Varanopidae is an extinctfamily ofamniotes known from theLate Carboniferous toMiddle Permian that resembledmonitor lizards (with the name of the group deriving from the monitor lizard genusVaranus) and may have filled a similar niche. Typically, they are considered to be relativelybasalsynapsids (and thus more closely related to mammals than to reptiles), although some studies from the late 2010s recovered them being taxonomically closer todiapsidreptiles,[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 late Middle PermianPristerognathus Assemblage Zone (Capitanian) is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.[7] Thus, Varanopidae vanishes from the fossil record at the same time as dinocephalians,[8] plausibly as a result of a major mass extinction event[9] that has been called the "Dinocephalian extinction event".[10]
No known varanopids developed a sail likeDimetrodon. The length of known varanopids, including the tail, varies from 1 to 2 metres (3 to 7 ft).[11] 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 bymammalian 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 inEarly Permian, varanopids were agile creatures.
The genusAscendonanus provides the first extensive skin impressions for ancient amniotes, revealing scales akin to those ofsquamates.[12] Parental care is known inHeleosaurus, suggesting that it is ancestral to synapsids as a whole.[13]
Varanopids are small to medium-sized possible synapsids that have been discovered throughout the supercontinentPangea. 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.[14][15] 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.[16]
At least some varanopids likeAscendonanus andEoscansor 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.[17]
The poorly knownBasicranodon andRuthiromia 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 toAerosaurus.Basicranodon was found to be a wildcard taxon due to its small amount of known materials, as it is based on a partialbraincase from the ?Kungurian stage Richards Spur locality inOklahoma. It occupies two possible positions, falling either as a mycterosaurine, or as the sister taxon ofPyozia. Although Reiszet al. (1997) consideredBasicranodon as a subjective junior synonym ofMycterosaurus, 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 acladogram modified from the analysis of Benson (2012), after the exclusion ofBasicranodon:[18]
^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.
^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
^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.Bibcode:2010JVPal..30..724C.doi:10.1080/02724631003762914.S2CID84949154.
^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.Bibcode:2010JVPal..30.1628R.doi:10.1080/02724634.2010.501441.S2CID129835335.
.jpg)
