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The Cenomanian-Turonian boundary event, also known as the Cenomanian-Turonian extinction, Cenomanian-Turonian Oceanic Anoxic Event ( OAE 2), and referred to also as the Bonarelli Event or Level, was an anoxic
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 fall in the diversity and abundance of multicellular organisms. It occ ...
in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. The Cenomanian-Turonian oceanic anoxic event is considered to be the most recent truly global oceanic anoxic event in Earth's geologic history. There was a large carbon cycle disturbance during this time period, signified by a large positive carbon isotope excursion. However, apart from the
carbon cycle The carbon cycle is a part of the biogeochemical cycle where carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of Earth. Other major biogeochemical cycles include the nitrogen cycle and the water cycl ...
disturbance, there were also large disturbances in the ocean's
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
,
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
,
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
, sulphur, and iron cycles.


Background

The Cenomanian and Turonian stages were first noted by
D'Orbigny Alcide Charles Victor Marie Dessalines d'Orbigny (6 September 1802 – 30 June 1857) was a French naturalist who made major contributions in many areas, including zoology (including malacology), palaeontology, geology, archaeology and anthropol ...
between 1843 and 1852. The global type section for this boundary is located in the Bridge Creek Limestone Member of the Greenhorn Formation near
Pueblo, Colorado Pueblo ( ) is a List of municipalities in Colorado#Home rule municipality, home rule municipality that is the county seat of and the List of municipalities in Colorado, most populous municipality in Pueblo County, Colorado, United States. The ...
, which are bedded with the Milankovitch orbital signature. Here, a positive carbon-isotope event is clearly shown, although none of the characteristic, organic-rich black
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
is present. It has been estimated that the isotope shift lasted approximately 850,000 years longer than the black shale event, which may be the cause of this anomaly in the Colorado type section. A significantly expanded OAE2 interval from southern Tibet documents a complete, more detailed, and finer-scale structures of the positive carbon isotope excursion that contains multiple shorter-term carbon isotope stages amounting to a total duration of 820 ±25 ka. The level is also known as the Bonarelli Event because of layer of thick, black shale that marks the boundary and was first studied by in 1891. It is characterized by interbedded black shales,
chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a prec ...
and radiolarian sands and is estimated to span a 400,000-year interval. Planktonic foraminifera do not exist in this Bonarelli Level, and the presence of radiolarians in this section indicates relatively high productivity and an availability of nutrients. In the Western Interior Seaway, the Cenomanian-Turonian boundary event is associated with the Benthonic Zone, characterised by a higher density of benthic foraminifera relative to planktonic foraminifera, although the timing of the appearance of the Benthonic Zone is not uniformly synchronous with the onset of the oceanic anoxic event and is thus cannot be used to consistently demarcate its beginning.


Timeline

Selby et al. in 2009 concluded the OAE 2 occurred approximately 91.5 ± 8.6 Ma, though estimates published by Leckie et al. (2002) are given as 93–94 Ma. The Cenomanian-Turonian boundary has been refined in 2012 to 93.9 ± 0.15 Ma. The total duration of OAE2 has been estimated at 0.9 Myr, 0.82 ± 0.025 Myr, or 0.71 ± 0.17 Myr. At high latitudes, the event lasted for a shorter time: only ~600 kyr. Biodiversity patterns of planktic foraminifera indicate that the Cenomanian-Turonian extinction occurred in five phases. Phase I, which took place from 313,000 to 55,000 years before the onset of the anoxic event, witnessed a stratified water column and high planktonic foraminiferal diversity, suggesting a stable marine environment. Phase II, characterised by significant environmental perturbations, lasted from 55,000 years before OAE2 until its onset and witnessed a decline in rotaliporids and heterohelicids, a zenith of schackoinids and hedbergellids, a 'large form eclipse' during which foraminifera exceeding 150 microns disappeared, and the start of a trend of dwarfism among many foraminifera. This phase also saw an enhanced oxygen minimum zone and increased productivity in surface waters. Phase III lasted for 100,000 to 900,000 years and was coincident with the Bonarelli Level's deposition and exhibited extensive proliferation of radiolarians, indicative of extremely eutrophic conditions. Phase IV lasted for around 35,000 years and was most notable for the increase in the abundance of hedbergellids and schackoinids, being extremely similar to Phase II, with the main difference being that rotaliporids were absent from Phase IV. Phase V was a recovery interval lasting 118,000 years and marked the end of the 'large form eclipse' that began in Phase II; heterohelicids and hedbergellids remained in abundance during this phase, pointing to continued environmental disturbance during this phase.


Causes


Climate change

Earth pronouncedly warmed just before the beginning of OAE2. The Cenomanian-Turonian interval represents one of the hottest intervals of the entire Phanerozoic eon, and it boasted the highest carbon dioxide concentrations of the Cretaceous period. Even before OAE2, during the late Cenomanian, tropical sea surface temperatures (SSTs) were very warm, about 27-29 Â°C. The onset of OAE2 was concurrent with a 4-5 °C rise in shelf sea temperatures. Mean tropical SSTs during OAE2 have been conservatively estimated to have been at least 30 Â°C, but may have reached as high as 36 Â°C. Minimum SSTs in mid-latitude oceans were >20 Â°C. This exceptional warmth persisted until the Turonian-Coniacian boundary. One possible cause of this hothouse was sub-oceanic volcanism. During the middle of the Cretaceous period, the rate of crustal production reached a peak, which may have been related to the rifting of the newly formed Atlantic Ocean. It was also caused by the widespread melting of hot mantle plumes under the
ocean crust Oceanic crust is the uppermost layer of the oceanic portion of the Plate tectonics, tectonic plates. It is composed of the upper oceanic crust, with pillow lavas and a dike (geology), dike complex, and the lower oceanic crust, composed of troct ...
, at the base of the
lithosphere A lithosphere () is the rigid, outermost rocky shell of a terrestrial planet or natural satellite. On Earth, it is composed of the crust and the lithospheric mantle, the topmost portion of the upper mantle that behaves elastically on time ...
, which may have resulted in the thickening of the oceanic crust in the
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
and
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
s. The resulting volcanism would have sent large quantities of carbon dioxide into the atmosphere, leading to an increase in global temperatures. Greenhouse gas release was further increased by the degassing of organic-rich sediments intruded into by volcanic sills. Several independent events related to large igneous provinces (LIPs) occurred around the time of OAE2. A multitude of LIPs were active during OAE2: the
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
,
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
, Gorgona, Ontong Java, and High Arctic LIPs. The abundance of LIPs at this time reflects a major overturning in mantle convection. Trace metals such as chromium (Cr),
scandium Scandium is a chemical element; it has Symbol (chemistry), symbol Sc and atomic number 21. It is a silvery-white metallic d-block, d-block element. Historically, it has been classified as a rare-earth element, together with yttrium and the lantha ...
(Sc),
copper Copper is a chemical element; it has symbol Cu (from Latin ) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkish-orang ...
(Cu) and
cobalt Cobalt is a chemical element; it has Symbol (chemistry), symbol Co and atomic number 27. As with nickel, cobalt is found in the Earth's crust only in a chemically combined form, save for small deposits found in alloys of natural meteoric iron. ...
(Co) have been found at the Cenomanian-Turonian boundary, which suggests that an LIP could have been one of the main basic causes involved in the contribution of the event. The timing of the peak in trace metal concentration coincides with the middle of the anoxic event, suggesting that the effects of the LIPs may have occurred during the event, but may not have initiated the event. Other studies linked the
lead Lead () is a chemical element; it has Chemical symbol, symbol Pb (from Latin ) and atomic number 82. It is a Heavy metal (elements), heavy metal that is density, denser than most common materials. Lead is Mohs scale, soft and Ductility, malleabl ...
(Pb)
isotope Isotopes are distinct nuclear species (or ''nuclides'') of the same chemical element. They have the same atomic number (number of protons in their Atomic nucleus, nuclei) and position in the periodic table (and hence belong to the same chemica ...
s of OAE-2 to the Caribbean-Colombian and the Madagascar LIPs. An osmium isotope excursion coeval with OAE2 strongly suggests submarine volcanism as its cause; in the Pacific, an unradiogenic osmium spike began about 350 kyr before the onset of OAE2 and terminated around 240 kyr after OAE2's beginning; the osmium isotope data from a highly expanded OAE2 interval in southern Tibet show multiple osmium excursions with the most pronounced one lagging the onset of OAE2 by ≈50 kyr that was probably related to the ocean connectivity change at ~94.5 Ma. Osmium data also reveal that three distinct pulses of intense volcanism occurred ~60, ~270, and ~400 kyr after OAE2's onset, prolonging it. Positive
neodymium Neodymium is a chemical element; it has Symbol (chemistry), 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 element, rare-earth metals. It is a hard (physics), hard, sli ...
isotope excursions provide additional indications of pervasive volcanism as a cause of OAE2. Enrichments in
zinc Zinc is a chemical element; it has symbol Zn and atomic number 30. It is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodic tabl ...
further bolster and reinforce the existence of extensive hydrothermal volcanism, as do extreme negative δ53Cr excursions. The absence of geographically widespread mercury (Hg) anomalies resulting from OAE2 has been suggested to be because of the limited dispersal range of this heavy metal by submarine volcanism. A modeling study performed in 2011 confirmed that it is possible that a LIP may have initiated the event, as the model revealed that the peak amount of carbon dioxide degassing from volcanic LIP degassing could have resulted in more than 90 percent global deep-ocean anoxia. Later on, when anoxia became widespread, the production of
nitrous oxide Nitrous oxide (dinitrogen oxide or dinitrogen monoxide), commonly known as laughing gas, nitrous, or factitious air, among others, is a chemical compound, an Nitrogen oxide, oxide of nitrogen with the Chemical formula, formula . At room te ...
, a greenhouse gas about 265 times more potent than carbon dioxide, drastically increased because of elevated nitrification and denitrification rates. This powerful positive feedback mechanism is what may have enabled extremely hot temperatures to persist in spite of the supercharged organic carbon burial associated with anoxic events.


Plenus Cool Event

Large-scale organic carbon burial acted as a negative feedback loop that partially mitigated the warming effects of volcanic discharge of carbon dioxide, resulting in the Plenus Cool Event during the ''Metoicoceras geslinianum'' European ammonite biozone. Global average temperatures fell to around 4 Â°C lower than they were pre-OAE2. Equatorial SSTs dropped by 2.5–5.5 Â°C. This cooling event was insufficient at completely stopping the rise in global temperatures. This
negative feedback Negative feedback (or balancing feedback) occurs when some function (Mathematics), function of the output of a system, process, or mechanism is feedback, fed back in a manner that tends to reduce the fluctuations in the output, whether caused ...
was ultimately overridden, as global temperatures continued to shoot up in sync with continued volcanic release of carbon dioxide following the Plenus Cool Event, although this theory has been criticised and the warming after the Plenus Cool Event attributed to decreased silicate weathering instead.


Ocean acidification

Within the oceans, the emission of SO2, H2S, CO2, and halogens would have increased the acidity of the water, causing the dissolution of carbonate, and a further release of carbon dioxide. Evidence of ocean acidification can be gleaned from δ44/40Ca increases coeval with the extinction event, as well as coccolith malformation and dwarfism. Lithologies characterised by low calcium carbonate concentrations predominated during intervals of carbonate compensation depth shoaling. Ocean acidification was exacerbated by a positive feedback loop of increased heterotrophic respiration in highly biologically productive waters, elevating seawater concentrations of carbon dioxide and further decreasing pH.


Anoxia and euxinia

When the volcanic activity declined, this run-away
greenhouse effect The greenhouse effect occurs when greenhouse gases in a planet's atmosphere insulate the planet from losing heat to space, raising its surface temperature. Surface heating can happen from an internal heat source (as in the case of Jupiter) or ...
would have likely been put into reverse. The increased CO2 content of the oceans could have increased organic productivity in the ocean surface waters. The consumption of this newly abundant organic life by aerobic bacteria would produce anoxia and mass extinction. An acceleration of the hydrological cycle induced by warmer global temperatures drove greater fluxes of nutrient runoff into the oceans, fuelling primary productivity. The global environmental disturbance that resulted in these conditions increased atmospheric and oceanic temperatures. Extreme hothouse conditions encouraged ocean stratification. Boundary sediments show an enrichment of trace elements, and contain elevated δ13C values. The positive δ13C excursion found at the Cenomanian-Turonian boundary is one of the main carbon isotope events of the Mesozoic. It represents one of the largest disturbances in the global carbon cycle from the past 110 million years. This δ13C excursion indicates a significant increase in the burial rate of organic carbon, indicating the widespread deposition and preservation of organic carbon-rich sediments and that the ocean was depleted of oxygen at the time. Depletion of
manganese Manganese is a chemical element; it has Symbol (chemistry), symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese was first isolated in the 1770s. It is a transition m ...
in sediments corresponding to OAE2 provides additional strong evidence of severe bottom water oxygen depletion. An increase in the abundance of the planktonic foraminifer '' Heterohelix'' provides further evidence still of anoxia. The resulting elevated levels of carbon burial would account for the black shale deposition in the ocean basins. The proto-North Atlantic in particular was a hotbed of carbon burial during OAE2 as it was in later, less severe anoxic events. Though anoxia was prevalent throughout the interval, there were transient periods of reoxygenation during OAE2. Sulphate reduction increased during OAE2, causing euxinia, a type of anoxia defined by sulphate reduction and hydrogen sulphide production, to occur during OAE2, as revealed by negative δ53Cr excursions, positive δ98Mo excursions, a drawdown of seawater
molybdenum Molybdenum is a chemical element; it has Symbol (chemistry), symbol Mo (from Neo-Latin ''molybdaenum'') and atomic number 42. The name derived from Ancient Greek ', meaning lead, since its ores were confused with lead ores. Molybdenum minerals hav ...
, and molecular biomarkers of green sulfur bacteria. Although euxinia was not uncommon in the latter part of the Cenomanian, it only expanded into the photic zone during OAE2 itself. OAE2 began on the southern margins of the proto-North Atlantic, from where anoxia spread across the rest of the proto-North Atlantic and then into the Western Interior Seaway (WIS) and the epicontinental seas of the Western Tethys. Anoxic waters spread rapidly throughout the WIS due to marine transgression and a powerful cyclonic circulation resulting from an imbalance between precipitation in the north and evaporation in the south. Anoxia was especially intense in the eastern North Sea, evidenced by its very positive δ13C values. Thanks to persistent upwelling, some marine regions, such as the South Atlantic, were able to remain partially oxygenated at least intermittently. Indeed, redox states of oceans vary geographically, bathymetrically and temporally during OAE2.


Milankovitch cycles

It has been hypothesised that the Cenomanian-Turonian boundary event occurred during a period of very low variability in Earth's insolation, which has been theorised to be the result of coincident nodes in all orbital parameters. Barring chaotic perturbations in Earth's and Mars' orbits, the simultaneous occurrence of nodes of
orbital eccentricity In astrodynamics, the orbital eccentricity of an astronomical object is a dimensionless parameter that determines the amount by which its orbit around another body deviates from a perfect circle. A value of 0 is a circular orbit, values be ...
, axial precession, and obliquity on Earth occurs approximately every 2.45 million years. Numerous other oceanic anoxic events occurred throughout the extremely warm greenhouse conditions of the Middle Cretaceous, and it has been suggested that these Middle Cretaceous ocean anoxic events occurred cyclically in accordance with orbital cycle patterns. The mid-Cenomanian Event (MCE), which occurred in the ''Rotalipora cushmani'' planktonic foraminifer biozone, has been argued to be another example supporting this hypothesis of regular oceanic anoxic events governed by Milankovitch cycles. The MCE took place approximately 2.4 million years before the Cenomanian-Turonian oceanic anoxic event, roughly at the time when an anoxic event would be expected to occur given such a cycle. Geochemical evidence from a sediment core in the Tarfaya Basin is indicative of the main positive carbon isotope excursion occurring during a prolonged eccentricity minimum. Carbon isotope shifts smaller in scale observed in this core likely reflected variability in obliquity. Ocean Drilling Program Site 1138 in the Kerguelen Plateau yields evidence of a 20,000 to 70,000 year periodicity in changes in sedimentation, suggesting that either obliquity or precession governed the large-scale burial of organic carbon. Within the OAE2 positive δ13C excursion, short eccentricity scale carbon isotope variability is documented in a significantly expanded OAE2 interval from southern Tibet; periodic negative δ13C excursions paced by the short eccentricity cycle are easily detectable in southwestern Utah too.


Enhanced phosphorus recycling

The phosphorus retention ability of seafloor sediments declined during OAE2, revealed by a decline in reactive phosphorus species within OAE2 sediments. The mineralisation of seafloor phosphorus into apatite was inhibited by the significantly lower pH of seawater and much warmer temperatures during the Cenomanian and Turonian compared to the present day, which meant that significantly more phosphorus was recycled back into ocean water after being deposited on the sea floor during this time. This would have intensified a positive feedback loop in which phosphorus is recycled faster into anoxic seawater compared to oxygen-rich water, which in turn fertilises the water, causes increased eutrophication, and further depletes the seawater of oxygen. The influx of volcanically erupted and chemically weathered sulphate into the ocean also inhibited phosphorus burial by increasing hydrogen sulphide production, which hinders the burial of phosphorus through sorption to iron oxyhydroxide phases. OAE2 may have occurred during a peak in a 5-6 Myr cycle governing phosphorus availability; at this and other peaks in this oscillation, an increase in chemical weathering would have increased the marine phosphorus inventory and sparked a positive feedback loop of increasing productivity, anoxia, and phosphorus recycling that was only ended by a negative feedback of increased atmospheric oxygenation and wildfire activity that decreased chemical weathering, a feedback which operated on a much longer timescale. Enhanced phosphorus recycling would have resulted in an abundance of nitrogen fixing bacteria, increasing the availability of yet another limiting nutrient and supercharging primary productivity through
nitrogen fixation Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
. The ratio of bioavailable nitrogen to bioavailable phosphorus, which is 16:1 in the present, fell precipitously as the ocean transitioned from being oxic and nitrate-dominated to anoxic and ammonium-dominated. A potent feedback loop of nitrogen fixation, productivity, deoxygenation, nitrogen removal, and phosphorus recycling was created. Bacterial hopanoids indicate populations of nitrogen fixing cyanobacteria were high during OAE2, providing a rich supply of nitrates and nitrites. Negative δ15N values reveal the dominance of ammonium through regenerative nutrient loops in the proto-North Atlantic.


Decreased sulphide oxidation

In the present day, sulphidic waters are generally prevented from spreading throughout the water column by the oxidation of sulphide with nitrate. However, during OAE2, the inventory of seawater nitrate was lower, meaning that chemolithoautotrophic oxidation of sulphides with nitrates was inefficient at preventing the spread of euxinia.


Sea level rise

A marine transgression in the latest Cenomanian resulted in an increase in average water depth, causing seawater to become less eutrophic in shallow, epicontinental seas. Turnovers in marine biota in such epicontinental seas have been suggested to be driven more so by changes in water depth rather than anoxia. Sea level rise also contributed to anoxia by transporting terrestrial plant matter from inundated lands seaward, providing an abundant source of sustenance for eutrophicating microorganisms.


Geological effects


Phosphate deposition

A phosphogenic event occurred in the Bohemian Cretaceous Basin during the peak of oceanic anoxia. Phosphorus liberation in the pore water environment, several centimetres below the interface between seafloor sediments and the water column, enabled the precipitation of phosphate through biological mediation by microorganisms.


Increase in weathering

Strontium and calcium isotope ratios both indicate that silicate weathering increased over the course of OAE2. Because of its effectiveness as a carbon sink on geologic timescales, the uptick in sequestration of carbon dioxide by the lithosphere may have helped to stabilise global temperatures after global temperatures soared. Particularly so at high latitudes, where the increase in weatherability was very pronounced.


Biotic effects


Changes in oceanic biodiversity and its implications

Although some early studies suggested the marine biodiversity decline observed during the Cenomanian-Turonian transition was not a real extinction but instead represented an artifact of preservation, recent work confirms that significant extinctions were experienced by vertebrates, invertebrates, and microbes. The event brought about the extinction of the pliosaurs, and most ichthyosaurs. Coracoids of
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age were once interpreted by some authors as belonging to ichthyosaurs, but these have since been interpreted as
plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
elements instead. Dolichosaurids became rare after OAE2, whereas mosasauroid diversity bloomed in its aftermath. Tethysuchians experienced a significant faunal turnover, and post-OAE2 tethysuchians tended to inhabit warmer environments compared to pre-OAE2 tethysuchians. Although the cause is still uncertain, the result starved the Earth's oceans of oxygen for nearly half a million years, causing the extinction of approximately 27 percent of marine invertebrates, including certain planktic and
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
, mollusks, bivalves, dinoflagellates and calcareous nannofossils. Planktonic foraminifera suffered from the expansion of oxygen minimum zones; those that dwelt in deeper waters were especially hard hit. In Whadi El Ghaib, a site in Sinai,
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
, the foraminiferal community during OAE2 was low in diversity and dominated by taxa that were extremely tolerant of low salinity, anoxic water. In the southeastern Indian Ocean, off the coast of Australia, the planktonic foraminifer ''Microhedbergella'' was highly abundant, while ''Heterohelix'' thrived in reducing waters in the South Atlantic, as well as in the Chalk Sea. Benthic foraminifera suffered noticeable losses. The benthic foraminifera ''Gavelella berthelini'' and ''Lingulogavelinella globosa'' dominated during deoxygenated conditions in Poland. The alterations in diversity of various marine invertebrate
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
such as calcareous nannofossils are reflective and characteristic of oligotrophy and ocean warmth in an environment with short spikes of productivity followed by long periods of low fertility. A study performed in the Cenomanian-Turonian boundary of Wunstorf, Germany, reveal the uncharacteristic dominance of a calcareous nannofossil species, ''Watznaueria'', present during the event. Unlike the ''Biscutum'' species, which prefer mesotrophic conditions and were generally the dominant species before and after the C/T boundary event; ''Watznaueria'' species prefer warm, oligotrophic conditions. In the Ohaba-Ponor section in
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
, the presence of ''Watznaueria barnesae'' indicates warm conditions, while the abundances of ''Biscutum constans'', ''Zeugrhabdotus erectus'', and ''Eprolithus floralis'' peak during cool intervals. Sites in
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
, and
Sicily Sicily (Italian language, Italian and ), officially the Sicilian Region (), is an island in the central Mediterranean Sea, south of the Italian Peninsula in continental Europe and is one of the 20 regions of Italy, regions of Italy. With 4. ...
show an inverse relationship between atmospheric carbon dioxide levels and the size of calcareous nannoplankton. Radiolarians also suffered heavy losses in OAE2, one of their highest diversity losses in the Cretaceous. Bivalves declined significantly in diversity during the leadup to the δ13Corg peak of OAE2. Rudist bivalves suffered high extinction rates combined with low origination rates during OAE2. Ammonoids suffered during the crisis, though anoxia was not the main driver of their declines in diversity. Ammonoid diversity losses were primarily concentrated in the seas around Europe; elsewhere, they were negligibly affected. The diversity of
trace fossils A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
sharply plummeted during the beginning of the Cenomanian-Turonian boundary event. The recovery interval after the anoxic event's conclusion features an abundance of ''Planolites'' and is characterised overall by a high degree of
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a ...
. At the time, there were also peak abundances of the green algal groups '' Botryococcus'' and prasinophytes, coincident with pelagic sedimentation. The abundances of these algal groups are strongly related to the increase of both the oxygen deficiency in the water column and the total content of organic carbon. The evidence from these algal groups suggest that there were episodes of halocline stratification of the water column during the time. A species of freshwater dinocyst—''Bosedinia''—was also found in the rocks dated to the time and these suggest that the oceans had reduced salinity.


Changes in terrestrial biodiversity

No major change in terrestrial ecosystems is known to have been synchronous with the marine transgression associated with OAE2, although the loss of freshwater
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
habitat has been speculated to have possibly resulted in the demise of some freshwater taxa. In fossiliferous rocks in southwestern
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
, a local
extirpation Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with extinction, global extinctions. Local extinctions ...
of some
metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as wel ...
ns and
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuary ...
vertebrates is associated with the later marine regression following OAE2 in the Turonian. Among mammals, diversity changes likely reflect shifting ranges and changes in ecology rather than a true extinction event. Whatever the nature and magnitude of terrestrial extinctions at or near the Cenomanian-Turonian boundary was, it was most likely caused mainly by other factors than eustatic sea level fluctuations. The effect of the ecological crisis on terrestrial plants has been concluded to have been inconsequential, in contrast to extinction events driven by terrestrial large igneous provinces. However, while terrestrial plants did persist even during the exceptional warmth, the Plenus Cool Event facilitated a notable expansion of angiosperm-dominated savanna ecosystems.


See also

* Biodiversity of the Cenomanian and Turonian ** Cenomanian life ** Turonian life ** Cenomanian extinctions ** Turonian extinctions *
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 fall in the diversity and abundance of multicellular organisms. It occ ...
* Timeline of extinctions in the Holocene ** Toarcian turnover **
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...


References


Further reading

* {{ExtEvent nav Extinction events Ceno Paleoclimatology Anoxic events Isotope excursions