The Permian[note 1] is a geologic period and system characterized among land vertebrates by the diversification of the early amniotes into the ancestral groups of the mammals, turtles, lepidosaurs and archosaurs. The Permian Period follows the Carboniferous and extends from 299.0 ± 0.8 to 251.0 ± 0.4 Ma (million years before the present). It is the last period of the Paleozoic Era and famous for its ending epoch event, the largest mass extinction known to science. The Permian Period was named after the kingdom of Permia in modern-day Russia by Scottish geologist Roderick Murchison in 1841.
Official (ICS, 2004, Subdivisions of the Permian System, from most recent to most ancient rock layers are:
Sea levels in the Permian remained generally low, and near-shore environments were limited by the collection of almost all major landmasses into a single continent -- Pangaea. This could have in part caused the widespread extinctions of marine species at the end of the period by severely reducing shallow coastal areas preferred by many marine organisms.
During the Permian, all the Earth's major land masses were collected into a single supercontinent known as Pangaea. 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 was between Asia and Gondwana. The Cimmeria continent rifted away from Gondwana and drifted north to Laurasia, causing the Paleo-Tethys to shrink. A new ocean was growing on its southern end, the Tethys Ocean, an ocean that would dominate much of the Mesozoic Era. Large continental landmasses create 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. 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, where the Permian Basin in the U.S. state of Texas is so named because it has one of the thickest deposits of Permian rocks in the world.
The climate in the Permian was quite varied. At the start of the Permian, the Earth was still at the grip of an Ice Age from the Carboniferous. Glaciers receded around the mid-Permian period as the climate gradually warmed, drying the continent's interiors. In the late Permian period, the drying continued although the temperature cycled between warm and cool cycles.
Permian marine deposits are rich in fossil mollusks, echinoderms, and brachiopods. Fossilized shells of two kinds of invertebrates are widely used to identify Permian strata and correlate them between sites: fusulinids, a kind of shelled amoeba-like protist that is one of the foraminiferans, and ammonoids, shelled cephalopods that are distant relatives of the modern nautilus. By the close of the Permian, trilobites and a host of other marine groups became extinct.
Terrestrial life in the Permian included diverse plants, fungi, arthropods, and various types of tetrapods. The period saw a massive desert covering the interior of the 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 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. At the close of the Permian, lycopod and equicete swamps reminiscent of Carboniferous flora was relegated to a series of equatorial islands in the Paleotethys Sea that later would become the 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 of the Permian
By the Pennsylvanian and well into the Permian, by far the most successful were primitive relatives of cockroaches. Six fast legs, two well developed folding wings, fairly good eyes, long, well developed antennae (olfactory), an omnivorous digestive system, a receptacle for storing sperm, a chitin skeleton that could support and protect, as well as form of gizzard and efficient mouth parts, gave it formidable advantages over other herbivorous animals. About 90% of insects were cockroach-like insects ("Blattopterans").
The dragonflies Odonata were the dominant aerial predator and probably dominated terrestrial insect predation as well. True Odonata appeared in the Permian and all are amphibious. Their prototypes are the oldest winged fossils, go back to the Devonian, and are different from other wings in every way. Their prototypes may have had the beginnings of many modern attributes even by late Carboniferous and it is possible that they even captured small vertebrates, for some species had a wing span of 71 cm. A number of important new insect groups appeared at this time, including the Coleoptera (beetles) and Diptera (flies).
Reptile and amphibian fauna
Early Permian terrestrial faunas were dominated by pelycosaurs and amphibians, the middle Permian by primitive therapsids such as the dinocephalia, and the late Permian by more advanced therapsids such as gorgonopsians and dicynodonts. Towards the very end of the Permian the first archosaurs appeared, a group that would give rise to the dinosaurs in the following period. Also appearing at the end of the Permian were the first cynodonts, which would go on to evolve into mammals during the Triassic. Another group of therapsids, the therocephalians (such as Trochosaurus), arose in the Middle Permian. There were no aerial vertebrates.
The Permian period saw the development of a fully terrestrial fauna and the appearance of the first large herbivores and carnivores. It was the high tide of the anapsides in the form of the massive Pareiasaurs and host of smaller, generally lizard-like groups. A group of small reptiles, the diapsids started to abound. These were the ancestors to most modern reptiles and the ruling dinosaurs as well as pterosaurs and crocodiles.
Thriving also, were the early ancestors to mammals, the synapsida, which included some large reptiles such as Dimetrodon. Reptiles grew to dominance among vertebrates, because their special adaptations enabled them to flourish in the drier climate.
Permian amphibians consisted of temnospondyli, lepospondyli and batrachosaurs.
Permian–Triassic extinction event
The Permian ended with 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. On an individual level, perhaps as many as 99.5% of separate organisms died as a result of the event. Recovery from the Permian-Triassic extinction event was protracted; on land ecosystems took 30M years to recover.
There is also significant evidence that massive flood basalt eruptions from magma output lasting thousands of years in what is now the Siberian Traps contributed to environmental stress leading to mass extinction. The reduced coastal habitat and highly increased aridity probably also contributed. Based on the amount of lava estimated to have been produced during this period, the worst-case scenario is an expulsion 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 deep ocean will periodically lose all of their dissolved oxygen allowing bacteria that live without oxygen to flourish and produce hydrogen sulfide gas. If enough hydrogen sulfide accumulates in an anoxic zone, 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 to change it. Hydrogen sulfide levels would increase dramatically over a few hundred years.
Modeling of such an event indicates that the gas would destroy ozone in the upper atmosphere allowing ultraviolet radiation to kill off species that had survived the toxic gas. Of course, there are species that can metabolize hydrogen sulfide.
Another hypothesis builds on the flood basalt eruption theory. Five degrees Celsius would not be enough increase in world temperatures 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 (a current target for a new energy source) melted, expelling enough methane, among 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 midway into 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 the third land-based again.
An even more speculative hypothesis is that intense radiation from a nearby supernova was responsible for the extinctions.
Trilobites, which had thrived since Cambrian times, finally became extinct before the end of the Permian.
Nautiluses, a species of cephalopods, surprisingly survived this occurrence.
In 2006, a group of American scientists from Ohio State University reported evidence for a possible huge meteorite crater (Wilkes Land crater) with a diameter of around 500 kilometers in Antarctica. The crater is located at a depth of 1.6 kilometers beneath the ice of Wilkes Land in eastern Antarctica. The scientists speculate that this impact may have caused the Permian–Triassic extinction event, although its age is bracketed only between 100 million and 500 million years ago. They also speculate that it may have contributed in some way to the separation of Australia from the Antarctic landmass, which were both part of a supercontinent called Gondwana. Levels of iridium and quartz fracturing in the Permian-Triassic layer do not approach those of the Cretaceous-Tertiary boundary layer. Given that a far greater proportion of species and individual organisms became extinct during the former, doubt is cast on the significance of a meteor impact in creating the latter. Further doubt has been cast on this theory based on fossils in Greenland showing the extinction to have been gradual, lasting about eighty thousand years, with three distinct phases.
Many scientists believe that the Permian-Triassic extinction event was caused by a combination of some or all of the hypotheses above and other factors; the formation of Pangaea decreased the number of coastal habitats and may have contributed to the extinction of many clades.
* List of fossil sites (with link directory)
1. ^ The term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian system" after the ancient kingdom of Permia, and not after the then small town of Perm, as usually assumed; see "Origin of the Permian"
1. ^ Image:Sauerstoffgehalt-1000mj.svg
* Ogg, Jim; June, 2004, Overview of Global Boundary Stratotype Sections and Points (GSSP's) http://www.stratigraphy.org/gssp.htm Accessed April 30, 2006.