Coccolithophore fluxes in the open tropical North Atlantic: influence of thermocline depth, Amazon water, and Saharan dust

Coccolithophores are calcifying phytoplankton and major contributors to both the organic and inorganic oceanic carbon pumps. Their export fluxes, species composition, and seasonal patterns were determined in two sediment trap moorings (M4 at 12 degrees N, 49 degrees W and M2 at 14 degrees N, 37 degrees W) collecting settling particles synchronously from October 2012 to November 2013 at 1200 m of water depth in the open equatorial North Atlantic. The two trap locations showed a similar seasonal pattern in total coccolith export fluxes and a predominantly tropical coccolithophore settling assemblage. Species fluxes were dominated throughout the year by lower photic zone (LPZ) taxa (Florisphaera profunda, Gladiolithus flabellatus) but also included upper photic zone (UPZ) taxa (Umbellosphaera spp., Rhabdosphaera spp., Umbilicosphaera spp., Helicosphaera spp.). The LPZ flora was most abundant during fall 2012, whereas the UPZ flora was more important during summer. In spite of these similarities, the western part of the study area produced persistently higher fluxes, averaging 241 x 10(7) +/- 76 x 10(7) coccoliths m(-2) d(-1) at station M4 compared to only 66 x 10(7) +/- 31 x 10(7) coccoliths m(-2) d(-1) at station M2. Higher fluxes at M4 were mainly produced by the LPZ species, favoured by the westward deepening of the thermocline and nutricline. Still, most UPZ species also contributed to higher fluxes, reflecting enhanced productivity in the western equatorial North Atlantic. Such was the case of two marked flux peaks of the more opportunistic species Gephyrocapsa muellerae and Emiliania huxleyi in January and April 2013 at M4, indicating a fast response to the nutrient enrichment of the UPZ, probably by wind-forced mixing. Later, increased fluxes of G. oceanica and E. huxleyi in October-November 2013 coincided with the occurrence of Amazon-River-affected surface waters. Since the spring and fall events of 2013 were also accompanied by two dust flux peaks, we propose a scenario in which atmospheric dust also provided fertilizing nutrients to this area. Enhanced surface buoyancy associated with the river plume indicates that the Amazon acted not only as a nutrient source, but also as a surface density retainer for nutrients supplied from the atmosphere. Nevertheless, lower total coccolith fluxes during these events compared to the maxima recorded in November 2012 and July 2013 indicate that transient productivity by opportunistic species was less important than "background" tropical productivity in the equatorial North Atlantic. This study illustrates how two apparently similar sites in the tropical open ocean actually differ greatly in ecological and oceanographic terms. The results presented here provide valuable insights into the processes governing the ecological dynamics and the downward export of coccolithophores in the tropical North Atlantic.