Biology:Araeoscelidia

Araeoscelidia or Araeoscelida is a clade of extinct tetrapods (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.

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, Petrolacosaurus Lane 1945 and Spinoaequalis, known from virtually complete skeletons. Zarcasaurus, Aphelosaurus and Kadaliosaurus 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) but this inclusion has been criticized e.g., by Evans (1988), 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, but lack a scientific description as of 2023.

Phylogeny

The majority of historical phylogenetic studies recovered araeoscelidians as the most basal group of diapsids:

Cladogram after Bickelmann et al., 2009 and Reisz et al., 2011:

Sauropsida †Parareptilia     †Millerettidae 50px           †Eunotosaurus     †Hallucicrania   †Lanthanosuchidae 50px   †Procolophonia   †Procolophonoidea 50px     †Pareiasauromorpha 60px           Eureptilia     †Captorhinidae 50px     Romeriida     †Paleothyris     Diapsida   †Araeoscelidia   Neodiapsida   †Orovenator       †Lanthanolania       †Tangasauridae       †Younginidae         †Claudiosaurus             †Palaeagama     †Saurosternon             †Coelurosauravus           †Thalattosauria       †Hupehsuchia     †Ichthyopterygia       Sauria   Lepidosauromorpha     Archosauromorpha

However, Simões et al. (2022) recover them as stem-amniotes instead, as the sister group to the clade including Captorhinidae and Protorothyris archeri.

Reptiliomorpha	Seymouriamorpha 100px       Diadectomorpha 100px         Araeoscelidia 100px       Protorothyris 100px     Captorhinidae 100px       Amniota   Synapsida 100px   Sauropsida   Acleistorhinidae 100px       Millerettidae 100px       Mesosauria 100px   Neoreptilia   Procolophonia 100px     Neodiapsida (including modern reptiles and birds)

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). Apart from araeoscelidans, only one other diapsid is known before the Late Permian: Orovenator from the Early Permian of Oklahoma.

Bibliography

• Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. New York: W.H. Freeman and Co. ISBN 0-7167-1822-7. https://archive.org/details/vertebratepaleon0000carr. 
• Benton, Michael J. (2000). Vertebrate Paleontology (2nd ed.). Oxford: Blackwell Science. ISBN 0-632-05614-2. 
• deBraga, M.; Reisz, R. R. (1995). "A new diapsid reptile from the uppermost Carboniferous (Stephanian) of Kansas". Palaeontology 38: 199–212. 
• deBraga, M.; Rieppel, O. (1997). "Reptile phylogeny and the interrelationships of turtles". Zoological Journal of the Linnean Society 120 (3): 281–354. doi:10.1111/j.1096-3642.1997.tb01280.x. 
• Brinkman, D. B.; Berman, D. S.; Eberth, D. Z. (1984). "A new araeoscelid reptile, Zarcasaurus tandyderus, from the Culter Formation, (Lower Permian) of north-central New Mexico". New Mexico Geology 6 (2): 34–39. https://geoinfo.nmt.edu/publications/periodicals/nmg/6/n2/nmg_v6_n2_p34.pdf. 
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• Evans, S. E. (1988). "The early history and relationships of the Diapsida". in Benton, M. J.. The phylogeny and classification of the tetrapods. 1: Amphibians, Reptiles, Birds. Systematics Association Special Volume 35 A. Oxford: Clarendon Press. pp. 221–260. ISBN 0-19-857705-2. 
• Falconnet, J.; Steyer, J.-S. (2007). "Revision, osteology and locomotion of Aphelosaurus, an enigmatic reptile from the Lower Permian of France". Journal of Morphology (abstract of the 8th International Congress of Vertebrate Morphology, Paris, July 2007): 38. 
• Gervais, P. (1859). Zoologie et paléontologie française (2nd ed.). Paris: Bertrand. 
• Laurin, M. (1991). "The osteology of a Lower Permian eosuchian from Texas and a review of a diapsid phylogeny". Zoological Journal of the Linnean Society 101: 59–95. doi:10.1111/j.1096-3642.1991.tb00886.x. 
• May, W. J.; Hall, J. D. (2002). "Geology and vertebrate fauna of a new site in the Wellington Formation (Lower Permian) of Northern Oklahoma". Oklahoma Geology Notes 62 (2): 63–66. 
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• Reisz, R. R.; Berman, D. S.; Scott, D. (1984). "The anatomy and relationships of the Lower Permian reptile Araeoscelis". Journal of Vertebrate Paleontology 4 (1): 57–67. doi:10.1080/02724634.1984.10011986. 
• Reisz, Robert R.; Modesto, Sean P.; Scott, Diane M. (2011). "A new Early Permian reptile and its significance in early diapsid evolution". Proceedings of the Royal Society B 278 (1725): 3731–3737. doi:10.1098/rspb.2011.0439. PMID 21525061. 
• Swanson, B. A.; Carlson, K. J. (2002). "Walk, Wade, or Swim? Vertebrate Traces on an Early Permian Lakeshore". PALAIOS 17 (2): 123–133. doi:10.1669/0883-1351(2002)017<0123:WWOSVT>2.0.CO;2. Bibcode: 2002Palai..17..123S. http://doc.rero.ch/record/14748/files/PAL_E1875.pdf. 
• Thévenin, A. (1910). "Les plus anciens quadrupèdes de France". Annales de Paléontologie 5: 1–65. 
• Vaughn, P. P. (1955). "The Permian reptile Araeoscelis restudied". Bulletin of the Harvard Museum of Comparative Zoology 113: 305–467. 

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