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The Influence of Environmental Variability on the Biogeography Of Biogeosciences, 14, 4905–4925, 2017 https://doi.org/10.5194/bg-14-4905-2017 © Author(s) 2017. This work is distributed under the Creative Commons Attribution 3.0 License. The influence of environmental variability on the biogeography of coccolithophores and diatoms in the Great Calcite Belt Helen E. K. Smith1,2, Alex J. Poulton1,a, Rebecca Garley3, Jason Hopkins4, Laura C. Lubelczyk4, Dave T. Drapeau4, Sara Rauschenberg4, Ben S. Twining4, Nicholas R. Bates2,3, and William M. Balch4 1National Oceanography Centre, European Way, Southampton, SO14 3ZH, UK 2Ocean and Earth Science, National Oceanography Centre Southampton, University of Southampton, Southampton, SO14 3ZH, UK 3Bermuda Institute of Ocean Sciences, 17 Biological Station, Ferry Reach, St. George’s GE 01, Bermuda 4Bigelow Laboratory for Ocean Sciences, 60 Bigelow Drive, P.O. Box 380, East Boothbay, Maine 04544, USA apresently at: The Lyell Centre, Heriot-Watt University, Edinburgh, EH14 4AS, UK Correspondence to: Helen E. K. Smith ([email protected]) Received: 27 March 2017 – Discussion started: 13 April 2017 Revised: 24 August 2017 – Accepted: 20 September 2017 – Published: 7 November 2017 Abstract. The Great Calcite Belt (GCB) of the Southern numerically dominant and widely distributed. A combination Ocean is a region of elevated summertime upper ocean cal- of SST, macronutrient concentrations, and pCO2 provided cite concentration derived from coccolithophores, despite the the best statistical descriptors of the biogeographic variabil- region being known for its diatom predominance. The over- ity in biomineralizing species composition between stations. lap of two major phytoplankton groups, coccolithophores Emiliania huxleyi occurred in silicic acid-depleted waters be- and diatoms, in the dynamic frontal systems characteristic of tween the Subantarctic Front and the Polar Front, a favorable this region provides an ideal setting to study environmental environment for this species after spring diatom blooms re- influences on the distribution of different species within these move silicic acid. Multivariate statistics identified a combi- taxonomic groups. Samples for phytoplankton enumeration nation of carbonate chemistry and macronutrients, covarying were collected from the upper mixed layer (30 m) during with temperature, as the dominant drivers of biomineralizing two cruises, the first to the South Atlantic sector (January– nanoplankton in the GCB sector of the Southern Ocean. February 2011; 60◦ W–15◦ E and 36–60◦ S) and the sec- ond in the South Indian sector (February–March 2012; 40– 120◦ E and 36–60◦ S). The species composition of coccol- ithophores and diatoms was examined using scanning elec- 1 Introduction tron microscopy at 27 stations across the Subtropical, Po- lar, and Subantarctic fronts. The influence of environmental The Great Calcite Belt (GCB), defined as an elevated partic- parameters, such as sea surface temperature (SST), salinity, ulate inorganic carbon (PIC) feature occurring alongside sea- carbonate chemistry (pH, partial pressure of CO2 (pCO2/, sonally elevated chlorophyll a in austral spring and summer alkalinity, dissolved inorganic carbon), macronutrients (ni- in the Southern Ocean (Fig. 1; Balch et al., 2005), plays an trate C nitrite, phosphate, silicic acid, ammonia), and mixed important role in climate fluctuations (Sarmiento et al., 1998, layer average irradiance, on species composition across the 2004), accounting for over 60 % of the Southern Ocean area ◦ GCB was assessed statistically. Nanophytoplankton (cells (30–60 S; Balch et al., 2011). The region between 30 and ◦ 2–20 µm) were the numerically abundant size group of 50 S has the highest uptake of anthropogenic carbon dioxide biomineralizing phytoplankton across the GCB, with the coc- (CO2/ alongside the North Atlantic and North Pacific oceans colithophore Emiliania huxleyi and diatoms Fragilariopsis (Sabine et al., 2004). Our knowledge of the impact of inter- nana, F. pseudonana, and Pseudo-nitzschia spp. as the most acting environmental influences on phytoplankton distribu- tion in the Southern Ocean is limited. For example, we do Published by Copernicus Publications on behalf of the European Geosciences Union. 4906 H. E. K. Smith et al.: Coccolithophore and diatom biogeography of the Great Calcite Belt Figure 1. Rolling 32-day composite from MODIS Aqua for both (a) chlorophyll a (mg m−3/ and (b) PIC (mol m−3/ for the South Atlantic sector (17 January to 17 February 2011) and the South Indian sector (18 February to 20 March 2012). Station number identifiers and averaged positions of fronts as defined by Orsi et al. (1995) are superimposed: Subtropical Front (STF), Subantarctic Front (SAF), Polar Front (PF), Southern Antarctic Circumpolar Current Front (SACCF), and southern boundary (SBDY). not yet fully understand how light and iron availability or atoms with higher silicic acid requirements (e.g., Fragilari- temperature and pH interact to control phytoplankton bio- opsis kerguelensis) are generally more abundant south of the geography (Boyd et al., 2010, 2012; Charalampopoulou et PF (Froneman et al., 1995). High abundances of nanoplank- al., 2016). Hence, if model parameterizations are to improve ton (coccolithophores, small diatoms, chrysophytes) have (Boyd and Newton, 1999) to provide accurate predictions of also been observed on the Patagonian Shelf (Poulton et al., biogeochemical change, a multivariate understanding of the 2013) and in the Scotia Sea (Hinz et al., 2012). Currently, full suite of environmental drivers is required. few studies incorporate small biomineralizing phytoplankton The Southern Ocean has often been considered as a to species level (e.g., Froneman et al., 1995; Bathmann et al., microplankton-dominated (20–200 µm) system with phyto- 1997; Poulton et al., 2007; Hinz et al., 2012). Rather, the fo- plankton blooms dominated by large diatoms and Phaeo- cus has often been on the larger and noncalcifying species cystis sp. (e.g., Bathmann et al., 1997; Poulton et al., 2007; in the Southern Ocean due to sample preservation issues Boyd, 2002). However, since the identification of the GCB as (i.e., acidified Lugol’s solution dissolves calcite, and light a consistent feature (Balch et al., 2005, 2016) and the recog- microscopy restricts accurate identification to cells > 10 µm; nition of picoplankton (< 2 µm) and nanoplankton (2–20 µm) Hinz et al., 2012). In the context of climate change and future importance in high-nutrient, low-chlorophyll (HNLC) wa- ecosystem function, the distribution of biomineralizing phy- ters (Barber and Hiscock, 2006), the dynamics of small toplankton is important to define when considering phyto- (bio)mineralizing plankton and their export need to be ac- plankton interactions with carbonate chemistry (e.g., Langer knowledged. The two dominant biomineralizing phytoplank- et al., 2006; Tortell et al., 2008) and ocean biogeochemistry ton groups in the GCB are coccolithophores and diatoms. (e.g., Baines et al., 2010; Assmy et al., 2013; Poulton et al., Coccolithophores are generally found north of the PF (e.g., 2013). Mohan et al., 2008), though Emiliania huxleyi has been ob- The GCB spans the major Southern Ocean circumpo- served as far south as 58◦ S in the Scotia Sea (Holligan et al., lar fronts (Fig. 1a): the Subantarctic Front (SAF), the Po- 2010), at 61◦ S across Drake Passage (Charalampopoulou et lar Front (PF), the Southern Antarctic Circumpolar Cur- al., 2016), and at 65◦ S south of Australia (Cubillos et al., rent Front (SACCF), and occasionally the southern boundary 2007). of the Antarctic Circumpolar Current (ACC; see Tsuchiya Diatoms are present throughout the GCB, with the Po- et al., 1994; Orsi et al., 1995; Belkin and Gordon, 1996). lar Front marking a strong divide between different size The Subtropical Front (STF; at approximately 10 ◦C) acts as fractions (Froneman et al., 1995). North of the PF, small the northern boundary of the GCB and is associated with diatom species, such as Pseudo-nitzschia spp. and Thalas- a sharp increase in PIC southwards (Balch et al., 2011). siosira spp., tend to dominate numerically, whereas large di- These fronts divide distinct environmental and biogeochem- Biogeosciences, 14, 4905–4925, 2017 www.biogeosciences.net/14/4905/2017/ H. E. K. Smith et al.: Coccolithophore and diatom biogeography of the Great Calcite Belt 4907 ical zones, making the GCB an ideal study area to examine dicating a potential change in future community structure. controls on phytoplankton communities in the open ocean Large phytoplankton species (> 50 µm) may also have phys- (Boyd, 2002; Boyd et al., 2010). A high PIC concentra- iological traits to withstand changes in ocean chemistry over tion observed in the GCB (1 µmol PIC L−1/ compared to the smaller-celled (< 50 µm) species (Flynn et al., 2012) and po- global average (0.2 µmol PIC L−1/ and significant quantities tentially be less susceptible to grazing pressure (Assmy et of detached E. huxleyi coccoliths (in concentrations > 20 000 al., 2013). Alternatively, there may be a shift towards small coccoliths mL−1; Balch et al., 2011) both characterize the phytoplankton groups due to the expansion of low-nutrient GCB. The GCB is clearly observed in satellite imagery (e.g., subtropical regions (Bopp et al., 2001, 2005). The response Balch et al., 2005; Fig. 1b;) spanning from the Patagonian of Southern Ocean phytoplankton biogeography to future cli- Shelf (Signorini et al., 2006; Painter et al., 2010) across the mate conditions, including ocean acidification, is complex Atlantic,
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