Diadectomorpha

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Diadectomorphs
Temporal range: Late Carboniferous–Wuchiapingian
Diadectes phaseolinus.JPG
Skeleton of Diadectes sideropelicus in the American Museum of Natural History
Limnoscelis paludis 2.JPG
Skeleton of Limnoscelis paludis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Reptiliomorpha
Order: Diadectomorpha
Watson, 1917
Subgroups

See text.

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), [1] They have typically been classified as advanced reptiliomorphs (transitional between "amphibians" sensu lato and amniotes) 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. [2] [3] [4] [5] They include both large (up to 2 meters long) carnivorous and even larger (to 3 meters) 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.

Contents

Anatomy

Life restoration of Limnoscelis Limnoscelis21DB.jpg
Life restoration of Limnoscelis

Diadectomorphs possessed both amphibian-like and amniote-like characteristics. Originally these animals were included under the order Cotylosauria, and were considered the most primitive and ancestral lineage of reptiles. More recently they have been reclassified either as non-amniote reptiliomorphs lying just outside the clade Amniota, or as early-diverging synapsids (members of the amniote group containing mammals and their extinct relatives). [2] [3] [5] Contrary to other non-amniote reptiliomorphs, the teeth of the Diadectomorpha lacked the infolding of the dentine and enamel that account for the name Labyrinthodontia for the non-amniote tetrapods. [6]

Classification

Diadectomorpha is most commonly given the rank of order when formal taxonomic ranks are applied. It is further divided into three families, representing specialization into different ecological niches. The exact phylogenetic relationship between the three is disputed. [7]

Below is a cladogram modified from the 2010 analysis: [1]

Diadectomorpha 

Limnoscelidae Limnoscelis21DB.jpg

Tseajaiidae Tseajaia BW.jpg

 Diadectidae

Ambedus pusillus ?

Oradectes sanmiguelensis

Orobates pabsti Orobates BW.jpg

Alveusdectes fenestralis

Desmatodon hesperis

Silvadectes absitus

Diadectes tenuitectes

Diadectes sideropelicus Diadectes1DB.jpg

Diasparactus zenos Diasparactus1DB.jpg

305 mya

Reproduction and the origin of Amniota

Life restoration of Diadectes Diadectes1DB.jpg
Life restoration of Diadectes

The reproduction of the Diadectomorphs has been the matter of some debate. [10] If their group lay within the Amniota as has at times been assumed, they would have laid an early version of the amniote egg. Current thinking favours the amniote egg being evolved in very small animals, like Westlothiana or Casineria , leaving the bulky Diadectomorphs just on the amphibian side of the divide. [11] [12] [13] [14]

This would indicate the large and bulky diadectomorphs laid anamniote eggs (in water). However, no unambiguously diadectomorph tadpole is known. Whether this is due to an actual lack of tadpole stage or taphonomy (many diadectomorphs were upland creatures where tadpoles would have a poor probability of being fossilized) is uncertain. Alfred Romer indicated that the anamniote/amniote divide might not have been very sharp, leaving the question of the actual mode of reproduction of these large animals unanswered. [15] Possible reproductive modes include full amphibian spawning with aquatic tadpoles, internal fertilization with or without ovoviviparity, aquatic eggs with direct development or some combination of these. The reproductive mode might also have varied within the group.

Lee and Spencer (1997) argued diadectomorphs probably laid amniote eggs because their adaptations to feed on terrestrial plants rich on fiber mean they were adapted to a niche not seen in unambiguous 'amphibians', and would have required an early acquisition of terrestrial endosymbionts necessary for this diet that supposedly could not have happened if young diadectomorphs were aquatic larvae. [16]

David Berman (2013) ran a phylogenetic analysis, and in this study the analysis resulted in Diadectomorpha being inside amniota as a sister taxon to Synapsida. [17]

Related Research Articles

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibian ancestors during the Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids. They are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration.

<span class="mw-page-title-main">Labyrinthodontia</span> Paraphyletic group of tetrapodomorphs

"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.

<span class="mw-page-title-main">Reptiliomorpha</span> Clade of reptile-like animals

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.

<span class="mw-page-title-main">Anthracosauria</span> Paraphyletic group of tetrapodomorphs

Anthracosauria is an order of extinct reptile-like amphibians that flourished during the Carboniferous and early Permian periods, although precisely which species are included depends on one's definition of the taxon. "Anthracosauria" is sometimes used to refer to all tetrapods more closely related to amniotes such as reptiles, mammals, and birds, than to lissamphibians such as frogs and salamanders. An equivalent term to this definition would be Reptiliomorpha. Anthracosauria has also been used to refer to a smaller group of large, crocodilian-like aquatic tetrapods also known as embolomeres.

<i>Diadectes</i> Extinct genus of reptiles

Diadectes is an extinct genus of large reptiliomorphs or synapsids that lived during the early Permian period. Diadectes was one of the first herbivorous tetrapods, and also one of the first fully terrestrial vertebrates to attain large size.

<span class="mw-page-title-main">Diadectidae</span> Extinct family of tetrapods

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.

<span class="mw-page-title-main">Captorhinidae</span> Extinct family of tetrapods

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.

<i>Seymouria</i> Extinct genus of reptile-like amphibians

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.

<span class="mw-page-title-main">Caseasauria</span> Extinct clade of synapsids

Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insectivorous or carnivorous Eothyrididae, and the large, herbivorous Caseidae. These two groups share a number of specialised features associated with the morphology of the snout and external naris.

<i>Westlothiana</i> Extinct genus of tetrapods

Westlothiana is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Viséan age of the Carboniferous. Members of the genus bore a superficial resemblance to modern-day lizards. The genus is known from a single species, Westlothiana lizziae. The type specimen was discovered in the East Kirkton Limestone at the East Kirkton Quarry, West Lothian, Scotland in 1984. This specimen was nicknamed "Lizzie the lizard" by fossil hunter Stan Wood, and this name was quickly adopted by other paleontologists and the press. When the specimen was formally named in 1990, it was given the specific name "lizziae" in homage to this nickname. However, despite its similar body shape, Westlothiana is not considered a true lizard. Westlothiana's anatomy contained a mixture of both "labyrinthodont" and reptilian features, and was originally regarded as the oldest known reptile or amniote. However, updated studies have shown that this identification is not entirely accurate. Instead of being one of the first amniotes, Westlothiana was rather a close relative of Amniota. As a result, most paleontologists since the original description place the genus within the group Reptiliomorpha, among other amniote relatives such as diadectomorphs and seymouriamorphs. Later analyses usually place the genus as the earliest diverging member of Lepospondyli, a collection of unusual tetrapods which may be close to amniotes or lissamphibians, or potentially both at the same time.

<i>Casineria</i> Species of tetrapodomorph (fossil)

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, 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.

<i>Limnoscelis</i> Genus of diadectomorphs

Limnoscelis was a genus of large diadectomorph tetrapods from the Late Carboniferous 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.

<i>Solenodonsaurus</i> Extinct genus of reptiles

Solenodonsaurus is an extinct genus of reptiliomorphs that lived in what is now Czech Republic, during the Westphalian stage.

<i>Tseajaia</i> Extinct genus of tetrapods

Tseajaia is an extinct genus of tetrapod. It was a basal diadectomorph that lived in the Permian of 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.

<i>Orobates</i> Extinct genus of reptiliomorphs

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.

<span class="mw-page-title-main">Limnoscelidae</span> Extinct family of tetrapods

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.

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.

<span class="mw-page-title-main">Organ Rock Formation</span>

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.

References

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