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Author's note: these pages were written some years ago. I am not planning to update them. For a more current coverage, see the link to palaeos com Palaeos website (to which many links on these pages point to anyway. More info here


Family Trithelodontidae

The Trithelodontids, also and better known (especuially in earlier literature) as the Ictidosaurs, were tiny latest Triassic and early Jurassic cynodonts that neatly bridge the gap between advanced theriodonts and the primitive mammals.

Although previously separated from other theriodonts as a distinct infraorder, the Ictidosauria, study of the type and only specimen of the genus the Trithelodon has determined its close affinities with the typical Ictidosaurs Pachygenelus and "Diarthrognathus". For this reason, Hopson and Kitching suggested using the family name Trithelodontidae Broom 1912 for all of the forms called "ictidosaurs", and included all these groups under the Cynodontia.

The Trithelodonts (Ictidosaurs) are thus very advanced, probably insectivorous, cynodonts of small size in which some incisors and in some species the upper and probably postcanines have a transversely-oriented cutting edge, in others the uppers have an oblique and the lowers a longitudinal cutting edge. These are features connected, as in the tritylodonts, with the more efficient chewing of food. As in the Chiniquodontidae the secondary palate is long (so the animal could eat and breathe at the same time, a mammalian feature) and the postorbital bar (the bar of bone behind the eyes) is absent, another feature found also in Tritylodonts and primitive mammals.

But what makes these animals unique is the new mammalian joint between the squamosal and dentary had come into functional being. Thus, the ictidosaur Pachygenelius (formerly known as Diarthrognathus) possesses both the reptilian quadrate-articular jaw joint and a newly developed mammalian squamosal-dentary jaw joint. Because of this Simpson (1959) suggested that it might better be classified as a mammal, but Hopson and Crompton retain it in the Therapsida. Although certain primitive features in the skull suggest a derivation from scaloposaurid bauriamorphs, indicating the posisbility that the mammalian condition may have evolved twoice (once from advanced therocephalians via the Ictidosaurs and once from cyndonts via Tritylodonts) But both the structure of the dentary and the braincase structure have independently confirm a cynodont ancestry, and the polyphyly of mammals is no longer considered a valid hypothesis.

Pachygenelius stands truly at the dividing line between reptile and mammal in so far as this important diagnostic feature of jaw articulation is concerned. This is the only reason why the ictidosaurs are classified as reptiles rather than mammals. In the mammals, the quadrate and articular bones have migrated from the articular region of the jaw to the middle ear where they have been transformed into two of the bones concerned with the transmission of vibrations from the eardrum to the inner ear. Because, in the ictidosaurs, the tta formation of the quadrate and articular bones had not taken place, these animals can be placed arbitrarily within the reptiles.

Family Trithelodontidae Broom 1912b.

Includes: Ictidosauridae Young 1947; Diarthrognathidae Crompton 1958.


Horizon: Upper Triassic?, Lower Jurassic: Stormberg Series (Red Beds and Cave Sandstone) of South Africa and Lesotho; Los Colorados Formation of Argentina.
Age: link to palaeos com Rhaetian/link to palaeos com Hettangian to link to palaeos com Sinemurian / early link to palaeos com Pliensbachian
Distribution: probably had a worldwide (link to palaeos com Pangea) distribution
Ecological community: Anchisaur-Plateosaur empire Ecological niche/Guild: small terrestrial insectivore
Modern equivalents: shrew, insectvore generally
preferred food: mostly invertebrates, very small vertebrates
length: about 10 or 20 cm long
Metabolism: endothermic
Potential Predators: small theropod dinosaurs (mostly Coelophysidae), large link to palaeos com sphenosuchid "lizards", and link to palaeos com protosuchid crodylomorphs
Ancestor: Chiniquodontidae
Replaced: small Chiniquodontidae
Replaced by: small Mesozoic mammals
Descendents: Class Mammalia (ancestral forms: Sinoconodontidae, Morganucodontidae, Megazostrodontidae)
Taxonomic status - valid Family

List of genera

Genus Trithelodon Broom 1912b

Trithelodon riconoi Broom 1912b


Horizon: Red Beds (Upper Elliot Formation), Orange Free State, South Africa
age: link to palaeos com Hettangian / link to palaeos com Sinemurian

Genus Pachygenelius Watson 1913

Synonym: Diarthrognathus Crompton 1958

Pachygenelius monus Watson 1913


Horizon: Red Beds and Cave Sandstone (Middle and Upper Elliot Formation), Orange Free State, South Africa and Lesotho.
age: link to palaeos com Hettangian to link to palaeos com Sinemurian
Remarks: Pachygenelius ?monus has also been reported from the Early Jurassic of Nova Scotia. But while it is not unlikely that this jaw might belong to Pachygenelius (or a similiar form), a species attribution, or even an unambigious genus attribution, is very unlikely

Pachygenelius broomi Crompton 1958

Synonym: Diarthrognathus
horizon: Clarens Formation, Orange Free State, South Africa
age: link to palaeos com Sinemurian / early link to palaeos com Pliensbachian
material: 2 juvenile specimens.
Remarks: Diarthrognathus broomi is based on two juvenile specimens which are possibly, though not certainly, referable to P. monus. This then is either a synonym of the earlier P. monus or a distinct but closely related species, probably a descendent. Although a specific distinction may prove to be valid, it is doubtful that generic differences exist.

Chaliminia musteloides Bonaparte

Horizon: Los Colorados Formation, Argentina
Age: late link to palaeos com Norian/link to palaeos com Rhaetian/link to palaeos com Hettangian
Place: west link to palaeos com Gondwana/South-west link to palaeos com Pangea

Comments: The only known Triassic Trithelodont, and part of a highly endemic fauna, which makes it difficult to correlate. However although the Los Colorados Formation is usually considered latest Triassic (Late Norian /Rhaetian), Jose Bonparte says somewhere (I can't find the exact passage...) that the fauna may not be isochronous. So it is possible that the upper part of the Los Colorados may be Hettangian (earliest Jurassic). That would fit in Chaliminia with the other Trithelodonts.


Theioherpeton

Comments: no further info at the moment



some printed references some Links and References Web links

Palaeos Trithelodontidae - Palaeos

web page JURASSIC CYNODONTS; Tritylodontidae and Trithelodontidae, an internet directory - by Trevor Dykes - the most complete listing (along with the present page) of genera and species

printed reference Carroll, R. L. Vertebrate paleontology and evolution. -W. H. Freeman and company, New York, 1988

printed reference Edwin H. Colbert, Evolution of the Vertebrates, 2nd edition, 1969, John Wiley and Sons

printed reference James A. Hopson, "The Origin and Adaptive Radiation of Mammal-Like Reptiles and Non-Therian Mammals, Annals of the New York Academy of Sciences 167:199-216, 1969

printed reference James A. Hopson and Herbert R. Barghusen, "An Analysis of Therapsid Relationships", in The Ecology and Biology of Mammal-Like Reptiles ed. by Nocholas Hotton III, Paul D. MacLean, Jan J. Roth and E. Carol Roth, Smithsonian Institute Press, Washington and London, 1986, pp.83-106

printed reference James A. Hopson and James W. Kitching, 1972, "A Revised Classification of the Cynodonts (Reptilia; Therapsida), Paleontologica Africa, 14. 17-85

web page Trithelodont jaw - a fine photo of a partial jaw (Nova Scotia Museum). Earliest Jurassic (early Hettangian) age.

web page The Parrsboro Fossil Site - more information, including a reconstruction of the live animal (scroll down)

web page Trithelodont jaw fragment with tooth - also from Nova Scotia, identified as Pachygenelus ?monus

web page cladogramEucynodontia - cladogram and genus list - Mikko K. Haaramo

web page 'La importancia turistica de Ischigualasto', preparado por Dr William Sill. a fossil inventory in Spanish - at the bottom it lists Chaliminia musteloides (under "cynodont"



Palaeos Palaeos Page (incorporates some of this material, plus a lot of additional material)




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link to palaeos com Kingdoms of Life > Wikipedia link Animalia > link to palaeos com Vertebrata > link to palaeos com Tetrapoda > link to palaeos com Theropsida > Therapsida


Terrestrial Evolutionary Biotas | link to palaeos com Pangea | link to palaeos com Triassic Period | link to palaeos com Jurassic Period link to palaeos com Early Jurassic | link to palaeos com Tetrapoda | Permo-Triassic Tetrapods | Anchisaur-Coelophysid fauna | Therapsida | Theriodontia


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page by M.Alan Kazlev
I am indebted to Trevor Dykes for helpful corrections and references.
page uploaded 8 October 1999. Reposted and last modified 1 September 2005