Terrestrial floras at the Triassic-Jurassic Boundary in Europe.

Proportions of range-through diversities of higher taxonomic categories of microfloral elements over the Middle Triassic–Early Jurassic interval (From Barbacka et al., 2017)

Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. Most of those events are associated with global warming and proximal killers such as marine anoxia. Volcanogenic-atmospheric kill mechanisms include ocean acidification, toxic metal poisoning, acid rain, increased UV-B radiation, volcanic darkness, cooling and photosynthetic shutdown. The mass extinction at the Triassic-Jurassic Boundary (TJB) has been linked to the eruption of the Central Atlantic Magmatic Province (CAMP), a large igneous province emplaced during the initial rifting of Pangea. Another theory is that a huge impact was the trigger of the extinction event. At least two craters impact were reported by the end of the Triassic. The Manicouagan Impact crater in the Côte-Nord region of Québec, Canada was caused by the impact of a 5km diameter asteroid, and it was suggested that could be part of a multiple impact event which also formed the Rochechouart crater in France, Saint Martin crater in Canada, Obolon crater in Ukraine, and the Red Wing crater in USA (Spray et al., 1998).

Photographs of some Rhaetian–Hettangian spores and pollen from the Danish Basin (From Lindström, 2015)

Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one. But there was no mass extinction of European terrestrial plants during the TJB. The majority of genera and a high percentage of species still existed in its later stages, and replacement seems to have been local, explainable as a typical reaction to an environmental disturbance. In Greenland, for example, the replacement of Triassic wide-leaved forms with Jurassic narrow-leaved forms was linked to the reaction of plants to increased wildfire. In Sweden, wildfire in the late Rhaetian and early Hettangian caused large-scale burning of conifer forests and ferns, and the appearance of new swampy vegetation. In Austria and the United Kingdom, conifers and seed ferns were replaced by ferns, club mosses and liverworts. In Hungary, there was a high spike of ferns and conifers at the TJB, followed by a sudden decrease in the number of ferns along with an increasing share of swamp-inhabiting conifers.

Although certain taxa/families indeed became extinct by the end of the Triassic (e.g. Peltaspermales), the floral changes across Europe were rather a consequence of local changes in topography.

References:

Maria Barbacka, Grzegorz Pacyna, Ádam T. Kocsis, Agata Jarzynka, Jadwiga Ziaja, Emese Bodor , Changes in terrestrial floras at the TriassicJurassic Boundary in Europe, Palaeogeography, Palaeoclimatology, Palaeoecology (2017), doi: 10.1016/j.palaeo.2017.05.024

S. Lindström, Palynofloral patterns of terrestrial ecosystem change during the end-Triassic event — a review, Geol. Mag., 1–23 (2015) https://doi.org/10.1017/S0016756815000552

Van de Schootbrugge, B., Quan, T.M., Lindström, S., Püttmann, W., Heunisch, C., Pross, J., Fiebig, J., Petschick, R., Röhling, H.-G., Richoz, S., Rosenthal, Y., Falkowski, P. G., 2009. Floral changes across the Triassic/Jurassic boundary linked to flood basalt volcanism. Nat. Geosci. 2, 589–594. doi: 10.1038/NGEO577.

N.R. Bonis, W.M. Kürschner, Vegetation history, diversity patterns, and climate change across the Triassic/Jurassic boundary, Paleobiology, 8 (2) (2012), pp. 240–264 https://doi.org/10.1666/09071.1

A Tale of Two Exctintions.

The permian triassic boundary at Meishan, China (Photo: Shuzhong Shen)

The Permian Triassic boundary at Meishan, China (Photo: Shuzhong Shen)

Extinction is the ultimate fate of all species. The fossil record indicates that more than 95% of all species that ever lived are now extinct. Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. In 1982, Jack Sepkoski and David M. Raup identified five major extinction events in Earth’s history: at the end of the Ordovician period, Late Devonian, End Permian, End Triassic and the End Cretaceous. These five events are know as the Big Five.

The end-Permian extinction is the most severe biotic crisis in the fossil record, with as much as 95% of the marine animal species and a similarly high proportion of terrestrial plants and animals going extinct . This great crisis occurred 252 million years ago (Ma) during an episode of global warming. The End-Triassic Extinction  is probably the least understood of the big five. Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. Although it’s almost impossible briefly summarize all the changes in biodiversity associated with both extinction events, we can describe their broad trends.

 

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

Both extinction events are commonly linked to the emplacement of the large igneous provinces of the Siberian Traps and the Central Atlantic Magmatic Province. Massive volcanic eruptions with lava flows, released large quantities of sulphur dioxide, carbon dioxide, thermogenic methane and large amounts of HF, HCl, halocarbons and toxic aromatics and heavy metals into the atmosphere. Furthermore, volcanism contribute gases to the atmosphere, such as Cl, F, and CH3Cl from coal combustion, that suppress ozone formation. Acid rain likely had an impact on freshwater ecosystems and may have triggered forest dieback. Mutagenesis observed in the Lower Triassic herbaceous lycopsid Isoetales has been attributed to increased levels of UV-radiation. Charcoal records point to forest fires as a common denominator during both events. Forest dieback was accompanied by the proliferation of opportunists and pioneers, including ferns and fern allies. Moreover, both events led to major schisms in the dominant terrestrial herbivores  and apex predators, including the late Permian extinction of the pariaeosaurs and many dicynodonts and the end-Triassic loss of crurotarsans (van de Schootbrugge and Wignall, 2016).

Aberrant pollen and spores from the end-Triassic extinction interval (scale bars are 20 μm). (a) Ricciisporites tuberculatus from the uppermost Rhaetian deposits at Northern Ireland (adapted from van de Schootbrugge and Wignall, 2016)

Aberrant pollen and spores from the end-Triassic extinction interval (scale bars are 20 μm). (a) Ricciisporites
tuberculatus and b) Kraeuselisporites reissingerii (adapted from van de Schootbrugge and Wignall, 2016)

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval. A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988). Palynological records from across Europe provide evidence for complete loss of tree-bearing vegetation reflected in a strong decline in pollen abundance at the end of the Triassic. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

Comparison of extinction rates for calcareous organisms during the end-Permian and end-Triassic extinction event (from van de Schootbrugge and Wignall, 2016)

Comparison of extinction rates for calcareous organisms during the end-Permian and end-Triassic extinction event (from van de Schootbrugge and Wignall, 2016)

Rapid additions of carbon dioxide during extreme events may have driven surface waters to undersaturation. Acidification affects the biogeochemical dynamics of calcium carbonate, organic carbon, nitrogen, and phosphorus in the ocean and interferes with a range of processes, including growth, calcification, development, reproduction and behaviour in a wide range of marine organisms like foraminifera, planktonic coccolithophores, pteropods and other molluscs,  echinoderms, corals, and coralline algae. Both extinction events led to near-annihilation of cnidarian clades and other taxa responsible for reef construction, resulting in ‘reef gaps’ that lasted millions of years. Black shales deposited across both extinction events also contain increased concentrations of the biomarker isorenieratane, a pigment from green sulphur bacteria, suggesting that the photic zone underwent prolonged periods of high concentrations of hydrogen sulphide. Following the end-Triassic extinction, Early Jurassic shallow seas witnessed recurrent euxinia over a time span of 25 million years, culminating in the Toarcian Oceanic Anoxic Event.

 

References:

BAS VAN DE SCHOOTBRUGGE and PAUL B. WIGNALL (2016). A tale of two extinctions: converging end-Permian and end-Triassic scenarios. Geological Magazine, 153, pp 332-354. doi:10.1017/S0016756815000643.

BACHAN, A. & PAYNE, J. L. 2015. Modelling the impact of pulsed CAMP volcanism on pCO2 and δ13C across the Triassic-Jurassic transition. Geological Magazine, published online

Retallack, G.J. 2013. Permian and Triassic greenhouse crises. Gondwana Research 24:90–103.

 

Unlocking the secrets of the Crater of Doom.

Luis and Walter Alvarez at the K-T Boundary in Gubbio, Italy, 1981 (From Wikimedia Commons)

Luis and Walter Alvarez at the K-T Boundary in Gubbio, Italy, 1981 (From Wikimedia Commons)

The noble and ancient city of Gubbio laid out along the ridges of Mount Ingino in Umbria, was founded by Etruscans between the second and first centuries B.C. The city has an exceptional artistic and monumental heritage which includes marvelous examples of Gothic architecture, like the Palazzo dei Consoli and the Palazzo del Bargello. The rich history of the city is recorded in those buildings. Outside the city, there are exposures of pelagic sedimentary rocks that recorded more than 50 million years of Earth’s history. In the 1970s it was recognized that these pelagic limestones carry a record of the reversals of the magnetic field. The  K-Pg boundary occurs within a portion of the sequence formed by pink limestone containing a variable amount of clay. This limestone, know as the “Scaglia rossa”, is composed by calcareous nannofossils and planktonic foraminifera.

In 1977, Walter Alvarez – an associate professor of geology University of California, Berkeley – was collecting samples of the limestone rock for a paleomagnetism study. He found that the foraminifera from the Upper Cretaceous (notably the genus Globotruncana) disappear abruptly and are replaced by Tertiary foraminifera. The extinction of most of the nannoplankton was simultaneus with the disappearance of the foraminifera (Alvarez et al., 1980).

Forams from the Upper Cretaceous vs. the post-impact foraminifera from the Paleogene. (Images from the Smithsonian Museum of Natural History)

Forams from the Upper Cretaceous vs. the post-impact foraminifera from the Paleogene. (Images from the Smithsonian Museum of Natural History)

At Caravaca on the southeast coast of Spain, Jan Smith, a Dutch geologist, had noticed a similar pattern of changes in forams in rocks around the K-T boundary. Looking for clues, Smith contacted to Jan Hertogen who found high iridium values at the clay boundary. At the same time, Walter Alvarez  gave his father, Luis Alvarez – an American physicist who won the  Nobel Prize in Physics in 1968 – a small polished cross-section of Gubbio  K-Pg boundary rock. The Alvarez gave some samples to Frank Asaro and Helen Michel, who had developed a new technique called neutron activation analysis (NAA). They also discovered the same iridium anomaly. The sea cliff of Stevns Klint, about 50 km south of Copenhagen, shows the same pattern of extinction and iridium anomaly. Another sample from New Zeland also exhibits a spike of iridium. The phenomenon was global.

Iridium is rare in the Earth’s crust but metal meteorites are often rich in iridium. Ten years before the iridium discovery, physicist Wallace Tucker and paleontologist Dale Russell proposed  that a supernova caused the mass extinction at the K-Pg boundary. Luis Alvarez realised that  a supernova would have also released plutonium-244, but there was no plutonium in the sample at all. They concluded that the anomalous iridium concentration at the K-Pg boundary is best interpreted as the result of an asteroid impact, which would explain the iridium and the lack of plutonium. In 1980, they published their seminal paper on Science, along with Asaro and Michel, and ignited a huge controversy. They even calculated the size of the asteroid (about 7 km in diameter) and the crater that this body might have caused (about 100–200 km across).

A paleogeographic map of the Gulf of Mexico at the end of the Cretaceous (From Vellekoop, 2014)

A paleogeographic map of the Gulf of Mexico at the end of the Cretaceous (From Vellekoop, 2014)

In 1981, Pemex (a Mexican oil company) identified Chicxulub as the site of a massive asteroid impact. In 1991, Alan Hildebrand, William Boynton, Glen Penfield and Antonio Camargo, published a paper entitled “Chicxulub crater: a possible Cretaceous/Tertiary boundary impact crater on the Yucatan Peninsula, Mexico.” They had found the long-sought K/Pg impact crater.

The crater is more than 180 km (110 miles) in diameter and 20 km (10 miles) in depth, making the feature one of the largest confirmed impact structures on Earth. The  Chicxulub impact released an estimated energy equivalent of 100 teratonnes of TNT and produced high concentrations of dust, soot, and sulfate aerosols in the atmosphere. Model simulations suggest that the amount of sunlight that reached Earth’s surface was reduced by approximately 20%.This decrease of sunlight caused a drastic short-term global reduction in temperature. This phenomenon is called “impact winter”. Cold and darkness lasted for a period of months to years.  Photosynthesis stopped and the food chain collapsed. This period of reduced solar radiation may only have lasted several months to decades. Three-quarters of the plant and animal species on Earth disappeared. Marine ecosystems lost about half of their species while freshwater environments shows low extinction rates, about 10% to 22% of genera. Additionally, the vapour produced by the impact  could have led to global acid rain and a dramatic acidification of marine surface waters.

The Chicxulub asteroid impact was the final straw that pushed Earth past the tipping point.  The K-Pg extinction that followed the impact was one of the five great Phanerozoic  mass extinctions. Currently about 170 impact craters are known on Earth; about one third of those structures are not exposed on the surface and can only be studied by geophysics or drilling. Now, a new drilling platform in the the Gulf of Mexico, sponsored by the International Ocean Discovery Program (IODP) and the International Continental Scientific Drilling Program, will looking rock cores from the site of the impact. The main object is learn more about the scale of the impact, and the environmental catastrophe that ensued.

References:

Alvarez, L., W. Alvarez, F. Asaro, and H.V. Michel. 1980. Extraterrestrial cause for the Cretaceous-Tertiary extinction: Experimental results and theoretical interpretation. Science 208:1095–1108.

Alvarez, W. (1997) T. rex and the Crater of Doom. Princeton University Press, Princeton, NJ.

Hildebrand, A.R., G.T. Penfield, D.A. Kring, M. Pilkington, A. Camargo, S.B. Jacobsen, and W.V. Boynton. 1991. Chicxulub crater: A possible Cretaceous/Tertiary boundary impact crater on the Yucatán Peninsula, Mexico. Geology 19:867–71.

 

 

A Brief Introduction to The Hell Creek Formation.

Hell Creek e Fort Union contact, as seen at Mountain Goat Lake Butte, southwestern North Dakota (Adapted from Fastovsky and Bercovici, 2015)

Hell Creek- Fort Union contact, as seen at Mountain Goat Lake Butte, southwestern North Dakota (Adapted from Fastovsky and Bercovici, 2015)

The Hell Creek Formation (HCF), in the northern Great Plains of the United States, is the most studied source for understanding the changes in the terrestrial biota across the Cretaceous-Paleogene boundary, because preserves an extraordinary record comprised of fossil flora, vertebrates, invertebrates, microfossils, a range of trace fossils, and critical geochemical markers such as multiple iridium anomalies associated with the Chicxulub impact event. The HCF is a fine-grained, fluvially derived, siliciclastic unit, that occupies part of the western Williston Basin, and overlies the Fox Hills Formation (Clemens and Hartman, 2014).
The history of research focused on the Hell Creek Formation and its biota started in October 1901, when William T. Hornaday, director of the New York Zoological Society, travelled to northeastern Montana and discovered three fragments of the nasal horn of a Triceratops in the valley of Hell Creek. He showed the fossils to Henry Fairfield Osborn who decided to include the valley of Hell Creek on the list of areas to be prospected by Barnum Brown the following year.
braun

Barnum Brown working in a quarry in 1902.

In July 1902, B. Brown arrived to Hell Creek. His field crew included Dr. Richard Swann Lull, and Phillip Brooks. Brown recounted that after their arrival, he found the partial skeleton that would become the type specimen of Tyrannosaurus rex. In 1904, William H. Utterback, preparator and collector for the Carnegie Museum of Natural History, collected a fragment of a jaw of Tyrannosaurus and two skulls of Triceratops. In the summer of 1906, B. Brown returned to Montana, and a year later he published a complete manuscript about the valley of Hell Creek. The field expeditions of 1908 and 1909 were crowned by the discovery of another skeleton of T-rex. Between 1902 and 1910, Osborn, Brown, and Lull published the analysis of some of the fossil vertebrates discovered in the Hell Creek Formation, including Tyrannosaurus rex, Triceratops, and Ankylosaurus.
Micrograph of Wodehouseia spinata and a specimenBisonia niemi, from the upper part of the Hell Creek Formation (Adapted from Fastovsky and Bercovici, 2015).

Micrograph of Wodehouseia spinata and a specimen Bisonia niemi, from the upper part of the Hell Creek Formation (Adapted from Fastovsky and Bercovici, 2015).

Plants are represented by fossil leaves, seeds and cones. Fossil wood is also commonly found in the HCF as permineralized fragments. The Hell Creek macroflora is largely dominated by angiosperms including palms, associated with several ferns, conifers, and single species of cycads and Ginkgo. The study of pollen and spores has played a very important role in the identification of the K/Pg boundary in the HCF. Palynologists were the first scientists to recognize that a major, abrupt change occurred at the end of the Cretaceous. Unlike the Permian-Triassic and Triassic-Jurassic boundaries, the palynologically defined K/Pg boundary is based on the extinction of Cretaceous taxa rather than the appearance of Paleocene taxa. Intimately associated with the K/Pg boundary globally, is the so-called “fern spike”, occurring exclusively at localities where the iridium anomaly is present. (Fastovsky and Bercovici, 2015; Vajda & Bercovici, 2014.)

 

References:

Fastovsky, D. E., & Bercovici, A., The Hell Creek Formation and its contribution to the CretaceousePaleogene extinction: A short primer, Cretaceous Research (2015), http://dx.doi.org/10.1016/j.cretres.2015.07.007
Clemens, W. A., Jr., & Hartman, J. H. (2014). From Tyrannosaurus rex to asteroid impact: early studies (1901- 1980) of the Hell Creek Formation in its type area. In J. Hartman, K. R. Johnson, & D. J. Nichols (Eds.), Geological society of America special paper: 361. The Hell Creek Formation and the Cretaceous-tertiary boundary in the northern great plains (pp. 217-245).
Husson, D., Galbrun, B., Laskar, J., Hinnov, L. A., Thibault, N., Gardin, S., & Locklair, R. E. (2011). “Astronomical calibration of the Maastrichtian (late Cretaceous)”. Earth and Planetary Science Letters 305 (3): 328–340.doi:10.1016/j.epsl.2011.03.008
Johnson, K. R., Nichols, D. J., & Hartman, J. H. (2002). Hell Creek Formation: A 2001 synthesis. The Hell Creek Formation and the Cretaceous-Tertiary Boundary in the northern Great Plains: Geological Society of America Special Paper, 361, 503-510.

The Middle Permian mass extinction.

The Kapp Starostin Formation, Festningen section, Spitsbergen. The uppermost of the 3 yellow limestone beds records the Middle Permian mass extinction (Credit: Photographer: Dierk Blomeier. For David P.G. Bond and colleagues, GSA Bulletin, 2015.)

The Kapp Starostin Formation, Festningen section, Spitsbergen. The uppermost of the 3 yellow limestone beds records the Middle Permian mass extinction (Photo Credit: Dierk Blomeier. For David P.G. Bond and colleagues, GSA Bulletin, 2015.)

Extinction is the ultimate fate of all species. The fossil record indicates that more than 95% of all species that ever lived are now extinct. Individuals better adapted to environments are more likely to survive and when a species does fail, it is called a background extinction. Occasionally extinction events reach a global scale, with many species of all ecological types dying out in a near geological instant. These are mass extinctions. They were originally identified in the marine fossil record and have been interpreted as a result of catastrophic events or major environmental changes that occurred too rapidly for organisms to adapt.  Mass extinctions are probably due to a set of different possible causes like basaltic super-eruptions, impacts of asteroids, global climate changes, or continental drift.

George Cuvier, the great French anatomist and paleontologist, was the first to suggested that periodic “revolutions” or catastrophes had befallen the Earth and wiped out a number of species. But under the influence of Lyell’s uniformitarianism, Cuvier’s ideas were rejected as “poor science”. The modern study of mass extinction did not begin until the middle of the twentieth century. One of the most popular of that time was “Revolutions in the history of life” written by Norman Newell in 1967.

The fossil record shows that biodiversity in the world has been increasing dramatically for 200 million years and is likely to continue. The two mass extinctions in that period (at 201 million and 66 million years ago) slowed the trend only temporarily. Genera are the next taxonomic level up from species and are easier to detect in fossils. The Phanerozoic is the 540-million-year period in which animal life has proliferated. Chart created by and courtesy of University of Chicago paleontologists J. John Sepkoski, Jr. and David M. Raup.

Biodiversity in the fossil record.  (From Wikimedia Commons)

Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. In 1982, Jack Sepkoski and David M. Raup used a simple form of time series analysis at the rank of family to distinguish between background extinction levels and mass extinctions in marine faunas, and identified five major extinction events in Earth’s history: at the end of the Ordovician period, Late Devonian, End Permian, End Triassic and the End Cretaceous. These five events are know as the Big Five. The most recently identified mass extinction occurred during the Middle Permian, about  262 million years ago, and it was first recognised in the marine realm as a turnover among foraminifera, with fusulinaceans among the principal casualties. The crisis also affected numerous other shallow-marine taxa, including corals, bryozoans, brachiopods, bivalves and ammonoids. Until now, all detailed studies have focused on equatorial sections, especially those of South China. That extinction coincide with the Emeishan large igneous province. But, new data indicates that at the same time there was two severe extinctions amongst brachiopods in northern boreal latitudes in the Kapp Starostin Formation of Spitsbergen, an island roughly 890 km north of the Norwegian mainland.

Fusulinids from the Topeka Limestone  (Upper Carboniferous of Kansas, USA) From Wikimedia Commons

Fusulinids from the Topeka Limestone (Upper Carboniferous of Kansas, USA) From Wikimedia Commons

The Kapp Starostin Formation contains cool-water boreal faunas that include abundant siliceous sponges, brachiopods, and bryozoans. The widespread and near-total loss of carbonates across the Boreal Realm also suggests a role for acidification in the crisis.  This extinction predates the end-Permian mass extinction, because a subsequent recovery of brachiopods and especially bivalves is seen in the Late Permian. This post-extinction fauna disappears 10 m below the top of the Kapp Starostin Formation and thus fails to survive until the end of the Permian (Bond et al., 2015). This is a true mass extinction because the new data suggest that about 50 per cent of all marine species died during the event.

Oceanic oxygen depletion represents a potent cause of extinction in marine settings, and is often linked with volcanic activity, warming, and transgression. However,  the role of anoxia in the wider Capitanian extinction scenario remains enigmatic. Volcanically induced effects are multiple and include acidification.

brachi

Brachiopods from the Kapp Starostin Formation (Image adapted from Bond et al., 2015)

Acidification affects the biogeochemical dynamics of calcium carbonate, organic carbon, nitrogen, and phosphorus in the ocean and interferes with a range of processes, including growth, calcification, development, reproduction and behaviour in a wide range of marine organisms like foraminifera, planktonic coccolithophores, pteropods and other molluscs,  echinoderms, corals, and coralline algae. Ocean acidification in the geological record, is often inferred from a decrease in the accumulation and preservation of CaCO3 in marine sediments, potentially indicated by an increased degree of fragmentation of foraminiferal shells. But, recently, a variety of trace-element and isotopic tools have become available to infer past seawater carbonate chemistry.

Undoubtedly, the proximity of the End Permian extinction, makes difficult to determine if these events are separate or are part of a the same event.

References:

David P.G. Bond, Paul B. Wignall, Michael M. Joachimski, Yadong Sun, Ivan Savov, Stephen E. Grasby, Benoit Beauchamp and Dierk P.G. Blomeier, 2015, An abrupt extinction in the Middle Permian (Capitanian) of the Boreal Realm (Spitsbergen) and its link to anoxia and acidification, Geological Society of America Bulletin, doi: 10.1130/B31216.1

Wignall, P.B., Bond, D.P.G., Kuwahara, K., Kakuwa, Y., Newton, R.J., and Poulton, S.W., 2010, An 80 million year oceanic redox history from Permian to Jurassic pelagic sediments of the Mino-Tamba terrane, SW Japan, and the origin of four mass extinctions: Global and Planetary Change, v. 71, p. 109–123, doi:10.1016/j.gloplacha .2010.01.022.

Wignall, P.B., Bond, D.P.G., Newton, R.J., Haas, J., Hips, K., Wang, W., Jiang, H.-S., Lai, X.-L., Sun, Y.-D., Altiner, D., Védrine, S., and Zajzon, N., 2012, The Capitanian (Middle Permian) mass extinction in western Tethys: A fossil, facies and δ13C study from Hungary and Hydra Island (Greece): Palaios, v. 27, p. 78–89, doi:10.2110/palo.2011.p11-058r.

Ocean acidification and the end-Permian mass extinction

 

Permian Seafloor Photograph by University of Michigan Exhibit Museum of Natural History.

Permian Seafloor
Photograph by University of Michigan Exhibit Museum of Natural History.

About one third of the carbon dioxide released by anthropogenic activity is absorbed by the oceans. But the CO2 uptake lowers the pH and alters the chemical balance of the oceans. This phenomenon is called ocean acidification, and is occurring at a rate faster than at any time in the last 300 million years (Gillings, 2014; Hönisch et al. 2012). Acidification affects the biogeochemical dynamics of calcium carbonate, organic carbon, nitrogen, and phosphorus in the ocean and interferes with a range of processes, including growth, calcification, development, reproduction and behaviour in a wide range of marine organisms like planktonic coccolithophores, foraminifera, pteropods and other molluscs,  echinoderms, corals, and coralline algae. Rapid additions of carbon dioxide during extreme events in Earth history, including the end-Permian mass extinction (252 million years ago) and the Paleocene-Eocene Thermal Maximum (PETM, 56 million years ago) may have driven surface waters to undersaturation.

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

The end-Permian extinction is the most severe biotic crisis in the fossil record, with as much as 95% of the marine animal species and a similarly high proportion of terrestrial plants and animals going extinct . This great crisis occurred about 252 million years ago (Ma) during an episode of global warming.  The cause or causes of the Permian extinction remain a mystery but new data indicates that the extinction had a duration of 60,000 years and may be linked to massive volcanic eruptions from the Siberian Traps. The same study found evidence that 10,000 years before the die-off, the ocean experienced a pulse of light carbon that most likely led to a spike of carbon dioxide in the atmosphere. Volcanism and coal burning also contribute gases to the atmosphere, such as Cl, F, and CH3Cl from coal combustion, that suppress ozone formation.

Image that shows field work in the United Arab Emirates. Credit: D. Astratti

Image that shows field work in the United Arab Emirates. Credit: D. Astratti

Ocean acidification in the geological record, is often inferred from a decrease in the accumulation and preservation of CaCO3 in marine sediments, potentially indicated by an increased degree of fragmentation of foraminiferal shells. But, recently, a variety of trace-element and isotopic tools have become available to infer past seawater carbonate chemistry. The boron isotope composition of carbonate samples obtained from a shallow-marine platform section at Wadi Bih on the Musandam Peninsula, United Arab Emirates, allowed to reconstruct seawater pH values and atmospheric pCO2 concentrations and obtain for the very first time, direct evidence of ocean acidification in the Permo-Triassic boundary. The evidence indicates that the first phase of extinction was coincident with a slow injection of carbon into the atmosphere, and ocean pH remained stable. During the second extinction pulse, however, a rapid and large injection of carbon caused an abrupt acidification event that drove the preferential loss of heavily calcified marine biota (Clarkson et al, 2015).

The increasing evidence that the end-Permian mass extinction was precipitated by rapid release of CO2 into Earth’s atmosphere is a valuable reminder for an immediate action on global carbon emission reductions.

 

References:

Clarkson MO, Kasemann SA, Wood RA, Lenton TM, Daines SJ, Richoz S, Ohnemueller F, Meixner A, Poulton SW, Tipper ET. Ocean acidification and the Permo-Triassic mass extinction. Science, 2015 DOI: 10.1126/science.aaa0193

Feng, Q., Algeo, T.J., Evolution of oceanic redox conditions during the Permo-Triassic transition: Evidence from deepwater radiolarian facies, Earth-Sci. Rev. (2014), http://dx.doi.org/10.1016/j.earscirev.2013.12.003

Hönisch, A. Ridgwell, D. N. Schmidt, E. Thomas, S. J. Gibbs, A. Sluijs, R. Zeebe, L. Kump, R. C. Martindale, S. E. Greene, W. Kiessling, J. Ries, J. C. Zachos, D. L. Royer, S. Barker, T. M. Marchitto Jr., R. Moyer, C. Pelejero, P. Ziveri, G. L. Foster, B. Williams, The geological record of ocean acidification. Science 335, 1058–1063 (2012).

Kump, L.R., T.J. Bralower, and A. Ridgwell (2009)  Ocean acidification in deep time. Oceanography 22(4):94–107, http://dx.doi.org/10.5670/oceanog.2009.100

Seth D. Burgess, Samuel Bowring, and Shu-zhong Shen, High-precision timeline for Earth’s most severe extinction, PNAS 2014, doi:10.1073/pnas.1317692111

 

The Anthropocene defaunation process.

 

Richard Owen stands next to the largest of all moa, Dinornis maximus (now D. novaezealandiae). From Wikimedia Commons.

Richard Owen stands next to the largest of all moa, Dinornis maximus (now D. novaezealandiae). From Wikimedia Commons.

In 2000,  Paul Crutzen proposed use the term Anthropocene to designate the last two hundred years of human history and to mark the end of the current Holocene geological epoch. Although there is no agreement on when the Anthropocene started, it has been defined, primarily, by significant and measurable increases in anthropogenic greenhouse gas emissions from ice cores and other geologic features including synthetic organic compounds, radionuclides and ocean acidification.

Another marker for the Anthropocene is the current biodiversity crisis. The term defaunation was created to designate the declining of top predators and herbivores triggered by human activity, that results in a lack of agents that control the components of the ecosystems vegetation.

Global population declines in mammals and birds represented in numbers of individuals per 10,000 km2 for mammals and birds (From Dirzo et al., 2014)

Global population declines in mammals and birds (From Dirzo et al., 2014).

Since the industrial revolution, the wave of animal and plant extinctions that began with the late Quaternary has accelerated. Calculations suggest that the current rates of extinction are 100–1000 times above normal, or background levels. We are in the midst of  the so called “Sixth Mass Extinction”.

Although anthropogenic climate change is playing a growing role, the primary drivers of modern extinctions seem to be habitat loss, human predation, and introduced species (Briggs, 2011). The same drivers that contributed to ancient megafaunal and island extinctions.

SConsequences of defaunation (From Dirzo et al., 2014)

The consequences of defaunation (From Dirzo et al., 2014)

 

One of the most famous and well-documented extinctions come from Madagascar. Pygmy hippos, giant tortoises, and large lemurs went extinct due to human hunting or habitat disturbance.  A very interesting study by Burney et al. (2003) tracked the decline of coprophilous Sporormiella fungus spores in sediments due to reduced megafaunal densities after the human arrival on the island. Another well documented case is the Moa extinction in New Zealand. Recent radiocarbon dating and population modeling suggests that their disappearance occurred within 100 years of first human arrival. A large number of  land birds across Oceania suffered a similar fate beginning about 3500 years ago.

Some biologist predict that the sixth extinction  may result in a 50% loss of the plants and animals on our planet by AD 2100, which would cause not only the collapse of ecosystems but also the loss of food economies, and medicinal resources.

References:

Richard N. Holdaway, Morten E. Allentoft, Christopher Jacomb, Charlotte L. Oskam, Nancy R. Beavan, Michael Bunce. An extremely low-density human population exterminated New Zealand moa. Nature Communications, 2014; 5: 5436 DOI: 10.1038/ncomms6436

Rodolfo Dirzo et al., Defaunation in the Anthropocene, Science 345, 401 (2014); DOI: 10.1126/science.1251817

Braje, T.J., Erlandson, J.M., Human acceleration of animal and plant extinctions: A Late Pleistocene, Holocene, and Anthropocene continuum. Anthropocene (2013), http://dx.doi.org/10.1016/j.ancene.2013.08.003

 

 

The plant fossil record and the extinction events.

Odontopteris lingulata, seed fern from the Late  Late Pennsylvanian to Early Permian. (Image Credit: Taylor et al, 2009)

Odontopteris lingulata, seed fern from the Late Late Pennsylvanian to Early Permian. (Image Credit: Taylor et al, 2009)

Mass extinctions has shaped the global diversity of our planet several times during the geological ages. They were originally identified in the marine fossil record and have been interpreted as a result of catastrophic events or major environmental changes that occurred too rapidly for organisms to adapt.

In 1982, Jack Sepkoski and David M. Raup used a simple form of time series analysis at the rank of family to distinguish between background extinction levels and mass extinctions in marine faunas, and identified five major extinction events in Earth’s history: at the end of the Ordovician period, Frasnian (Late Devonian), Permian, Triassic and Cretaceous. But the plant fossil record reveals a different pattern of major taxonomic extinctions compared with marine organisms. The first of them took place at the Carboniferous-Permian transition, which is interpreted as result of the collapse of the tropical wetlands in Euramerica. The second mass extinction corresponds to the end-Permian event.

Glossopteris sp., seed ferns, Permian - Triassic - Houston Museum of Natural Science (From Wikimedia Commons)

Glossopteris sp., seed ferns, Permian – Triassic – Houston Museum of Natural Science (From Wikimedia Commons)

At the late Carboniferous the characteristic wetland families disappeared (e.g. Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Alethopteridaceae, Cyclopteridaceae, Neurodontopteridaceae). The downfall of rainforests probably reflects the complexity of the environmental changes that were taking place during the late Moscovian-early Sakmarian time interval (DiMichele et al., 2006, Sahney et al., 2010). This collapse probably drove the rapid diversification of Carboniferous tetrapods (amphibians and reptiles) in Euramerica (Sahney et al., 2010).

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval.

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

At the end-Triassic event,  the vegetation turnover in the Southern Hemisphere  consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

The end-Cretaceous biotic crisis had a significant effect on marine and terrestrial faunas, and caused localized loss of species diversity in vegetation. Patagonia shows a reduction in diversity and relative abundance in almost all plant groups from the latest Maastrichtian to the Danian, although only a few true extinctions occurred (Barreda et al, 2013). The nature of vegetational change in the south polar region suggests that terrestrial ecosystems were already responding to relatively rapid climate change prior to the K–Pg catastrophe.

Two examples of grains pollen from the Lopez de Bertodano Formation: Podocarpidites sp. (left) and Nothofagidites asperus (right)

Two examples of grains pollen: Podocarpidites sp. (left) and Nothofagidites asperus (right)

A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988).

References:

Cascales-Miñana, B., and C. J. Cleal, 2014, The plant fossil record reflects just two great extinction events. Terra Nova. vol. 26, no. 3, pp. 195–200. DOI: 10.1111/ter.12086

Bambach, R.K., Knoll, A.H. and Wang, S.C., 2004. Origination, extinction, and mass depletions of marine diversity. Paleobiology, 30, 522–542.

Mayhew, Peter J.; Gareth B. Jenkins, Timothy G. Benton (January 7, 2008). “A long-term association between global temperature and biodiversity, origination and extinction in the fossil record”. Proceedings of the Royal Society B: Biological Sciences 275 (1630): 47–53.

Sahney, S., Benton, M.J. & Falcon-Lang, H.J. (2010). “Rainforest collapse triggered Pennsylvanian tetrapod diversification in Euramerica” (PDF). Geology 38 (12): 1079–1082. doi:10.1130/G31182.1.

 

The palynological record and the extinction events.

The main palynological provinces at the end of the Cretaceous (From Vajda and Bercovici, 2014)

The main palynological provinces at the end of the Cretaceous (From Vajda and Bercovici, 2014)

Pollen and other palynomorphs proved to be an extraordinary tool to palaeoenvironmental reconstruction. In 1921, Gunnar Erdtman, a Swedish botanist, was the first to suggest this application for fossil pollen study. Like spores, pollen grains reflects the ecology of their parent plants and their habitats and provide a continuous record of their evolutionary history. Pollen analysis involves the quantitative examination of spores and pollen at successive horizons through a core, specially in lake, marsh or delta sediments. The morphology of pollen grains is diverse. Gymnosperm pollen often is saccate (grains with two or three air sacs attached to the central body), while Angiosperm pollen shows more variation and covers a multitude of combinations of features: they could be  in groups of four (tetrads),  in pairs (dyads),  or single (monads). The individual grains can be inaperturate, or have one or more pores, or slit-like apertures or colpi (monocolpate, tricolpate).

Since the 1980s, many fossil pollen data sets were developed specifically to reconstruct past climate change.

Aquilapollenites quadricretaeus and Nothofagidites kaitangata

Aquilapollenites quadricretaeus and Nothofagidites kaitangata

 

The palynological record across the Cretaceous–Paleogene (K–Pg) boundary  is a unique global  marker that can be use as template to asses the causal mechanism behind other major extinction events in Earths history. Four major palynological provinces have been recognized based on distinctive angiosperm pollen and fern spores of restricted geographic and stratigraphic distribution. The Aquilapollenites Province had a northern circumpolar distribution that extended from Siberia, northern China, Japon and the western North America. The Normapolles Province occupied eastern North America,  Europe and western Asia. The Palmae Province occupied equatorial regions in the Late Cretacic and included SouthAmerica, Africa and India. Finally, the Notofagidites Province that extended across southern South America, Antartica, New Zeland and Australia.

During the Late Cretaceous the global climate change has been associated with episodes of outgassing from major volcanic events, orbital cyclicity and tectonism before ending with the cataclysm caused by a large bolide impact at Chicxulub, on the Yucatán Peninsula, Mexico. Although, during the middle Maastrichtian, there was a short-lived warming event related to an increase in atmospheric carbon dioxide from the first Deccan eruption phase, the global climate cooled during the latest Maastrichtian and across the K–Pg boundary (Wang et al., 2014; Brusatte et al., 2014). The variations in floral composition reflect these paleoclimatic changes.

Fern spike adapted from Bercovicci

Fern spike adapted from Bercovicci

Mainly angiosperms, disappear at the boundary, as evidenced the palynofloral records of North America and New Zealand. Patagonia shows a reduction in diversity and relative abundance in almost all plant groups from the latest Maastrichtian to the Danian, although only a few true extinctions occurred (Barreda et al, 2013).  The nature of vegetational change in the south polar region suggests that terrestrial ecosystems were already responding to relatively rapid climate change prior to the K–Pg catastrophe.

The earliest Paleocene vegetation shows an anomalous concentration of fern spores just above the level of palynological extinction. R. H. Tschudy, in 1984,  was the first to recognize this very distinctive pattern when he analyzed samples from the K/PG boundary and observed that just after the extinction event, the palynological assemblages were dominated by a high abundance of fern spores.

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. At the end-Triassic event,  the vegetation turnover in the Southern Hemisphere  consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

Despite their difference, these three extinction events are consequences of dramatic environmental upheavals that generated comparable extinction patterns, and similar phases of vegetation recovery but at different temporal scales. First, all these events share a similar pattern of a short-term bloom of opportunistic “crisis” taxa proliferating in the devastated environment. Second, there’s a pulse in pioneer communities (spore spike). Third , a recovery in diversity including the evolution of new taxa. Furthermore, the longer the extreme environmental conditions last the greater is the extinction rate and the extinction patterns between autotrophs and heterotrophs, and between terrestrial and marine faunas become more similar (Vajda and Bercovici, 2014).

 

References:

Vivi Vajda & Antoine Bercovici (2014); The global vegetation pattern across the Cretaceous–Paleogene mass extinction interval: A template for other extinction events; Global and Planetary Change (advance online publication) Open Access DOI: 10.1016/j.gloplacha.2014.07.014, http://www.sciencedirect.com/science/article/pii/S0921818114001477

Vajda, V., Raine, J.I., 2003. Pollen and spores in marine Cretaceous/Tertiary boundary sediments at mid–Waipara River, North Canterbury, New Zealand. New Zealand Journal of Geology and Geophysics 46, 255–273

Wang, Y., Huang, C., Sun, B., Quan, C., Wu, J., Lin, Z., 2014. Paleo-CO2 variation trends and the Cretaceous greenhouse climate. Earth-Science Reviews 129, 136–147.

Vanessa C. Bowman, Jane E. Francis, Rosemary A. Askinb, James B. Riding, Graeme T. Swindles, Latest Cretaceous–earliest Paleogene vegetation and climate change at the high southern latitudes: palynological evidence fromSeymour Island, Antarctic Peninsula, Palaeogeography, Palaeoclimatology, Palaeoecology, 408. 26-47. DOI 10.1016/j.palaeo.2014.04.018

Barreda VD, Cúneo NR, Wilf P, Currano ED, Scasso RA, et al. (2012) Cretaceous/Paleogene Floral Turnover in Patagonia: Drop in Diversity, Low Extinction, and a Classopollis Spike. PLoS ONE 7(12): e52455. doi: 10.1371/journal.pone.0052455

Brusatte, S. L., Butler, R. J., Barrett, P. M., Carrano, M. T., Evans, D. C., Lloyd, G. T., Mannion, P. D., Norell, M. A., Peppe, D. J., Upchurch, P., and Williamson, T. E. In press. The extinction of the dinosaurs.Biological Reviews

The Winter of Our Discontent: short-term cooling following the Chicxulub impact.

ooimpact.2

The K-T impact by Don Davis.

The Cretaceous–Paleogene extinction that followed the  Chicxulub impact was one of the five great Phanerozoic  mass extinctions. The impact released an estimated energy equivalent of 100 teratonnes of TNT and produced high concentrations of dust, soot, and sulfate aerosols in the atmosphere. Model simulations suggest that the amount of sunlight that reached Earth’s surface was reduced by approximately 20%.This decrease of sunlight caused a drastic short-term global reduction in temperature. This phenomenon is called “impact winter”.

Cold and darkness lasted for a period of months to years.  Photosynthesis stopped and the food chain collapsed. This period of reduced solar radiation may only have lasted several months to decades. Three-quarters of the plant and animal species on Earth disappeared. Marine ecosystems lost about half of their species while freshwater environments shows low extinction rates, about 10% to 22% of genera.

Three factors can be associated with the impact winter in marine and fresh water enviroments. First, starvation caused by the stop of photosynthesis. Second, the loss of dissolved oxygen. Third, the low temperatures. The flux of organic detritus to the sea floor also

A paleogeographic map of the Gulf of Mexico at the end of the Cretaceous (From Vellekoop, 2014)

Three factors can be associated with the impact winter in marine and fresh water environments. First, starvation caused by the stop of photosynthesis. Second, the loss of dissolved oxygen. Third, the low temperatures. Because the late Cretaceous climate was warm, a major challenge for aquatic organisms, especially in inland waters, may have been the persistence of low temperatures. Additionally, the vapour produced by the impact  could have led to global acid rain and a dramatic acidification of marine surface waters.

Fossil evidence for this impact winter was recovered in the Brazos River region of Texas.  The biostratigraphy of the section presents the Ir anomaly, and impact-related tsunami beds. The age of the outcrops was updated using  planktonic foraminifera and  dinocysts.

The “impact winter”  model is supported by a migration of cool, boreal dinoflagellate species into the subtropic Tethyan realm directly across the K–Pg boundary interval and the ingression of boreal benthic foraminifera into the deeper parts of the Tethys Ocean, interpreted to reflect millennial timescale changes in ocean circulation following the impact (Vellekoop, 2014).

References:

Johan Vellekoop, Appy Sluijs, Jan Smit, Stefan Schouten, Johan W. H. Weijers, Jaap S. Sinningh Damsté, and Henk Brinkhuis, Rapid short-term cooling following the Chicxulub impact at the Cretaceous–Paleogene boundary, PNAS (2014) doi: 10.1073/pnas.1319253111

Douglas S. Robertson, William M. Lewis, Peter M. Sheehan and Owen B. Toon, K-Pg extinction patterns in marine and freshwater environments: The impact winter model, Journal of Geophysical Research: Biogeosciences, JUL 2013, DOI: 10.1002/jgrg.20086.