Interval of geologic time, 490–443 million years ago, the second oldest period of the Paleozoic Era.It follows the Cambrian and precedes the Silurian. During the Ordovician, many of the landmasses were aligned in the tropics. Life was dominated by marine invertebrates, but some forms of land plants may have appeared during the middle of the period. Spores suggesting a tropical terrestrial environment have been found in rocks of that age.
* * *Introductionin geologic time, the second period of the Paleozoic Era. It began 488.3 million years ago, following the Cambrian Period, and ended 443.7 million years ago, when the Silurian Period began. Ordovician rocks have the distinction of occurring at the highest elevation on Earth—the top of Mount Everest (Everest, Mount).The Ordovician Period ushered in significant changes in plate tectonics, climate, and biological systems. Rapid seafloor spreading (seafloor spreading hypothesis) at oceanic ridges (ocean) fostered some of the highest global sea levels in the Phanerozoic Eon. As a result, continents were flooded to an unprecedented level, with North America almost entirely underwater at times. These seas deposited widespread blankets of sediment that preserved the extraordinarily abundant fossil remains of marine animals. Numerical models of the Ordovician atmosphere estimate that levels of carbon dioxide were several times higher than today. This would have created warm climates from the Equator to the poles; however, extensive glaciation (glacier) did occur for a brief time over much of the Southern Hemisphere at the end of the period.The Ordovician Period was also characterized by the intense diversification (an increase in the number of species) of marine animal life in what became known as the Ordovician radiation. This event precipitated the appearance of almost every modern phylum (a group of organisms having the same body plan) of marine invertebrate by the end of the period, as well as the rise of fish. Ordovician seas were filled with a diverse assemblage of invertebrates, dominated by brachiopods ( lamp shells), bryozoans (moss animals (moss animal)), trilobites (trilobite), mollusks (mollusk), echinoderms (echinoderm) (a group of spiny-skinned marine invertebrates), and graptolites (graptolite) (small, colonial, planktonic animals). On land the first plants appeared, as well as possibly the first invasion of terrestrial arthropods (arthropod). The end of the Ordovician was heralded by a mass extinction, the second largest in Earth history. The largest mass extinction took place at the end of the Permian Period and resulted in the loss of about 90 percent of then-existing species.The Ordovician was demarcated in the late 19th century as a compromise in a dispute over the boundaries of the Cambrian and Silurian systems. Studying the rock succession from northwest to southeast within Wales, English geologist Adam Sedgwick (Sedgwick, Adam) named the Cambrian System in 1835. At the same time and working in the opposite direction, Scottish geologist Roderick Murchison (Murchison, Sir Roderick Impey) named the Silurian System. Both geologists expanded their systems until they overlapped, triggering a scientific feud. English geologist Charles Lapworth (Lapworth, Charles) proposed the Ordovician System (named for an ancient Celtic tribe of northern Wales called the Ordovices) in 1879 to define the disputed overlapping interval. Lapworth's proposal was resisted in Britain into the 1890s and, despite subsequent widespread international usage, was not officially adopted there until 1960.The Ordovician is divided into three epochs: Early Ordovician (488.3–471.8 million years ago), Middle Ordovician (471.8–460.9 million years ago), and Late Ordovician (460.9–443.7 million years ago).The Ordovician environmentDuring the Ordovician Period, four major continents (continent) were present and separated by three major oceans. Although the positions of these continents are frequently updated with new evidence, current understanding of their position is based on paleomagnetic evidence, fossil markers, and climatically sensitive sediments, such as evaporite minerals. The craton (stable interior portion of a continent) of Laurentia—made up of most of present-day North America, Greenland, and part of Scotland—straddled the Equator and was rotated approximately 45° clockwise from its present orientation. The craton of Siberia-Kazakhstan lay east of Laurentia, along and slightly north of the Equator. The Iapetus Ocean separated these two landmasses on the south from the Baltica craton, which included present-day Scandinavia and north-central Europe. The microcontinent of Avalonia—made up of England, New England, and maritime Canada—was positioned to the west of Baltica and also faced Laurentia across the Iapetus Ocean. The Paleotethys Sea separated England, Baltica, and Kazakhstan from the supercontinent of Gondwana, which consisted of Africa, South America, India, Arabia, China, Australia, Antarctica, Western Europe, the southeastern United States, and the Yucatán Peninsula of Mexico. This immense supercontinent straddled both the South Pole, located then in what is now northwest Africa, and the Equator, which then crossed present-day Australia and Antarctica. In this position, Africa and South America were rotated nearly 180° from their present orientation. A single body of water, the Panthalassic Ocean, covered almost the entire Northern Hemisphere and was as wide at the Equator as the modern Pacific Ocean.Following the breakup of the supercontinent Rodinia during the Cambrian Period, the rate of seafloor spreading (seafloor spreading hypothesis) reached a maximum during the Ordovician Period. Tall oceanic ridges (ocean) produced by this activity raised the average elevation of the seafloor and flooded parts of many continents, creating vast shallow seas within their interiors and at their margins. During the Early and Late Ordovician epochs, almost all of North America was submerged under these epicontinental seas.Sea level fluctuated continuously throughout the Ordovician. At the broadest scale, sea level was highest during the Early and Late Ordovician epochs and may have been up to 200 metres (about 660 feet) lower during the Middle Ordovician Epoch. Numerous shorter-term fluctuations were superimposed on this broad rise and fall, with each fluctuation typically lasting one to five million years. Additional minor fluctuations of a few metres or less, occurring on the order of tens to hundreds of thousands of years, have also been identified. The causes of all of these changes in sea level are difficult to identify. Some may have been driven by variations in the rates of plate motion, some by glaciation (glacier), some by local tectonic uplift or subsidence, and some by changes in groundwater storage capacity. The end of the Ordovician Period is marked by a pronounced fall in sea level of nearly 160 metres (525 feet), which was triggered by the rapid expansion of continental ice sheets on Gondwana.Although it is impossible to observe Ordovician oceanic currents (ocean current) directly, major circulation patterns can be inferred from basic oceanographic principles. Circulation within the Panthalassic Ocean was unimpeded by continents and followed a relatively simple zonal system, with westward circumpolar flow north of 60° N. The northward projections of Laurentia and Gondwana into the Panthalassic Ocean would have created two large clockwise-spinning gyres (eddies) between the Equator and 60° N. In the Southern Hemisphere, Gondwana blocked this zonal flow by spanning the area from the South Pole to the Equator. Smaller gyres were set up within the Paleotethys Sea and Iapetus Ocean, with generally counterclockwise flow south of 60° S and clockwise flow between the Equator and 60° S. The tectonic movements of small continents and oceanic islands considerably modified these relatively simple gyres. Monsoonal (monsoon) circulation, characterized by seasonal alterations in ocean currents, would have been fostered in subtropical latitudes, particularly along the margins of Gondwana.Upwelling (marine ecosystem) (the movement of colder, nutrient-rich water from the depths to the surface) would have developed along the west coasts of continents, along the Equator and 60° S. Unlike the present time, in which the water in the deep ocean is generated from the sinking of cold, dense water at the poles, bottom water in the Ordovician would have originated in the warm, saline conditions of tropical epicontinental seas. Relatively rapid evaporation in these warm locations would have increased the density of a volume of water because of the high concentrations of dissolved salts. Similar conditions also existed during the Cretaceous Period and led in both cases to sluggish ocean circulation and resulting anoxia (lack of dissolved oxygen) in deep marine environments. The widespread black shales (shale) found in deepwater settings in the Ordovician are evidence of anoxic conditions. Only at the very end of Ordovician times during the extended glacial period would ocean circulation patterns accelerate. Chilled by continental glaciation, surface waters would have descended and displaced the warmer, deep ocean water. Because the sinking of cold polar water occurs at a faster rate than the sinking of warm, saline tropical water, the strength of cold ocean currents, and thus upwelling, would have increased.Following the breakup of the Rodinia supercontinent in the Cambrian, extensive oceanic ridges were established, circling the globe and creating the Paleotethys Sea and Iapetus and Panthalassic oceans. The production of oceanic lithosphere (Earth) at these ridges was accommodated through its destruction at subduction zones (subduction zone). The Panthalassic Ocean was apparently bordered on all sides by subduction zones, much like the Pacific Ocean is today. A subduction zone also separated Laurentia from both Siberia and Baltica.All of the major tectonic plates were in motion during the Ordovician Period. Laurentia gradually rotated counterclockwise as Siberia-Kazakhstan approached it from the east. Baltica drew toward Laurentia from the southeast, gradually closing the Iapetus Ocean and ultimately colliding with Laurentia in the Silurian Period. The Paleotethys Sea also gradually narrowed as a subduction zone on the west side of Gondwana consumed the ocean floor. Gondwana itself underwent a gradual clockwise rotation in the Ordovician, bringing Africa over the South Pole and Australia and North China across the Equator and into the Northern Hemisphere.Ordovician volcanism (volcano) was extensive and was primarily generated at subduction zones and oceanic ridges. Globally, volcanism appears to have peaked twice during the early Paleozoic Era, first near the end of the Cambrian Period through earliest parts of the Ordovician Period and later within the Middle to Late Ordovician epochs. The progressive closing of ocean basins as continents collided during the Middle Paleozoic is thought to reflect an overall slowing of rates of plate motion and therefore a slowdown in volcanicity, which is observed after the Ordovician. Ordovician volcanic deposits produced at subduction zones consist mostly of ashes as well as basaltic (basalt) to andesitic (andesite) flows, much as seen in modern subduction zones. Other Ordovician volcanic deposits record eruptions in extensional basins (rift zones) and are characterized by thick accumulations of basalt. Volcanic rocks of Ordovician origin are particularly common in Great Britain, Kazakhstan, the Baltic region and Scandinavia, eastern North America, and Argentina.Unique mineralogical and geochemical features of many Ordovician volcanic ashes allow them to be correlated over long distances. One such ash bed, the Millbrig K- bentonite of eastern North America, has been correlated to the “Big Bentonite” of Scandinavia, which was separated from North America in the Ordovician by the 1,000-km-wide (620-mile-wide) Iapetus Ocean. Reconstruction of the volume of this eruption suggests that over 1,000 cubic km (about 240 cubic miles) of ash was erupted, making it one of the largest recorded ashfalls in Earth history. In contrast, the eruption of Mount St. Helens (Saint Helens, Mount) in 1980 produced a mere 0.2 cubic km (about 0.05 cubic mile) of ash.Orogenic (orogeny) (mountain-building) belts formed in the Ordovician wherever plates converged—at subduction zones (subduction zone) and at collisions between continents and terranes, such as microplates (smaller fragments of continental plates), oceanic arcs (chains of volcanic islands), and oceanic plateaus. Subduction zones have been recognized along the Panthalassic margin of Tasmania, Trans-Antarctica, western South America, western North America, Ellesmere Island, Mongolia, Kazakhstan, and the Qin (Tsinling)-Qilian ranges in China. Collisions with terranes are also well known. One well-studied example is the Taconic orogeny, which occurred during the Middle and Late Ordovician epochs in the eastern United States. This event includes at least three separate collisional events from Maine to Alabama. The Taconic orogeny created a series of deep basins along the eastern edge of Laurentia, some of which are now filled with over 3,000 metres (about 9,900 feet) of sedimentary (sedimentary rock) rock. The thick accumulation of sediment filling one of these basins in present-day New York and Pennsylvania is known as the Queenston Delta.Paleoclimate (paleoclimatology)Numerical climate models as well as carbon isotope measurements from preserved Ordovician soils suggest that atmospheric levels of carbon dioxide during the period were 14–16 times higher than today. These high levels were driven by widespread volcanic activity, which would have released large volumes of carbon dioxide into the atmosphere. The extensive flooding of continents due to high sea levels, combined with the lack of widespread vegetation on land, would have suppressed the weathering of silicate (silicate mineral) rocks, a major mechanism for removing carbon dioxide from the atmosphere. In short, the rate at which carbon dioxide was added to the atmosphere increased during the Ordovician Period, whereas its rate of removal decreased.Because of the greenhouse effect, high levels of atmospheric carbon dioxide would have caused temperatures to rise everywhere from the Equator to the poles. Clear evidence of warm temperatures in the tropics can be seen in the extensive Ordovician limestone deposits with features similar to those found in modern tropical carbonate areas, such as the coral reefs of the Bahamas. Other evidence suggesting warm low-latitude climates in the Ordovician include bauxite deposits in Kazakhstan and evaporite minerals in North America, Australia, China, Kazakhstan, and Siberia.Warm tropical regions would have fostered the development of tropical cyclones (tropical cyclone); there is a rich record of such storms in the Ordovician. In subtropical areas of North America, sedimentary (sedimentary rock) deposits from Ordovician continental shelves commonly occur in alternating beds of shale and either sandstone or limestone. These sandstone and limestone beds display evidence of erosion at their bases, become finer-grained upward within the beds, and contain distinctive sedimentary structures known to form from the combination of strong currents and large waves. These storm-driven structures and processes also occur on the continental shelves of the present day. In addition, storm deposits known as tempestites can be quite common in Ordovician rocks. For example, an interval of Late Ordovician strata near Cincinnati, Ohio, is 65 metres (213 feet) thick. This interval represents an estimated 1.5 million years of deposition and contains over 300 tempestites. Because the erosion of the seafloor during the early phases of a hurricane has the potential to remove previously deposited tempestites, the interval near Cincinnati suggests that storms left a preserved sedimentary record at least once every 5,000 years during the period.Despite high carbon dioxide levels in the Ordovician Period, evidence of cooler climates in higher latitudes is seen by the presence of unweathered mica in sedimentary rocks from North Africa, central and southern Europe, and much of South America. Their abundance, coupled with the presence of faunas interpreted as cold-water forms, as well as paleomagnetic evidence, suggests that northwestern Africa was located over the South Pole. Furthermore, glacial deposits dating back to the Ordovician are also known from much of Africa, southern Asia, and parts of Europe. It had been thought the South Pole was glaciated for much of the Ordovician, but more-recent dating of these glacial deposits, plus isotopic evidence, suggests that major continental glaciation was limited to the last half-million years of the Late Ordovician Epoch and a shorter portion of the Early Silurian Epoch.How continental glaciation could have formed when carbon dioxide levels were so high has been a paradox. Recently it has been proposed that the terminal Ordovician glaciation was triggered through a combination of the placement of Africa over the South Pole and a short-lived drawdown of atmospheric carbon dioxide. This drawdown was favoured by declining volcanism during the Ordovician, which would have introduced progressively less carbon dioxide into the atmosphere. Increasing orogeny and uplift during later Ordovician times would have accelerated the removal of carbon dioxide from the atmosphere through the weathering of silicate rocks. Once snow began to accumulate in North Africa, the increased albedo (surface reflectance) of the snow would have fostered lower temperatures and increased the accumulation of greater amounts of snow in Gondwana, thereby bringing about a glacial period. The cooling of the oceans may also have fostered the increased productivity of photosynthetic (photosynthesis) organisms, moving nutrients from the depths to the surface through the process of upwelling. As the populations of photosynthesizers increased, additional carbon dioxide was removed from the atmosphere. At the end of this glacial period, it is now believed, as the production of carbon dioxide from volcanoes continued, the proportion of atmospheric carbon dioxide removal due to the weathering of silicate rocks declined. The silicate rocks of Gondwana had no access to the atmosphere, because many were covered by thick layers of glacial ice.Ordovician lifeMarine organismsMicrofossils and planktonOrdovician seas were characterized by a rich and diverse assemblage of species. Calcified microbial mats, known as stromatolites (stromatolite), are found in Ordovician rocks, although they are not as common there as in strata from the Cambrian Period and the Proterozoic Eon. Chitinozoans or acritarchs, microfossils with a hollow cavity and organic walls, represent the phytoplankton (small, free-floating, photosynthetic organisms). Ordovician foraminiferans (foraminiferan) include both agglutinated (glued or cemented) and calcareous (containing calcium carbonate) forms, including the first fusulinids (single-celled, amoeba-like organisms with complex shells). The siliceous radiolarians (radiolarian) (spherically symmetrical organisms with fine, sculptured skeletons) that lived as zooplankton (groups of small animals that feed on phytoplankton) are also found in Ordovician rocks.Invertebrates (invertebrate)Invertebrate life became increasingly diverse and complex through the Ordovician. Both calcareous and siliceous sponges (sponge) are known; among other types, the stromatoporoids first appeared in the Ordovician. Tabulata (platform) and rugosa (horn coral) (horn) corals (coral) also first appeared in the Ordovician, the solitary or horn corals being especially distinctive. Bryozoans (moss animal) (moss animals) and brachiopods (lamp shells) (lamp shells) were a dominant component of many assemblages. Mollusks (mollusk) were also common and included the gastropods (gastropod), monoplacophorans (limpet-shaped, segmented mollusks), bivalves (bivalve), cephalopods (cephalopod), chitons (chiton), scaphopods (tusk shell) (tusk shells), and rostroconchs (single-shelled mollusks).The fossil record of Ordovician annelids (annelid) (worms and leeches) consists chiefly of small, calcareous tubes, tiny jaws made up of phosphate material, and trace fossils. Trilobites (trilobite) are common and diverse in Ordovician strata but do not dominate assemblages as they did in the Cambrian Period. Ordovician arthropods (arthropod) are also represented by the ostracods (crustacean) (tiny crustaceans) as well as by much rarer forms such as branchiopods, barnacles (barnacle), phyllocarid (“leaf”) shrimp, aglaspids (primitive horseshoe crabs), and eurypterids (eurypterid) (sea scorpions).Echinoderms (echinoderm) reached their peak diversity of 20 classes during the Ordovician, with crinoids (crinoid) (a group related to present-day sea lilies and feather stars), cystoids (spherical, stalked echinoderms), asteroids (starfishes), edrioasteroids (sessile, plate-covered echinoderms), and homalozoans (asymmetrical echinoderms) being the most common. Graptolites (graptolite) (small, colonial, planktonic animals) and conodonts (conodont) (toothlike remains of primitive chordates (chordate)) are among the most important fossils in the Ordovician for correlating, or demonstrating age equivalence between, different layers of rock.Early fishes (fish)Until the discovery in 1999 of Early Cambrian vertebrates in South China, the oldest generally accepted vertebrates (vertebrate) were known from the Ordovician. The first examples were two genera of primitive fishes (fish), described by American geologist Charles Doolittle Walcott in the late 19th century, from the Upper Ordovician Harding Sandstone of Colorado. Subsequently, fish fragments have been described from the later part of the Early Ordovician Epoch to the early Middle Ordovician of Australia and from the first part of the Late Ordovician of Bolivia. Unconfirmed fish remains have also been reported from the Late Cambrian to Early Ordovician epochs of Spitsbergen, Greenland, Germany, and many localities in North America.The complete morphology of these fishes is unknown, and only fragments and individual scales have been found. Most specimens are made up of individual bony plates covered by ridges and nodes. Enough fragmentary material has been found, however, to allow a reconstruction of these animals. Their heads appear to have been covered by densely interlocking plates and their bodies by rows of thinner plates. The tail structure is unknown.All of these fossils are interpreted to be agnathans (agnathan), or jawless fishes. The environment in which they lived continues to be disputed, although the interpreted environment at all of the localities is similar. At all three locations, sediments were laid down in very shallow marine to marginal marine environments, possibly with low salinity as found in lagoons and estuaries. The fishes are interpreted to have fed on organic matter on the seafloor.ReefsReefs are known throughout the Ordovician Period, although the types of organisms that built them changed over time. Most Ordovician reefs are rather small, typically up to 5 metres (16 feet) thick, though some are up to 20 to 30 metres (65 to 100 feet) thick. Most Ordovician reefs were built on continental shelves or in shallow seas within the continental interior, although some were built at the edge of continental shelves or on the continental slopes. Earliest Ordovician reefs were most commonly mud mounds built by microbes, algae, and sponges. Microbial mounds are also known from the latest Ordovician times. Stromatoporoid and coral patch reefs first appeared in the Middle Ordovician Epoch and persisted after the close of the Ordovician Period, whereas the first appearance of bryozoan mounds and red algal reefs occurred during the Middle Ordovician Epoch. Large carbonate mud mounds containing a diverse suite of reef-building organisms first appeared in the latest part of the Early Ordovician Epoch and are notable in that they attained thicknesses exceeding 250 metres (820 feet).Terrestrial organismsEarliest land plants (plant)The Ordovician contains the oldest generally accepted remains of land plants, in the form of cuticle fragments and spores (spore). Fragments of cuticle lack stomata and other structures and have eluded identification. The spores are more diagnostic, and two types are known. Cryptospores (sporelike structures predating land plants) first appeared in the early part of the Middle Ordovician Epoch and rapidly spread to all continents. Through this time their morphology changed little and shows minute evolution.Trilete spores, which divide via meiosis to form a tetrad of cells, first appeared in the late Late Ordovician and form a rare, geographically isolated component in cryptospore assemblages. Although spores are known from older rocks, cryptospore and trilete spores are thought to be the direct ancestors of land plants because of their size, gross morphology, and wall structure, their abundance in nonmarine and marginal marine deposits, and their association later in the geologic record with definite land plants. Cryptospores were limited to life in damp habitats. Their distribution over a wide range of latitudes suggests their adaptability to many environments, as opposed to the much more narrowly distributed triletes.These plant fossils suggest two phases of evolution. Cryptospores are thought to be the precursors of nonvascular plants (plant) of the liverwort or bryophyte grade of organization. Triletes are thought to be the originators of vascular plants (plant) and may have been similar to the Late Silurian and Early Devonian rhyniophytes (Rhynie plant). Both forms are thought to have arisen from green algae. Although both the cryptospore- and trilete-producing plants gave rise to other plants, both are now thought to be extinct.AnimalsAlthough no fossils of land animals are known from the Ordovician, burrows from the Late Ordovician of Pennsylvania have been interpreted as produced by animals similar to millipedes (millipede). A millipede-like organism is inferred because the burrows occur in discrete size classes, are bilaterally symmetrical, and were backfilled by the burrowing organism. The burrows are found in a preserved soil and are associated with carbonate concretions that precipitated within the soil, indicating that the burrows were produced at the time of soil formation. The presence of plants and possibly arthropods suggests that Ordovician terrestrial ecosystems may have been more extensive and complex than generally thought.Ordovician radiationDuring the Ordovician Period, life diversified to an unprecedented degree, undergoing a fourfold increase in the number of genera. This unique period, known as the Ordovician radiation, unfolded over tens of millions of years and produced organisms that would dominate marine ecosystems for the remainder of the Paleozoic Era. The Ordovician radiation was an extension of the Cambrian explosion, an event during which all modern marine phyla appeared (with the exception of the bryozoans, which emerged during the Ordovician). The Ordovician continued this diversification at lower levels of taxonomy and saw a rapid increase in the amount of habitats and ecological niches exploited by living things, as well as an increase in the complexity of biological communities.The number of marine genera in most of the Early Ordovician Epoch was comparable to that seen in the Cambrian Period and had comparable rates of species turnover or extinction. By the latest age of the Early Ordovician Epoch, trilobites and other organisms dominant in the Cambrian were replaced by a wide range of other marine invertebrates, including corals, bryozoans, brachiopods, mollusks, echinoderms, graptolites (graptolite), and conodonts. One theory posits that diversification reached a peak by the first age of the Late Ordovician Epoch, with minor fluctuations. On the other hand, it has also been argued that this early Late Ordovician “peak” only represents a higher-quality fossil record than that of later Ordovician times. When this difference is accounted for, diversity is seen to rise to a plateau by the Middle Ordovician, after which it changes little.The timing of diversification differs for each group of organisms and on each of the Ordovician continents. For example, graptolites reached their peak diversity in the Early Ordovician Epoch, whereas gastropods continued to diversify steadily through the entire Ordovician Period. Similarly, overall diversity on the cratons of Laurentia and Baltica peaked in the early Late Ordovician Epoch, whereas diversity peaked in South China in the Early Ordovician Epoch. These intercontinental differences suggest that global diversification was driven by changes unique to each continent rather than by a single global factor.Exploitation of habitatsThe Ordovician radiation began in shallow marine environments and proceeded into deeper water. Newer fauna intermingled with older Cambrian fauna, which was primarily made up of various trilobites (trilobite) and inarticulate (unjointed) brachiopods (lamp shells) living in a wide range of environments between the shore and the continental slope. In the Early Ordovician Epoch, articulate (jointed) brachiopods, gastropods (gastropod), and cephalopods appeared in shallow-water habitats as inarticulate brachiopods and trilobites declined in those habitats. Through the remainder of the Ordovician Period, articulate brachiopods and gastropods continued to spread farther offshore as trilobites and inarticulate brachiopods became rarer in all but deepwater habitats. Finally, in the Late Ordovician Epoch, bivalve communities appeared in shallow-water habitats and displaced the brachiopod-gastropod communities offshore.Much of the increase in diversity occurring during the Ordovician Period took place within biological communities formed during the Cambrian Period. New species made use of unexploited niches within these communities. Another large portion of this new diversity came from increased provinciality—that is, the differences in the species present between one continent and another. Since most species did not expand beyond their own local regions, the species assemblages of many areas were unique, and few species were distributed globally. Diversity was also increased because of the expansion of life into new habitats not present in the Cambrian, such as reefs, hardgrounds, bryozoan thickets, and crinoid gardens.Ordovician communities were ecologically more complicated than Cambrian ones. The Ordovician saw the rise of several new life habits, including deep-deposit feeders, mobile epifaunal (superficially attached) carnivores, and pelagic (open-water) carnivores. In contrast to Cambrian communities that lived very close to the sediment surface, Ordovician communities also grew up to 50 cm (1.5 feet) above the seafloor and established distinct tiers, or levels, similar to those present in modern forests. Also, invertebrates burrowed into the seafloor more intensely during the Ordovician Period than in the Cambrian Period, reaching depths of up to 1 metre (3 feet) below the seafloor.CausesThe causes of the Ordovician radiation remain unclear. One view points to the Middle Ordovician fall in sea level, although this event has also been coupled to a global drop in diversity. Another view posits that biological interactions or an inherently higher rate of speciation in some groups fostered the diversification. Others have noted the correlation between the Ordovician diversification and the increase in global orogenic and volcanic activity. Indeed, on continents affected by orogenic activity, diversity proceeded at a faster pace than on other continents, suggesting that an increase in the supply of some nutrients, such as phosphorous and potassium, during the process of uplift may have fueled the diversification.Extinctions (extinction)Mass extinction at the end of the OrdovicianThe Ordovician Period was terminated by an interval of mass extinction. This extinction interval ranks second in severity to the one that occurred at the boundary between the Permian (Permian Period) and Triassic (Triassic Period) periods in terms of the percentage of marine families affected, and it was almost twice as severe as the extinction event that occurred at the end of the Cretaceous Period, which is famous for bringing an end to the dinosaurs. An estimated 85 percent of all Ordovician species became extinct during the end-Ordovician extinction in the two-million-year-long Hirnantian Age and the subsequent Rhuddanian Age of the Silurian Period.Brachiopods (lamp shells) display the effects of this extinction well. Laurentian brachiopods were hit hard, particularly those that lived in the broad and shallow seas both within and near the continent. Many of these brachiopods were endemic (confined to a particular region) to Laurentia, as opposed to the more cosmopolitan (globally distributed) forms that lived at the edges of the continent. Following the extinction, Laurentian seas were repopulated with brachiopod genera previously found only on other continents. As a result, Silurian brachiopods were far more widely distributed than their Ordovician predecessors. Other groups of organisms—including conodonts, acritarchs (a catchall group of various small microfossils), bryozoans, and trilobites—that showed this pattern of regional, but not global, distribution were similarly affected by this extinction event. Despite the intensity of the extinction and the loss of many endemic species, Silurian ecosystems were remarkably similar to those in the Ordovician.The extinction appears to have occurred in several phases. An early phase affecting graptolites, brachiopods, and trilobites occurred prior to the end of the Ordovician Period, before the major fall in sea level. A second phase of extinction occurred as sea levels fell because of the onset of glaciation over the African and South American portions of Gondwana. In many areas the interval of glaciation was accompanied by the invasion of cool-water brachiopod fauna, even into tropical latitudes, suggesting the onset of significant global cooling. A third phase of extinction occurred with the rise of sea level that took place during the Rhuddanian Age of the Silurian Period.The cause of the end-Ordovician extinction is generally attributed to two factors: the first wave of extinction may be related to rapid cooling at the end of the Ordovician Period, and the second phase is widely regarded as having been caused by the sea-level fall associated with the glaciation. The drop in sea level would have drained the large epicontinental seas and reduced the available habitat for organisms that favoured those settings. No concentration of iridium has been identified near the extinction that would suggest a bolide (meteoroid or asteroid) impact like the one identified at the boundary between the Cretaceous and Tertiary (Tertiary Period) periods.Regional extinctions within the OrdovicianIn addition to this mass extinction, smaller-scale or background extinctions occurred during the Ordovician Period. Most of these are poorly understood, but one that has been studied occurred in the eastern United States during the early Late Ordovician Epoch. This extinction involved a wide range of organisms in a wide variety of life habits, including brachiopods, corals, trilobites, echinoderms, and mollusks. Especially at risk were species that were restricted to the eastern United States. Many surviving species experienced a contraction in their geographic ranges and were driven out from the eastern regions to the western United States and Canada.This particular extinction is very closely timed with a major tectonic event that affected the North American region. The collision of a terrane with the eastern edge of the continent during the Taconic orogeny caused the continental margin to flex and the sedimentary basin to deepen. This deepening allowed cool, nutrient-rich, turbid water to spread from the Appalachians to Indiana and central Tennessee. These changes appear in the strata as a relatively rapid switch from tropical limestones with little shale to cool-water phosphatic limestones with abundant shale. Several new species adapted to these new conditions invaded the eastern United States at this time.As this sedimentary basin filled and the source of cool, nutrient-rich water was cut off, conditions gradually returned to those prior to the extinction. Many of the organisms that had retreated from the eastern United States returned to this region later in the Ordovician and into the Silurian.Faunal (faunal region) provincesOrdovician faunas were strikingly provincial, much more so than those in the Silurian Period. The main control on the distribution of Ordovician faunas was temperature. The planktonic graptolites are commonly divided into the Pacific and Atlantic (or European) provinces. The Pacific Province corresponds to regions that were tropical during the Ordovician, such as North America, Siberia, Kazakhstan, and parts of China. The Atlantic Province is found in more-temperate regions of the Ordovician Period, including the British Isles, the Baltic region and Scandinavia, and northern Africa. Some modern regions, such as South America, straddled both of these provinces. Acritarchs follow a distribution similar to graptolites, with their faunas limited to tropical, polar, and intermediate regions in the Ordovician.Conodont provinces include the North American Midcontinent and the North Atlantic provinces. The North American Midcontinent Province is also found in Scotland, Siberia, and temporarily in parts of the Baltic region and Scandinavia. The North Atlantic Province is known from the Baltic region and Scandinavia, the British Isles, and the margins of North America. Australian faunas bear similarities to both of these provinces yet have some distinctive elements, suggesting that they may represent another province.Provinces have also been recognized within single continents for other groups. Late Ordovician corals from Laurentia have been divided into a Red River–Stony Mountain Province in the western United States and Canada and a Richmond Province, extending in a narrow belt from Ontario to Michigan to central Tennessee. In addition to all these temperature-controlled provinces, Ordovician faunas are known to change with water depth, nutrient levels, and sediment type. This regional isolation of many faunas contributed to the intense pace of the Ordovician radiation, but it has also made the global correlation of Ordovician rocks difficult.Ordovician geologySignificant geologic eventsA wide variety of rock types is found in the Ordovician System. limestone and dolomite are the predominant rock types from the tropical epicontinental sea regions in the Ordovician, such as Laurentia, Siberia, Kazakhstan, and South China. These limestones formed principally from the accumulation of calcareous skeletons of organisms, from carbonate mud, and from ooids (a type of seafloor sand) and peloids (small sand-sized grains formed from fecal matter, algae, and mud). Reefs are also commonly found in these limestone regions.Ordovician evaporite deposits, such as anhydrite, are found in areas that were situated in arid belts near 30° N and 30° S. Anhydrite is known from the Early and Late Ordovician epochs of Laurentia and Kazakhstan.Ordovician siliciclastic (made from broken parts of silica rocks) rocks can be divided into coarse-grained and fine-grained types. Relatively coarse-grained siliciclastic rocks such as sandstones (sandstone) and conglomerates (conglomerate) tended to accumulate in high latitudes where significant carbonate accumulation was precluded and in low latitudes where orogenic uplift supplied abundant sediment. In North America, thick accumulations of coarse siliciclastic sediment can be found in the Appalachians. The Ordovician Taconic orogeny supplied voluminous amounts of sediment to the Queenston Delta, which spread from the Appalachians westward to Ontario and Ohio. Many of these sandstones are resistant to weathering and form the tops of ridges in the Ridge and Valley Province (Ridge and Valley) of the Appalachians. Additional thick sandstones are known from the Middle Ordovician Saint Peter Sandstone, which accumulated as a vast coastal sand dune system spread across much of North America. On other continents, thick successions of sandstones may be found in the Gondwana-associated regions of Antarctica, Australia, Great Britain, southern Europe, South America, and Africa. Coarse siliciclastic rocks also occur as glacial deposits in the late Late Ordovician of Africa and South America. These deposits consist of tilloids (gravity-driven sediments) and dropstones (rocks dropped into fine-grained sediments as they melted out of floating glacial ice).Fine-grained siliciclastic rocks, represented by mudstones (mudstone) and shales (shale), accumulated for the most part in deepwater settings of the Ordovician Period. As a result, thick accumulations of shale tend to occur around the margins of Ordovician continents and in the deepwater portions of epicontinental seas. Finely laminated black shales record deposition in anoxic water, whereas burrowed gray or red mudstones indicate oxygenated conditions. Bedded chert (chert and flint) is sometimes associated with these basinal shales, as in China.Volcanic ashes are known from many regions in the Ordovician Period, attesting to the widespread volcanism of the times. These ashes are commonly weathered to form K-bentonites (a type of clay useful in correlation because its particles are easily dated) that are interbedded with other rock types, such as limestone, shale, and sandstone. Other volcanic and igneous rocks are found principally near Ordovician subduction zones and back-arc basins (marine basins that form behind subduction zones (subduction zone)), as in the British Isles, Kazakhstan, and China. Rock types include volcanic breccia, tuff, basalt, andesite, and rhyolite.Economic significance of Ordovician depositsSeveral types of Ordovician deposits are economically important. petroleum and natural gas have been extracted from Middle and Upper Ordovician carbonates from the United States and Canada. Oil was first produced from a well that was drilled into Ordovician limestone in western Pennsylvania. Oil and natural gas continue to be produced from Middle and Upper Ordovician limestone and dolomite in the United States and Canada. Unusual oil-rich shales called kukersites are major sources of petroleum in Estonia. Limestone deposits in the Mississippi valley, central Tennessee, and the southern Appalachians are an important source of lead and zinc. Limestone itself is quarried in many places as a building stone, for use in roads, and for use in reducing sulfur emissions from coal-burning power plants. Phosphates have been mined for fertilizer from Ordovician limestones, particularly where weathering has concentrated the phosphate, as in central Tennessee. Ordovician sandstones, such as the Saint Peter Sandstone in the Midwestern United States, are important for glassmaking. Ordovician shales have been used to manufacture bricks. Black cephalopod-bearing limestones from Africa are widely sold as decorative pieces and tabletops.Major subdivisions of the Ordovician SystemThe rocks that originated during the Ordovician compose the Ordovician System. The Ordovician Period is divided into seven stages—two each in the early and middle epochs and three in the late epoch. Of these, only three stages are named. The intense provincialism of Ordovician faunas has long hindered the establishment of a single global succession of stages and zones. Existing names and descriptions useful at regional levels are inconsistent and do not correlate well at the global level.The International Commission on Stratigraphy (ICS) has formed several working groups to define new stages based on a more accurate global correlation of stratigraphic and fossil markers. Thus far, the Tremadocian (Tremadocian Stage) (Stage 1), Floian (Stage 2), Darriwilian (Stage 4), Sandbian (Stage 5), Katian (Stage 6), and Hirnantian (Stage 7) have been formally designated, and one unnamed stage remains. Both the Tremadocian and Floian stages date to the Early Ordovician Epoch, which occurred between 488.3 and 471.8 million years ago; Stage 3 and the Darriwilian Stage date to the Middle Ordovician Epoch, which occurred between 471.8 and 460.9 million years ago; and the Sandbian, Katian, and Hirnantian stages date to the Late Ordovician Epoch, which occurred between 460.9 and 443.7 million years ago.Correlation of Ordovician strataCorrelation, or the demonstration of the age equivalence, of strata in the Ordovician System has traditionally relied on fossils. Shelly fossils, such as brachiopods (lamp shells) (lamp shells) and trilobites (trilobite), have proved most useful for correlation within individual continents because of their tendency to be endemic. Other shelly groups, including bryozoans (moss animal) (moss animals), crinoids (crinoid), and corals (coral), have also been used for correlation but to a lesser extent. The Cambrian-Ordovician boundary is marked by the first appearance of the conodont Iapetognathus fluctivagus and the trilobites Jujuyaspis borealis and Symphysurina bulbosa.Graptolites (graptolite) (small, colonial, planktonic animals) and conodonts (conodont) (toothlike remains of primitive chordates (chordate)) are the most widely used organisms for the intercontinental correlation of Ordovician strata. Graptolites in particular are now used to define most of the new global Ordovician stage boundaries. Yet, even for this purpose, the effects of provinciality can limit their geographic ranges and thus their usefulness.More recently, packages of rocks bounded by unconformities (interruptions in the deposition of sedimentary rock) have been used for correlation within continents. The field of sequence stratigraphy recognizes these units, termed depositional sequences. Many of these units may have formed from changes in global sea level and may ultimately be useful for global correlation.Geochemical correlation of Ordovician rocks has also been successful. Broad correlations have been achieved with oxygen and carbon isotopes (isotope). Highly precise correlations of individual volcanic ash beds have been made using their major, minor, and trace element content.Steven M. HollandAdditional ReadingIntroductory treatments of the geology and paleontology of the Ordovician can be found in Steven M. Stanley, Earth and Life Through Time (1989); and Harold L. Levin, The Earth Through Time, 8th ed. (1996). A complete treatment of extinction events through geologic time, including the Ordovician mass extinction event, can be found in A. Hallam and P.B. Wignall, Mass Extinctions and Their Aftermath (1997).Steven M. Holland
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