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108-492: The Permian witnessed the diversification of the two groups of amniotes , the synapsids and the sauropsids ( reptiles ). The world at the time was dominated by the supercontinent Pangaea , which had formed due to the collision of Euramerica and Gondwana during the Carboniferous. Pangaea was surrounded by the superocean Panthalassa . The Carboniferous rainforest collapse left behind vast regions of desert within

216-463: A crown group definition, Amniota has a slightly different content than the biological amniotes as defined by an apomorphy. Though traditionally considered reptiliomorphs, some recent research has recovered diadectomorphs as the sister group to Synapsida within Amniota, based on inner ear anatomy. The cladogram presented here illustrates the phylogeny (family tree) of amniotes, and follows

324-494: A stratigraphic set of smaller units called stages , each formed during corresponding time intervals called ages. Stages can be defined globally or regionally. For global stratigraphic correlation, the International Commission on Stratigraphy (ICS) ratify global stages based on a Global Boundary Stratotype Section and Point (GSSP) from a single formation (a stratotype ) identifying the lower boundary of

432-482: A (typically) terrestrial form with limbs and a thick stratified epithelium (rather than first entering a feeding larval tadpole stage followed by metamorphosis , as amphibians do). In amniotes, the transition from a two-layered periderm to a cornified epithelium is triggered by thyroid hormone during embryonic development, rather than by metamorphosis. The unique embryonic features of amniotes may reflect specializations for eggs to survive drier environments; or

540-493: A calcified shell, were not essential and probably evolved later. It has been suggested that shelled terrestrial eggs without extraembryonic membranes could still not have been more than about 1 cm (0.4-inch) in diameter because of diffusion problems, like the inability to get rid of carbon dioxide if the egg was larger. The combination of small eggs and the absence of a larval stage, where posthatching growth occurs in anamniotic tetrapods before turning into juveniles, would limit

648-809: A chronostratigraphic term as a substage of the Guadalupian Stage. The GSSP for the base of the Wordian is located in Guadalupe Pass, Texas, within the sediments of the Getaway Limestone Member of the Cherry Canyon Formation , which was ratified in 2001. The base of the Wordian is defined by the first appearance of the conodont Jinogondolella aserrata. The Capitanian is named after the Capitan Reef in

756-687: A defined GSSP. Recent proposals have suggested the appearance of Neostreptognathodus pnevi as the lower boundary. The Guadalupian Series is named after the Guadalupe Mountains in Texas and New Mexico, where extensive marine sequences of this age are exposed. It was named by George Herbert Girty in 1902. The Roadian was named in 1968 in reference to the Road Canyon Member of the Word Formation in Texas. The GSSP for

864-611: A geologic formation goes back to the beginnings of modern scientific geology. The term was used by Abraham Gottlob Werner in his theory of the origin of the Earth, which was developed over the period from 1774 to his death in 1817. The concept became increasingly formalized over time and is now codified in such works as the North American Stratigraphic Code and its counterparts in other regions. Geologic maps showing where various formations are exposed at

972-468: A hypersaline epicontinental sea , existed in what is now northwestern Europe. Large continental landmass interiors experience climates with extreme variations of heat and cold (" continental climate ") and monsoon conditions with highly seasonal rainfall patterns. Deserts seem to have been widespread on Pangaea. Such dry conditions favored gymnosperms , plants with seeds enclosed in a protective cover, over plants such as ferns that disperse spores in

1080-588: A lower canopy consisting of Marattialean tree ferns, and Noeggerathiales. Early conifers appeared in the Late Carboniferous, represented by primitive walchian conifers, but were replaced with more derived voltzialeans during the Permian. Permian conifers were very similar morphologically to their modern counterparts, and were adapted to stressed dry or seasonally dry climatic conditions. The increasing aridity, especially at low latitudes, facilitated

1188-650: A major group during the Early-Mid Permian, but declined during the Late Permian. Members of the order Prolecanitida were less diverse. The Ceratitida originated from the family Daraelitidae within Prolecanitida during the mid-Permian, and extensively diversified during the Late Permian. Only three families of trilobite are known from the Permian, Proetidae , Brachymetopidae and Phillipsiidae . Diversity, origination and extinction rates during

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1296-441: A massive desert covering the interior of Pangaea . The warm zone spread in the northern hemisphere, where extensive dry desert appeared. The rocks formed at that time were stained red by iron oxides, the result of intense heating by the sun of a surface devoid of vegetation cover. A number of older types of plants and animals died out or became marginal elements. The Permian began with the Carboniferous flora still flourishing. About

1404-627: A megamonsoon would occur given the continental arrangement of the Permian. The aforementioned increasing equatorial aridity was likely driven by the development and intensification of this Pangaean megamonsoon. Permian marine deposits are rich in fossil mollusks , brachiopods , and echinoderms . Brachiopods were highly diverse during the Permian. The extinct order Productida was the predominant group of Permian brachiopods, accounting for up to about half of all Permian brachiopod genera. Brachiopods also served as important ecosystem engineers in Permian reef complexes. Amongst ammonoids , Goniatitida were

1512-542: A newly designated formation could not be named the Kaibab Formation, since the Kaibab Limestone is already established as a formation name. The first use of a name has precedence over all others, as does the first name applied to a particular formation. As with other stratigraphic units, the formal designation of a formation includes a stratotype which is usually a type section . A type section

1620-628: A particularly great magnitude. In the late Kungurian, cooling resumed, resulting in a cool glacial interval that lasted into the early Capitanian, though average temperatures were still much higher than during the beginning of the Cisuralian. Another cool period began around the middle Capitanian. This cool period, lasting for 3-4 Myr, was known as the Kamura Event. It was interrupted by the Emeishan Thermal Excursion in

1728-480: A peak of diversity in the Cisuralian, with a substantial decline during the Guadalupian-Lopingian following Olson's extinction, with the family diversity dropping below Carboniferous levels. Embolomeres , a group of aquatic crocodile-like limbed vertebrates that are reptilliomorphs under some phylogenies. They previously had their last records in the Cisuralian, are now known to have persisted into

1836-617: A permanent natural or artificial feature of the geographic area in which they were first described. The name consists of the geographic name plus either "Formation" or a descriptive name. Examples include the Morrison Formation , named for the town of Morrison, Colorado , and the Kaibab Limestone , named after the Kaibab Plateau of Arizona. The names must not duplicate previous formation names, so, for example,

1944-417: A robust, air-breathing, respiratory system , allow amniotes to live on land as true terrestrial animals . Amniotes have the ability to procreate without water bodies . Because the amnion and the fluid it secretes shields the embryo from environmental fluctuations, amniotes can reproduce on dry land by either laying shelled eggs (reptiles, birds and monotremes ) or nurturing fertilized eggs within

2052-671: A series of equatorial islands in the Paleo-Tethys Ocean that later would become South China . The Permian saw the radiation of many important conifer groups, including the ancestors of many present-day families. Rich forests were present in many areas, with a diverse mix of plant groups. The southern continent saw extensive seed fern forests of the Glossopteris flora. Oxygen levels were probably high there. The ginkgos and cycads also appeared during this period. Insects, which had first appeared and become abundant during

2160-1713: A simplified version of the relationships found by Laurin & Reisz (1995), with the exception of turtles, which more recent morphological and molecular phylogenetic studies placed firmly within diapsids . The cladogram covers the group as defined under Gauthier's definition. † Diadectomorpha [REDACTED] Synapsida (mammals and their extinct relatives) [REDACTED] † Mesosauridae [REDACTED] † Millerettidae [REDACTED] † Pareiasauria [REDACTED] † Procolophonoidea [REDACTED] † Captorhinidae [REDACTED] † Protorothyrididae [REDACTED] Diapsida (lizards, snakes, turtles , crocodiles , dinosaurs , birds, etc.) [REDACTED] Following studies in 2022 and 2023, with Drepanosauromorpha placed sister to Weigeltisauridae ( Coelurosauravus ) in Avicephala based on Senter (2004): † Seymouriamorpha [REDACTED] † Diadectomorpha [REDACTED] † Araeoscelida [REDACTED] † Captorhinidae [REDACTED] † Protorothyris [REDACTED] † Vaughnictis [REDACTED] † Eothyris [REDACTED] † Caseidae [REDACTED] † Oedaleops [REDACTED] † Varanopsidae [REDACTED] † Ophiacodontidae [REDACTED] † Edaphosauridae [REDACTED] † Haptodus [REDACTED] † Sphenacodontidae [REDACTED] Therapsida [REDACTED] [REDACTED] [REDACTED] † Acleistorhinidae [REDACTED] Formation (geology) A geological formation , or simply formation ,

2268-463: A single supercontinent known as Pangaea , with the microcontinental terranes of Cathaysia to the east. Pangaea straddled the equator and extended toward the poles, with a corresponding effect on ocean currents in the single great ocean (" Panthalassa ", the "universal sea"), and the Paleo-Tethys Ocean , a large ocean that existed between Asia and Gondwana. The Cimmeria continent rifted away from Gondwana and drifted north to Laurasia , causing

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2376-634: A vote by the Subcommission on Permian Stratigraphy of the ICS adopted the Lopingian as an international standard chronostratigraphic unit. The Wuchiapinginan and Changhsingian were first introduced in 1962, by J. Z. Sheng as the "Wuchiaping Formation" and "Changhsing Formation" within the Lopingian series. The GSSP for the base of the Wuchiapingian is located at Penglaitan, Guangxi , China and

2484-531: A wetter environment. The first modern trees ( conifers , ginkgos and cycads ) appeared in the Permian. Three general areas are especially noted for their extensive Permian deposits—the Ural Mountains (where Perm itself is located), China, and the southwest of North America, including the Texas red beds. The Permian Basin in the U.S. states of Texas and New Mexico is so named because it has one of

2592-449: Is not a valid lithological basis for defining a formation. The contrast in lithology between formations required to justify their establishment varies with the complexity of the geology of a region. Formations must be able to be delineated at the scale of geologic mapping normally practiced in the region; the thickness of formations may range from less than a meter to several thousand meters. Geologic formations are typically named after

2700-412: Is a body of rock having a consistent set of physical characteristics ( lithology ) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exposed in a geographical region (the stratigraphic column ). It is the fundamental unit of lithostratigraphy , the study of strata or rock layers. A formation must be large enough that it can be mapped at

2808-637: Is a global hiatus in the terrestrial fossil record during the late Kungurian and early Roadian , referred to as "Olson's Gap" that obscures the nature of the transition. Other proposals have suggested that the North American and Russian records overlap, with the latest terrestrial North American deposition occurring during the Roadian, suggesting that there was an extinction event, dubbed " Olson's Extinction ". The Middle Permian faunas of South Africa and Russia are dominated by therapsids, most abundantly by

2916-560: Is central to the geologic discipline of stratigraphy , and the formation is the fundamental unit of stratigraphy. Formations may be combined into groups of strata or divided into members . Members differ from formations in that they need not be mappable at the same scale as formations, though they must be lithologically distinctive where present. The definition and recognition of formations allow geologists to correlate geologic strata across wide distances between outcrops and exposures of rock strata . Formations were at first described as

3024-590: Is defined by the first appearance of Streptognathodus postfusus . The Sakmarian is named in reference to the Sakmara River in the southern Urals, and was coined by Alexander Karpinsky in 1874. The GSSP for the base of the Sakmarian is located at the Usolka section in the southern Urals, which was ratified in 2018. The GSSP is defined by the first appearance of Sweetognathus binodosus . The Artinskian

3132-568: Is facilitated by their astragalus. Basal amniotes resembled small lizards and evolved from semiaquatic reptiliomorphs during the Carboniferous period. After the Carboniferous rainforest collapse , amniotes spread around Earth's land and became the dominant land vertebrates. They almost immediately diverged into two groups, namely the sauropsids (including all reptiles and birds ) and synapsids (including mammals and extinct ancestors like " pelycosaurs " and therapsids ). Among

3240-455: Is ideally a good exposure of the formation that shows its entire thickness. If the formation is nowhere entirely exposed, or if it shows considerably lateral variation, additional reference sections may be defined. Long-established formations dating to before the modern codification of stratigraphy, or which lack tabular form (such as volcanic formations), may substitute a type locality for a type section as their stratotype. The geologist defining

3348-450: Is presented in simplified form below. With the advent of cladistics, other researchers have attempted to establish new classes, based on phylogeny , but disregarding the physiological and anatomical unity of the groups. Unlike Benton, for example, Jacques Gauthier and colleagues forwarded a definition of Amniota in 1988 as "the most recent common ancestor of extant mammals and reptiles, and all its descendants". As Gauthier makes use of

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3456-512: Is thought to have existed during the Early Permian. Though the fossil record is fragmentary, lungfish appear to have undergone an evolutionary diversification and size increase in freshwater habitats during the Early Permian, but subsequently declined during the middle and late Permian. Conodonts experienced their lowest diversity of their entire evolutionary history during the Permian. Permian chondrichthyan faunas are poorly known. Members of

3564-861: The Capitanian Stage of the Permian was marked by the major Capitanian mass extinction event , associated with the eruption of the Emeishan Traps . The Permian (along with the Paleozoic) ended with the Permian–Triassic extinction event (colloquially known as the Great Dying), the largest mass extinction in Earth's history (which is the last of the three or four crises that occurred in the Permian), in which nearly 81% of marine species and 70% of terrestrial species died out, associated with

3672-481: The Geological Society of London , after extensive Russian explorations undertaken with Édouard de Verneuil in the vicinity of the Ural Mountains in the years 1840 and 1841. Murchison identified "vast series of beds of marl , schist , limestone , sandstone and conglomerate" that succeeded Carboniferous strata in the region. Murchison, in collaboration with Russian geologists, named the period after

3780-590: The Greek tortoise , and the Nile crocodile . Amniotes are distinguished from the other living tetrapod clade — the non-amniote lissamphibians ( frogs / toads , salamanders , newts and caecilians ) — by the development of three extraembryonic membranes ( amnion for embryonic protection, chorion for gas exchange , and allantois for metabolic waste disposal or storage), thicker and keratinized skin , costal respiration (breathing by expanding/constricting

3888-456: The amnion , which derives from Greek ἀμνίον ( amnion ), which denoted the membrane that surrounds a fetus. The term originally described a bowl in which the blood of sacrificed animals was caught, and derived from ἀμνός ( amnos ), meaning "lamb". Zoologists characterize amniotes in part by embryonic development that includes the formation of several extensive membranes, the amnion , chorion , and allantois . Amniotes develop directly into

3996-473: The placenta . The ancestors of true amniotes, such as Casineria kiddi , which lived about 340 million years ago, evolved from amphibian reptiliomorphs and resembled small lizards. At the late Devonian mass extinction (360 million years ago), all known tetrapods were essentially aquatic and fish-like. Because the reptiliomorphs were already established 20 million years later when all their fishlike relatives were extinct, it appears they separated from

4104-514: The pseudosuchians , dinosaurs , and pterosaurs in the following Triassic, first appeared and diversified during the Late Permian, including the first appearance of the Archosauriformes during the latest Permian. Cynodonts , the group of therapsids ancestral to modern mammals , first appeared and gained a worldwide distribution during the Late Permian. Another group of therapsids, the therocephalians (such as Lycosuchus ), arose in

4212-430: The rib cage ), the presence of adrenocortical and chromaffin tissues as a discrete pair of glands near their kidneys , more complex kidneys , the presence of an astragalus for better extremity range of motion , the diminished role of skin breathing , and the complete loss of metamorphosis , gills , and lateral lines . The presence of an amniotic buffer, of a water-impermeable skin , and of

4320-641: The "Uralian Series", but the subsequent inconsistent usage of this term meant that it was later abandoned. The Asselian was named by the Russian stratigrapher V.E. Ruzhenchev in 1954, after the Assel River in the southern Ural Mountains. The GSSP for the base of the Asselian is located in the Aidaralash River valley near Aqtöbe , Kazakhstan, which was ratified in 1996. The beginning of the stage

4428-650: The Asselian and Sakmarian, during which the LPIA peaked. By 287 million years ago, temperatures warmed and the South Pole ice cap retreated in what was known as the Artinskian Warming Event (AWE), though glaciers remained present in the uplands of eastern Australia, and perhaps also the mountainous regions of far northern Siberia. Southern Africa also retained glaciers during the late Cisuralian in upland environments. The AWE also witnessed aridification of

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4536-548: The Carboniferous, declined during the Permian. This is likely due to competition by Hemiptera , due to their similar mouthparts and therefore ecology. Primitive relatives of damselflies and dragonflies ( Meganisoptera ), which include the largest flying insects of all time, also declined during the Permian. Holometabola , the largest group of modern insects, also diversified during this time. " Grylloblattidans ", an extinct group of winged insects thought to be related to modern ice crawlers , reached their apex of diversity during

4644-599: The Carboniferous, reached their maximum height during the early Permian around 295 million years ago, comparable to the present Himalayas , but became heavily eroded as the Permian progressed. The Kazakhstania block collided with Baltica during the Cisuralian, while the North China Craton , the South China Block and Indochina fused to each other and Pangea by the end of the Permian. The Zechstein Sea ,

4752-663: The Changhsingian. The Permian was cool in comparison to most other geologic time periods, with modest pole to Equator temperature gradients. At the start of the Permian, the Earth was still in the Late Paleozoic icehouse (LPIA), which began in the latest Devonian and spanned the entire Carboniferous period, with its most intense phase occurring during the latter part of the Pennsylvanian epoch. A significant trend of increasing aridification can be observed over

4860-448: The Cisuralian. Permian synapsids included some large members such as Dimetrodon . The special adaptations of synapsids enabled them to flourish in the drier climate of the Permian and they grew to dominate the vertebrates. A faunal turnover occurred around the transition between the Cisuralian and Guadalupian, with the decline of amphibians and the replacement of pelycosaurs (a paraphyletic group) with more advanced therapsids , although

4968-498: The Early Permian Chemnitz petrified forest of Germany demonstrates that they had complex branching patterns similar to modern angiosperm trees. By the Late Permian, high thin forests had become widespread across the globe, as evidenced by the global distribution of weigeltisaurids. The oldest likely record of Ginkgoales (the group containing Ginkgo and its close relatives) is Trichopitys heteromorpha from

5076-578: The Early Permian were low. Trilobites underwent a diversification during the Kungurian-Wordian, the last in their evolutionary history, before declining during the Late Permian. By the Changhsingian, only a handful (4-6) genera remained. Corals exhibited a decline in diversity over the course of the Middle and Late Permian. Terrestrial life in the Permian included diverse plants, fungi , arthropods , and various types of tetrapods . The period saw

5184-565: The Early Triassic, but a Permian origin is suspected. The diversity of coelacanths is relatively low throughout the Permian in comparison to other marine fishes, though there is an increase in diversity during the terminal Permian (Changhsingian), corresponding with the highest diversity in their evolutionary history during the Early Triassic. Diversity of freshwater fish faunas was generally low and dominated by lungfish and "Paleopterygians". The last common ancestor of all living lungfish

5292-538: The Guadalupe Mountains of Texas, named by George Burr Richardson in 1904, and first used in a chronostratigraphic sense by Glenister and Furnish in 1961 as a substage of the Guadalupian Stage. The Capitanian was ratified as an international stage by the ICS in 2001. The GSSP for the base of the Capitanian is located at Nipple Hill in the southeast Guadalupe Mountains of Texas, and was ratified in 2001,

5400-468: The Late Permian. Members of the modern orders Archostemata and Adephaga are known from the Late Permian. Complex wood boring traces found in the Late Permian of China suggest that members of Polyphaga , the most diverse group of modern beetles, were also present by the Late Permian. The terrestrial fossil record of the Permian is patchy and temporally discontinuous. Early Permian records are dominated by equatorial Europe and North America, while those of

5508-562: The Lopingian in China. Modern amphibians ( lissamphibians ) are suggested to have originated during Permian, descending from a lineage of dissorophoid temnospondyls or lepospondyls . The diversity of fish during the Permian is relatively low compared to the following Triassic. The dominant group of bony fishes during the Permian were the " Paleopterygii " a paraphyletic grouping of Actinopterygii that lie outside of Neopterygii . The earliest unequivocal members of Neopterygii appear during

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5616-528: The Meishan D section, Zhejiang, China and was ratified in 2005, the boundary is defined by the first appearance of Clarkina wangi. The GSSP for the base of the Triassic is located at the base of Bed 27c at the Meishan D section, and was ratified in 2001. The GSSP is defined by the first appearance of the conodont Hindeodus parvus . The Russian Tatarian Stage includes the Lopingian, Capitanian and part of

5724-508: The Middle Permian. There were no flying vertebrates, though the extinct lizard-like reptile family Weigeltisauridae from the Late Permian had extendable wings like modern gliding lizards , and are the oldest known gliding vertebrates. Permian stem-amniotes consisted of lepospondyli and batrachosaurs , according to some phylogenies; according to others, stem-amniotes are represented only by diadectomorphs . Temnospondyls reached

5832-594: The Middle and Late Permian are dominated by temperate Karoo Supergroup sediments of South Africa and the Ural region of European Russia. Early Permian terrestrial faunas of North America and Europe were dominated by primitive pelycosaur synapsids including the herbivorous edaphosaurids , and carnivorous sphenacodontids , diadectids and amphibians . Early Permian reptiles, such as acleistorhinids , were mostly small insectivores. Synapsids (the group that would later include mammals) thrived and diversified greatly during

5940-577: The Palaeozoic and the Triassic. The Cisuralian Series is named after the strata exposed on the western slopes of the Ural Mountains in Russia and Kazakhstan. The name was proposed by J. B. Waterhouse in 1982 to comprise the Asselian, Sakmarian, and Artinskian stages. The Kungurian was later added to conform to the Russian "Lower Permian". Albert Auguste Cochon de Lapparent in 1900 had proposed

6048-595: The Paleo-Tethys Ocean to shrink. A new ocean was growing on its southern end, the Neotethys Ocean , an ocean that would dominate much of the Mesozoic Era. A magmatic arc, containing Hainan on its southwesternmost end, began to form as Panthalassa subducted under the southeastern South China. The Central Pangean Mountains , which began forming due to the collision of Laurasia and Gondwana during

6156-580: The Permian, representing up to a third of all insects at some localities. Mecoptera (sometimes known as scorpionflies) first appeared during the Early Permian, going on to become diverse during the Late Permian. Some Permian mecopterans, like Mesopsychidae have long proboscis that suggest they may have pollinated gymnosperms. The earliest known beetles appeared at the beginning of the Permian. Early beetles such as members of Permocupedidae were likely xylophagous , feeding on decaying wood. Several lineages such as Schizophoridae expanded into aquatic habitats by

6264-504: The Permian. Xenacanthiformes , another extinct group of shark-like chondrichthyans, were common in freshwater habitats, and represented the apex predators of freshwater ecosystems. Four floristic provinces in the Permian are recognised, the Angaran , Euramerican, Gondwanan, and Cathaysian realms. The Carboniferous Rainforest Collapse would result in the replacement of lycopsid -dominated forests with tree-fern dominated ones during

6372-801: The Wordian, while the underlying Kazanian includes the rest of the Wordian as well as the Roadian. In North America, the Permian is divided into the Wolfcampian (which includes the Nealian and the Lenoxian stages); the Leonardian (Hessian and Cathedralian stages); the Guadalupian; and the Ochoan, corresponding to the Lopingian. During the Permian, all the Earth 's major landmasses were collected into

6480-449: The amniote ancestors, the next major breakthrough appears to have involved a gradual replacement of the gelatinous coating covering the amphibian egg with a fibrous shell membrane. This allowed the egg to increase both its size and in the rate of gas exchange, permitting a larger, metabolically more active embryo to reach full development before hatching. Further developments, like extraembryonic membranes (amnion, chorion, and allantois) and

6588-505: The amount of lava estimated to have been produced during this period, the worst-case scenario is the release of enough carbon dioxide from the eruptions to raise world temperatures five degrees Celsius. Another hypothesis involves ocean venting of hydrogen sulfide gas. Portions of the deep ocean will periodically lose all of its dissolved oxygen allowing bacteria that live without oxygen to flourish and produce hydrogen sulfide gas. If enough hydrogen sulfide accumulates in an anoxic zone ,

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6696-614: The base of the Roadian is located 42.7m above the base of the Cutoff Formation in Stratotype Canyon, Guadalupe Mountains, Texas, and was ratified in 2001. The beginning of the stage is defined by the first appearance of Jinogondolella nankingensis . The Wordian was named in reference to the Word Formation by Johan August Udden in 1916, Glenister and Furnish in 1961 was the first publication to use it as

6804-535: The beginning of the stage is defined by the first appearance of Jinogondolella postserrata. The Lopingian was first introduced by Amadeus William Grabau in 1923 as the "Loping Series" after Leping , Jiangxi , China. Originally used as a lithostraphic unit, T.K. Huang in 1932 raised the Lopingian to a series, including all Permian deposits in South China that overlie the Maokou Limestone. In 1995,

6912-501: The chondrichthyan clade Holocephali , which contains living chimaeras , reached their apex of diversity during the Carboniferous-Permian, the most famous Permian representative being the "buzz-saw shark" Helicoprion , known for its unusual spiral shaped spiral tooth whorl in the lower jaw. Hybodonts , a group of shark-like chondrichthyans, were widespread and abundant members of marine and freshwater faunas throughout

7020-555: The class Reptilia is paraphyletic —it has given rise to two other classes not included in Reptilia. Most species described as microsaurs , formerly grouped in the extinct and prehistoric amphibian group lepospondyls , has been placed in the newer clade Recumbirostra , and shares many anatomical features with amniotes which indicates they were amniotes themselves. A different approach is adopted by writers who reject paraphyletic groupings. One such classification, by Michael Benton ,

7128-446: The continental interior. Amniotes, which could better cope with these drier conditions, rose to dominance in place of their amphibian ancestors. Various authors recognise at least three, and possibly four extinction events in the Permian. The end of the Early Permian ( Cisuralian ) saw a major faunal turnover, with most lineages of primitive " pelycosaur " synapsids becoming extinct, being replaced by more advanced therapsids . The end of

7236-458: The course of the Cisuralian. Early Permian aridification was most notable in Pangaean localities at near-equatorial latitudes. Sea levels also rose notably in the Early Permian as the LPIA slowly waned. At the Carboniferous-Permian boundary, a warming event occurred. In addition to becoming warmer, the climate became notably more arid at the end of the Carboniferous and beginning of the Permian. Nonetheless, temperatures continued to cool during most of

7344-456: The decline of early synapsid clades was apparently a slow event that lasted about 20 Ma, from the Sakmarian to the end of the Kungurian . Predator-prey interactions among terrestrial synapsids became more dynamic. If terrestrial deposition ended around the end of the Cisuralian in North America and began in Russia during the early Guadalupian, a continuous record of the transition is not preserved. Uncertain dating has led to suggestions that there

7452-402: The diapsid line of descent. Post-cranial remains of amniotes can be identified from their Labyrinthodont ancestors by their having at least two pairs of sacral ribs , a sternum in the pectoral girdle (some amniotes have lost it) and an astragalus bone in the ankle. Amniota was first formally described by the embryologist Ernst Haeckel in 1866 on the presence of the amnion , hence

7560-411: The diverse Dinocephalia . Dinocephalians become extinct at the end of the Middle Permian, during the Capitanian mass extinction event . Late Permian faunas are dominated by advanced therapsids such as the predatory sabertoothed gorgonopsians and herbivorous beaked dicynodonts , alongside large herbivorous pareiasaur parareptiles . The Archosauromorpha , the group of reptiles that would give rise to

7668-492: The earliest Permian of France. The oldest known fossils definitively assignable to modern cycads are known from the Late Permian. In Cathaysia, where a wet tropical frost-free climate prevailed, the Noeggerathiales , an extinct group of tree fern-like progymnosperms were a common component of the flora The earliest Permian (~ 298 million years ago) Cathyasian Wuda Tuff flora, representing a coal swamp community, has an upper canopy consisting of lycopsid tree Sigillaria , with

7776-609: The earliest known crown group amniotes, the oldest known sauropsid is Hylonomus and the oldest known synapsid is Asaphestera , both of which are from Nova Scotia during the Bashkirian age of the Late Carboniferous around 318  million years ago . This basal divergence within Amniota has also been dated by molecular studies at 310–329 Ma, or 312–330 Ma, and by a fossilized birth–death process study at 322–340 Ma. The term amniote comes from

7884-400: The early amniotes resembled their amphibian ancestors in many respects, a key difference was the lack of an otic notch at the back margin of the skull roof . In their ancestors, this notch held a spiracle , an unnecessary structure in an animal without an aquatic larval stage. There are three main lines of amniotes, which may be distinguished by the structure of the skull and in particular

7992-415: The embryonic membrane. Evolution of the amniote egg required increased exchange of gases and wastes between the embryo and the atmosphere. Structures to permit these traits allowed further adaption that increased the feasible size of amniote eggs and enabled breeding in progressively drier habitats. The increased size of eggs permitted increase in size of offspring and consequently of adults. Further growth for

8100-494: The end of the Permian. Nautiloids , a subclass of cephalopods, surprisingly survived this occurrence. There is evidence that magma , in the form of flood basalt , poured onto the Earth's surface in what is now called the Siberian Traps , for thousands of years, contributing to the environmental stress that led to mass extinction. The reduced coastal habitat and highly increased aridity probably also contributed. Based on

8208-699: The eruption of the Siberian Traps . It took well into the Triassic for life to recover from this catastrophe; on land, ecosystems took 30 million years to recover. Prior to the introduction of the term Permian , rocks of equivalent age in Germany had been named the Rotliegend and Zechstein , and in Great Britain as the New Red Sandstone . The term Permian was introduced into geology in 1841 by Sir Roderick Impey Murchison , president of

8316-550: The essential geologic time markers, based on their relative ages and the law of superposition . The divisions of the geological time scale were described and put in chronological order by the geologists and stratigraphers of the 18th and 19th centuries. Geologic formations can be usefully defined for sedimentary rock layers, low-grade metamorphic rocks , and volcanic rocks . Intrusive igneous rocks and highly metamorphosed rocks are generally not considered to be formations, but are described instead as lithodemes . "Formation"

8424-579: The extreme magnitude of this climatic shift. This extremely rapid interval of greenhouse gas release caused the Permian-Triassic mass extinction, as well as ushering in an extreme hothouse that persisted for several million years into the next geologic epoch, the Triassic. The Permian climate was also extremely seasonal and characterised by megamonsoons , which produced high aridity and extreme seasonality in Pangaea's interiors. Precipitation along

8532-507: The finale of the Late Palaeozoic Ice Age, when the last Australian glaciers melted. The end of the Permian is marked by a temperature excursion, much larger than the Emeishan Thermal Excursion, at the Permian-Triassic boundary, corresponding to the eruption of the Siberian Traps , which released more than 5 teratonnes of CO 2 , more than doubling the atmospheric carbon dioxide concentration. A -2% δ O excursion signifies

8640-522: The formation is expected to describe the stratotype in sufficient detail that other geologists can unequivocally recognize the formation. Although formations should not be defined by any criteria other than primary lithology, it is often useful to define biostratigraphic units on paleontological criteria, chronostratigraphic units on the age of the rocks, and chemostratigraphic units on geochemical criteria, and these are included in stratigraphic codes. The concept of formally defined layers or strata

8748-429: The gas can rise into the atmosphere. Oxidizing gases in the atmosphere would destroy the toxic gas, but the hydrogen sulfide would soon consume all of the atmospheric gas available. Hydrogen sulfide levels might have increased dramatically over a few hundred years. Models of such an event indicate that the gas would destroy ozone in the upper atmosphere allowing ultraviolet radiation to kill off species that had survived

8856-515: The increase in size and yolk content of eggs may have permitted, and coevolved with, direct development of the embryo to a large size. Features of amniotes evolved for survival on land include a sturdy but porous leathery or hard eggshell and an allantois that facilitates respiration while providing a reservoir for disposal of wastes. Their kidneys (metanephros) and large intestines are also well-suited to water retention. Most mammals do not lay eggs, but corresponding structures develop inside

8964-746: The late Carboniferous in Euramerica, and result in the differentiation of the Cathaysian floras from those of Euramerica. The Gondwanan floristic region was dominated by Glossopteridales , a group of woody gymnosperm plants, for most of the Permian, extending to high southern latitudes. The ecology of the most prominent glossopterid, Glossopteris , has been compared to that of bald cypress , living in mires with waterlogged soils. The tree-like calamites , distant relatives of modern horsetails , lived in coal swamps and grew in bamboo -like vertical thickets. A mostly complete specimen of Arthropitys from

9072-523: The late part of the Capitanian, around 260 million years ago, corresponding to the eruption of the Emeishan Traps . This interval of rapid climate change was responsible for the Capitanian mass extinction event. During the early Wuchiapingian, following the emplacement of the Emeishan Traps, global temperatures declined as carbon dioxide was weathered out of the atmosphere by the large igneous province's emplaced basalts. The late Wuchiapingian saw

9180-410: The latter, however, was limited by their position in the terrestrial food-chain , which was restricted to level three and below, with only invertebrates occupying level two. Amniotes would eventually experience adaptive radiations when some species evolved the ability to digest plants and new ecological niches opened up, permitting larger body-size for herbivores, omnivores and predators. While

9288-515: The middle of the Permian a major transition in vegetation began. The swamp -loving lycopod trees of the Carboniferous, such as Lepidodendron and Sigillaria , were progressively replaced in the continental interior by the more advanced seed ferns and early conifers as a result of the Carboniferous rainforest collapse . At the close of the Permian, lycopod and equisete swamps reminiscent of Carboniferous flora survived only in Cathaysia ,

9396-511: The most extensive extinction event recorded in paleontology : the Permian–Triassic extinction event . 90 to 95% of marine species became extinct , as well as 70% of all land organisms. It is also the only known mass extinction of insects. Recovery from the Permian–Triassic extinction event was protracted; on land, ecosystems took 30 million years to recover. Trilobites , which had thrived since Cambrian times, finally became extinct before

9504-473: The most potent greenhouse gases ) into the atmosphere to raise world temperatures an additional five degrees Celsius. The frozen methane hypothesis helps explain the increase in carbon-12 levels found midway in the Permian–Triassic boundary layer. It also helps explain why the first phase of the layer's extinctions was land-based, the second was marine-based (and starting right after the increase in C-12 levels), and

9612-432: The mother ( marsupial and placental mammals ). This distinguishes amniotes from anamniotes ( fish and amphibians) that have to spawn in aquatic environments . Most amniotes still require regular access to drinking water for rehydration, like the semiaquatic amphibians do. They have better homeostasis in drier environments, and more efficient non-aquatic gas exchange to power terrestrial locomotion , which

9720-448: The name. A problem with this definition is that the trait ( apomorphy ) in question does not fossilize , and the status of fossil forms has to be inferred from other traits. Older classifications of the amniotes traditionally recognised three classes based on major traits and physiology : This rather orderly scheme is the one most commonly found in popular and basic scientific works. It has come under critique from cladistics , as

9828-408: The number of holes behind each eye. In anapsids , the ancestral condition, there are none; in synapsids (mammals and their extinct relatives) there is one; and in diapsids (including birds, crocodilians , squamates , and tuataras ), there are two. Turtles have secondarily lost their fenestrae, and were traditionally classified as anapsids because of this. Molecular testing firmly places them in

9936-399: The other tetrapods somewhere during Romer's gap , when the adult tetrapods became fully terrestrial (some forms would later become secondarily aquatic). The modest-sized ancestors of the amniotes laid their eggs in moist places, such as depressions under fallen logs or other suitable places in the Carboniferous swamps and forests; and dry conditions probably do not account for the emergence of

10044-483: The preceding Carboniferous, experienced a dramatic increase in diversification during the Early Permian. Towards the end of the Permian, there was a substantial drop in both origination and extinction rates. The dominant insects during the Permian Period were early representatives of Paleoptera , Polyneoptera , and Paraneoptera . Palaeodictyopteroidea , which had represented the dominant group of insects during

10152-729: The size of the adults. This is supported by the fact that extant squamate species that lay eggs less than 1 cm in diameter have adults whose snout-vent length is less than 10 cm. The only way for the eggs to increase in size would be to develop new internal structures specialized for respiration and for waste products. As this happened, it would also affect how much the juveniles could grow before they reached adulthood. A similar pattern can be seen in modern amphibians. Frogs that have evolved terrestrial reproduction and direct development have both smaller adults and fewer and larger eggs compared to their relatives that still reproduce in water. Fish and amphibian eggs have only one inner membrane,

10260-401: The soft shell. Indeed, many modern-day amniotes require moisture to keep their eggs from desiccating . Although some modern amphibians lay eggs on land, all amphibians lack advanced traits like an amnion. The amniotic egg formed through a series of evolutionary steps. After internal fertilization and the habit of laying eggs in terrestrial environments became a reproduction strategy amongst

10368-529: The spread of conifers and their increasing prevalence throughout terrestrial ecosystems. Bennettitales , which would go on to become in widespread the Mesozoic, first appeared during the Cisuralian in China. Lyginopterids , which had declined in the late Pennsylvanian and subsequently have a patchy fossil record, survived into the Late Permian in Cathaysia and equatorial east Gondwana. The Permian ended with

10476-609: The stage. The ages of the Permian, from youngest to oldest, are: For most of the 20th century, the Permian was divided into the Early and Late Permian, with the Kungurian being the last stage of the Early Permian. Glenister and colleagues in 1992 proposed a tripartite scheme, advocating that the Roadian-Capitanian was distinct from the rest of the Late Permian, and should be regarded as a separate epoch. The tripartite split

10584-643: The surface are fundamental to such fields as structural geology , allowing geologists to infer the tectonic history of a region or predict likely locations for buried mineral resources. The boundaries of a formation are chosen to give it the greatest practical lithological consistency. Formations should not be defined by any criteria other than lithology. The lithology of a formation includes characteristics such as chemical and mineralogical composition, texture, color, primary depositional structures , fossils regarded as rock-forming particles, or other organic materials such as coal or kerogen . The taxonomy of fossils

10692-432: The surface or traced in the subsurface. Formations are otherwise not defined by the thickness of their rock strata, which can vary widely. They are usually, but not universally, tabular in form. They may consist of a single lithology (rock type), or of alternating beds of two or more lithologies, or even a heterogeneous mixture of lithologies, so long as this distinguishes them from adjacent bodies of rock. The concept of

10800-671: The surrounding Russian region of Perm, which takes its name from the medieval kingdom of Permia that occupied the same area hundreds of years prior, and which is now located in the Perm Krai administrative region. Between 1853 and 1867, Jules Marcou recognised Permian strata in a large area of North America from the Mississippi River to the Colorado River and proposed the name Dyassic , from Dyas and Trias , though Murchison rejected this in 1871. The Permian system

10908-418: The thickest deposits of Permian rocks in the world. Sea levels dropped slightly during the earliest Permian (Asselian). The sea level was stable at several tens of metres above present during the Early Permian, but there was a sharp drop beginning during the Roadian, culminating in the lowest sea level of the entire Palaeozoic at around present sea level during the Wuchiapingian, followed by a slight rise during

11016-413: The third land-based again. Amniote Amniotes are tetrapod vertebrate animals belonging to the clade Amniota , a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the Carboniferous period . Those of Amniota are defined as the smallest crown clade containing humans ,

11124-412: The toxic gas. There are species that can metabolize hydrogen sulfide. Another hypothesis builds on the flood basalt eruption theory. An increase in temperature of five degrees Celsius would not be enough to explain the death of 95% of life. But such warming could slowly raise ocean temperatures until frozen methane reservoirs below the ocean floor near coastlines melted, expelling enough methane (among

11232-628: The western margins of the Palaeo-Tethys Ocean was very high. Evidence for the megamonsoon includes the presence of megamonsoonal rainforests in the Qiangtang Basin of Tibet, enormous seasonal variation in sedimentation, bioturbation, and ichnofossil deposition recorded in sedimentary facies in the Sydney Basin , and palaeoclimatic models of the Earth's climate based on the behaviour of modern weather patterns showing that such

11340-514: Was adopted after a formal proposal by Glenister et al. (1999). Historically, most marine biostratigraphy of the Permian was based on ammonoids ; however, ammonoid localities are rare in Permian stratigraphic sections, and species characterise relatively long periods of time. All GSSPs for the Permian are based around the first appearance datum of specific species of conodont , an enigmatic group of jawless chordates with hard tooth-like oral elements. Conodonts are used as index fossils for most of

11448-547: Was controversial for over a century after its original naming, with the United States Geological Survey until 1941 considering the Permian a subsystem of the Carboniferous equivalent to the Mississippian and Pennsylvanian . The Permian Period is divided into three epochs , from oldest to youngest, the Cisuralian, Guadalupian, and Lopingian. Geologists divide the rocks of the Permian into

11556-611: Was named after the city of Arti in Sverdlovsk Oblast , Russia. It was named by Karpinsky in 1874. The Artinskian currently lacks a defined GSSP. The proposed definition for the base of the Artinskian is the first appearance of Sweetognathus aff. S. whitei. The Kungurian takes its name after Kungur , a city in Perm Krai. The stage was introduced by Alexandr Antonovich Stukenberg in 1890. The Kungurian currently lacks

11664-526: Was ratified in 2004. The boundary is defined by the first appearance of Clarkina postbitteri postbitteri The Changhsingian was originally derived from the Changxing Limestone, a geological unit first named by the Grabau in 1923, ultimately deriving from Changxing County , Zhejiang .The GSSP for the base of the Changhsingian is located 88 cm above the base of the Changxing Limestone in

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