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The Ordovician ( ) is a geologic period and
system A system is a group of interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment, is described by its boundaries, structure and purpose and express ...
, the second of six periods of the
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
Era. The Ordovician spans 41.6 million years from the end of the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
Period million years ago (Mya) to the start of the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
Period Mya. The Ordovician, named after the
Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peopl ...
tribe of the
Ordovices The Ordovīcēs (Common Brittonic: *''Ordowīcī'') were one of the Celtic tribes living in Great Britain before the Roman invasion. Their tribal lands were located in present-day North Wales and England, between the Silures to the south and the ...
, was defined by Charles Lapworth in 1879 to resolve a dispute between followers of
Adam Sedgwick Adam Sedgwick (; 22 March 1785 – 27 January 1873) was a British geologist and Anglican priest, one of the founders of modern geology. He proposed the Cambrian and Devonian period of the geological timescale. Based on work which he did on ...
and
Roderick Murchison Sir Roderick Impey Murchison, 1st Baronet, (19 February 1792 – 22 October 1871) was a Scottish geologist who served as director-general of the British Geological Survey from 1855 until his death in 1871. He is noted for investigating and ...
, who were placing the same rock beds in
North Wales North Wales ( cy, Gogledd Cymru) is a regions of Wales, region of Wales, encompassing its northernmost areas. It borders Mid Wales to the south, England to the east, and the Irish Sea to the north and west. The area is highly mountainous and rural, ...
in the Cambrian and Silurian systems, respectively. Lapworth recognized that the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...
in the disputed
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
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 was adopted as an official period of the Paleozoic Era by the
International Geological Congress The International Union of Geological Sciences (IUGS) is an international non-governmental organization devoted to international cooperation in the field of geology. About The IUGS was founded in 1961 and is a Scientific Union member of the Int ...
. Life continued to flourish during the Ordovician as it did in the earlier Cambrian Period, although the end of the period was marked by the
Ordovician–Silurian extinction events The Late Ordovician mass extinction (LOME), sometimes known as the end-Ordovician mass extinction or the Ordovician-Silurian extinction, is the first of the "big five" major mass extinction events in Earth's history, occurring roughly 443 Mya. It ...
. Invertebrates, namely molluscs and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s, dominated the oceans, with members of the latter group probably starting their establishment on land during this time, becoming fully established by the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
. The first
land plants The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
are known from this period. The Great Ordovician Biodiversification Event considerably increased the diversity of life.
Fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
, the world's first true
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, 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.


Subdivisions

A number of regional terms have been used to subdivide the Ordovician Period. In 2008, the ICS erected a formal international system of subdivisions. There exist Baltoscandic, British, Siberian, North American, Australian, Chinese Mediterranean and North-
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
n regional stratigraphic schemes. The Ordovician Period in Britain was traditionally broken into Early (Tremadocian and Arenig), Middle ( Llanvirn (subdivided into Abereiddian and Llandeilian) and Llandeilo) and Late (
Caradoc Caradoc Vreichvras (; Modern cy, Caradog Freichfras, ) was a semi-legendary ancestor to the kings of Gwent. He may have lived during the 5th or 6th century. He is remembered in the Matter of Britain as a Knight of the Round Table, under the ...
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 faunal stages (subdivisions of epochs) from youngest to oldest are: Late Ordovician * Hirnantian/Gamach (Ashgill) * Rawtheyan/Richmond (Ashgill) * Cautleyan/Richmond (Ashgill) * Pusgillian/Maysville/Richmond (Ashgill) Middle Ordovician * Trenton (Caradoc) * Onnian/Maysville/Eden (Caradoc) * Actonian/Eden (Caradoc) * Marshbrookian/Sherman (Caradoc) * Longvillian/Sherman (Caradoc) * Soudleyan/Kirkfield (Caradoc) * Harnagian/Rockland (Caradoc) * Costonian/Black River (Caradoc) * Chazy (Llandeilo) * Llandeilo (Llandeilo) * Whiterock (Llanvirn) * Llanvirn (Llanvirn) Early Ordovician * Cassinian (Arenig) * Arenig/Jefferson/Castleman (Arenig) * Tremadoc/Deming/Gaconadian (Tremadoc)


British stages

The Tremadoc corresponds to the (modern) Tremadocian. The Floian corresponds to the lower Arenig; the Arenig continues until the early Darriwilian, subsuming the Dapingian. The Llanvirn occupies the rest of the Darriwilian, and terminates with it at the base of the Late Ordovician. The Sandbian represents the first half of the Caradoc; the Caradoc ends in the mid-Katian, and the Ashgill represents the last half of the Katian, plus the Hirnantian.


Paleogeography and tectonics

During the Ordovician, the southern continents were assembled into
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
, which reached from north of the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
to the
South Pole The South Pole, also known as the Geographic South Pole, Terrestrial South Pole or 90th Parallel South, is one of the two points where Earth's axis of rotation intersects its surface. It is the southernmost point on Earth and lies antipod ...
. The Panthalassic Ocean, centered in the northern hemisphere, covered over half the globe. At the start of the period, the continents of
Laurentia Laurentia or the North American Craton is a large continental craton that forms the ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of North America, althoug ...
(in present-day
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
),
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part ...
, and Baltica (present-day northern Europe) were separated from Gondwana by over 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 Avalonia was a microcontinent in the Paleozoic era. Crustal fragments of this former microcontinent underlie south-west Great Britain, southern Ireland, and the eastern coast of North America. It is the source of many of the older rocks of West ...
had just rifted from Gondwana and began to move north towards Baltica and Laurentia, opening 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 The Mongol-Okhotsk Ocean also known as the Khangai-Khantey Ocean was an ancient ocean, present during the Mesozoic. Oceanic rocks from the ancient ocean floor are preserved in the Mongol-Okhotsk suture zone extending through Mongolia to the Sea of ...
formed a deep embayment between Siberia and the Central Mongolian
terrane In geology, a terrane (; in full, a tectonostratigraphic terrane) is a crust fragment formed on a tectonic plate (or broken off from it) and accreted or " sutured" to crust lying on another plate. The crustal block or fragment preserves its ow ...
s. 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 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 The Taconic orogeny was a mountain building period that ended 440 million years ago and affected most of modern-day New England. A great mountain chain formed from eastern Canada down through what is now the Piedmont of the East coast of the Unit ...
, a major mountain-building episode, was well under way in Cambrian times. This continued into the Ordovician, when at least two volcanic island arcs collided with Laurentia to form the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. The ...
. 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 a dense rock equivalent volume of as much as . 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 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 The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
. Towards the end of the period, Gondwana began to drift across the South Pole. This contributed to the Hibernian glaciation and the associated extinction event.


Ordovician meteor 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.


Geochemistry

The Ordovician was a time of
calcite sea A calcite sea is a sea in which low-magnesium calcite is the primary inorganic marine calcium carbonate precipitate. An aragonite sea is the alternate seawater chemistry in which aragonite and high-magnesium calcite are the primary inorganic ca ...
geochemistry in which low-magnesium
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
was the primary inorganic marine precipitate of calcium carbonate.
Carbonate hardgrounds Carbonate hardgrounds are surfaces of synsedimentarily cemented carbonate layers that have been exposed on the seafloor (Wilson and Palmer, 1992). A hardground is essentially, then, a lithified seafloor. Ancient hardgrounds are found in limestone ...
were thus very common, along with calcitic ooids, calcitic cements, and invertebrate faunas with dominantly calcitic skeletons. Biogenic aragonite, like that composing 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.


Climate and sea level

The early Ordovician climate was very hot, with intense
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
conditions giving way to a more temperate climate in the Middle Ordovician. Further cooling 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 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 30 million years leading up to the Hirnantian glaciation. During this icy stage, sea level seems to have risen and dropped somewhat. Despite much study, 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 also evidence that global temperatures rose briefly in the early Katian (Boda Event), depositing bioherms and radiating fauna across Europe. As with
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
and
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
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 Panthalassa, also known as the Panthalassic Ocean or Panthalassan Ocean (from Greek "all" and "sea"), was the superocean that surrounded the supercontinent Pangaea, the latest in a series of supercontinents in the history of Earth. During the ...
covered much of the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
, and other minor oceans included Proto-Tethys,
Paleo-Tethys The Paleo-Tethys or Palaeo-Tethys Ocean was an ocean located along the northern margin of the paleocontinent Gondwana that started to open during the Middle Cambrian, grew throughout the Paleozoic, and finally closed during the Late Triassic; exi ...
, Khanty Ocean, which was closed off by the Late Ordovician,
Iapetus Ocean The Iapetus Ocean (; ) was an ocean that existed in the late Neoproterozoic and early Paleozoic eras of the geologic timescale (between 600 and 400 million years ago). The Iapetus Ocean was situated in the southern hemisphere, between the paleo ...
, and the new Rheic Ocean. As the Ordovician progressed, there is evidence of
glacier A glacier (; ) is a persistent body of dense ice that is constantly moving under its own weight. A glacier forms where the accumulation of snow exceeds its ablation over many years, often centuries. It acquires distinguishing features, such a ...
s on the land we now know as Africa and South America, which were near the
South Pole The South Pole, also known as the Geographic South Pole, Terrestrial South Pole or 90th Parallel South, is one of the two points where Earth's axis of rotation intersects its surface. It is the southernmost point on Earth and lies antipod ...
at the time, resulting in the
ice cap In glaciology, an ice cap is a mass of ice that covers less than of land area (usually covering a highland area). Larger ice masses covering more than are termed ice sheets. Description Ice caps are not constrained by topographical feat ...
s of the Late Ordovician glaciation.


Life

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
conodont Conodonts ( Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whi ...
s and
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cruc ...
ic forms like graptolites. The
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
s
Agnostida Agnostida is an order of arthropod which have classically been seen as a group of highly modified trilobites, though some recent research has doubted this placement. Regardless, they appear to be close relatives as part of the Artiopoda. They a ...
and Ptychopariida completely died out, and the
Asaphida Asaphida is a large, morphologically diverse order of trilobites found in marine strata dated from the Middle Cambrian until their extinction during the Silurian. Asaphida contains six superfamilies (Anomocaroidea, Asaphoidea, Cyclopygoidea, ...
were much reduced. Brachiopods,
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
ns and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s were also heavily affected, and the endocerid
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s 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.


Fauna

On the whole, 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 Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
, the Ordovician radiation (also known as the Great Ordovician Biodiversification Event) was no less remarkable; marine faunal
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
increased fourfold, resulting in 12% of all known
Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anim ...
marine fauna. Another change in the fauna was the strong increase in filter-feeding organisms. The trilobite, inarticulate brachiopod, archaeocyathid, and
eocrinoid The Eocrinoidea are an extinct class of echinoderms that lived between the Early Cambrian and Late Silurian periods. They are the earliest known group of stalked, arm-bearing echinoderms, and were the most common echinoderms during the Cambrian. ...
faunas of the Cambrian were succeeded by those that dominated the rest of the Paleozoic, such as articulate brachiopods,
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, and crinoids. Articulate brachiopods, in particular, largely replaced trilobites in shelf communities. Their success epitomizes the greatly increased diversity of
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate ...
shell-secreting organisms in the Ordovician compared to the Cambrian. Ordovician geography had its effect on the diversity of fauna. The widely separated continents of Laurentia and Baltica, then positioned close to the tropics and boasting many shallow seas rich in life, developed a distinct trilobite fauna from the trilobite fauna of Gondwana, and Gondwana developed distinct fauna in its tropical and temperature zones. However, tropical articulate brachiopods had a more
cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The extr ...
, with less diversity on different continents. Faunas become less provincial later in the Ordovician, though they were still distinguishable into the late Ordovician. Trilobites in particular were rich and diverse. Trilobites in 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 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
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
s date back to at least the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
,
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes—deposition of sand, wave erosion planing down rock ...
-forming corals appeared in the early Ordovician, including the earliest known
octocorals Octocorallia (also known as Alcyonaria) is a class of Anthozoa comprising around 3,000 species of water-based organisms formed of colonial polyps with 8-fold symmetry. It includes the blue coral, soft corals, sea pens, and gorgonians (sea fans ...
, 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. Molluscs, which appeared during the Cambrian or even the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and t ...
, became common and varied, especially
bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
, 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 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 Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
(fish —
Ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and ...
s) appeared in the Ordovician, but recent discoveries in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
reveal that they probably originated in the Early
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
. The first
gnathostome Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...
(jawed fish) may have appeared in the Late Ordovician epoch. Chitinozoans, which first appeared late in 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 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 the first rugose corals appeared. The planktonic graptolites remained diverse, with the Diplograptina making their appearance. One of the earliest known armoured agnathan ("
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and ...
") vertebrates, ''
Arandaspis ''Arandaspis prionotolepis'' is an extinct species of jawless fish that lived in the Ordovician period, about 480 to 470 million years ago. Its remains were found in the Stairway Sandstone near Alice Springs, Australia in 1959, but it was not d ...
'', 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 is marked by a sudden abundance of hard substrate trace fossils such as '' Trypanites'', ''Palaeosabella'', '' Petroxestes'' and '' Osprioneides''.
Bioerosion Bioerosion describes the breakdown of hard ocean substrates – and less often terrestrial substrates – by living organisms. Marine bioerosion can be caused by mollusks, polychaete worms, phoronids, sponges, crustaceans, echinoids, and ...
became an important process, particularly in the thick calcitic skeletons of corals, bryozoans and brachiopods, and on the extensive
carbonate hardgrounds Carbonate hardgrounds are surfaces of synsedimentarily cemented carbonate layers that have been exposed on the seafloor (Wilson and Palmer, 1992). A hardground is essentially, then, a lithified seafloor. Ancient hardgrounds are found in limestone ...
that appear in abundance at this time. File:OrdovicianEdrio.jpg, Upper Ordovician
edrioasteroid Edrioasteroidea is an extinct class of echinoderms. The living animal would have resembled a pentamerously symmetrical disc or cushion. They were obligate encrusters and attached themselves to inorganic or biologic hard substrates (frequently ha ...
''Cystaster stellatus'' on a cobble from the Kope Formation in northern Kentucky with the cyclostome bryozoan ''Corynotrypa'' in the background File:FossilMtnUT.jpg, Middle Ordovician fossiliferous shales and limestones at Fossil Mountain, west-central Utah File:Outcrop of Upper Ordovician rubbly limestone and shale, southern Indiana.jpg, Outcrop of Upper Ordovician rubbly limestone and shale, southern Indiana File:OrdOutcropTN.JPG, Outcrop of Upper Ordovician limestone and minor shale, central Tennessee File:LibertyBorings.jpg, '' Trypanites'' borings in an Ordovician hardground, southeastern Indiana File:Petroxestes borings Ordovician.jpg, '' Petroxestes'' borings in an Ordovician hardground, southern Ohio File:OilShaleEstonia.jpg, Outcrop of Ordovician kukersite oil shale, northern
Estonia Estonia, formally the Republic of Estonia, is a country by the Baltic Sea in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the sea across from Sweden, to the south by Latvia, an ...
File:OilShaleFossilsEstonia.jpg, Bryozoan fossils in Ordovician kukersite oil shale, northern
Estonia Estonia, formally the Republic of Estonia, is a country by the Baltic Sea in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the sea across from Sweden, to the south by Latvia, an ...
File:OrdFossilsMN.JPG, Brachiopods and
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
ns in an Ordovician limestone, southern Minnesota File:PlatystrophiaOrdovician.jpg, ''Vinlandostrophia ponderosa'', Maysvillian (Upper Ordovician) near Madison, Indiana (scale bar is 5.0 mm) File:Echinosphaerites.JPG, The Ordovician cystoid ''
Echinosphaerites ''Echinosphaerites'' is a genus of rhombiferan cystoid echinoderms that lived in the Early to Middle Ordovician of North America and Europe (Bockelie, 1981). Biology Echinosphaerites had branched biserial brachioles which is rare for species ...
'' (an extinct
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
) from northeastern Estonia; approximately 5 cm in diameter File:Prasopora.JPG, ''Prasopora'', a trepostome bryozoan from the Ordovician of Iowa File:EncrustedStroph.JPG, An Ordovician strophomenid brachiopod with encrusting inarticulate brachiopods and a bryozoan File:Protaraea.jpg, The heliolitid coral ''Protaraea richmondensis'' encrusting a gastropod; Cincinnatian (Upper Ordovician) of southeastern Indiana File:ZygospiraAttached.jpg, ''Zygospira modesta'', atrypid brachiopods, preserved in their original positions on a trepostome bryozoan from the Cincinnatian (Upper Ordovician) of southeastern Indiana File:DiplograptusCaneySprings.jpg, Graptolites (''Amplexograptus'') from the Ordovician near Caney Springs, Tennessee


Flora

Green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga ...
were common in the Late Cambrian (perhaps earlier) and in the Ordovician. Terrestrial plants probably evolved from green algae, first appearing as tiny non-
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
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 A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
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
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e and
spore In biology, a spore is a unit of sexual 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, ...
s from 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.


End of the period

The Ordovician came to a close in a series of
extinction event An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
s that, taken together, comprise the second largest of the five major extinction events in Earth's history in terms of percentage of
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
that became extinct. The only larger one was the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
. The extinctions occurred approximately 447–444 million years ago and mark the boundary between the Ordovician and the following
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
Period. At that time all complex multicellular organisms lived in the sea, and about 49% of genera of fauna disappeared forever; brachiopods and bryozoans were greatly reduced, along with many
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
,
conodont Conodonts ( Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whi ...
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 An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
, in the Hirnantian faunal stage, that ended the long, stable
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
conditions typical of the Ordovician. The ice age was possibly not long-lasting. Oxygen
isotopes Isotopes are two or more types of atoms that have the same atomic number (number of protons in their nuclei) and position in the periodic table (and hence belong to the same chemical element), and that differ in nucleon numbers ( mass numbers ...
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).Jeff Hecht
High-carbon ice age mystery solved
''
New Scientist ''New Scientist'' is a magazine covering all aspects of science and technology. Based in London, it publishes weekly English-language editions in the United Kingdom, the United States and Australia. An editorially separate organisation publish ...
'', 8 March 2010 (retrieved 30 June 2014)
The dip may have been caused by a burst of volcanic activity that deposited new silicate rocks, which draw CO2 out of the air as they erode. Another possibility is that
bryophytes The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants ( embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited ...
and lichens, which colonized land in the middle to late Ordovician, may have increased weathering enough to draw down levels. The drop in selectively affected the shallow seas where most organisms lived. As the southern supercontinent
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
drifted over the South Pole, ice caps formed on it, which have been detected in Upper Ordovician rock strata of
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
and then-adjacent northeastern South America, which were south-polar locations at the time. As glaciers grew, 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 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 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 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 The ozone layer or ozone shield is a region of Earth's stratosphere that absorbs most of the Sun's ultraviolet radiation. It contains a high concentration of ozone (O3) in relation to other parts of the atmosphere, although still small in rel ...
and exposed terrestrial and marine surface-dwelling life to deadly ultraviolet
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
and initiated global cooling. Recent work considering the sequence stratigraphy of 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.


References


External links

* * An Ordovician reef in Vermont.
Ordovician fossils of the famous Cincinnatian Group
{{Authority control Geological periods 1879 in paleontology