Ordovician Geology Of North Dakota
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The Ordovician ( ) is a
geologic period The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochronol ...
and
system A system is a group of Interaction, 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 (systems), environment, is described by its boundaries, ...
, the second of six periods of the Paleozoic
Era An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy, a calendar era used for a given calendar, or the geological eras defined for the history of Earth. Compa ...
. The Ordovician spans 41.6 million years from the end of the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, 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 ...
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 Paleozo ...
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 peop ...
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 D ...
, 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 W ...
and
Roderick Murchison Sir Roderick Impey Murchison, 1st Baronet, (19 February 1792 – 22 October 1871) was a Scotland, Scottish geologist who served as director-general of the British Geological Survey from 1855 until his death in 1871. He is noted for investigat ...
, who were placing the same
rock Rock most often refers to: * Rock (geology), a naturally occurring solid aggregate of minerals or mineraloids * Rock music, a genre of popular music Rock or Rocks may also refer to: Places United Kingdom * Rock, Caerphilly, a location in Wales ...
beds in North Wales in the Cambrian and Silurian systems, respectively. Lapworth recognized that the fossil fauna 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 ei ...
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 Inte ...
. 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 Extinction event, mass extinction events in Earth's history, occurring ro ...
. 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 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, whe ...
. 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 The Great Ordovician Biodiversification Event (GOBE), was an evolutionary radiation of animal life throughout the Ordovician period, 40 million years after the Cambrian explosion, whereby the distinctive Cambrian fauna fizzled out to be replaced wi ...
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.


Subdivisions

A number of regional terms have been used to subdivide the Ordovician Period. In 2008, the
ICS ICS may refer to: Computing * Image Cytometry Standard, a digital multidimensional image file format used in life sciences microscopy * Industrial control system, computer systems and networks used to control industrial plants and infrastructu ...
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 stages ...
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 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 Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * S ...
(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 stages ...
, 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 als ...
to the South Pole. The Panthalassic Ocean, centered in the northern hemisphere, covered over half the globe. At the start of the period, the continents of Laurentia (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 the Car ...
), Siberia, 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 Wester ...
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 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 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. 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. 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 minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
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 ceramics, transparent material, such as glass, in which plants requiring regulated climatic condit ...
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 the Car ...
and Europe,
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 stages ...
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 The Proto-Tethys or Theic Ocean was an ancient ocean that existed from the latest Ediacaran to the Carboniferous (550–330 Ma). History of concept The name "Proto-Tethys" has been used inconsistently for several concepts for a supposed predece ...
,
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; ex ...
,
Khanty Ocean Khanty or Khanty-Mansi Ocean was a small ocean located between Baltica and an island arc (the Kipchaks, Kipchak Arc) near Siberia (continent), Siberia, from near the end of the Precambrian time to the Silurian. The ocean is named after two indige ...
, 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 paleoco ...
, and the new Rheic Ocean. As the Ordovician progressed, there is evidence of glaciers on the land we now know as Africa and South America, which were near the South Pole 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 features ...
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, which ...
s and planktonic forms like graptolites. The trilobites
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, bryozoans and echinoderms 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 recor ...
, 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 anima ...
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 Shelf ( : shelves) may refer to: * Shelf (storage), a flat horizontal surface used for display and storage Geology * Continental shelf, the extended perimeter of a continent, usually covered by shallow seas * Ice shelf, a thick platform of ice f ...
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 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 ext ...
, 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 corals date back to at least the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, 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 ...
, reef-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 th ...
, 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 (fish — Ostracoderms) 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 most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
reveal that they probably originated in the Early
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, 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 ...
. The first gnathostome (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 The taxonomic order Rhynchonellida is one of the two main groups of living articulate brachiopods, the other being the order Terebratulida. They are recognized by their strongly ribbed wedge-shaped or nut-like shells, and the very short hing ...
, 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 The rugosa, also called the tetracorallia or horn coral, are an extinct order of solitary and colonial corals that were abundant in Middle Ordovician to Late Permian seas. Solitary rugosans (e.g., '' Caninia'', '' Lophophyllidium'', '' Neoza ...
appeared. The planktonic graptolites remained diverse, with the Diplograptina making their appearance. One of the earliest known armoured agnathan (" ostracoderm") 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 ...
'', 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 ''Petroxestes'' is a shallow, elongate boring (a type of trace fossil) originally found excavated in carbonate skeletons and hardgrounds of the Upper Ordovician of North America (Wilson and Palmer, 1988, 2006). These Ordovician borings were li ...
'' 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 (biology), 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 substrate ...
''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 ''Petroxestes'' is a shallow, elongate boring (a type of trace fossil) originally found excavated in carbonate skeletons and hardgrounds of the Upper Ordovician of North America (Wilson and Palmer, 1988, 2006). These Ordovician borings were li ...
'' borings in an Ordovician hardground, southern Ohio File:OilShaleEstonia.jpg, Outcrop of Ordovician
kukersite Kukersite is a light-brown marine type oil shale of Ordovician age. It is found in the Baltic Oil Shale Basin in Estonia and North-West Russia. It is of the lowest Upper Ordovician formation, formed some 460 million years ago. It was nam ...
oil shale, northern Estonia File:OilShaleFossilsEstonia.jpg, Bryozoan fossils in Ordovician kukersite oil shale, northern Estonia File:OrdFossilsMN.JPG, Brachiopods and bryozoans 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) 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 as ...
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 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   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
, 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, f ...
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 The history of Earth concerns the development of planet Earth from its formation to the present day. Nearly all branches of natural science have contributed to understanding of the main events of Earth's past, characterized by constant geologic ...
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 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 Paleozo ...
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,
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, which ...
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 A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of Transparent ceramics, transparent material, such as glass, in which plants requiring regulated climatic condit ...
conditions typical of 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).Jeff Hecht
High-carbon ice age mystery solved
'' New Scientist'', 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 i ...
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 stages ...
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, 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 rela ...
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