Araeoscelidia

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Araeoscelidans
Temporal range: CarboniferousPermian 302–275.6  Ma
Petrolacosaurus BW.jpg
Petrolacosaurus skull diagram.png
Life restoration (top) and skull reconstruction (bottom) of Petrolacosaurus kansensis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Diapsida
Order: Araeoscelidia
Williston, 1913
Genera

Araeoscelidia or Araeoscelida is a clade of extinct amniotes (traditionally classified as diapsid reptiles) superficially resembling lizards, extending from the Late Carboniferous to the Early Permian. The group contains the genera Araeoscelis , Petrolacosaurus , the possibly aquatic Spinoaequalis , and less well-known genera such as Kadaliosaurus and Zarcasaurus . This clade is usually considered to be the sister group to all (currently known) later diapsids.

Contents

Description

Araeoscelidians were small animals (less than one meter in length) looking somewhat like lizards, though they are only distantly related to true lizards. They differ from other, earlier sauropsids by their slender limbs, their elongated tail, and of course by the presence of two temporal openings, the feature defining the diapsid condition. In Araeoscelis , only the upper temporal opening remains, thus resulting in a derived euryapsid condition.

Genera

Araeoscelidia includes well-known genera such as Araeoscelis Williston 1910, [1] [2] Petrolacosaurus Lane 1945 [3] [4] and Spinoaequalis , [5] [6] known from virtually complete skeletons. Zarcasaurus , [7] Aphelosaurus [8] [9] [10] and Kadaliosaurus [11] belong to this clade but are known only from post-cranial remains and a mandible fragment for Zarcasaurus.

The genus Dictybolos has been included in Araeoscelidia by Olson (1970) [12] but this inclusion has been criticized e.g., by Evans (1988), [13] especially since Olson also included distantly related groups such as protorosaurs and mesosaurs.

New specimens have been discovered in the United States state of Oklahoma, [14] [15] but lack a scientific description as of 2023.

Phylogeny

The majority of phylogenetic studies recover araeoscelidians as the most basal group of diapsids; however, Simões et al. (2022) recover them as stem-amniotes instead, as the sister group to the clade including Captorhinidae and Protorothyris archeri. [16]

Stratigraphic and geographic distribution

Araeoscelidia are known from the Late Carboniferous in the United States (Petrolacosaurus, Spinoaequalis) to the Early Permian in France (Aphelosaurus), Germany (Kadaliosaurus) and the United States (Dictybolos, Zarcasaurus, Araeoscelis, Halgaitosaurus [17] ). Apart from araeoscelidans, only one other diapsid is known before the Late Permian: Orovenator from the Early Permian of Oklahoma. [18]

Related Research Articles

<span class="mw-page-title-main">Diapsid</span> Clade of reptiles with two holes in each side of their skulls

Diapsids are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago during the late Carboniferous period. All diapsids other than the most primitive ones in the clade Araeoscelidia are sometimes placed into the clade Neodiapsida. The diapsids are extremely diverse, and include birds and all modern reptile groups, including turtles, which were historically thought to lie outside the group. Although some diapsids have lost either one hole (lizards), or both holes, or have a heavily restructured skull, they are still classified as diapsids based on their ancestry. At least 17,084 species of diapsid animals are extant: 9,159 birds, and 7,925 snakes, lizards, tuatara, turtles, and crocodiles.

<span class="mw-page-title-main">Mesosaur</span> Extinct family of reptiles

Mesosaurs were a group of small aquatic reptiles that lived during the early Permian period (Cisuralian), roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to an aquatic lifestyle from more terrestrial ancestors. It is uncertain which and how many terrestrial traits these ancestors displayed; recent research cannot establish with confidence if the first amniotes were fully terrestrial, or only amphibious. Most authors consider mesosaurs to have been aquatic, although adult animals may have been amphibious, rather than completely aquatic, as indicated by their moderate skeletal adaptations to a semiaquatic lifestyle. Similarly, their affinities are uncertain; they may have been among the most basal sauropsids or among the most basal parareptiles.

<span class="mw-page-title-main">Eureptilia</span> Taxon of tetrapods

Eureptilia is one of the two major subgroups of the clade Sauropsida, the other one being Parareptilia. Eureptilia includes Diapsida, as well as a number of primitive Permo-Carboniferous forms previously classified under Anapsida, in the old order "Cotylosauria".

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

Youngina is an extinct genus of diapsid reptile from the Late Permian Beaufort Group of the Karoo Red Beds of South Africa. This, and a few related forms, make up the family Younginidae, within the order Eosuchia. Eosuchia, having become a wastebasket taxon for many probably distantly-related primitive diapsid reptiles ranging from the Late Carboniferous to the Eocene, Romer proposed that it be replaced by Younginiformes.

<span class="mw-page-title-main">Protorothyrididae</span> Family of reptiles

Protorothyrididae is an extinct family of small, lizard-like reptiles belonging to Eureptilia. Their skulls did not have fenestrae, like the more derived diapsids. Protorothyridids lived from the Late Carboniferous to Early Permian periods, in what is now North America. Many genera of primitive reptiles were thought to be protorothyridids. Brouffia, Coelostegus, Paleothyris and Hylonomus, for example, were found to be more basal eureptiles in Muller and Reisz (2006), making the family as historically defined paraphyletic, though three genera, Protorothyris, Anthracodromeus, and Cephalerpeton, were recovered as a monophyletic group. Anthracodromeus, Paleothyris, and Protorothyris were recovered as a monophyletic group in Ford and Benson (2020), who recovered them as more derived than captorhinids and Hylonomus, but less so than araeoscelidians. Anthracodromeus is the earliest known reptile to display adaptations to climbing. The majority of phylogenetic studies recover protorothyridids as basal members of Eureptilia; however, Simões et al. (2022) recover them as stem-amniotes instead.

<span class="mw-page-title-main">Dissorophidae</span> Extinct family of temnospondyls

Dissorophidae is an extinct family of medium-sized temnospondyls that flourished during the late Carboniferous and early Permian periods. The clade is known almost exclusively from North America.

<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>Petrolacosaurus</i> Genus of tetrapods

Petrolacosaurus is an extinct genus of diapsid reptile from the late Carboniferous period. It was a small, 40-centimetre (16 in) long reptile, and one of the earliest known reptile with two temporal fenestrae. This means that it was at the base of Diapsida, the largest and most successful radiation of reptiles that would eventually include all modern reptile groups, as well as dinosaurs and other famous extinct reptiles such as plesiosaurs, ichthyosaurs, and pterosaurs. However, Petrolacosaurus itself was part of Araeoscelida, a short-lived early branch of the diapsid family tree which went extinct in the mid-Permian.

Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an Archaeothyris-like synapsid in the Late Carboniferous. Although some studies from the late 2010s have recovered them being taxonomically closer to diapsid reptiles, recent studies from the early 2020s support their traditional placement as synapsids on the basis of high degree of bone labyrinth ossification, maxillary canal morphology and phylogenetic analyses. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.

<span class="mw-page-title-main">Procolophonia</span> Extinct suborder of reptiles

Procolophonia is an extinct suborder (clade) of herbivorous reptiles that lived from the Middle Permian till the end of the Triassic period. They were originally included as a suborder of the Cotylosauria but are now considered a clade of Parareptilia. They are closely related to other generally lizard-like Permian reptiles such as the Millerettidae, Bolosauridae, Acleistorhinidae, and Lanthanosuchidae, all of which are included under the Anapsida or "Parareptiles".

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

Araeoscelis is an extinct genus of tetrapods from the Early Permian of what is now Texas. Fossils have been found in the Nocona, Arroyo and Waggoner Ranch Formations. Two species have been described, A. casei and A. gracilis.

<span class="mw-page-title-main">Trematopidae</span> Extinct family of temnospondyls

Trematopidae is a family of dissorophoid temnospondyls spanning the late Carboniferous to the early Permian. Together with Dissorophidae, the family forms Olsoniformes, a clade comprising the medium-large terrestrial dissorophoids. Trematopids are known from numerous localities in North America, primarily in New Mexico, Oklahoma, and Texas, and from the Bromacker quarry in Germany.

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

Acleistorhinus (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian of Oklahoma. It is notable for being the earliest known anapsid reptile yet discovered. The morphology of the lower temporal fenestra of the skull of Acleistorhinus bears a superficial resemblance to that seen in early synapsids, a result of convergent evolution. Only a single species, A. pteroticus, is known, and it is classified in the Family Acleistorhinidae, along with Colobomycter.

<i>Ianthodon</i> Extinct genus of synapsids

Ianthodon is an extinct genus of basal haptodontiform synapsids from the Late Carboniferous about 304 million years ago. The taxon was discovered and named by Kissel & Reisz in 2004. The only species in the taxon, Ianthodon schultzei, was found by separating it from a block that also contained the remains of Petrolacosaurus and Haptodus. The evolutionary significance of the taxon wasn't realized until a publication in 2015. The fossil of this organism was discovered in Garnett, Kansas.

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 2011.

<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.

<i>Orovenator</i>

Orovenator is an extinct genus of diapsid from Lower Permian deposits of Oklahoma, United States. It is known from two partial skulls from the Richards Spur locality in Oklahoma. The holotype OMNH 74606 consists of a partial skull preserving snout and mandible, and the referred specimen, OMNH 74607, a partial skull preserving the skull roof, vertebrae and palatal elements. It was first named by Robert R. Reisz, Sean P. Modesto and Diane M. Scott in 2011 and the type species is Orovenator mayorum. The generic name means "mountain", oro, in Greek in reference to the Richards Spur locality, which was mountainous during the Permian period and "hunter", venator, in Latin. The specific name honours Bill and Julie May. Orovenator is the oldest and most basal neodiapsid to date.

<span class="mw-page-title-main">Younginidae</span> Extinct family of reptiles

Younginidae is an extinct family of diapsid reptiles from the Late Permian and Early Triassic. In a phylogenetic context, younginids are near the base of the clade Neodiapsida. Younginidae includes the species Youngina capensis from the Late Permian of South Africa and Thadeosaurus colcanapi from the Late Permian and Early Triassic of Madagascar. Heleosuchus griesbachi from the Late Permian of South Africa may also be a member of the family.

<span class="mw-page-title-main">Richards Spur</span> Permian fossil locality in Oklahoma

Richards Spur is a Permian fossil locality located at the Dolese Brothers Limestone Quarry north of Lawton, Oklahoma. The locality preserves clay and mudstone fissure fills of a karst system eroded out of Ordovician limestone and dolomite, with the infilling dating to the Artinskian stage of the early Permian (Cisuralian), around 289 to 286 million years ago. Fossils of terrestrial animals are abundant and well-preserved, representing one of the most diverse Paleozoic tetrapod communities known. A common historical name for the site is Fort Sill, in reference to the nearby military base. Fossils were first reported at the quarry by workers in 1932, spurring a wave of collecting by local and international geologists. Early taxa of interest included the abundant reptile Captorhinus and microsaurs such as Cardiocephalus and Euryodus. Later notable discoveries include Doleserpeton, the most diverse assortment of parareptiles in the Early Permian, and the rare early diapsid Orovenator.

References

  1. Vaughn 1955
  2. Reisz, Berman & Scott 1984
  3. Peabody 1952
  4. Reisz 1981
  5. deBraga & Reisz 1995
  6. deBraga & Rieppel 1997
  7. Brinkman, Berman & Eberth 1984
  8. Gervais 1859
  9. Thévenin 1910
  10. Falconnet & Steyer 2007
  11. Credner 1889
  12. Olson 1970
  13. Evans 1988
  14. May & Hall 2002
  15. Swanson & Carlson 2002
  16. Simões, T. R.; Kammerer, C. F.; Caldwell, M. W.; Pierce, S. E. (2022). "Successive climate crises in the deep past drove the early evolution and radiation of reptiles". Science Advances. 8 (33): eabq1898. doi: 10.1126/sciadv.abq1898 . PMC   9390993 . PMID   35984885.
  17. Henrici, Amy C.; Berman, David S; Sumida, Stuart S.; Huttenlocker, Adam K. (2023-11-15). "Halgaitosaurus gregarius, a New Upper Carboniferous Araeoscelidian (Reptilia: Diapsida) from the Halgaito Formation, Bears Ears National Monument, Utah, USA". Annals of Carnegie Museum. 88 (3). doi:10.2992/007.088.0301. ISSN   0097-4463.
  18. Reisz, Modesto & Scott 2011

Further reading