Green Algae Blooms Research Articles

Morphological and Molecular Identification of Ulva spp. (Ulvophyceae; Chlorophyta) from Algarrobo Bay, Chile: Understanding the Composition of Green Tides.

Green algae blooms of the genus Ulva are occurring globally and are primarily attributed to anthropogenic factors. At Los Tubos beach in Algarrobo Bay along the central Chilean coast, there have been blooms of these algae that persist almost year-round over the past 20 years, leading to environmental, economic, and social issues that affect the local government and communities. The objective of this study was to characterize the species that form these green tides based on a combination of ecological, morpho-anatomical, and molecular information. For this purpose, seasonal surveys of beached algal fronds were conducted between 2021 and 2022. Subsequently, the sampled algae were analyzed morphologically and phylogenetically using the molecular markers ITS1 and tufA, allowing for the identification of at least five taxa. Of these five taxa, three (U. stenophylloides, U. uncialis, U. australis) have laminar, foliose, and distromatic morphology, while the other two (U. compressa, U. aragoensis) have tubular, filamentous, and monostromatic fronds. Intertidal surveys showed that U. stenophylloides showed the highest relative coverage throughout the seasons and all intertidal levels, followed by U. uncialis. Therefore, we can establish that the green tides on the coast of Algarrobo in Chile are multispecific, with differences in relative abundance during different seasons and across the intertidal zone, opening opportunities for diverse future studies, ranging from ecology to algal biotechnology.

Dynamics of dissolved organic phosphorus in the nearshore of eastern Lake Erie

Abstract The east basin of Lake Erie continues to suffer from blooms of filamentous green algae (primarily Cladophora). Potential management action through reduced phosphorus loadings have and continue to focus on the dissolved inorganic phosphorus pool but other potentially bioavailable phosphorus pools are not always considered. In this study, we describe the dissolved organic phosphorus pool in eastern Lake Erie, with an extensive sampling of four transects along the north shore of the eastern basin from May to September 2019. The dissolved organic phosphorus pool was characterized and quantified using sequential enzymatic hydrolysis to provide information on the enzymatically hydrolysable phosphorus fraction and component monoester P, diester P and a phytase hydrolysable component. These dissolved organic phosphorus fractions were compared to the soluble reactive phosphorus and total dissolved phosphorus pools. Results from this study revealed that a significant fraction (up to 63%) of the dissolved organic phosphorus pool is potentially bioavailable via enzymatic hydrolysis. Vertical differences in soluble reactive phosphorus, total dissolved phosphorus, and phosphate diester pools were also observed when comparing water column with near lake-bed samples, suggesting that the new benthic ecosystem since dreissenid colonization may be a greater source of dissolved inorganic phosphorus to the east basin than prior to colonization.

Signals of intense primary production in response to Ulva prolifera bloom in the Yellow Sea during summer 2021

The variability in changing climate has been predicted to lead to significant changes in the world's oceans primary productivity. Recent years have experienced more intense algal blooms during the spring-summer months in many coastal waters. Here we present results from the analysis of satellite ocean color products from MODIS-Terra datasets during June 2 to July 20, 2021, corresponding to the most intense period of the Enteromorpha (Ulva) prolifera algae (U. prolifera) bloom in the Yellow Sea. Because the studies on sea surface temperature (SST) variability over the U. prolifera bloom have not yet been fully explored by satellite measurements in the offshore waters of Qingdao, China, we analyzed the MODIS SST and their relationship with the U. prolifera blooms in the 2021 summer months. Analysis of SST showed a variation in the range of 18–24 °C in near-shore waters in response to the bloom of green algae. We find that a significant thermal front with a gradient magnitude of 0.03 °C/km was developed over the shelf and offshore regions during the algal bloom period. The observed changes in thermal fronts are driven by high-wind speed and Ekman pumping-velocity processes. We concluded that the changes in SSTs with a range of 20–23 °C, high-magnitude thermal fronts and wind speed, and Ekman pumping velocity resulted in the intensification of the largest bloom of U. prolifera in the Yellow Sea during the study period.

Are mussels and oysters capable of reducing the abundances of Picochlorum sp., responsible for a massive green algae bloom in Thau lagoon, France?

After an exceptional hydroclimatic year, a “massive green algae bloom”, dominated by the autotrophic picoeukaryote Picochlorum sp, was observed in 2018–2019 in the Thau lagoon. Oyster farmers informed decision-makers and scientists about an alarming halt to growth, loss of flesh weight and abnormal mortality in commercial size oysters, whereas mussel farmers observed no adverse effects. Two hypotheses were tested. First, Mediterranean mussels are capable of filtering Picochlorum spp. whereas Pacific and flat oysters are not. Second, picophytoplankton filtration rates vary with the size of the oyster, spats having greater ability than juveniles and commercial size oysters. A series of experiments was conducted in February 2019 using a series of beakers containing (i) three different types of solutions (1 L): water from the Thau lagoon containing Picochlorum at a rate of 45.2 million cells.L−1 (P) or diluted to 20.7 million cells. L−1 (D) and a solution of Isochrysis galbana containing 9.4 million cells. L−1; (ii) in the absence or presence of five types of bivalves: Pacific oysters of three different sizes (spat, juvenile and commercial size), flat oysters, and Mediterranean mussels of commercial size. Four separate water samples were taken during a 40-min incubation period to measure fluctuations in picophytoplankton and nanophytoplankton abundances using flow cytometry. Retention efficiency (%) was compared according to species and size. In contrast to Pacific and flat oysters, mussels with an average weight of 21.4 g, were able to drastically reduce the abundances of Picochlorum and depleted nearly one million Picochlorum cells per minute at 11.4 °C. These results suggest that the 1228 t of mussels that died in 2018 before the Picochlorum bloom could have helped limit the Picochlorum bloom if they had survived the heat and anoxia.

Effect of Anthropogenic Factors on the Ecosystem of the Yenisei River Anabranch within the City of Krasnoyarsk

In recent decades, freshwater ecosystems have regularly experienced blooms of green algae (green tides), causing environmental and economic damage. Blooms of green alga Spirogyra sp. occur in the ‘Abakanskaya’ anabranch of the Yenisei River in summer months. The anabranch is influenced by several anthropogenic factors: construction of a dam in the upper reaches; heated water discharge from a thermal power plant; fish farming. The aim of this study was to assess the impact of anthropogenic factors on the ecosystem of the Abakanskaya anabranch and to identify the main factors favoring Spirogyra sp. blooms. Samples were taken in May-September 2018-2019 at four sites: 1 – upstream of the dam (control), 2 – downstream of the dam, 3 – near the point of heated water discharge, 4 – downstream of the fish farm. Physical and chemical parameters, biomass and species composition of phytoplankton, phytoperiphyton, zooplankton, zoobenthos, and higher water plants, gross primary production (GPP), nitrogen content in biomass of periphyton, higher plants, and zoobenthos were studied. The main factor that caused changes in the community was that there was no water flow in the anabranch because of the low capacity of the dam drainage pipes. While upstream of the dam most ecosystem features were within the limits typical for this part of the Yenisei River, downstream of the dam, the system seemed to change from lotic to lentic: percentage of motile forms of phytoplankton, biomass of zooplankton, percentage of true zooplankton species, taxon number and taxonomic diversity of zooplankton and zoobenthos became higher; phytoperiphyton primary production rose (because of Spirogyra sp. bloom); higher water plants thrived. The other investigated factors, namely, thermal, nutrient, and organic pollution evidently produced combined effect on the ecosystem because of the absence of water flow, and their specific effects were not obvious

Hindcast and Near Real-Time Monitoring of Green Macroalgae Blooms in Shallow Coral Reef Lagoons Using Sentinel-2: A New-Caledonia Case Study

Despite the necessary trade-offs between spatial and temporal resolution, remote sensing is an effective approach to monitor macroalgae blooms, understand their origins and anticipate their developments. Monitoring of small tropical lagoons is challenging because they require high resolutions. Since 2017, the Sentinel-2 satellites has provided new perspectives, and the feasibility of monitoring green algae blooms was investigated in this study. In the Poé-Gouaro-Déva lagoon, New Caledonia, recent Ulva blooms are the cause of significant nuisances when beaching. Spectral indices using the blue and green spectral bands were confronted with field observations of algal abundances using images concurrent with fieldwork. Depending on seabed compositions and types of correction applied to reflectance data, the spectral indices explained between 1 and 64.9% of variance. The models providing the best statistical fit were used to revisit the algal dynamics using Sentinel-2 data from January 2017 to December 2019, through two image segmentation approaches: unsupervised and supervised. The latter accurately reproduced the two algal blooms that occurred in the area in 2018. This paper demonstrates that Sentinel-2 data can be an effective source to hindcast and monitor the dynamics of green algae in shallow lagoons.

Effects of microcystin on the motility of planarians

Cyanobacteria (blue green algae) blooms formed in natural water resources due to the environmental pollution produce toxic compounds as secondary metabolites, causing health hazards to both humans and other living beings. Microcystin is a well-known toxin produced by cynobacteria. The present study was undertaken to evaluate varying concentrations and exposure times of two different forms of microcystin, viz., -LR (MCLR) and -LA (MCLA), on the motility and seizure-like behavior of planarian (Dugesia japonica). Compared to control, reduced motility was observed in both the MCLR or MCLA treated groups, but did not differ significantly with increasing concentrations of microcystin. However, the number of seizure-like behaviors were increased dose-dependently in planarian exposed to MLCR or MCLA. Exposure time to microcystine also affected the motility and seizure-like behaviors of planarians; 24 hrs incubation with MCLR, and 48 and 96 hrs exposure to MCLA, showed significantly (p<0.05) lower motility, as compared to the control. Assessing regeneration of the planarians revealed the simultaneous completion of eye formation at day 9 in planarians incubated in the absence or presence of MCLR or MCLA, thereby indicating that exposure to microcystin has no effect on the process. In conclusion, we determined that exposure to microcystins resulted in decrease in the number of motility, and induced abnormal behavior pattern in planarians. Further studies are required to identify the toxicity of microcystin that affects aquatic ecosystems.

Filamentous green algae Spirogyra regulates methane emissions from eutrophic rivers.

Excessive growth of filamentous green algae in rivers has attracted much attention due to their functional importance to primary production and carbon cycling. However, comprehensive knowledge of how filamentous green algae affect carbon cycling, especially the CH4 emissions from river ecosystems, remains limited. In this study, incubation experiments were conducted to examine the factors regulating CH4 emissions from a eutrophic river with dense growth of filamentous green algae Spirogyra through combinations of biogeochemical, molecular biological, and stable carbon isotope analyses. Results showed that although water dissolved oxygen (DO) in the algae+sediment (A+S) incubation groups increased up to 19 mg L-1, average CH4 flux of the groups was 13.09 μmol m-2 day-1, nearly up to two times higher than that from sediments without algae (S groups). The significant increase of sediment CH4 oxidation potential and methanotroph abundances identified the enhancing sediment CH4 oxidation during Spirogyra bloom. However, the increased water CH4 concentration was consistent with depleted water [Formula: see text] and decreased apparent fractionation factor (αapp), suggesting the important contribution of Spirogyra to the oxic water CH4 production. It can thus be concluded that high DO concentration during the algal bloom promoted the CH4 consumption by enhancing sediment CH4 oxidation, while algal-linked oxic water CH4 production as a major component of water CH4 promoted the CH4 emissions from the river. Our study highlights the regulation of Spirogyra in aquatic CH4 fluxes and will help to estimate accurately CH4 emissions from eutrophic rivers with dense blooms of filamentous green algae. Graphical abstract.

Dynamic Diurnal Changes in Green Algae Biomass in the Southern Yellow Sea Based on GOCI Images

Macroalgae blooms of Ulva prolifera have occurred every summer in the southern Yellow Sea since 2007, inducing severe ecological problems and huge economic losses. Genesis and secular movement of green algae blooms have been well monitored by using remote sensing and other methods. In this study, green algae were detected and traced by using Geostationary Ocean Color Imager (GOCI), and a novel biomass estimation model was developed from the relationship between biomass measurements and previously published satellite-derived biomass indexes. The results show that the green algae biomass can be determined most accurately with the biomass index of green algae for GOCI (BIGAG), which is calculated from the Rsurf data that had been atmospherically corrected by ENVI/QUAC method. For the first time, dynamic changes in green algae biomass were studied over an hourly scale. Short-term biomass changes were highly influenced by Photosynthetically Available Radiation (PAR) and tidal phases, but less by sea surface temperature variations on a daily timescale. A new parameter of biomass changes (PBC), calculated by the ratio of the biomass growth rate to movement velocity, could provide an effective way to assess and forecast green tide in the southern Yellow Sea and similar areas.

Remotely-Observed Early Spring Warming in the Southwestern Yellow Sea Due to Weakened Winter Monsoon

The seasonal warming over the southwestern Yellow Sea (YS) in the spring is of vital importance to the local ecologic environment, especially to the massive green algae blooms of the YS in late spring and early summer. Based on daily optimum interpolation sea surface temperature (SST) data consisting of satellite derived SST from Advanced Very High Resolution Radiometer (AVHRR) and in situ measurements, this study analyzed the spring SST variation over the southwestern YS (SWYS) from 1982 to 2018. The results show that the recent warming trend of spring SST over the SWYS is four-to-six times that of the global average, and as a result, sea water over the Subei Shoal (SBS) shifts about 10–13 days earlier to reach 10 °C in early April. This implies that, accordingly, the micro-propagules of green algae over the SBS may have the chance to germinate earlier. SST variability in early April significantly correlates with northerly wind and exhibits a general warming over the SWYS with an intensified warming anchored along the axis of the submarine canyon off the Yangtze estuary. The Moderate Resolution Imaging Spectroradiometer (MODIS) red–green–blue composite images captured the intrusion of the Taiwan Warm Current (TWC) into the SWYS through the submarine canyon during northerly wind relaxation in early April. Ocean remote sensing provides important clues for understanding the regional SST variability in the SWYS. Following this clue, this study finds that the weakening of winter monsoon in the spring leads to northward migration of the TWC and results in enhanced spring warming over the SWYS. The attendant advanced warming in spring, resulting in a favorable temperature condition for early development of green alga, may have contributed to the green tide blooms in the Yellow Sea in the recent decade.

Water Quality Measurement using Amphibious Drones and Development of Chlorophyll-a Prediction Formula Through Temperature and DO

As the eutrophication in rivers becomes serious, the green algae bloom has also a great influence on the water quality in recent years. In South Korea, since most of the water resources come from rivers, it is important to predict and prevent the causative phenomena beforehand in order to prevent contamination of water resources. In order to know the number of Chl-a in the current stream, the process is complex and time-consuming. To solve these problems, this study compared the growth rate of Chl-a by temperature and DO and developed a formula that can predict the number of Chl-a by using the UV absorbance value. Culturing for developing formula were conducted inside a constant temperature chamber, and the temperature and DO conditions were determined by referring to river water quality of 2016. Culturing showed that Chl-a grew equally well at a temperature of 27oC under all DO conditions. Also, by developing formula that can predict the number of Chl-a using DO, temperature and UV absorbance values, so that can know the number of Chl-a without doing complex experiment.

Analysis of Water Quality Conditions of Lake Hachiroko Using Fuzzy C-Means

Lake Hachiroko, Japan, has many water quality issues, evident from phenomena such as green algae blooms. Understanding the details of the surface water quality of the lake, and the effect of seasons on the quality, is important. In our previous studies, we conducted fuzzy regression analysis of remote sensing data and direct measurements of water quality. The results showed that estimation maps of water quality were well created, using only five data points of the water quality parameters. To obtain maps that are in good agreement with the experimental data, remote sensing data and water quality values should be acquired simultaneously. However, performing such simultaneous observations can affect the preparation of the water quality estimation maps. We overcame this obstacle by using fuzzy c-means clustering (FCM), and considered the effect of specific disturbances and uncertainties on the remote sensing data. Furthermore, FCM using only remote sensing data creates estimation maps in which relative water surface conditions are classified. Therefore, determining the relationship between FCM results and water quality facilitates the creation of low-cost, high-frequency water quality estimation maps. Our results indicated that FCM was particularly effective in determining the presence of suspended solids (SS) during water quality analysis. However, the relationship between FCM results and water quality has not been determined in detail. In this study, we analyzed the water quality conditions of Lake Hachiroko with FCM using the data collected by the Advanced Space-borne Thermal Emission and Reflection Radiometer on Terra and, the Operational Land Imager on Landsat-8. In addition, FCM results were compared with the maps created by fuzzy regression analysis and the actual conditions of water pollution. The results indicated that (i) the maps created using FCM are effective in determining the water surface conditions, (ii) the FCM maps using data obtained during August and September have a strong relationship with biochemical oxygen demand (BOD) and SS, and (iii) the FCM maps using data obtained during May and June have a strong relationship with chemical oxygen demand (COD), SS, and total nitrogen (T-N).

Dual bloom of green algae and purple bacteria in an extremely shallow soda pan

In April 2014, dual bloom of green algae and purple bacteria occurred in a shallow, alkaline soda pan (Kiskunság National Park, Hungary). The water was only 5 cm deep, in which an upper green layer was clearly separated from a near-sediment purple one. Based on microscopy and DNA-based identification, the upper was inhabited by a dense population of the planktonic green alga, Oocystis submarina Lagerheim, while the deeper layer was formed by purple, bacteriochlorophyll-containing bacteria, predominated by Thiorhodospira and Rhodobaca. Additional bacterial taxa with a presumed capability of anoxygenic phototrophic growth belonged to the genera Loktanella and Porphyrobacter. Comparing the bacterial community of the purple layer with a former blooming event in a nearby soda pan, similar functional but different taxonomic composition was revealed. Members from many dominant bacterial groups were successfully cultivated including potentially new species, which could be the result of the application of newly designed media.

Characteristics of spring green algae blooms and their influencing factors in an urban lake, Moon Lake in Ningbo City, China

Characteristics of spring green algae blooms and their influencing factors in an urban lake, Moon Lake in Ningbo City, China

Physiological changes of submerged macrophytes in response to a floating filamentous green algae bloom in clear-water conditions

The aim of this study was to evaluate the physiological responses of a submerged macrophyte to a floating filamentous green algal bloom in clear-water conditions. Elodea nuttallii was grown with floating Cladophora sp. at four different levels (0, control; 140, 280, 560 g FW/m(2)) in an outdoor experimental system, and its photosynthetic and antioxidant systems were evaluated. The presence of floating Cladophora sp. significantly changed the water environment by decreasing light intensity and increasing dissolved oxygen and the pH value. The photosynthetic parameters of E. nuttallii (e.g. Delta F/F-m',F-v/F-m, total chlorophyll) were higher in the presence of floating Cladophora sp. than in the control at the beginning of experiment. Because of the increasing dissolved oxygen concentration and pH value, the values of these indicators decreased (except for photosynthetic pigments) during the experiment. Compared with E. nuttallii in the control, E. nuttallii growing in the presence of floating Cladophora sp. showed higher malondialdehyde content, catalase activity, and peroxidase activity. The biomass of E. nuttallii was decreased by about 30% in the presence of high biomasses of floating Cladophora sp. (280 and 560 g FW/m(2)). These results suggest that floating Cladophora had complex effects on the biomass of E. nuttallii and that changes in water quality resulting from floating Cladophora sp. may be more important than its direct shading effect.

Interaction between simulated dense Scenedesmus dimorphus (Chlorophyta) bloom and freshwater meta-zooplankton community

Algal bloom has been a subject of much research, especially the occurrence of blue-green algae (cyanobacteria) blooms and their effects on aquatic ecosystems. However, the interaction between green algae blooms and zooplankton community was rarely investigated. In the present study, the effects exerted by Scenedesmus dimorphus (green alga) bloom on the community structure of zooplankton and the top-down control of the bloom process mediated by the zooplankton were evaluated using a series of laboratory cultures. The results showed that a dense S. dimorphus bloom could change the zooplankton community structure by decreasing its diversity indices, leading to the enrichment of a particular zooplankton species, Brachionus calyciflorus. In the presence of mixed species of zooplankton, the density of S. dimorphus in the culture was decreased as determined by a change in total chlorophyll a (Chl a) concentration, which was about 200 μg L-1 lower than that of the zooplankton-free culture. Furthermore, the number of species belonging to Cladocera, Copepoda and Rotifera all decreased, with all the cladocerans disappeared in the co-culture within 2 weeks of culturing, while the density of rotifers increased from 818 (±243) ind L-1 at the time of inoculation to 40733 (±2173) ind L-1 on the 14th day post-inoculation. Grazing of S. dimorphus by the rotifer B. calyciflorus neutralized its growth, and the gradual increase in B. calyciflorus density eventually led to the collapse of the bloom. Furthermore, grazing by B. calyciflorus also led to a decrease in the maximal photochemical efficiency (Fv/Fm) of photosystem II (PSII). The combined changes occurring in the zooplankton community structure during the process of S. dimorphus bloom and the negative effects of grazing on algal growth, morphology and photosynthetic activities confirmed the key role of zooplankton in the control of algal bloom. The results of the study therefore indicated that dense algal blooms caused by non-toxic algae could still remain a threat to aquatic ecosystems.

A Novel Spectral-Unmixing-Based Green Algae Area Estimation Method for GOCI Data

Geostationary Ocean Color Imager (GOCI) data have been widely used in the detection and area estimation of green algae blooms. However, due to the low spatial resolution of GOCI data, pixels in an image are usually “mixed,” which means that the region a pixel covers may include many different materials. Traditional index-based methods can detect whether there are green algal blooms in each pixel, whereas it is still challenging to determine the proportion that green algae blooms occupy in a pixel. In this paper, we propose a novel subpixel-level area estimation method for green algae blooms based on spectral unmixing, which can not only detect the existence of green algae but also determine their proportion in each pixel. A fast endmember extraction method is proposed to automatically calculate the endmember spectral matrix, and the abundance map of green algae which could be regarded as the area estimation is obtained by nonnegatively constrained least squares. This new fast endmembers extraction technique outperforms the classical N-FINDR method by applying two models: candidates location and distance-based vertices determination. In the first model, we propose a medium-distance-based candidates location strategy, which could reduce the searching space during vertices selection. In the second model, we replace the simplex volume measure with a more simple distance measure, thus complex matrix determinant calculation is avoided. We have theoretically proven the equivalency of volume and distance measure. Experiments on GOCI data and synthetic data demonstrate the superiority of the proposed method compared with some state-of-art approaches.

Examining Recreation Demand for Lakeshore Parks in Oklahoma: What is Causing the Downward Trend in Attendance?

EXECUTIVE SUMMARY: This paper develops a demand model to examine attendance trends at individual state parks in Oklahoma between 1998 and 2014. The model tests whether visitation and thus the decision to visit is affected by reservoir water levels, blue green algae blooms, weather conditions and gas prices, in addition to other controls. The results show that visitation declines by less than 1% for every foot that reservoir water levels move away from normal surface elevation, and 7% for every $1 increase in the price of gas. Since 1998, visitation to these parks has persistently declined, which the model attributes to gas price increases and other time varying (site-invariant) conditions, rather than changes in water or weather conditions. Subscribe to JPRA

Coagulation performance and floc characteristics of polytitanium tetrachloride (PTC) compared with titanium tetrachloride (TiCl 4 ) and ferric chloride (FeCl 3 ) in algal turbid water

Abstract Seasonal green algae blooms in freshwaters have raised attention on the need to develop novel effective treatment processes for the removal of algae in water. In the present study, the performance of newly developed polytitanium tetrachloride (PTC) coagulant for the removal of freshwater microalga Chlorella vulgaris has been investigated and compared with titanium tetrachloride (TiCl 4 ) coagulant and the conventional ferric chloride (FeCl 3 ) coagulant. The main benefit of using titanium-based coagulants is that the sludge produced after flocculation may be recycled into a valuable product: titanium dioxide photocatalyst. Both titanium-based coagulants achieved good flocculation over a broader pH range and coagulant dose compared to conventional FeCl 3 coagulant. All three coagulants achieved comparable performance in terms of turbidity removal (i.e. turbidity removal efficiency >97%); although TiCl 4 performed slightly better at the lower tested dose (i.e. 3 and also grew at a faster rate. This study indicates that Ti-based coagulants are effective and promising coagulants for algae removal in water.

The effect of bloom of filamentous green algae on the reproduction of yellowfin sculpin Cottocomephorus grewingkii (Dybowski, 1874) (Cottoidae) during ecological crisis in Lake Baikal.

In shallow water areas of open Lake Baikal, filamentous green alga of the genus Spirogyra grows abundantly. Together with alga of the genus Ulothrix, it forms algal mats. According to our observations from 2010 to 2013, the spawning habitat conditions for the yellowfin sculpin Cottocomephorus grewingkii (Dybowski, 1874) (Cottidae) proved to be significantly disturbed in the littoral zone of Listvennichnyi Bay (southern Baikal), which, in turn, reduced the number of egg layings. With a 100% projective cover of the floor and a high density of green filamentous algae, the shallow-water stony substrate becomes completely inaccessible for spawning of the August population.