1 Melting Glaciers Help Fuel Productivity Hot-spots Around Antarctica 2 3 Kevin. R. Arrigo and Gert L. van Dijken 4 5 Department of Environmental Earth System Science, Stanford University, Stanford, California, 6 USA 7 8 Abstract 9 Antarctic coastal polynyas are biologically rich ecosystems that support large populations of 10 mammals and birds and are strong sinks of atmospheric carbon dioxide. To support local 11 phytoplankton blooms, these ecosystems require a large input of iron, the source of which is 12 poorly known. Satellite data suggest that melting glaciers are a primary supplier of iron to 13 coastal polynyas, with basal melt rates explaining 58% of the between-polynya variance in 14 annual mean chlorophyll a concentration. Iron upwelled from sediments, which is partly 15 controlled by continental shelf width, is also important. Given the sensitivity of these biological 16 hotspots to iron released from melting ice shelves, future changes in glacial melt rates could 17 dramatically impact coastal ecosystems and the ability of continental shelf waters to sequester 18 atmospheric carbon dioxide. 19 20 1. Introduction 21 Antarctic coastal polynyas are areas of open water surrounded by sea ice that are maintained 22 throughout the year through the upwelling of warm water or as continental winds blow ice away 23 from shore [Bromwich, 1989]. Because polynya waters are already ice-free in early spring when 24 solar elevation is increasing rapidly, they often harbor extremely large phytoplankton 25 concentrations compared to surrounding ice covered waters; these blooms persist even after sea 26 ice has disappeared in summer [Arrigo and Van Dijken, 2003]. This enhanced biological 27 productivity enables polynyas to support the highest densities of upper trophic level organisms in 1 28 the Southern Ocean [Karnovsky et al., 2007] and increases the efficiency of the biological pump, 29 making areas like the Ross Sea polynya disproportionately large sinks of anthropogenic carbon 30 dioxide [Arrigo et al., 2008a]. 31 Phytoplankton growth and abundance in most Southern Ocean waters is limited by the 32 availability of iron, which is usually in short supply due to the lack of an efficient supply 33 mechanism [Boyd et al., 2012]. While many parts of the global ocean receive large iron fluxes 34 through atmospheric dust deposition, the isolation of high latitude Antarctic waters from 35 terrestrial iron sources limits its supply by this means [Boyd et al., 2012]. Consequently, some 36 of the most productive waters of the Southern Ocean are in the Scotia Sea, where iron from 37 shallow shelves is mixed into surface waters during advection of the Antarctic Circumpolar 38 Current [Ardelan et al., 2010; Chever et al., 2010; Planquette et al., 2011; Frants et al., 2013]. 39 In other parts of the Southern Ocean, iron is also brought into surface waters through convective 40 and turbulent mixing as well as upwelling [Fitzwater et al., 2000; Planquette et al., 2013; 41 Measures et al., 2013], from melting of drifting icebergs [Smith et al., 2007; Raiswell et al., 42 2008; Schwartz and Schlodlok, 2009; Lin et al., 2011] and sea ice [Fitzwater et al., 2000; Grotti 43 et al., 2005; Lannuzel et al., 2010], and a small amount from atmospheric deposition [Boyd et al., 44 2012]. 45 Like the rest of the Southern Ocean, the growth of phytoplankton in coastal polynyas 46 depends on the availability of iron [Coale et al., 2005; Alderkamp et al., 2012; Gerringa et al., 47 2012]. However, despite the ecological and biogeochemical importance of coastal polynyas, 48 little is known about the processes that fuel the growth of local phytoplankton populations. Of 49 the dozens of Antarctic coastal polynyas that have been identified [Arrigo and Van Dijken, 50 2003], only a handful has been studied in any detail. A recent study of polynyas in the 51 Amundsen Sea showed that iron introduced from melting at the base of the Pine Island Glacier 52 was the dominant dissolved iron pool on the continental shelf, despite relatively high 53 concentrations of sediment-derived iron [Gerringa et al., 2012]. This glacially-derived iron, 54 produced primarily as ice streams abrade continental crust while flowing to the coast [Shaw et al 2 55 2011], was needed to satisfy the iron requirement of the intense phytoplankton bloom that 56 formed in the Pine Island Bay polynya [Gerringa et al., 2012; Alderkamp et al., 2012; Mills et 57 al., 2012]. 58 59 2. Data 60 In the present study, we quantify the amount of phytoplankton biomass and rates of net 61 primary production (NPP) in all coastal polynyas around the Antarctic continent between 1998 62 and 2013 and relate these to local environmental conditions. We identified and mapped 46 63 coastal polynyas (Fig. 1) using satellite-derived sea ice concentrations from the Special Sensor 64 Microwave/Imager (SSM/I). For each polynya, we used satellite data to quantify daily open 65 water area (from SSM/I), weekly sea surface temperature (SST, from the Advanced Very High 66 Resolution Radiometer, AVHRR, OISST), 8-day chlorophyll a (Chl a) concentrations (from 67 SeaWiFS and MODIS-Aqua), and calculated the daily rate of net primary production (NPP) 68 [Arrigo et al., 2008]. At each polynya location, we used bathymetric data to calculate the width 69 of the continental shelf (the shortest distance between the local coastline and the 1000 m 70 isobath). Because 80% of Antarctic coastal polynyas are adjacent to ice shelves, we obtained ice 71 shelf basal melt rates calculated from recent satellite-based estimates of the balance between ice 72 accumulation and thinning [Rignot et al., 2013]. Basal melt rates were considered a proxy for 73 the amount of iron introduced into adjacent polynyas. To determine what factors control 74 phytoplankton abundance within the polynyas, we regressed annual mean Chl a concentration 75 for each polynya against 1) local SST, 2) the length of the phytoplankton growing season (the 76 number of days per year that open water area in the polynya exceeded 50% of its maximum 77 value), 3) the continental shelf width, and 4) the total basal melt rate (Gt yr-1) of any ice shelves 78 located in the vicinity of each polynya. For all data, means are presented with standard 79 deviations. 80 81 3. Results 3 82 While coastal polynyas, by definition, have either reduced or no ice cover during winter, all 83 46 polynyas expanded greatly in size in October and November due to intensified rates of sea ice 84 melt, and contracted in March as sea ice began to freeze. The annual mean open water area 85 varied by approximately three orders of magnitude between polynyas, ranging from 954±196 86 km2 in the West Lazarev Sea polynya to 239,603±39,132 km2 in the Ross Ice Shelf polynya 87 (Table 1). Although the annual mean size of all polynyas was 14,063±35,803 km2, most 88 polynyas were substantially smaller than the mean, with 31 polynyas having annual mean open 89 water areas <10,000 km2 and 24 with open water areas <5,000 km2. 90 Surface Chl a concentrations in coastal polynyas began to increase in October and peaked 91 around January. By March, low solar elevation precluded net phytoplankton growth and Chl a 92 concentrations fell back to pre-bloom levels. The annual mean Chl a concentration calculated 93 over the phytoplankton growing season in the 46 polynyas ranged from 0.17±0.10 mg m-3 94 (Lutzoh-Holm Bay polynya) to 2.28±0.63 mg m-3 (Amundsen Sea polynya), averaging 95 0.60±0.42 mg m-3 (Table 1). The mean Chl a concentration for coastal polynyas was more than 96 double the concentration in non-shelf waters of the Southern Ocean [Arrigo et al., 2008b]. 97 The annual rate of NPP varied by a factor of 16 between coastal polynyas, ranging from 6.6 g 98 C m-2 yr-1 in the Lutzoh-Holm Bay polynya to 106 g C m-2 yr-1 in the Amundsen Sea polynya 99 (Table 1). The annual mean Chl a concentration was the most important factor controlling 100 annual NPP, explaining 91% of the variance between polynyas. Total NPP in each polynya is 101 calculated as a product of the annual NPP rate and the polynya area. Because of the large 102 difference in area between polynyas, total NPP was far more variable than annual NPP, ranging 103 over four orders of magnitude between polynyas (0.01-22.2 Tg C yr-1). Of the 46 polynyas, 40 104 had total NPP values below 1.0 Tg C yr-1 and 20 had values below 0.1 Tg C yr-1 (Table 1). 105 Although temperature and light availability can be important regulators of phytoplankton 106 populations in polar waters, linear regression analysis demonstrated that neither SST nor the 107 length of the phytoplankton growing season had a significant impact on annual mean Chl a 108 concentrations within the polynyas (p >0.05). Because Antarctic coastal polynyas are located in 4 109 very high latitude waters, SST varies little throughout the year and variability between polynyas 110 is small, minimizing its effect. The lack of a relationship between the length of the open water 111 season and either annual mean Chl a concentration or NPP is likely because coastal polynyas 112 remain near their annual maximum size for an average of 126 days, more than twice as long as 113 the average phytoplankton bloom (62 days, defined as the number of days that Chl a 114 concentration exceeded 50% of its annual maximum, Table 1). As a result, light availability did 115 not limit phytoplankton growth in the polynyas. It should be noted, however, that we were 116 unable to assess the impact of mixed layer depth, which also controls the amount of light 117 available to phytoplankton, on either Chl a or NPP. While existing data from a small number of 118 polynyas suggests that mixed layers are relatively shallow and do not limit phytoplankton growth 119 rates [Arrigo et al., 1999; Alderkamp et al., 2012], we cannot rule out the possibility that MLD 120 may be responsible for some of the unexplained variance in Chl a concentration between 121 polynyas. 122 In contrast to the negligible impact of temperature and light, the width of the continental 123 shelf explained 40% of the variance in annual mean Chl a concentration between polynyas [Fig. 124 2a). This high correlation is consistent with a sediment source of iron to coastal polynyas. 125 Dissolved iron concentrations increase with proximity to the Antarctic coast [Sedwick et al., 126 2008] and shelf waters are especially enriched in iron [Fitzwater et al., 2000; Gerringa et al., 127 2012]. As waters upwell onto the continental shelf [Gerringa et al., 2012] or during winter 128 convection [Blain et al., 2008], wider continental shelves increase the contact between upwelling 129 water and iron-rich sediments, allowing more iron to be entrained into surface waters, and 130 support greater phytoplankton abundance [Bruland et al., 2005; Biller and Bruland, 2013; Biller 131 et al., 2013]. 132 Although continental shelf sediments are an important iron source, iron from melting glaciers 133 appears to be even more critical in controlling phytoplankton abundance in coastal polynyas. In 134 our study, 59% of the variance in annual mean Chl a concentration was explained by the basal 135 melt rate of nearby ice shelves (Fig. 2b). The y-intercept in Fig. 2b, which provides an estimate 5 136 of the annual mean Chl a concentration to be expected under conditions of no basal melt, was 137 0.39±0.37 mg m-3, indistinguishable from the mean Chl a concentration in the nine polynyas that 138 are not located near an ice shelf (0.44±0.19 mg m-3). Therefore, including the nine polynyas not 139 associated with an ice sheet and giving them a basal melt rate of zero had no impact in the 140 amount of variance explained. A multiple linear regression that included both continental shelf 141 width and basal melt rate of nearby ice shelves was able to explain 70% of the variance in mean 142 annual Chl a concentration among the 46 coastal polynyas. 143 144 4. Conclusions 145 From the high correlation between basal melt rates and phytoplankton biomass in Antarctic 146 polynyas, we infer that iron released from melting glaciers stimulates phytoplankton growth in 147 polynyas throughout the Antarctic. We recognize, though, that there may be alternative, albeit 148 less likely, explanations for this correlation. First, sea ice in the vicinity of expanding polynyas 149 can also release iron in spring and summer when it melts [Grotti et al., 2005]. However, sea ice 150 is not a true source of iron since it accumulates iron from other sources (mostly from the water 151 column, Fitzwater et al., 2000]. Second, intrusions of warm Circumpolar Deep Water can 152 resuspend sediment iron and also accelerate basal melting [Dutrieux et al., 2014]. Thus, high 153 phytoplankton abundance in polynyas near ice shelves experiencing enhanced basal melting 154 could be driven by increased upwelling rates and a greater flux of sediment-derived iron, rather 155 than a glacial iron source. We consider this unlikely given that melting ice shelves are known 156 sources of dissolved iron to local waters [Gerringa et al., 2012] and our observation that basal 157 melt rates explained 50% more of the variance in Chl a between polynyas than shelf width. 158 Finally, melting glaciers could reduce the density of local waters, increasing stratification and 159 enhancing surface light levels, thereby increasing phytoplankton growth, irrespective of any iron 160 input. However, the limited available data suggest that as meltwater upwells at the face of the 161 glaciers, they promote deep mixing, rather than stratification, and that phytoplankton abundance 162 in these upwelled waters is very low [Gerringa et al., 2012, Alderkamp et al., 2012]. 6 163 Annual rates of NPP in Antarctic coastal polynyas are among the highest in the Southern 164 Ocean [Arrigo and Van Dijken, 2003] due in large part to the relaxation of nutrient limitation by 165 iron released from melting glaciers. Our results also show that 50% of total NPP in Antarctic 166 polynyas, which are important but often overlooked carbon sinks, is associated with the Ross Sea 167 polynya and 50% is associated with the other 45 polynyas (Table 1). If we assume that the ratio 168 of NPP to the magnitude of the anthropogenic carbon dioxide sink measured in the Ross Sea 169 (0.24) [Arrigo et al., 2008] is applicable to other polynyas, all Antarctic polynyas combined 170 would represent an anthropogenic carbon dioxide sink of about 0.019 Pg C yr-1. Thus, despite 171 that fact that coastal Antarctic polynyas represent only 0.06% of the ocean area south of 62°S, 172 accounting for them in analyses of oceanic carbon dioxide sequestration would shift these waters 173 from being a 0.01 Pg C yr-1 source [Takahashi et al., 2009] to a 0.01 Pg C yr-1 sink of 174 anthropogenic carbon dioxide. 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Vernet, Free-drifting icebergs: Hot spots of chemical and biological 272 enrichment in the Weddell Sea, Science, 317(5837), 478-482, doi: 10.1126/science.1142834. 273 274 11 275 Figure legends 276 277 Figure 1. Location of the 46 coastal polynyas included in this study. 278 279 Figure 2. Regression of annual mean chlorophyll a concentration against (A) continental shelf 280 width and (B) basal melt rate (Gt yr-1) of nearby ice shelves for the 37 polynyas that were 281 located near an ice shelf (Rignot et al., 2013). 12 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 Table 1. Mean (±SD) statistics for 46 Antarctic polynyas for the years 1998-2013. The column heading #days>50%max OW gives the number of days per year that the open water area in each polynya exceeded 50% of the annual maximum open water area. The column heading #days>50%max NPP gives the number of days per year that NPP in each polynya exceeded 50% of the annual maximum NPP. The basal melt rates and corresponding glacier names are from Rignot et al. [2013]. Name Polynya Sulzberger Bay 1 Hull Bay 2 Wrigley Gulf 3 Amundsen Sea 4 Pine Island Bay 5 Eltanin Bay 6 Ronne Entrance 7 Marguerite Bay 8 Larsen Ice Shelf 9 Ronne Ice Shelf 10 Brunt Ice Shelf 11 Lyddan Island 12 Seal Bay 13 Cape Norvegia 14 Jelbart Ice Shelf 15 Fimbul Ice Shelf 16 W. Lazarev Ice Shelf 17 E. Lazarev Ice Shelf 18 Breid Bay 19 Vestvika Bay 20 Lutzoh-Holm Bay 21 Amundsen Bay 22 Cape Borle 23 Stefansson Bay 24 Holme Bay 25 Bjerkø Peninsula 26 MacKenzie Bay 27 Prydz Bay 28 West Ice Shelf 29 Farr Bay 30 Mill Island 31 Cape Nutt 32 Open water area (km2) 6473±1689 6359±828 11100±1151 31844±4679 19688±5448 11283±1483 15120±1986 4348±550 4727±2774 44680±56686 15189±4153 6003±1434 4340±865 2337±297 2195±371 3493±402 954±196 1995±292 1129±175 2360±440 1657±1881 1845±721 4258±980 2087±438 4650±1060 12110±899 1867±237 54541±5655 2148±584 26723±4260 2491±1139 1463±353 Annual NPP (g C m-2 yr-1) 33.0±22.9 30.2±15.8 43.0±19.1 105.4±21.9 72.2±28.2 46.1±14.1 61.7±18.3 67.1±24.2 20.3±29.6 43.8±32.5 35.5±18.2 27.9±17.8 23.0±17.1 16.4±9.3 16.5±10.5 17.6±6.9 10.7±7.1 17.0±5.8 15.3±8.9 17.7±9.3 6.7±10.8 8.6±4.6 21.3±12.8 26.9±9.1 36.8±13.4 47.7±11.9 44.1±18.5 88.3±19.4 12.7±6.3 52.6±15.1 17.3±16.3 13.8±11.9 Total NPP (Tg C yr-1) 0.24±0.19 0.20±0.12 0.49±0.26 3.38±0.90 1.55±0.96 0.53±0.22 0.95±0.38 0.30±0.13 0.15±0.28 3.41±7.04 0.59±0.38 0.19±0.13 0.11±0.09 0.04±0.02 0.04±0.03 0.06±0.03 0.01±0.01 0.03±0.01 0.02±0.01 0.04±0.03 0.03±0.07 0.02±0.01 0.10±0.08 0.06±0.03 0.18±0.08 0.58±0.17 0.08±0.04 4.84±1.29 0.03±0.02 1.45±0.52 0.06±0.07 0.02±0.02 Chl a (mg m-3) 0.68±0.55 0.60±0.45 0.97±0.61 2.28±0.63 1.25±0.60 0.74±0.25 1.10±0.47 1.23±0.51 0.50±0.54 0.85±0.79 0.55±0.27 0.52±0.25 0.46±0.28 0.36±0.15 0.32±0.11 0.33±0.12 0.25±0.14 0.25±0.08 0.28±0.10 0.28±0.11 0.17±0.10 0.25±0.13 0.29±0.20 0.33±0.12 0.49±0.19 0.64±0.17 0.83±0.36 1.43±0.43 0.28±0.12 0.72±0.27 0.34±0.21 0.29±0.21 Shelf # days> # days> Basal melt width (km) 50%max OW 50%max NPP (Gt yr-1) 163.5 90 69 18.2 71.3 101 64 4.2 139.3 105 53 144.9 264.2 114 45 181.2 444.7 108 51 101.2 372.8 139 62 24.5 346.6 135 70 56.8 266.2 211 83 89 196.2 128 69 20.7 453.9 64 76 113.5 176.6 95 65 1 174.6 102 71 8.7 109.9 84 72 8.7 53.0 106 52 5.7 53.0 113 59 0 130.2 120 54 23.5 77.8 138 40 3.2 85.0 176 77 6.3 45.1 184 63 7.5 109.2 143 77 14.1 128.8 104 52 5.7 67.4 94 53 6.7 44.5 140 89 92.2 115 62 88.9 107 61 96.8 136 68 215.2 217 65 270.8 217 65 37 130.2 124 74 27.2 158.9 129 68 72.6 168.7 144 61 3 117.1 138 61 3.6 Glacier name Sulzberger Land Getz Dotson/Crosson/Thwaites Pine Island Venable/Ferrigno Strange/Bach/Wilkins George VI Larsen C Ronne Brunt/Stancomb Riiser-Larsen Riiser-Larsen Quar/Ekstrom Jelbart Fimbul Vigrid Lazarev Borchgrevink Baudouin Shirase Rayner-Thyer Amery/Publications West Shackleton Tracy/Tremenchus Conger/Glenzer 13 321 322 323 324 325 326 327 Vincennes Bay Cape Poinsett Totten Glacier Henry Bay Paulding Bay Porpoise Bay Davis Bay 33 34 35 36 37 38 39 12339±1906 3986±744 1414±493 6512±1225 1568±412 1653±758 11608±981 37.4±14.4 18.9±10.1 10.4±7.7 20.7±9.5 7.6±7.7 13.2±12.9 29.8±8.6 0.48±0.23 0.08±0.05 0.02±0.01 0.14±0.09 0.01±0.02 0.03±0.04 0.35±0.12 0.46±0.19 0.36±0.30 0.36±0.24 0.31±0.18 0.25±0.15 0.32±0.30 0.40±0.16 124.3 181.2 71.9 142.6 160.2 113.8 209.3 138 138 141 134 113 126 142 69 56 36 65 48 30 72 5 Vincennes 27.4 Moscow University 6.7 Holmes 14