Summer Bloom Research Articles

Phytoplankton blooms in summer and autumn in the northwestern subarctic Pacific detected by the mooring and float systems

Phytoplankton blooms in summer and autumn in the northwestern subarctic Pacific detected by the mooring and float systems

Nutrient dynamics in a eutrophic blackwater urban lake

Iraola ND, Mallin MA, Cahoon LB, Gamble DW, Zamora PB. 2022. Nutrient dynamics in a eutrophic blackwater urban lake. Lake Reserv Manage. 38:28–46. Greenfield Lake is a eutrophic blackwater urban lake in Wilmington, North Carolina, and in 2014 was declared by the state as impaired waters due to excessive chlorophyll a concentrations. Phytoplankton production is strongly nitrogen (N) limited, and the lake supports filamentous green algal blooms in spring and N-fixing cyanobacteria blooms in summer. To inform future lake restoration efforts, we quantified nutrient loads from 5 perennial streams that drain the highly impervious Greenfield Lake watershed to determine their impact on the lake’s eutrophic state. The 5 streams were sampled monthly from 2016 to 2017 during periods of dry weather and again after rain events of at least 1.2 cm to compare stormwater runoff to baseflow conditions. Two streams alone accounted for 76% of inorganic N and 63% of inorganic P surface inflow, whereas 2 other streams, despite high (53–74%) impervious coverage, had low loading. A lake-wide sediment phosphorus survey also revealed P accumulations mirrored streams of high nutrient load, while low P accumulations were in areas draining streams with large wetland and stormwater treatment areas. Drainage area was positively correlated with nitrate concentration and nitrate and ammonium loading. Unconventionally, percent impervious coverage was negatively correlated with nutrient concentrations and loads; we suspect that the presence of a large golf course in the watershed with the lowest impervious cover was a key driving factor. Our work also demonstrates the significance of catchment features: Drainages with large stormwater retention structures and significant natural wetlands in the lower reaches yielded low nutrient loadings.

Species-specific monitoring of Skeletonema blooms in the coastal waters of Ariake Sound, Japan

Skeletonema is a cosmopolitan diatom observed in coastal waters that forms extensive blooms, largely underpinning coastal ecosystems and fisheries. Recent revisions to the phylogeny of Skeletonema have shown multiple pseudocryptic species within the type species S. costatum. We developed a novel species-specific quantitative PCR (qPCR) method to track the annual dynamics of 7 Skeletonema species, including the pseudocryptic species, in the coastal waters of Ariake Sound, Japan. Fortnightly monitoring revealed that each Skeletonema species had different seasonality and occurrence patterns. The significant findings related to: (1) the species-specific ecological strategies and (2) the ecophysiology of the 5 pseudocryptic species assigned as S. costatum (sensu lato). Three Skeletonema species (S. costatum [sensu stricto], S. menzelii, and S. tropicum) formed blooms in summer, while the copy number of other species increased in winter (S. ardens, S. dohrnii/marinoi, S. grevillei, and S. japonicum). Bloom dynamics also differed among species, with either single intense bloom events or sustained moderate blooms. Summer species bloomed successively under the highly dynamic summer conditions (high freshwater input and anticyclonic tropical typhoons). By contrast, in winter, multi-species blooms were observed under conditions of low sea surface temperature and solar insolation. The single-peak species intensely bloomed during an ephemeral optimal time whereas others sustained their biomass in the pax between the intense blooms. Our in situ species-specific monitoring study connects ex situ physiology and in situ distributions, highlighting the diverse ecophysiology within this cosmopolitan diatom genus.

Impact of Shifting Subpolar Front on Phytoplankton Dynamics in the Western Margin of East/Japan Sea

A subpolar front (SPF) generated between the East Korea Warm Current (EKWC) and the North Korea Cold Current (NKCC) in the western margin of the East/Japan Sea has shifted northward in recent decades. This study investigated the biomass and composition of the phytoplankton assemblage in relation to hydrological and biogeochemical features in the shallow shelf and slope off the Korean coast from January to June in 2016 and 2017, to determine the mechanistic effects of SPF on spring–summer phytoplankton bloom dynamics. Monthly average depth-integrated chlorophylla(Chla) levels and the contribution of phytoplankton classes revealed bimodal diatom blooms in early spring and summer in the frontal zone. Canonical correspondence analysis showed that the distribution of high Chlawas associated with cold, low-salinity NKCC water in March 2016. No Chlapeak was observed in March 2017 when the warm saline EKWC water mass invaded. These results suggest that the NKCC intrusion acts as a forcing mechanism leading to enhanced phytoplankton biomass in the frontal zone. In contrast, positive correlations of Chlaconcentration with water density and nutrient concentrations suggest that summer blooms were fed by the subsurface chlorophyll maximum (SCM) driven by shoaling of the pycnocline and nitracline. Varying water-column stratification determined the thickness of the SCM layer, driving year-to-year variability in the magnitude of diatom blooms. These findings further suggest that seasonal/interannual variability in the timing of algal blooms affects regional trophodynamics and hence could be an important factor in explaining ecosystem changes in this region.

Linking population dynamics models with empirically derived models through phytoplankton primary production.

There are alternative methods for estimation of phytoplankton primary production (PP) that are fundamentally different in the calculation approach. The process‐oriented PP model is a mechanistic, empirically derived method based on the photosynthesis–light relationships. The population dynamics‐based PP calculation, which is a synthetic method, provides a production estimate based on population dynamics of phytoplankton. These alternative methods were here compared with regard to production estimates and linked to enhance the performance of the existing models of population dynamics applied to a wide variety of lakes worldwide in terms of morphometry, nutrient status, and light environments. Estimates of PP were shown to be sensitive to changes in phytoplankton sinking and zooplankton grazing rates in both methods. Production estimates in the process‐oriented PP model were also sensitive to light‐associated parameters such as day length. Although the production estimated from the population dynamics‐based PP calculation tended to be lower than that from the process‐oriented PP model irrespective of lake morphometry, production estimates calculated from both methods with standard parameterization were comparable when production was estimated on an annual timescale. However, it was also shown that the alternative methods could produce different production estimates when estimated on shorter timescales such as cyanobacterial blooms in summer. Cyanobacteria with low mortality due to grazing and sinking losses have been considered as trophic bottlenecks, but there is increasing evidence that their mortality is, to a considerable extent, due to parasitic pathogens. In the case of cyanobacterial blooms, an addition of parasite‐related loss term (19%–33% of standing stock) resulted in a resolution of the difference in production estimates between the methods. These analyses theoretically support the critical role of parasitism and resolve the bottleneck problem in aquatic ecosystem metabolism.

Twenty-One Years of Phytoplankton Bloom Phenology in the Barents, Norwegian, and North Seas

Phytoplankton blooms provide biomass to the marine trophic web, contribute to the carbon removal from the atmosphere and can be deadly when associated with harmful species. This points to the need to understand the phenology of the blooms in the Barents, Norwegian, and North seas. We use satellite chlorophyll-a from 2000 to 2020 to assess robust climatological and the interannual trends of spring and summer blooms onset, peak day, duration and intensity. Further, we also correlate the interannual variability of the blooms with mixed layer depth (MLD), sea surface temperature (SST), wind speed and suspended particulate matter (SPM) retrieved from models and remote sensing. The climatological spring blooms start on March 10th and end on June 19th. The climatological summer blooms begin on July 13th and end on September 17th. In the Barents Sea, years of shallower mixed layer (ML) driven by both calm waters and higher freshwaters input keeps the phytoplankton in the euphotic zone, causing the spring bloom to start earlier and reach higher biomass but end sooner due to the lack of nutrients upwelling from the deep. In the Norwegian Sea, a correlation between SST and the spring blooms is found. Here, warmer waters are correlated to earlier and stronger blooms in most regions but with later and weaker blooms in the eastern Norwegian Sea. In the North Sea, years of shallower ML reduces the phytoplankton sinking below the euphotic zone and limits the SPM increase from the bed shear stress, creating an ideal environment of stratified and clear waters to develop stronger spring blooms. Last, the summer blooms onset, peak day and duration have been rapidly delaying at a rate of 1.25-day year–1, but with inconclusive causes based on the parameters assessed in this study.

The response of coastal macrobenthic food-web structure to seasonal and regional variability in organic matter properties

The quantity and quality of organic matter (OM) available for macrobenthos vary on temporal and spatial scale, especially in coastal areas, where the seafloor is supplied by both seasonal pulses of primary production and terrestrial carbon transported by rivers. Human-induced environmental changes will alter OM transport to the seafloor, thus, it is of crucial importance to recognize the impacts of spatio-temporal OM variability on macrobenthic food webs. In this study, the seasonal variability of quantity and quality of OM and its effect on benthic food-web structure were studied at four locations along the southern coast of the Baltic Sea, characterized by similar species pools but different environmental settings and various scales of anthropogenic and riverine influence. Our study shows that temporal changes in the isotopic niche (δ13C and δ15N) of benthic invertebrates follow the patterns of quantity and quality of OM related to seasonal changes of pelagic primary production, riverine discharge of terrestrial OM and the input of anthropogenically enriched sources. In the semi-enclosed bay, exposed to intensive summer blooms, all feeding groups expressed lower δ15N values in autumn, suggesting the incorporation of δ15N-depleted cyanobacteria. At a location supplied by a higher proportion of OM from allochthonous sources (riverine discharge), omnivores occupied higher trophic levels, probably due to a higher abundance of meiobenthic prey. In contrast, at the open Polish coast, where the main food supply occurs during diatom spring bloom, the isotopic niche of omnivorous species showed much higher seasonal variability, underlining the importance of food availability for food-web diversity. Quick adaptation of benthic food-web structure to seasonal variations in OM properties and its modification by local conditions may indicate high plasticity of species feeding behavior that suggests a high capacity of benthic communities to mitigate ongoing changes in the marine environment, at least within the range observed today. Our results confirm that combination of spatial and temporal scales in sampling of the food-web structure is a good indicator of the ecosystem stability and function.

Regionalization of the Red Sea Based on Phytoplankton Phenology: A Satellite Analysis

AbstractThe current average state of Red Sea phytoplankton phenology needs to be resolved in order to study future variations that could be induced by climate change. Moreover, a regionalization of the Red Sea could help to identify areas of interest and guide in situ sampling strategies. Here, a clustering method used 21 years of satellite surface chlorophyll‐a concentration observations to characterize similar regions of the Red Sea. Four relevant phytoplankton spatiotemporal patterns (i.e., bio‐regions) were found and linked to biophysical interactions occurring in their respective areas. Two of them, located in the northern part the Red Sea, were characterized by a distinct winter‐time phytoplankton bloom induced by mixing events or associated with a convergence zone. The other two, located in the southern regions, were characterized by phytoplankton blooms in summer and winter which might be under the influence of water advected into the Red Sea from the Gulf of Aden in response to the seasonal monsoon winds. Some observed inter‐annual variabilities in these bio‐regions suggested that physical mechanisms could be highly variable in response to variations in air‐sea heat fluxes and ENSO phases in the northern and southern half of the Red Sea, respectively. This study reveals the importance of sustaining in situ measurements in the Red Sea to build a full understanding about the physical processes that contribute to phytoplankton production in this basin.

Deep particle stocks following the summer bloom around the Kerguelen islands: Insights into diatoms physiological state, community structure and mortality modes

Particles located at the interface between the surface ocean layer and the top of the mesopelagic domain are the initial vector of the biological pump yet, their nature is still largely unknown. During the MOBYDICK cruise in the vicinity of the Kerguelen Islands (Indian sector of the Southern Ocean) we deployed a recently available device that concentrates and collects deep particles over a predetermined layer of water. In this paper, we present a detailed description of the collected particles and individual planktonic cells, including their taxonomy, carbon and lipid content, as well as cell viability, in order to characterize the particle stocks present at depth. The cruise was carried out at the end of the summer bloom, a period characterized by declining stocks of biogenic material associated with various mortality processes of planktonic organisms. Unexpectedly, the majority of the collected particles consisted of single empty diatom frustules, while fecal pellets and aggregates accounted for only a minor fraction. Distinct mortality processes, from parasitic infection to mesozooplankton grazing, and distinct silicification degrees as well as different life stages could be identified in relation to diatom taxa suggesting the occurrence of several export modes to intermediate and deep layers within the diatom community. We observed a dominant contribution of single cell diatoms (93 ± 6%) to the deep particle stocks but a very small contribution of intact diatom cells (~0.3%) to C content in the intermediate layer (125–500 m), together with a very small fecal pellet contribution, that was dominated by the minipellet size-class. Taxonomical analyses revealed distinct communities west of Kerguelen in the HNLC area compared to the island's fertilized plateau and its eastern flank. Differences in silicification degrees as well as distinct mortality/export processes linked to surface nutrient depletion and trophic interactions (such as parasitic infection or grazing by phaeodarians) were identified in the upper layer, leading to distinct contributions of major diatom taxa to deep suspended particles.

Prey selectivity and feeding rates of the scyphozoan Catostylus tagi (Haeckel, 1869)

Abstract The occurrence of scyphozoan blooms is frequently reported in coastal areas, suggesting a need to assess its impacts on trophic webs. However, many scyphozoan species remain understudied with regard to their feeding ecology. Here, feeding selectivity experiments were conducted to learn about the possible prey preference of Catostylus tagi among mesozooplankton prey and estimate feeding rates. In general, C. tagi demonstrated a relative positive selection on small prey as cyclopoid copepods (Oithona spp. and Oncaea spp.), copepod nauplii as well as non-swimming prey (fish eggs), and ingestion rates increased with prey concentration. Further in situ studies must be conducted during summer blooms, to determine the predatory impact, retrieving data of C. tagi and prey abundance, particularly in important nursery areas such as estuaries.

Lake water dissolved inorganic carbon dynamics revealed from monthly measurements of radiocarbon in the Fuji Five Lakes, Japan

Lakes are sensitive recorders of anthropogenic activities, as human society often develops in their vicinity. Lake sediments thus have been widely used to reconstruct the history of environmental changes in the past, anthropogenic, or otherwise, and radiocarbon dating provides chronological control of the samples. However, specific values of radiocarbon in different carbon reservoirs due to the different pathways of radiocarbon from the upper atmosphere to the lake, called the radiocarbon reservoir age, is always difficult to evaluate because of dynamic processes in and around lakes. There are few systematic studies on radiocarbon reservoir ages for lakes owing to the complex radiocarbon transfer processes for lakes. Here, we investigate lake waters of the Fuji Five Lakes with monthly monitoring of the radiocarbon reservoir effects. Radiocarbon from dissolved inorganic carbon (DIC) for groundwater and river water is also measured, with resulting concentrations (Δ14C) at their lowest at Lake Kawaguchi in August 2018 (–122.4 ± 3.2‰), and at their highest at Lake Motosu in January 2019 (–22.4 ± 2.5‰), despite a distance of 25 km. However, winter values in both lakes show similar trends of rising Δ14C (about 20‰). Our lake water DIC Δ14C results are compared to previously published records obtained from sediments in Lake Motosu and Lake Kawaguchi. These suggest that total organic carbon and compound-specific radiocarbon found in sediments are heavily influenced by summer blooms of aquatic organisms that fix DIC in water. Thus, future studies to conduct similar analyses at the various lakes would be able to provide further insights into the carbon cycle around inland water, namely understanding the nature of radiocarbon reservoir ages.

Depth-Dependent Spatiotemporal Dynamics of Overwintering Pelagic Microcystis in a Temperate Water Body.

Cyanobacteria in the genus Microcystis are dominant components of many harmful algal blooms worldwide. Their pelagic–benthic life cycle helps them survive periods of adverse conditions and contributes greatly to their ecological success. Many studies on Microcystis overwintering have focused on benthic colonies and suggest that sediment serves as the major inoculum for subsequent summer blooms. However, the contemporaneous overwintering pelagic population may be important as well but is understudied. In this study, we investigated near-surface and near-bottom pelagic population dynamics of both microcystin-producing Microcystis and total Microcystis over six weeks in winter at Dog Lake (South Frontenac, ON, Canada). We quantified relative Microcystis concentrations using real-time PCR. Our results showed that the spatiotemporal distribution of overwintering pelagic Microcystis was depth dependent. The abundance of near-bottom pelagic Microcystis declined with increased depth with no influence of depth on near-surface Microcystis abundance. In the shallow region of the lake (<10 m), most pelagic Microcystis was found near the lake bottom (>90%). However, the proportion of near-surface Microcystis rose sharply to over 60% as the depth increased to approximately 18 m. The depth-dependent distribution pattern was found to be similar in both microcystin-producing Microcystis and total Microcystis. Our results suggest the top of the water column may be a more significant contributor of Microcystis recruitment inoculum than previously thought and merits more attention in early CHAB characterization and remediation.

Effects of Multiple Karenia brevis Red Tide Blooms on a Common Bottlenose Dolphin (Tursiops truncatus) Prey Fish Assemblage: Patterns of Resistance and Resilience in Sarasota Bay, Florida

Red tide blooms caused by the toxic dinoflagellate Karenia brevis are natural disturbance events that occur regularly along Florida’s west coast, often resulting in massive fish kills and marine mammal, seabird, and sea turtle mortalities. Limited prior work on the ecological effects of red tides suggests they play an important role in structuring ecosystem dynamics and regulating communities, however specific effects on prey populations and potential alterations to predator-prey interactions are unknown. We surveyed the prey fish assemblage of a top marine predator, the common bottlenose dolphin (Tursiops truncatus), in shallow seagrass habitat in Sarasota Bay, Florida, during 2004–2019, collecting data on prey density, species composition, K. brevis cell densities, and environmental variables. Across eight distinct red tide bloom events, resistance, resilience, and the ecological effects on the prey assemblage varied depending on bloom intensity, season, and frequency. Prey assemblage structure showed significant and distinct short-term shifts during blooms independent of the normal seasonal shifts in prey structure seen during non-bloom conditions. Canonical correspondence analysis indicated a strong influence of K. brevis density on assemblage structure. Blooms occurring primarily in the summer were associated with less initial prey resistance and higher than average annual catch per unit effort (CPUE) 1–3 years following bloom cessation, with bloom frequency prolonging the time needed to reach higher than average annual CPUE. Regardless of season, recovery to pre-bloom prey abundances occurred within 1 year. Sample-based rarefaction and extrapolation indicated significant differences in prey diversity among summer bloom events. This study is a first step in identifying differences in resistance, resilience, and the ecological effects of multiple red tide bloom events of various temporal scales and intensity on a dolphin prey assemblage. Improved understanding of the influence of red tides on estuarine structural dynamics and function can better inform management, and potentially guide mitigation efforts post-bloom.

Oceanography time series reveals annual asynchrony input between oceanic and estuarine waters in Patagonian fjords

The postglacial Patagonian fjord system along the west coast of southern South America is one of the largest stretches of the southern hemisphere (SH) fjord belt, influenced by the SH westerly wind belt and continental freshwater input. This study reports a 3-year monthly time series (2017–2020) of physical and biogeochemical parameters obtained from the Reloncaví Marine Observatory (OMARE, Spanish acronym) at the northernmost embayment and fjord system of Patagonia. The main objective of this work was to understand the land–atmosphere–ocean interactions and to identify the mechanisms that modulate the density of phytoplankton. A key finding of this study was the seasonally varying asynchronous input of oceanic and estuarine water. Surface lower salinity and warmer estuarine water arrived in late winter to summer, contributing to water column stability, followed by subsurface higher salinity and less warmer oceanic water during fall–winter. In late winter 2019, an interannual change above the picnocline due to the record-high polarity of the Indian Ocean Dipole inhibited water column stability. The biogeochemical parameters (NO3−, NO2−, PO43−, Si(OH)4, pH, and dissolved oxygen) responded to the surface annual salinity variations, and oceanic water mass contributed greatly to the subsurface inorganic nutrient input. The water column N/P ratio indicated that no eutrophication occurred, even under intense aquaculture activity, likely because of the high ventilation dynamics of the Reloncaví Sound. Finally, a shift in phytoplankton composition, characterized by surface chlorophyll-a maxima in late winter and deepening of spring–summer blooms related to the physicochemical conditions of the water column, was observed. Our results support the ecosystem services provided by local oceanography processes in the north Patagonian fjords. Here, the anthropogenic impact caused by economic activities could be, in part, chemically reduced by the annual ventilation cycle mediated by the exchange of oceanic water masses into Patagonian fjords.

Assessing energy return on investment for harvest of wild Nodularia spumigena during blooms in the Baltic Sea

Recurring summer cyanobacteria blooms in the Baltic Sea has gained academic interests for decades. The harvest of wild cyanobacteria, for example, Nodularia spumigena, during summer blooms in the Baltic Sea has been studied in the past but lacked evaluation for environmental and economic performances. This study provides a first-hand assessment of environmental and economic performance from an energy perspective, using energy return on investment (EROI) as evaluation method where harvest of biomass and the downstream conversion of biomass to biogas and biofertilizer are considered for Gotland, Sweden. Energy analysis results indicate fuel consumption during harvest and transport operations to be the major energy consumer. Traditional sailing boats have been suggested as an alternative. Overall, when considering only biogas yield and usage of sailing boats, a break-even EROI of 1 is achieved. When including biofertilizer as product, a breakeven EROI of 1 is also achieved. Depending upon the biomass concentration in the Baltic Sea at the time of harvest, an EROI > 6 is possible, surpassing the economic viability EROI benchmark of 3, indicating the importance of nutrient recovery as the driver for harvest of wild cyanobacteria biomass during blooms in the Baltic Sea. This article met the requirements for a gold-gold JIE data openness badge described at http://jie.click/badges.

Exploiting high frequency monitoring and satellite imagery for assessing chlorophyll-a dynamics in a shallow eutrophic lake

Freshwater ecosystems are challenged by cultural eutrophication across the globe, and it is a priority for water managers to implement water quality monitoring at different spatio-temporal scales to control and mitigate the eutrophication process. Phytoplankton abundance is a key indicator of the trophic and water quality status of lakes. Phytoplankton dynamics are characterized by high spatio-temporal variation, driven by physical, chemical and biological factors, that challenge the capacity of routine monitoring with conventional sampling techniques (i.e., boat based sampling) to characterise these complex relationships. In this study, high frequency in situ measurements and multispectral satellite data were used in a synergistic way to explore temporal (diurnal and seasonal) dynamics and spatial distribution of Chlorophyll-a (Chl-a) concentration, a proxy of phytoplankton abundance, together with physico-chemical water parameters in a shallow fluvial-lake system (Mantua Lakes). A good agreement was found between Chl-a retrieved by remote sensing data and Chl-a fluorescence data recorded by multi-parameters probes (R2 = 0.94). The Chl-a maps allowed a seasonal classification of the Mantua Lakes system as eutrophic or hypertrophic. Along the Mantua lakes system an increasing gradient in Chl-a concentration was recorded following the transition from a fluvial to lacustrine system. There was significant seasonal heterogeneity among the sub-basins, probably due to different hydrodynamics, influenced also by macrophyte stands. High-frequency data revealed the importance of rainfall events in the timing and growth dynamics of phytoplankton, particularly for spring and late summer blooms. Combining temporal and spatial data at high resolution improves the understanding of complex fluvial-lake systems. This technique can allow managers to target blooms in near-real time as they move through a system and guide them to localized hot spots enabling timely management action in ecosystems of high conservation and recreational value.

Daytime intra-estuary movements and harmful algal bloom avoidance by Mugil cephalus (family Mugilidae)

The estuarine movement patterns of Mugil cephalus (Mugilidae) in response to Harmful algal blooms (HABs) was investigated in the eutrophied Sundays Estuary on the warm temperate coast of South Africa. The movements of six tagged M. cephalus (31–35 cm FL) were manually tracked using a Vemco VR100 acoustic receiver over 27 days during the typical summer bloom period. Fish were released above, within and below the mesohaline zone where Heterosigma akashiwo bloom maxima are known to occur. Physico-chemical variables and chlorophyll-a biomass were collected at each fish location and at four bloom monitoring sites, which locations were based on surface water salinity of 18, 10, 8 and 5. Mugil cephalus mostly occupied the upper mesohaline to lower oligohaline zones moving with the tidal ebb and flow. Phytoplankton biomass peaked at sites with a salinity of 10 and 8, with tagged fish only briefly (<0.63% of total time free roaming) venturing into bloom maxima areas. This is the first account of small scale intra-estuary movements of a mugilid worldwide and also the first to suggest avoidance behaviour of estuarine fishes in response to HABs. More work is needed to further understand how anthropogenic threats may impact the movements of important estuarine-dependent fish species in order to prevent the possible loss of ecosystem services induced by eutrophication.

Understanding controls on Margalefidinium polykrikoides blooms in the lower Chesapeake Bay

A time-dependent model of Margalefidinium polykrikoides, a mixotrophic dinoflagellate, cell growth was implemented to assess controls on blooms in the Lafayette River, a shallow, tidal sub-tributary of the lower Chesapeake Bay. Simulated cell growth included autotrophic and heterotrophic contributions. Autotrophic cell growth with no nutrient limitation resulted in a bloom but produced chlorophyll concentrations that were 45% less than observed bloom concentrations (~80 mg Chl m−3 vs. 145 mg Chl m−3) and a bloom progression that did not match observations. Excystment (cyst germination) was important for bloom initiation, but did not influence the development of algal biomass or bloom duration. Encystment (cyst formation) resulted in small losses of biomass throughout the bloom but similarly, did not influence M. polykrikoides cell density or the duration of blooms. In contrast, the degree of heterotrophy significantly impacted cell densities achieved and bloom duration. When heterotrophy contributed a constant 30% to cell growth, and dissolved inorganic nitrogen was not limiting, simulated chlorophyll concentrations were within those observed during blooms (maximum ~140 mg Chl m−3). However, nitrogen limitation quenched the maximum chlorophyll concentration by a factor of three. Specifying heterotrophy as an increasing function of nutrient limitation, allowing it to contribute up to 50% and 70% of total growth, resulted in simulated maximum chlorophyll concentrations of 90 mg Chl m−3 and 180 mg Chl m−3, respectively. This suggested that blooms of M. polykrikoides in the Lafayette River are fortified and maintained by substantial heterotrophic nutritional inputs. The timing and progression of the simulated bloom was controlled by the temperature range, 23 °C to 28 °C, that supports M. polykrikoides growth. Temperature increases of 0.5 °C and 1.0 °C, consistent with current warming trends in the lower Chesapeake Bay due to climate change, shifted the timing of bloom initiation to be earlier and extended the duration of blooms; maximum bloom magnitude was reduced by 50% and 65%, respectively. Warming by 5 °C suppressed the summer bloom. The simulations suggested that the timing of M. polykrikoides blooms in the Lafayette River is controlled by temperature and the bloom magnitude is determined by trade-offs between the severity of nutrient limitation and the relative contribution of mixotrophy to cell growth.

Different Roles of Top-Down and Bottom-Up Processes in the Distribution of Size-Fractionated Phytoplankton in Gwangyang Bay

The interactive roles of zooplankton grazing (top-down) and nutrient (bottom-up) processes on phytoplankton distribution in a temperate estuary were investigated via dilution and nutrient addition experiments. The responses of size-fractionated phytoplankton and major phytoplankton groups, as determined by flow cytometry, were examined in association with zooplankton grazing and nutrient availability. The summer bloom was attributed to nanoplankton, and microplankton was largely responsible for the winter bloom, whereas the picoplankton biomass was relatively consistent throughout the sampling periods, except for the fall. The nutrient addition experiments illustrated that nanoplankton responded more quickly to phosphate than the other groups in the summer, whereas microplankton had a faster response to most nutrients in the winter. The dilution experiments ascribed that the grazing mortality rates of eukaryotes were low compared to those of the other groups, whereas autotrophic cyanobacteria were more palatable to zooplankton than cryptophytes and eukaryotes. Our experimental results indicate that efficient escape from zooplankton grazing and fast response to nutrient availability synergistically caused the microplankton to bloom in the winter, whereas the bottom-up process (i.e., the phosphate effect) largely governed the nanoplankton bloom in the summer.

Summer‐flowering trees and shrubs in landscaping for the south‐eastern coast of the Crimea

Aim. To study the species composition, form diversity, bioecological features of summer‐ flowering trees and shrubs in green spaces of the south‐eastern coast of the Crimea and suggest ways to expand the range. Materials and Methods. The green spaces of Feodosia, Sudak and the localities of Semidvorye, Solnechnogorskoye, Malorechenskoye, Rybachye, Morskoye, Novy Svet, Solnechnaya Dolina, Shchebetovka, Kurortnoye, Koktebel and Primorskiy were investigated. Bioecological and phenological studies of ornamental trees and shrubs were undertaken according to generally accepted methods. Results. The species composition and shape diversity of summer‐flowering trees and shrubs in green plantings of Crimea’s south‐eastern coast were identified. It was found that summer‐ flowering plants make up almost half of the species composition (48.6%) used in landscaping. However, only 23 species are widespread throughout the region, such as Ailanthus altissima (Mill.) Swingle, Albizia julibrissin Durazz., Buddleja davidii Franch., Campsis radicans (L.) Seem., Elaeagnus angustifolia L. etc. The greatest number of species and forms of arboreal plants bloom in early summer, their number decreasing significantly from the second decade of June. Conclusion. An assortment is proposed for increasing the number of trees and shrubs which flower during the summer, taking into account both their decorative visual appeal and their environmental requirements, especially drought and frost resistance. Of great interest are species that bloom from June onwards, especially those which are highly decorative and have a long flowering period, such as Albizia julibrissin, Buddleja davidi and forms, Hydrangea arborescens L., Campsis radicans, Kerria japonica f. plena C.K. Schneid, Robinia viscosа Vent., Vitex angus‐castus L.