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
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
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
Charles Lapworth FRS FGS (20 September 1842 – 13 March 1920) was a headteacher and an English geologist who pioneered faunal analysis using index fossils and identified the Ordovician period.
Biography
Charles Lapworth was born at Faring ...
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
, area_land_km2 = 6,172
, postal_code_type = Postcode
, postal_code = LL, CH, SY
, image_map1 = Wales North Wales locator map.svg
, map_caption1 = Six principal areas of Wales common ...
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''. Zoo ...
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
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
and
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
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, 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
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
, 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, ...
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 stages ...
n regional stratigraphic schemes.
The Ordovician Period in Britain was traditionally broken into Early (Tremadocian and
Arenig
In geology, the Arenig (or Arenigian) is a time interval during the Ordovician period and also the suite of rocks which were deposited during this interval.
History
The term was first used by Adam Sedgwick in 1847 with reference to the "Areni ...
), Middle (
Llanvirn
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
The ...
(subdivided into Abereiddian and Llandeilian) and
Llandeilo
Llandeilo () is a town and community in Carmarthenshire, Wales, situated at the crossing of the River Towy by the A483 on a 19th-century stone bridge. Its population was 1,795 at the 2011 Census. It is adjacent to the westernmost point of the B ...
) 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 na ...
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
The Hirnantian is the final internationally recognized stage of the Ordovician Period of the Paleozoic Era. It was of short duration, lasting about 1.4 million years, from to Ma (million years ago). The early part of the Hirnantian was charact ...
/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
![Ordovician 470](https://upload.wikimedia.org/wikipedia/commons/0/0e/Ordovician_470.svg)
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 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, although ...
(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
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 of ...
, and
Baltica
Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains.
The thick core of Baltica, the East European Craton, is mo ...
(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
The Rheic Ocean was an ocean which separated two major palaeocontinents, Gondwana and Laurussia (Laurentia- Baltica-Avalonia). One of the principal oceans of the Palaeozoic, its sutures today stretch from Mexico to Turkey and its closure result ...
between Gondwana and Avalonia. Avalonia collided with Baltica towards the end of Ordovician.
Other geographic features of the Ordovician world included the
Tornquist Sea
The Tornquist Sea or Tornquist Ocean was a sea located between the palaeocontinents Avalonia and Baltica about . The remains of the sea today form a suture stretching across northern Europe (Tornquist Zone).
Geologic evolution
...
, 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 own ...
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
Orogeny is a mountain building process. An orogeny is an event that takes place at a convergent plate margin when plate motion compresses the margin. An ''orogenic belt'' or ''orogen'' develops as the compressed plate crumples and is uplifted t ...
(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. They ...
. 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
Volcanic ash consists of fragments of rock, mineral crystals, and volcanic glass, created during volcanic eruptions and measuring less than 2 mm (0.079 inches) in diameter. The term volcanic ash is also often loosely used to refer t ...
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
Dense-rock equivalent (DRE) is a volcanologic calculation used to estimate volcanic eruption volume. One of the widely accepted measures of the size of a historic or prehistoric eruption is the volume of magma ejected as pumice and volcanic ash, k ...
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 period ...
.
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
The Ordovician meteor event was a dramatic increase in the rate at which L chondrite meteorites fell to Earth during the Middle Ordovician period, about 467.5±0.28 million years ago. This is indicated by abundant fossil L chondrite meteorites in ...
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
The L type ordinary chondrites are the second most common group of meteorites, accounting for approximately 35% of all those catalogued, and 40% of the ordinary chondrites. The ordinary chondrites are thought to have originated from three parent a ...
parent body.
It is not associated with any major extinction event.
Geochemistry
![Anomalodonta gigantea Waynesville Franklin Co IN](https://upload.wikimedia.org/wikipedia/commons/a/af/Anomalodonta_gigantea_Waynesville_Franklin_Co_IN.JPG)
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
ooid
Ooids are small (commonly ≤2 mm in diameter), spheroidal, "coated" (layered) sedimentary grains, usually composed of calcium carbonate, but sometimes made up of iron- or phosphate-based minerals. Ooids usually form on the sea floor, m ...
s, calcitic cements, and invertebrate faunas with dominantly calcitic skeletons. Biogenic
aragonite
Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including prec ...
, like that composing the shells of most
molluscs
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
, 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 Andean-Saharan glaciation, also known as the Early Palaeozoic Icehouse, the Early Palaeozoic Ice Age, the Late Ordovician glaciation, the end-Ordovician glaciation, or the Hirnantian glaciation, occurred during the Paleozoic from approximately ...
. 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
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 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
The Rheic Ocean was an ocean which separated two major palaeocontinents, Gondwana and Laurussia (Laurentia- Baltica-Avalonia). One of the principal oceans of the Palaeozoic, its sutures today stretch from Mexico to Turkey and its closure result ...
.
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#Glaciology, ablation over many years, often Century, centuries. It acquires dis ...
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 features ...
s of the Late Ordovician glaciation
The Andean-Saharan glaciation, also known as the Early Palaeozoic Icehouse, the Early Palaeozoic Ice Age, the Late Ordovician glaciation, the end-Ordovician glaciation, or the Hirnantian glaciation, occurred during the Paleozoic from approximately ...
.
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 plankton
Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
ic forms like graptolites
Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding organisms are known chiefly from fossils found from the Middle Cambrian (Miaolingian, Wuliuan) through the L ...
. 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 At ...
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
Ptychopariida is a large, heterogeneous order of trilobite containing some of the most primitive species known. The earliest species occurred in the second half of the Lower Cambrian, and the last species did not survive the Ordovician–Silurian ...
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. Brachiopod
Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s, 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 l ...
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 sea ...
s were also heavily affected, and the endocerid
Endocerida is an extinct Nautiloidea, nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. Endocerida was a diverse group of cephalopods that lived from the Early Ordovician possibly to the ...
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
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 ...
(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
Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
organisms. The trilobite, inarticulate brachiopod, archaeocyathid
Archaeocyatha (or archaeocyathids 'ancient cups') is a taxon of extinct, sessile, reef-building marine sponges that lived in warm tropical and subtropical waters during the Cambrian Period. It is believed that the centre of the Archaeocyatha ...
, 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 crinoid
Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
s. 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
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 g ...
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 eurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 Myr, million yea ...
s 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 sec ...
s 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
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 out ...
-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
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
, 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
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, and nautiloid
Nautiloids are a group of marine cephalopods ( Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species ...
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, ...
(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 th ...
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 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
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
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
The ...
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
Tabulata, commonly known as tabulate corals, are an order of extinct forms of coral. They are almost always colonial, forming colonies of individual hexagonal cells known as corallites defined by a skeleton of calcite, similar in appearance to a ...
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
Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
. Nevertheless, the arthropods remained abundant; all the Late Cambrian orders continued, and were joined by the new group Phacopida
Phacopida ("lens-face") is an order of trilobites that lived from the Late Cambrian to the Late Devonian. It is made up of a morphologically diverse assemblage of taxa in three related suborders.
Characteristics
Phacopida had 8 to 19 thorac ...
. The first evidence of land plants also appeared (see evolutionary history of life
The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest #Origins of life on Earth, emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', fo ...
).
In the Middle Ordovician, the trilobite-dominated Early Ordovician communities were replaced by generally more mixed ecosystems, in which brachiopods, bryozoans, molluscs, cornulitids
Cornulitida is an extinct order of encrusting animals from class Tentaculita, which were common around the globe in the Ordovician to Devonian oceans, and survived until the Carboniferous.
The organisms had shells, and were subject to predation ...
, 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 agnatha
Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct ( conodonts and ostracoderms) species. Among recent animals, cyclosto ...
n ("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 th ...
") 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
''Trypanites'' is a narrow, cylindrical, unbranched boring which is one of the most common trace fossils in hard substrates such as rocks, carbonate hardgrounds and shells (Bromley, 1972). It appears first in the Lower Cambrian (James et al ...
'', ''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
''Osprioneides'' is an ichnogenus of unbranched, elongate borings (a type of trace fossil) in lithic substrate with oval cross−section, single−entrance and straight, curved or irregular course. ''Osprioneides kampto'' Beuck and Wisshak, 200 ...
''. 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
''Trypanites'' is a narrow, cylindrical, unbranched boring which is one of the most common trace fossils in hard substrates such as rocks, carbonate hardgrounds and shells (Bromley, 1972). It appears first in the Lower Cambrian (James et al ...
'' borings in an Ordovician hardground
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 ...
, 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
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 ...
, 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
Oil shale is an organic-rich fine-grained sedimentary rock containing kerogen (a solid mixture of organic chemical compounds) from which liquid hydrocarbons can be produced. In addition to kerogen, general composition of oil shales constitute ...
, 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, a ...
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, a ...
File:OrdFossilsMN.JPG, Brachiopod
Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s 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 l ...
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 sea ...
) 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
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 liverwort
The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
s, 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 from ...
may have been arbuscular mycorrhiza
An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. (N ...
fungi (Glomerales
Glomerales is an order of symbiotic fungi within the phylum Glomeromycota.
Biology
These fungi are all biotrophic mutualists. Most employ the arbuscular mycorrhizal method of nutrient exchange with plants. They produce large (.1-.5mm) spores ...
), 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 bryophyte
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 in ...
s.
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 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, as ...
.
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
Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, whi ...
and bryozoans
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 l ...
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 At ...
, 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
Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding
Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and ...
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 gree ...
, 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
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
The Andean-Saharan glaciation, also known as the Early Palaeozoic Icehouse, the Early Palaeozoic Ice Age, the Late Ordovician glaciation, the end-Ordovician glaciation, or the Hirnantian glaciation, occurred during the Paleozoic from approximately ...
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 publishe ...
'', 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
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
In gamma-ray astronomy, gamma-ray bursts (GRBs) are immensely energetic explosions that have been observed in distant galaxies. They are the most energetic and luminous electromagnetic events since the Big Bang. Bursts can last from ten milli ...
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
Sequence stratigraphy is a branch of geology, specifically a branch of stratigraphy, that attempts to discern and understand historic geology through time by subdividing and linking sedimentary deposits into unconformity bounded units on a variety ...
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