Latitudinal phytoplankton distribution and the neutral theory of biodiversity

Aim Recent studies have suggested that global diatom distributions are not limited by dispersal, in the case of both extant species and fossil species, but rather that environmental filtering explains their spatial patterns. Hubbell's neutral theory of biodiversity provides a framework in which to test these alternatives. Our aim is to test whether the structure of marine phytoplankton (diatoms, dinoflagellates and coccolithophores) assemblages across the Atlantic agrees with neutral theory predictions. We asked: (1) whether intersite variance in phytoplankton diversity is explained predominantly by dispersal limitation or by environmental conditions; and (2) whether species abundance distributions are consistent with those expected by the neutral model. Location Meridional transect of the Atlantic (50 degrees N50 degrees S). Methods We estimated the relative contributions of environmental factors and geographic distance to phytoplankton composition using similarity matrices, Mantel tests and variation partitioning of the species composition based upon canonical ordination methods. We compared the species abundance distribution of phytoplankton with the neutral model using Etienne's maximum-likelihood inference method. Results Phytoplankton communities are slightly more determined by niche segregation (24%), than by dispersal limitation and ecological drift (17%). In 60% of communities, the assumption of neutrality in species' abundance distributions could not be rejected. In tropical zones, where oceanic gyres enclose large stable water masses, most communities showed low species immigration rates; in contrast, we infer that communities in temperate areas, out of oligotrophic gyres, have higher rates of species immigration. Conclusions Phytoplankton community structure is consistent with partial niche assembly and partial dispersal and drift assembly (neutral processes). The role of dispersal limitation is almost as important as habitat filtering, a fact that has been largely overlooked in previous studies. Furthermore, the polewards increase in immigration rates of species that we have discovered is probably caused by water mixing conditions and productivity.

Long-term responses of North Atlantic calcifying plankton to climate change

The global increase in atmospheric carbon dioxide concentration is potentially threatening marine biodiversity in two ways. First, carbon dioxide and other greenhouse gases accumulating in the atmosphere are causing global warming1. Second, carbon dioxide is altering sea water chemistry, making the ocean more acidic2. Although temperature has a cardinal influence on all biological processes from the molecular to the ecosystem level3, acidification might impair the process of calcification or exacerbate dissolution of calcifying organisms4. Here, we show however that North Atlantic calcifying plankton primarily responded to climate-induced changes in temperatures during the period 1960–2009, overriding the signal from the effects of ocean acidification. We provide evidence that foraminifers, coccolithophores, both pteropod and nonpteropod molluscs and echinoderms exhibited an abrupt shift circa 1996 at a time of a substantial increase in temperature5 and that some taxa exhibited a poleward movement in agreement with expected biogeographical changes under sea temperature warming6,7. Although acidification may become a serious threat to marine calcifying organisms, our results suggest that over the study period the primary driver of North Atlantic calcifying planktonwas oceanic temperature.

Identifying four phytoplankton functional types from space: An ecological approach

Deriving maps of phytoplankton taxa based on remote sensing data using bio-optical properties of phytoplankton alone is challenging. A more holistic approach was developed using artificial neural networks, incorporating ecological and geographical knowledge together with ocean color, bio-optical characteristics, and remotely sensed physical parameters. Results show that the combined remote sensing approach could discriminate four major phytoplankton functional types (diatoms, dinoflagellates, coccolithophores, and silicoflagellates) with an accuracy of more than 70%. Models indicate that the most important information for phytoplankton functional type discrimination is spatio-temporal information and sea surface temperature. This approach can supply data for large-scale maps of predicted phytoplankton functional types, and an example is shown.

Development and validation of a molecular technique for the analysis of archived formalin-preserved phytoplankton samples permits retrospective assessment of Emiliania huxleyi communities

The coccolithophores, particularly the species Emiliania huxleyi (Lohmann) Hay & Mohler, account for the bulk of global calcium carbonate production and as such play a fundamental role in global CO2 cycling and the carbonate chemistry of the oceans. To evaluate the response of this functional group to the effects of climate change, we undertook a feasibility study to determine whether a retrospective approach could be used on archived coccolithophore datasets. We demonstrate for the first time a technique for the extraction of E. huxleyi nucleic acids from archived formalin-fixed samples of the long-term Continuous Plankton Recorder. Molecular analysis of a nine year old formalin-fixed sample reveals the presence of a diverse population of E. huxleyi genotypes within a developing coccolithophore bloom. In addition, E. huxleyi sequences were amplified from a number of formalin-fixed samples, the earliest of which was collected in August 1972. This molecular assay promises the possibility of studying global variations in the distribution and genetic make-up of E. huxleyi communities over extensive periods of time. (c) 2008 Elsevier B.V. All rights reserved.

Coccolithophore bloom detection in the north east Atlantic using SeaWiFS: Algorithm description, application and sensitivity analysis

Coccolithophores are the largest source of calcium carbonate in the oceans and are considered to play an important role in oceanic carbon cycles. Current methods to detect the presence of coccolithophore blooms from Earth observation data often produce high numbers of false positives in shelf seas and coastal zones due to the spectral similarity between coccolithophores and other suspended particulates. Current methods are therefore unable to characterise the bloom events in shelf seas and coastal zones, despite the importance of these phytoplankton in the global carbon cycle. A novel approach to detect the presence of coccolithophore blooms from Earth observation data is presented. The method builds upon previous optical work and uses a statistical framework to combine spectral, spatial and temporal information to produce maps of coccolithophore bloom extent. Validation and verification results for an area of the north east Atlantic are presented using an in situ database (N = 432) and all available SeaWiFS data for 2003 and 2004. Verification results show that the approach produces a temporal seasonal signal consistent with biological studies of these phytoplankton. Validation using the in situ coccolithophore cell count database shows a high correct recognition rate of 80% and a low false-positive rate of 0.14 (in comparison to 63% and 0.34 respectively for the established, purely spectral approach). To guide its broader use, a full sensitivity analysis for the algorithm parameters is presented.

Coccolithophore surface distributions in the North Atlantic and their modulation of the air-sea flux of CO2 from 10 years of satellite Earth observation data

Coccolithophores are the primary oceanic phytoplankton responsible for the production of calcium carbonate (CaCO3). These climatically important plankton play a key role in the oceanic carbon cycle as a major contributor of carbon to the open ocean 5 carbonate pump (�50%) and their formation can affect the atmosphere-to-ocean (airsea) uptake of carbon dioxide (CO2) through increasing the seawater partial pressure of CO2 (pCO2). Here we document variations in the areal extent of surface blooms of the globally important coccolithophore, Emiliania huxleyi, in the North Atlantic over a 10-year period (1998–2007), using Earth observation data from the Sea-viewing Wide 10 Field of view Sensor (SeaWiFS).We calculate the annual mean surface areal coverage of E. huxleyi in the North Atlantic to be 474 000±119 000km2 yr−1, which results in a net CaCO3 production of 0.62±0.15 Tg CaCO3 carbon per year. However, this surface coverage and net production can fluctuate by −54/+81% about these mean values and are strongly correlated with the El Ni˜no/Southern Oscillation (ENSO) climate os15 cillation index (r =0.75, p<0.02). Our analysis evaluates the spatial extent over which the E. huxleyi blooms in the North Atlantic can increase the pCO2 and thus decrease the localised sink of atmospheric CO2. In regions where the blooms are prevalent, the average reduction in the monthly CO2 sink can reach 12 %. The maximum reduction of the monthly CO2 sink in the time series is 32 %. This work suggests that the high 20 variability, frequency and distribution of these calcifying plankton and their impact on pCO2 should be considered within modelling studies of the North Atlantic if we are to fully understand the variability of its air-to-sea CO2 flux.

Exploiting satellite earth observation to quantify current global oceanic DMS flux and its future climate sensitivity

We used coincident Envisat RA2 and AATSR temperature and wind speed data from 2008/2009 to calculate the global net sea-air flux of dimethyl sulfide (DMS), which we estimate to be 19.6 Tg S a21. Our monthly flux calculations are compared to open ocean eddy correlation measurements of DMS flux from 10 recent cruises, with a root mean square difference of 3.1 lmol m22 day21. In a sensitivity analysis, we varied temperature, salinity, surface wind speed, and aqueous DMS concentration, using fixed global changes as well as CMIP5 model output. The range of DMS flux in future climate scenarios is discussed. The CMIP5 model predicts a reduction in surface wind speed and we estimate that this will decrease the global annual sea-air flux of DMS by 22% over 25 years. Concurrent changes in temperature, salinity, and DMS concentration increase the global flux by much smaller amounts. The net effect of all CMIP5 modelled 25 year predictions was a 19% reduction in global DMS flux. 25 year DMS concentration changes had significant regional effects, some positive (Southern Ocean, North Atlantic, Northwest Pacific) and some negative (isolated regions along the Equator and in the Indian Ocean). Using satellite-detected coverage of coccolithophore blooms, our estimate of their contribution to North Atlantic DMS emissions suggests that the coccolithophores contribute only a small percentage of the North Atlantic annual flux estimate, but may be more important in the summertime and in the northeast Atlantic.

Nanoplankton and picoplankton in the Western English Channel: abundance and seasonality from 2007–2013

The nano- and picoplankton community at Station L4 in the Western English Channel was studied between 2007 and 2013 by flow cytometry to quantify abundance and investigate seasonal cycles within these communities. Nanoplankton included both photosynthetic and heterotrophic eukaryotic single-celled organisms while the picoplankton included picoeukaryote phytoplankton, Synechococcus sp. cyanobacteria and heterotrophic bacteria. A Box–Jenkins Transfer Function climatology analysis of surface data revealed that Synechococcus sp., cryptophytes, and heterotrophic flagellates had bimodal annual cycles. Nanoeukaryotes and both high and low nucleic acid-containing bacteria (HNA and LNA, respectively) groups exhibited unimodal annual cycles. Phaeocystis sp., whilst having clearly defined abundance maxima in spring was not detectable the rest of the year. Coccolithophores exhibited a weak seasonal cycle, with abundance peaks in spring and autumn. Picoeukaryotes did not exhibit a discernable seasonal cycle at the surface. Timings of maximum group abundance varied through the year. Phaeocystis sp. and heterotrophic flagellates peaked in April/May. Nanoeukaryotes and HNA bacteria peaked in June/July and had relatively high abundance throughout the summer. Synechococcus sp., cryptophytes and LNA bacteria all peaked from mid to late September. The transfer function model techniques used represent a useful means of identifying repeating annual cycles in time series data with the added ability to detect trends and harmonic terms at different time scales from months to decades.

Feeding selectivity of bivalve larvae on natural plankton assemblages in the Western English Channel

Meroplankton, including bivalve larvae, are an important and yet understudied component of coastal marine food webs. Understanding the baseline of meroplankton ecology is imperative to establish and predict their sensitivity to local and global marine stressors. Over an annual cycle (October 2009–September 2010), bivalve larvae were collected from the Western Channel Observatory time series station L4 (50°15.00′N, 4°13.02′W). The morphologically similar larvae were identified by analysis of the 18S nuclear small subunit ribosomal RNA gene, and a series of incubation experiments were conducted to determine larval ingestion rates on natural plankton assemblages. Complementary gut content analysis was performed using a PCR-based method for detecting prey DNA both from field-collected larvae and those from the feeding experiments. Molecular identification of bivalve larvae showed the community composition to change over the course of the sampling period with domination by Phaxas in winter and higher diversity in autumn. The larvae selected for nanoeukaryotes (2–20 µm) including coccolithophores (<20 µm) which together comprised >75 % of the bivalve larvae diet. Additionally, a small percentage of carbon ingested originated from heterotrophic ciliates (<30 µm). The molecular analysis of bivalve larvae gut content provided increased resolution of identification of prey consumed and demonstrated that the composition of prey consumed established through bottle incubations conferred with that established from in situ larvae. Despite changes in bivalve larvae community structure, clearance rates of each prey type did not change significantly over the course of the experiment, suggesting different bivalve larvae species may consume similar prey.

Coccolithophore biomineralization: New questions, new answers.

Coccolithophores are unicellular phytoplankton that are characterized by the presence intricately formed calcite scales (coccoliths) on their surfaces. In most cases coccolith formation is an entirely intracellular process - crystal growth is confined within a Golgi-derived vesicle. A wide range of coccolith morphologies can be found amongst the different coccolithophore groups. This review discusses the cellular factors that regulate coccolith production, from the roles of organic components, endomembrane organization and cytoskeleton to the mechanisms of delivery of substrates to the calcifying compartment. New findings are also providing important information on how the delivery of substrates to the calcification site is co-ordinated with the removal of H(+) that are a bi-product of the calcification reaction. While there appear to be a number of species-specific features of the structural and biochemical components underlying coccolith formation, the fluxes of Ca(2+) and a HCO3(-) required to support coccolith formation appear to involve spatially organized recruitment of conserved transport processes.

Coccolithophore biomineralization: New questions, new answers.

Coccolithophores are unicellular phytoplankton that are characterized by the presence intricately formed calcite scales (coccoliths) on their surfaces. In most cases coccolith formation is an entirely intracellular process - crystal growth is confined within a Golgi-derived vesicle. A wide range of coccolith morphologies can be found amongst the different coccolithophore groups. This review discusses the cellular factors that regulate coccolith production, from the roles of organic components, endomembrane organization and cytoskeleton to the mechanisms of delivery of substrates to the calcifying compartment. New findings are also providing important information on how the delivery of substrates to the calcification site is co-ordinated with the removal of H(+) that are a bi-product of the calcification reaction. While there appear to be a number of species-specific features of the structural and biochemical components underlying coccolith formation, the fluxes of Ca(2+) and a HCO3(-) required to support coccolith formation appear to involve spatially organized recruitment of conserved transport processes.

Haptophyta

Haptophyta are predominantly planktonic and phototrophic organisms that have their main distribution in marine environments worldwide. They are a major component of the microbial ecosystem, some form massive blooms and some are toxic. Haptophytes are significant players in the global carbonate cycle through photosynthesis and calcification. They are characterized by the haptonema, a third appendage used for attachment and food handling, two similar flagella, two golden-brown chloroplasts, and organic body scales that serve in species identification. Coccolithophores have calcified scales termed coccoliths. Phylogenetically Haptophyta form a well-defined group and are divided into two classes Pavlovophyceae and Coccolithophyceae (Prymnesiophyceae). Currently, about 330 species are described. Environmental DNA sequencing shows high haptophyte diversity in the marine pico- and nanoplankton, of which many likely represent novel species and lineages. Haptophyte diversity is believed to have peaked in the past and their presence is documented in the fossil record back to the Triassic, approximately 225 million years ago. Some biomolecules of haptophyte origin are extraordinarily resistant to decay and are thus used by geologists as sedimentary proxies of past climatic conditions.

Haptophyta

Haptophyta are predominantly planktonic and phototrophic organisms that have their main distribution in marine environments worldwide. They are a major component of the microbial ecosystem, some form massive blooms and some are toxic. Haptophytes are significant players in the global carbonate cycle through photosynthesis and calcification. They are characterized by the haptonema, a third appendage used for attachment and food handling, two similar flagella, two golden-brown chloroplasts, and organic body scales that serve in species identification. Coccolithophores have calcified scales termed coccoliths. Phylogenetically Haptophyta form a well-defined group and are divided into two classes Pavlovophyceae and Coccolithophyceae (Prymnesiophyceae). Currently, about 330 species are described. Environmental DNA sequencing shows high haptophyte diversity in the marine pico- and nanoplankton, of which many likely represent novel species and lineages. Haptophyte diversity is believed to have peaked in the past and their presence is documented in the fossil record back to the Triassic, approximately 225 million years ago. Some biomolecules of haptophyte origin are extraordinarily resistant to decay and are thus used by geologists as sedimentary proxies of past climatic conditions.

Why marine phytoplankton calcify

Calcifying marine phytoplankton - coccolithophores - are some of the most successful yet enigmatic organisms in the ocean, and are at risk from global change. In order to better understand how they will be affected we need to know 'why' coccolithophores calcify. Here we review coccolithophorid evolutionary history, cell biology, and insights from recent experiments to provide a critical assessment of the costs and benefits of calcification. We conclude that calcification has high energy demands, and that coccolithophores might have calcified initially to reduce grazing pressure, but that additional benefits such as protection from photo-damage and viral-bacterial attack further explain their high diversity and broad spectrum ecology. The cost-versus-benefit of these traits is illustrated by novel ecosystem modeling, although conclusive observations are still limited. In the future ocean, the trade-off between changing ecological and physiological costs of calcification and their benefits will ultimately decide how this important group is affected by ocean acidification and global warming.

Why marine phytoplankton calcify

Calcifying marine phytoplankton - coccolithophores - are some of the most successful yet enigmatic organisms in the ocean, and are at risk from global change. In order to better understand how they will be affected we need to know 'why' coccolithophores calcify. Here we review coccolithophorid evolutionary history, cell biology, and insights from recent experiments to provide a critical assessment of the costs and benefits of calcification. We conclude that calcification has high energy demands, and that coccolithophores might have calcified initially to reduce grazing pressure, but that additional benefits such as protection from photo-damage and viral-bacterial attack further explain their high diversity and broad spectrum ecology. The cost-versus-benefit of these traits is illustrated by novel ecosystem modeling, although conclusive observations are still limited. In the future ocean, the trade-off between changing ecological and physiological costs of calcification and their benefits will ultimately decide how this important group is affected by ocean acidification and global warming.