Stereospondyls Temporal range: | |
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Life restoration of Siderops kehli , a chigutisaurid | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Temnospondyli |
Clade: | † Stereospondylomorpha |
Suborder: | † Stereospondyli |
Subgroups [1] | |
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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.
The group was first defined by Zittel (1888) [2] on the recognition of the distinctive vertebral anatomy of the best known stereospondyls of the time, such as Mastodonsaurus and Metoposaurus . The term 'stereospondylous' as a descriptor of vertebral anatomy was coined the following year by Fraas, [3] referring to a vertebral position consisting largely or entirely of the intercentrum in addition to the neural arch. While the name 'Stereospondyli' is derived from the stereospondylous vertebral condition, there is a diversity of vertebral morphologies among stereospondyls, including the diplospondylous ('tupilakosaurid') condition, [4] where the arch sits between the corresponding intercentrum and pleurocentrum, and the plagiosaurid condition, where a single large centrum ossification (identity unknown) is present, and the arch sits between subsequent vertebral positions. [5] The concept of Stereospondyli has thus undergone repeated and frequent revisions by different workers. Defining features include a tight articulation between the parasphenoid and the pterygoid and a stapedial groove. [1]
Stereospondyls first definitively appeared during the early Permian, as represented by fragmentary remains of a rhinesuchid from the Pedra de Fogo Formation of Brazil. [6] Rhinesuchids are one of the earliest groups of stereospondyls to appear in the fossil record and are predominantly a late Permian clade, with only one species, Broomistega putterilli, from the Early Triassic of South Africa. However, almost all other groups of stereospondyls are not known from any Paleozoic deposits, which remained dominated by non-stereospondyl stereospondylomorphs. The taxonomically unresolved Peltobatrachus pustulatus, which has historically been regarded as a stereospondyl, is also known from the late Permian of Tanzania. [7] Several more fragmentary records are known from horizons spanning the Permo-Triassic boundary in South America, such as the rhinesuchid-like Arachana nigra from Uruguay [8] and an indeterminate mastodonsaurid [9] from Uruguay.
Following the Permo-Triassic mass extinction, stereospondyls are abundantly represented in the fossil record, particularly from Russia, South Africa, and Australia. [10] This led Yates & Warren (2000) to propose that stereospondyls had sheltered in a high-latitude refugium that would have been somewhat shielded from the global effects of the extinction, and that they subsequently radiated from present-day Australia or Antarctica. [11] Recent discoveries of a diverse rhinesuchid community in South America alongside non-stereospondyl stereospondylomorphs have led to an alternative hypothesis for a radiation from western Gondwana in South America. [1] By the end of the Early Triassic, virtually all major clades of stereospondyls had appeared in the fossil record, although some were more geographically localized (e.g., lapillopsids, rhytidosteids) than those with cosmopolitan distributions (e.g., capitosauroids, trematosauroids).
Stereospondyls were the latest-surviving temnospondyl group. With the diversification of crocodile-like archosaurs and an extinction event at the end of the Triassic, most other temnospondyls disappeared. Chigutisaurid brachyopoids persisted into the Jurassic in Asia and Australia, [12] [13] [14] including Koolasuchus, the youngest known stereospondyl (late Early Cretaceous) from what is now Australia. [15] There is also sparse evidence for the persistence of some trematosauroids into the Jurassic of Asia. [16] If the recent hypothesis that Chinlestegophis, a Late Triassic stereospondyl from North America, is indeed a stem caecilian is correct, then stereospondyls would survive to the present day. [17]
Stereospondyls were particularly diverse during the Early Triassic, with small-bodied taxa such as lapillopsids and lydekkerinids that were likely more terrestrially capable present alongside larger taxa that would continue into the Middle Triassic, such as brachyopoids and trematosauroids. The vast majority of stereospondyls, particularly the large-bodied taxa, have been inferred to have been obligately aquatic based on features of the external anatomy such as a well-developed lateral line system, [18] poorly ossified postcranial skeleton, [19] and occasional preservation of proxies of external gills. [20] Many taxa also reflect adaptations for an aquatic lifestyle as evidence in bone histology, which is pachyostotic in many taxa, [21] although some studies suggest a greater terrestrial ability than historically inferred. [22] [23] [24] Most of the aquatic taxa resided in freshwater environments, but some trematosauroids in particular are thought to have been euryhaline based on their preservation in marine sediments with marine organisms. [25] [26] While stereospondyls are often compared to modern crocodilians, the presence of multiple temnospondyls in some environments and the range of morphologies across Stereospondyli indicates that at least some clades occupied drastically different ecological niches, such as benthic ambush predators. [27] Some groups, such as metoposaurids, are often recovered from large monotaxic bone beds interpreted as evidence of aggregation prior to mass death. [28] [29]
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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 large armour-like bony plates (osteoderms) that generally distinguish them from the modern soft-bodied lissamphibians.
Koolasuchus is an extinct genus of brachyopoid temnospondyl in the family Chigutisauridae. Fossils have been found from Victoria, Australia and date back 125-120 million years ago to Barremian-Aptian stages of the Early Cretaceous. Koolasuchus is the youngest known temnospondyl. It is known from several fragments of the skull and other bones such as vertebrae, ribs, and pectoral elements. The type species Koolasuchus cleelandi was named in 1997. K. cleelandi was adopted as the fossil emblem for the state of Victoria, Australia on 13 January 2022.
Trematosauria is one of two major groups of temnospondyl amphibians that survived the Permian-Triassic extinction event, the other being the Capitosauria. The trematosaurs were a diverse and important group that included many medium-sized to large forms that were semi-aquatic to fully aquatic. The group included long-snouted forms such as the trematosauroids and short, broad-headed forms such as the metoposaurs. Although most groups did not survive beyond the Triassic, one lineage, the brachyopoids, continued until the Cretaceous period. Trematosauria is defined as all stereospondyls more closely related to Trematosaurus than to Parotosuchus, a capitosaurian.
Metoposaurus meaning "front lizard" is an extinct genus of stereospondyl temnospondyls, known from the Late Triassic of Germany, Italy, Poland, and Portugal. This mostly aquatic animal possessed small, weak limbs, sharp teeth, and a large, flat head. This highly flattened creature mainly fed on fish, which it captured with its wide jaws lined with needle-like teeth. Many Metoposaurus mass graves have been found, probably from creatures that grouped together in drying pools during drought.
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.
Siderops is an extinct genus of chigutisaurid temnospondyl from Early Jurassic of Australia, containing the species S. kehli.
Dvinosaurus is an extinct genus of amphibious temnospondyls localized to regions of western and central Russia during the middle and late Permian, approximately 265-254 million years ago. Its discovery was first noted in 1921 by Russian paleontologist Vladimir Prokhorovich Amalitskii in a posthumously published paper that documents the findings of a site in Russia's Arkhangelsk District. Its name is derived from the proximity of this site to the Northern Dvina River.
Brachyopoidea is a superfamily of temnospondyls that lived during the Mesozoic. It contains the families Brachyopidae and Chigutisauridae. The earliest records of brachyopids are from the Lower Triassic in Australia. The latest-surviving member of the superfamily is the chigutisaurid Koolasuchus from the Early Cretaceous of Australia.
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.
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. In contrast to most other stereospondyls, lydekkerinids were relatively small-bodied. The type genus is Lydekkerina, the namesake of the family and the best-known lydekkerinid.
Mastodonsauridae is a family of capitosauroid temnospondyls. Fossils belonging to this family have been found in North America, Greenland, Europe, Asia, and Australia. The family Capitosauridae is synonymous with Mastodonsauridae.
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.
Calamops is an extinct genus of large temnospondyl amphibian known from the base of the Solebury Member of the Late Triassic Newark Supergroup of Pennsylvania, United States. Calamops was first named by Sinclair in 1917 and the type species is Calamops paludosus. It was usually thought to be a metoposaurid of questionable validity as its holotype and only known specimen, which comprises three pieces of a left mandibular ramus, had never been prepared. Following a preparation and casting of the specimen, Hans-Dieter Sues and Rainer R. Schoch found in 2013 Calamops to represent a valid taxon of trematosauroid temnospondyls that can be diagnosed by several autapomorphies. It represents one of the geologically youngest known long-snouted trematosaurs and the first record of these temnospondyls from the Late Triassic of North America. It is also the oldest known tetrapod fossil from the Triassic of the Newark basin.
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.
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.
Rileymillerus is an extinct genus of temnospondyl amphibian from the Late Triassic Post Quarry in the Dockum Group of Texas that was described by John Bolt and Sankar Chatterjee in 2000. The holotype, a nearly complete skull with articulated jaws, is housed at the Museum of Texas Tech University. The genus is named for Riley Miller, who allowed Chatterjee to work on the Post Quarry, and the species is named for the paleontologist John Cosgriff.
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.
This list of fossil amphibians described in 2018 is a list of new taxa of fossil amphibians that were described during the year 2018, as well as other significant discoveries and events related to amphibian paleontology that occurred in 2018.