Diadectes Tenuitecus
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''Diadectes'' (meaning ''crosswise-biter'') is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of large
reptiliomorph Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians ( lissamphibians). It was de ...
s or
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s that lived during the early
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
period (
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of the ...
-
Kungurian In the geologic timescale, the Kungurian is an age or stage of the Permian. It is the latest or upper of four subdivisions of the Cisuralian Epoch or Series. The Kungurian lasted between and million years ago (Ma). It was preceded by the Artin ...
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 A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s, 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
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e and ribs, massive limb
girdle A belt, especially if a cord or rope, is called a girdle if it is worn as part of Christian liturgical vestments, or in certain historical, literary or sports contexts. Girdles are used to close a cassock in Christian denominations, including th ...
s, 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 a ...
(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
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s that were used to nip off mouthfuls of vegetation. The broad, blunt cheek teeth show extensive wear associated with
occlusion Occlusion may refer to: Health and fitness * Occlusion (dentistry), the manner in which the upper and lower teeth come together when the mouth is closed * Occlusion miliaria, a skin condition * Occlusive dressing, an air- and water-tight trauma ...
, and would have functioned as molars, grinding up the food. It also had a partial
secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...
, 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 Analogy (from Greek ''analogia'', "proportion", from ''ana-'' "upon, according to" lso "against", "anew"+ ''logos'' "ratio" lso "word, speech, reckoning" is a cognitive process of transferring information or meaning from a particular subject ...
rather than homologous to those of early amniotes such as
pelycosaurs Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is now ...
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 ''Casineria'' is an extinct genus of tetrapod which lived about 340-334 million years ago in the Mississippian epoch of the Carboniferous period. Its generic name, ''Casineria'', is a latinization of Cheese Bay, the site near Edinburgh, Scotla ...
'' and ''
Westlothiana ''Westlothiana'' ("animal from West Lothian") is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Visean age of the Carboniferous. Members of the genus bore a superficial resemblance to modern ...
''. 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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.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 Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among h ...
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 The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
(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