The Eocene Arctic Azolla bloom: environmental conditions, productivity and carbon drawdown

Enormous quantities of the free‐floating freshwater fern Azolla grew and reproduced in situ in the Arctic Ocean during the middle Eocene, as was demonstrated by microscopic analysis of microlaminated sediments recovered from the Lomonosov Ridge during Integrated Ocean Drilling Program (IODP) Expedition 302. The timing of the Azolla phase (~48.5 Ma) coincides with the earliest signs of onset of the transition from a greenhouse towards the modern icehouse Earth. The sustained growth of Azolla, currently ranking among the fastest growing plants on Earth, in a major anoxic oceanic basin may have contributed to decreasing atmospheric pCO2 levels via burial of Azolla‐derived organic matter. The consequences of these enormous Azolla blooms for regional and global nutrient and carbon cycles are still largely unknown. Cultivation experiments have been set up to investigate the influence of elevated pCO2 on Azolla growth, showing a marked increase in Azolla productivity under elevated (760 and 1910 ppm) pCO2 conditions. The combined results of organic carbon, sulphur, nitrogen content and 15N and 13C measurements of sediments from the Azolla interval illustrate the potential contribution of nitrogen fixation in a euxinic stratified Eocene Arctic. Flux calculations were used to quantitatively reconstruct the potential storage of carbon (0.9–3.5 1018 gC) in the Arctic during the Azolla interval. It is estimated that storing 0.9 1018 to 3.5 1018 g carbon would result in a 55 to 470 ppm drawdown of pCO2 under Eocene conditions, indicating that the Arctic Azolla blooms may have had a significant effect on global atmospheric pCO2 levels through enhanced burial of organic matter.

[1]  A. Ridgwell,et al.  Ocean‐atmosphere partitioning of anthropogenic carbon dioxide on multimillennial timescales , 2010 .

[2]  J. Damsté,et al.  Biomarker lipids of the freshwater fern Azolla and its fossil counterpart from the Eocene Arctic Ocean , 2009 .

[3]  B. Popp,et al.  Surface water productivity and paleoceanographic implications in the Cenozoic Arctic Ocean , 2008 .

[4]  J. King,et al.  Age model and core-seismic integration for the Cenozoic Arctic Coring Expedition sediments from the Lomonosov Ridge , 2008 .

[5]  T. C. Moore,et al.  Salinity of the Eocene Arctic Ocean from oxygen isotope analysis of fish bone carbonate , 2008 .

[6]  Stefan Schouten,et al.  Cyclicity in the middle Eocene central Arctic Ocean sediment record: Orbital forcing and environmental response , 2008 .

[7]  R. Jordan,et al.  A Siliceous Microfossil View of Middle Eocene Arctic Paleoenvironments , 2008 .

[8]  Gerald R. Dickens,et al.  An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics , 2008, Nature.

[9]  J. King,et al.  The early Miocene onset of a ventilated circulation regime in the Arctic Ocean , 2007, Nature.

[10]  M. Follows,et al.  Ocean‐atmosphere partitioning of anthropogenic carbon dioxide on centennial timescales , 2007 .

[11]  R. Stein,et al.  Anoxia and high primary production in the Paleogene central Arctic Ocean: First detailed records from Lomonosov Ridge , 2006 .

[12]  S. A. Merkur’ev,et al.  Formation of the Eurasia Basin in the Arctic Ocean as inferred from geohistorical analysis of the anomalous magnetic field , 2006 .

[13]  M. Huber,et al.  Episodic fresh surface waters in the Eocene Arctic Ocean , 2006, Nature.

[14]  M. Huber,et al.  Subtropical Arctic Ocean temperatures during the Palaeocene/Eocene thermal maximum , 2006, Nature.

[15]  David C. Smith,et al.  The Cenozoic palaeoenvironment of the Arctic Ocean , 2006, Nature.

[16]  David Archer,et al.  Fate of fossil fuel CO2 in geologic time , 2005 .

[17]  J. Zachos,et al.  Marked Decline in Atmospheric Carbon Dioxide Concentrations During the Paleogene , 2005, Science.

[18]  H. Elderfield,et al.  Eocene bipolar glaciation associated with global carbon cycle changes , 2005, Nature.

[19]  Stefan Schouten,et al.  N2-fixing cyanobacteria supplied nutrient N for Cretaceous oceanic anoxic events , 2004 .

[20]  J. Montoya,et al.  High rates of N2 fixation by unicellular diazotrophs in the oligotrophic Pacific Ocean , 2004, Nature.

[21]  Stefan Schouten,et al.  Crenarchaeotal membrane lipids in lake sediments : a new paleotemperature proxy for continental paleoclimate reconstruction? , 2004 .

[22]  C. Yapp Fe(CO3)OH in goethite from a mid-latitude North American Oxisol: estimate of atmospheric CO2 concentration in the Early Eocene “climatic optimum” , 2004 .

[23]  A. Roberts,et al.  Magnetostratigraphic calibration of Eocene–Oligocene dinoflagellate cyst biostratigraphy from the Norwegian–Greenland Sea ☆ , 2004 .

[24]  J. Zachos,et al.  Early Cenozoic decoupling of the global carbon and sulfur cycles , 2003 .

[25]  L. Sternberg,et al.  Humidity estimate for the middle Eocene Arctic rain forest , 2003 .

[26]  J. G. Kuenen,et al.  Anaerobic ammonium oxidation by anammox bacteria in the Black Sea , 2003, Nature.

[27]  Stefan Schouten,et al.  Distributional variations in marine crenarchaeotal membrane lipids: a new tool for reconstructing ancient sea water temperatures? , 2002 .

[28]  J. Knies,et al.  Depositional environment and source rock potential of Miocene strata from the central Fram Strait: introduction of a new computing tool for simulating organic facies variations , 2002 .

[29]  M. Collinson The ecology of Cainozoic ferns , 2002 .

[30]  Christopher J. Nicholas,et al.  Warm tropical sea surface temperatures in the Late Cretaceous and Eocene epochs , 2001, Nature.

[31]  Vandna Rai,et al.  Effect of NaCl on nitrogen fixation of unadapted and NaCl-adapted Azolla pinnata–Anabaena azollae , 2001 .

[32]  D. Beerling,et al.  Paleobotanical evidence for near present-day levels of atmospheric Co2 during part of the tertiary. , 2001, Science.

[33]  H. Koizumi,et al.  Effect of free‐air CO2 enrichment (FACE) on CO2 exchange at the flood‐water surface in a rice paddy field , 2001 .

[34]  L. Sloan,et al.  Trends, Rhythms, and Aberrations in Global Climate 65 Ma to Present , 2001, Science.

[35]  C. Schubert,et al.  Tracking nutrient and productivity variations over the Last Deglaciation in the Arctic Ocean , 2001 .

[36]  P. Pearson,et al.  Atmospheric carbon dioxide concentrations over the past 60 million years , 2000, Nature.

[37]  A. J. Kaufman,et al.  THE ABUNDANCE OF 13C IN MARINE ORGANIC MATTER AND ISOTOPIC FRACTIONATION IN THE GLOBAL BIOGEOCHEMICAL CYCLE OF CARBON DURING THE PAST 800 MA , 1999 .

[38]  K. Grice,et al.  Biosynthetic effects on the stable carbon isotopic compositions of algal lipids: implications for deciphering the carbon isotopic biomarker record , 1998 .

[39]  Rachel Janes Growth and survival of Azolla filiculoides in Britain. II. Sexual reproduction. , 1998, The New phytologist.

[40]  J. Gale,et al.  Differentiating Day from Night Effects of High Ambient [CO2] on the Gas Exchange and Growth ofXanthium strumariumL. Exposed to Salinity Stress☆ , 1997 .

[41]  D. Greenwood,et al.  Eocene continental climates and latitudinal temperature gradients: Comment and Reply , 1996 .

[42]  D. Greenwood,et al.  Eocene continental climates and latitudinal temperature gradients , 1995 .

[43]  Ellery D. Ingall,et al.  Benthic phosphorus regeneration, net primary production, and ocean anoxia: A model of the coupled marine biogeochemical cycles of carbon and phosphorus , 1994 .

[44]  S. Wakeham,et al.  Molecular evidence for degradation and preservation of organic matter in the anoxic Black Sea Basin , 1994 .

[45]  M. Altabet,et al.  Sedimentary nitrogen isotopic ratio as a recorder for surface ocean nitrate utilization , 1994 .

[46]  S. Bunn,et al.  What sources of organic carbon drive food webs in billabongs? A study based on stable isotope analysis , 1993, Oecologia.

[47]  R. Saunders,et al.  The supraspecific taxonomy and evolution of the fern genusAzolla (Azollaceae) , 1993, Plant Systematics and Evolution.

[48]  P. R. Cary,et al.  Growth and nutrient composition of Azolla pinnata R. Brown and Azolla filiculoides Lamarck as affected by water temperature, nitrogen and phosphorus supply, light intensity and pH , 1992 .

[49]  H. Gunnarsdóttir,et al.  Eocene to Miocene Palynology of the Norwegian Sea (ODP Leg 104) , 1989 .

[50]  S. Idso,et al.  Atmospheric CO2 enrichment enhances survival of Azolla at high temperatures , 1989 .

[51]  S. Goyal,et al.  Interactive effects of exogenous combined nitrogen and phosphorus on growth and nitrogen fixation by azolla , 1989, Plant and Soil.

[52]  L. Gahagan,et al.  Plate tectonic reconstructions of the Cretaceous and Cenozoic ocean basins , 1988 .

[53]  H. Marschner Mineral Nutrition of Higher Plants , 1988 .

[54]  Joseph S. Meyer,et al.  Contribution of bacteria to release and fixation of phosphorus in lake sediments , 1988 .

[55]  A. Moretti,et al.  Influence of light and pH on growth and nitrogenase activity on temperate-grown Azolla , 1988, Biology and Fertility of Soils.

[56]  S. Idso,et al.  Interactive effects of Co2 and environment on photosynthesis of Azolla , 1988 .

[57]  M. Minagawa,et al.  Nitrogen isotope ratios of red tide organisms in the East China Sea: A characterization of biological nitrogen fixation , 1986 .

[58]  R. Berner Sedimentary pyrite formation: An update , 1984 .

[59]  I. Kaplan,et al.  Isotopic fractionation of dissolved nitrate during denitrification in the eastern tropical north pacific ocean , 1975 .

[60]  V. P. Skipski,et al.  Separation of lipid classes by thin-layer chromatography. , 1965, Biochimica et biophysica acta.

[61]  Roger Revelle,et al.  Carbon Dioxide Exchange Between Atmosphere and Ocean and the Question of an Increase of Atmospheric CO2 during the Past Decades , 1957 .

[62]  G. M. Wagner Azolla: A review of its biology and utilization , 2008, The Botanical Review.

[63]  L. Sloan,et al.  Early Paleogene oceans and climate: A fully coupled modeling approach using the NCAR CCSM , 2003 .

[64]  J. Hayes Fractionation of Carbon and Hydrogen Isotopes in Biosynthetic Processes , 2001 .

[65]  S. Manabe Early Development in the Study of Greenhouse Warming: The Emergence of Climate Models , 1997 .

[66]  H. M. Kingston,et al.  Microwave-Enhanced Chemistry, Fundamentals, Sample Preparation, and Applications, ACS professional r , 1997 .

[67]  G. A. Peters,et al.  The Azolla-Anabaena Symbiosis: Basic Biology , 1989 .

[68]  J. Hutchison,et al.  Eocene lower vertebrates from Ellesmere Island, Canadian Arctic Archipelago , 1980 .