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Varanopidae

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Varanopidae is an extinct family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus Varanus) and may have filled a similar niche. Typically, they are considered to be relatively basal synapsids (and thus more closely related to mammals than to reptiles), although some studies from the late 2010s recovered them being taxonomically closer to diapsid reptiles,[1][2][3] 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.[4][5][6] 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.[7]

Varanopidae
Temporal range: Late Carboniferous - Middle Permian, 309–260 Ma
Fossil skeleton of Varanops brevirostris in the University of Michigan Museum of Natural History
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Family: †Varanopidae
Romer and Price, 1940
Genera

See below

Description

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Fossil of Aerosaurus wellesi

No known varanopids developed a sail like Dimetrodon. The length of known varanopids, including the tail, varies from 1 to 2 metres (3 to 7 ft).[8] Varanopids already showed some advanced characteristics of true pelycosaurs such as their deep, narrow, elongated skulls. Their jaws were long and their teeth were sharp. However, they were still primitive by mammalian standards. They had long tails, lizard-like bodies, and thin legs. The varanopids were mostly carnivorous, but as they were reduced in size, their diets changed from a carnivorous to an insectivorous lifestyle. Compared to the other animals in Early Permian, varanopids were agile creatures.

 
Varanodon agilis
 
Skulls of Mesenosaurus (top) and Varanodon (bottom), showing variation in skull shape.

The genus Ascendonanus provides the first extensive skin impressions for ancient amniotes, revealing scales akin to those of squamates.[9] Parental care is known in Heleosaurus, suggesting that it is ancestral to synapsids as a whole.[10]

Varanopids are small to medium-sized possible synapsids that have been discovered throughout the supercontinent Pangea. Varanopids are found in formerly areas of North America, Russia, Europe, and South Africa. The authors Romer and Price (1940) discussed the original positioning of Varanopidae within Synapsida and considered them as the suborder Sphenacodontia. Most phylogenetic analyses have place Varanopidae as a basal member of Synapsida and due to their positioning, a better understanding of the morphology and phylogeny of varanopids is needed for synapsid evolution. The phylogeny of varanopids is based mostly on cranial morphology.[11][12] The atlas−axis complex can be described with little effort with variation of this structure within a small clade. Varanopids, members of synapsid predators have well preserved atlas−axes permitting a descriptions and examination of morphological variation between taxon. The size of the transverse processes on the axis and the shape of the axial neural spine can be variable. For the small mycterosaurine varanopids, they have a small transverse processes that point posteroventrally, and the axial spine is dorsoventrally short, with a flattened dorsal margin in lateral view. The larger varanodontine varanopids have large transverse processes with a broad base, and a much taller axial spine with a rounded dorsal margin in lateral view. Using outgroup comparisons, the morphology of the transverse processes is considered a derived trait in varanodontines, while in mycterosaurines the morphology of the axial spine is the derived trait.[13]

Ecology

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At least some varanopids like Ascendonanus and Eoscansor are amongst the oldest known tree climbing (arboreal) animals, with limbs and digits adapted for grasping. Other varanopids lacked these adaptations and were probably terrestrial.[14]

Classification

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Archaeovenator hamiltonensis

Family Varanopidae

Apsisaurus was formerly assigned as an "eosuchian" diapsid. In 2010, it was redescribed by Robert R. Reisz, Michel Laurin and David Marjanović; their phylogenetic analysis found it to be a basal varanopid synapsid. The cladogram below is modified after Reisz, Laurin and Marjanović, 2010.[18]

Varanopidae

The poorly known Basicranodon and Ruthiromia were tentatively assigned to Varanopidae by Reisz (1986), but have been neglected in more recent studies. They were included for the first time in a phylogenetic analysis by Benson (2012). Ruthiromia was found to be most closely related to Aerosaurus. Basicranodon was found to be a wildcard taxon due to its small amount of known materials, as it is based on a partial braincase from the ?Kungurian stage Richards Spur locality in Oklahoma. It occupies two possible positions, falling either as a mycterosaurine, or as the sister taxon of Pyozia. Although Reisz et al. (1997) considered Basicranodon as a subjective junior synonym of Mycterosaurus, Benson (2012) found some differences in the distribution of teeth and shape of the dentigerous ventral platform medial to the basipterygoid processes that may indicate taxonomic distinction. Below is a cladogram modified from the analysis of Benson (2012), after the exclusion of Basicranodon:[15]

Varanopidae

References

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  1. ^ Ford, David P.; Benson, Roger B. J. (2018). "A redescription of Orovenator mayorum (Sauropsida, Diapsida) using high‐resolution μCT, and the consequences for early amniote phylogeny". Papers in Palaeontology. 5 (2): 197–239. doi:10.1002/spp2.1236. S2CID 92485505.
  2. ^ Modesto, Sean P. (December 23, 2019). "Rooting about reptile relationships". Nature Ecology & Evolution. 4 (1): 10–11. doi:10.1038/s41559-019-1074-0. ISSN 2397-334X. PMID 31900449. S2CID 209672518.
  3. ^ MacDougall, Mark J.; Modesto, Sean P.; Brocklehurst, Neil; Verrière, Antoine; Reisz, Robert R.; Fröbisch, Jörg (2018). "Commentary: A Reassessment of the Taxonomic Position of Mesosaurs, and a Surprising Phylogeny of Early Amniotes". Frontiers in Earth Science. 6. doi:10.3389/feart.2018.00099. ISSN 2296-6463.
  4. ^ Bazzana, K. D.; Evans, D. C.; Bevitt, J. J.; Reisz, R. R. (2021). "Neurosensory anatomy of Varanopidae and its implications for early synapsid evolution". Journal of Anatomy. 240 (5): 833–849. doi:10.1111/joa.13593. PMC 9005680. PMID 34775594. S2CID 244116303.
  5. ^ Benoit, J.; Ford, D. P.; Miyamae, J. A.; Ruf, I. (2021). "Can maxillary canal morphology inform varanopid phylogenetic affinities?". Acta Palaeontologica Polonica. 66 (2): 389–393. doi:10.4202/app.00816.2020. S2CID 237333701.
  6. ^ Simões, T.; Kammerer, C. (August 2022). "Successive climate crises in the deep past drove the early evolution and radiation of reptiles". Science Advances. 08 (33): eabq1898. Bibcode:2022SciA....8.1898S. doi:10.1126/sciadv.abq1898. PMC 9390993. PMID 35984885. S2CID 251694019.
  7. ^ Modesto, S.P.; Smith, R.M.H.; Campione, N.E.; Reisz, R.R. (2011). "The last "pelycosaur": a varanopid synapsid from the Pristerognathus Assemblage Zone, Middle Permian of South Africa". Naturwissenschaften. 98 (12): 1027–1034. Bibcode:2011NW.....98.1027M. doi:10.1007/s00114-011-0856-2. PMID 22009069. S2CID 27865550.
  8. ^ Reisz, R.R. & Laurin, M. (2004). "A reevaluation of the enigmatic Permian synapsid Watongia and of its stratigraphic significance". Canadian Journal of Earth Sciences. 41 (4): 377–396. Bibcode:2004CaJES..41..377R. doi:10.1139/e04-016.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Frederik Spindler; Ralf Werneburg; Joerg W. Schneider; Ludwig Luthardt; Volker Annacker; Ronny Rößler (2018). "First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny". PalZ. in press. doi:10.1007/s12542-018-0405-9.
  10. ^ Botha-Brink, Jennifer. "A Mixed-Age Classed 'Pelycosaur' Aggregation from South Africa: Earliest Evidence of Parental Care in Amniotes?" Proceedings of the Royal Society B: Biological Sciences 274.1627 (2007): 2829-834. JSTOR. Web. 06 Mar. 2017
  11. ^ Maddin, H. C.; Evans, D. C.; Reisz, R. R. (2006). "An Early Permian Varanodontine Varanopid (Synapsida: Eupelycosauria) from the Richards Spurs Locality, Oklahoma". Journal of Vertebrate Paleontology. 26 (4): 957–966. doi:10.1671/0272-4634(2006)26[957:AEPVVS]2.0.CO;2. JSTOR 4524646. S2CID 130455511.
  12. ^ Campione, N.; Reisz, R. (2010). "Varanops brevirostris (Eupelycosauria: Varanopidae) from the Lower Permian of Texas, with Discussion of Varanopid Morphology and Interrelationships". Journal of Vertebrate Paleontology. 30 (3): 724–746. doi:10.1080/02724631003762914. S2CID 84949154.
  13. ^ Campione, N. E.; Reisz, R. R. (2011). "Morphology and Evolutionary Significance of the Atlas-axis Complex in Varanopid Synapsids" (PDF). Acta Palaeontologica Polonica. 56 (4): 739–748. doi:10.4202/app.2010.0071.
  14. ^ Lucas, Spencer G.; Rinehart, Larry F.; Celeskey, Matthew D.; Berman, David S.; Henrici, Amy C. (June 2022). "A Scansorial Varanopid Eupelycosaur from the Pennsylvanian of New Mexico". Annals of Carnegie Museum. 87 (3): 167–205. doi:10.2992/007.087.0301. ISSN 0097-4463. S2CID 250015681.
  15. ^ a b Benson, R.J. (2012). "Interrelationships of basal synapsids: cranial and postcranial morphological partitions suggest different topologies". Journal of Systematic Palaeontology. 10 (4): 601–624. doi:10.1080/14772019.2011.631042. S2CID 84706899.
  16. ^ a b Spindler, F.; Werneburg, R.; Schneider, J. W.; Luthardt, L.; Annacker, V.; Rößler, R. (2018). "First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny". PalZ. 92 (2): 315–364. doi:10.1007/s12542-018-0405-9. S2CID 133846070.
  17. ^ Spindler, Frederik; Werneburg, Ralf; Schneider, Jörg W. (2019-06-01). "A new mesenosaurine from the lower Permian of Germany and the postcrania of Mesenosaurus: implications for early amniote comparative osteology". PalZ. 93 (2): 303–344. doi:10.1007/s12542-018-0439-z. ISSN 1867-6812. S2CID 91871872.
  18. ^ Robert R. Reisz, Michel Laurin and David Marjanović (2010). "Apsisaurus witteri from the Lower Permian of Texas: yet another small varanopid synapsid, not a diapsid". Journal of Vertebrate Paleontology. 30 (5): 1628–1631. doi:10.1080/02724634.2010.501441. S2CID 129835335.
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