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<title>Spring bloom</title>
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<p>The <b>spring bloom</b> is a strong increase in <a href="Phytoplankton" title="Phytoplankton">phytoplankton</a> abundance (i.e. stock) that typically occurs in the early spring and lasts until late spring or early summer. This seasonal event is characteristic of temperate North Atlantic, sub-polar, and coastal waters.<sup id="cite_ref-Mann_1-0" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-0" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Phytoplankton blooms occur when growth exceeds losses, however there is no universally accepted definition of the magnitude of change or the threshold of abundance that constitutes a bloom. The magnitude, spatial extent and duration of a bloom depends on a variety of abiotic and biotic factors. Abiotic factors include light availability, nutrients, temperature, and physical processes that influence light availability,<sup id="cite_ref-Mann_1-1" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-1" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Winder_3-0" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Oviatt_4-0" class="reference"><a href="#cite_note-Oviatt-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smayda_5-0" class="reference"><a href="#cite_note-Smayda-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> and biotic factors include <a href="Grazing" title="Grazing">grazing</a>, viral <a href="Lysis" title="Lysis">lysis</a>, and phytoplankton physiology.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The factors that lead to bloom initiation are still actively debated (see <a href="Critical_depth" title="Critical depth">Critical depth</a>).
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<div class="mw-heading mw-heading2"><h2 id="Classical_mechanism">Classical mechanism</h2></div>
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</style><table class="sidebar sidebar-collapse nomobile nowraplinks"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#82C3D8; padding:0.2em; font-size:160%; font-weight:bold;"><a href="Plankton" title="Plankton">Plankton</a></th></tr><tr><td class="sidebar-image"><span typeof="mw:File"></span></td></tr><tr><td class="sidebar-content">
<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">By habitat</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist">
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<ul><li><a href="Marine_plankton" class="mw-redirect" title="Marine plankton">Marine plankton</a>
<ul><li><a href="Marine_prokaryotes" title="Marine prokaryotes">prokaryotes</a></li>
<li><a href="Marine_protists" title="Marine protists">protists</a></li></ul></li></ul>
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<ul><li><a href="Freshwater_plankton" class="mw-redirect" title="Freshwater plankton">Freshwater plankton</a></li></ul>
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<ul><li><a href="Calcareous_nannofossils" title="Calcareous nannofossils">calcareous</a></li></ul></li>
<li><a href="Photosynthetic_picoplankton" title="Photosynthetic picoplankton">Photosynthetic picoplankton</a></li>
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<li>Decomposition
<ul><li><a href="Microbial_loop" title="Microbial loop">microbial loop</a></li>
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<ul><li><a href="Algal_bloom" title="Algal bloom">Algal bloom</a></li>
<li><a href="Critical_depth" title="Critical depth">Critical depth</a></li>
<li><a href="Cyanobacterial_bloom" class="mw-redirect" title="Cyanobacterial bloom">Cyanobacterial bloom</a></li>
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<li><a href="Eutrophication" title="Eutrophication">Eutrophication</a></li>
<li><a href="Great_Atlantic_Sargassum_Belt" title="Great Atlantic Sargassum Belt">Great Atlantic Sargassum Belt</a></li>
<li><a href="Great_Calcite_Belt" title="Great Calcite Belt">Great Calcite Belt</a></li>
<li><a href="Milky_seas_effect" title="Milky seas effect">Milky seas effect</a></li></ul>
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<ul><li><a href="Algaculture" title="Algaculture">Algaculture</a></li>
<li><a href="CLAW_hypothesis" title="CLAW hypothesis">CLAW hypothesis</a></li>
<li><a href="Continuous_Plankton_Recorder" title="Continuous Plankton Recorder">CPR</a></li>
<li><a href="Diel_vertical_migration" title="Diel vertical migration">Diel vertical migration</a></li>
<li><a href="F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li>
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<ul><li><a href="Iron_fertilization" title="Iron fertilization">iron</a></li></ul></li>
<li><a href="Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li>
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<li><a href="North_Atlantic_Aerosols_and_Marine_Ecosystems_Study" title="North Atlantic Aerosols and Marine Ecosystems Study">NAAMES</a></li></ul>
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<p>In the spring, more light becomes available and stratification of the water column occurs as increasing temperatures warm the surface waters (referred to as thermal stratification). As a result, vertical mixing is inhibited and phytoplankton and nutrients are entrained in the <a href="Euphotic_zone" class="mw-redirect" title="Euphotic zone">euphotic zone</a>.<sup id="cite_ref-Mann_1-2" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-2" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> This creates a comparatively high nutrient and high light environment that allows rapid phytoplankton growth.<sup id="cite_ref-Mann_1-3" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-3" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hunt_7-0" class="reference"><a href="#cite_note-Hunt-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
</p><p>Along with thermal stratification, spring blooms can be triggered by salinity stratification due to freshwater input, from sources such as high river runoff. This type of stratification is normally limited to coastal areas and estuaries, including Chesapeake Bay.<sup id="cite_ref-Harding_8-0" class="reference"><a href="#cite_note-Harding-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Freshwater influences primary productivity in two ways. First, because freshwater is less dense, it rests on top of seawater and creates a stratified water column.<sup id="cite_ref-Mann_1-4" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Second, freshwater often carries nutrients <sup id="cite_ref-Winder_3-1" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> that phytoplankton need to carry out processes, including photosynthesis.
</p><p>Rapid increases in phytoplankton growth, that typically occur during the spring bloom, arise because phytoplankton can reproduce rapidly under optimal growth conditions (i.e., high nutrient levels, ideal light and temperature, and minimal losses from grazing and vertical mixing). In terms of reproduction, many species of phytoplankton can double at least once per day, allowing for exponential increases in phytoplankton stock size. For example, the stock size of a population that doubles once per day will increase 1000-fold in just 10 days.<sup id="cite_ref-Miller_2-4" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In addition, there is a lag in the grazing response of herbivorous <a href="Zooplankton" title="Zooplankton">zooplankton</a> at the start of blooms, which minimize phytoplankton losses. This lag occurs because there is low winter zooplankton abundance and many zooplankton, such as <a href="Copepod" title="Copepod">copepods</a>, have longer generation times than phytoplankton.<sup id="cite_ref-Miller_2-5" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p><p>Spring blooms typically last until late spring or early summer, at which time the bloom collapses due to nutrient depletion in the stratified water column and increased grazing pressure by zooplankton.<sup id="cite_ref-Mann_1-5" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-6" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Winder_3-2" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smayda_5-1" class="reference"><a href="#cite_note-Smayda-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The most limiting nutrient in the marine environment is typically <a href="Nitrogen" title="Nitrogen">nitrogen</a> (N). This is because most organisms are unable to fix atmospheric nitrogen into usable forms (i.e. <a href="Ammonium" title="Ammonium">ammonium</a>, <a href="Nitrite" title="Nitrite">nitrite</a>, or <a href="Nitrate" title="Nitrate">nitrate</a>). However, with the exception of coastal waters, it can be argued, that <a href="Iron" title="Iron">iron</a> (Fe) is the most limiting nutrient because it is required to fix nitrogen, but is only available in small quantities in the marine environment, coming from dust storms and leaching from rocks.<sup id="cite_ref-Miller_2-7" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> <a href="Phosphorus" title="Phosphorus">Phosphorus</a> can also be limiting, particularly in freshwater environments and tropical coastal regions.<sup id="cite_ref-Miller_2-8" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p><p>During winter, wind-driven <a href="Turbulence" title="Turbulence">turbulence</a> and cooling water temperatures break down the stratified <a href="Water_column" title="Water column">water column</a> formed during the summer. This breakdown allows vertical mixing of the water column and replenishes nutrients from deep water to the surface waters and the rest of the <a href="Euphotic_zone" class="mw-redirect" title="Euphotic zone">euphotic zone</a>. However, vertical mixing also causes high losses, as phytoplankton are carried below the euphotic zone (so their <a href="Cellular_respiration" title="Cellular respiration">respiration</a> exceeds primary production). In addition, reduced illumination (intensity and daily duration) during winter limits growth rates.
</p>
<div class="mw-heading mw-heading2"><h2 id="Alternative_mechanisms">Alternative mechanisms</h2></div>
<p>Historically, blooms have been explained by Sverdrup's <a href="Critical_depth" title="Critical depth">critical depth</a> hypothesis, which says blooms are caused by shoaling of the mixed layer. Similarly, Winder and Cloern (2010) described spring blooms as a response to increasing temperature and light availability.<sup id="cite_ref-Winder_3-3" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> However, new explanations have been offered recently, including that blooms occur due to:
</p>
<ul><li>Coupling between phytoplankton growth and zooplankton grazing.<sup id="cite_ref-Behrenfeld_9-0" class="reference"><a href="#cite_note-Behrenfeld-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></li>
<li>The onset of near surface stratification in the spring.<sup id="cite_ref-Chiswell_10-0" class="reference"><a href="#cite_note-Chiswell-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></li>
<li>Mixing of the water column, rather than stratification<sup id="cite_ref-Townsend_11-0" class="reference"><a href="#cite_note-Townsend-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup></li>
<li>Low turbulence<sup id="cite_ref-Huisman_12-0" class="reference"><a href="#cite_note-Huisman-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup></li>
<li>Increasing light intensity (in shallow water environments).<sup id="cite_ref-Miller_2-9" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></li>
<li>Eddies (see ‘The role of eddies in the onset of the North Atlantic spring bloom’)<sup id="cite_ref-Mahadevan_13-0" class="reference"><a href="#cite_note-Mahadevan-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup></li></ul>
<div class="mw-heading mw-heading2"><h2 id="The_role_of_eddies_in_the_onset_of_the_North_Atlantic_spring_bloom">The role of eddies in the onset of the North Atlantic spring bloom</h2></div>
<p>A 2012 study showed that the onset of the North Atlantic bloom is due to eddies. Eddies, or circular currents of water, are ubiquitous throughout the world’s ocean and play an important role in ocean mixing.<sup id="cite_ref-Currents_14-0" class="reference"><a href="#cite_note-Currents-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> In the North Atlantic, surface water is colder and denser farther north and warmer and lighter in the south. This sets up a horizontal density gradient. Earth’s rotation maintains this gradient by preventing the dense water from slipping underneath the light water. Eddies, however, can mix dense water underneath the lighter water, setting up a vertical stratification that limits the depth of vertical mixing (leading to a shallower mixed layer).<sup id="cite_ref-Fox-Kemper_15-0" class="reference"><a href="#cite_note-Fox-Kemper-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>
</p><p>Mechanisms that limit the depth of vertical mixing can be referred to as ‘restratifying mechanisms’ (e.g. eddies, solar heating), which compete against mechanisms that increase vertical mixing (and deepen the mixed layer). This includes convection and down-front winds. Convection is strongest in the winter when surface cooling is strongest. Convection increases the depth of vertical mixing, which can move phytoplankton away from the light they need to grow.<sup id="cite_ref-Taylor_16-0" class="reference"><a href="#cite_note-Taylor-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup>
</p><p>When convection weakens and wind switches direction in the spring, the re-stratifying effect of eddies becomes dominant. Phytoplankton are trapped closer to the surface, increasing their exposure to light. This spurs phytoplankton growth, leading to the onset of the North Atlantic spring bloom 20-30 days earlier than would occur with thermal stratification alone.<sup id="cite_ref-Mahadevan_13-1" class="reference"><a href="#cite_note-Mahadevan-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Northward_progression">Northward progression</h2></div>
<p>At greater <a href="Latitudes" class="mw-redirect" title="Latitudes">latitudes</a>, spring blooms take place later in the year. This northward progression is because spring occurs later, delaying thermal stratification and increases in illumination that promote blooms. A study by Wolf and Woods (1988) showed evidence that spring blooms follow the northward migration of the 12 °C isotherm, suggesting that blooms may be controlled by temperature limitations, in addition to stratification.<sup id="cite_ref-Mann_1-6" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p><p>At high latitudes, the shorter warm season commonly results in one mid-summer bloom. These blooms tend to be more intense than spring blooms of temperate areas because there is a longer duration of daylight for photosynthesis to take place. Also, grazing pressure tends to be lower because the generally cooler temperatures at higher latitudes slow zooplankton metabolism.<sup id="cite_ref-Mann_1-7" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Species_succession">Species succession</h2></div>
<p>The spring bloom often consists of a series of sequential blooms of different phytoplankton species. Succession occurs because different species have optimal nutrient uptake at different ambient concentrations and reach their growth peaks at different times. Shifts in the dominant phytoplankton species are likely caused by biological and physical (i.e. environmental) factors.<sup id="cite_ref-Miller_2-10" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> For instance, <a href="Diatom" title="Diatom">diatom</a> growth rate becomes limited when the supply of <a href="Silicate" title="Silicate">silicate</a> is depleted.<sup id="cite_ref-Mann_1-8" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-11" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kristiansen_17-0" class="reference"><a href="#cite_note-Kristiansen-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Since silicate is not required by other phytoplankton, such as <a href="Dinoflagellate" title="Dinoflagellate">dinoflagellates</a>, their growth rates continue to increase.
</p><p>For example, in oceanic environments, diatoms (cells diameter greater than 10 to 70 μm or larger) typically dominate first because they are capable of growing faster. Once silicate is depleted in the environment, diatoms are succeeded by smaller dinoflagellates.<sup id="cite_ref-Mann_1-9" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-12" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kristiansen_17-1" class="reference"><a href="#cite_note-Kristiansen-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> This scenario has been observed in Rhode Island,<sup id="cite_ref-Smayda1957_18-0" class="reference"><a href="#cite_note-Smayda1957-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Nixon_19-0" class="reference"><a href="#cite_note-Nixon-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pratt_20-0" class="reference"><a href="#cite_note-Pratt-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> as well as Massachusetts and Cape Cod Bay.<sup id="cite_ref-Hunt_7-1" class="reference"><a href="#cite_note-Hunt-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> By the end of a spring bloom, when most nutrients have been depleted, the majority of the total phytoplankton <a href="Biomass_(ecology)" title="Biomass (ecology)">biomass</a> is very small phytoplankton, known as ultraphytoplankton (cell diameter <5 to 10 μm).<sup id="cite_ref-Miller_2-13" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Ultraphytoplankton can sustain low, but constant stocks, in nutrient depleted environments because they have a larger <a href="Surface_area_to_volume_ratio" class="mw-redirect" title="Surface area to volume ratio">surface area to volume ratio</a>, which offers a much more effective rate of <a href="Diffusion" title="Diffusion">diffusion</a>.<sup id="cite_ref-Mann_1-10" class="reference"><a href="#cite_note-Mann-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller_2-14" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The types of phytoplankton comprising a bloom can be determined by examination of the varying photosynthetic pigments found in <a href="Chloroplast" title="Chloroplast">chloroplasts</a> of each species.<sup id="cite_ref-Miller_2-15" class="reference"><a href="#cite_note-Miller-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Variability_and_the_influence_of_climate_change">Variability and the influence of climate change</h2></div>
<p>Variability in the patterns (e.g., timing of onset, duration, magnitude, position, and spatial extent) of annual spring bloom events has been well documented.<sup id="cite_ref-Winder_3-4" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smayda_5-2" class="reference"><a href="#cite_note-Smayda-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> These variations occur due to fluctuations in environmental conditions, such as wind intensity, temperature, freshwater input, and light. Consequently, spring bloom patterns are likely sensitive to global <a href="Climate_change" title="Climate change">climate change</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup>
</p><p>Links have been found between temperature and spring bloom patterns. For example, several studies have reported a correlation between earlier spring bloom onset and temperature increases over time.<sup id="cite_ref-Winder_3-5" class="reference"><a href="#cite_note-Winder-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Furthermore, in Long Island Sound and the Gulf of Maine, blooms begin later in the year, are more productive, and last longer during colder years, while years that are warmer exhibit earlier, shorter blooms of greater magnitude.<sup id="cite_ref-Smayda_5-3" class="reference"><a href="#cite_note-Smayda-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
</p><p>Temperature may also regulate bloom sizes. In Narragansett Bay, Rhode Island, a study by Durbin et al. (1992)<sup id="cite_ref-Durbin_22-0" class="reference"><a href="#cite_note-Durbin-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> indicated that a 2 °C increase in water temperature resulted in a three-week shift in the maturation of the copepod, <i><a href="Acartia_hudsonica" title="Acartia hudsonica">Acartia hudsonica</a></i>, which could significantly increase zooplankton grazing intensity. Oviatt et al. (2002)<sup id="cite_ref-Oviatt_4-1" class="reference"><a href="#cite_note-Oviatt-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> noted a reduction in spring bloom intensity and duration in years when winter water temperatures were warmer. Oviatt et al. suggested that the reduction was due to increased grazing pressure, which could potentially become intense enough to prevent spring blooms from occurring altogether.
</p><p>Miller and Harding (2007)<sup id="cite_ref-MillerWD_23-0" class="reference"><a href="#cite_note-MillerWD-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> suggested climate change (influencing winter weather patterns and freshwater influxes) was responsible for shifts in spring bloom patterns in the Chesapeake Bay. They found that during warm, wet years (as opposed to cool, dry years), the spatial extent of blooms was larger and was positioned more seaward. Also, during these same years, biomass was higher and peak biomass occurred later in the spring.
</p>
<div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div>
<ul><li><a href="Algal_bloom" title="Algal bloom">Algal bloom</a></li>
<li><a href="Critical_depth" title="Critical depth">Critical depth</a></li>
<li><a href="Gordon_Arthur_Riley" title="Gordon Arthur Riley">Gordon Arthur Riley</a></li>
<li><a href="Plankton" title="Plankton">Plankton</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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<div class="mw-references-wrap mw-references-columns"><ol class="references">
<li id="cite_note-Mann-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Mann_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Mann_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Mann_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Mann_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Mann_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Mann_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Mann_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Mann_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Mann_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Mann_1-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-Mann_1-10"><sup><i><b>k</b></i></sup></a></span> <span class="reference-text">Mann, K.H., Lazier, J.R.N. (2006). <i>Dynamics of Marine Ecosystems: Biological-Physical Interactions in the Oceans</i>. Oxford: Blackwell Publishing Ltd. <style data-mw-deduplicate="TemplateStyles:r1238218222">
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.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("./mw/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("./mw/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("./mw/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("./mw/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}
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</style><a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>1-4051-1118-6</bdi></span>
</li>
<li id="cite_note-Miller-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Miller_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Miller_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Miller_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Miller_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Miller_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Miller_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Miller_2-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Miller_2-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Miller_2-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Miller_2-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-Miller_2-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-Miller_2-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-Miller_2-12"><sup><i><b>m</b></i></sup></a> <a href="#cite_ref-Miller_2-13"><sup><i><b>n</b></i></sup></a> <a href="#cite_ref-Miller_2-14"><sup><i><b>o</b></i></sup></a> <a href="#cite_ref-Miller_2-15"><sup><i><b>p</b></i></sup></a></span> <span class="reference-text">Miller, C.B. (2004). "Biological Oceanography" Oxford: Blackwell Publishing Ltd. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-632-05536-4</bdi></span>
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<li id="cite_note-Oviatt-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Oviatt_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Oviatt_4-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Oviatt, C., Keller, A., and Reed, L. (2002). "Annual Primary Production in Narragansett Bay with no Bay-Wide Winter–Spring Phytoplankton Bloom". <a href="Estuarine%2C_Coastal_and_Shelf_Science" title="Estuarine, Coastal and Shelf Science">Estuarine, Coastal and Shelf Science</a> 54: 1013–1026. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Fecss.2001.0872">10.1006/ecss.2001.0872</a></span>
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<li id="cite_note-Smayda-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-Smayda_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Smayda_5-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Smayda_5-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Smayda_5-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text">Smayda, T.J. (1998). "Patterns of variability characterizing marine phytoplankton, with examples from Narragansett Bay". ICES Journal of Marine Science 55: 562–573</span>
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<li id="cite_note-Townsend-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-Townsend_11-0">^</a></b></span> <span class="reference-text">Townsend, D.W., Cammen, L.M., Holligan, P.M., Campbell, D.E., Pettigrew, N.R. (1994). "Causes and consequences of variability in the timing of spring phytoplankton blooms". <i>Deep-Sea Research</i> 41: 747–765</span>
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<li id="cite_note-Mahadevan-13"><span class="mw-cite-backlink">^ <a href="#cite_ref-Mahadevan_13-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Mahadevan_13-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Mahadevan, A., D’Asaro, E., Lee, C., & Perry, M. J. (2012). Eddy-driven stratification initiates North Atlantic spring phytoplankton blooms. Science, 337(6090), 54–58. <a rel="nofollow" class="external free" href="https://doi.org/10.1126/science.1218740">https://doi.org/10.1126/science.1218740</a> </span>
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<li id="cite_note-Durbin-22"><span class="mw-cite-backlink"><b><a href="#cite_ref-Durbin_22-0">^</a></b></span> <span class="reference-text">Durbin, A.G. and Durbin, E.G. (1992). "Seasonal changes in size frequency distribution and estimated age in the marine copepod Acartia hudsortica during a winter-spring diatom bloom in Narragansett Bay". Limnol. Oceanogr., 37(2): 379–392</span>
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</ol></div></div>
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</style></div><div role="navigation" class="navbox" aria-labelledby="Plankton118" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Plankton118" style="font-size:114%;margin:0 4em"><a href="Plankton" title="Plankton">Plankton</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">About <a href="Plankton" title="Plankton">plankton</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Algal_bloom" title="Algal bloom">Algal bloom</a></li>
<li><a href="CLAW_hypothesis" title="CLAW hypothesis">CLAW hypothesis</a></li>
<li>High lipid content microalgae</li>
<li><a href="Holoplankton" title="Holoplankton">Holoplankton</a></li>
<li><a href="Marine_microorganisms" title="Marine microorganisms">Marine microorganisms</a></li>
<li><a href="Meroplankton" title="Meroplankton">Meroplankton</a></li>
<li><a href="Mycoplankton" title="Mycoplankton">Mycoplankton</a></li>
<li><a href="Milky_seas_effect" title="Milky seas effect">Milky seas effect</a></li>
<li><a href="Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li>
<li><a href="Planktivore" title="Planktivore">Planktivore</a></li>
<li><a href="Planktology" title="Planktology">Planktology</a></li>
<li><a href="Red_tide" class="mw-redirect" title="Red tide">Red tide</a></li>
<li><a href="Thin_layers_(oceanography)" title="Thin layers (oceanography)">Thin layers</a></li>
<li>More...</li></ul>
</div></td><td class="noviewer navbox-image" rowspan="7" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By size</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="List_of_eukaryotic_picoplankton_species" title="List of eukaryotic picoplankton species">Eukaryotic picoplankton</a></li>
<li><a href="Heterotrophic_picoplankton" title="Heterotrophic picoplankton">Heterotrophic picoplankton</a></li>
<li><a href="Marine_microplankton" class="mw-redirect" title="Marine microplankton">Marine microplankton</a></li>
<li><a href="Microalgae" title="Microalgae">Microphyte (microalgae)</a></li>
<li><a href="Nanophytoplankton" title="Nanophytoplankton">Nanophytoplankton</a></li>
<li><a href="Photosynthetic_picoplankton" title="Photosynthetic picoplankton">Photosynthetic picoplankton</a></li>
<li><a href="Picobiliphyte" class="mw-redirect" title="Picobiliphyte">Picobiliphyte</a></li>
<li><a href="Picoeukaryote" title="Picoeukaryote">Picoeukaryote</a></li>
<li><a href="Picoplankton" title="Picoplankton">Picoplankton</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Bacterioplankton" title="Bacterioplankton">Bacterioplankton</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><i><a href="Aeromonas_salmonicida" title="Aeromonas salmonicida">Aeromonas salmonicida</a></i></li>
<li><a href="Cyanobacteria" title="Cyanobacteria">Cyanobacteria</a></li>
<li><a href="Cyanobiont" title="Cyanobiont">Cyanobiont</a></li>
<li><a href="Cyanotoxin" title="Cyanotoxin">Cyanotoxin</a></li>
<li><a href="Enteric_redmouth_disease" title="Enteric redmouth disease">Enteric redmouth disease</a></li>
<li><i><a href="Flavobacterium" title="Flavobacterium">Flavobacterium</a></i></li>
<li><i><a href="Flavobacterium_columnare" title="Flavobacterium columnare">Flavobacterium columnare</a></i></li>
<li><i><a href="Pelagibacter_ubique" class="mw-redirect" title="Pelagibacter ubique">Pelagibacter ubique</a></i></li>
<li><a href="Marine_bacteriophage" class="mw-redirect" title="Marine bacteriophage">Marine bacteriophage</a></li>
<li><a href="Pelagibacterales" class="mw-redirect" title="Pelagibacterales">SAR11 clade</a></li>
<li><i><a href="Streptococcus_iniae" title="Streptococcus iniae">Streptococcus iniae</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Phytoplankton" title="Phytoplankton">Phytoplankton</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Auxospore" title="Auxospore">Auxospore</a></li>
<li><a href="Axodine" title="Axodine">Axodine</a></li>
<li><i><a href="Bacteriastrum" title="Bacteriastrum">Bacteriastrum</a></i></li>
<li><i><a href="Chaetoceros" title="Chaetoceros">Chaetoceros</a></i></li>
<li><a href="Chaetocerotaceae" title="Chaetocerotaceae">Chaetocerotaceae</a></li>
<li><a href="Coccolithophore" title="Coccolithophore">Coccolithophore</a></li>
<li><i><a href="Emiliania_huxleyi" class="mw-redirect" title="Emiliania huxleyi">Emiliania huxleyi</a></i></li>
<li><a href="Eustigmatophyte" title="Eustigmatophyte">Eustigmatophyte</a></li>
<li><a href="Frustule" title="Frustule">Frustule</a></li>
<li><a href="Stramenopile" title="Stramenopile">Stramenopile</a></li>
<li><i><a href="Nannochloropsis" title="Nannochloropsis">Nannochloropsis</a></i></li>
<li><i><a href="Navicula" title="Navicula">Navicula</a></i></li>
<li><a href="Prasinophyceae" class="mw-redirect" title="Prasinophyceae">Prasinophyceae</a></li>
<li><a href="Raphidophyte" title="Raphidophyte">Raphidophyte</a></li>
<li><i><a href="Thalassiosira_pseudonana" title="Thalassiosira pseudonana">Thalassiosira pseudonana</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Diatom" title="Diatom">Diatom orders</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Coscinodiscophyceae" title="Coscinodiscophyceae">Centrales</a></li>
<li><a href="Pennales" title="Pennales">Pennales</a>
<ul><li>Classes: <a href="Coscinodiscophyceae" title="Coscinodiscophyceae">Coscinodiscophyceae</a></li>
<li><a href="Fragilariophyceae" title="Fragilariophyceae">Fragilariophyceae</a></li>
<li><a href="Diatom" title="Diatom">Bacillariophyceae</a></li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Flagellate" title="Flagellate">Flagellates</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Brevetoxin" title="Brevetoxin">Brevetoxin</a></li>
<li><a href="Choanoflagellate" title="Choanoflagellate">Choanoflagellates</a></li>
<li><a href="Dinoflagellate" title="Dinoflagellate">Dinoflagellates</a></li>
<li><a href="Flagellum" title="Flagellum">Flagellum</a></li>
<li><i><a href="Pfiesteria_piscicida" title="Pfiesteria piscicida">Pfiesteria piscicida</a></i></li>
<li><a href="Saxitoxin" title="Saxitoxin">Saxitoxin</a></li>
<li><i><a href="Symbiodinium" title="Symbiodinium">Symbiodinium</a></i></li>
<li><a href="Velvet_(fish_disease)" title="Velvet (fish disease)">Velvet (fish disease)</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Zooplankton" title="Zooplankton">Zooplankton</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Chaetognatha" title="Chaetognatha">Chaetognatha</a></li>
<li><a href="Ciguatera" class="mw-redirect" title="Ciguatera">Ciguatera</a></li>
<li><a href="Ctenophora" title="Ctenophora">Ctenophora</a></li>
<li><a href="Gelatinous_zooplankton" title="Gelatinous zooplankton">Gelatinous zooplankton</a></li>
<li><a href="Ichthyoplankton" title="Ichthyoplankton">Ichthyoplankton</a></li>
<li><a href="Jellyfish" title="Jellyfish">Jellyfish</a></li>
<li><a href="Marine_larval_ecology" title="Marine larval ecology">Marine larvae</a></li>
<li><a href="Crustacean_larva" title="Crustacean larva">Crustacean larvae</a></li>
<li><a href="Salmon_louse" title="Salmon louse">Salmon louse</a></li>
<li><a href="Sea_louse" title="Sea louse">Sea louse</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Copepod" title="Copepod">Copepod orders</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Calanoida" title="Calanoida">Calanoida</a></li>
<li><a href="Canuelloida" class="mw-redirect" title="Canuelloida">Canuelloida</a></li>
<li><a href="Cyclopoida" title="Cyclopoida">Cyclopoida</a></li>
<li><a href="Gelyelloida" class="mw-redirect" title="Gelyelloida">Gelyelloida</a></li>
<li><a href="Harpacticoida" title="Harpacticoida">Harpacticoida</a></li>
<li><a href="Misophrioida" title="Misophrioida">Misophrioida</a></li>
<li><a href="Monstrilloida" class="mw-redirect" title="Monstrilloida">Monstrilloida</a></li>
<li><a href="Mormonilloida" class="mw-redirect" title="Mormonilloida">Mormonilloida</a></li>
<li><a href="Platycopioida" class="mw-redirect" title="Platycopioida">Platycopioida</a></li>
<li><a href="Siphonostomatoida" title="Siphonostomatoida">Siphonostomatoida</a></li>
<li>More...</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Aeroplankton" title="Aeroplankton">Aeroplankton</a></li>
<li><a href="Algaculture" title="Algaculture">Algaculture</a></li>
<li><a href="Algal_mat" title="Algal mat">Algal mat</a></li>
<li><a href="Algal_nutrient_solution" title="Algal nutrient solution">Algal nutrient solutions</a></li>
<li><a href="Artificial_seawater" title="Artificial seawater">Artificial seawater</a></li>
<li><a href="Autotroph" title="Autotroph">Autotrophs</a></li>
<li><a href="Biological_pump" title="Biological pump">Biological pump</a></li>
<li><a href="Diel_vertical_migration" title="Diel vertical migration">Diel vertical migration</a></li>
<li><a href="Dimethylsulfoniopropionate" title="Dimethylsulfoniopropionate">Dimethylsulfoniopropionate</a></li>
<li><a href="F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li>
<li><a href="Fish_diseases_and_parasites" title="Fish diseases and parasites">Fish diseases and parasites</a></li>
<li><a href="Heterotroph" title="Heterotroph">Heterotroph</a></li>
<li><a href="High-nutrient%2C_low-chlorophyll_regions" title="High-nutrient, low-chlorophyll regions">HNLC</a></li>
<li><a href="Seaweed" title="Seaweed">Macroalgae</a></li>
<li><a href="Manta_trawl" title="Manta trawl">Manta trawl</a></li>
<li><a href="Sea_snot" class="mw-redirect" title="Sea snot">Marine mucilage</a></li>
<li><a href="Microbial_mat" title="Microbial mat">Microbial mat</a></li>
<li><a href="Ocean_acidification" title="Ocean acidification">Ocean acidification</a></li>
<li><a href="Marine_microorganisms" title="Marine microorganisms">Marine microorganisms</a></li>
<li><a href="Marine_primary_production" title="Marine primary production">Marine primary production</a></li>
<li><a href="Pseudoplankton" title="Pseudoplankton">Pseudoplankton</a></li>
<li><a href="Stromatolite" title="Stromatolite">Stromatolite</a></li>
<li><a href="Tychoplankton" title="Tychoplankton">Tychoplankton</a></li>
<li><a href="Zoid" title="Zoid">Zoid</a></li>
<li><a href="Center_for_Microbial_Oceanography%3A_Research_and_Education" title="Center for Microbial Oceanography: Research and Education">C-MORE</a></li>
<li><a href="Continuous_Plankton_Recorder" title="Continuous Plankton Recorder">CPR</a></li>
<li><a href="Australian_Continuous_Plankton_Recorder_Survey" title="Australian Continuous Plankton Recorder Survey">AusCPR</a></li>
<li><a href="MOCNESS" title="MOCNESS">MOCNESS</a></li>
<li><a href="SCAR_Southern_Ocean_Continuous_Plankton_Recorder_Survey" title="SCAR Southern Ocean Continuous Plankton Recorder Survey">SCAR</a></li></ul>
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