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123-511: Climacograptus is an Ordovician genus of graptolites . This hemichordate -related article is a stub . You can help Misplaced Pages by expanding it . This article related to a Cambrian animal is a stub . You can help Misplaced Pages by expanding it . Ordovician The Ordovician, named after the Welsh tribe of the Ordovices , was defined by Charles Lapworth in 1879 to resolve

246-600: 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

369-603: 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

492-468: 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

615-642: A complex series of orogenic events assembled the eastern parts of Gondwana (eastern Africa, Arabian-Nubian Shield, Seychelles, Madagascar, India, Sri Lanka, East Antarctica, and Australia) c. 750 to 530 Ma . First, the Arabian-Nubian Shield collided with eastern Africa (in the Kenya-Tanzania region) in the East African Orogeny c. 750 to 620 Ma . Then Australia and East Antarctica were merged with

738-599: A continuous arc chain, the direction of subduction was different between the Australian-Tasmanian and New Zealand-Antarctica arc segments. Many terranes were accreted to Eurasia during Gondwana's existence, but the Cambrian or Precambrian origin of many of these terranes remains uncertain. For example, some Palaeozoic terranes and microcontinents that now make up Central Asia, often called the "Kazakh" and "Mongolian terranes", were progressively amalgamated into

861-590: 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

984-583: 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

1107-607: 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

1230-689: A diversified assemblage of true insects. In Gondwana, in contrast, ice and, in Australia, volcanism decimated the Devonian flora to a low-diversity seed fern flora – the pteridophytes were increasingly replaced by the gymnosperms which were to dominate until the Mid-Cretaceous. Australia, however, was still located near the Equator during the Early Carboniferous, and during this period, temnospondyl and lepospondyl amphibians and

1353-472: A few million years, reached its peak at c. 200 Ma , and coincided with the Triassic–Jurassic extinction event . The reformed Gondwanan continent was not precisely the same as that which had existed before Pangaea formed; for example, most of Florida and southern Georgia and Alabama is underlain by rocks that were originally part of Gondwana, but this region stayed attached to North America when

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1476-464: 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

1599-594: A gradual decline during the Triassic while ferns, though never dominant, managed to diversify. The brief period of icehouse conditions during the Triassic–Jurassic extinction event had a dramatic impact on dinosaurs but left plants largely unaffected. The Jurassic was mostly one of hot-house conditions and, while vertebrates managed to diversify in this environment, plants have left little evidence of such development, apart from Cheiroleidiacean conifers and Caytoniales and other groups of seed ferns. In terms of biomass,

1722-693: 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

1845-518: A north-east motion about 90 million years ago. While subduction direction changed, it remained oblique (and not perpendicular) to the coast of South America, and the direction change affected several subduction zone -parallel faults including Atacama , Domeyko and Liquiñe-Ofqui . Insular India began to collide with Asia circa 70 Ma , forming the Indian subcontinent , since which more than 1,400 km (870 mi) of crust has been absorbed by

1968-639: A number of species related to those of the laurissilva of Valdivia, through the connection of the Antarctic flora . These include gymnosperms and the deciduous species of Nothofagus , as well as the New Zealand laurel, Corynocarpus laevigatus , and Laurelia novae-zelandiae . New Caledonia and New Zealand became separated from Australia by continental drift 85 million years ago. The islands still retain plants that originated in Gondwana and spread to

2091-544: A pattern that reflects the Jurassic break-up of Pangaea. The Cretaceous saw the arrival of the angiosperms , or flowering plants, a group that probably evolved in western Gondwana (South America–Africa). From there the angiosperms diversified in two stages: the monocots and magnoliids evolved in the Early Cretaceous, followed by the hammamelid dicots . By the Mid-Cretaceous, angiosperms constituted half of

2214-521: A peak in biodiversity – the end-Permian extinction was enormous and so was the radiation that followed. Two families of conifers, Podocarpaceae and Araucariaceae , dominated Gondwana in the Early Triassic, but Dicroidium , an extinct genus of fork-leaved seed ferns, dominated woodlands and forests of Gondwana during most of the Triassic. Conifers evolved and radiated during the period, with six of eight extant families already present before

2337-728: 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

2460-853: A series of events severally restricted the Proto-ACC: change to shallow marine conditions along the North Scotia Ridge; closure of the Fuegan Seaway, the deep sea that existed in Tierra del Fuego; and uplift of the Patagonian Cordillera. This, together with the reactivated Iceland plume , contributed to global warming. During the Miocene, the Drake Passage began to widen, and as water flow between South America and

2583-541: 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

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2706-694: A single jaw from Australia. The closure of the Rheic Ocean and the formation of Pangaea in the Carboniferous resulted in the rerouting of ocean currents that initiated an Ice House period. As Gondwana began to rotate clockwise, Australia shifted south to more temperate latitudes. An ice cap initially covered most of southern Africa and South America but spread to eventually cover most of the supercontinent, save for northernmost Africa-South America and eastern Australia. Giant lycopod and horsetail forests continued to evolve in tropical Laurasia together with

2829-489: 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

2952-818: Is now the Transantarctic Mountains ): the Antarctic Peninsula , Marie Byrd Land , Zealandia , and Thurston Island ; the Falkland Islands and Ellsworth–Whitmore Mountains (in Antarctica) were rotated 90° in opposite directions; and South America south of the Gastre Fault (often referred to as Patagonia ) was pushed westward. The history of the Africa-Antarctica break-up can be studied in great detail in

3075-576: Is now the southern Weddell Sea where initial break-up occurred during the Jurassic c. 180 to 160 Ma . Gondwana began to break up in the early Jurassic following the extensive and fast emplacement of the Karoo-Ferrar flood basalts c. 184 Ma . Before the Karoo plume initiated rifting between Africa and Antarctica , it separated a series of smaller continental blocks from Gondwana's southern, Proto-Pacific margin (along what

3198-541: Is presumed to have been linked to the subduction of cold oceanic lithosphere . During the mid to Late Cretaceous ( c.  90 million years ago ), the Andean orogeny changed significantly in character. Warmer and younger oceanic lithosphere is believed to have started to be subducted beneath South America around this time. Such kind of subduction is held responsible not only for the intense contractional deformation that different lithologies were subject to, but also

3321-787: The Antarctic Peninsula increased, the renewed ACC resulted in cooler global climate. Since the Eocene, the northward movement of the Australian Plate has resulted in an arc-continent collision with the Philippine and Caroline plates and the uplift of the New Guinea Highlands . From the Oligocene to the late Miocene, the climate in Australia, dominated by warm and humid rainforests before this collision, began to alternate between open forest and rainforest before

3444-1075: The Australian Plate are now separated by the Capricorn Plate and its diffuse boundaries. During the opening of the Indian Ocean, the Kerguelen hotspot first formed the Kerguelen Plateau on the Antarctic Plate c. 118 to 95 Ma and then the Ninety East Ridge on the Indian Plate at c. 100 Ma . The Kerguelen Plateau and the Broken Ridge , the southern end of the Ninety East Ridge, are now separated by

3567-518: 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

3690-595: The Cambrian explosion occurred. Laurentia was docked against the western shores of a united Gondwana for a brief period near the Precambrian/Cambrian boundary, forming the short-lived and still disputed supercontinent Pannotia . The Mozambique Ocean separated the Congo – Tanzania – Bangweulu Block of central Africa from Neoproterozoic India (India, the Antongil Block in far eastern Madagascar,

3813-844: The Campbell Plateau , Chatham Rise , Lord Howe Rise , Norfolk Ridge , and New Caledonia , from West Antarctica c. 84 Ma . The opening of the South Atlantic Ocean divided West Gondwana (South America and Africa), but there is considerable debate over the exact timing of this break-up. Rifting propagated from south to north along Triassic–Early Jurassic lineaments, but intra-continental rifts also began to develop within both continents in Jurassic–Cretaceous sedimentary basins, subdividing each continent into three sub-plates. Rifting began c. 190 Ma at Falkland latitudes, forcing Patagonia to move relative to

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3936-550: The Central Atlantic opened . Antarctica, the centre of the supercontinent, shared boundaries with all other Gondwana continents and the fragmentation of Gondwana propagated clockwise around it. The break-up was the result of the eruption of the Karoo-Ferrar igneous province , one of the Earth's most extensive large igneous provinces (LIP) c. 200 to 170 Ma , but the oldest magnetic anomalies between South America, Africa, and Antarctica are found in what

4059-455: 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

4182-834: The Drake Passage and the deepening of the Tasman Gateway. The oldest oceanic crust in the Drake Passage, however, is 34 to 29 Ma -old which indicates that the spreading between the Antarctic and South American plates began near the Eocene/Oligocene boundary. Deep sea environments in Tierra del Fuego and the North Scotia Ridge during the Eocene and Oligocene indicate a "Proto-ACC" opened during this period. Later, 26 to 14 Ma ,

4305-757: The Great American Interchange . The break-up of Gondwana can be said to continue in eastern Africa at the Afar Triple Junction , which separates the Arabian , Nubian , and Somali plates, resulting in rifting in the Red Sea and East African Rift . In the Early Cenozoic , Australia was still connected to Antarctica c. 35–40° south of its current location and both continents were largely unglaciated. A rift between

4428-803: The Himalayan - Tibetan orogen. During the Cenozoic, the orogen resulted in the construction of the Tibetan Plateau between the Tethyan Himalayas in the south and the Kunlun and Qilian mountains in the north. Later, South America was connected to North America via the Isthmus of Panama , cutting off a circulation of warm water and thereby making the Arctic colder, as well as allowing

4551-440: The Indian Subcontinent . Gondwana was formed by the accretion of several cratons (large stable blocks of the Earth's crust), beginning c.   800 to 650 Ma with the East African Orogeny , the collision of India and Madagascar with East Africa, and culminating in c.   600 to 530 Ma with the overlapping Brasiliano and Kuunga orogenies, the collision of South America with Africa, and

4674-414: The Malagasy Orogeny in southern Madagascar) ( 550 Ma ), the collision between East Gondwana and East Africa in two steps, and the Brasiliano orogeny ( 660 to 530 Ma ), the successive collision between South American and African cratons . The last stages of Gondwanan assembly overlapped with the opening of the Iapetus Ocean between Laurentia and western Gondwana. During this interval,

4797-422: The Mozambique Belt , formed 800 to 650 Ma and was originally interpreted as the suture between East (India, Madagascar, Antarctica, and Australia) and West Gondwana (Africa and South America). Three orogenies were recognised during the 1990s as a result of data sets compiled on behalf of oil and mining companies: the East African Orogeny ( 650 to 800 Ma ) and Kuunga orogeny (including

4920-481: The Ordovician . This is the Cuyania or Precordillera terrane of the Famatinian orogeny in northwest Argentina which may have continued the line of the Appalachians southwards. Chilenia terrane accreted later against Cuyania. The collision of the Patagonian terrane with the southwestern Gondwanan occurred in the late Paleozoic. Subduction-related igneous rocks from beneath the North Patagonian Massif have been dated at 320–330 million years old, indicating that

5043-481: The Pangaea supercontinent during the Carboniferous. Pangaea began to break up in the Mid-Jurassic when the Central Atlantic opened . In the western end of Pangaea, the collision between Gondwana and Laurasia closed the Rheic and Palaeo-Tethys oceans. The obliquity of this closure resulted in the docking of some northern terranes in the Marathon , Ouachita , Alleghanian , and Variscan orogenies, respectively. Southern terranes, such as Chortis and Oaxaca , on

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5166-455: 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

5289-483: The Seychelles , and the Napier and Rayner Complexes in East Antarctica ). The Azania continent (much of central Madagascar , the Horn of Africa and parts of Yemen and Arabia) was an island in the Mozambique Ocean. The continent Australia/ Mawson was still separated from India, eastern Africa, and Kalahari by c. 600 Ma , when most of western Gondwana had already been amalgamated. By c. 550 Ma, India had reached its Gondwanan position, which initiated

5412-499: The Southeast Indian Ridge . Separation between Australia and East Antarctica began c. 132 Ma with seafloor spreading occurring c. 96 Ma . A shallow seaway developed over the South Tasman Rise during the Early Cenozoic and as oceanic crust started to separate the continents during the Eocene c. 35.5 Ma global ocean temperature dropped significantly. A dramatic shift from arc- to rift magmatism c. 100 Ma separated Zealandia , including New Zealand ,

5535-404: The Triassic , and started to fragment during the Early Jurassic (around 180 million years ago). The final stages of break-up, involving the separation of Antarctica from South America (forming the Drake Passage ) and Australia, occurred during the Paleogene (from around 66 to 23 million years ago (Ma)). Gondwana was not considered a supercontinent by the earliest definition, since

5658-442: The Uralian orogeny and Laurasia . Pangaea was finally amalgamated in the Late Carboniferous-Early Permian, but the oblique forces continued until Pangaea began to rift in the Triassic. In the eastern end, collisions occurred slightly later. The North China , South China , and Indochina blocks rifted from Gondwana during the middle Paleozoic and opened the Proto-Tethys Ocean . North China docked with Mongolia and Siberia during

5781-422: The Variscan orogeny close to the Carboniferous–Permian boundary. South-east Asia was made of Gondwanan and Cathaysian continental fragments that were assembled during the Mid-Palaeozoic and Cenozoic. This process can be divided into three phases of rifting along Gondwana's northern margin: first, in the Devonian, North and South China , together with Tarim and Quidam (north-western China) rifted, opening

5904-406: 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,

6027-403: The region in central India of the same name , which is derived from Sanskrit for "forest of the Gonds ". The name had been previously used in a geological context, first by H. B. Medlicott in 1872, from which the Gondwana sedimentary sequences ( Permian - Triassic ) are also described. Some scientists prefer the term "Gondwanaland" for the supercontinent to make a clear distinction between

6150-403: The uplift and erosion known to have occurred from the Late Cretaceous onward. Plate tectonic reorganisation since the mid-Cretaceous might also have been linked to the opening of the South Atlantic Ocean . Another change related to mid-Cretaceous plate tectonic rearrangement was the change of subduction direction of the oceanic lithosphere that went from having south-east motion to having

6273-404: 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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6396-415: 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

6519-502: The Carboniferous–Permian, followed by South China. The Cimmerian blocks then rifted from Gondwana to form the Palaeo-Thethys and Neo-Tethys oceans in the Late Carboniferous, and docked with Asia during the Triassic and Jurassic. Western Pangaea began to rift while the eastern end was still being assembled. The formation of Pangaea and its mountains had a tremendous impact on global climate and sea levels, which resulted in glaciations and continent-wide sedimentation. In North America,

6642-421: 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

6765-405: The Early Cretaceous, and West Burma and Woyla during the Late Cretaceous. Gondwana's long, northern margin remained a mostly passive margin throughout the Palaeozoic. The Early Permian opening of the Neo-Tethys Ocean along this margin produced a long series of terranes, many of which were and still are being deformed in the Himalaya Orogeny . These terranes are, from Turkey to north-eastern India:

6888-567: 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

7011-406: 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,

7134-448: The Equator during this period and remained a lifeless and barren landscape. West Gondwana drifted north during the Devonian , bringing Gondwana and Laurasia close together. Global cooling contributed to the Late Devonian extinction (19% of marine families and 50% of genera went extinct) and glaciation occurred in South America. Before Pangaea had formed, terrestrial plants, such as pteridophytes , began to diversify rapidly resulting in

7257-400: The Equator on landmasses then limited to Laurasia and, in Gondwana, to Australia. In the late Silurian, two distinctive lineages, zosterophylls and rhyniophytes , had colonised the tropics. The former evolved into the lycopods that were to dominate the Gondwanan vegetation over a long period, whilst the latter evolved into horsetails and gymnosperms . Most of Gondwana was located far from

7380-459: 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

7503-437: 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

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7626-446: The Jurassic flora was dominated by conifer families and other gymnosperms that had evolved during the Triassic. The Pteridophytes that had dominated during the Palaeozoic were now marginalised, except for ferns. In contrast to Laurentia, very few insect fossils have been found in Gondwana, to a considerable extent because of widespread deserts and volcanism. While plants had a cosmopolitan distribution, dinosaurs evolved and diversified in

7749-406: The Kuunga orogeny (also known as the Pinjarra orogeny). Meanwhile, on the other side of the newly forming Africa, Kalahari collided with Congo and Rio de la Plata which closed the Adamastor Ocean . c. 540–530 Ma, the closure of the Mozambique Ocean brought India next to Australia–East Antarctica, and both North and South China were in proximity to Australia. As the rest of Gondwana formed,

7872-547: The Late Ordovician argues that the mass extinction was a single protracted episode lasting several hundred thousand years, with abrupt changes in water depth and sedimentation rate producing two pulses of last occurrences of species. Gondwana Gondwana ( / ɡ ɒ n d ˈ w ɑː n ə / ) was a large landmass, sometimes referred to as a supercontinent . The remnants of Gondwana make up around two-thirds of today's continental area, including South America , Africa , Antarctica , Australia , Zealandia , Arabia , and

7995-422: The Late Permian, many known from South Africa and Australia. Beetles and cockroaches remained minor elements in this fauna. Tetrapod fossils from the Early Permian have only been found in Laurasia but they became common in Gondwana later during the Permian. The arrival of the therapsids resulted in the first plant-vertebrate-insect ecosystem. During the Mid- to Late Triassic, hot-house conditions coincided with

8118-405: The Miocene, a warm and humid climate developed with pockets of rainforests in central Australia, but before the end of the period, colder and drier climate severely reduced this rainforest. A brief period of increased rainfall in the Pliocene was followed by drier climate which favoured grassland. Since then, the fluctuation between wet interglacial periods and dry glacial periods has developed into

8241-443: The Neoproterozoic to Palaeozoic phase of the Terra Australis Orogen , a series of terranes were rafted from the proto-Andean margin when the Iapteus Ocean opened, to be added back to Gondwana during the closure of that ocean. During the Paleozoic, some blocks which helped to form parts of the Southern Cone of South America, include a piece transferred from Laurentia when the west edge of Gondwana scraped against southeast Laurentia in

8364-409: 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

8487-421: 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

8610-435: 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

8733-428: 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

8856-557: 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

8979-703: The Palaeo-Tethys behind them. These terranes accreted to Asia during Late Devonian and Permian. Second, in the Late Carboniferous to Early Permian, Cimmerian terranes opened Meso-Tethys Ocean; Sibumasu and Qiangtang were added to south-east Asia during Late Permian and Early Jurassic. Third, in the Late Triassic to Late Jurassic, Lhasa , West Burma , Woyla terranes opened the Neo-Tethys Ocean; Lhasa collided with Asia during

9102-575: The South Atlantic (Brazil and Cameroon ) dating to around 120  million years ago , suggesting that some form of land connection still existed between Africa and South America as recently as the early Aptian . The first phases of Andean orogeny in the Jurassic and Early Cretaceous were characterised by extensional tectonics , rifting , the development of back-arc basins and the emplacement of large batholiths . This development

9225-613: 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

9348-595: The Southern Hemisphere, has a "Gondwanan distribution" and is often described as an archaic, or relict , lineage. The distributions in the Proteaceae is, nevertheless, the result of both Gondwanan rafting and later oceanic dispersal. During the Silurian, Gondwana extended from the Equator (Australia) to the South Pole (North Africa and South America) whilst Laurasia was located on the Equator opposite to Australia. A short-lived Late Ordovician glaciation

9471-1113: The Taurides in southern Turkey; the Lesser Caucasus Terrane in Georgia; the Sanand, Alborz, and Lut terranes in Iran; the Mangysglak or Kopetdag Terrane in the Caspian Sea; the Afghan Terrane; the Karakorum Terrane in northern Pakistan; and the Lhasa and Qiangtang terranes in Tibet. The Permian–Triassic widening of the Neo-Tethys pushed all these terranes across the Equator and over to Eurasia. During

9594-631: 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

9717-526: The addition of Australia and Antarctica, respectively. Eventually, Gondwana became the largest piece of continental crust of the Palaeozoic Era, covering an area of some 100,000,000 km (39,000,000 sq mi), about one-fifth of the Earth's surface. It fused with Euramerica during the Carboniferous to form Pangea . It began to separate from northern Pangea ( Laurasia ) during

9840-779: The base of the Absaroka sequence coincides with the Alleghanian and Ouachita orogenies and are indicative of a large-scale change in the mode of deposition far away from the Pangaean orogenies. Ultimately, these changes contributed to the Permian–Triassic extinction event and left large deposits of hydrocarbons, coal, evaporite, and metals. The breakup of Pangaea began with the Central Atlantic magmatic province (CAMP) between South America, Africa, North America, and Europe. CAMP covered more than seven million square kilometres over

9963-540: The colonisation of Gondwana. The Baragwanathia Flora, found only in the Yea Beds of Victoria, Australia, occurs in two strata separated by 1,700 m (5,600 ft) or 30 Ma; the upper assemblage is more diverse and includes Baragwanathia, the first primitive herbaceous lycopod to evolve from the zosterophylls. During the Devonian, giant club mosses replaced the Baragwanathia Flora, introducing

10086-753: The continent Kazakhstania in the late Silurian . Whether these blocks originated on the shores of Gondwana is not known. In the Early Palaeozoic, the Armorican terrane , which today form large parts of France, was part of either Peri-Gondwana or core Gondwana; the Rheic Ocean closed in front of it and the Palaeo-Tethys Ocean opened behind it. Precambrian rocks from the Iberian Peninsula suggest that it, too, formed part of core Gondwana before its detachment as an orocline in

10209-424: The continent became the arid or semiarid landscape it is today. The adjective "Gondwanan" is in common use in biogeography when referring to patterns of distribution of living organisms, typically when the organisms are restricted to two or more of the now-discontinuous regions that were once part of Gondwana, including the Antarctic flora . For example, the plant family Proteaceae , known from all continents in

10332-456: 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

10455-532: 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

10578-833: The development of rifts systems on both continents, including the Central African Rift System and the Central African Shear Zone which lasted until c. 85 Ma . At Brazilian latitudes spreading is more difficult to assess because of the lack of palaeo-magnetic data, but rifting occurred in Nigeria at the Benue Trough c. 118 Ma . North of the Equator the rifting began after 120.4 Ma and continued until c. 100 to 96 Ma . Dinosaur footprints representing identical species assemblages are known from opposite sides of

10701-422: 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

10824-616: 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

10947-493: The end of it. Bennettitales and Pentoxylales , two now extinct orders of gymnospermous plants, evolved in the Late Triassic and became important in the Jurassic and Cretaceous. It is possible that gymnosperm biodiversity surpassed later angiosperm biodiversity and that the evolution of angiosperms began during the Triassic but, if so, in Laurasia rather than in Gondwana. Two Gondwanan classes, lycophytes and sphenophytes , saw

11070-996: The eruption of the Deccan basalts , whose eruption site may survive as the Réunion hotspot . The Seychelles and the Maldives are now separated by the Central Indian Ridge . During the initial break-up in the Early Jurassic a marine transgression swept over the Horn of Africa covering Triassic planation surfaces with sandstone , limestone , shale , marls and evaporites . East Gondwana, comprising Antarctica, Madagascar, India, and Australia, began to separate from Africa. East Gondwana then began to break up c. 132.5 to 96 Ma when India moved northwest from Australia-Antarctica. The Indian Plate and

11193-563: The evolution of Voltziales , one of the few plant orders to survive the end-Permian extinction (57% of marine families and 83% of genera went extinct) and which came to dominate in the Late Permian and from whom true conifers evolved. Tall lycopods and horsetails dominated the wetlands of Gondwana in the Early Permian. Insects co-evolved with glossopterids across Gondwana and diversified with more than 200 species in 21 orders by

11316-425: 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

11439-485: 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

11562-564: The first amniote reptilians evolved, all closely related to the Laurasian fauna, but spreading ice eventually drove these animals away from Gondwana entirely. The Gondwana ice sheet melted, and sea levels dropped during the Permian and Triassic global warming. During this period, the extinct glossopterids colonised Gondwana and reached peak diversity in the Late Permian when coal-forming forests covered much of Gondwana. The period also saw

11685-578: 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

11808-522: The first trees, and by the Late Devonian this first forest was accompanied by the progymnosperms , including the first large trees Archaeopteris . The Late Devonian extinction probably also resulted in osteolepiform fishes evolving into the amphibian tetrapods , the earliest land vertebrates, in Greenland and Russia. The only traces of this evolution in Gondwana are amphibian footprints and

11931-526: The flora in northeastern Australia. There is, however, no obvious connection between this spectacular angiosperm radiation and any known extinction event nor with vertebrate/insect evolution. Insect orders associated with pollination, such as beetles , flies , butterflies and moths , and wasps, bees, and ants , radiated continuously from the Permian-Triassic, long before the arrival of the angiosperms. Well-preserved insect fossils have been found in

12054-694: The fracture zones and magnetic anomalies flanking the Southwest Indian Ridge . The Madagascar block and the Mascarene Plateau , stretching from the Seychelles to Réunion , were broken off India, causing Madagascar and Insular India to be separate landmasses : elements of this break-up nearly coincide with the Cretaceous–Paleogene extinction event . The India–Madagascar–Seychelles separations appear to coincide with

12177-502: 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

12300-861: The lake deposits of the Santana Formation in Brazil, the Koonwarra Lake fauna in Australia, and the Orapa diamond mine in Botswana. Dinosaurs continued to prosper but, as the angiosperm diversified, conifers, bennettitaleans and pentoxylaleans disappeared from Gondwana c. 115 Ma together with the specialised herbivorous ornithischians , whilst generalist browsers, such as several families of sauropodomorph Saurischia , prevailed. The Cretaceous–Paleogene extinction event killed off all dinosaurs except birds, but plant evolution in Gondwana

12423-512: The landmasses of Baltica , Laurentia , and Siberia were separated from it. To differentiate it from the Indian region of the same name (see § Name ), it is also commonly called Gondwanaland . Regions that were part of Gondwana shared floral and zoological elements that persist to the present day. The continent of Gondwana was named by the Austrian scientist Eduard Suess , after

12546-548: 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

12669-600: The other hand, remained largely unaffected by the collision along the southern shores of Laurentia. Some Peri-Gondwanan terranes, such as Yucatán and Florida , were buffered from collisions by major promontories. Other terranes, such as Carolina and Meguma , were directly involved in the collision. The final collision resulted in the Variscan- Appalachian Mountains , stretching from present-day Mexico to southern Europe. Meanwhile, Baltica collided with Siberia and Kazakhstania which resulted in

12792-561: The present arid regime. Australia has thus experienced various climate changes over a 15-million-year period with a gradual decrease in precipitation. The Tasman Gateway between Australia and Antarctica began to open c. 40 to 30 Ma . Palaeontological evidence indicates the Antarctic Circumpolar Current (ACC) was established in the Late Oligocene c. 23 Ma with the full opening of

12915-526: The region and the supercontinent. The assembly of Gondwana was a protracted process during the Neoproterozoic and Paleozoic , which remains incompletely understood because of the lack of paleo-magnetic data. Several orogenies , collectively known as the Pan-African orogeny , caused the continental fragments of a much older supercontinent, Rodinia , to amalgamate. One of those orogenic belts,

13038-760: The remaining Gondwana c. 570 to 530 Ma in the Kuunga Orogeny. The later Malagasy orogeny at about 550–515 Mya affected Madagascar, eastern East Africa and southern India. In it, Neoproterozoic India collided with the already combined Azania and Congo–Tanzania–Bangweulu Block, suturing along the Mozambique Belt. The 18,000 km-long (11,000 mi) Terra Australis Orogen developed along Gondwana's western, southern, and eastern margins. Proto-Gondwanan Cambrian arc belts from this margin have been found in eastern Australia, Tasmania, New Zealand, and Antarctica. Though these belts formed

13161-400: 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

13284-426: 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

13407-454: 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

13530-425: 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

13653-665: The still static remainder of South America and Africa, and this westward movement lasted until the Early Cretaceous 126.7 Ma . From there rifting propagated northward during the Late Jurassic c. 150 Ma or Early Cretaceous c. 140 Ma most likely forcing dextral movements between sub-plates on either side. South of the Walvis Ridge and Rio Grande Rise the Paraná and Etendeka magmatics resulted in further ocean-floor spreading c. 130 to 135 Ma and

13776-399: The subduction process initiated in the early Carboniferous. This was relatively short-lived (lasting about 20 million years), and initial contact of the two landmasses occurred in the mid-Carboniferous, with broader collision during the early Permian. In the Devonian, an island arc named Chaitenia accreted to Patagonia in what is now south-central Chile. Gondwana and Laurasia formed

13899-492: 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,

14022-577: 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

14145-399: 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

14268-636: The two developed but remained an embayment until the Eocene-Oligocene boundary when the Circumpolar Current developed and the glaciation of Antarctica began. Australia was warm and wet during the Palaeocene and dominated by rainforest. The opening of the Tasman Gateway at the Eocene-Oligocene boundary ( 33 Ma ) resulted in abrupt cooling but the Oligocene became a period of high rainfall with swamps in southeast Australia. During

14391-693: 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

14514-602: 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

14637-610: 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

14760-424: Was followed by a Silurian Hot House period. The End-Ordovician extinction , which resulted in 27% of marine invertebrate families and 57% of genera going extinct, occurred during this shift from Ice House to Hot House. By the end of the Ordovician, Cooksonia , a slender, ground-covering plant, became the first known vascular plant to establish itself on land. This first colonisation occurred exclusively around

14883-475: Was hardly affected. Gondwanatheria is an extinct group of non- therian mammals with a Gondwanan distribution (South America, Africa, Madagascar, India, Zealandia and Antarctica) during the Late Cretaceous and Palaeogene. Xenarthra and Afrotheria , two placental clades, are of Gondwanan origin and probably began to evolve separately c. 105 Ma when Africa and South America separated. The laurel forests of Australia, New Caledonia, and New Zealand have

15006-440: 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

15129-412: 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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