| Khmeria Temporal range: | |
|---|---|
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Subphylum: | Tunicata |
| Class: | Ascidiacea |
| Order: | † Khmeriamorpha |
| Family: | † Khmeriidae |
| Genus: | † Khmeria Mansuy, 1914 |
| Species | |
| |
Khmeria is a fossil genus of controversial affiliation, known from the Middle Permian to Late Triassic. Originally described as putative anthozoan corals, it is now believed to be a genus of ascidian tunicates. [1]
Khmeria problematica was first described in 1914 by Henri Mansuy from Productus -bearing limestone in Cambodia. It was stated to be similar in morphology to operculate rugose corals, sharing features such as a curved, wrinkled body shape with a thin pedicle. However, such an affiliation was rejected due to the morphological variability exhibited by Khmeria and its less defined internal structure compared to these corals, and Khmeria was left unclassified. [2] In 1932, specimens of K. problematica found in Nagato, Japan, provided visible evidence of internal deposits, including tabulae. [3]
The genus Osium was described in 1930 from specimens discovered in the Sosio Valley of Sicily. It was later found to be a subjective junior synonym of Khmeria, and its only described species was reclassified as Khmeria importans. [4] The name has since been argued to be invalid due to the poor quality of the description and the lack of surviving material. [5]
The eponymous family Khmeriidae was erected in 1954 inside the order Rugosa. [6] The rugosan affiliation of the genus was supported by the presence of tabulae and vesiculae, [5] and a relationship with "cystiphylloid" corals was suggested. [6]
In 2018, Khmeria minima was described from the Upper Triassic San Cassiano Formation in central Italy, alongside the putatively related Zardinisoma . At the same time, specimens of Khmeria problematica exhibiting colonial growth were reclassified as the separate species Khmeria stolonifera. [7] Khmeria importans was reclassified as a synonym of K. problematica. [5]
The same 2018 study argued for a tunicate identity for Khmeria, and established the order Khmeriamorpha as part of the class Ascidiacea. [8] The structures previously identified as vesiculae were stated to be misidentified cement crystals. [5] A rugosan affiliation was deemed unlikely based on the lack of septa and vesiculae, [9] as well as the presence of multiple opercula and an aragonitic mineralogy in some species. [10]
Khmeria is a bilaterally symmetric, sessile organism. It has a bivalved shell, made of a conical body covered by an operculum. Known specimens vary from 4 to 35 millimetres (0.16 to 1.38 in) in height and 4 to 17 millimetres (0.16 to 0.67 in) in width. [11]
The shell grows along the margins, displaying concentric growth lines along both the body and operculum. The margins are covered in indented grooves, called crenellae by analogy with the similar structures present in crinoid ossicles. Internal compartments, or tabulae, are rare, but may form through endoskeletal biomineralization. [11]
Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.
Helicoprion is an extinct genus of shark-like eugeneodont fish. Almost all fossil specimens are of spirally arranged clusters of the individuals' teeth, called "tooth whorls", which in life were embedded in the lower jaw. As with most extinct cartilaginous fish, the skeleton is mostly unknown. Fossils of Helicoprion are known from a 20 million year timespan during the Permian period from the Artinskian stage of the Cisuralian to the Roadian stage of the Guadalupian. The closest living relatives of Helicoprion are the chimaeras, though their relationship is very distant. The unusual tooth arrangement is thought to have been an adaption for feeding on soft bodied prey, and may have functioned as a deshelling mechanism for hard bodied cephalopods such as nautiloids and ammonoids. In 2013, systematic revision of Helicoprion via morphometric analysis of the tooth whorls found only H. davisii, H. bessonowi and H. ergassaminon to be valid, with some of the larger tooth whorls being outliers.
Velocipes is a saurischian dinosaur genus from the Late Triassic that may have been a theropod; its fossils were found in the Norian-age Lissauer Breccia of southern Poland.
Ericiolacerta is an extinct genus of small therocephalian therapsids from the early Triassic of South Africa and Antarctica. Ericiolacerta, meaning "hedgehog lizard", was named by D.M.S. Watson in 1931. The species E. parva is known from the holotype specimen which consists of a nearly complete skeleton found in the Lystrosaurus Assemblage Zone within the Katberg Formation of the Beaufort Group in South Africa, and from a partial jaw found in the Lower Triassic Fremouw Formation in Antarctica. Ericiolacerta was around 20 centimetres (7.9 in) in length, with long limbs and relatively small teeth. It probably ate insects and other small invertebrates. The therocephalians – therapsids with mammal-like heads – were abundant in Permian times, but only a few made it into the Triassic. Ericiolacerta was one of those. It is possible that they gave rise to the cynodonts, the only therapsid group to survive into post-Triassic times. Cynodonts gave rise to mammals.
Diplocaulus is an extinct genus of lepospondyl amphibians which lived from the Late Carboniferous to the Late Permian of North America and Africa. Diplocaulus is by far the largest and best-known of the lepospondyls, characterized by a distinctive boomerang-shaped skull. Remains attributed to Diplocaulus have been found from the Late Permian of Morocco and represent the youngest-known occurrence of a lepospondyl.
Proterosuchus is an extinct genus of archosauriform reptiles that lived during the Early Triassic. It contains three valid species: the type species P. fergusi and the referred species P. alexanderi and P. goweri. All three species lived in what is now South Africa. The genus was named in 1903 by the South African paleontologist Robert Broom. The genus Chasmatosaurus is a junior synonym of Proterosuchus.
Sauroctonus is an extinct genus of gorgonopsian therapsids who lived during the end of the Middle Permian in what is now European Russia. The first fossils, discovered in Tatarstan, were initially believed to belong to a new species of the South African genus Arctognathus. The taxon was designated as such until 1940, when it was assigned to the genus Inostrancevia by Ivan Yefremov, before being definitively classified in a separate genus erected by Alexey Bystrow. The most complete, known fossils of S. progressus include cranial and postcranial elements, currently all recorded from Tatarstan. These elements show that the animal was a mid-sized gorgonopsian.
Campylocephalus is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of Campylocephalus have been discovered in deposits ranging from the Carboniferous period in the Czech Republic to the Permian period of Russia. The generic name is composed of the Greek words καμπύλος (kampýlos), meaning "curved", and κεφαλή (kephalē), meaning "head".
Venyukovia is an extinct genus of venyukovioid therapsid, a basal anomodont from the Middle Permian of Russia. The type and sole species, V. prima, is known only by a partial lower jaw with teeth. Venyukovia has often been incorrectly spelt as 'Venjukovia' in English literature. This stems from a spelling error made by Russian palaeontologist Ivan Efremov in 1940, who mistakenly replaced the 'y' with a 'j', which subsequently permeated through therapsid literature before the mistake was caught and corrected. Venyukovia is the namesake for the Venyukovioidea, a group of small Russian basal anomodonts also including the closely related Otsheria, Suminia, Parasuminia and Ulemica, although it itself is also one of the poorest known. Like other venyukovioids, it had large projecting incisor-like teeth at the front and lacked canines, although the remaining teeth are simple compared to some other venyukovioids, but may resemble those of Otsheria.
Pelanomodon is an extinct genus of dicynodont therapsids that lived in the Late Permian period. Fossil evidence of this genus is principally found in the Karoo Basin of South Africa, in the Dicynodon Assemblage Zone. Lack of fossil record after the Late Permian epoch suggests that Pelanomodon fell victim to the Permian-Triassic extinction event.
Akidnognathidae is an extinct family of therocephalian therapsids from the Late Permian and Early Triassic of South Africa, Russia and China. The family includes many large-bodied therocephalians that were probably carnivorous, including Moschorhinus and Olivierosuchus. One akidnognathid, Euchambersia, may even have been venomous. Akidnognathids have robust skulls with a pair of large caniniform teeth in their upper jaws. The family is morphologically intermediate between the more basal therocephalian group Scylacosauridae and the more derived group Baurioidea.
Homodontosaurus is an extinct genus of therocephalian therapsids from the Late Permian of South Africa. The type species Homodontosaurus kitchingi was named by South African paleontologist Robert Broom in 1949. Broom based his description on a small skull found in the Cistecephalus Assemblage Zone near Graaff-Reinet. The skull is very small, at about 55 millimetres (2.2 in) long and 20 millimetres (0.79 in) wide. Homodontosaurus has large eye sockets and an elongated snout. The lower jaw is long, thin, and curved. Numerous small teeth line the upper jaw and are long, pointed, and round in cross-section.
Paleontology in Michigan refers to paleontological research occurring within or conducted by people from the U.S. state of Michigan. During the Precambrian, the Upper Peninsula was home to filamentous algae. The remains it left behind are among the oldest known fossils in the world. During the early part of the Paleozoic Michigan was covered by a shallow tropical sea which was home to a rich invertebrate fauna including brachiopods, corals, crinoids, and trilobites. Primitive armored fishes and sharks were also present. Swamps covered the state during the Carboniferous. There are little to no sedimentary deposits in the state for an interval spanning from the Permian to the end of the Neogene. Deposition resumed as glaciers transformed the state's landscape during the Pleistocene. Michigan was home to large mammals like mammoths and mastodons at that time. The Holocene American mastodon, Mammut americanum, is the Michigan state fossil. The Petoskey stone, which is made of fossil coral, is the state stone of Michigan.
The Gnishik Formation is a geologic formation in Armenia, Azerbaijan and Iran. It preserves fossils dated to the Wordian age of the Permian period.
This timeline of Permian research is a chronological listing of events in the history of geology and paleontology focused on the study of earth during the span of time lasting from 298.9–252.17 million years ago and the legacies of this period in the rock and fossil records.
Adelophthalmidae is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily Adelophthalmoidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina.
Pectodens is an extinct genus of archosauromorph reptile which lived during the Middle Triassic in China. The type and only species of the genus is P. zhenyuensis, named by Chun Li and colleagues in 2017. It was a member of the Archosauromorpha, specifically part of the unnatural grouping Protorosauria. However, an unusual combination of traits similar and dissimilar to other protorosaurs initially led to confusion over its evolutionary relationships. In 2021, it was placed in a newly-established group, Dinocephalosauridae, along with its closest relative Dinocephalosaurus.
Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 2018.
Counillonia is an extinct genus of dicynodont therapsid from the area of Luang Prabang in Laos, Southeast Asia that lived at around the time of the Permian-Triassic boundary and possibly dates to the earliest Early Triassic. Its type and only known species is C. superoculis. Counillonia was related to the Triassic dicynodonts such as Lystrosaurus and the Kannemeyeriiformes that survived the Permian mass extinction, but it was more closely related to the Permian genus Dicynodon than to either of these lineages. Counillonia may then possibly represent another line of dicynodonts that survived the Permian mass extinction into the Triassic period, depending on its age. The discovery of Counillonia in Laos and its unexpected evolutionary relationships hint at the less well understood geographies of dicynodont diversity across the Permo-Triassic boundary outside of well explored regions like the Karoo Basin in South Africa.
Repelinosaurus is an extinct genus of dicynodont from the Purple Claystone Formation of Luang Prabang in Laos, Southeast Asia that lived at around the time of the Permian-Triassic boundary and possibly dates to the earliest Early Triassic. Its type and only known species is R. robustus. Repelinosaurus was originally described as the earliest known kannemeyeriiform dicynodont, supporting the idea of a more rapid radiation of the Triassic kannemeyeriiform dicynodonts during the Early Triassic following the Permian mass extinction. However, it may alternatively be more closely related to the Permian Dicynodon. The discovery of a potential early kannemeyeriiform in an understudied locality like Laos highlights the importance of such places in dicynodont research, which has been largely focused on historically important localities such as the Karoo Basin of South Africa.
