Abstract. The global distribution pattern of coccolithophorid blooms was mapped in order to ascertain the prevalence of these blooms in the world's oceans and to estimate their worldwide production of CaCO 3 and dimethyl sulfide (DMS). Mapping was accomplished by classifying pixels of 5-day global composites of coastal zone color scanner imagery into bloom and nonbloom classes using a supervised, multispectral classification scheme. Surface waters with the spectral signature of coccolithophorid blooms annually covered an average of 1.4 x 10 6 km 2 in the world oceans from 1979 to 1985, with the subpolar latitudes accounting for 71% of this surface area. Classified blooms were most extensive in the Subarctic North Atlantic. Large expanses of the bloom signal were also detected in the North Pacific, on the Argentine shelf and slope, and in numerous lower latitude marginal seas and shelf regions. The greatest spatial extent of classified blooms in subpolar oceanic regions occurred in the months from summer to early autumn, while those in lower latitude marginal seas occurred in midwinter to early spring. Though the classification scheme was efficient in separating bloom and nonbloom classes during test simulations, and biogeographical literature generally confirms the resulting distribution pattern of blooms in the subpolar regions, the cause of the bloom signal is equivocal in some geographic areas, particularly on shelf regions at lower latitudes. Standing stock estimates suggest that the presumed Emiliania huxleyi blooms act as a significant source of calcite carbon and DMS sulfur on a regional scale. On a global scale, however, the satellite-detected coccolithophorid blooms are estimated to play only a minor role in the annual production of these two compounds and their flux from the surface mixed layer.

[1] Carbonates are the largest reservoirs of carbon on Earth. From mid-Mesozoic time, the biologically catalyzed precipitation of calcium carbonates by pelagic phytoplankton has been primarily due to the production of calcite by coccolithophorids. In this paper we address the physical and chemical processes that select for coccolithophorid blooms detected in Sea-viewing Wide Field-of-view Sensor (SeaWiFS) ocean color imagery. Our primary goal is to develop both diagnostic and prognostic models that represent the spatial and temporal dynamics of coccolithophorid blooms in order to improve our knowledge of the role of these organisms in mediating fluxes of carbon between the ocean, the atmosphere, and the lithosphere. On the basis of monthly composite images of classified coccolithophorid blooms and global climatological maps of physical variables and nutrient fields, we developed a probability density function that accounts for the physical chemical variables that predict the spatiotemporal distribution of coccolithophorids in the world oceans. Our analysis revealed that areas with sea surface temperatures (SST) between 3°and 15°C, a critical irradiance between 25 and 150 mmol quanta m À2 s À1 , and decreasing nitrate concentrations (DN/Dt < 0) are selective for upper ocean large-scale coccolithophorid blooms. While these conditions favor both Northern and Southern Hemisphere blooms of the most abundant coccolithophorid in the modern oceans, Emiliania huxleyi, the Northern and Southern Hemisphere populations of this organism are genetically distinct. Applying amplified fragment length polymorphism as a marker of genetic diversity, we identified two major taxonomic clades of E. huxleyi; one is associated with the Northern Hemisphere blooms, while the other is found in the Southern Hemisphere. We suggest a rule of ''universal distribution and local selection'': that is, coccolithophorids can be considered cosmopolitan taxa, but their genetic plasticity provides physiological accommodation to local environmental selection pressure. Sea surface temperature, critical irradiance, and DN/Dt were predicted for the years 2060-2070 using the NCAR Community Climate System Model to generate future monthly probability distributions of coccolithophorids based upon the relationships observed between the environmental variables and coccolithophorid blooms in modern oceans. Our projected probability distribution analysis suggests that in the North Atlantic, the largest habitat for coccolithophorids on Earth, the areal extent of blooms will decrease by up to 50% by the middle of this century. We discuss how the magnitude of carbon fluxes may be affected by the evolutionary success of coccolithophorids in future climate scenarios.

The North Atlantic spring bloom is one of the largest annual biological events in the ocean, and is characterized by dominance transitions from siliceous (diatoms) to calcareous (coccolithophores) algal groups. To study the effects of future global change on these phytoplankton and the biogeochemical cycles they mediate, a shipboard continuous culture experiment (Ecostat) was conducted in June 2005 during this transition period. Four treatments were examined: (1) 12°C and 390 ppm CO 2 (ambient control), (2) 12°C and 690 ppm CO 2 (high pCO 2 ), (3) 16°C and 390 ppm CO 2 (high temperature), and (4) 16°C and 690 ppm CO 2 ('greenhouse'). Nutrient availability in all treatments was designed to reproduce the low silicate conditions typical of this late stage of the bloom. Both elevated pCO 2 and temperature resulted in changes in phytoplankton community structure. Increased temperature promoted whole community photosynthesis and particulate organic carbon (POC) production rates per unit chlorophyll a. Despite much higher coccolithophore abundance in the greenhouse treatment, particulate inorganic carbon production (calcification) was significantly decreased by the combination of increased pCO 2 and temperature. Our experiments suggest that future trends during the bloom could include greatly reduced export of calcium carbonate relative to POC, thus providing a potential negative feedback to atmospheric CO 2 concentration. Other trends with potential climate feedback effects include decreased community biogenic silica to POC ratios at higher temperature. These shipboard experiments suggest the need to examine whether future pCO 2 and temperature increases on longer decadal timescales will similarly alter the biological and biogeochemical dynamics of the North Atlantic spring bloom.

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In this paper, we review the state of the art and major challenges in current efforts to incorporate biogeochemical functional groups into models that can be applied on basin-wide and global scales, with an emphasis on models that might ultimately be used to predict how biogeochemical cycles in the ocean will respond to global warming. We define the term ''biogeochemical functional group'' to refer to groups of organisms that mediate specific chemical reactions in the ocean. Thus, according to this definition, ''functional groups'' have no phylogenetic meaning-these are composed of many different species with common biogeochemical functions. Substantial progress has been made in the last decade toward quantifying the rates of these various functions and understanding the factors that control them. For some of these groups, we have developed fairly sophisticated models that incorporate this understanding, e.g. for diazotrophs (e.g. Trichodesmium), silica producers (diatoms) and calcifiers (e.g. coccolithophorids and specifically Emiliania huxleyi). However, current representations of nitrogen fixation and calcification are incomplete, i.e., based primarily upon models of Trichodesmium and E. huxleyi, respectively, and many important functional groups have not yet been considered in open-ocean biogeochemical models. Progress has been made over the last decade in efforts to simulate dimethylsulfide (DMS) production and cycling (i.e., by dinoflagellates and prymnesiophytes) and denitrification, but these efforts are still in their infancy, and many significant problems remain. ARTICLE IN PRESSOne obvious gap is that virtually all functional group modeling efforts have focused on autotrophic microbes, while higher trophic levels have been completely ignored. It appears that in some cases (e.g., calcification), incorporating higher trophic levels may be essential not only for representing a particular biogeochemical reaction, but also for modeling export. Another serious problem is our tendency to model the organisms for which we have the most validation data (e.g., E. huxleyi and Trichodesmium) even when they may represent only a fraction of the biogeochemical functional group we are trying to represent.When we step back and look at the paleo-oceanographic record, it suggests that oxygen concentrations have played a central role in the evolution and emergence of many of the key functional groups that influence biogeochemical cycles in the present-day ocean. However, more subtle effects are likely to be important over the next century like changes in silicate supply or turbulence that can influence the relative success of diatoms versus dinoflagellates, coccolithophorids and diazotrophs. In general, inferences drawn from the paleo-oceanographic record and theoretical work suggest that global warming will tend to favor the latter because it will give rise to increased stratification. However, decreases in pH and Fe supply could adversely impact coccolithophorids and diazotrophs in the future.It may be necessary...

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Coccolithophores are one of the most abundant eukaryotic phytoplankton in the oceans and are distinguished by their ability to build calcitic platelets (coccoliths). Of the numerous species, Emiliania huxleyi is considered one of the major calcifiers in the pelagic ocean. There is growing concern that increasing levels of CO 2 in the atmosphere and the subsequent acidification of the ocean may disrupt the production of coccoliths. Furthermore, any change in the global distribution and abundance of E. huxleyi relative to non-calcifying groups of phytoplankton (e.g. diatoms) will have important effects on the biogeochemical cycling of carbon and climatic feedbacks. We review different lines of evidence that suggest E. huxleyi is increasingly expanding its range into the polar oceans. These observations contribute to the debate on the climatic effects on natural coccolithophore populations. We postulate that E. huxleyi may be more sensitive to recent environmental changes such as increasing sea surface temperature and salinity than to changing ocean carbonate chemistry, partly because increased availability of CO 2 (aq) likely alleviates a carbon limitation for the inefficient Rubisco enzyme in these algae. Any potentially important climatic feedbacks of coccolithophores need a better knowledge of the mechanisms and rates of