Ordovician-Silurian Extinction Events
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The Late Ordovician mass extinction (LOME), sometimes known as the end-Ordovician mass extinction or the Ordovician-Silurian extinction, is the first of the "big five" major
mass extinction events 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 ...
in Earth's history, occurring roughly 443 Mya. It is often considered to be the second-largest known extinction event, in terms of the 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. Extinction was global during this interval, eliminating 49–60% of marine genera and nearly 85% of marine species. Under most tabulations, only the Permian-Triassic mass extinction exceeds the Late Ordovician mass extinction in
biodiversity loss Biodiversity loss includes the worldwide extinction of different species, as well as the local reduction or loss of species in a certain habitat, resulting in a loss of biological diversity. The latter phenomenon can be temporary or permanent, de ...
. The extinction event abruptly affected all major taxonomic groups and caused the disappearance of one third of all
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 ...
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 ...
n families, as well as numerous groups of
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,
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,
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,
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, bivalves, 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 ...
s. Despite its taxonomic severity, the Late Ordovician mass extinction did not produce major changes to ecosystem structures compared to other mass extinctions, nor did it lead to any particular morphological innovations. Diversity gradually recovered to pre-extinction levels over the first 5 million years of the Silurian Period. The Late Ordovician mass extinction is traditionally considered to occur in two distinct pulses. The first pulse began at the boundary between the
Katian The Katian is the second stage of the Upper Ordovician. It is preceded by the Sandbian and succeeded by the Hirnantian Stage. The Katian began million years ago and lasted for about 7.8 million years until the beginning of the Hirnantian millio ...
and
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 ...
stages of the Late Ordovician Period. This extinction pulse is typically attributed 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 approximatel ...
, which abruptly expanded over
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 ...
at the beginning of the Hirnantian and shifted the Earth from a greenhouse to icehouse climate. Cooling and a falling sea level brought on by the glaciation led to habitat loss for many organisms along the continental shelves, especially endemic taxa with restricted temperature tolerance and latitudinal range. During this extinction pulse there were also several marked changes in biologically responsive
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
and
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
isotope 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) ...
s. Marine life partially rediversified during the cold period and a new cold-water ecosystem, the "''
Hirnantia 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 charac ...
'' fauna", was established. The second pulse of extinction occurred in the later half of the Hirnantian as the glaciation abruptly receded and warm conditions returned. The second pulse was associated with intense worldwide anoxia (oxygen depletion) and
euxinia Euxinia or euxinic conditions occur when water is both anoxic and sulfidic. This means that there is no oxygen (O2) and a raised level of free hydrogen sulfide (H2S). Euxinic bodies of water are frequently strongly stratified, have an oxic, highly ...
(toxic sulfide production), which persisted into the subsequent
Rhuddanian In the geologic timescale, the Rhuddanian is the first age of the Silurian Period and of the Llandovery Epoch. The Silurian is in the Paleozoic Era of the Phanerozoic Eon. The Rhuddanian Age began 443.8 ± 1.5 Ma and ended 440.8 ± 1.2 Ma (mi ...
stage of the
Silurian Period 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 ...
. Some researchers have proposed the existence of a third distinct pulse of the mass extinction during the early Rhuddanian, evidenced by a negative carbon isotope excursion and a pulse of anoxia into shelf environments amidst already low background oxygen levels. Others, however, have argued that Rhuddanian anoxia was simply part of the second pulse, which according to this view was longer and more drawn out than most authors suggest.


Impact on life


Ecological impacts

The Late Ordovician mass extinction followed 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 ...
(GOBE), one of the largest surges of increasing biodiversity in the geological and biological history of the Earth. At the time of the extinction, most complex
multicellular organism A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
s lived in the sea, and the only evidence of life on land are rare spores from small early
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 sist ...
. At the time of the extinction, around 100 marine
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
became extinct, covering about 49% 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 ...
(a more reliable estimate than species). The
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 ...
ns were strongly impacted, along with many of 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 ...
,
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 extinction was divided into two major extinction pulses. The first pulse occurred at the base of the global ''Metabolograptus extraordinarius''
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 ...
biozone In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...
, which marks the end of the Katian stage and the start of the Hirnantian stage. The second pulse of extinction occurred in the later part of the Hirnantian stage, coinciding with the ''Metabolograptus persculptus'' zone. Each extinction pulse affected different groups of animals and was followed by a rediversification event. Statistical analysis of marine losses at this time suggests that the decrease in diversity was mainly caused by a sharp increase in extinctions, rather than a decrease in
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. Following such a major loss of diversity, Silurian communities were initially less complex and broader niched. Highly endemic faunas, which characterized the Late Ordovician, were replaced by faunas that were amongst the most cosmopolitan in the Phanerozoic, biogeographic patterns that persisted throughout most of the Silurian. The Late Ordovician mass extinction had few of the long-term ecological impacts associated with the Permian–Triassic and Cretaceous–Paleogene extinction events. Nevertheless, a large number of taxa disappeared from the Earth over a short time interval, eliminating and altering the relative diversity and abundance of certain groups. The Cambrian-type evolutionary fauna nearly died out, and was unable to rediversify after the extinction.


Biodiversity changes in marine invertebrates

Brachiopod diversity and composition was strongly affected, with the Cambrian-type inarticulate brachiopods ( linguliforms and craniiforms) never recovering their pre-extinction diversity. Articulate (
rhynchonelliform Rhynchonelliformea is a major subphylum and clade of brachiopods. It is equivalent to the former class Articulata, which was used previously in brachiopod taxonomy. Articulate brachiopods have many anatomical differences relative to "inarticulate" ...
) brachiopods, part of the Paleozoic evolutionary fauna, were more variable in their response to the extinction. Some early rhynchonelliform groups, such as the
Orthida Orthida is an extinct order of brachiopods which appeared during the Early Cambrian period and became very diverse by the Ordovician, living in shallow-shelf seas. Orthids are the oldest member of the subphylum Rhynchonelliformea, and is the ord ...
and
Strophomenida Strophomenida is an extinct order of articulate brachiopods which lived from the lower Ordovician period to the mid Carboniferous period. Strophomenida is part of the extinct class Strophomenata, and was the largest known order of brachiopods, e ...
, declined significantly. Others, including the
Pentamerida Pentamerida is an order of biconvex, impunctate shelled, articulate brachiopods that are found in marine sedimentary rocks that range from the Middle Cambrian through the Devonian.Moore, Lalcker and Fischer, 1952, Invertebrate Fossils, McGraw-Hil ...
,
Athyridida Athyridida is an order of Paleozoic brachiopods included in the Rhynchonellata, which makes up part of the articulate brachiopods. The Athyridida are the Rostrospracea of R.C Moore, 1952,Moore, Lalcker and Fischer, 1952, Invertebrate Fossils, McG ...
,
Spiriferida Spiriferida is an order of extinct articulate brachiopod fossils which are known for their long hinge-line, which is often the widest part of the shell. In some genera (e.g. ''Mucrospirifer'') it is greatly elongated, giving them a wing-like app ...
, and Atrypida, were less affected and took the opportunity to diversify after the extinction. The extinction pulse at the end of the Katian was selective in its effects, disproportionally affecting deep-water species and tropical endemics inhabiting epicontinental seas. The ''Foliomena'' fauna, an assemblage of thin-shelled species adapted for deep dysoxic (low oxygen) waters, went extinct completely in the first extinction pulse. The ''Foliomena'' fauna was formerly widespread and resistant to background extinction rates prior to the Hirnantian, so their unexpected extinction points towards the abrupt loss of their specific habitat. During the glaciation, a high-latitude brachiopod assemblage, the ''
Hirnantia 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 charac ...
'' fauna, established itself along outer shelf environments in lower latitudes, probably in response to cooling. However, the ''Hirnantia'' fauna would meet its demise in the second extinction pulse, replaced by Silurian-style assemblages adapted for warmer waters. Trilobites were hit hard by both phases of the extinction, with about 70% of genera and 50% of families going extinct between the Katian and Silurian. The extinction disproportionately affected deep water species and groups with fully planktonic larvae or adults. The order Agnostida was completely wiped out, and the formerly diverse
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, ...
survived with only a single genus, ''Raphiophorus''. A cool-water trilobite assemblage, the ''Mucronaspis'' fauna, coincides with the ''Hirnantia'' brachiopod fauna in the timing of its expansion and demise. Over a third of bryozoan genera went extinct, but most families survived the extinction interval and the group as a whole recovered in the Silurian. The hardest-hit subgroups were the cryptostomes and trepostomes, which never recovered the full extent of their Ordovician diversity. Bryozoan extinctions started in coastal regions of Laurentia, before high extinction rates shifted to Baltica by the end of the Hirnantian. Bryozoan biodiversity loss appears to have been a prolonged process which partially preceded the Hirnantian extinction pulses. Extinction rates among Ordovician bryozoan genera were actually higher in the early and late Katian, and origination rates sharply dropped in the late Katian and Hirnantian. About 70% of
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 ...
genera died out, with most extinctions occurring in the first pulse. However, they rediversified quickly in tropical areas and reacquired their pre-extinction diversity not long into the Silurian. Many other echinoderms became very rare after the Ordovician, such as the
cystoids Cystoidea is a class of extinct crinozoan echinoderms, termed cystoids, that lived attached to the sea floor by stalks. They existed during the Paleozoic Era, in the Middle Ordovician and Silurian Periods, until their extinction in the Devo ...
, edrioasteroids, and other early crinoid-like groups.


Possible causes


Glaciation

The first pulse of the Late Ordovician Extinction has typically been attributed 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 approximatel ...
. Although there was a longer cooling trend in Middle and Lower Ordovician, the most severe and abrupt period of glaciation occurred in the Hirnantian stage, which was bracketed by both pulses of the extinction. The rapid continental glaciation was centered on
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 was located at 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 ...
in the Late Ordovician. The Hirnantian glaciation is considered one of the most severe
ice ages 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 ...
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 ' ...
, which previously maintained the relatively warm climate conditions of a greenhouse earth. The cause of the glaciation is heavily debated. The appearance and development of terrestrial plants and microphytoplankton, which consumed atmospheric carbon dioxide, may have diminished the greenhouse effect and promoting the transition of the climatic system to the glacial mode. Though more commonly associated with greenhouse gasses and warming, volcanism may have induced cooling. Volcanoes can supply cooling sulfur
aerosols An aerosol is a suspension of fine solid particles or liquid droplets in air or another gas. Aerosols can be natural or anthropogenic. Examples of natural aerosols are fog or mist, dust, forest exudates, and geyser steam. Examples of anthrop ...
to the atmosphere or deposit basalt flows which accelerate
carbon sequestration Carbon sequestration is the process of storing carbon in a carbon pool. Carbon dioxide () is naturally captured from the atmosphere through biological, chemical, and physical processes. These changes can be accelerated through changes in land ...
in a tropical environment. In addition, volcanic fertilisation of the oceans with phosphorus may have increased populations of photosynthetic algae and enhanced biological sequestration of carbon dioxide from the atmosphere. Increased burial of organic carbon is another method of drawing down carbon dioxide from the air that may have played a role in the Late Ordovician. Two environmental changes associated with the
glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate betw ...
were responsible for much of the Late Ordovician extinction. First, the cooling global climate was probably especially detrimental because the biota were adapted to an intense greenhouse, especially because most shallow sea habitats in the Ordovician were located in the tropics. Second, sea level decline, caused by sequestering of water in the ice cap, drained the vast epicontinental seaways and eliminated the habitat of many endemic communities. Falling sea levels may have acted as a positive feedback loop accelerating further cooling; as shallow seas receded, carbonate-shelf production declined and atmospheric carbon dioxide levels correspondingly decreased, fostering even more cooling. As the southern
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
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 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 ...
,
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 formed on it. Correlating rock strata have been detected in Late Ordovician
rock 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 ...
of North Africa and then-adjacent northeastern South America, which were south-polar locations at the time.
Glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate betw ...
locks up water from the world-ocean, and the
interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...
s free it, causing sea levels repeatedly to drop and rise; the vast shallow mediterranean Ordovician seas withdrew, which eliminated many
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s, then returned, carrying diminished founder populations lacking many whole families of organisms. Then they withdrew again with the next pulse of glaciation, eliminating biological diversity at each change. In the North African strata, five pulses of glaciation from
seismic Seismology (; from Ancient Greek σεισμός (''seismós'') meaning "earthquake" and -λογία (''-logía'') meaning "study of") is the scientific study of earthquakes and the propagation of elastic waves through the Earth or through other ...
sections are recorded. This incurred a shift in the location of bottom-water formation, shifting from low
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...
s, characteristic of greenhouse conditions, to high latitudes, characteristic of icehouse conditions, which was accompanied by increased deep-ocean currents and oxygenation of the bottom-water. An opportunistic fauna briefly thrived there, before anoxic conditions returned. The breakdown in the oceanic circulation patterns brought up nutrients from the abyssal waters. Surviving species were those that coped with the changed conditions and filled the ecological niches left by the extinctions.


Anoxia and euxinia

Another heavily-discussed factor in the Late Ordovician mass extinction is anoxia, the absence of dissolved oxygen in seawater. Anoxia not only deprives most life forms of a vital component of
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
, it also encourages the formation of toxic metal ions and other compounds. One of the most common of these poisonous chemicals is
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
, a biological waste product and major component of the sulfur cycle. Oxygen depletion when combined with high levels of sulfide is called
euxinia Euxinia or euxinic conditions occur when water is both anoxic and sulfidic. This means that there is no oxygen (O2) and a raised level of free hydrogen sulfide (H2S). Euxinic bodies of water are frequently strongly stratified, have an oxic, highly ...
. Though less toxic,
ferrous iron In chemistry, iron(II) refers to the element iron in its +2 oxidation state. In ionic compounds (salts), such an atom may occur as a separate cation (positive ion) denoted by Fe2+. The adjective ferrous or the prefix ferro- is often used to ...
(Fe2+) is another substance which commonly forms in anoxic waters. Anoxia is the most common culprit for the second pulse of the Late Ordovician mass extinction and is connected to many other mass extinctions throughout geological time. It may have also had a role in the first pulse of the Late Ordovician mass extinction, though support for this hypothesis is inconclusive and contradicts other evidence for high oxygen levels in seawater during the glaciation.


Early Hirnantian anoxia

Some geologists have argued that anoxia played a role in the first extinction pulse, though this hypothesis is controversial. In the early Hirnantian, shallow-water sediments throughout the world experience a large positive excursion in the δ34S ratio of buried
pyrite The mineral pyrite (), or iron pyrite, also known as fool's gold, is an iron sulfide with the chemical formula Iron, FeSulfur, S2 (iron (II) disulfide). Pyrite is the most abundant sulfide mineral. Pyrite's metallic Luster (mineralogy), lust ...
. This ratio indicates that shallow-water pyrite which formed at the beginning of the glaciation had a decreased proportion of 32S, a common lightweight isotope of sulfur. 32S in the seawater could hypothetically be used up by extensive deep-sea pyrite deposition. The Ordovician ocean also had very low levels of
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ...
, a nutrient which would otherwise resupply 32S from the land. Pyrite forms most easily in anoxic and euxinic environments, while better oxygenation encourages the formation of
gypsum Gypsum is a soft sulfate mineral composed of calcium sulfate dihydrate, with the chemical formula . It is widely mined and is used as a fertilizer and as the main constituent in many forms of plaster, blackboard or sidewalk chalk, and drywall. ...
instead. As a result, anoxia and euxinia would need to be common in the deep sea to produce enough pyrite to shift the δ34S ratio. A more direct proxy for anoxic conditions is FeHR/FeT. This ratio describes the comparative abundance of highly reactive
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
compounds which are only stable without oxygen. Most geological sections corresponding to the beginning of the Hirnantian glaciation have FeHR/FeT below 0.38, indicating oxygenated waters. However, higher FeHR/FeT values are known from a few deep-water early Hirnantian sequences found in
Nevada Nevada ( ; ) is a U.S. state, state in the Western United States, Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. N ...
and China. Glaciation could conceivably trigger anoxic conditions, albeit indirectly. If continental shelves are exposed by falling sea levels, then organic surface runoff flows into deeper oceanic basins. The organic matter would have more time to leach out
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
and other nutrients before being deposited on the seabed. Increased phosphate concentration in the seawater would lead to
eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytopla ...
and then anoxia. Deep-water anoxia and euxinia would impact deep-water benthic fauna, as expected for the first pulse of extinction. Chemical cycle disturbances would also steepen the
chemocline A chemocline is a type of cline, a layer of fluid with different properties, characterized by a strong, vertical chemistry gradient within a body of water. In bodies of water where chemoclines occur, the cline separates the upper and lower layers ...
, restricting the habitable zone of planktonic fauna which also go extinct in the first pulse. This scenario is congruent with both organic carbon isotope excursions and general extinction patterns observed in the first pulse. However, data supporting deep-water anoxia during the glaciation contrasts with more extensive evidence for well-oxygenated waters. Black shales, which are indicative of an anoxic environment, become very rare in the early Hirnantian compared to surrounding time periods. Although early Hirnantian black shales can be found in a few isolated ocean basins (such as the Yangtze platform of China), from a worldwide perspective these correspond to local events. Some Chinese sections record an early Hirnantian increase in the abundance of Mo-98, a heavy isotope of
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
. This shift can correspond to a balance between minor local anoxia and well-oxygenated waters on a global scale. Other trace elements point towards increased deep-sea oxygenation at the start of the glaciation. Oceanic current modelling suggest that glaciation would have encouraged oxygenation in most areas, apart from the
Paleo-Tethys ocean The Paleo-Tethys or Palaeo-Tethys Ocean was an ocean located along the northern margin of the paleocontinent Gondwana that started to open during the Middle Cambrian, grew throughout the Paleozoic, and finally closed during the Late Triassic; exi ...
. Deep-sea anoxia is not the only explanation for the δ34S excursion of pyrite.
Carbonate-associated sulfate Carbonate-associated sulfates (CAS) are sulfate species found in association with carbonate minerals, either as inclusions, adsorbed phases, or in distorted sites within the carbonate mineral lattice. It is derived primarily from dissolved sulfa ...
maintains high 32S levels, indicating that seawater in general did not experience 32S depletion during the glaciation. Even if pyrite burial did increase at that time, its chemical effects would have been far too slow to explain the rapid excursion or extinction pulse. Instead, cooling may lower the metabolism of warm-water
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cel ...
bacteria, reducing decomposition of organic matter. Fresh organic matter would eventually sink down and supply nutrients to sulfate-reducing microbes living in the seabed. Sulfate-reducing microbes prioritize 32S during
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing r ...
, leaving behind heavier isotopes. A bloom of sulfate-reducing microbes can quickly account for the δ34S excursion in marine sediments without a corresponding decrease in oxygen. A few studies have proposed that the first extinction pulse did not begin with the Hirnantian glaciation, but instead corresponds to an interglacial period or other warming event. Anoxia would be the most likely mechanism of extinction in a warming event, as evidenced by other extinctions involving warming. However, this view of the first extinction pulse is controversial and not widely accepted.


Late Hirnantian anoxia

The late Hirnantian experienced a dramatic increase in the abundance of black shales. Coinciding with the retreat of the Hirnantian glaciation, black shale expands out of isolated basins to become the dominant oceanic sediment at all latitudes and depths. The worldwide distribution of black shales in the late Hirnantian is indicative of a global anoxic event, which has been termed the Hirnantian ocean anoxic event (HOAE).
Molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
,
uranium Uranium is a chemical element with the symbol U and atomic number 92. It is a silvery-grey metal in the actinide series of the periodic table. A uranium atom has 92 protons and 92 electrons, of which 6 are valence electrons. Uranium is weak ...
, and
neodymium Neodymium is a chemical element with the symbol Nd and atomic number 60. It is the fourth member of the lanthanide series and is considered to be one of the rare-earth metals. It is a hard, slightly malleable, silvery metal that quickly tarnishe ...
isotope excursions found in many different regions also correspond to widespread anoxia. At least in European sections, late Hirnantian anoxic waters were originally ferruginous (dominated by ferrous iron) before gradually becoming more euxinic. In the Yangtze Sea, located on the western margins of the South China microcontinent, the second extinction pulse occurred alongside intense euxinia which spread out from the middle of the continental shelf. However, some evidence suggests that the top of the water column remained well oxygenated even as the seafloor became deoxygenated. On a global scale, euxinia was probably one or two orders of magnitude more prevalent than in the modern day. Global anoxia may have lasted more than 3 million years, persisting through the entire
Rhuddanian In the geologic timescale, the Rhuddanian is the first age of the Silurian Period and of the Llandovery Epoch. The Silurian is in the Paleozoic Era of the Phanerozoic Eon. The Rhuddanian Age began 443.8 ± 1.5 Ma and ended 440.8 ± 1.2 Ma (mi ...
stage of the
Silurian period 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 ...
. This would make the Hirnantian-Rhuddanian anoxia one of the longest-lasting anoxic events in geologic time. The cause of the Hirnantian-Rhuddanian anoxic event is uncertain. Like most global anoxic events, an increased supply of nutrients (such as nitrates and
phosphates In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
) would encourage algal or microbial blooms that deplete oxygen levels in the seawater. The most likely culprits are
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
, which can use
nitrogen fixation Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmo ...
to produce usable nitrogen compounds in the absence of nitrates. Nitrogen isotopes during the anoxic event record high rates of
denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denitr ...
, a biological process which depletes nitrates. The Nitrogen-fixing ability of cyanobacteria would give them an edge over inflexible competitors like
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
. At
Anticosti Island ; moe, Notiskuan; mic, Natigostec , sobriquet = , image_name = RiviereHuileAnticosti.jpg , image_caption = Salmon fisherman on Rivière à l'Huile , image_map ...
, a uranium isotope excursion consistent with anoxia actually occurs prior to indicators of receding glaciation. This may suggest that the Hirnantian-Rhuddanian anoxic event (and its corresponding extinction) began during the glaciation, not after it. Cool temperatures can lead to
upwelling Upwelling is an oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water from deep water towards the ocean surface. It replaces the warmer and usually nutrient-depleted surface water. The nut ...
, cycling nutrients into productive surface waters via air and ocean cycles. Upwelling could instead be encouraged by increasing oceanic stratification through an input of freshwater from melting glaciers. This would be more reasonable if the anoxic event coincided with the end of glaciation, as supported by most other studies. However, oceanic models argue that marine currents would recover too quickly for freshwater disruptions to have a meaningful effect on nutrient cycles. Retreating glaciers could expose more land to weathering, which would be a more sustained source of phosphates flowing into the ocean. There were few clear patterns of extinction associated with the second extinction pulse. Every region and marine environment experienced the second extinction pulse to some extent. Many taxa which survived or diversified after the first pulse were finished off in the second pulse. These include the ''
Hirnantia 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 charac ...
'' brachiopod fauna and ''Mucronaspis'' trilobite fauna, which previously thrived in the cold glacial period. Other taxa such as
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 th ...
and warm-water reef denizens were less affected. Sediments from China 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, ...
seemingly show a more gradual replacement of the ''Hirnantia'' fauna after glaciation. Although this suggests that the second extinction pulse may have been a minor event at best, other paleontologists maintain that an abrupt ecological turnover accompanied the end of glaciation. There may be a correlation between the relatively slow recovery after the second extinction pulse, and the prolonged nature of the anoxic event which accompanied it.


Early Rhuddanian anoxia

Deposition of black graptolite shales continued to be common into the earliest Rhuddanian, indicating that anoxia persisted well into the
Llandovery Llandovery (; cy, Llanymddyfri ) is a market town and community in Carmarthenshire, Wales. It lies on the River Tywi and at the junction of the A40 and A483 roads, about north-east of Carmarthen, north of Swansea and west of Brecon. Hi ...
. A sharp reduction in the average size of many organisms, likely attributable to the Lilliput effect, and the disappearance of many relict taxa from the Ordovician indicate a third extinction interval linked to an expansion of anoxic conditions into shallower shelf environments, particularly in Baltica. This sharp decline in dissolved oxygen concentrations was likely linked to a period of global warming documented by a negative carbon isotope excursion preserved in Baltican sediments.


Metal poisoning

Toxic metals Metal toxicity or metal poisoning is the toxic effect of certain metals in certain forms and doses on life. Some metals are toxic when they form poisonous soluble compounds. Certain metals have no biological role, i.e. are not essential minerals, ...
on the ocean floor may have dissolved into the water when the oceans' oxygen was depleted. An increase in available nutrients in the oceans may have been a factor, and decreased ocean circulation caused by global cooling may also have been a factor. Hg/TOC values from the Peri-Baltic region indicate noticeable spikes in mercury concentrations during the lower late Katian, the Katian-Hirnantian boundary, and the late Hirnantian. The toxic metals may have killed life forms in lower trophic levels of the
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), det ...
, causing a decline in population, and subsequently resulting in starvation for the dependent higher feeding life forms in the chain.


Gamma-ray burst

A minority hypothesis to explain the first burst has been proposed by Philip Ball, Adrian L. Melott, and Brian C. Thomas, suggesting that the initial extinctions could have been caused by a
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 ...
originating from a
hypernova A hypernova (sometimes called a collapsar) is a very energetic supernova thought to result from an extreme core-collapse scenario. In this case, a massive star (>30 solar masses) collapses to form a rotating black hole emitting twin energetic je ...
in a nearby arm of the
Milky Way The Milky Way is the galaxy that includes our Solar System, with the name describing the galaxy's appearance from Earth: a hazy band of light seen in the night sky formed from stars that cannot be individually distinguished by the naked eye ...
galaxy A galaxy is a system of stars, stellar remnants, interstellar gas, dust, dark matter, bound together by gravity. The word is derived from the Greek ' (), literally 'milky', a reference to the Milky Way galaxy that contains the Solar System. ...
, within 6,000
light-year A light-year, alternatively spelled light year, is a large unit of length used to express astronomical distances and is equivalent to about 9.46 trillion kilometers (), or 5.88 trillion miles ().One trillion here is taken to be 1012 ...
s of Earth. A ten-second burst would have stripped the Earth's atmosphere of half of its
ozone Ozone (), or trioxygen, is an inorganic molecule with the chemical formula . It is a pale blue gas with a distinctively pungent smell. It is an allotrope of oxygen that is much less stable than the diatomic allotrope , breaking down in the lo ...
almost immediately, exposing surface-dwelling organisms, including those responsible for planetary
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
, to high levels of
extreme ultraviolet Extreme ultraviolet radiation (EUV or XUV) or high-energy ultraviolet radiation is electromagnetic radiation in the part of the electromagnetic spectrum spanning wavelengths from 124  nm down to 10 nm, and therefore (by the Planck–E ...
radiation. Under this hypothesis, several groups of marine organisms with a planktonic lifestyle were more exposed to UV radiation than groups that lived on the seabed. It is estimated that 20% to 60% of the total phytoplankton biomass on Earth would have been killed in such an event because the oceans were mostly oligotrophic and clear during the Late Ordovician. This is consistent with observations that planktonic organisms suffered severely during the first extinction pulse. In addition,
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
dwelling in shallow water were more likely to become
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
than species dwelling in deep water, also consistent with the hypothetical effects of a galactic gamma-ray burst. A gamma-ray burst could also explain the rapid expansion of glaciers, since the high energy rays would cause
ozone Ozone (), or trioxygen, is an inorganic molecule with the chemical formula . It is a pale blue gas with a distinctively pungent smell. It is an allotrope of oxygen that is much less stable than the diatomic allotrope , breaking down in the lo ...
, a
greenhouse gas A greenhouse gas (GHG or GhG) is a gas that Absorption (electromagnetic radiation), absorbs and Emission (electromagnetic radiation), emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse ...
, to dissociate and its dissociated oxygen atoms to then react with nitrogen to form
nitrogen dioxide Nitrogen dioxide is a chemical compound with the formula . It is one of several nitrogen oxides. is an intermediate in the industrial synthesis of nitric acid, millions of tons of which are produced each year for use primarily in the producti ...
, a darkly-coloured
aerosol An aerosol is a suspension (chemistry), suspension of fine solid particles or liquid Drop (liquid), droplets in air or another gas. Aerosols can be natural or Human impact on the environment, anthropogenic. Examples of natural aerosols are fog o ...
which cools the planet. It would also cohere with the major
δ13C In geochemistry, paleoclimatology, and paleoceanography ''δ''13C (pronounced "delta c thirteen") is an isotopic signature, a measure of the ratio of stable isotopes 13C : 12C, reported in parts per thousand (per mil, ‰). The measure is also ...
isotopic excursion indicating increased sequestration of carbon-12 out of the atmosphere, which would have occurred as a result of nitrogen dioxide, formed after the reaction of nitrogen and oxygen atoms dissociated by the gamma-ray burst, reacting with hydroxyl and raining back down to Earth as
nitric acid Nitric acid is the inorganic compound with the formula . It is a highly corrosive mineral acid. The compound is colorless, but older samples tend to be yellow cast due to decomposition into oxides of nitrogen. Most commercially available nitri ...
, precipitating large quantities of nitrates that would have enhanced wetland productivity and sequestration of carbon dioxide. Although the gamma-ray burst hypothesis is consistent with some patterns at the onset of extinction, there is no unambiguous evidence that such a nearby gamma-ray burst ever happened.


Volcanism and weathering

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 approximatel ...
was preceded by a fall in atmospheric
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
(from 7,000 ppm to 4,400 ppm).Seth A. Young, Matthew R. Saltzman, William I. Ausich, André Desrochers, and Dimitri Kaljo, "Did changes in atmospheric CO2 coincide with latest Ordovician glacial–interglacial cycles?", ''Palaeogeography, Palaeoclimatology, Palaeoecology'', Vol. 296, No. 3–4, 15 October 2010, Pages 376–388.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 is correlated with a burst of volcanic activity that deposited new silicate rocks, which draw CO2 out of the air as they erode. A major role of CO2 is implied by a 2009 paper. Atmospheric and oceanic CO2 levels may have fluctuated with the growth and decay of Gondwanan glaciation. Increased volcanic activity during the early late Katian and around the Katian-Hirnantian boundary is also implied by heightened mercury concentrations relative to total organic carbon. Marine bentonite layers associated with the subduction of the Junggar Ocean underneath the Yili Block have been dated to the late Katian, close to the Katian-Hirnantian boundary. Through the Late Ordovician, outgassing from major volcanism was balanced by heavy weathering of the uplifting
Appalachians 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 ...
and
Caledonides The Caledonian orogeny was a mountain-building era recorded in the northern parts of the British Isles, the Scandinavian Mountains, Svalbard, eastern Greenland and parts of north-central Europe. The Caledonian orogeny encompasses events that o ...
, which sequestered CO2. In the
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 ...
stage the volcanism diminished, and the continued weathering caused a significant and rapid draw down of CO2. This coincides with the rapid and short ice age. More recently, in May 2020, a study suggested the first pulse of mass extinction was caused by volcanism which induced
global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
and anoxia, rather than cooling and glaciation. Higher resolution of species diversity patterns in the Late Ordovician suggest that extinction rates rose significantly in the early or middle Katian stage, several million years earlier than the Hirnantian glaciation. This early phase of extinction is associated with
large igneous province A large igneous province (LIP) is an extremely large accumulation of igneous rocks, including intrusive (sills, dikes) and extrusive (lava flows, tephra deposits), arising when magma travels through the crust towards the surface. The formation ...
(LIP) activity, possibly that of the Alborz LIP of northern Iran, as well as a warming phase known as the Boda event. Other papers have criticised the volcanism hypothesis, claiming that volcanic activity was relatively low in the Ordovician and that superplume and LIP volcanic activity is especially unlikely to have caused the mass extinction at the end of the Ordovician. A 2022 study argued against a volcanic cause of LOME, citing the lack of mercury anomalies and the discordance between deposition of bentonites and
redox Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate (chemistry), substrate change. Oxidation is the loss of Electron, electrons or an increase in the oxidation state, while reduction ...
changes in drillcores from South China straddling the Ordovician-Silurian boundary.


See also

*
Global catastrophic risk A global catastrophic risk or a doomsday scenario is a hypothetical future event that could damage human well-being on a global scale, even endangering or destroying modern civilization. An event that could cause human extinction or permanen ...
*
Near-Earth supernova A near-Earth supernova is an explosion resulting from the death of a star that occurs close enough to the Earth (roughly less than 10 to 300 parsecs (30 to 1000 light-years) away) to have noticeable effects on Earth's biosphere. An estima ...
*
Anoxic event Oceanic anoxic events or anoxic events ( anoxia conditions) describe periods wherein large expanses of Earth's oceans were depleted of dissolved oxygen (O2), creating toxic, euxinic (anoxic and sulfidic) waters. Although anoxic events have not ...
*
Late Devonian extinction The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major ex ...
*
Capitanian mass extinction event The Capitanian mass extinction event, also known as the end-Guadalupian extinction event or the pre-Lopingian crisis was an extinction event that predated the end-Permian extinction event and occurred around 260 million years ago during a period ...
*
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 ...
*
Triassic–Jurassic extinction event The Triassic–Jurassic (Tr-J) extinction event, often called the end-Triassic extinction, marks the boundary between the Triassic and Jurassic periods, , and is one of the top five major extinction events of the Phanerozoic eon, profoundly affect ...
*
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
*
Andean-Saharan 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 ...


Sources


Further reading

* * *


External links


Jacques Veniers, "The end-Ordovician extinction event"
abstract of Hallam and Wignall, 1997. {{DEFAULTSORT:Ordovician-Silurian extinction event Extinction events History of climate variability and change Late Ordovician extinctions Silurian events