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''Diadectes'' (meaning ''crosswise-biter'') is an extinct genus of large reptiliomorphs or synapsids that lived during the early Permian period ( Artinskian- Kungurian stages of the
Cisuralian The Cisuralian is the first Series (stratigraphy), series/Epoch (geology), epoch of the Permian. The Cisuralian was preceded by the Pennsylvanian (geology), Pennsylvanian and followed by the Guadalupian. The Cisuralian Epoch is named after the w ...
epoch, between 290 and 272 million years ago). ''Diadectes'' was one of the first herbivorous tetrapods, and also one of the first fully terrestrial vertebrates to attain large size.


Description

''Diadectes'' was a heavily built animal, up to long, with a thick-boned skull, heavy vertebrae and
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s, massive limb girdles, and short, robust limbs. The nature of the limbs and vertebrae clearly indicates a terrestrial animal. The rib cage was assumed to be barrel-shaped, but new fossils show the ribs were actually sticking out to the sides.


Paleobiology

It possesses some characteristics of reptilians and amphibians, combining a reptile-like skeleton with a more primitive,
seymouriamorph Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are s ...
-like skull. ''Diadectes'' has been classified as belonging to the sister group of the
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s. Among its primitive features, ''Diadectes'' has a large
otic notch Otic notches are invaginations in the posterior margin of the skull roof, one behind each orbit. Otic notches are one of the features lost in the evolution of amniotes from their tetrapod ancestors. The notches have been interpreted as part of an ...
(a feature found in all
labyrinthodonts "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consi ...
, but not in
reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
) with an ossified tympanum. At the same time, its teeth show advanced specialisations for an herbivorous diet that are not found in any other type of early Permian animal. The eight front teeth are spatulate and peg-like, and served as incisors that were used to nip off mouthfuls of vegetation. The broad, blunt cheek teeth show extensive wear associated with occlusion, and would have functioned as molars, grinding up the food. It also had a partial secondary palate, which meant it could chew its food and breathe at the same time, something many even more advanced reptiles were unable to do. These traits are likely adaptations related to the animals' high-fiber, herbivorous diet, and evolved independently of similar traits seen in some reptilian groups. Many of the reptile-like details of the postcranial skeleton are possibly related to carrying the substantial trunk; these may be independently derived traits on ''Diadectes'' and their relatives. Though very similar, they would be analogous rather than
homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...
to those of early amniotes such as pelycosaurs and
pareiasaur Pareiasaurs (meaning "cheek lizards") are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with scutes which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, ...
s, as the first reptiles evolved from small, swamp-dwelling animals like '' Casineria'' and '' Westlothiana''. Carroll R.L. (1991): The origin of reptiles. In: Schultze H.-P., Trueb L., (ed) ''Origins of the higher groups of tetrapods — controversy and consensus''. Ithaca: Cornell University Press, pp 331-353. Laurin, M. (2004): The Evolution of Body Size, Cope's Rule and the Origin of Amniotes. ''Systematic Biology'' no 53 (4): pp 594-622.
article
/ref> The phenomenon of unrelated animals evolving similarly is known as convergent evolution.Mayr, Ernst, and Peter D. Ashlock (1991): ''Principles of systematic zoology''. New York: McGraw-Hill


Discovery

''Diadectes'' was first named and described by American paleontologist
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
in 1878, based on part of a lower jaw (AMNH 4360) from the Permian of Texas. Cope noted: "Teeth with short and much compressed crowns, whose long axis is transverse to that of the jaws," the feature expressed in the generic name ''Diadectes'' "crosswise biter" (from Greek ''dia'' "crosswise" + Greek ''dēktēs'' "biter"). He described the animal as "in all probability, herbivorous." Cope's neo-Latin type species name ''sideropelicus'' (from Greek ''sidēros'' "iron" + Greek ''pēlos'' "clay" + -''ikos'') "of iron clay" alluded to the Wichita beds in Texas, where the fossil was found. ''Diadectes'' fossil remains are known from a number of locations across North America, especially the Texas Red Beds ( Wichita and Clear Fork).


Classification and species

Numerous species have been assigned to ''Diadectes'', though most of those have proven to be synonyms of one another. Similarly, many supposed separate genera of diadectids have been shown to be junior synonyms of ''Diadectes''. One of these, ''Nothodon'', was actually published by Othniel Charles Marsh five days before the name ''Diadectes'' was published by his rival Cope. Despite this fact, in 1912, Case synonymized the two names and treated ''Diadectes'' as the senior synonym, which has been followed by other paleontologists since, despite the fact that it violates the rules of International Code of Zoological Nomenclature (ICZN).Kissel, R. (2010). "Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha)." Thesis (Graduate Department of Ecology & Evolutionary Biology University of Toronto).


Phylogeny

A phylogenetic analysis of ''Diadectes'' and related diadectids was presented in an unpublished PhD thesis by Richard Kissel in 2010. Previous phylogenetic analyses of diadectids had found ''D. absitus'' to be more basal than other species of ''Diadectes'', outside the derived
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, ...
composed of these species. In these analyses, '' Diasparactus zenos'' was more closely related to the other species of ''Diadectes'' than was ''D. absitus'', making ''Diadectes''
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. Kissel recovered this paraphyly in his analysis and proposed the new genus name "Silvadectes" for ''D. absitus''. Below is the cladogram from Kissel's thesis: However, according to the ICZN, a name presented in an initially unpublished thesis such as Kissel's is not valid. Because the name "Silvadectes" has not yet been formally erected in a published paper, it was not, as of 2010, considered valid.


References

* Parker, Steve. Dinosaurus: the complete guide to dinosaurs. Firefly Books Inc, 2003. Pg. 83 * Benton, M. J. (2000), ''Vertebrate Paleontology'', 2nd ed. Blackwell Science Ltd * Carroll, R. L. (1988), ''Vertebrate Paleontology and Evolution'', WH Freeman & Co. * Colbert, E. H., (1969), ''Evolution of the Vertebrates'', John Wiley & Sons Inc (2nd ed.) * Reisz, Robert, (no date),
Biology 356 - Major Features of Vertebrate Evolution - Anthracosaurs and Diadectomorphs
{{Taxonbar, from=Q132762 Permian tetrapods Diadectids Permian tetrapods of North America