Volume 26, Issue 19, Pages (October 2016)

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Volume 26, Issue 19, Pages 2667-2673 (October 2016) Zooplankton Gut Passage Mobilizes Lithogenic Iron for Ocean Productivity  Katrin Schmidt, Christian Schlosser, Angus Atkinson, Sophie Fielding, Hugh J. Venables, Claire M. Waluda, Eric P. Achterberg  Current Biology  Volume 26, Issue 19, Pages 2667-2673 (October 2016) DOI: 10.1016/j.cub.2016.07.058 Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 1 Glacial Supply of Iron, Phytoplankton Distribution, and Krill Grazing at South Georgia (A) The Southern Ocean with the study area at South Georgia (red box) overlaying a chlorophyll a (chl a) climatology derived from MODIS-Aqua (July 2002–February 2015). (B–H) Results from our study period: December 25, 2010–January 19, 2011. (B) Salinity in surface waters. Cumberland Bay (CB) is a major glacial outlet. (C) Distribution of total dissolvable iron (unfiltered, TDFe). Information on the distribution of dissolved iron (<0.2 μm, DFe) and correlations between salinity and both TDFe and DFe are given in Figure S1. (D) Distribution of chl a concentrations (μg L−1). (E) Distribution of krill density (g wet mass m−2). (F) Proportion of diatoms in the suspended particulate matter (PM) at 20 m water depth. (G) Proportion of fecal pellets in the suspended particulate matter at 20 m water depth. (H) Stomach content of freshly caught krill. Current Biology 2016 26, 2667-2673DOI: (10.1016/j.cub.2016.07.058) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 2 Krill Iron Cycling (A–D) Changes in krill characteristics with increasing distance from the major glacial outlet (Cumberland Bay). (A) Volume of lithogenic particles in krill stomachs. (B) Total particulate iron content in krill muscle tissue. (C) Total particulate iron content in krill fecal pellets. (D) Labile iron content in krill fecal pellets and in suspended particulate matter (PM) at 20 m water depth. Dm, dry mass; TPFe, total particulate iron. (E and F) Results from nine short-term shipboard incubations of freshly caught krill. (E) DFe excretion rates in relation to the volume of diatoms in krill stomachs. (F) TDFe release rates in relation to the dry mass of fecal pellets produced during 3 hr incubations. Table S1 includes calculations of total DFe supply by krill. Current Biology 2016 26, 2667-2673DOI: (10.1016/j.cub.2016.07.058) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 3 Schema of Iron Flux through Krill and Pathways of DFe Supply The numbers (nmol Fe g−1 krill dm d−1) indicate the partitioning of ingested iron between body tissue, fecal pellets and the ambient pool of dissolved iron (details are given in Table S2). On a daily basis, ∼5% of the krill body iron content is assimilated, >200% is egested via fecal pellets, and ∼10% is directly excreted as DFe. This indicates the high iron content of the ingested food compared to krill’s own requirements. The iron that is ingested and mobilized by krill can reach the DFe pool via several pathways (indicated in A–D). (A) Direct excretion by krill. (B) Release from krill fecal pellets due to microbial degradation and zooplankton coprophagy. (C) Release from krill fecal pellets due to the dissolution of particulate iron via complexation with ligands or photochemical reactions. (D) Digestion of krill tissue by predators. All of these processes can occur in the upper mixed layer but may also happen in deeper water due to krill vertical migration and the sinking of their pellets. Black and open arrows represent the relative fractions sourced from lithogenic and biogenic iron, respectively. Gray arrows indicate processes that remain to be quantified. In Table S3, DFe fluxes from krill are compared to those from physical sources. Current Biology 2016 26, 2667-2673DOI: (10.1016/j.cub.2016.07.058) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 4 The Potential Role of Krill in Phytoplankton Fertilization and Grazing Losses (A) Spatial distribution of negative (blue-purple) and positive (yellow-red) slope values for the regression between median chl a concentration and summer krill abundance on the South Georgia shelf for the years 2002–2013 (details in Figure S2). Chl a concentrations were derived from ocean color radiometry (MODIS 2002–2013, mid-August to mid-April, 8-day composites). The black lines are drifter trajectories that indicate that surface currents link the northern shelf of South Georgia to the main bloom area downstream, with a transit time of 20–50 days [42]. (B) The time course of the average annual chl a development downstream of South Georgia (yellow-red area in A) and on the shelf (blue-purple area in A) during high- and low-krill years. The left plot indicates that the blooms downstream extend into autumn when the krill abundance on the shelf is high, but they are restricted to spring and summer when the krill abundance is low. In the area that benefits most from fertilization, the median chl a concentration increased from 1.0 to 1.3 μg L−1 and the bloom duration from 12 to 16 weeks in high-krill years. A bloom was defined as ≥1 μg chl a L−1. Low-krill years at South Georgia: 2002/2003, 2003/2004, 2004/2005, 2008/2009, 2010/2011, 2012/2013. High-krill years: 2005/2006, 2007/2008, 2009/2010, 2011/2012. Current Biology 2016 26, 2667-2673DOI: (10.1016/j.cub.2016.07.058) Copyright © 2016 Elsevier Ltd Terms and Conditions