Lydekkerinidae Temporal range: Triassic, | |
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Life restoration of Lyddekerina huxleyi | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Temnospondyli |
Suborder: | † Stereospondyli |
Family: | † Lydekkerinidae Watson, 1919 |
Synonyms | |
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Lydekkerinidae is a family of stereospondyl temnospondyls that lived in the Early Triassic period. During this time period, lydekkerinids were widely distributed, with putative remains reported from Russia, Greenland, India, South Africa, Madagascar, Australia, and Antarctica. [1] [2] [3] [4] [5] [6] In contrast to most other stereospondyls, lydekkerinids were relatively small-bodied (most with skulls less than 10 cm in length). The type genus is Lydekkerina , the namesake of the family and the best-known lydekkerinid.
The identification of features shared among lydekkerinids (synapomorphies) necessarily varies depending on which taxa are considered to belong to this group (see further in next section). In the most expansive concept, the family includes the eponymous Lydekkerina (and junior synonyms like 'Broomulus' and 'Limnoiketes'), Eolydekkerina from South Africa, Deltacephalus from Madagascar, Luzocephalus from Russia and Greenland (which includes the 'Aquiloniferus' of Bjerring (1999), [7] which is largely refuted by other workers), Chomatobatrachus from Australia, and indeterminate records from Antarctica and India (' Indobenthosuchus ' and ' Cryobatrachus '). However, most previous workers have not considered all of these taxa to be true lydekkerinds; [4] in particular, the affinities of non-South African taxa have been challenged.
For example, Schoch & Milner (2000) considered all nominal lydekkerinids to belong to this family and listed featured like longitudinally oval, unpaired anterior palatal openings, with a pointed posterior end; and a broad and laterally extensive postorbital and prefrontal as synapomorphies. [3] Jeannot et al. (2006) considered most nominal lydekkerinids to indeed be lydekkerinids except for Deltacephalus and the indeterminate records. [4] They list features like a step-shaped contact between the nasal and prefrontal; indentation of anterolateral margin of interpterygoid vacuity; and a straight cheek margin when viewed in occipital view as synapomorphies. Conversely, Hewison (2007) did not consider Chomatobatrachus or Luzocephalus to be lydekkerinids and therefore listed a different set of features, such as palatine lacking denticles, but having an elongated postero-mesial process extending behind the most anterior ectopterygoid tooth; ectopterygoid lacking denticles; and septomaxillary with an ornamented roofing portion and an unornamented, plate- like intranarial portion. [8] Many of Hewison's features (of which there are more than two dozen) are not synapomorphies but rather are part of a unique combination of features or are symplesiomorphies, and they are not mutually exclusive with the autapomorphies of other workers.
Most recently, it has also been suggested that another small-bodied Early Triassic clade, Lapillopsidae, might nest within Lydekkerinidae, rendering the latter paraphyletic. [9] If so, this would introduce further uncertainty regarding diagnostic features of Lydekkerinidae.
Lydekkerinids are usually classified as basal stereospondyls. Schoch and Milner (2000) placed them in the clade Rhytidostea along with brachyopoids and rhytidosteids, [3] but this broader grouping is not widely employed today given the higher nesting position of brachyopoids and the uncertain position and monophyly of rhytidosteids. [10] [11] [12] [13] Lydekkerinidae was also sometimes historically placed within the largely defunct Rhinesuchoidea [14] [15] or within the still valid Capitosauroidea or Capitosauria, [2] but this too has not been supported by more recent work. Most phylogenetic studies that sampled only certain nominal lydekkerinids within a larger temnospondyl sample have found lydekkerinids to be paraphyletic or polyphyletic. [16] [12] [13] [17] [9] Furthermore, no complete phylogenetic analysis of all nominal lydekkerinids that would support monophyly in at least a restricted framework has been formally published. [18] Luzocephalus, which would be the largest of the lydekkerinids with a skull length over 15 cm, [19] has often been found to be more closely related to a family called Trematosauridae, such as in the study by Yates and Warren (2000). Chomatobatrachus has also been frequently dissociated from other nominal lydekkerinids. [2] [8] Below is a cladogram from Yates and Warren (2000) showing the polyphyly: [10]
The phylogenetic analysis of Damiani (2001) resulted in a monophyletic Lydekkerinidae, although it was only weakly supported and included what is now recognized as the small-bodied Early Triassic capitosaur Edingerella madagascariensis . Below is a cladogram from that analysis: [20]
Temnospondyli or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carboniferous, Permian and Triassic periods, with fossils being found on every continent. A few species continued into the Jurassic and Early Cretaceous periods, but all had gone extinct by the Late Cretaceous. During about 210 million years of evolutionary history, they adapted to a wide range of habitats, including freshwater, terrestrial, and even coastal marine environments. Their life history is well understood, with fossils known from the larval stage, metamorphosis and maturity. Most temnospondyls were semiaquatic, although some were almost fully terrestrial, returning to the water only to breed. These temnospondyls were some of the first vertebrates fully adapted to life on land. Although temnospondyls are amphibians, many had characteristics such as scales and armour-like bony plates that distinguish them from the modern soft-bodied lissamphibians.
Dvinosaurs are one of several new clades of temnospondyls named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic temnospondyls, and are known from the Late Carboniferous to the Early Triassic, being most common in the Permian period. Their distinguishing characteristics are a reduction of the otic notch; the loss of a flange on the rear side of the pterygoid; and 28 or more presacral vertebrae.
The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecological niche to modern crocodilians prior to the diversification of pseudosuchian archosaurs.
Eryosuchus is an extinct genus of capitosauroid temnospondyl from the Middle Triassic of northern Russia. It was a very large predator: the largest specimen known could reach up to 3.5 m (11.5 ft) in length, with a skull over 1 m long.
Cherninia is an extinct genus of mastodonsaurid temnospondyl. The type species, Cherninia denwai, is known from the Denwa Formation of India. It is based on a massive skull, ISI A 54, which was originally considered a species of Parotosuchus in 1998 before being given its own genus in 2001.
Rhinesuchidae is a family of tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians are probably descended from. Rhinesuchids can be differentiated from other temnospondyls by details of their skulls, most notably the interior structure of their otic notches at the back of the skull. They were among the earliest-diverging members of the Stereospondyli, a subgroup of temnospondyls with flat heads and aquatic habits. Although more advanced stereospondyls evolved to reach worldwide distribution in the Triassic period, rhinesuchids primarily lived in the high-latitude environments of Gondwana during the Guadalupian and Lopingian epochs of the Permian. The taxonomy of this family has been convoluted, with more than twenty species having been named in the past; a 2017 review recognized only eight of them to be valid. While several purported members of this group have been reported to have lived in the Triassic period, most are either dubious or do not belong to the group. However, at least one valid genus of rhinesuchid is known from the early Triassic, a small member known as Broomistega. The most recent formal definition of Rhinesuchidae, advocated by Mariscano et al. (2017) is that of a stem-based clade containing all taxa more closely related to Rhinesuchus whaitsi than to Lydekkerina huxleyi or Peltobatrachus pustulatus. A similar alternate definition is that Rhinesuchidae is a stem-based clade containing all taxa more closely related to Uranocentrodon senekalensis than to Lydekkerina huxleyi, Trematosaurus brauni, or Mastodonsaurus giganteus.
Limnarchia is a clade of temnospondyls. It includes the mostly Carboniferous-Permian age Dvinosauria and the mostly Permian-Triassic age Stereospondylomorpha. The clade was named in a 2000 phylogenetic analysis of stereospondyls and their relatives. Limnarchia means "lake rulers" in Greek, in reference to their aquatic lifestyles and long existence over a span of approximately 200 million years from the Late Carboniferous to the Early Cretaceous. In phylogenetic terms, Limnarchia is a stem-based taxon including all temnospondyls more closely related to Parotosuchus than to Eryops. It is the sister group of the clade Euskelia, which is all temnospondyls more closely related to Eryops than to Parotosuchus. Limnarchians represent an evolutionary radiation of temnospondyls into aquatic environments, while euskelians represent a radiation into terrestrial environments. While many euskelians were adapted to life on land with strong limbs and bony scutes, most limnarchians were better adapted for the water with poorly developed limbs and lateral line sensory systems in their skulls.
Acroplous is an extinct genus of dvinosaurian Temnospondyli within the family Eobrachyopidae.
Bukobaja is an extinct genus of mastodonsaurid temnospondyl from the middle Triassic of Russia. It contains a single species, Bukobaja enigmatica. Bukobaja mainly occurs in the Bukobay Svita as part of the Ladinian?-age "Mastodonsaurus fauna", a section of Russian Triassic biostratigraphy characterized by "Mastodonsaurus" torvus. It was also present in the underlying Donguz Svita. Bukobaja appears to be a valid genus similar to, yet distinct from, Mastodonsaurus. Despite appearing to possess several unique features, Bukobaja is still known from very few remains. This has led to difficulties in determining its relations more precisely than "Mastodonsauridae incertae sedis". It has also been compared to trematosaurids.
Lydekkerina is an extinct genus of stereospondyl temnospondyl. It is the type genus of the family Lydekkerinidae. Fossils have been collected from Early Triassic deposits in South Africa and Australia. The type species is L. huxleyi, first described in 1889. While most other stereospondyls were semiaquatic, Lydekkerina was exclusively terrestrial.
Luzocephalus is an extinct genus of temnospondyl from the Early Triassic of Russia and Greenland. It is usually regarded as a member of the family Lydekkerinidae, although it has also been placed in the family Trematosauridae.
Lapillopsis is an extinct genus of stereospondyl temnospondyl within the family Lapillopsidae. Fossils belonging to the genus have been found in the Arcadia Formation of Queensland, Australia.
Keratobrachyops is an extinct genus of trematosaurian temnospondyl found in the Arcadia Formation of Queensland, Australia. It had been thought to be a basal chigutisaurid but is now thought to be a basal brachyopomorph closely related to the genus Bothriceps, and may even be a synonym of it.
Rhineceps is an extinct genus of temnospondyl amphibian in the family Rhinesuchidae. Rhineceps was found in Northern Malawi in Southern Africa known only from its type species R. nyasaensis. Rhineceps was a late Permian semi-aquatic carnivore that lived in streams, rivers, lakes or lagoons. Rhineceps is an early divergent Stereopondyl within the family Rhinesuchidae, which only existed in the late Permian (Lopingian) and failed to survive the Permian-Triassic extinction unlike other stereospondyl families.
Wellesaurus is an extinct genus of mastodonsauroid temnospondyl. They were amphibious carnivores that lived in freshwater environments.
Micropholis is an extinct genus of dissorophoid temnospondyl. Fossils have been found from the Lystrosaurus Assemblage Zone of the Karoo Basin in South Africa and are dated to the Induan. Fossils have also been found from the lower Fremouw of Antarctica.Micropholis is the only post-Permian dissorophoid and the only dissorophoid in what is presently the southern hemisphere and what would have been termed Gondwana during the amalgamation of Pangea.
Capitosauria is an extinct group of large temnospondyl amphibians with simplified stereospondyl vertebrae. Mainly living as piscivores in lakes and rivers, the Capitosauria and its sister taxon Trematosauria were the only major labyrinthodonts that existed during the Mesozoic in ecological niches broadly similar to those of modern crocodiles, and some grew to very large sizes. At 6 meters in length, the Mid-Triassic Mastodonsaurus giganteus is not only thought to have been the largest capitosaur, but possibly also the largest amphibian to have lived. The latest known remains are from the Rhaetian of Germany and are referred to Cyclotosaurus.
Rhytidostea is a clade of stereospondyl temnospondyls. It was erected in 2000 to include several temnospondyl groups distinct from the "higher" group of capitosaurs, including lydekkerinids, brachyopoids, and rhytidosteids. Rhytidosteans first appeared in the Permian period and underwent an evolutionary radiation during the Induan stage of the Early Triassic. Along with capitosaurs, rhytidosteans comprise much of the larger suborder Stereospondyli. Rhytidostea has often been considered the sister group of the clade Capitosauria, but has been placed in various other phylogenetic positions. In many studies, members of Rhytidostea are split, with lydekkerinids having a more basal position among stereospondyls while rhytidosteids and brachyopoids form a group placed among the more derived trematosaurian stereospondyls.
Manubrantlia was a genus of lapillopsid temnospondyls from the Early Triassic Panchet Formation of India. This genus is only known from a single holotype left jaw, given the designation ISI A 57. Despite the paucity of remains, the jaw is still identifiable as belonging to a relative of Lapillopsis. For example, all three of its coronoid bones possessed teeth, the articular bone is partially visible in lateral (outer) view, and its postsplenial does not contact the posterior meckelian foramen. However, the jaw also possesses certain unique features which justify the erection of a new genus separate from Lapillopsis. For example, the mandible is twice the size of any jaws referred to other lapillopsids. The most notable unique feature is an enlarged "pump-handle" shaped arcadian process at the back of the jaw. This structure is responsible for the generic name of this genus, as "Manubrantlia" translates from Latin to the English expression "pump-handle". The type and only known species of this genus is Manubrantlia khaki. The specific name refers to the greenish-brown mudstones of the Panchet Formation, with a color that had been described as "khaki" by the first British geologists who studied the formation.
Latiscopus disjunctus is a small Late Triassic temnospondyl collected in 1940 by a Works Projects Administration crew working near Otis Chalk, Texas that was described by John Wilson in 1948.
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