Varanopidae

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Varanopidae
Temporal range: Late Carboniferous - Middle Permian, 309–260  Ma
Varanops brevirostris Exhibit Museum of Natural History cropped.jpg
Fossil skeleton of Varanops brevirostris in the University of Michigan Museum of Natural History
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Clade: Amniota
Family: Varanopidae
Romer and Price, 1940
Genera

See below

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. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. [1] [2] [3] 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. [4]

Contents

Description

Fossil of Aerosaurus wellesi Aerosaurus wellesi 4.jpg
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). [5] 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 Varanodon1DB.jpg
Varanodon agilis
Skulls of Mesenosaurus (top) and Varanodon (bottom), showing variation in skull shape. Mesenosaurus and Varanodon.jpg
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. [6] Parental care is known in Heleosaurus , suggesting that it is ancestral to synapsids as a whole. [7]

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. [8] [9] 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. [10]

Ecology

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. [11]

Classification

Archaeovenator hamiltonensis ArchaeovenatorDB.jpg
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. [14]

Varanopidae

Archaeovenator

Apsisaurus

Mesenosaurus

Mycterosaurus

Varanodontinae

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: [12]

Varanopidae

Archaeovenator hamiltonensis

Pyozia mesenensis

Mycterosaurinae

Mycterosaurus longiceps

? Elliotsmithia longiceps (BP/1/5678)

Heleosaurus scholtzi

Mesenosaurus romeri

Varanodontinae

Varanops brevirostris

Watongia meieri

Varanodon agilis

Ruthiromia elcobriensis

Aerosaurus wellesi

Aerosaurus greenleorum

Related Research Articles

<span class="mw-page-title-main">Synapsid</span> Clade of tetrapods

Synapsids are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptiles and birds. The group includes mammals and every animal more closely related to mammals than to sauropsids. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.

<span class="mw-page-title-main">Diapsid</span> Clade of amniote tetrapods 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">Pelycosaur</span> Informal grouping composed of basal or primitive Late Paleozoic synapsids

Pelycosaur is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile had been used, and pelycosaur was considered an order, but this is now thought to be incorrect, and seen as outdated.

<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">Sphenacodontia</span> Clade of synapsids

Sphenacodontia is a stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes Haptodus baylei, Haptodus garnettensis and Sphenacodon ferox, but not Edaphosaurus pogonias". They first appear during the Late Pennsylvanian epoch. From the end of the Carboniferous to the end of the Permian, most of them remained large, with only some secondarily becoming small in size.

<span class="mw-page-title-main">Eupelycosauria</span> Clade of synapsids

Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils of Echinerpeton and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.

<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.

<span class="mw-page-title-main">Ophiacodontidae</span> Extinct family of synapsids

Ophiacodontidae is an extinct family of early eupelycosaurs from the Carboniferous and Permian. Archaeothyris, and Clepsydrops were among the earliest ophiacodontids, appearing in the Late Carboniferous. Ophiacodontids are among the most basal synapsids, an offshoot of the lineage which includes therapsids and their descendants, the mammals. The group became extinct by the Middle Permian, replaced by anomodonts, theriodonts, and the diapsid reptiles.

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

Mycterosaurus is an extinct genus of synapsids belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.

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

Varanops is an extinct genus of Early Permian varanopid synapsids known from Texas and Oklahoma of the United States. It was first named by Samuel Wendell Williston in 1911 as a second species of Varanosaurus, Varanosaurus brevirostris. In 1914, Samuel W. Williston reassigned it to its own genus and the type species is Varanops brevirostris.

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

Varanodon is an extinct genus of amniotes from the family Varanopidae. It has been found in the Chickasha Formation of Oklahoma, which dates to the Roadian stage of the Middle Permian. The largest varanopid known at the time of its description, with a skull length of 17.5 centimetres (6.9 in), it was closely related to and lived alongside its much larger relative Watongia. The two may represent growth stages of a single animal.

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

Mesenosaurus is an extinct genus of amniote. It belongs to the family Varanopidae. This genus includes two species: the type species Mesenosaurus romeri from the middle Permian Mezen River Basin of northern Russia, and Mesenosaurus efremovi from the early Permian (Artinskian) Richards Spur locality. M. romeri’s stratigraphic range is the middle to late Guadalupian while M. efremovi’s stratigraphic range is the Cisuralian.

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

Watongia is an extinct genus of non-mammalian synapsids from Middle Permian of Oklahoma. Only one species has been described, Watongia meieri, from the Chickasha Formation. It was assigned to family Gorgonopsidae by Olson and to Eotitanosuchia by Carroll. Reisz and collaborators assigned the genus in Varanopidae. Based on comparisons of its vertebrae with other varanopids, it was the largest varanopid with a body length of approximately 2 metres. It was a contemporary of its closest relative, the much smaller Varanodon; the two may possibly represent growth stages of a single animal.

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

Heleosaurus scholtzi is an extinct species of basal synapsids, known as pelycosaurs, in the family of Varanopidae during the middle Permian. At first H. scholtzi was mistakenly classified as a diapsid. Members of this family were carnivorous and had dermal armor, and somewhat resembled monitor lizards. This family was the most geologically long lived, widespread, and diverse group of early amniotes. To date only two fossils have been found in the rocks of South Africa. One of these fossils is an aggregation of five individuals.

<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.

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

Apsisaurus is an extinct genus of Early Permian varanopid synapsids known from Texas of the United States. It was first named by Michel Laurin in 1991 and the type species is Apsisaurus witteri. Apsisaurus witteri is known from the holotype MCZ 1474, a three-dimensionally preserved partial skeleton including an incomplete skull and mandibles. The skull roof of Archeria is also articulated to the postcranial skeleton. It was collected in the Archer City Bonebed 1 site, from the Archer City Formation of the Wichita Group, dating to the Early Permian epoch. 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.

Ascendonanus is an extinct genus of varanopid amniote from the Early Permian of Germany. It is one of the earliest specialized arboreal (tree-living) tetrapods currently known and outwardly resembled a small lizard. The animal was about 40 cm long, with strongly curved claws, short limbs, a slender, elongated trunk, and a long tail. It would have preyed on insects and other small arthropods.

Dendromaia is an extinct genus of varanopid from the Carboniferous of Nova Scotia. It contains a single species, Dendromaia unamakiensis. Dendromaia is the oldest known varanopid, likely the oldest known synapsid, and the only member of the family Varanopidae to be discovered in Nova Scotia. Known from a large partial skeleton preserved with its tail wrapped around a much smaller partial skeleton, Dendromaia may also represent the oldest known occurrence of parental care in the fossil record. While the larger skeleton possessed certain mycterosaurine-like features, the smaller skeleton resembled basal varanopids such as Archaeovenator and Pyozia, creating uncertainty over whether characteristics at the base of Varanopidae have legitimate phylogenetic significance or instead reflect the immaturity of basal varanopid specimens.

Anningia is an extinct genus in Varanopidae, a family of monitor lizard-like amniotes. It contains a single species, Anningia megalops.

References

  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. (January 2020). "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. 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.
  5. 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)
  6. 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.
  7. 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
  8. 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.
  9. 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.
  10. 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 .
  11. 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.
  12. 1 2 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.
  13. 1 2 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.
  14. 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.