Varanopidae Temporal range: Late Carboniferous - Middle Permian, | |
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Fossil skeleton of Varanops brevirostris in the University of Michigan Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
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. Although some studies from the late 2010s have 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]
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.
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]
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]
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 |
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Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida. The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. 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.
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.
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.
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.
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.
Ophiacodontidae is an extinct family of early synapsids 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 Kungurian or the Roadian, replaced by anomodonts, theriodonts, and the diapsid reptiles.
Mycterosaurus is an extinct genus of amniotes 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.
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.
Asaphestera is an extinct genus of a synapsid described on the basis of fossils from the Carboniferous of the Joggins locality in Nova Scotia, Canada. It was originally described as an undetermined lepospondyl and subsequently classified as a microsaur within the family Tuditanidae. A study published in May 2020 found that specimens referred to Asaphestera represented several unrelated species. Steen (1934)'s original species name Asaphestera platyris was retained for a skull which has been re-evaluated as the earliest known synapsid.
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.
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.
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.
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.
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.
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.
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.
Microvaranops is a Middle Permian synapsid of the family Varanopidae from the Abrahamskraal Formation of South Africa. It includes one species, Microvaranops parentis, which was probably arboreal. A slab containing five specimens of Microvaranops indicates that it gathered or lived in groups.
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, the only member of which 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.
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