Diadectes Temporal range: Artinskian-Kungurian | |
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Mounted skeleton of D. sideropelicus, American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Diadectomorpha |
Family: | † Diadectidae |
Genus: | † Diadectes Cope, 1878 |
Type species | |
†Diadectes sideropelicus Cope, 1878 | |
Species | |
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Synonyms | |
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Diadectes (meaning crosswise-biter) is an extinct genus of large reptiliomorphs or synapsids [1] that lived during the early Permian period (Artinskian-Kungurian stages of the Cisuralian epoch, between 290 and 272 million years ago). [2] Diadectes was one of the first herbivorous tetrapods, and also one of the first fully terrestrial vertebrates to attain large size.
Diadectes was a heavily built animal, up to 3 m (10 ft) long, [3] [4] with a thick-boned skull, heavy vertebrae and ribs, 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. [5]
It possesses some characteristics of reptilians and amphibians, combining a reptile-like skeleton with a more primitive, seymouriamorph-like skull. Diadectes has been classified as belonging to the sister group of the amniotes.
Among its primitive features, Diadectes has a large otic notch (a feature found in all labyrinthodonts, but not in reptiles) 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 to those of early amniotes such as pelycosaurs and pareiasaurs, as the first reptiles evolved from small, swamp-dwelling animals like Casineria and Westlothiana . [6] [7] The phenomenon of unrelated animals evolving similarly is known as convergent evolution. [8]
Diadectes was first named and described by American paleontologist Edward Drinker Cope in 1878, [9] 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).
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). [2]
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. sanmiguelensis and D. absitus to be more basal than other species of Diadectes, outside the derived clade composed of these species. In these analyses, Diasparactus zenos was more closely related to the other species of Diadectes than was D. sanmiguelensis and D. absitus, making Diadectes paraphyletic. Kissel recovered this paraphyly in his analysis and proposed the new genus name "Oradectes" for D. sanmiguelensis, and "Silvadectes" for D. absitus. [10] Below is the cladogram from Kissel's thesis:
Diadectomorpha |
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However, according to the ICZN, a name presented in an initially unpublished thesis such as Kissel's is not valid. Because the names "Oradectes" and "Silvadectes" have not yet been formally erected in a published paper, they were not, as of 2010, considered valid. A 2024 paper formally erected the genus Kuwavaatakdectes for D. sanmiguelensis. The same paper also named another Diadectes species, D. dreigleichenensis, which coexisted with D. absitus in the Tambach Formation of Germany. [11]
"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco, lepospondyls lived from the Visean stage of the Early Carboniferous to the Early Permian and were geographically restricted to what is now Europe and North America. Five major groups of lepospondyls are known: Adelospondyli; Aïstopoda; Lysorophia; Microsauria; and Nectridea. Lepospondyls have a diverse range of body forms and include species with newt-like, eel- or snake-like, and lizard-like forms. Various species were aquatic, semiaquatic, or terrestrial. None were large, and they are assumed to have lived in specialized ecological niches not taken by the more numerous temnospondyl amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by Karl Alfred von Zittel, who coined the name to include some tetrapods from the Paleozoic that shared some specific characteristics in the notochord and teeth. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota. It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive reptiles.
Reptiliomorpha 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 defined by Michel Laurin (2001) and Vallin and Laurin (2004) as the largest clade that includes Homo sapiens, but not Ascaphus truei. Laurin and Reisz (2020) defined Pan-Amniota as the largest total clade containing Homo sapiens, but not Pipa pipa, Caecilia tentaculata, and Siren lacertina.
Diadectidae is an extinct family of early tetrapods that lived in what is now North America and Europe during the Late Carboniferous and Early Permian, and in Asia during the Late Permian. They were the first herbivorous tetrapods, and also the first fully terrestrial animals to attain large sizes. Footprints indicate that diadectids walked with an erect posture. They were the first to exploit plant material in terrestrial food chains, making their appearance an important stage in both vertebrate evolution and the development of terrestrial ecosystems.
Diadectomorpha is a clade of large tetrapods that lived in Euramerica during the Carboniferous and Early Permian periods and in Asia during Late Permian (Wuchiapingian), They have typically been classified as advanced reptiliomorphs positioned close to, but outside of the clade Amniota, though some recent research has recovered them as the sister group to the traditional Synapsida within Amniota, based on inner ear anatomy and cladistic analyses. They include both large carnivorous and even larger herbivorous forms, some semi-aquatic and others fully terrestrial. The diadectomorphs seem to have originated during late Mississippian times, although they only became common after the Carboniferous rainforest collapse and flourished during the Late Pennsylvanian and Early Permian periods.
Captorhinidae is an extinct family of tetrapods, typically considered primitive reptiles, known from the late Carboniferous to the Late Permian. They had a cosmopolitan distribution across Pangea.
Seymouria is an extinct genus of seymouriamorph from the Early Permian of North America and Europe. Although they were amphibians, Seymouria were well-adapted to life on land, with many reptilian features—so many, in fact, that Seymouria was first thought to be a primitive reptile. It is primarily known from two species, Seymouria baylorensis and Seymouria sanjuanensis. The type species, S. baylorensis, is more robust and specialized, though its fossils have only been found in Texas. On the other hand, Seymouria sanjuanensis is more abundant and widespread. This smaller species is known from multiple well-preserved fossils, including a block of six skeletons found in the Cutler Formation of New Mexico, and a pair of fully grown skeletons from the Tambach Formation of Germany, which were fossilized lying next to each other.
Casineria is an extinct genus of tetrapodomorph 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, Scotland, where the holotype fossil was found. When originally described in 1999, it was identified as a transitional fossil noted for its mix of basal (amphibian-like) and advanced (reptile-like) characteristics, putting it at or very near the origin of the amniotes, the group containing all mammals, birds, modern reptiles, and other descendants of their reptile-like common ancestor. However, the sole known fossil is lacking key elements such as a skull, making exact analysis difficult. As a result, the classification of Casineria has been more controversial in analyses conducted since 1999. Other proposed affinities include a placement among the lepospondyls, seymouriamorphs, "gephyrostegids", or as a synonym of Caerorhachis, another controversial tetrapod which may have been an early temnospondyl.
Ichniotherium is an ichnogenus of tetrapod footprints from between the Late Carboniferous period to the Early Permian period attributed to diadectomorph track-makers. These footprints are commonly found in Europe, and have also been identified in North America and Morocco. Three ichnospecies of Ichniotherium have been proposed as valid: I. cotta, I. sphaerodactylum, and I. praesidentis.
Otsheria is an extinct genus of anomodont, in the infraorder venyukovioidea. It lived in modern-day Russia during the Permian.
Limnoscelis was a genus of large diadectomorph tetrapods from the Late Carboniferous to early Permian of western North America. It includes two species: the type species Limnoscelis paludis from New Mexico, and Limnoscelis dynatis from Colorado, both of which are thought to have lived concurrently. No specimens of Limnoscelis are known from outside of North America. Limnoscelis was carnivorous, and likely semiaquatic, though it may have spent a significant portion of its life on land. Limnoscelis had a combination of derived amphibian and primitive reptilian features, and its placement relative to Amniota has significant implications regarding the origins of the first amniotes.
Captorhinus is an extinct genus of captorhinid reptiles that lived during the Permian period. Its remains are known from North America and possibly South America.
Tseajaia is an extinct genus of diadectomorph tetrapod from the Early Permian of western North America. The skeleton is that of a medium-sized, rather advanced reptile-like animal. In life it was about 1 metre (3 ft) long and may have looked vaguely like an iguana. The dentition was somewhat blunt, indicating herbivory or possibly omnivory. It contains a single known species, Tseajaia campi.
Orobates is an extinct genus of diadectid reptiliomorphs that lived during the Early Permian. Its fossilised remains were found in Germany. A combination of primitive and derived traits distinguish it from all other well-known members of Diadectidae, a family of herbivorous reptiliomorphs. It weighed about 4 kg and appears to have been part of an upland fauna, browsing on high fibre plants.
Limnoscelidae is a family of carnivorous diadectomorphs. They would have been the largest terrestrial carnivores of their day, the other large carnivores being aquatic or semi aquatic labyrinthodont amphibians. The Limnoscelidae themselves, being close to the ancestry of amniotes, would have been well adapted land animals, but still dependent on anamniote eggs, and possibly having a tadpole stage. Contrary to the more advanced herbivorous diadectids, the teeth retained labyrinthodont infolding of the enamel, and were pointed and slightly recurved at the tip.
Ambedus is an extinct genus of possible diadectid reptiliomorph. Fossils have been found from the Early Permian Dunkard Group of Monroe County, Ohio. The type species A. pusillus was named in 2004. The genus name comes from the Latin word ambedo meaning "to nibble", in reference to its herbivorous diet. The specific name pusillus means "tiny" in Latin.
Kuwavaatakdectes is a genus of diadectid from the early Permian of Colorado.
Desmatodon is an extinct genus of diadectid reptiliomorph. With fossils found from the Kasimovian (Missourian) stage of the Late Carboniferous of Pennsylvania, Colorado, and New Mexico in the United States, Desmatodon is the oldest known diadectid. Two species are currently recognized: the type species D. hollandi and the species D. hesperis.
The Tambach Formation is an Early Permian-age geologic formation in central Germany. It consists of red to brown-colored sedimentary rocks such as conglomerate, sandstone, and mudstone, and is the oldest portion of the Upper Rotliegend within the Thuringian Forest Basin.
The Organ Rock Formation or Organ Rock Shale is a formation within the late Pennsylvanian to early Permian Cutler Group and is deposited across southeastern Utah, northwestern New Mexico, and northeastern Arizona. This formation notably outcrops around Canyonlands National Park, Natural Bridges National Monument, and Monument Valley of northeast Arizona, southern Utah. The age of the Organ Rock is constrained to the latter half of the Cisuralian epoch by age dates from overlying and underlying formations. Important early terrestrial vertebrate fossils have been recovered from this formation in northern Arizona, southern Utah, and northern New Mexico. These include the iconic Permian terrestrial fauna: Seymouria, Diadectes, Ophiacodon, and Dimetrodon. The fossil assemblage present suggests arid environmental conditions. This is corroborated with paleoclimate data indicative of global drying throughout the early Permian.