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77-530: Spores are dispersed by many different environmental factors, such as wind, water or animals. In suitable conditions, the sporopollenin-rich walls of pollen grains and spores can persist in the fossil record for hundreds of millions of years, since sporopollenin is resistant to chemical degradation by organic and inorganic chemicals. The chemical composition of sporopollenin has long been elusive due to its unusual chemical stability, insolubility and resistance to degradation by enzymes and strong chemical reagents. It

154-600: 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

231-457: A megasporangium that produces megaspores or a microsporangium that produces microspores. In flowering plants, these sporangia occur within the carpel and anthers, respectively. Fungi commonly produce spores during sexual and asexual reproduction. Spores are usually haploid and grow into mature haploid individuals through mitotic division of cells ( Urediniospores and Teliospores among rusts are dikaryotic). Dikaryotic cells result from

308-403: A "spore" and a " gamete " is that a spore will germinate and develop into a sporeling , while a gamete needs to combine with another gamete to form a zygote before developing further. The main difference between spores and seeds as dispersal units is that spores are unicellular, the first cell of a gametophyte, while seeds contain within them a developing embryo (the multicellular sporophyte of

385-603: 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

462-468: 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

539-437: A center pole. This shows that four spores shared a common origin and were initially in contact with each other forming a tetrahedron. A wider aperture in the shape of a groove may be termed a colpus . The number of colpi distinguishes major groups of plants. Eudicots have tricolpate spores (i.e. spores with three colpi). Envelope-enclosed spore tetrads are taken as the earliest evidence of plant life on land, dating from

616-560: A complete lack of aromatic structures and the presence of a macrocyclic backbone of polyhydroxylated tetraketide-like monomers with pseudo-aromatic 2-pyrone rings. These were crosslinked to a poly(hydroxy acid) chain by ether linkages to form the polymer. Electron microscopy shows that the tapetal cells that surround the developing pollen grain in the anther have a highly active secretory system containing lipophilic globules. These globules are believed to contain sporopollenin precursors. Tracer experiments have shown that phenylalanine

693-590: 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

770-639: 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

847-607: 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

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924-464: 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

1001-693: 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

1078-728: 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

1155-595: A putrid odour, for dispersal of fungal spores is yet another strategy, most prominently used by the stinkhorns . In Common Smoothcap moss ( Atrichum undulatum ), the vibration of sporophyte has been shown to be an important mechanism for spore release. In the case of spore-shedding vascular plants such as ferns, wind distribution of very light spores provides great capacity for dispersal. Also, spores are less subject to animal predation than seeds because they contain almost no food reserve; however they are more subject to fungal and bacterial predation. Their chief advantage

1232-463: A rigid structure. There is also good evidence that the chemical composition of sporopollenin is not the same in all plants, indicating it is a class of compounds rather than having one constant structure. In 2019, thioacidolysis degradation and solid-state NMR was used to determine the molecular structure of pitch pine sporopollenin, finding it primarily composed of polyvinyl alcohol units alongside other aliphatic monomers, all crosslinked through

1309-401: A series of acetal linkages. Its complex and heterogeneous chemical structure give some protection from the biodegradative enzymes of bacteria, fungi and animals. Some aromatic structures based on p -coumarate and naringenin were also identified within the sporopollenin polymer. These can absorb ultraviolet light and thus prevent it penetrating further into the spore. This has relevance to

1386-541: 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

1463-423: A sexual cycle, but are resistant structures used for survival under unfavourable conditions. Myxozoan spores release amoeboid infectious germs ("amoebulae") into their hosts for parasitic infection, but also reproduce within the hosts through the pairing of two nuclei within the plasmodium, which develops from the amoebula. In plants, spores are usually haploid and unicellular and are produced by meiosis in

1540-789: 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

1617-407: Is a major precursor, but other carbon sources also contribute. The biosynthetic pathway for phenylpropanoid is very active in tapetal cells, supporting the idea that its products are needed for sporopollenin synthesis. Chemical inhibitors of pollen development and many male sterile mutants have effects on the secretion of these globules by the tapetal cells. Spore In biology , a spore

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1694-424: Is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants , algae , fungi and protozoa . They were thought to have appeared as early as the mid-late Ordovician period as an adaptation of early land plants. Bacterial spores are not part of

1771-489: 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

1848-478: Is that spores were an adaptation of early land plant species, such as embryophytes , that allowed for plants to easily disperse while adapting to their non-aquatic environment. This is particularly supported by the observation of a thick spore wall in cryptospores . These spore walls would have protected potential offspring from novel weather elements. The second more recent hypothesis is that spores were an early predecessor of land plants and formed during errors in

1925-451: Is that, of all forms of progeny, spores require the least energy and materials to produce. In the spikemoss Selaginella lepidophylla , dispersal is achieved in part by an unusual type of diaspore , a tumbleweed . Spores have been found in microfossils dating back to the mid-late Ordovician period. Two hypothesized initial functions of spores relate to whether they appeared before or after land plants. The heavily studied hypothesis

2002-518: 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

2079-455: 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

2156-766: 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

2233-471: 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,

2310-491: The meiosis of algae , a hypothesized early ancestor of land plants. Whether spores arose before or after land plants, their contributions to topics in fields like paleontology and plant phylogenetics have been useful. The spores found in microfossils, also known as cryptospores, are well preserved due to the fixed material they are in as well as how abundant and widespread they were during their respective time periods. These microfossils are especially helpful when studying

2387-496: The sporangium of a diploid sporophyte . In some rare cases, diploid spore is also produced in some algae, or fungi. Under favourable conditions, the spore can develop into a new organism using mitotic division, producing a multicellular gametophyte , which eventually goes on to produce gametes. Two gametes fuse to form a zygote , which develops into a new sporophyte. This cycle is known as alternation of generations . The spores of seed plants are produced internally, and

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2464-460: 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

2541-472: 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

2618-967: 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

2695-848: 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

2772-532: 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,

2849-559: 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

2926-507: 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

3003-469: 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

3080-421: 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

3157-582: 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

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3234-428: 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

3311-557: 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

3388-613: 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

3465-564: 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

3542-456: 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

3619-473: 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

3696-422: 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

3773-559: 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

3850-456: The early periods of earth as macrofossils such as plants are not common nor well preserved. Both cryptospores and modern spores have diverse morphology that indicate possible environmental conditions of earlier periods of Earth and evolutionary relationships of plant species. 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 )

3927-425: 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

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4004-485: 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

4081-578: 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

4158-569: The fluids of the ascus that lead to explosive discharge of the ascospores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water ( Buller's drop ), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g . Other fungi rely on alternative mechanisms for spore release, such as external mechanical forces, exemplified by puffballs . Attracting insects, such as flies, to fruiting structures, by virtue of their having lively colours and

4235-735: The fusion of two haploid gamete cells. Among sporogenic dikaryotic cells, karyogamy (the fusion of the two haploid nuclei) occurs to produce a diploid cell. Diploid cells undergo meiosis to produce haploid spores. Spores can be classified in several ways such as by their spore producing structure, function, origin during life cycle, and mobility. Below is a table listing the mode of classification, name, identifying characteristic, examples, and images of different spore species. Under high magnification , spores often have complex patterns or ornamentation on their exterior surfaces. A specialized terminology has been developed to describe features of such patterns. Some markings represent apertures, places where

4312-502: 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

4389-408: The megaspores (formed within the ovules) and the microspores are involved in the formation of more complex structures that form the dispersal units, the seeds and pollen grains. The term spore derives from the ancient Greek word σπορά spora , meaning " seed , sowing", related to σπόρος sporos , "sowing", and σπείρειν speirein , "to sow". In common parlance, the difference between

4466-474: The mid-Ordovician (early Llanvirn, ~ 470  million years ago ), a period from which no macrofossils have yet been recovered. Individual trilete spores resembling those of modern cryptogamic plants first appeared in the fossil record at the end of the Ordovician period. In fungi, both asexual and sexual spores or sporangiospores of many fungal species are actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of

4543-443: The next generation), produced by the fusion of the male gamete of the pollen tube with the female gamete formed by the megagametophyte within the ovule. Spores germinate to give rise to haploid gametophytes, while seeds germinate to give rise to diploid sporophytes. Vascular plant spores are always haploid . Vascular plants are either homosporous (or isosporous) or heterosporous . Plants that are homosporous produce spores of

4620-548: 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

4697-553: The role of pollen and spores in transporting and dispersing the gametes of plants. The DNA of the gametes is readily damaged by the ultraviolet component of daylight. Sporopollenin thus provides some protection from this damage as well as a physically robust container. Analysis of sporopollenin from the clubmoss Lycopodium in the late 1980s have shown distinct structural differences from that of flowering plants. In 2020, more detailed analysis of sporopollenin from Lycopodium clavatum provided more structural information. It showed

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4774-399: The same size and type. Heterosporous plants, such as seed plants , spikemosses , quillworts , and ferns of the order Salviniales produce spores of two different sizes: the larger spore (megaspore) in effect functioning as a "female" spore and the smaller (microspore) functioning as a "male". Such plants typically give rise to the two kind of spores from within separate sporangia, either

4851-400: 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

4928-426: 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

5005-512: 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

5082-425: 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

5159-407: The spores from the reproductive structures as well as travelling through the air over long distances. Many fungi thereby possess specialized mechanical and physiological mechanisms as well as spore-surface structures, such as hydrophobins , for spore ejection. These mechanisms include, for example, forcible discharge of ascospores enabled by the structure of the ascus and accumulation of osmolytes in

5236-492: 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,

5313-577: 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

5390-418: The tough outer coat of the spore can be penetrated when germination occurs. Spores can be categorized based on the position and number of these markings and apertures. Alete spores show no lines. In monolete spores , there is a single narrow line (laesura) on the spore. Indicating the prior contact of two spores that eventually separated. In trilete spores , each spore shows three narrow lines radiating from

5467-399: 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

5544-693: 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

5621-537: 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

5698-554: 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

5775-495: 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

5852-442: Was once thought to consist of polymerised carotenoids but the application of more detailed analytical methods since the 1980s has shown that this is not correct. Analyses have revealed a complex biopolymer , containing mainly long-chain fatty acids , phenylpropanoids , phenolics and traces of carotenoids in a random co-polymer. It is likely that sporopollenin derives from several precursors that are chemically cross-linked to form

5929-532: 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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