in the Cynognathus Assemblage Zone of South Africa. [5] The Cynognathus Assemblage Zone encompasses the boundary between the late Early and early Middle Triassic period, and has been subdivided into three distinct subzones (Subzone A, Subzone B, and Subzone C) [6] based primarily on the spatial and temporal ranges of key temnospondyl index taxa. [5] Cricodon fossils have been found in the youngest of the three subzones, Subzone C. [5] However, upper postcanines resembling those of Cricodon are also known from deposits corresponding to Subzone B. [5]

The trirachodontid "Trirachodon" kannemeyeri Seeley, 1895 is now referred to Cricodon as C. kannemeyeri. [7]

Description

Skull

A lower right dentary of Cricdon metabolus with tool provided for size reference Right Dentary of C. metabolus.jpg
A lower right dentary of Cricdon metabolus with tool provided for size reference

Cranial bones recovered from the Manda Beds consist of a badly crushed orbito-ethmoidal region, a practically complete right mandible, two fragments of the left mandible, several loose teeth, a portion of the occiput, and several unidentified fragments. [3]

In the upper jaw, only the posterior ends of the maxillae, a portion of the palate, and the floor of the orbits are well preserved. [3] The maxillary postcanines are transversely ovate and have three main cusps arranged upon the same transverse plane. [3] The three main cusps are composed of the lingual, central, and labial cusp. [3] There are also small peripheral cuspules present on the anterior and posterior borders of the crown. [3] Roots of the upper postcanines are long, transversely flattened near the crown and taper away to a short point distally. [3] The maximum known transverse diameter of the maxillary postcanines is 13mm. [3]

A characteristic feature of the mandible is the sharp angle formed by the junction of the lower margin of the dentary and the anterior surface of the dental symphysis. [3] A diagnostic difference between the upper jaw postcanine teeth and mandibular postcanine teeth is that mandibular postcanine teeth tend to be squarer in horizontal sections in contrast to the transversely ovate maxillary postcanines. [3] A mammalian feature that can be seen in gomphodont cynodonts is that each root is surrounded by a pocket of cancellous bone. [3] The maximum transverse diameter of the mandibular postcanines is 9mm. [3]

The main cusps of the mandibular and maxillary postcanines form a transverse ridge across the center of the crown. [3] The transverse ridge of a tooth would fit into the depression formed between the transverse ridges of two adjoining teeth. [3] Recent research conducted by Hendrickx, Abdala, and Choiniere (2016) [8] has revealed new information in regard to the distribution of enamel microstructure in non-mammaliform cynodonts, specifically in Cricodon metabolus. Their research uncovered the presence of columnar divergence units in both the sectorial and gomphodont teeth of a trirachodontid along with the consistent presence of synapsid columnar enamel in cynognathians. [8]

Reconstruction of Cricodon kannemeyeri. Cricodon.jpg
Reconstruction of Cricodon kannemeyeri.

The newfound discovery in regards to the thickened enamel has many ecological implications. [8] In Cricodon metabolus, the enamel layer of the gomphodont tooth is around 11.5 times thicker than the sectorial tooth. [8] The postcanine gomphodont teeth (labiolingually expanded teeth with large occlusal surfaces) were used for chewing, crushing, and grinding fibrous plant material, meaning that they were under higher loads and apically oriented stresses. [8] Sectorial teeth on the other hand were used to shear plant material [3] and were not subjected to the same types of occlusal stresses, therefore the enamel thickness was not maintained. [8] Another reason proposed by Hendrickx, Abdala, and Choiniere (2016) [8] to explain the thin enamel in sectorial teeth is due to replacement timing and patterns, as they will be sequentially replaced by gomphodont teeth. [3] Another characteristic that was observed in the gomphodont teeth were enamel and dentine incremental lines. [8] The enamel and dentine incremental lines, odontoblast tubules in dentine, and discontinuous columnar divergence units in enamel support the consistent presence of synapsid columnar enamel in Cynognathia. [8]

Postcranial skeleton

An array of various postcranial bones of Cricodon metabolus Postcranial Bones of C. metabolus.jpg
An array of various postcranial bones of Cricodon metabolus

Twenty-five vertebrae were discovered which belong to the dorsal and sacral regions with only one vertebrae having a well preserved neural arch from the sacral region. [3] There are additional apophyses below the posterior zygapophysis which articulate with concavities on the lateral surface of the neural arch, posterior and inferior to the anterior zygapophysis. [3] Cricodon metabolus has the typical cynodont expanded ribs of which thirteen dorsal ribs were discovered. [3]

The entire humerus was 12.5 cm long and had typical cynodont characterizations such as the twisted bone and the plane of the distal end forming an angle of 40 degrees with that of the proximal. [3] The discovered femur was 12.5 cm long with the capitulum directed at an angle of 45 to 50 degrees to the main axis of the slender shaft. [3] Another characteristic of the femur noted is that it projects well forward from the main body of the shaft. [3] Although little is known of the hand of cynodonts, it is believed that Cricodon metabolus has a phalangeal formula of 23443, with the second phalange reduced in digits three and four. [3]

Classification

Below is a cladogram from Gao et al. (2010) [9] showing the phylogenetic relationships of one part of the Cynodontia relative to Cricodon:

Cynognathia  

Cynognathus

  Gomphodontia  

Diademodon

  Trirachodontidae  
 Trirachodontinae 

Trirachodon

Langbergia

Cricodon

 Sinognathinae 

Beishanodon

Sinognathus

Traversodontidae

See also

Related Research Articles

<span class="mw-page-title-main">Cynognathia</span> Clade of cynodonts

Cynognathia is one of two major clades of cynodonts, the other being Probainognathia. Cynognathians included the large carnivorous genus Cynognathus and the herbivorous or omnivorous gomphodonts such as traversodontids. Cynognathians can be identified by several synapomorphies including a very deep zygomatic arch that extends above the middle of the orbit.

<i>Massetognathus</i> Extinct genus of cynodonts

Massetognathus is an extinct genus of plant-eating traversodontid cynodonts. They lived during the Triassic Period about 235 million years ago, and are known from the Chañares Formation in Argentina and the Santa Maria Formation in Brazil.

<span class="mw-page-title-main">Traversodontidae</span> Extinct family of cynodonts

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 North America. Traversodonts first appeared in the Middle Triassic and diversified in the Late Triassic before going extinct at the end of the epoch. The family Traversodontidae was erected by Friedrich von Huene in 1936 for cynodonts first found in São Pedro do Sul in Paleorrota, Brazil.

<i>Cynognathus</i> Assemblage Zone Biozone which correlates to the Burgersdorp Formation of the Beaufort Group

The Cynognathus Assemblage Zone is a tetrapod biozone utilized in the Karoo Basin of South Africa. It is equivalent to the Burgersdorp Formation, the youngest lithostratigraphic formation in the Beaufort Group, which is part of the fossiliferous and geologically important Karoo Supergroup. The Cynognathus Assemblage Zone is the youngest of the eight biozones found in the Beaufort Group, and is considered to be late Early Triassic (Olenekian) to early Middle Triassic (Anisian) in age. The name of the biozone refers to Cynognathus crateronotus, a large and carnivorous cynodont therapsid which occurs throughout the entire biozone.

<i>Trirachodon</i> Extinct genus of cynodonts

Trirachodon is an extinct genus of cynodonts. Fossils have been found in the Cynognathus Assemblage Zone of the Beaufort Group in South Africa and the Omingonde Formation of Namibia, dating back to the Early and Middle Triassic.

<i>Lumkuia</i> Extinct genus of cynodonts

Lumkuia is an extinct genus of cynodonts, fossils of which have been found in the Cynognathus Assemblage Zone of the Beaufort Group in the South African Karoo Basin that date back to the early Middle Triassic. It contains a single species, Lumkuia fuzzi, which was named in 2001 on the basis of the holotype specimen BP/1/2669, which can now be found at the Bernard Price Institute in Johannesburg, South Africa. The genus has been placed in its own family, Lumkuiidae. Lumkuia is not as common as other cynodonts from the same locality such as Diademodon and Trirachodon.

<i>Langbergia</i> Extinct genus of cynodonts

Langbergia is an extinct genus of trirachodontid cynodont from the Early Triassic of South Africa. The type and only species L. modisei was named in 2006 after the farm where the holotype was found, Langberg 566. Langbergia was found in the Burgersdorp Formation in the Beaufort Group, a part of the Cynognathus Assemblage Zone. The closely related trirachodontids Trirachodon and Cricodon were found in the same area.

Neotrirachodon is an extinct genus of therapsids which existed in Russia during the Middle Triassic period. Its type and only species is Neotrirachodon expectatus.

<span class="mw-page-title-main">Trirachodontidae</span> Extinct family of cynodonts

Trirachodontidae is an extinct, possibly paraphyletic family of cynognathian cynodonts from the Triassic of China and southern Africa. Trirachodontids appeared during the Early Triassic soon after the Permian-Triassic extinction event and quickly spread over a wide geographic area in a comparatively brief amount of time from 250 to 237 million years ago.

<span class="mw-page-title-main">Gomphodontia</span> Clade of cynodonts

Gomphodontia is a clade of cynognathian cynodonts that includes the families Diademodontidae, Trirachodontidae, and Traversodontidae. Gomphodonts are distinguished by wide and closely spaced molar-like postcanine teeth, which are convergent with those of mammals. Other distinguishing characteristics of gomphodonts include deep zygomatic arches, upper postcanines with three or more cusps spanning their widths and lower postcanines with two cusps spanning their widths. They are thought to have been herbivorous or omnivorous. Gomphodonts first appeared in the Early Triassic and became extinct at the end of the Late Triassic. Fossils are known from southern Africa, Argentina and southern Brazil, eastern North America, Europe, China, and Antarctica.

<span class="mw-page-title-main">Diademodontidae</span> Family of gomphodontian cynodonts

Diademodontidae is an extinct family of Triassic gomphodonts. The best-known genus is Diademodon from South Africa. Titanogomphodon from Namibia may also be a member of Diademodontidae. The Chinese genera Hazhenia and Ordosiodon have also been included in the family, but were more recently identified as baurioid therocephalians. Remains of a diademodontid were reported in the Early-Middle Triassic Fremouw Formation in Antarctica, but that specimen was later referred to the trirachodontid Impidens

Titanogomphodon is an extinct genus of diademodontid cynodonts from the Middle Triassic Omingonde Formation of Namibia. It is known from a single partial skull that was described in 1973 from the Omingonde Formation. The type and only species is Titanogomphodon crassus. At about 40 centimetres (16 in), the skull of Titanogomphodon was significantly larger than that of its closest relative, Diademodon. Its teeth are similar to those of another group of cynodonts called Traversodontidae, but the similarities are likely the result of convergent evolution. Aside from its larger size, Titanogomphodon differs from Diademodon in having a bony projection on the postorbital bar behind the eye socket.

<i>Aleodon</i> Extinct genus of cynodonts

Aleodon is an extinct genus of cynodonts that lived from the Middle to the Late Triassic. Relatively few analyses have been conducted to identify the phylogenetic placement of Aleodon, however those that have place Aleodon as a sister taxon to Chiniquodon. Two species of Aleodon are recognized: A. brachyramphus which was discovered in Tanzania, and A. cromptoni which was discovered most recently in Brazil.

<i>Abdalodon</i> Extinct genus of cynodonts

Abdalodon is an extinct genus of late Permian cynodonts, known by its only species A. diastematicus.Abdalodon together with the genus Charassognathus, form the clade Charassognathidae. This clade represents the earliest known cynodonts, and is the first known radiation of Permian cynodonts.

<i>Ufudocyclops</i> Extinct genus of dicynodonts

Ufudocyclops is an extinct genus of stahleckeriid dicynodont from the Middle Triassic of South Africa. It was found in the Burgersdorp Formation, part of the uppermost Cynognathus Assemblage Zone of the Beaufort Group in the Karoo Basin. The type and only known species is U. mukanelai. It was a large, beaked herbivore like other Triassic dicynodonts, lacking tusks, and is mostly characterised by unique features of the skull. It is known from three specimens, two of which were previously referred to the Tanzanian dicynodont Angonisaurus. The separation of Ufudocyclops from Angonisaurus indicates that the Middle Triassic fauna of the Beaufort Group in South Africa was not part of a larger shared fauna with those of the Manda Beds in Tanzania, as was previously supposed, and suggests that they were separated as more localised faunas, possibly by geographic barriers or in time. Ufudocyclops then would have been a unique part of the uppermost Cynognathus Assemblage Zone in South Africa. It is also the oldest known member of the family Stahleckeriidae, and implies that the family was already diversifying in the Middle Triassic alongside other kannemeyeriiforms, not just in the Late Triassic after other families died out.

<i>Etjoia</i> Extinct genus of cynodonts

Etjoia is an extinct genus of traversodontid cynodonts that lived during the Middle Triassic or Late Triassic period in southern Africa. This medium-sized omnivorous cynognathian provides important information on the dental evolution of early diverging gomphodonts and traversodontids.

Vetusodon is an extinct genus of cynodonts belonging to the clade Epicynodontia. It contains one species, Vetusodon elikhulu, which is known from four specimens found in the Late Permian Daptocephalus Assemblage Zone of South Africa. With a skull length of about 18 centimetres (7.1 in), Vetusodon is the largest known cynodont from the Permian. Through convergent evolution, it possessed several unusual features reminiscent of the contemporary therocephalian Moschorhinus, including broad, robust jaws, large incisors and canines, and small, single-cusped postcanine teeth.

Arctotraversodontinae is a subfamily of Late Triassic cynodonts belonging to the family Traversodontidae. Members of the subfamily include Arctotraversodon, Boreogomphodon and Plinthogomphodon from North America, and Habayia, Maubeugia, Microscalenodon and Rosieria from Europe.

Scalenodontoides is an extinct genus of Traversodontidae, a family of herbivorous cynodonts. It lived during the Late Triassic in what is now South Africa. Its type species is Scalenodontoides macrodontes. It was named in 1957 by A. W. Crompton and F. Ellenberger. Arctotraversodon plemmyrodon was originally classified as a species of Scalenodontoides, but was given its own genus in 1992. It is found in the Scalenodontoides Assemblage Zone of the Elliot Formation, which is named for it. It is one of the geologically youngest traversodontids, alongside the putative traversodontid Boreogomphodon. It is closely related to Exaeretodon and Siriusgnathus, but is distinguished by the presence of a shelf-like expansion of its parietal called the nuchal table. Though the largest known complete skull is only 248 millimetres (9.8 in) long, it may have been the largest non-mammalian cynodont, as an incomplete snout would have belonged to a specimen with an estimated skull length of 617 millimetres (24.3 in).

<i>Impidens</i> Extinct genus of cynodonts

Impidens is an extinct genus of large omnivorous cynodont from the Triassic of South Africa and Antarctica. Its type and only species is Impidens hancoxi. Impidens inhabited high-latitude environments of southern Gondwana during the Middle Triassic, where it was probably the apex predator.

References

  1. 1 2 Abdala, F., J. Neveling, and J. Welman. 2006. A new trirachodontid cynodont from the lower levels of the Burgersdorp Formation (Lower Triassic) of the Beaufort Group, South Africa and the cladistic relationships of Gondwanan gomphodonts. Zoological Journal of the Linnean Society 147:383–413.
  2. Hopson, J. A. 2005. A juvenile gomphodont cynodont specimen from the Cynognathus Assemblage Zone of South Africa: implications for the origin of gomphodont postcanine morphology. Palaeontologia Africana 41:53–66.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Crompton, A. W. 1955. On some Triassic cynodonts from Tanganyika. Proceedings of the Zoological Society of London 125(3–4):617–669.
  4. Broili, F. (1935). Beobachtungen an Wirbeltieren der Karrooformation; VIII, Ein Dinocephalen-Rest aus den unteren Beaufort-Schichten; IX, Ueber den Schaedel von Gomphognathus Seeley. Sitzungsberichte - Bayerische Akademie Der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 93-114.
  5. 1 2 3 4 Abdala, F., P. J. Hancox, and J. Neveling. 2005. Cynodonts from the uppermost Burgersdorp Formation, South Africa, and their bearing on the biostratigraphy and correlation of the Triassic Cynognathus Assemblage Zone. Journal of Vertebrate Paleontology 25:192–199.
  6. Hancox, P., Shishkin, M., Rubidge, B., & Kitching, J. (1995). A threefold subdivision of the Cynognathus assemblage zone (Beaufort Group, South Africa) and its palaeogeographical implications. South African Journal of Science,91(3), 143-144.
  7. Sidor, C. A., and J. A. Hopson. 2018. Cricodon metabolus (Cynodontia: Gomphodontia) from the Triassic Ntawere Formation of northeastern Zambia: patterns of tooth replacement and a systematic review of the Trirachodontidae; pp. 39–64 in C. A. Sidor and S. J. Nesbitt (eds.), Vertebrate and Climatic Evolution in the Triassic Rift Basins of Tanzania and Zambia. Society of Vertebrate Paleontology Memoir 17. Journal of Vertebrate Paleontology 37(6,Supplement).
  8. 1 2 3 4 5 6 7 8 9 Hendrickx, C., Abdala, F., & Choiniere, J. (2016). Postcanine microstructure in Cricodon metabolus, a Middle Triassic gomphodont cynodont from south-eastern Africa. Palaeontology, 59(6), 851-861.
  9. Gao, K., Fox, R., Zhou, C., & Li, D. (2010). A New Nonmammalian Eucynodont (Synapsida: Therapsida) from the Triassic of Northern Gansu Province, China, and its Biostratigraphic and Biogeographic Implications. American Museum Novitates, 1-25.