Detritivores known as detrivores, detritus feeders, or detritus eaters, are heterotrophs that obtain nutrients by consuming detritus. There are many kinds of invertebrates and plants that carry out coprophagy. By doing so, all these detritivores contribute to the nutrient cycles, they should be distinguished from other decomposers, such as many species of bacteria and protists, which are unable to ingest discrete lumps of matter, but instead live by absorbing and metabolizing on a molecular scale. However, the terms detritivore and decomposer are used interchangeably. Detritivores are an important aspect of many ecosystems, they can live on any type of soil with an organic component, including marine ecosystems, where they are termed interchangeably with bottom feeders. Typical detritivorous animals include millipedes, woodlice, dung flies, many terrestrial worms, sea stars, sea cucumbers, fiddler crabs, some sedentary polychaetes such as amphitrites and other terebellids. Scavengers are not thought to be detritivores, as they eat large quantities of organic matter, but both detritivores and scavengers are same type of cases of consumer-resource systems.
The consumption of wood, whether dead, is known as xylophagy. Τhe activity of animals feeding only on dead wood is called sapro-xylophagy and those animals, sapro-xylophagous. In food webs, detritivores play the roles of decomposers. Detritivores are eaten by consumers and therefore play important roles as recyclers in ecosystem energy flow and biogeochemical cycles. Many detritivores live in mature woodland, though the term can be applied to certain bottom-feeders in wet environments; these organisms play a crucial role in benthic ecosystems, forming essential food chains and participating in the nitrogen cycle. Fungi, acting as decomposers, are important in today's terrestrial environment. During the Carboniferous period and bacteria had yet to evolve the capacity to digest lignin, so large deposits of dead plant tissue accumulated during this period becoming the fossil fuels. By feeding on sediments directly to extract the organic component, some detritivores incidentally concentrate toxic pollutants.
Decomposer Saprotrophic nutrition Nepenthes ampullaria Consumer-resource systems
The Precambrian is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of the Phanerozoic eon, named after Cambria, the Latinised name for Wales, where rocks from this age were first studied; the Precambrian accounts for 88% of the Earth's geologic time. The Precambrian is an informal unit of geologic time, subdivided into three eons of the geologic time scale, it spans from the formation of Earth about 4.6 billion years ago to the beginning of the Cambrian Period, about 541 million years ago, when hard-shelled creatures first appeared in abundance. Little is known about the Precambrian, despite it making up seven-eighths of the Earth's history, what is known has been discovered from the 1960s onwards; the Precambrian fossil record is poorer than that of the succeeding Phanerozoic, fossils from the Precambrian are of limited biostratigraphic use. This is because many Precambrian rocks have been metamorphosed, obscuring their origins, while others have been destroyed by erosion, or remain buried beneath Phanerozoic strata.
It is thought that the Earth coalesced from material in orbit around the Sun at 4,543 Ma, may have been struck by a large planetesimal shortly after it formed, splitting off material that formed the Moon. A stable crust was in place by 4,433 Ma, since zircon crystals from Western Australia have been dated at 4,404 ± 8 Ma; the term "Precambrian" is recognized by the International Commission on Stratigraphy as the only "supereon" in geologic time. "Precambrian" is still used by geologists and paleontologists for general discussions not requiring the more specific eon names. As of 2010, the United States Geological Survey considers the term informal, lacking a stratigraphic rank. A specific date for the origin of life has not been determined. Carbon found in 3.8 billion-year-old rocks from islands off western Greenland may be of organic origin. Well-preserved microscopic fossils of bacteria older than 3.46 billion years have been found in Western Australia. Probable fossils 100 million years older have been found in the same area.
However, there is evidence. There is a solid record of bacterial life throughout the remainder of the Precambrian. Excluding a few contested reports of much older forms from North America and India, the first complex multicellular life forms seem to have appeared at 1500 Ma, in the Mesoproterozoic era of the Proterozoic eon. Fossil evidence from the Ediacaran period of such complex life comes from the Lantian formation, at least 580 million years ago. A diverse collection of soft-bodied forms is found in a variety of locations worldwide and date to between 635 and 542 Ma; these are referred to as Vendian biota. Hard-shelled creatures appeared toward the end of that time span, marking the beginning of the Phanerozoic eon. By the middle of the following Cambrian period, a diverse fauna is recorded in the Burgess Shale, including some which may represent stem groups of modern taxa; the increase in diversity of lifeforms during the early Cambrian is called the Cambrian explosion of life. While land seems to have been devoid of plants and animals and other microbes formed prokaryotic mats that covered terrestrial areas.
Tracks from an animal with leg like appendages have been found in what was mud 551 million years ago. Evidence of the details of plate motions and other tectonic activity in the Precambrian has been poorly preserved, it is believed that small proto-continents existed prior to 4280 Ma, that most of the Earth's landmasses collected into a single supercontinent around 1130 Ma. The supercontinent, known as Rodinia, broke up around 750 Ma. A number of glacial periods have been identified going as far back as the Huronian epoch 2400–2100 Ma. One of the best studied is the Sturtian-Varangian glaciation, around 850–635 Ma, which may have brought glacial conditions all the way to the equator, resulting in a "Snowball Earth"; the atmosphere of the early Earth is not well understood. Most geologists believe it was composed of nitrogen, carbon dioxide, other inert gases, was lacking in free oxygen. There is, evidence that an oxygen-rich atmosphere existed since the early Archean. At present, it is still believed that molecular oxygen was not a significant fraction of Earth's atmosphere until after photosynthetic life forms evolved and began to produce it in large quantities as a byproduct of their metabolism.
This radical shift from a chemically inert to an oxidizing atmosphere caused an ecological crisis, sometimes called the oxygen catastrophe. At first, oxygen would have combined with other elements in Earth's crust iron, removing it from the atmosphere. After the supply of oxidizable surfaces ran out, oxygen would have begun to accumulate in the atmosphere, the modern high-oxygen atmosphere would have developed. Evidence for this lies in older rocks that contain massive banded iron formations that were laid down as iron oxides. A terminology has evolved covering the early years of the Earth's existence, as radiometric dating has allowed real dates to be assigned to specific formations and features; the Precambrian is divided into
The Silurian is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago, to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the shortest period of the Paleozoic Era; as with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by several million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when 60% of marine species were wiped out. A significant evolutionary milestone during the Silurian was the diversification of jawed fish and bony fish. Multi-cellular life began to appear on land in the form of small, bryophyte-like and vascular plants that grew beside lakes and coastlines, terrestrial arthropods are first found on land during the Silurian. However, terrestrial life would not diversify and affect the landscape until the Devonian; the Silurian system was first identified by British geologist Roderick Murchison, examining fossil-bearing sedimentary rock strata in south Wales in the early 1830s.
He named the sequences for a Celtic tribe of Wales, the Silures, inspired by his friend Adam Sedgwick, who had named the period of his study the Cambrian, from the Latin name for Wales. This naming does not indicate any correlation between the occurrence of the Silurian rocks and the land inhabited by the Silures. In 1835 the two men presented a joint paper, under the title On the Silurian and Cambrian Systems, Exhibiting the Order in which the Older Sedimentary Strata Succeed each other in England and Wales, the germ of the modern geological time scale; as it was first identified, the "Silurian" series when traced farther afield came to overlap Sedgwick's "Cambrian" sequence, provoking furious disagreements that ended the friendship. Charles Lapworth resolved the conflict by defining a new Ordovician system including the contested beds. An early alternative name for the Silurian was "Gotlandian" after the strata of the Baltic island of Gotland; the French geologist Joachim Barrande, building on Murchison's work, used the term Silurian in a more comprehensive sense than was justified by subsequent knowledge.
He divided the Silurian rocks of Bohemia into eight stages. His interpretation was questioned in 1854 by Edward Forbes, the stages of Barrande, F, G and H, have since been shown to be Devonian. Despite these modifications in the original groupings of the strata, it is recognized that Barrande established Bohemia as a classic ground for the study of the earliest fossils; the Llandovery Epoch lasted from 443.8 ± 1.5 to 433.4 ± 2.8 mya, is subdivided into three stages: the Rhuddanian, lasting until 440.8 million years ago, the Aeronian, lasting to 438.5 million years ago, the Telychian. The epoch is named for the town of Llandovery in Wales; the Wenlock, which lasted from 433.4 ± 1.5 to 427.4 ± 2.8 mya, is subdivided into the Sheinwoodian and Homerian ages. It is named after Wenlock Edge in England. During the Wenlock, the oldest-known tracheophytes of the genus Cooksonia, appear; the complexity of later Gondwana plants like Baragwanathia, which resembled a modern clubmoss, indicates a much longer history for vascular plants, extending into the early Silurian or Ordovician.
The first terrestrial animals appear in the Wenlock, represented by air-breathing millipedes from Scotland. The Ludlow, lasting from 427.4 ± 1.5 to 423 ± 2.8 mya, comprises the Gorstian stage, lasting until 425.6 million years ago, the Ludfordian stage. It is named for the town of Ludlow in England; the Přídolí, lasting from 423 ± 1.5 to 419.2 ± 2.8 mya, is the final and shortest epoch of the Silurian. It is named after one locality at the Homolka a Přídolí nature reserve near the Prague suburb Slivenec in the Czech Republic. Přídolí is the old name of a cadastral field area. In North America a different suite of regional stages is sometimes used: Cayugan Lockportian Tonawandan Ontarian Alexandrian In Estonia the following suite of regional stages is used: Ohessaare stage Kaugatuma stage Kuressaare stage Paadla stage Rootsiküla stage Jaagarahu stage Jaani stage Adavere stage Raikküla stage Juuru stage With the supercontinent Gondwana covering the equator and much of the southern hemisphere, a large ocean occupied most of the northern half of the globe.
The high sea levels of the Silurian and the flat land resulted in a number of island chains, thus a rich diversity of environmental settings. During the Silurian, Gondwana continued a slow southward drift to high southern latitudes, but there is evidence that the Silurian icecaps were less extensive than those of the late-Ordovician glaciation; the southern continents remained united during this period. The melting of icecaps and glaciers contributed to a rise in sea level, recognizable from the fact that Silurian sediments overlie eroded Ordovician sediments, forming an unconformity; the continents of Avalonia and Laurentia drifted together near the equator, starting the formation of a second supercontinent known as Euramerica. When the proto-Europe coll
The Ordovician is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.2 million years from the end of the Cambrian Period 485.4 million years ago to the start of the Silurian Period 443.8 Mya. The Ordovician, named after the Celtic tribe of the Ordovices, was defined by Charles Lapworth in 1879 to resolve a dispute between followers of Adam Sedgwick and Roderick Murchison, who were placing the same rock beds in northern Wales into the Cambrian and Silurian systems, respectively. Lapworth recognized that the fossil fauna in the disputed strata were different from those of either the Cambrian or the Silurian systems, placed them in a system of their own; the Ordovician received international approval in 1960, when it was adopted as an official period of the Paleozoic Era by the International Geological Congress. Life continued to flourish during the Ordovician as it did in the earlier Cambrian period, although the end of the period was marked by the Ordovician–Silurian extinction events.
Invertebrates, namely molluscs and arthropods, dominated the oceans. The Great Ordovician Biodiversification Event increased the diversity of life. Fish, the world's first true vertebrates, continued to evolve, those with jaws may have first appeared late in the period. Life had yet to diversify on land. About 100 times as many meteorites struck the Earth per year during the Ordovician compared with today; the Ordovician Period began with a major extinction called the Cambrian–Ordovician extinction event, about 485.4 Mya. It lasted for about 42 million years and ended with the Ordovician–Silurian extinction events, about 443.8 Mya which wiped out 60% of marine genera. The dates given are recent radiometric dates and vary from those found in other sources; this second period of the Paleozoic era created abundant fossils that became major petroleum and gas reservoirs. The boundary chosen for the beginning of both the Ordovician Period and the Tremadocian stage is significant, it correlates well with the occurrence of widespread graptolite and trilobite species.
The base of the Tremadocian allows scientists to relate these species not only to each other, but to species that occur with them in other areas. This makes it easier to place many more species in time relative to the beginning of the Ordovician Period. A number of regional terms have been used to subdivide the Ordovician Period. In 2008, the ICS erected a formal international system of subdivisions. There exist Baltoscandic, Siberian, North American, Chinese Mediterranean and North-Gondwanan regional stratigraphic schemes; the Ordovician Period in Britain was traditionally broken into Early and Late epochs. The corresponding rocks of the Ordovician System are referred to as coming from the Lower, Middle, or Upper part of the column; the faunal stages from youngest to oldest are: Late Ordovician Hirnantian/Gamach Rawtheyan/Richmond Cautleyan/Richmond Pusgillian/Maysville/Richmond Middle Ordovician Trenton Onnian/Maysville/Eden Actonian/Eden Marshbrookian/Sherman Longvillian/Sherman Soudleyan/Kirkfield Harnagian/Rockland Costonian/Black River Chazy Llandeilo Whiterock Llanvirn Early Ordovician Cassinian Arenig/Jefferson/Castleman Tremadoc/Deming/Gaconadian The Tremadoc corresponds to the Tremadocian.
The Floian corresponds to the lower Arenig. The Llanvirn occupies the rest of the Darriwilian, terminates with it at the base of the Late Ordovician; the Sandbian represents the first half of the Caradoc. During the Ordovician, the southern continents were collected into Gondwana. Gondwana started the period in equatorial latitudes and, as the period progressed, drifted toward the South Pole. Early in the Ordovician, the continents of Laurentia and Baltica were still independent continents, but Baltica began to move towards Laurentia in the period, causing the Iapetus Ocean between them to shrink; the small continent Avalonia separated from Gondwana and began to move north towards Baltica and Laurentia, opening the Rheic Ocean between Gondwana and Avalonia. The Taconic orogeny, a major mountain-building episode, was well under way in Cambrian times. In the early and middle Ordovician, temperatures were mild, but at the beginning of the Late Ordovician, from 460 to 450 Ma, volcanoes along the margin of the Iapetus Ocean spewed massive amounts of carbon dioxide, a greenhouse gas, into the atmosphere, turning the planet into a hothouse.
Sea levels were high, but as Gondwana moved south, ice accumulated into glaciers and sea levels dropped. At first, low-lying sea beds increased diversity, but glaciation led to mass extinctions as the seas drained and continental shelves became dry land. During the Ordovician, in fact during the Tremadocian, marine transgressions worldwide were the greatest for which evidence is preserved; these volcanic island arcs collided with proto North America to form the Appalachian mountains. By the end of the Late Ordovician the volcanic emissions had stopped. Gondwana had by that time neared the South Pole and was glaciated
Hemocyanins are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms, they are second only to hemoglobin in frequency of use as an oxygen transport molecule. Unlike the hemoglobin in red blood cells found in vertebrates, hemocyanins are not bound to blood cells but are instead suspended directly in the hemolymph. Oxygenation causes a color change between the colorless Cu deoxygenated form and the blue Cu oxygenated form. Hemocyanins are found only in the Mollusca and Arthropoda: the earliest discoveries of hemocyanins were in the snail Helix pomatia and in the horseshoe crab, they were subsequently found to be common among crustaceans and are utilized by some land arthropods such as the tarantula Eurypelma californicum, the emperor scorpion, the centipede Scutigera coleoptrata. Larval storage proteins in many insects appear to be derived from hemocyanins; the arthropod hemocyanin superfamily is composed of phenoloxidases, pseudohemocyanins or cryptocyanins, hexamerin receptors.
Phenoloxidase are copper containing tyrosinases. These proteins are involved in the process of sclerotization of arthropod cuticle, in wound healing, humoral immune defense. Phenoloxidase are activated by cleaving a N-terminal peptide. Hexamerins are storage proteins found in insects; these proteins are synthesized by the larval fat body and are associated with molting cycles or nutritional conditions. Pseudohemocyanin and cryptocyanins genetic sequences are related to hemocyanins in crustaceans; these proteins have a similar structure and function, but lack the copper binding sites. The evolutionary changes within the phylogeny of the hemocyanin superfamily are related to the emergence of these different proteins in various species; the understanding of proteins within this superfamily would not be well understood without the extensive studies of hemocyanin in arthropods. Although the respiratory function of hemocyanin is similar to that of hemoglobin, there are a significant number of differences in its molecular structure and mechanism.
Whereas hemoglobin carries its iron atoms in porphyrin rings, the copper atoms of hemocyanin are bound as prosthetic groups coordinated by histidine residues. The active site of hemocyanin is composed of a pair of copper cations which are directly coordinated to the protein through the driving force of imidazolic rings of six histidine residues, it has been noted that species using hemocyanin for oxygen transportation include crustaceans living in cold environments with low oxygen pressure. Under these circumstances hemoglobin oxygen transportation is less efficient than hemocyanin oxygen transportation. There are terrestrial arthropods using hemocyanin, notably spiders and scorpions, that live in warm climates; the molecule is conformationally stable and functioning at temperatures up to 90 degrees C. Most hemocyanins bind with oxygen non-cooperatively and are one-fourth as efficient as hemoglobin at transporting oxygen per amount of blood. Hemoglobin binds oxygen cooperatively due to steric conformation changes in the protein complex, which increases hemoglobin's affinity for oxygen when oxygenated.
In some hemocyanins of horseshoe crabs and some other species of arthropods, cooperative binding is observed, with Hill coefficients of 1.6–3.0. Hill coefficients vary depending on laboratory measurement settings. Hemoglobin, for comparison, has a Hill coefficient of 2.8–3.0. In these cases of cooperative binding hemocyanin was arranged in protein sub-complexes of 6 subunits each with one oxygen binding site; each hexamer complex was arranged together to form a larger complex of dozens of hexamers. In one study, cooperative binding was found to be dependent on hexamers being arranged together in the larger complex, suggesting cooperative binding between hexamers. Hemocyanin oxygen-binding profile is affected by dissolved salt ion levels and pH. Hemocyanin is made of many individual subunit proteins, each of which contains two copper atoms and can bind one oxygen molecule; each subunit weighs about 75 kilodaltons. Subunits may be arranged in hexamers depending on species; the subunits are homogeneous, or heterogeneous with two variant subunit types.
Because of the large size of hemocyanin, it is found free-floating in the blood, unlike hemoglobin. Hexamers are characteristic of arthropod hemocyanins. A hemocyanin of the tarantula Eurypelma californicum is made up of 24 peptide chains. A hemocyanin from the house centipede Scutigera coleoptrata is made up of 36 chains. Horseshoe crabs have an 8-hexamer hemocyanin. Simple hexamers are found in the spiny lobster Panulirus interruptus and the isopod Bathynomus giganteus. Peptide chains in crustaceans are about 660 amino acid residues long, in chelicerates they are about 625. In the large complexes there is a variety of all about the same length. Hemocyanin is homologous to the phenol oxidases since both proteins share type 3 Cu active site coordination. In both cases inactive proenzymes such as hemocyanin and catcholoxidase must be activated first; this is done by removing the amino acid that blocks the entrance channel to the active site when the proenzyme is not active. Th
The Cambrian Period was the first geological period of the Paleozoic Era, of the Phanerozoic Eon. The Cambrian lasted 55.6 million years from the end of the preceding Ediacaran Period 541 million years ago to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, its base, are somewhat in flux; the period was established by Adam Sedgwick, who named it after Cambria, the Latin name of Wales, where Britain's Cambrian rocks are best exposed. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells; as a result, our understanding of the Cambrian biology surpasses that of some periods. The Cambrian marked a profound change in life on Earth. Complex, multicellular organisms became more common in the millions of years preceding the Cambrian, but it was not until this period that mineralized—hence fossilized—organisms became common; the rapid diversification of life forms in the Cambrian, known as the Cambrian explosion, produced the first representatives of all modern animal phyla.
Phylogenetic analysis has supported the view that during the Cambrian radiation, metazoa evolved monophyletically from a single common ancestor: flagellated colonial protists similar to modern choanoflagellates. Although diverse life forms prospered in the oceans, the land is thought to have been comparatively barren—with nothing more complex than a microbial soil crust and a few molluscs that emerged to browse on the microbial biofilm. Most of the continents were dry and rocky due to a lack of vegetation. Shallow seas flanked the margins of several continents created during the breakup of the supercontinent Pannotia; the seas were warm, polar ice was absent for much of the period. Despite the long recognition of its distinction from younger Ordovician rocks and older Precambrian rocks, it was not until 1994 that the Cambrian system/period was internationally ratified; the base of the Cambrian lies atop a complex assemblage of trace fossils known as the Treptichnus pedum assemblage. The use of Treptichnus pedum, a reference ichnofossil to mark the lower boundary of the Cambrian, is difficult since the occurrence of similar trace fossils belonging to the Treptichnids group are found well below the T. pedum in Namibia and Newfoundland, in the western USA.
The stratigraphic range of T. pedum overlaps the range of the Ediacaran fossils in Namibia, in Spain. The Cambrian Period was followed by the Ordovician Period; the Cambrian is divided into ten ages. Only three series and six stages are named and have a GSSP; because the international stratigraphic subdivision is not yet complete, many local subdivisions are still used. In some of these subdivisions the Cambrian is divided into three series with locally differing names – the Early Cambrian, Middle Cambrian and Furongian. Rocks of these epochs are referred to as belonging to Upper Cambrian. Trilobite zones allow biostratigraphic correlation in the Cambrian; each of the local series is divided into several stages. The Cambrian is divided into several regional faunal stages of which the Russian-Kazakhian system is most used in international parlance: *Most Russian paleontologists define the lower boundary of the Cambrian at the base of the Tommotian Stage, characterized by diversification and global distribution of organisms with mineral skeletons and the appearance of the first Archaeocyath bioherms.
The International Commission on Stratigraphy list the Cambrian period as beginning at 541 million years ago and ending at 485.4 million years ago. The lower boundary of the Cambrian was held to represent the first appearance of complex life, represented by trilobites; the recognition of small shelly fossils before the first trilobites, Ediacara biota earlier, led to calls for a more defined base to the Cambrian period. After decades of careful consideration, a continuous sedimentary sequence at Fortune Head, Newfoundland was settled upon as a formal base of the Cambrian period, to be correlated worldwide by the earliest appearance of Treptichnus pedum. Discovery of this fossil a few metres below the GSSP led to the refinement of this statement, it is the T. pedum ichnofossil assemblage, now formally used to correlate the base of the Cambrian. This formal designation allowed radiometric dates to be obtained from samples across the globe that corresponded to the base of the Cambrian. Early dates of 570 million years ago gained favour, though the methods used to obtain this number are now considered to be unsuitable and inaccurate.
A more precise date using modern radiometric dating yield a date of 541 ± 0.3 million years ago. The ash horizon in Oman from which this date was recovered corresponds to a marked fall in the abundance of carbon-13 that correlates to equivalent excursions elsewhere in the world, to the disappearance of distinctive Ediacaran fossils. There are arguments that the dated horizon in Oman does not correspond to the Ediacaran-Cambrian boundary, but represents a facies change from marine to evaporite-dominated strata — which w
The Permian is a geologic period and system which spans 47 million years from the end of the Carboniferous Period 298.9 million years ago, to the beginning of the Triassic period 251.902 Mya. It is the last period of the Paleozoic era; the concept of the Permian was introduced in 1841 by geologist Sir Roderick Murchison, who named it after the city of Perm. The Permian witnessed the diversification of the early amniotes into the ancestral groups of the mammals, turtles and archosaurs; the world at the time was dominated by two continents known as Pangaea and Siberia, surrounded by a global ocean called Panthalassa. The Carboniferous rainforest collapse left behind vast regions of desert within the continental interior. Amniotes, who could better cope with these drier conditions, rose to dominance in place of their amphibian ancestors; the Permian ended with the Permian–Triassic extinction event, the largest mass extinction in Earth's history, in which nearly 96% of marine species and 70% of terrestrial species died out.
It would take well into the Triassic for life to recover from this catastrophe. Recovery from the Permian–Triassic extinction event was protracted; the term "Permian" was introduced into geology in 1841 by Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Édouard de Verneuil; the region now lies in the Perm Krai of Russia. Official ICS 2017 subdivisions of the Permian System from most recent to most ancient rock layers are: Lopingian epoch Changhsingian Wuchiapingian Others: Waiitian Makabewan Ochoan Guadalupian epoch Capitanian stage Wordian stage Roadian stage Others: Kazanian or Maokovian Braxtonian stage Cisuralian epoch Kungurian stage Artinskian stage Sakmarian stage Asselian stage Others: Telfordian Mangapirian Sea levels in the Permian remained low, near-shore environments were reduced as all major landmasses collected into a single continent—Pangaea; this could have in part caused the widespread extinctions of marine species at the end of the period by reducing shallow coastal areas preferred by many marine organisms.
During the Permian, all the Earth's major landmasses 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, 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 the Paleo-Tethys Ocean 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 landmass interiors experience climates with extreme variations of heat and cold and monsoon conditions with 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 a wetter environment. The first modern trees appeared in the Permian. Three general areas are noted for their extensive Permian deposits—the Ural Mountains 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 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 in an ice age. Glaciers receded around the mid-Permian period as the climate 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 and brachiopods. Fossilized shells of two kinds of invertebrates are used to identify Permian strata and correlate them between sites: fusulinids, a kind of shelled amoeba-like protist, one of the foraminiferans, 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 and various types of tetrapods; the period saw a massive desert covering the interior of Pangaea.
The warm zone spread in the northern hemisphere. 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 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