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

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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.

 

Palynology of the Ischigualasto Formation.

Ischigualasto-perfil-gusano-montañas

Image from Ischigualasto Park (http://www.ischigualasto.gob.ar/)

Ischigualasto is an arid, sculpted valley, in northwest Argentina (San Juan Province), limiting to the north with the Talampaya National Park, in La Rioja Province. Both areas belong to the same geological formation: the Ischigualasto-Villa Unión Basin which is centered on a rift zone that accumulated thick terrestrial deposits during the Triassic. This basin preserves a complete and continuous fossiliferous succession of continental Triassic rocks.

The Ischigualasto Formation is known worldwide for its tetrapod assemblage, which included the oldest known record of dinosaurs. Adolf Stelzner in 1889 published the first data on the geology of Ischigualasto, but it was not until 1911, that Bondenbender briefly refers to the fossils of the site. Several thin volcanic ash horizons, indicates that the deposition of the Ischigualasto Formation began at the Carnian Stage (approximately 228 mya), and consists of four lithostratigraphic members which in ascending order include the La Peña Member, the Cancha de Bochas Member, the Valle de la Luna Member, and the Quebrada de la Sal Member.

1–3. Retusotriletes herbstii sp. nov; 4–5. Rogalskaisporites cicatricosus; 6. Rugulatisporites

1–3. Retusotriletes herbstii sp. nov; 4–5. Rogalskaisporites cicatricosus; 6. Rugulatisporites

During the Late Triassic two distinct microfloras have been recognised in the southern hemisphere: the Ipswich microflora and the Onslow microflora. The Ipswich province, characterized by the abundance of bisaccate pollen, monosulcate pollen and trilete spores, evolved in southern and eastern Australia, Transantarctic Mountains region, South Africa and Argentina. The Onslow province is a mixture of Gondwanan and European taxa recognized in of north-western Australia, Madagascar, East Africa, Indian, and East Antarctic (Cesari and Colombi; 2013).

The recognition of Carnian European species in the Valle de la Luna Member of the Ishchigualasto Formation expands the distribution of the Onslow-type palynofloras. This assemblage was recovered from the site known as “El Hongo” in the Provincial Park, and contain the diagnostic “Onslow” species: Samaropollenites speciosus, Enzonalasporites vigens, Patinasporites densus, Vallatisporites ignacii, Ovalipollis pseudoalatus and Cycadopites stonei. This assemblage indicates that the Valle de la Luna Member was likely deposited under more humid conditions. It also implies the existence of a latitudinal floral belt from Timor (through the Circum-Mediterranean area) to western Argentina.

 

References:

Cesari, Silvia N., Colombi, Carina, Palynology of the late Triassic ischigualasto formation, Argentina: Paleoecological and paleogeographic implications, Palaeogeography, Palaeoclimatology, Palaeoecology (2016), doi: 10.1016/j.palaeo.2016.02.023

Césari, S. N., Colombi, C. E., 2013. A new Late Triassic phytogeographical scenario in westernmost Gondwana. Nature communications, 4.

Spalletti, L. A. Artabe, A. E. & Morel, E. M. Geological factors and evolution of southwestern Gondwana Triassic plants. Gondwana Res. 6, 119–134 (2003).

 

Darwin and the flowering plant evolution in South America.

pollen

Retimonocolpites sp. (Adapted from Llorens and Loinaze, 2015)

Charles Darwin’s fascination and frustration with the evolutionary events associated with the origin and early radiation of flowering plants are legendary. In a letter to Oswald Heer, a famous Swiss botanist, and paleontologist, Darwin wrote: “the sudden appearance of so many Dicotyledons in the Upper Chalk appears to me a most perplexing phenomenon to all who believe in any form of evolution, especially to those who believe in extremely gradual evolution, to which view I know that you are strongly opposed”. Heer discussed about the early angiosperm fossil record with Darwin, in a letter dated 1 March 1875: “if we say that the Dicotyledons begin with the Upper Cretaceous, we must still concede that this section of the vegetable kingdom, which forms the bulk of modern vegetation, appears relatively late and that, in geological terms, it underwent a substantial transformation within a brief period of time.” 

Darwin’s defense of a gradualist perspective led him to suggest that prior to the Cretaceous record of flowering plants, angiosperms had slowly evolved and diversified on a remote landmass. On 22 July 1879, in a letter to Joseph Dalton Hooker, Darwin refers to the early evolution of flowering plants as an “abominable mystery”. Nearing the end of his life, he wrote to Hooker another letter about a lost fossil record in the earliest phases of angiosperm diversification:  “Nothing is more extraordinary in the history of the Vegetable Kingdom, as it seems to me, than the apparently very sudden or abrupt development of the higher plants. I have sometimes speculated whether there did not exist somewhere during long ages an extremely isolated continent, perhaps near the South Pole.”

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

While the oldest records of the different groups of angiosperms are still in discussion, the outcrops of the Baquero Group, located in Argentinean Patagonia, contain one of the richest and most diverse Early Cretaceous floras in the Southern Hemisphere. The unit comprises three formations: Anfiteatro de Tico, Bajo Tigre and Punta del Barco. The first reports of angiosperm remains for the Anfiteatro de Tico Formation were made in 1967. The dominant types are Clavatipollenites, and Retimonocolpites.

Pollen grains  could enter into the fossil record by falling directly into swamps or lakes, or being washed into them or into the rivers and seas. The ones which are not buried in reducing sediments will tend to become oxidized and be destroyed. They reflects the ecology of their parent plants and their habitats and provide a continuous record of their evolutionary history. Gymnosperms 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).

pollen

Clavatipollenites sp. SEM (Adapted from Archangelsky 2013)

Clavatipollenites pollen grains are interpreted as related to the modern family Chloranthaceae. The genus was established by Couper for dispersed monosulcate pollen grains recovered from the Early Cretaceous of Britain. Currently, the genus has a very broad definition. The genus Retimonocolpites include elongated to subcircular semitectate, columellate and microreticulate pollen grains with well defined monocolpate aperture (Llorens and Loinaze, 2015). The new species Jusinghipollisticoensis sp. nov. represents one of the oldest records of trichotomosulcate, and extends the geographical distribution of Early Cretaceous trichotomosulcate pollen grains to southern South America.

The data also indicates strong similarities between the Baquero Group assemblages and other coeval units from Argentina, Australia and United States.

References:

M. Llorens, V.S. Perez Loinaze, Late Aptian angiosperm pollen grains from Patagonia: Earliest steps in flowering plant evolution at middle latitudes in southern South America, Cretaceous Research 57 (2016) 66-78

Archangelsky, S.,et al. (2009). Early angiosperm diversification: evidence from southern South America. Cretaceous Research, 30, 1073-1082.

Doyle, J. A., & Endress, P. K. (2014). Integrating Early Cretaceous fossils into the phylogeny of living Angiosperms: ANITA. Lines and relatives of Chloranthaceae. International Journal of Plant Sciences, 175, 555-560.

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.

Ecosystem instability in the Late Triassic and the early evolution of dinosaurs.

The Late Triassic Petrified Forest Member of the Chinle Formation (Photo from AASG)

The Late Triassic Petrified Forest Member of the Chinle Formation (Photo from AASG)

Dinosaurs likely originated in the Middle Triassic and the first unequivocal dinosaur fossils are known from the late Carnian, but much about the geological and temporal backdrop of early dinosaur history remains poorly understood. A key question is why early dinosaurs were rare and species-poor at low paleolatitudes throughout the Late Triassic Period, for at least 30 million years after their origin.

The oldest well-dated identified dinosaurs are from the late Carnian (approx. 230 Ma) of the lower Ischigualasto Formation in northwestern Argentina. Similarly, the Santa Maria and Caturrita formations in southern Brazil preserve basal dinosauromorphs, basal saurischians, and early sauropodomorphs. In North America, the oldest dated occurrences of vertebrate assemblages with dinosaurs are from the Chinle Formation, but are less abundant and species rich compared to those from South America. The fact that those assemblages were at moderately high paleolatitudes during the Late Triassic, and the North American assemblages were near the paleoequator supports the hypotheses for a diachronous rise of dinosaurs across paleolatitudes (Irmis et al., 2011).

A reconstructed scene from the Late Triassic (Norian) of central Pangea. (Credit: image from Brusatte, S. L. 2008)

A reconstructed scene from the Late Triassic (Norian) of central Pangea. (Image from Brusatte, S. L. 2008, Dinosaurs, Quercus Publishing, London).

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. Now, a multiproxy study  suggests  that fluctuating aridity in tropical and subtropical Pangea could explain why Triassic dinosaur faunas at low latitudes are restricted to small, slower growing carnivorous forms, whereas large-bodied herbivores, including sauropodomorph dinosaurs, are absent at low paleolatitudes during the Late Triassic “hothouse.” The palynomorphs recovered from sediments of the Chinle Formation indicate a major change from a seed fern-dominated (Alisporites) assemblage with accessory gymnosperms to one dominated by conifers and seed ferns in the lower portion of the Petrified Forest Member. In addition, the extensive charcoal record in the Petrified Forest Member provides evidence of paleo-environmental variability and aridity. 

 

References:

Jessica H. Whiteside, Sofie Lindström, Randall B. Irmis, Ian J. Glasspool, Morgan F. Schaller, Maria Dunlavey, Sterling J. Nesbitt, Nathan D. Smith, and Alan H. Turner. 2015. Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years. PNAS: doi:10.1073/pnas.1505252112

Brusatte, S. L., Nesbitt, S. J., Irmis, R. B., Butler, R. J., Benton, M. J., and Norell, M. A. 2010. The origin and early radiation of dinosaurs. Earth-Science Reviews, 101, 68-100

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

Nesbitt,  S. J., Irmis,  R. B, Parker,  W. G. (2007) A critical re-evaluation of the Late Triassic di-nosaur taxa of North America. J Syst Palaeontology 5(2):209243

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

 

Response of marine ecosystems to the PETM.

Biotic change among foraminifera during and after the PETM. (From Speijer et al., 2012)

Biotic change during and after the PETM. (From Speijer et al., 2012)

The Paleocene-Eocene Thermal Maximum (PETM; 55.8 million years ago), was a short-lived (~ 200,000 years) global warming event attributed to a rapid rise in the concentration of greenhouse gases in the atmosphere. It was suggested that this warming was initiated by the melting of methane hydrates on the seafloor and permafrost at high latitudes. During the PETM, around 5 billion tons of CO2 was released into the atmosphere per year, and temperatures increased by 5 – 9°C. This event was accompanied by other large-scale changes in the climate system, for example, the patterns of atmospheric circulation, vapor transport, precipitation, intermediate and deep-sea circulation and a rise in global sea level.

Dinoflagellate cysts: Adnatosphaeridium robustum and Apectodinium augustum (From Sluijs and Brinkhuis, 2009)

Dinoflagellate cysts: Adnatosphaeridium robustum and Apectodinium augustum (From Sluijs and Brinkhuis, 2009)

The rapid warming at the beginning of the Eocene has been inferred from the widespread distribution of dinoflagellate cysts. One taxon in particularly, Apectodinium, spans the entire carbon isotope excursion (CIE) of the PETM. The distribution of Apectodinium is linked to high temperatures and increased food availability.

The most disruptive impact during the PETM was likely the exceptional ocean acidification and the rise of the calcite dissolution depth, affecting marine organisms with calcareous shells (Zachos et al., 2005). When CO2 dissolves in seawater, it produce carbonic acid. The carbonic acid dissociates in the water releasing hydrogen ions and bicarbonate. The formation of bicarbonate then removes carbonate ions from the water, making them less available for use by organisms.

Nannofossil abundance changes during the PETM. (From Kump, 2009.)

Nannofossil abundance changes during the PETM. (From Kump, 2009.)

The PETM onset is also marked by the largest deep-sea mass extinction among calcareous benthic foraminifera (including calcareous agglutinated taxa) in the last 93 million years. Similarly, planktonic foraminifera communities at low and high latitudes show reductions in diversity, while larger foraminifera are the most common constituents of late Paleocene–early Eocene carbonate platforms.

The response of most marine invertebrates (mollusks, echinoderms, brachiopods) to paleoclimatic change during the PETM is poorly documented.

Coccolithus bownii and Toweius pertusus

Coccolithus bownii and Toweius pertusus (Adapted from Bown and Pearson, 2009)

The PETM is also associated with dramatic changes among the calcareous plankton,characterized by the appearance of transient nanoplankton taxa of heavily calcified forms of Rhomboaster spp., Discoaster araneus, and D. anartios as well as Coccolithus bownii, a more delicate form.

The combination of global warming and the release of large amounts of carbon to the ocean-atmosphere system during the PETM has encouraged analogies to be drawn with modern anthropogenic climate change. The current rate of the anthropogenic carbon input  is probably greater than during the PETM, causing a more severe decline in ocean pH and saturation state. Also the biotic consequences of the PETM were fairly minor, while the current rate of species extinction is already 100–1000 times higher than would be considered natural. This underlines the urgency for immediate action on global carbon emission reductions.

Comparison of the effects of anthropogenic emissions (total of 5000 Pg C over 500 years) and PETM carbon release (3000 Pg C over 6 kyr) on the surface ocean saturation state of calcite. From Zeebe, 2013

Comparison of the effects of anthropogenic emissions (total of 5000 Pg C over 500 years) and PETM carbon release (3000 Pg C over 6 kyr) on the surface ocean saturation state of calcite. From Zeebe, 2013

 

References:

Maria Rose Petrizzo, The onset of the Paleocene–Eocene Thermal Maximum (PETM) at Sites 1209 and 1210 (Shatsky Rise, Pacific Ocean) as recorded by planktonic foraminifera, Marine Micropaleontology, Volume 63, Issues 3–4, 13 June 2007, Pages 187-200

Zeebe RE and Zachos JC. 2013 Long-term legacy ofmassive carbon input to the Earth system: Anthropocene versus Eocene. Phil Trans R Soc A 371: 20120006. http://dx.doi.org/10.1098/rsta.2012.0006.

Wright JD, Schaller MF (2013) Evidence for a rapid release of carbon at the Paleocene-Eocene thermal maximum. Proc Natl Acad Sci USA 110(40):15908–15913.

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

Raffi, I. and de Bernardi, B.: Response of calcareous nannofossils to the Paleocene-Eocene Thermal Maximum: observations on composition, preservation and calcification in sediments from ODP Site 1263 (Walvis Ridge – SW Atlantic), Mar. Micropaleontol., 69, 119–138, 2008.

Robert P. SPEIJER, Christian SCHEIBNER, Peter STASSEN & Abdel-Mohsen M. MORSI; Response of marine ecosystems to deep-time global warming: a synthesis of biotic patterns across the Paleocene-Eocene thermal maximum (PETM), Austrian Journal of Earth Sciences. Vienna. 2012. Volume 105/1.

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

Palynological reconstruction of the Antarctic Cretaceous-Paleocene climate.

Artist’s impression of the eastern flank of the Antarctic Peninsula during theMaastrichtian (Artist: James McKay, University of Leeds.)

Artist’s impression of the eastern flank of the Antarctic Peninsula during the Maastrichtian (From Bowman et al, 2014, Artist: James McKay, University of Leeds.)

Past fluctuations in global temperatures are crucial to understand Earth’s climatic evolution. 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.
The Antarctic Peninsula is an area of specific interest to modern and past climatic studies, as it seems particularly sensitive to change (Kemp et al., 2014). Most of the studies are focused on Seymour Island which has one of the most expanded Cretaceous–Paleogene successions known. The K-Pg boundary occurs in the uppermost part of the López de Bertodano Formation, where it is marked by a minor iridium anomaly.

The terrestrial palynomorph record at the López de Bertodano Formation was divided into six phases. The first one contains an assemblage dominated by Nothofagidites spp. and Podocarpidites spp., with aquatic fern spores (Azolla spp., Grapnelispora sp.) and rare freshwater algal spores, suggesting a cool and relatively humid period.

 

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

Two examples of grains pollen from the Lopez de Bertodano Formation: Podocarpidites sp. (left) and Nothofagidites asperus (right). From Bowman et al, 2014.

In the phase two the increased abundance of Phyllocladidites mawsonii implies a gradual increase in humidity. During phase three, bryophytes began to increase. The phase four is characterised by relatively high abundances of Podocarpidites spp. and relatively low levels of Nothofagidites spp.
The phase 5 is characterised by a rapidly changing sequence of abundance peaks of different taxa, which may indicate a successional turnover in forest composition. The phase six suggests a return to a cool climatic conditions with high abundances of Araucariacites australis and Nothofagidites at the top of the section. It seems that Araucariaceae were capable of surviving long periods of adverse climatic conditions during the Early Pleistocene, but most modern araucarians have subtropical to mesothermal climatic preferences.

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 composition of the terrestrial palynoflora indicates that the Maastrichtian climate fluctuated from cool, humid conditions, through a rapid warming about 2 million years prior to the K–Pg event – which is consistent with the evidence from the marine palynomorph record –  followed by cooling conditions in the earliest Danian.

 

Two examples of spores from the  Lopez de Bertodano Formation: Grapnelispora sp. (left) and Azolla sp.(right).

Two examples of spores from the Lopez de Bertodano Formation: Grapnelispora sp. (left) and Azolla sp.(right). From Bowman et al, 2014.

 

Reference:

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. 10.1016/j.palaeo.2014.04.018
David B. Kemp, Stuart A. Robinson, J. Alistair Crame, Jane E. Francis, Jon Ineson, Rowan J. Whittle, Vanessa Bowman, and Charlotte O’Brien, A cool temperate climate on the Antarctic Peninsula through the latest Cretaceous to early Paleogene, Geology (2014) doi: 10.1130/G35512.1

 

Isabel Clifton Cookson, the first Australian palynologist.

Isabel Clifton Cookson (1893-1973). From Wikimedia Commons

Isabel Clifton Cookson (1893-1973). From Wikimedia Commons

Isabel Clifton Cookson was one of Australia’s first professional women scientists, but unlike Adele V. Vicent, who studied  the importance Silurian-Devonian floras in Victoria,  her scientific work is well recognized.  She was one of the most prominent palynologist of the twenty century. She described a total of 110 genera, 557 species and 32 sub especific taxa of palynomorphs and plants, and published 93 scientific papers (some of them in collaboration with other prominent scientists).

She was born on December 25, 1893 in Melbourne, Australia. After graduating in Zoology and Botany at the University of Melbourne in 1916, she worked for a brief time at the National Museum of Victoria and became interested in fossil plants. 

Between 1916 and 1917 she received the Government Research Scholarship, for work on the flora of the Northern Territory of Australia and was awarded with the McBain Research Scholarship in biology. She also collaborated with some illustrations to the  book The Flora of the Northern Territory by Alfred J. Ewart and O. B. Davies.

Cooksonia pertoni, one of the earliest land plants (Credit: Hans Steur, The Netherlands.)

Cooksonia pertoni, one of the earliest land plants (Credit: Hans Steur, The Netherlands.)

In 1925, she went to England  to study with Professor Le Rayner  and with Professor Sir A. C. Seward, an authority on  fossil plants. She returned a year later as a mycologist in cotton research in the University of Manchester, where she met Professor W. H. Lang.  She started an important and  productive academic relationship with Lang, who named the genus Cooksonia in her honour.

In 1932, she returned to Melbourne and became mentor of many female researchers like Lorna Medwell and Mary E. Dettmann.

During the 1940s , she began to conduct detailed palaeobotanical studies, with emphasis on pollen analysis and demonstrated the importance of plant microfossils  in biostratigraphy  and in oil exploration.

Lingulodinium machaerophorum is a dinoflagellate cyst first described by Deflandre and Cookson. From UCL.

Lingulodinium machaerophorum is a dinoflagellate cyst first described by Deflandre and Cookson. From UCL.

In the early 1950s, she was a pioneer in the study of marine palynomorphs: dinoflagellate cysts, acritarchs and chitinozoans from Australian Tertiary and Mesozoic sediments. She also worked with George Deflandre and Alfred Eisenack.

Although her important work, she only reached the senior lecturer status in the department of botany and officially retired in 1959.

After her retirement, she  continued doing active research work  mainly by self-funding thanks to her skills as an investor on the stock exchange.

Isabel Clifton Cookson died on 1 July 1973 at her Hawthorn home. In her honor,  the Botanical Society of America gives the Isabel Cookson Award since 1976,  to the best paper on palaeobotany presented at their annual meeting.

References:

Riding, James B.; Dettmann, Mary E.. 2013 The first Australian palynologist: Isabel Clifton Cookson (1893–1973) and her scientific work. Alcheringa: An Australasian Journal of Palaeontology. 1-33. 10.1080/03115518.2013.828252

Mary E. Dettmann, ‘Cookson, Isabel Clifton (1893–1973)’, Australian Dictionary of Biography, National Centre of Biography, Australian National University