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Picophytoplankton Biomass Distribution in the Global Ocean

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Picophytoplankton Biomass Distribution in the Global Ocean CMYK RGB History of Geo- and Space Open Access Open Sciences Advances in Science & Research Open Access Proceedings Drinking Water Drinking Water Engineering and Science Engineering and Science Open Access Access Open Discussions Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussions Earth Syst. Sci. Data Discuss., 5, 221–242, Earth 2012 System Earth System www.earth-syst-sci-data-discuss.net/5/221/2012/ Science Science doi:10.5194/essdd-5-221-2012 ESSDD © Author(s) 2012. CC Attribution 3.0 License. 5, 221–242, 2012 Open Access Open Open Access Open Data Data This discussion paper is/has been under review for the journal Earth System Science Discussions Picophytoplankton Data (ESSD). Please refer to the corresponding final paper in ESSD if available. Social Social biomass distribution in the global ocean Open Access Open Geography Open Access Open Geography Picophytoplankton biomass distribution E. T. Buitenhuis et al. in the global ocean Title Page E. T. Buitenhuis1, W. K. W. Li2, D. Vaulot3, M. W. Lomas4, M. Landry5, F. Partensky3, D. M. Karl6, O. Ulloa7, L. Campbell8, S. Jacquet9, F. Lantoine10, Abstract Instruments F. Chavez11, D. Macias12, M. Gosselin13, and G. B. McManus14 Data Provenance & Structure 1Tyndall Centre for Climate Change Research and School of Environmental Sciences, Tables Figures University of East Anglia, Norwich NR4 7TJ, UK 2Fisheries and Oceans Canada, Bedford Institute of Oceanography, Dartmouth, Nova Scotia, Canada J I 3UMR 7144 (CNRS and UPMC, Paris 06), Station Biologique, 29680 Roscoff, France 4Bermuda Institute of Ocean Sciences, St. George’s GE01, Bermuda, USA J I 5 Scripps Institution of Oceanography, University of California San Diego, La Jolla, Back Close California, USA 6 Department of Oceanography, University of Hawaii, Honolulu, HI, 96822, USA Full Screen / Esc 7Department of Oceanography, University of Concepcion,´ Casilla 160-C, Concepcion,´ Chile 8 Department of Oceanography, Texas A&M University, College Station, TX 77843, USA Printer-friendly Version 9INRA, UMR CARRTEL, 75 Avenue de Corzent, 74200 Thonon-les-Bains, France 10UPMC Univ Paris 06, CNRS, LECOB, Observatoire Oceanologique,´ 66650, Interactive Discussion Banyuls/Mer, France 11MBARI, 7700 Sandholdt Rd, Moss Landing, CA 95039, USA 221 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 12Department of Coastal Ecology and Management, Instituto de Ciencias Marinas de Andaluc´ıa (ICMAN-CSIC), Avd. Republica Saharaui s/n, CP11510, Puerto Real, Cadiz,´ Spain ESSDD 13Institut des sciences de la mer de Rimouski, Universite´ du Quebec´ a` Rimouski, 310 Allee´ des Ursulines, Rimouski, Quebec´ G5L 3A1, Canada 5, 221–242, 2012 14Department of Marine Sciences, University of Connecticut, Groton CT 06340, USA Received: 13 February 2012 – Accepted: 10 April 2012 – Published: 27 April 2012 Picophytoplankton biomass distribution Correspondence to: E. T. Buitenhuis (http://tinyurl.com/contacterik) in the global ocean Published by Copernicus Publications. E. T. Buitenhuis et al. Title Page Abstract Instruments Data Provenance & Structure Tables Figures J I J I Back Close Full Screen / Esc Printer-friendly Version Interactive Discussion 222 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Abstract ESSDD The smallest marine phytoplankton, collectively termed picophytoplankton, have been routinely enumerated by flow cytometry since the late 1980s, during cruises throughout 5, 221–242, 2012 most of the world ocean. We compiled a database of 40 946 data points, with separate 5 abundance entries for Prochlorococcus, Synechococcus and picoeukaryotes. We use Picophytoplankton average conversion factors for each of the three groups to convert the abundance data biomass distribution ◦ to carbon biomass. After gridding with 1 spacing, the database covers 2.4 % of the in the global ocean ocean surface area, with the best data coverage in the North Atlantic, the South Pacific and North Indian basins. The average picophytoplankton biomass is 12 ± 22 µg C l−1 or E. T. Buitenhuis et al. −2 10 1.9 g C m . We estimate a total global picophytoplankton biomass of 0.53–0.74 Pg C (17–39 % Prochlorococcus, 12–15 % Synechococcus and 49–69 % picoeukaryotes). Future efforts in this area of research should focus on reporting calibrated cell size, Title Page and collecting data in undersampled regions. Abstract Instruments Data Provenance & Structure 1 Introduction Tables Figures 15 Picophytoplankton are usually defined as phytoplankton less than 2 or 3 µm diameter (e.g. Sieburth et al., 1978; Takahashi and Hori, 1984; Vaulot et al., 2008). They are the J I smallest class of phytoplankton, and are composed of both prokaryotes and eukary- otes. The eukaryotes (0.8–3 µm) are a taxonomically diverse group that include rep- J I resentatives from four algal phyla: the Chlorophyta, Haptophyta, Cryptophyta and Het- Back Close 20 erokontophyta (Vaulot et al., 2008). The prokaryotes belong to the phylum Cyanobac- teria, and are subdivided into the genera Prochlorococcus (∼0.6 µm) and Synechococ- Full Screen / Esc cus (∼1 µm), although with each group having many ecotypes that dominate in different ocean regions (Johnson et al., 2006). Printer-friendly Version Picophytoplankton tend to dominate the phytoplankton biomass under oligotrophic Interactive Discussion 25 conditions such as in the subtropical gyres (Alvain et al., 2005), where their high sur- face to volume ratio makes them the best competitors for low nutrient concentrations 223 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | (Raven, 1998). The abundance of the prokaryotes is often inversely related with the eu- karyotes, which are favoured by more physically active mixed layers (e.g. Boumann et ESSDD al., 2011). Furthermore, as the temperate to subpolar North Atlantic and the Canadian 5, 221–242, 2012 high Arctic warm, picophytoplankton (specifically picoeukaryotes) have been found to 5 become an increasingly large fraction of the total chlorophyll (Li et al., 2009; Moran et al., 2010). Picophytoplankton As part of the marine ecology data synthesis effort (MAREDAT, this special issue), biomass distribution we compiled a database on picophytoplankton in the global ocean. MAREDAT is a in the global ocean community effort to synthesise abundance and carbon biomass data for the major lower E. T. Buitenhuis et al. 10 trophic level taxonomic groups in the marine ecosystem. It addresses both autotrophs and heterotrophs and covers the size range from bacteria to macrozooplankton. Title Page 2 Data Abstract Instruments We compiled data for picophytoplankton abundance in three taxonomic groups: Data Provenance & Structure Prochlorococcus, Synechococcus, and picoeukaryotes (Table 1). We used the size 15 range of picoeukaryotes as defined by the contributing researchers. The size range Tables Figures has a large impact on the resulting biomass (see Discussion). All of the data were obtained by flow cytometry. Both the raw data and the gridded data are available J I from PANGAEA (http://doi.pangaea.de/10.1594/PANGAEA.777385) and the MARE- DAT webpage (http://lgmacweb.env.uea.ac.uk/maremip/data/essd.shtml). J I Back Close 20 2.1 Conversion factors Full Screen / Esc Conversion factors from cell abundance to carbon biomass for the three picophyto- plankton groups were compiled from the literature (Table 2). Conversion factors were Printer-friendly Version either measured directly on unialgal cultures in the laboratory, or derived from indi- rect methods on in situ samples. Most of the indirect measures were derived from cell Interactive Discussion 25 sizes that were estimated from average forward angle light scatter (FALS) multiplied 224 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | by a carbon content per biovolume. The conversion factors of Veldhuis et al. (1997) were based on nitrate uptake in incubated in situ samples and assuming a C:N ratio of ESSDD 6. Since the biggest source of variability in the other indirect measures is the carbon 5, 221–242, 2012 content per biovolume, which was measured on laboratory cultures, the advantage of 5 using in situ biovolume to determine conversion factors does not seem to improve the local applicability of these data and we therefore used the directly measured conversion Picophytoplankton factors as the standard. biomass distribution in the global ocean 2.2 Quality control E. T. Buitenhuis et al. Contributed data were assumed to have undergone the contributing researchers own 10 internal quality control procedures. As a statistical filter for outliers, we applied the Chauvenet criterion (Buitenhuis et al., 2012) to the total carbon data. The data were Title Page not normally distributed, so we log transformed them, excluding zero values. No high Abstract Instruments outliers were found by this criterion. The highest picophytoplankton biomass in the −1 database is 575 µg C l , measured near the coast of Oman (Indian Ocean). Data Provenance & Structure Tables Figures 15 3 Results J I The database contains 40 946 data points. Data are included from a number of sta- tions that have been sampled repeatedly over many years, or programs where mea- J I surements have been made on a fine resolution grid. Therefore, after gridding, we Back Close obtained 10 747 data points on the World Ocean Atlas grid (1◦ × 1◦ × 33 vertical lay- 20 ers × 12 months), representing a coverage of vertically integrated and annually aver- Full Screen / Esc aged biomass for 2.4 % of the ocean surface. To limit the overrepresentation of well sampled locations, we present results of the gridded data. Only 15 % of the data are Printer-friendly Version from the Southern Hemisphere (Fig. 1a), 33 % are from the tropics (43 % of the ocean surface), while 13 % are from the polar oceans (5 % of the ocean surface). Observa- Interactive Discussion 25 tions in the top 112.5 m make up 81 % of the data (Fig. 1b). Zero values make up 1.6 % 225 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | of the data, and 95 % of those are from below 62.5 m depth. There is some sampling bias towards the growing season, with 67 % of the data sampled in the spring and ESSDD summer months (Fig.
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