Mutagenesis in land plants during the end-Triassic mass extinction

 

A basaltic lava flow section from the Middle Atlas, Morocco. From Wikimedia Commons.

During the last 540 million years five mass extinction events shaped the history of the Earth. The End-Triassic Extinction at 201.51 million years (Ma) is probably the least understood of these events. 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.

The mass extinction event was likely caused by the eruption of the Central Atlantic Magmatic Province (CAMP), a large igneous province emplaced during the initial rifting of Pangea. Data indicates that magmatic activity started c. 100,000 years before the endTriassic event and continued in pulses for 700,000 years. The CO2 emissions caused global warming. The SO2 emissions on mixing with water vapour in the atmosphere, caused acid rain, which in turn killed land plants and caused soil erosion.

A normal fern spore compared with mutated ones from the end-Triassic mass extinction event. Image credit: S LINDSTRÖM, GEUS

Volcanoes are also a primary source of mercury (Hg) in the global atmosphere. Mercury can cause morphologically visible abnormalities in plants and their reproductive cells (spores and pollen). A new study led by Sofie Lindström of the Geological Survey of Denmark and Greenland analized various types of abnormalities in the reproductive cells of ferns, with focus in two morphogroups: LTT-spores (laevigate, trilete fern spores with thick exine), and LCT-spores (laevigate, circular, trilete spores). The LTT-spores were produced primarily by the fern families Dipteridaceae, Dicksoniaceae, and Matoniaceae, while LCT spores were primarily produced by ferns belonging to Osmundaceae and Marattiales.

The elevated concentrations of mercury (Hg) in sedimentary rocks in North America, Greenland, England, Austria, Morocco, and Peru are linked to CAMP eruptions. This pulse of mercury also correlate with high occurrences of abnormal fern spores, indicating severe environmental stress and genetic disturbance in the parent plants. Three negative organic C-isotope excursions (CIEs) have being recognized at the end-Triassic: the Marshi, the Spelae, and the top-Tilmanni CIEs. Malformations in LTT-spores first occur sporadically in the lower pre-Marshi interval. LCT-spores are present but are generally rare in this interval. During the Spelae CIE, the occurrences of moderate to severe malformations increased and aberrant forms can encompass as much as 56% of the counted LTT-spores. This interval is associated with marked global warming, recorded by stomatal proxy data.

 

 

References:

Sofie Lindström et al. Volcanic mercury and mutagenesis in land plants during the end-Triassic mass extinction, Science Advances (2019). DOI: 10.1126/sciadv.aaw4018}

Grasby, S. E., Them, T. R., Chen, Z., Yin, R., & Ardakani, O. H. (2019). Mercury as a proxy for volcanic emissions in the geologic record. Earth-Science Reviews, 102880. doi:10.1016/j.earscirev.2019.102880

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.

 

The Triassic Paleoclimate.

Early to Middle Triassic (240Ma) From Wikimedia Commons.

Early to Middle Triassic (240Ma) From Wikimedia Commons.

There are three basic states for Earth climate: Icehouse, Greenhouse (subdivided into Cool and Warm states), and Hothouse (Kidder & Worsley, 2010). The “Hothouse” condition is relatively short-lived and is consequence from the release of anomalously large inputs of CO2 into the atmosphere during the formation of Large Igneous Provinces (LIPs), when atmospheric CO2 concentrations may rise above 16 times (4,800 ppmv), while the “Icehouse” is characterized by polar ice, with alternating glacial–interglacial episodes in response to orbital forcing. The ‘Cool Greenhouse” displays  some polar ice and alpine glaciers,  with global average temperatures between 21° and 24°C. Finally, the ‘Warm Greenhouse’ lacked of any polar ice, and global average temperatures might have ranged from 24° to 30°C.

The rifting of Pangea during the Mesozoic modified the paleoposition and shoreline configuration of the land masses and generated huge epicontinental seas. This altered significantly the oceanic circulation and caused profound consequences for paleoclimates and for the evolution of life.

Siberian flood-basalt flows in Putorana, Taymyr Peninsula.(From Earth science: Lethal volcanism, Paul B. Wignall, 2011,  Nature 477, 285–286 )

Siberian flood-basalt flows in Putorana, Taymyr Peninsula.(From Earth science: Lethal volcanism, Paul B. Wignall, 2011, Nature 477, 285–286 )

During the Late Permian, massive volcanic eruptions in Siberia covered more than 2 millions of km 2 with lava flows, releasing more carbon in the atmosphere and high amounts of fluorine and chlorine increasing the climatic instability, which means that the Mesozoic began under extreme hothouse conditions.

The Early Triassic transition is marked by a moderate oxygen depletion and by mass extinction of glossopterids, gigantopterids, tree lycopsids and cordaites, as major contributors to coal deposits in the southern hemisphere. That was accompanied  by unusually anoxic swamp soils (Retallack, 2013). The rifting of Gondwana began during the Early Triassic with the opening of the Indian Ocean and the separation of India and Australia, that modified shoreline configuration and enhanced platform areas inducing intense marine biodiversification. It was suggested that during that time there was a moderate oxygen depletion that caused the low body size of the amphibian and reptilian life-forms found in those rocks.

Fossil of Lystrosaurus, one of the few survivors of the Late Permian shows a variety of adaptations to low oxygen atmosphere. It was by far the most common terrestrial vertebrate of the Early Triassic (Staatliches Museum für Naturkunde Stuttgart; from Wikimedia Commons).

Fossil of Lystrosaurus, one of the few survivors of the Late Permian shows a variety of adaptations to low oxygen atmosphere. It was by far the most common terrestrial vertebrate of the Early Triassic (Staatliches Museum für Naturkunde Stuttgart; from Wikimedia Commons).

By the Mid Triassic, global temperature was still high – between 20°C and 30°C – and the atmospheric CO2 began to increase. There are reported episodes of humid climate registered by fossil vertebrates from the Molteno Formation in South Africa, and from Los Rastros Formation in central western Argentina.

The Late Triassic is marked by a return to the hothouse condition of the Early Triassic, with two greenhouse crisis that may also have played a role in mass extinctions and long-term evolutionary trends (Retallack, 2013). The paleoclimate was a very arid with intense evaporation rate. Although there was at least one time of significant increase in rainfall known as the “Carnian Pluvial Event”, possibly related to the rifting of Pangea. Massive volcanic eruptions from a large region known as the Central Atlantic Magmatic Province (CAMP) release huge amounts of lava and gas, including carbon dioxide, sulfur and methane into the atmosphere which led to global warming and acidification of the oceans.

Light-microscope photographs of Classopollis pollen from the Late Triassic (Image adapted from Kürschner et al., 2013).

Light-microscope photographs of Classopollis pollen from the Late Triassic (Image adapted from Kürschner et al., 2013).

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

Most of scientists agree that the extinctions were caused by massive volcanic activity associated with the break-up of the super-continent Pangaea. 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).

References:

Holz, M., Mesozoic paleogeography and paleoclimates – a discussion of the diverse greenhouse and hothouse conditions of an alien world, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.01.001

Sellwood, B.W. & Valdes, P.J. 2006. Mesozoic climates: General circulation models and the rock Record. Sedimentary Geology 190:269–287.

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

Retallack, G.J. 2009. Greenhouse crises of the past 300 million years. Geological Society of America Bulletin, 121:1441–1455.