Ankyramorphs Temporal range: | |
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Skull of Lanthanosuchus watsoni , a lanthanosuchoid | |
Skeleton of Scutosaurus karpinskii , a procolophonian | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | † Parareptilia |
Order: | † Procolophonomorpha |
Node: | † Ankyramorpha DeBraga & Rieppel, 1996 |
Subgroups | |
Ankyramorpha ("anchor forms") is an extinct clade of procolophonomorph parareptiles which lived between the early Cisuralian epoch (middle Sakmarian stage) and the latest Triassic period (latest Rhaetian stage) of Africa, Antarctica, Asia, Australia, Europe, North America and South America. [1] [2]
This clade was named in a 1996 parareptile study by Michael deBraga and Robert R. Reisz. They provided the name Ankyramorpha for a newly recognized clade encompassing "the most recent common ancestor of Procolophonia and Lanthanosuchoidea and all its descendants", and this clade name sees continued use among modern parareptile studies. [3] [4] [5] A similar name, Hallucicrania, was provided in an earlier 1995 study by Michael S. Y. Lee, who defined it as the node-based taxon formed by the most recent common ancestor of lanthanosuchids and "pareiasauroids" (pareiasaurs + Sclerosaurus ), and all its descendants. Unlike Ankyramorpha, which explicitly included procolophonoids, Hallucicrania was originally designed to exclude procolophonoids, which Lee's analysis argued to have split off prior to the divergence between lanthanosuchids and pareiasaurs. [6] Nevertheless, purely considering the taxa encompassed by their definitions, Hallucicrania and Ankyramorpha refer to an identical grouping.
The following cladogram is simplified after the phylogenetic analysis of MacDougall and Reisz (2014) and shows the placement of Ankyramorpha within Parareptilia. Relationships within emboldened terminal clades are not shown. [5]
Parareptilia |
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An anapsid is an amniote whose skull lacks one or more skull openings near the temples. Traditionally, the Anapsida are the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolved, making anapsids paraphyletic. It is however doubtful that all anapsids lack temporal fenestra as a primitive trait, and that all the groups traditionally seen as anapsids truly lacked fenestra.
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.
Pareiasaurs are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with osteoderms which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, before becoming globally distributed during the Late Permian. Pareiasaurs were the largest reptiles of the Permian, reaching sizes equivalent to those of contemporary therapsids. Pareiasaurs became extinct at the end of the Permian during the Permian-Triassic extinction event.
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.
The Procolophonia are a suborder of herbivorous reptiles that lived from the Middle Permian till the end of the Triassic period. They were originally included as a suborder of the Cotylosauria but are now considered a clade of Parareptilia. They are closely related to other generally lizard-like Permian reptiles such as the Millerettidae, Bolosauridae, Acleistorhinidae, and Lanthanosuchidae, all of which are included under the Anapsida or "Parareptiles".
Procolophonomorpha is an order or clade containing most parareptiles. Many papers have applied various definitions to the name, though most of these definitions have since been considered synonymous with modern parareptile clades such as Ankyramorpha and Procolophonia. The current definition of Procolophonomorpha, as defined by Modesto, Scott, & Reisz (2009), is that of as a stem-based group containing Procolophon and all taxa more closely related to it than to Milleretta. It constitutes a diverse assemblage that includes a number of lizard-like forms, as well as more diverse types such as the pareiasaurs. Lee 1995, 1996, 1997 argues that turtles evolved from pareiasaurs, but this view is no longer considered likely. Rieppel and deBraga 1996 and deBraga and Rieppel, 1997 argue that turtles evolved from sauropterygians, and there is both molecular and fossil (Pappochelys) evidence for the origin of turtles among diapsid reptiles.
Procolophonidae is an extinct family of small, lizard-like parareptiles known from the Late Permian to Late Triassic that were distributed across Pangaea, having been reported from Europe, North America, China, South Africa, South America, Antarctica and Australia. The most primitive procolophonids were likely insectiovous or omnivorous, more derived members of the clade developed bicusped molars, and were likely herbivorous feeding on high fiber vegetation or durophagous omnivores. Many members of the group are noted for spines projecting from the quadratojugal bone of the skull, which likely served a defensive purpose as well as possibly also for display. At least some taxa were likely fossorial burrowers. While diverse during the Early and Middle Triassic, they had very low diversity during the Late Triassic, and were extinct by the beginning of the Jurassic.
Nyctiphruretus is an extinct genus of nyctiphruretid parareptile known from the Guadalupian series of European Russia.
Macroleter is an extinct genus of nycteroleterid parareptile which existed in Oklahoma and Russia during the upper Permian period. It was a quite generalized primitive reptile, in many ways resembling their amphibian ancestors. It was first named by paleontologists Tverdochlebova and Ivachnenko in 1984. According to classification by Michel Laurin and Robert R. Reisz, the genus is a parareptile, belonging to the same branch as Millerettidae, Procolophonidae and other generalized anapsid reptiles. The type species is Macroleter poezicus from Upper Permian of Russia.
Rhipaeosaurus is an extinct genus of nycteroleterid parareptile known from an articulated skeleton from the mid Middle Permian of European Russia. It contained a single species, Rhipaeosaurus tricuspidens. A bayesian analysis suggests that it is more closely related to pareiasaurs than to the other nycteroleterids, due to skull and tooth features. For this reason, "Nycteroleteridae" may be a grade rather than a clade, unless redefined to exclude Rhipaeosaurus.
Colobomycter is an extinct genus of lanthanosuchoid parareptile known from the Early Permian of Oklahoma.
Delorhynchus is an extinct genus of lanthanosuchoid parareptile known from the late Early Permian Garber Formation of Comanche County, Oklahoma. It contains three species: the type species D. priscus is based on a series of maxillae. The second species to be described, D. cifellii, is known from a larger number of well-preserved skulls and skeletal material. The third species, D. multidentatus, is based on a fragmentary skull with several rows of teeth on its jaw.
Acleistorhinidae is an extinct family of Late Carboniferous and Early Permian-aged parareptiles. Acleistorhinids are most diverse from the Richards Spur locality of the Early Permian of Oklahoma. Richards Spur acleistorhinids include Acleistorhinus, Colobomycter, and possibly Delorhynchus and Feeserpeton. Other taxa include Carbonodraco from the Late Carboniferous of Ohio and Karutia from the Early Permian of Brazil. Acleistorhinidae is commonly considered a subgroup of lanthanosuchoids, related to taxa such as Chalcosaurus, Lanthaniscus and Lanthanosuchus. However, a re-examination of parareptile phylogeny conducted by Cisneros et al. (2021) argued that lanthanosuchids were not closely related to acleistorhinids. The phylogenetic analysis conducted by these authors recovered acleistorhinids as the sister group of the clade Procolophonia, while lanthanosuchids were recovered within the procolophonian subgroup Pareiasauromorpha.
Acleistorhinus (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian of Oklahoma. It is notable for being the earliest known anapsid reptile yet discovered. The morphology of the lower temporal fenestra of the skull of Acleistorhinus bears a superficial resemblance to that seen in early synapsids, a result of convergent evolution. Only a single species, A. pteroticus, is known, and it is classified in the Family Acleistorhinidae, along with Colobomycter.
Owenetta is an extinct genus of owenettid procolophonian parareptile. Fossils have been found from the Beaufort Group in the Karoo Basin of South Africa. Although most procolophonians lived during the Triassic, Owenetta existed during the Wuchiapingian and Changhsingian stages of the Late Permian as well as the early Induan stage of the Early Triassic. It is the type genus of the family Owenettidae, and can be distinguished from other related taxa in that the posterior portion of the supratemporal bears a lateral notch and that the pineal foramen is surrounded by a depressed parietal surface on the skull table.
Microleter is an extinct genus of basal procolophonomorph parareptiles which lived in Oklahoma during the Early Permian period. The type and only known species is Microleter mckinzieorum. Microleter is one of several parareptile taxa described from the Richards Spur fissure fills, and can be characterized from its high tooth count, lacrimal/narial contact, short postfrontal, and slit-like temporal emargination edged by the postorbital, jugal, squamosal, and quadratojugal. Contrary to Australothyris, which had a similar phylogenetic position as a basal procolophonomorph, Microleter suggests that early parareptile evolution occurred in Laurasia and that multiple lineages developed openings or emarginations in the temporal region.
Lanthanosuchoidea is an extinct superfamily of ankyramorph parareptiles from the middle Pennsylvanian to the middle Guadalupian epoch of Europe, North America and Asia. It was named by the Russian paleontologist Ivachnenko in 1980, and it contains two families Acleistorhinidae and Lanthanosuchidae.
Nyctiphruretidae is an extinct family of hallucicranian parareptiles known from the late Early to the late Middle Permian of European Russia and south-central United States.
Lanthaniscus is an extinct genus of lanthanosuchoid ankyramorph parareptile known from the Guadalupian epoch of Eastern Europe, Russia. Lanthaniscus was first named by M. F. Ivakhnenko in 1980 and the type species is Lanthaniscus efremovi. L. efremovi was originally described on the basis of the holotype PIN 3706/9 from Peza-1 locality, Krasnoshchel' Formation, of Arkhangelsk. Various authors had assigned it to the family Lanthanosuchidae; however, Ivakhnenko, who described an additional specimen of L. efremovi in 2008, assigned Lanthaniscus to its own family, the Lanthaniscidae. The additional specimen PIN 4543/2, was collected from the same formation as the holotype, from the Nisogora locality, which is slightly younger in age.
Pareiasauromorpha is a group of parareptilian amniotes from the Permian. It includes genera found all over the world, with many genera from Asia and South Africa. The clade was first used as a group by Linda A. Tsuji in 2011, in order to group the family Nycteroleteridae (nycteroleters) and the superfamily Pareiasauroidea (pareiasaurs). Pareiasauromorpha is considered to be a monophyletic node, the sister group to procolophonoids.