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90-823: The Eramosa Member is a Silurian stratigraphic unit of the Lockport Formation exposed along the Niagara Escarpment in Ontario and western New York State. In the late nineteenth century it was an important source of building stone in Hamilton , Ancaster and Waterdown , and in the late twentieth century quarries in a similar unit, also called the Eramosa, near Wiarton in the Bruce Peninsula , became an important source of dimension stone at

180-600: A dense rock equivalent volume of as much as 1,140 cubic kilometres (270 cu mi). Remarkably, this appears to have had little impact on life. There was vigorous tectonic activity along northwest margin of Gondwana during the Floian, 478 Ma, recorded in the Central Iberian Zone of Spain. The activity reached as far as Turkey by the end of Ordovician. The opposite margin of Gondwana, in Australia, faced

270-603: A biogeographic affinity with Gondwana, and the Alborz margin of Gondwana was linked biogeographically to South China. Southeast Asia's fauna also maintained strong affinities to Gondwana's. North China was biogeographically connected to Laurentia and the Argentinian margin of Gondwana. A Celtic biogeographic province also existed, separate from the Laurentian and Baltican ones. However, tropical articulate brachiopods had

360-468: A burst of volcanic activity that deposited new silicate rocks, which draw CO 2 out of the air as they erode. Another possibility is that bryophytes and lichens, which colonized land in the middle to late Ordovician, may have increased weathering enough to draw down CO 2 levels. The drop in CO 2 selectively affected the shallow seas where most organisms lived. It has also been suggested that shielding of

450-590: A deep embayment between Siberia and the Central Mongolian terranes . Most of the terranes of central Asia were part of an equatorial archipelago whose geometry is poorly constrained by the available evidence. The period was one of extensive, widespread tectonism and volcanism. However, orogenesis (mountain-building) was not primarily due to continent-continent collisions. Instead, mountains arose along active continental margins during accretion of arc terranes or ribbon microcontinents. Accretion of new crust

540-639: A dispute between followers of Adam Sedgwick and Roderick Murchison , who were placing the same rock beds in North Wales in 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, and placed them in a system of their own. The Ordovician received international approval in 1960 (forty years after Lapworth's death), when it

630-607: A distinct band around the Earth, and that the breakup of the parent body may have formed a ring system for a period of about 40 million years, with frequent falling debris causing these craters. The Ordovician was a time of calcite sea geochemistry in which low-magnesium calcite was the primary inorganic marine precipitate of calcium carbonate . Carbonate hardgrounds were thus very common, along with calcitic ooids , calcitic cements, and invertebrate faunas with dominantly calcitic skeletons. Biogenic aragonite , like that composing

720-535: A diverse range of epibionts, including certain hederelloids as aforementioned. Photosymbiotic scleractinians made their first appearance during the Middle Silurian. Reef abundance was patchy; sometimes, fossils are frequent, but at other points, are virtually absent from the rock record. Ordovician The Ordovician ( / ɔːr d ə ˈ v ɪ ʃ i . ə n , - d oʊ -, - ˈ v ɪ ʃ ən / or-də- VISH -ee-ən, -⁠doh-, -⁠ VISH -ən )

810-411: A food web based on as-yet-undiscovered detritivores and grazers on micro-organisms. Millipedes from Cowie Formation such as Cowiedesmus and Pneumodesmus were considered as the oldest millipede from the middle Silurian at 428–430 million years ago, although the age of this formation is later reinterpreted to be from the early Devonian instead by some researchers. Regardless, Pneumodesmus

900-551: A fossil arthropod found at the site that shares similarities to Parioscorpio , an enigmatic arthropod known from the Waukesha biota site. In addition, a lobopodian with preserved patterning has been found. The exceptionally preserved biota are present in a stratum of rock about 8 m thick, which extends for around 16 km (9.9 mi) in the South Bruce Peninsula , Ontario . So-called "Eramosa marble"

990-464: A given landmass were severely affected. Tropical lifeforms were hit particularly hard in the first wave of extinction, while cool-water species were hit worst in the second pulse. Those species able to adapt to the changing conditions survived to fill the ecological niches left by the extinctions. For example, there is evidence the oceans became more deeply oxygenated during the glaciation, allowing unusual benthic organisms (Hirnantian fauna) to colonize

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1080-462: A high degree of development in relation to the age of its fossil remains. Fossils of this plant have been recorded in Australia, Canada, and China. Eohostimella heathana is an early, probably terrestrial, "plant" known from compression fossils of Early Silurian (Llandovery) age. The chemistry of its fossils is similar to that of fossilised vascular plants, rather than algae. Fossils that are considered as terrestrial animals are also known from

1170-434: A large ocean occupied most of the northern half of the globe. The high sea levels of the Silurian and the relatively flat land (with few significant mountain belts) resulted in a number of island chains, and 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

1260-423: A minor mass extinction and associated with rapid sea-level change. Each one leaves a similar signature in the geological record, both geochemically and biologically; pelagic (free-swimming) organisms were particularly hard hit, as were brachiopods , corals , and trilobites , and extinctions rarely occur in a rapid series of fast bursts. The climate fluctuations are best explained by a sequence of glaciations, but

1350-693: A more cosmopolitan distribution , with less diversity on different continents. During the Middle Ordovician, beta diversity began a significant decline as marine taxa began to disperse widely across space. Faunas become less provincial later in the Ordovician, partly due to the narrowing of the Iapetus Ocean, though they were still distinguishable into the late Ordovician. Trilobites in particular were rich and diverse, and experienced rapid diversification in many regions. Trilobites in

1440-728: A period known as the Ordovician meteor event . It has been theorized that this increase in impacts may originate from a ring system that formed around Earth at the time. In 2008, the ICS erected a formal international system of subdivisions for the Ordovician Period and System. Pre-existing Baltoscandic, British, Siberian, North American, Australian, Chinese, Mediterranean and North- Gondwanan regional stratigraphic schemes are also used locally. The Ordovician Period in Britain

1530-469: A second supercontinent known as Euramerica . When the proto-Europe collided with North America, the collision folded coastal sediments that had been accumulating since the Cambrian off the east coast of North America and the west coast of Europe. This event is the Caledonian orogeny , a spate of mountain building that stretched from New York State through conjoined Europe and Greenland to Norway. At

1620-541: A set of island arcs. The accretion of these arcs to the eastern margin of Gondwana was responsible for the Benambran Orogeny of eastern Australia. Subduction also took place along what is now Argentina (Famatinian Orogeny) at 450 Ma. This involved significant back arc rifting. The interior of Gondwana was tectonically quiet until the Triassic . Towards the end of the period, Gondwana began to drift across

1710-515: A time when most of the other resources of similar stone were depleted. Work in these quarries led to the discovery of exceptionally well preserved fossils (the Eramosa lagerstätte ). On the east Mountain at Hamilton, a well-developed cave system was discovered in the Eramosa and has now been designated as the Eramosa Karst Conservation Area. The term was first used for a stratigraphic unit by Williams (1915) who named

1800-789: Is a geologic period and system , the second of six periods of the Paleozoic Era , and the second of twelve periods of the Phanerozoic Eon . The Ordovician spans 41.6 million years from the end of the Cambrian Period 485.4 Ma (million years ago) to the start of the Silurian Period 443.8 Ma. The Ordovician, named after the Welsh tribe of the Ordovices , was defined by Charles Lapworth in 1879 to resolve

1890-537: Is actually a bituminous dolomite (see list ). Silurian The Silurian ( / s ɪ ˈ lj ʊər i . ən , s aɪ -/ sih- LURE -ee-ən, sy- ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago ( Mya ), to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the third and shortest period of

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1980-424: Is generally a minor component. The most characteristic feature is bituminous shaley interbeds. Probably the Eramosa was deposited in several related environments, most likely including shallow, restricted (and poorly oxygenated) marine waters. Among the fossils, perhaps the most spectacular are the eurypterids . Some species lived in fresh or brackish water, but Silurian examples were marine. There also seems to be

2070-489: Is marked by a sudden abundance of hard substrate trace fossils such as Trypanites , Palaeosabella , Petroxestes and Osprioneides . Bioerosion became an important process, particularly in the thick calcitic skeletons of corals, bryozoans and brachiopods, and on the extensive carbonate hardgrounds that appear in abundance at this time. Green algae were common in the Late Cambrian (perhaps earlier) and in

2160-561: Is still an important fossil as the oldest definitive evidence of spiracles to breath in the air. The first bony fish, the Osteichthyes , appeared, represented by the Acanthodians covered with bony scales. Fish reached considerable diversity and developed movable jaws , adapted from the supports of the front two or three gill arches. A diverse fauna of eurypterids (sea scorpions)—some of them several meters in length—prowled

2250-518: The Cambrian , reef -forming corals appeared in the early Ordovician, including the earliest known octocorals , corresponding to an increase in the stability of carbonate and thus a new abundance of calcifying animals. Brachiopods surged in diversity, adapting to almost every type of marine environment. Even after GOBE, there is evidence suggesting that Ordovician brachiopods maintained elevated rates of speciation. Molluscs , which appeared during

2340-455: The Devonian . The first land plants are known from this period. The Great Ordovician Biodiversification Event considerably increased the diversity of life. Fish , the world's first true vertebrates , continued to evolve, and those with jaws may have first appeared late in the period. About 100 times as many meteorites struck the Earth per year during the Ordovician compared with today in

2430-550: The Paleozoic Era, and the third of twelve periods of the Phanerozoic Eon. 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 a few million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when up to 60% of marine genera were wiped out. One important event in this period

2520-766: The Rheic Ocean between Gondwana and Avalonia. Avalonia collided with Baltica towards the end of Ordovician. Other geographic features of the Ordovician world included the Tornquist Sea , which separated Avalonia from Baltica; the Aegir Ocean, which separated Baltica from Siberia; and an oceanic area between Siberia, Baltica, and Gondwana which expanded to become the Paleoasian Ocean in Carboniferous time. The Mongol-Okhotsk Ocean formed

2610-580: The South Pole until they almost disappeared in the middle of Silurian. Layers of broken shells (called coquina ) provide strong evidence of a climate dominated by violent storms generated then as now by warm sea surfaces. The climate and carbon cycle appear to be rather unsettled during the Silurian, which had a higher frequency of isotopic excursions (indicative of climate fluctuations) than any other period. The Ireviken event , Mulde event , and Lau event each represent isotopic excursions following

2700-471: The endocerid cephalopods died out completely, except for possible rare Silurian forms. The Ordovician–Silurian extinction events may have been caused by an ice age that occurred at the end of the Ordovician Period, due to the expansion of the first terrestrial plants , as the end of the Late Ordovician was one of the coldest times in the last 600 million years of Earth's history. On the whole,

2790-460: The Cambrian or even the Ediacaran , became common and varied, especially bivalves , gastropods , and nautiloid cephalopods. Cephalopods diversified from shallow marine tropical environments to dominate almost all marine environments. Graptolites, which evolved in the preceding Cambrian period, thrived in the oceans. This includes the distinctive Nemagraptus gracilis graptolite fauna, which

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2880-472: The Cambrian were succeeded by those that dominated the rest of the Paleozoic, such as articulate brachiopods, cephalopods , and crinoids . Articulate brachiopods, in particular, largely replaced trilobites in shelf communities. Their success epitomizes the greatly increased diversity of carbonate shell-secreting organisms in the Ordovician compared to the Cambrian. Ordovician geography had its effect on

2970-967: The Dapingian and the early Darriwilian. The Llanvirn corresponds to the late Darriwilian. The Caradoc covers the Sandbian and the first half of the Katian. The Ashgill represents the second half of the Katian, plus the Hirnantian . The Ashgill Epoch, the last epoch of the British Ordovician, is made of four ages: the Hirnantian Age, the Rawtheyan Age, the Cautleyan Age, and the Pusgillian Age. These ages make up

3060-848: The Early Eocene Climatic Optimum. Carbon dioxide levels were very high at the Ordovician period's beginning. By the late Early Ordovician, the Earth cooled, giving way to a more temperate climate in the Middle Ordovician, with the Earth likely entering the Early Palaeozoic Ice Age during the Sandbian, and possibly as early as the Darriwilian or even the Floian. The Dapingian and Sandbian saw major humidification events evidenced by trace metal concentrations in Baltoscandia from this time. Evidence suggests that global temperatures rose briefly in

3150-532: The Early Ordovician, leveling off somewhat during the middle of the period. Locally, some regressions occurred, but the sea level rise continued in the beginning of the Late Ordovician. Sea levels fell steadily due to the cooling temperatures for about 3 million years leading up to the Hirnantian glaciation. During this icy stage, sea level seems to have risen and dropped somewhat. Despite much study,

3240-594: The Eramosa Member of the Lockport Formation for the bituminous dolomites exposed below the Guelph Formation along the Eramosa River , northeast of Guelph , Ontario. No detailed description of the type section has ever been published, and the status of the unit has been subject to many different interpretations. Until recently, the Eramosa in Ontario was regarded as the highest Member of

3330-514: The Eramosa are still uncertain. Conodonts at first indicated that the Eramosa age was Ludlovian (c.420 Ma), but recent studies of both conodonts and the Eramosa lagerstätte suggest an earlier, late Wenlockian (c.425 Ma) age. The rocks forming the Eramosa are mostly dolomites, but the composition varies from almost pure, grey-weathering, fine-grained dolomite (at Ancaster) to shaley , bituminous, brown-weathering, bioturbated dolomites. Fossils are common at some localities, scarce at others, and chert

3420-559: The Great Ordovician Biodiversification Event) was no less remarkable; marine faunal genera increased fourfold, resulting in 12% of all known Phanerozoic marine fauna. Several animals also went through a miniaturization process, becoming much smaller than their Cambrian counterparts. Another change in the fauna was the strong increase in filter-feeding organisms. The trilobite, inarticulate brachiopod, archaeocyathid , and eocrinoid faunas of

3510-507: The Hirnantian glaciation. As with North America and Europe , Gondwana was largely covered with shallow seas during the Ordovician. Shallow clear waters over continental shelves encouraged the growth of organisms that deposit calcium carbonates in their shells and hard parts. The Panthalassic Ocean covered much of the Northern Hemisphere , and other minor oceans included Proto-Tethys , Paleo-Tethys , Khanty Ocean , which

3600-624: The Llandovery and Wenlock. Trilobites started to recover in the Rhuddanian, and they continued to enjoy success in the Silurian as they had in the Ordovician despite their reduction in clade diversity as a result of LOME. The Early Silurian was a chaotic time of turnover for crinoids as they rediversified after LOME. Members of Flexibilia, which were minimally impacted by LOME, took on an increasing ecological prominence in Silurian seas. Monobathrid camerates, like flexibles, diversified in

3690-490: The Llandovery, whereas cyathocrinids and dendrocrinids diversified later in the Silurian. Scyphocrinoid loboliths suddenly appeared in the terminal Silurian, shortly before the Silurian-Devonian boundary, and disappeared as abruptly as they appeared very shortly after their first appearance. Endobiotic symbionts were common in the corals and stromatoporoids. Rugose corals especially were colonised and encrusted by

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3780-926: The Lockport Formation, with gradational contacts with the Goat Island Member (below) and the Guelph Formation (above). In 1995, the US Geological Survey proposed a revised stratigraphy based on studies in the Niagara region. It extended the Lockport to Group status and included, from base to top, the Gasport , Goat Island, Eramosa, and Guelph as Formations within the Lockport Group. In a "reference section" in

3870-745: The Niagara River Gorge, the Eramosa was divided into six "units" recognized throughout the Niagara region, but a shaley lower unit previously named for the Hamilton area (the Vinemount Member) was transferred to the Goat Island Formation. Brunton (2009) has proposed a revision for Ontario which restores the Vinemount Member to the Eramosa and recognizes two other members. The exact age and correlation of

3960-596: The Order in which the Older Sedimentary Strata Succeed each other in England and Wales, which was the germ of the modern geological time scale . As it was first identified, the "Silurian" series when traced farther afield quickly came to overlap Sedgwick's "Cambrian" sequence, however, provoking furious disagreements that ended the friendship. The English geologist Charles Lapworth resolved

4050-469: The Ordovician of Wisconsin have been found with an age of about 460 million years ago, a time when the land flora most likely only consisted of plants similar to non-vascular bryophytes . Though stromatolites had declined from their peak in the Proterozoic, they continued to exist in localised settings. The Ordovician came to a close in a series of extinction events that, taken together, comprise

4140-421: The Ordovician were very different from their predecessors in the Cambrian. Many trilobites developed bizarre spines and nodules to defend against predators such as primitive eurypterids and nautiloids while other trilobites such as Aeglina prisca evolved to become swimming forms. Some trilobites even developed shovel-like snouts for ploughing through muddy sea bottoms. Another unusual clade of trilobites known as

4230-582: The Ordovician, when at least two volcanic island arcs collided with Laurentia to form the Appalachian Mountains . Laurentia was otherwise tectonically stable. An island arc accreted to South China during the period, while subduction along north China (Sulinheer) resulted in the emplacement of ophiolites. The ash fall of the Millburg/Big Bentonite bed, at about 454 Ma, was the largest in the last 590 million years. This had

4320-428: The Ordovician. The ice age was possibly not long-lasting. Oxygen isotopes in fossil brachiopods show its duration may have been only 0.5 to 1.5 million years. Other researchers (Page et al.) estimate more temperate conditions did not return until the late Silurian. The late Ordovician glaciation event was preceded by a fall in atmospheric carbon dioxide (from 7000 ppm to 4400 ppm). The dip may have been caused by

4410-557: The Ordovician. Terrestrial plants probably evolved from green algae, first appearing as tiny non- vascular forms resembling liverworts , in the middle to late Ordovician. Fossil spores found in Ordovician sedimentary rock are typical of bryophytes. Among the first land fungi may have been arbuscular mycorrhiza fungi ( Glomerales ), playing a crucial role in facilitating the colonization of land by plants through mycorrhizal symbiosis , which makes mineral nutrients available to plant cells; such fossilized fungal hyphae and spores from

4500-475: The Silures show little correlation ( cf . Geologic map of Wales , Map of pre-Roman tribes of Wales ), Murchison conjectured that their territory included Caer Caradoc and Wenlock Edge exposures - and that if it did not there were plenty of Silurian rocks elsewhere 'to sanction the name proposed'. In 1835 the two men presented a joint paper, under the title On the Silurian and Cambrian Systems, Exhibiting

4590-486: The Silurian was the diversification of jawed fish , which include placoderms , acanthodians (which gave rise to cartilaginous fish ) and osteichthyan ( bony fish , further divided into lobe-finned and ray-finned fishes )， although this corresponded to sharp decline of jawless fish such as conodonts and ostracoderms . The Silurian system was first identified by the Scottish geologist Roderick Murchison , who

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4680-518: The Silurian. The definitive oldest record of millipede ever known is Kampecaris obanensis and Archidesmus sp. from the late Silurian (425 million years ago) of Kerrera . There are also other millipedes, centipedes , and trigonotarbid arachnoids known from Ludlow (420 million years ago). Predatory invertebrates would indicate that simple food webs were in place that included non-predatory prey animals. Extrapolating back from Early Devonian biota, Andrew Jeram et al. in 1990 suggested

4770-421: The South Pole. This contributed to the Hibernian glaciation and the associated extinction event. The Ordovician meteor event is a proposed shower of meteors that occurred during the Middle Ordovician Epoch, about 467.5 ± 0.28 million years ago, due to the break-up of the L chondrite parent body. It is not associated with any major extinction event. A 2024 study found that craters from this event cluster in

4860-416: The Tethys, the Proto-Tethys and Paleo-Tethys , the Rheic Ocean , the Iapetus Ocean (a narrow seaway between Avalonia and Laurentia), and the newly formed Ural Ocean . The Silurian period was once believed to have enjoyed relatively stable and warm temperatures, in contrast with the extreme glaciations of the Ordovician before it and the extreme heat of the ensuing Devonian; however, it is now known that

4950-449: The Wuliuan, exploded in diversity during the Tremadocian, quickly becoming globally widespread. Several groups of endobiotic symbionts appeared in the Ordovician. In the Early Ordovician, trilobites were joined by many new types of organisms, including tabulate corals, strophomenid , rhynchonellid , and many new orthid brachiopods, bryozoans, planktonic graptolites and conodonts, and many types of molluscs and echinoderms, including

5040-444: The conflict by defining a new Ordovician system including the contested beds. An 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

5130-456: The depths. These organisms were cosmopolitan in distribution and present at most latitudes. At the end of the second event, melting glaciers caused the sea level to rise and stabilise once more. The rebound of life's diversity with the permanent re-flooding of continental shelves at the onset of the Silurian saw increased biodiversity within the surviving Orders. Recovery was characterized by an unusual number of "Lazarus taxa", disappearing during

5220-415: The details remain unresolved. In particular, some researches interpret the fluctuations in sea level as pre-Hibernian glaciation, but sedimentary evidence of glaciation is lacking until the end of the period. There is evidence of glaciers during the Hirnantian on the land we now know as Africa and South America, which were near the South Pole at the time, facilitating the formation of the ice caps of

5310-422: The diversity of fauna; Ordovician invertebrates displayed a very high degree of provincialism. The widely separated continents of Laurentia and Baltica, then positioned close to the tropics and boasting many shallow seas rich in life, developed distinct trilobite faunas from the trilobite fauna of Gondwana, and Gondwana developed distinct fauna in its tropical and temperature zones. The Tien Shan terrane maintained

5400-495: The early Katian (Boda Event), depositing bioherms and radiating fauna across Europe. The early Katian also witnessed yet another humidification event. Further cooling during the Hirnantian, at the end of the Ordovician, led to the Late Ordovician glaciation . The Ordovician saw the highest sea levels of the Paleozoic, and the low relief of the continents led to many shelf deposits being formed under hundreds of metres of water. The sea level rose more or less continuously throughout

5490-408: The edge of the continental shelf) can be identified, and the highest Silurian sea level was probably around 140 metres (459 ft) higher than the lowest level reached. During this period, the Earth entered a warm greenhouse phase, supported by high CO 2 levels of 4500 ppm, and warm shallow seas covered much of the equatorial land masses. Early in the Silurian, glaciers retreated back into

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5580-461: The end of the Silurian, sea levels dropped again, leaving telltale basins of evaporites extending from Michigan to West Virginia, and the new mountain ranges were rapidly eroded. The Teays River , flowing into the shallow mid-continental sea, eroded Ordovician Period strata, forming deposits of Silurian strata in northern Ohio and Indiana. The vast ocean of Panthalassa covered most of the northern hemisphere. Other minor oceans include two phases of

5670-425: The extinction and reappearing well into the Silurian, which suggests that the taxa survived in small numbers in refugia . An alternate extinction hypothesis suggested that a ten-second gamma-ray burst could have destroyed the ozone layer and exposed terrestrial and marine surface-dwelling life to deadly ultraviolet radiation and initiated global cooling. Recent work considering the sequence stratigraphy of

5760-426: The fauna that emerged in the Ordovician were the template for the remainder of the Palaeozoic. The fauna was dominated by tiered communities of suspension feeders, mainly with short food chains. The ecological system reached a new grade of complexity far beyond that of the Cambrian fauna, which has persisted until the present day. Though less famous than the Cambrian explosion , the Ordovician radiation (also known as

5850-427: The first rugose corals appeared. The planktonic graptolites remained diverse, with the Diplograptina making their appearance. One of the earliest known armoured agnathan (" ostracoderm ") vertebrates, Arandaspis , dates from the Middle Ordovician. During the Middle Ordovician there was a large increase in the intensity and diversity of bioeroding organisms. This is known as the Ordovician Bioerosion Revolution. It

5940-448: The first deep-boring bivalves are known from this period. Chitons saw a peak in diversity during the middle of the Silurian. Hederelloids enjoyed significant success in the Silurian, with some developing symbioses with the colonial rugose coral Entelophyllum . The Silurian was a heyday for tentaculitoids , which experienced an evolutionary radiation focused mainly in Baltoscandia, along with an expansion of their geographic range in

6030-488: The first period to see megafossils of extensive terrestrial biota in the form of moss -like miniature forests along lakes and streams and networks of large, mycorrhizal nematophytes , heralding the beginning of the Silurian-Devonian Terrestrial Revolution. However, the land fauna did not have a major impact on the Earth until it diversified in the Devonian. The first fossil records of vascular plants , that is, land plants with tissues that carry water and food, appeared in

6120-451: The global climate underwent many drastic fluctuations throughout the Silurian, evidenced by numerous major carbon and oxygen isotope excursions during this geologic period. Sea levels rose from their Hirnantian low throughout the first half of the Silurian; they subsequently fell throughout the rest of the period, although smaller scale patterns are superimposed on this general trend; fifteen high-stands (periods when sea levels were above

6210-502: The globe. At the start of the period, the continents of Laurentia (in present-day North America ), Siberia , and Baltica (present-day northern Europe) were separated from Gondwana by over 5,000 kilometres (3,100 mi) of ocean. These smaller continents were also sufficiently widely separated from each other to develop distinct communities of benthic organisms. The small continent of Avalonia had just rifted from Gondwana and began to move north towards Baltica and Laurentia, opening

6300-400: The lack of tillites in the middle to late Silurian make this explanation problematic. The Silurian period has been viewed by some palaeontologists as an extended recovery interval following the Late Ordovician mass extinction (LOME), which interrupted the cascading increase in biodiversity that had continuously gone on throughout the Cambrian and most of the Ordovician. The Silurian was

6390-402: 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 , Baltica , and Laurentia drifted together near the equator , starting the formation of

6480-561: The mass extinction's aftermath, but expanded their range afterwards. The most abundant brachiopods were atrypids and pentamerides; atrypids were the first to recover and rediversify in the Rhuddanian after LOME, while pentameride recovery was delayed until the Aeronian. Bryozoans exhibited significant degrees of endemism to a particular shelf. They also developed symbiotic relationships with cnidarians and stromatolites. Many bivalve fossils have also been found in Silurian deposits, and

6570-548: The ophiuroids ("brittle stars") and the first sea stars . Nevertheless, the arthropods remained abundant; all the Late Cambrian orders continued, and were joined by the new group Phacopida . The first evidence of land plants also appeared (see evolutionary history of life ). In the Middle Ordovician, the trilobite-dominated Early Ordovician communities were replaced by generally more mixed ecosystems, in which brachiopods, bryozoans, molluscs, cornulitids , tentaculitids and echinoderms all flourished, tabulate corals diversified and

6660-400: The sea level dropped, and the vast shallow intra-continental Ordovician seas withdrew, which eliminated many ecological niches. When they returned, they carried diminished founder populations that lacked many whole families of organisms. They then withdrew again with the next pulse of glaciation, eliminating biological diversity with each change. Species limited to a single epicontinental sea on

6750-426: The sea, and about 49% of genera of fauna disappeared forever; brachiopods and bryozoans were greatly reduced, along with many trilobite , conodont and graptolite families. The most commonly accepted theory is that these events were triggered by the onset of cold conditions in the late Katian, followed by an ice age , in the Hirnantian faunal stage, that ended the long, stable greenhouse conditions typical of

6840-495: The second half of the Silurian Period. The earliest-known representatives of this group are Cooksonia . Most of the sediments containing Cooksonia are marine in nature. Preferred habitats were likely along rivers and streams. Baragwanathia appears to be almost as old, dating to the early Ludlow (420 million years) and has branching stems and needle-like leaves of 10–20 centimetres (3.9–7.9 in). The plant shows

6930-512: The second largest of the five major extinction events in Earth's history in terms of percentage of genera that became extinct. The only larger one was the Permian–Triassic extinction event . The extinctions occurred approximately 447–444 million years ago and mark the boundary between the Ordovician and the following Silurian Period. At that time all complex multicellular organisms lived in

7020-516: The shallow Silurian seas and lakes of North America; many of their fossils have been found in New York state . Brachiopods were abundant and diverse, with the taxonomic composition, ecology, and biodiversity of Silurian brachiopods mirroring Ordovician ones. Brachiopods that survived the LOME developed novel adaptations for environmental stress, and they tended to be endemic to a single palaeoplate in

7110-425: The shells of most molluscs , dissolved rapidly on the sea floor after death. Unlike Cambrian times, when calcite production was dominated by microbial and non-biological processes, animals (and macroalgae) became a dominant source of calcareous material in Ordovician deposits. The Early Ordovician climate was very hot, with intense greenhouse conditions and sea surface temperatures comparable to those during

7200-551: The sun's rays from the proposed Ordovician ring system, which also caused the Ordovician meteor event , may have also led to the glaciation. As the southern supercontinent Gondwana drifted over the South Pole, ice caps formed on it, which have been detected in Upper Ordovician rock strata of North Africa and then-adjacent northeastern South America, which were south-polar locations at the time. As glaciers grew,

7290-577: The time period from c. 450 Ma to c. 443 Ma. The Rawtheyan, the second last of the Ashgill ages, was from c. 449 Ma to c. 445 Ma. It is in the Katian Age of the ICS's Geologic Time Scale . During the Ordovician, the southern continents were assembled into Gondwana , which reached from north of the equator to the South Pole . The Panthalassic Ocean, centered in the northern hemisphere, covered over half

7380-399: The trinucleids developed a broad pitted margin around their head shields. Some trilobites such as Asaphus kowalewski evolved long eyestalks to assist in detecting predators whereas other trilobite eyes in contrast disappeared completely. Molecular clock analyses suggest that early arachnids started living on land by the end of the Ordovician. Although solitary corals date back to at least

7470-693: Was adopted as an official period of the Paleozoic Era by the International Geological Congress . Life continued to flourish during the Ordovician as it had 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, with members of the latter group probably starting their establishment on land during this time, becoming fully established by

7560-537: Was closed off by the Late Ordovician, Iapetus Ocean , and the new Rheic Ocean . For most of the Late Ordovician life continued to flourish, but at and near the end of the period there were mass-extinction events that seriously affected conodonts and planktonic forms like graptolites . The trilobites Agnostida and Ptychopariida completely died out, and the Asaphida were much reduced. Brachiopods , bryozoans and echinoderms were also heavily affected, and

7650-618: Was distributed widely during peak sea levels in the Sandbian. Some new cystoids and crinoids appeared. It was long thought that the first true vertebrates (fish — Ostracoderms ) appeared in the Ordovician, but recent discoveries in China reveal that they probably originated in the Early Cambrian . The first gnathostome (jawed fish) may have appeared in the Late Ordovician epoch. Chitinozoans, which first appeared late in

7740-601: Was 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 a Latin name for Wales. Whilst the British rocks now identified as belonging to the Silurian System and the lands now thought to have been inhabited in antiquity by

7830-495: Was limited to the Iapetus margin of Laurentia; elsewhere, the pattern was of rifting in back-arc basins followed by remerger. This reflected episodic switching from extension to compression. The initiation of new subduction reflected a global reorganization of tectonic plates centered on the amalgamation of Gondwana. The Taconic orogeny , a major mountain-building episode, was well under way in Cambrian times. This continued into

7920-409: Was questioned in 1854 by Edward Forbes , and the later 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 Silurian fossils. With the supercontinent Gondwana covering the equator and much of the southern hemisphere,

8010-520: Was the initial establishment of terrestrial life in what is known as the Silurian-Devonian Terrestrial Revolution : vascular plants emerged from more primitive land plants, dikaryan fungi started expanding and diversifying along with glomeromycotan fungi, and three groups of arthropods ( myriapods , arachnids and hexapods ) became fully terrestrialized. Another significant evolutionary milestone during

8100-532: Was traditionally broken into Early (Tremadocian and Arenig ), Middle ( Llanvirn (subdivided into Abereiddian and Llandeilian) and Llandeilo ) and Late ( Caradoc and Ashgill) epochs. The corresponding rocks of the Ordovician System are referred to as coming from the Lower, Middle, or Upper part of the column. The Tremadoc corresponds to the ICS's Tremadocian. The Arenig corresponds to the Floian, all of

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