Younginiformes Temporal range: | |
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Skull of Youngina | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Neodiapsida |
Order: | † Younginiformes Romer, 1945 |
Families | |
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Younginiformes is a group of diapsid reptiles known from the Permian-Triassic of Africa and Madagascar. It has been used as a replacement for "Eosuchia". [1] Younginiformes (including Acerosodontosaurus , Hovasaurus , Kenyasaurus , Tangasaurus , Thadeosaurus and Youngina ) were historically suggested to be lepidosauromorphs, but were later suggested to be basal non-saurian neodiapsids. [2] [3] The group is sometimes divided into two families, Tangasauridae and Younginidae. The monophyly of the group is disputed. A 2009 study found them to be an unresolved polytomy at the base of Neodiapsida, [4] while a 2011 study recovered the group as paraphyletic. [5] A 2022 study recovered the Younginiformes as a monophyletic group of basal neodiapsid reptiles, also including Claudiosaurus and Saurosternon as part of the group. [6] Some younginiforms like Hovasaurus and Acerosodontosaurus are thought to have had an amphibious lifestyle, while others like Kenyasaurus, Thadeosaurus and Youngina were probably terrestrial. [4]
Included genera:
2011 phylogeny showing a paraphyletic Younginiformes: [5]
Neodiapsida |
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Phylogeny of Younginiformes from the 2022 study: [6]
Younginiformes |
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Sauria is the clade containing the most recent common ancestor of Archosauria and Lepidosauria, and all its descendants. Since most molecular phylogenies recover turtles as more closely related to archosaurs than to lepidosaurs as part of Archelosauria, Sauria can be considered the crown group of diapsids, or reptiles in general. Depending on the systematics, Sauria includes all modern reptiles or most of them as well as various extinct groups.
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.
Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade Sauria.
Youngina is an extinct genus of diapsid reptile from the Late Permian Beaufort Group of the Karoo Red Beds of South Africa. This, and a few related forms, make up the family Younginidae, within the order Eosuchia. Eosuchia, having become a wastebasket taxon for many probably distantly-related primitive diapsid reptiles ranging from the Late Carboniferous to the Eocene, Romer proposed that it be replaced by Younginiformes.
Protorothyrididae is an extinct family of small, lizard-like reptiles belonging to Eureptilia. Their skulls did not have fenestrae, like the more derived diapsids. Protorothyridids lived from the Late Carboniferous to Early Permian periods, in what is now North America. Many genera of primitive reptiles were thought to be protorothyridids. Brouffia, Coelostegus, Paleothyris and Hylonomus, for example, were found to be more basal eureptiles in Muller and Reisz (2006), making the family as historically defined paraphyletic, though three genera, Protorothyris, Anthracodromeus, and Cephalerpeton, were recovered as a monophyletic group. Anthracodromeus, Paleothyris, and Protorothyris were recovered as a monophyletic group in Ford and Benson (2020), who recovered them as more derived than captorhinids and Hylonomus, but less so than araeoscelidians. Anthracodromeus is the earliest known reptile to display adaptations to climbing. The majority of phylogenetic studies recover protorothyridids as basal members of Eureptilia; however, Simões et al. (2022) recover them as stem-amniotes instead.
Avicephala is a potentially polyphyletic grouping of extinct diapsid reptiles that lived during the Late Permian and Triassic periods characterised by superficially bird-like skulls and arboreal lifestyles. As a clade, Avicephala is defined as including the gliding weigeltisaurids and the arboreal drepanosaurs to the exclusion of other major diapsid groups. This relationship is not recovered in the majority of phylogenetic analyses of early diapsids and so Avicephala is typically regarded as an unnatural grouping. However, the clade was recovered again in 2021 in a redescription of Weigeltisaurus, raising the possibility that the clade may be valid after all.
Parareptilia ("near-reptiles") is a subclass or clade of basal sauropsids/reptiles, typically considered the sister taxon to Eureptilia. Parareptiles first arose near the end of the Carboniferous period and achieved their highest diversity during the Permian period. Several ecological innovations were first accomplished by parareptiles among reptiles. These include the first reptiles to return to marine ecosystems (mesosaurs), the first bipedal reptiles, the first reptiles with advanced hearing systems, and the first large herbivorous reptiles. The only parareptiles to survive into the Triassic period were the procolophonoids, a group of small generalists, omnivores, and herbivores. The largest family of procolophonoids, the procolophonids, rediversified in the Triassic, but subsequently declined and became extinct by the end of the period.
Eosuchians are an extinct order of diapsid reptiles. Depending on which taxa are included the order may have ranged from the late Carboniferous to the Eocene but the consensus is that eosuchians are confined to the Permian and Triassic.
Claudiosaurus is an extinct genus of diapsid reptiles from the Late Permian Sakamena Formation of the Morondava Basin, Madagascar. It has been suggested to be semi-aquatic.
Thadeosaurus is an extinct genus of diapsid reptile belonging to the family Younginidae. Fossils have been found in the Lower Sakamena Formation of the Morondava Basin, Madagascar in 1981, and date to the late Permian to the early Triassic period.
Hovasaurus is an extinct genus of basal diapsid reptile. It lived in what is now Madagascar during the Late Permian and Early Triassic, being a survivor of the Permian–Triassic extinction event and the paleontologically youngest member of the Tangasauridae. Fossils have been found in the Permian Lower and Triassic Middle Sakamena Formations of the Sakamena Group, where it is amongst the commonest fossils. Its morphology suggests an aquatic ecology.
Acerosodontosaurus is an extinct genus of neodiapsid reptiles that lived during the Late Permian of Madagascar. The only species of Acerosodontosaurus, A. piveteaui, is known from a natural mold of a single partial skeleton including a crushed skull and part of the body and limbs. The fossil was discovered in deposits of the Lower Sakamena Formation. Based on skeletal characteristics, it has been suggested that Acerosodontosaurus individuals were at least partially aquatic.
Tangasauridae is an extinct family of diapsids known from fossil specimens from Madagascar, Kenya and Tanzania that are Late Permian to Early Triassic in age. Fossils have been found of numerous specimens of common members of this family such as Hovasaurus in different stages of ontogenic development. Recent material from the Middle Sakamena Formation of the Morondava Basin of Madagascar that dates back to the early Triassic period suggests that the Tangasauridae were relatively unaffected by the Permian-Triassic extinction event.
Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.
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.
Tangasaurus is an extinct genus of aquatic basal tangasaurid neodiapsid known from the Late Permian period of Tanga, northeastern Tanzania. It contains a single species, Tangasaurus mennelli.
Kenyasaurus is an extinct genus of basal tangasaurid known from the Early Triassic period of Coast Province, southeastern Kenya. It contains a single species, Kenyasaurus mariakaniensis.
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.
Palaeagama is an extinct genus of neodiapsid reptile from the Late Permian or Early Triassic of South Africa. It is based on an articulated skeleton which was probably found in the Early Triassic Lystrosaurus Assemblage Zone, or potentially the Late Permian Daptocephalus Assemblage Zone. Despite the completeness of the specimen, Palaeagama is considered as a "wildcard" taxon of uncertain affinities due to poor preservation. It was originally considered an "eosuchian", and later reinterpreted as a lizard ancestor closely related to Paliguana and Saurosternon. Modern studies generally consider it an indeterminate neodiapsid, though a few phylogenetic analyses tentatively support a position at the base of Lepidosauromorpha.