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Diversity plays a key role in the adaptive capacity of marine ecosystems to environmental changes. However, modelling the adaptive dynamics of phytoplankton traits remains challenging due to the competitive exclusion of sub-optimal phenotypes and the complexity of evolutionary processes leading to optimal phenotypes. Trait diffusion (TD) is a recently developed approach to sustain diversity in plankton models by introducing mutations, therefore allowing the adaptive evolution of functional traits to occur at ecological timescales. In this study, we present a model called Simulating Plankton Evolution with Adaptive Dynamics (SPEAD) that resolves the eco-evolutionary processes of a multi-trait plankton community. The SPEAD model can be used to evaluate plankton adaptation to environmental changes at different timescales or address ecological issues affected by adaptive evolution. Phytoplankton phenotypes in SPEAD are characterized by two traits, the nitrogen half-saturation constant and optimal temperature, which can mutate at each generation using the TD mechanism. SPEAD does not resolve the different phenotypes as discrete entities, instead computing six aggregate properties: total phytoplankton biomass, the mean value of each trait, trait variances, and the inter-trait covariance of a single population in a continuous trait space. Therefore, SPEAD resolves the dynamics of the population's continuous trait distribution by solving its statistical moments, wherein the variances of trait values represent the diversity of ecotypes. The ecological model is coupled to a vertically resolved (1D) physical environment, and therefore the adaptive dynamics of the simulated phytoplankton population are driven by seasonal variations in vertical mixing, nutrient concentration, water temperature, and solar irradiance. The simulated bulk properties are validated by observations from Bermuda Atlantic Time-series Studies (BATS) in the Sargasso Sea. We find that moderate mutation rates sustain trait diversity at decadal timescales and soften the almost total inter-trait correlation induced by the environment alone, without reducing the annual primary production or promoting permanently maladapted phenotypes, as occur with high mutation rates. As a way to evaluate the performance of the continuous trait approximation, we also compare the solutions of SPEAD to the solutions of a classical discrete entities approach, with both approaches including TD as a mechanism to sustain trait variance. We only find minor discrepancies between the continuous model SPEAD and the discrete model, with the computational cost of SPEAD being lower by 2 orders of magnitude. Therefore, SPEAD should be an ideal eco-evolutionary plankton model to be coupled to a general circulation model (GCM) of the global ocean. [ABSTRACT FROM AUTHOR]

Diversity plays a key role in the adaptive capacities of marine ecosystems to environmental changes. However, modeling phytoplankton trait diversity remains challenging due to the strength of the competitive exclusion of sub-optimal phenotypes. Trait diffusion (TD) is a recently developed approach to sustain diversity in plankton models by allowing the evolution of functional traits at ecological timescales. In this study, we present a model for Simulating Plankton Evolution with Adaptive Dynamics (SPEAD), where phytoplankton phenotypes characterized by two traits, nitrogen half-saturation constant and optimal temperature, can mutate at each generation using the TD mechanism. SPEAD does not resolve the different phenotypes as discrete entities, computing instead six aggregate properties: total phytoplankton biomass, mean value of each trait, trait variances, and inter-trait covariance of a single population in a continuous trait space. Therefore SPEAD resolves the dynamics of the population's continuous trait distribution by solving its statistical moments, where the variances of trait values represent the diversity of ecotypes. The ecological model is coupled to a vertically-resolved (1D) physical environment, and therefore the adaptive dynamics of the simulated phytoplankton population are driven by seasonal variations in vertical mixing, nutrient concentration, water temperature, and solar irradiance. The simulated bulk properties are validated by observations from BATS in the Sargasso Sea. We find that moderate mutation rates sustain trait diversity at decadal timescales and soften the almost total inter-trait correlation induced by the environment alone, without reducing the annual primary production or promoting permanently maladapted phenotypes, as occur with high mutation rates. As a way to evaluate the performance of the continuous-trait approximation, we also compare the solutions of SPEAD to the solutions of a classical discrete entities approach, both approaches including TD as a mechanism to sustain trait variance. We only find minor discrepancies between the continuous model SPEAD and the discrete model, the computational cost of SPEAD being lower by two orders of magnitude. Therefore SPEAD should be an ideal eco-evolutionary plankton model to be coupled to a general circulation model (GCM) at the global ocean. [ABSTRACT FROM AUTHOR]

The effect of inorganic nutrients on planktonic assemblages has traditionally relied on concentrations rather than estimates of nutrient supply. We combined a novel dataset of hydrographic properties, turbulent mixing, nutrient concentration, and picoplankton community composition with the aims of (i) quantifying the role of temperature, light, and nitrate fluxes as factors controlling the distribution of autotrophic and heterotrophic picoplankton subgroups, as determined by flow cytometry, and (ii) describing the ecological niches of the various components of the picoplankton community. Data were collected at 97 stations in the Atlantic Ocean, including tropical and subtropical open-ocean waters, the northwestern Mediterranean Sea, and the Galician coastal upwelling system of the northwest Iberian Peninsula. A generalized additive model (GAM) approach was used to predict depth-integrated biomass of each picoplankton subgroup based on three niche predictors: sea surface temperature, averaged daily surface irradiance, and the transport of nitrate into the euphotic zone, through both diffusion and advection. In addition, niche overlap among different picoplankton subgroups was computed using nonparametric kernel density functions. Temperature and nitrate supply were more relevant than light in predicting the biomass of most picoplankton subgroups, except for Prochlorococcus and lownucleic-acid (LNA) prokaryotes, for which irradiance also played a significant role. Nitrate supply was the only factor that allowed the distinction among the ecological niches of all autotrophic and heterotrophic picoplankton subgroups. Prochlorococcus and LNA prokaryotes were more abundant in warmer waters ( > 20 °C) where the nitrate fluxes were low, whereas Synechococcus and high-nucleic-acid (HNA) prokaryotes prevailed mainly in cooler environments characterized by intermediate or high levels of nitrate supply. Finally, the niche of picoeukaryotes was defined by low temperatures and high nitrate supply. These results support the key role of nitrate supply, as it not only promotes the growth of large phytoplankton, but it also controls the structure of marine picoplankton communities. [ABSTRACT FROM AUTHOR]

The biological production of calcium carbonate (CaCO3), a process termed calcification, is a key term in the marine carbon cycle. A major planktonic group responsible for such pelagic CaCO3 production (CP) is the coccolithophores, single-celled haptophytes that inhabit the euphotic zone of the ocean. Satellite-based estimates of areal CP are limited to surface waters and open-ocean areas, with current algorithms utilising the unique optical properties of the cosmopolitan bloom-forming species Emiliania huxleyi , whereas little understanding of deep-water ecology, optical properties or environmental responses by species other than E. huxleyi is currently available to parameterise algorithms or models. To aid future areal estimations and validate future modelling efforts we have constructed a database of 2765 CP measurements, the majority of which were measured using 12 to 24 h incorporation of radioactive carbon (14C) into acid-labile inorganic carbon (CaCO3). We present data collated from over 30 studies covering the period from 1991 to 2015, sampling the Atlantic, Pacific, Indian, Arctic and Southern oceans. Globally, CP in surface waters (<20 m) ranged from 0.01 to 8398 µ mol C m -3 d -1 (with a geometric mean of 16.1 µ mol C m -3 d -1). An integral value for the upper euphotic zone (herein surface to the depth of 1 % surface irradiance) ranged from <0.1 to 6 mmol C m -2 d -1 (geometric mean 1.19 mmol C m -2 d -1). The full database is available for download from PANGAEA at https://doi.org/10.1594/PANGAEA.888182. [ABSTRACT FROM AUTHOR]

The relative contribution of coccolithophores and diatoms to export production and burial influences the mechanisms of carbon (C) removal from Earth's surface reservoirs. Whereas the supply of phosphorus (P) to the ocean basins controls export production over geological timescales, the availability of dissolved silica (Si) determines the relative contribution of diatoms to export production and increases the efficiency of the ocean's biologically-driven C sequestration. Here we combine analyses of data from the sedimentary record and Earth system model simulations to investigate the coupling of P and Si weathering fluxes over the last 40 million years (My). Although both weathering fluxes increased on average through time, the relative increase of Si weathering over the last 20 My exceeded that of P. Our analysis suggests that the increase in P weathering under a regime of increasingly ocean turbulence allowed full utilization of increased Si supplies and links the ecological success of marine diatoms, and consequential decline of coccolithophores, to changes in the Si / P weathering ratio. These results allow us to envisage that ocean fertilization initiatives would be more effective if they were integrated within the framework of geo-engineering programs aimed at enhancing silicate weathering. [ABSTRACT FROM AUTHOR]