Tritylodontidae

Tritylodonts Temporal range: Late Triassic - Early Cretaceous
Life restoration of Oligokyphus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Synapsida
Order:	Therapsida
Suborder:	Cynodontia
Clade:	Mammaliamorpha
Family:	†Tritylodontidae Cope, 1884
Genera
See below

Tritylodontids ("three knob teeth", named after the shape of animal's teeth) were small to medium-sized, highly specialized and extremely mammal-like cynodonts. They were the last known family of the non-mammalian synapsids. One of the last cynodont lines to appear, the Tritylodontidae descended from a Cynognathus-like cynodont. The Tritylodontids were herbivorous, chewing through vegetation, such as stems, leaves, and roots.

Description

Skull of Kayentatherium wellesi

The tritylodont's skull had a high flat crest. They retained the reptilian joint between the quadrate bone of the skull and the auricular bone of the lower jaw, but they were reduced. It is only through the retention of the vestigial reptilian jawbones that they are technically regarded to not be mammals. The back of the skull had huge zygomatic arches for the attachment of its large jaw muscles. They also had a very well-developed secondary palate. The tritylodont dentition was very different from that of other cynodonts. They did not have canines. The front pair of incisors were enlarged that were very similar to rodents of today. Traversodonts had a large gap, the diastema, that separated the incisors from the square-shaped cheek. Each of the cheek teeth in the upper jaw had three rows of cusps running along its length that had grooves in between. The lower teeth had two rows of cusps which fitted into the grooves in the upper teeth. The matching of the cusps allowed the teeth to meet in a precise bite. It would grind its food between the teeth in somewhat the same way that a modern rodent would with their food. The teeth were well suited for shredding plants matter. Like Mammaliformes, tritylodontids have epipubic bones, a possible synampomorphy between both clades, and as such were forced to produce undeveloped fetus-like young like modern monotremes and marsupials.

The Tritylodonts can very much be seen as Mesozoic rodents. Tritylodonts were active animals that were probably warm blooded and burrowed like modern day rodents. For example, Oligokyphus could be compared to a weasel or mink, with a long, slim body and tail.

Discovery

Tritylodonts were first discovered in the Upper Triassic rocks of South Africa in the late 1800s and were initially thought to be amongst the very earliest mammals. The first genus was named in 1854, Stereognathus ooliticus, and the family name erected by Cope in 1884.

Evolutionary history

Tritylodontids appeared in the late Triassic period, and persisted through the Jurassic until the early Cretaceous. Yuanotherium was one of the last known species of tritylodont. Since then a genus from Cretaceous called Xenocretosuchus has been found, which appears to be a member of the clade as well. Some authors argue Chronoperates is a tritylodontid member that persisted into the Paleocene after the Cretaceous–Paleogene extinction event, but this is not widely accepted. If true, then tritylodontids were elusive and rare during the Late Cretaceous, because no Tritylodonts were found by that time. However, the Chronoperates 's anatomy closely resembles to that of symmetrodonts - a mammalian lineage. Tritylodontids were likely to have been warm-blooded. Tritylodontid fossils are found in the Americas, South Africa, and Eurasia - they appear to have had an almost global distribution, including Antarctica.

Phylogeny

Because of their extremely mammal-like appearance, tritylodontids were originally placed within Mammalia. Starting with the work of British palaeontologist D. M. S. Watson in 1942, a close relationship was favored between tritylodontids and cynodonts. Watson and other palaeontologists noted that tritylodontids lacked the dentary and squamosal jaw articulation that was characteristic of early mammals. Haughton and Brink (1954) were the first to classify tritylodontids within Cynodontia. Later studies identified close similarities between the teeth of tritylodontids and traversodontids, and tritylodontids were eventually thought to be descendants of traversodontids. Under this classification, which was widely accepted in the following decades, Tritylodontidae was previous considered to be part of Gomphodontia, a larger group within Cynognathia. The name Tritylodontoidea was previously used for the group, which traditionally included the families Diademodontidae, Trirachodontidae, Traversodontidae, and Tritylodontidae.

More recently, tritylodontids have been reinterpreted as close relatives of mammals. Beginning with Kemp (1983), Tritylodontidae has been proposed by numerous studies as a member of Probainognathia, the closest sister group to Mammalia. Gomphodontia is still used for the cynognathian group containing traversodontids and is preferred over Tritylodontoidea now that tritylodontoids are not part of it. A phylogenetic analysis performed by Liu and Olsen (2010) places Tritylodontidae very closely to Mammalia, as the sister taxon of the clade formed by Brasilodontidae and Mammalia. Ruta et al. (2013) phylogenetic analysis which is partially based on Liu and Olsen (2010) places Tritylodontidae in a more derived position than Brasilodontidae. Below is a cladogram from this analysis.

It should be noted that the exact position of Tritylodontidae in relation to Mammalia is still debated, but most researchers agree they are closely related, although Tritylodontidae is considered a non-mammaliamorph cynodont.

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  Sinoconodon Morganucodon

Genera

Kayentatherium

• Bienotherium
• Bienotheroides
• Bocatherium
• Dianzhongia
• Dinnebitodon
• Kayentatherium
• Lufengia
• Oligokyphus
• Polistodon
• Stereognathus
• Tritylodon
• Xenocretosuchus
• Yuanotherium
• Yunnanodon
• Montirictus

See also

• Mammalian embryonic diapause