Impact of submerged macrophytes on fish-zooplanl phytoplankton interactions: large-scale enclosure experiments in a shallow eutrophic lake

1. The impact of changes in submerged macrophyte abundance on fish-zooplankton-phytoplankton interactions was studied in eighteen large-scale (100 m²) enclosures in a shallow eutrophic take. The submerged macrophytes comprised Potamategon pectinatus L., P. pusillus L. and Callitriche hermaphroditica L. while the fish fry stock comprised three-spined sticklebacks, Gasterosteus acuteatus L., and roach, Rutilus rutilus L. 2. In the absence of macrophytes zooplankton biomass was low and dominated by cyclopoid copepods regardless of fish density, while the phytoplankton biovolume was high (up to 38 mm³1) and dominated by small pennate diatoms and chlorococcales. When the lake volume infested by submerged macrophytes (PVI) exceeded 15–20% and the fish density was below a catch per unit effort (CPUE) of 10 (approx. 2 fry m^?2), planktonic cladoceran biomass was high and dominated by relatively large-sized specimens, while the phytoplankton biovolume was low and dominated by small fast-growing flagellates. At higher fish densities, zooplankton biomass and average biomass of cladocerans decreased and a shift to cyclopoids occurred, while phytoplankton biovolume increased markedly and became dominated by cyanophytes and dinoflagellates. 3. Stepwise multiple linear regressions on log-transformed data revealed that the biomass of Daphnia, Bosmina, Ceriodaphmia and Chydorus were all significantly positively related to PVI and negatively to the abundance of fish or PVI x fish. The average individual biomass of cladocerans was negatively related to fish, but unrelated to PVI. Calculated zooplankton grazing pressure on phytoplankton was positively related to PVI and negatively to PVI x fish. Accordingly the phytoplankton biovolume was negatively related to PVI and to PVI x zooplankton biomass. Cyanophytes and chryptophytes (% of biomass) were positively and Chlorococcales and diatoms negatively related to PVI, while cyanophytes and Chlorococcales were negatively related to PVI x zooplankton biomass. In contrast diatoms and cryptophytes were positively related to the zooplankton biomass or PVI x zooplankton. 4. The results suggest that fish predation has less impact on the zooplankton community in the more structured environment of macrophyte beds, particularly when the PVI exceeds 15–20%. They further suggest that the refuge capacity of macrophytes decreases markedly with increasing fish density (in our study above approximately 10 CPUE). Provided that the density of planktivorous fish is not high, even small improvements in submerged macrophyte abundance may have a substantial positive impact on the zooplankton, leading to a lower phytoplankton biovolume and higher water transparency. However, at high fish densities the refuge effect seems low and no major zooplankton mediated effects of enhanced growth of macrophytes are to be expected.     

Seasonal phytoplankton variations in the shallow Pahlavi Mordab,Iran

Pelagic phytoplankton variations in the shallow, western basin of the Pahlavi Mordab, Iran were monitored by the chlorophyll a, gross oxygen productivity and cell enumeration techniques. Annual maxima were 29.4 mg/m³, 0.98 mg O₂/l/hr and 32, 639, 467 cells/liter respectively and were recorded from late summer until early autumn. During the spring and summer, dense growths of Ceratophyllum, Hydrilla and Myriophyllum flourished. Competition between the phytoplankton and submerged aquatic macrophytes was minimal following late summer, when water depths increased following heavy seasonal precipitation and the die-back of submerged aquatic macrophytes occurred.     

The structure of the plankton community of the Öregrundsgrepen (southwest Bothnian Sea)

Taxonomic composition and variations in density and biomass of the plankton community in the Öregrundsgrepen, a shallow coastal area, were investigated from June 1972 to November 1973. The phytoplankton biomass was large in spring but small during the rest of the year. The spring bloom was dominated by diatoms and dinoflagellates, especially byThalassiosira spp. which were also important during other seasons. Small forms, such asCryptomonas spp.,Rhodomonas spp. and monads, dominated during summer. Blue-green algae were never of any major importance. During the summer, the trophogenic layer exceeded 10 m in thickness. The metazoan fauna was of lower diversity than the plankton flora. The dominating species, the copepodsAcartia bifilosa andEurytemora affinis, constituted on the average 83% of the standing crop. The low salinities, 5–6 S, were regarded as the principal pertinent limiting factor. The metazoan fauna reached large biomass values from July to October. The protozoan fauna (in the case of ciliates), obtained biomass maxima during the spring bloom. It is suggested that the Öregrundsgrepen represents an area of elevated productivity within a region of low overall production, presumably due to local upwelling. From June 1972 to May 1973, the average biomasses were: phytoplankton 0.464 g C m^–2, ciliates 0.040 g C m^–2, copepod nauplii 0.010 g C m^–2, micro-rotifers 0.004 g C m^–2, and mesozooplankton (larger than 0.2 mm) 0.312 g C m^–2. It is estimated that about than 60% of the phytoplankton production is consumed by the microzooplankton (<0.2 mm).     

Composition and Productivity of the Benthic Macroinvertebrate Community of a Subtropical Reservoir

Physical-chemical conditions, phytoplankton productivity, community structure and productivity of the macroinvertebrate benthic community were determined during 1976–77 in a subtropical reservoir. Physical-chemical results revealed high nitrate and phosphate concentrations with highest values in the riverine segment. Large phytoplankton populations were present during most of the year. Phytoplankton productivity was high, producing an annual mean of 87 mg C · m⁻³ · h⁻¹ (12 hours light day). High turbidity in the riverine segment limited phytoplankton productivity during winter and spring. Macrobenthos was dominated by chironomids (Chironomus, Procladius, Coelotanypus and Tanypus) and oligochaetes (Limnodrilus). The annual mean benthic population was estimated at 1,626 · m⁻² with a mean dry weight of 0.66 g · m⁻². Mean benthic species diversity was 1.80. A lacustrine-riverine community gradient was revealed. Benthic productivity was 6.8 g · m⁻² · yr⁻¹ (dry weight) with a P: B ratio of 10. A low correlation was observed between benthic and phytoplankton productivity, and between phytoplankton standing crop and benthic macroinvertebrate numbers throughout the reservoir. Algal food supplies had little impact on the benthic community which was composed predominately of species which fed mostly on organic detritus. Stressful conditions caused by low dissolved oxygen concentrations probably inhibited development of the benthic community throughout the reservoir during summer months, while high sedimentation rates limited development in the head waters.     

Seasonal dynamics and composition of cladoceran and copepod assemblages in ponds of a Hungarian cutaway peatland

Seasonal dynamics of cladoceran and copepod zooplankton were studied over a one‐year period in five permanent ponds of a cutaway peatland, situated in the Danube−Tisza Interfluve, Central Hungary. 17 cladoceran, 11 cyclopoid copepod and 6 harpacticoid copepod species were identified and most of them were typical of small lowland ponds. Nevertheless, some taxa like Cyclops insignis, Ceriodaphnia setosa and Macrocyclops distinctus are considered to be rare in Hungary. The microcrustacean assemblages exhibited apparent seasonal succession with typical seasonal species. There appears to be at least two main successional patterns in the five ponds. After general cyclopoid copepod dominance in winter (Cyclops strenuus and Cyclops insignis), at sites with higher proportion of open water and submerged vegetation, spring was characterized by the dominance of the large cladoceran Daphnia curvirostris, which declined during summer, when microcrustacean assemblages composed mainly of smaller, littoral cladocerans. At these sites, species richness and diversity reached their maximum in autumn. In the case of duckweed covered ponds, succession led to less diverse autumn assemblages with fewer species, dominated by Simocephalus exspinosus. Our results draw the attention to the importance of long‐term investigations and the often neglected winter sampling in the accurate evaluation of species richness (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

A Seasonal Quantitative Study of the Littoral Cladocera and Copepoda in a Bog Pond and an Acid Marsh in Newfoundland

A quantitative study was made of the littoral microcrustaceans in a bog pond and an acid marsh located on the Avalon Peninsula, Newfoundland, from May, 1972 to May, 1973. A total of 17 species of Cladocera and 7 species of Copepoda were found in La Manche Marsh, while 21 species of Cladocera and 5 species of Copepoda were collected in Round Pond. The maximum standing stocks were similar in the two localities. The dominant forms in the summer and fall in La Manche Marsh were Cladocera: Acantholeberis curvirostris, Biapertura intermedia, Chydorus sphaericus, and Ilyocryptus spinifer. In the winter the copepod, Macrocyclops albidus, was dominant, while Cyclops varicans rubellus was the most abundant species in the spring. Round Pond also was dominated by Cladocera in the summer and fall; namely, Acroperus alonoides, Alona rustica, Alonella excisa, Chydorus sphaericus, and Sida crystallina. The copepods, Eucyclops agilis and Macrocyclops albidus, were dominant in the winter and early spring.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Plant community structure determines primary productivity in shallow,eutrophic lakes

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Phytoplankton productivity in a pond of brownish-colored water in a Japanese lowland marsh, Naka-ikemi

To understand the characteristics of the ecosystem in Japanese lowland marsh, we investigated chlorophyll-a (Chl. a), photosynthesis and respiration of a phytoplankton community in a brownish-colored pond in Naka-ikemi marsh, Tsuruga, Fukui Prefecture. Chl. a concentrations and volumetric gross primary production rates ranged between 1.3–57.0 μg Chl. a l⁻¹ and 148–1619 μg C l⁻¹ day⁻¹ during the study period. Higher values of Chl. a and primary production rates were clearly observed from June to September, when the dominant algae were the phytoflagellates, Peridinium (Dinophyceae) and Cryptomonas (Cryptophyceae), with swimming ability. The trophic status of the pond water of Naka-ikemi marsh was defined as being in eutrophic condition based on the biomass and productivity of phytoplankton. However, depths of Z _1% showing the productive layer in this study site were relatively narrower than those observed in the hyper-eutrophic Lake Suwa with frequent cyanobacterial water bloom. Factor-attenuating underwater light intensity in Naka-ikemi marsh was presumed to be colored dissolved organic matter. Thus, not only phytoplankton primary production, but also allochthonous organic matter supplied from the catchment area seems to be the dominant factor in the whole energy budget of the pond. In conclusion, we regarded the pond ecosystem in Naka-ikemi marsh to be in a eutrophic–dystrophic condition.     

Eutrophication and black swan (Cygnus atratus Latham) populations: tests of two simple relationships

In an earlier study, regression models were developed which related black swan populations to macrophyte biomass and to phytoplankton productivity in Tomahawk Lagoon No 2, a small South Island, New Zealand, lake in which phytoplankton and macrophytes vary widely from year to year, in an irregular inverse cycle. Two years of further study confirmed that winter swan populations are directly correlated with macrophytes, for samples taken on a single day in winter each year. They also confirmed that winter maximum swan populations can be predicted from measurements of phytoplankton productivity in the previous summer. This relationship is inverse. For 6 other lakes, 9 of the 11 data points lay within the 95% confidence intervals of the macrophyte-swan relationship, with the only outlier being a lake in which the macrophytic vegetation is dominated by filamentous algae. Further analysis, giving equal weight to each of the 7 lakes in the sample, confirmed that there is a significant correlation in winter between swan populations and macrophyte biomass for these lakes which show a wide range of size and trophic status. There was no such significant correlation in summer. All of the 8 data points from 7 other lakes lay within the rather broad 95% confidence intervals of the phytoplankton- swan relationship. The 4 lakes which fitted most closely to the predictions of this relationship have silty sediment. The other 3, which all had lower swan populations than predicted, have sediment predominantly of sand or fine clay, and macrophyte biomass and swan populations may be constrained by the nature of the sediment, rather than by shading effects from phytoplankton.     

Variability of the microbial community in the western Antarctic Peninsula from late fall to spring during a low ice cover year

Although winter conditions play a major role in determining the productivity of the western Antarctic Peninsula (WAP) waters for the following spring and summer, a few studies have dealt with the seasonal variability of microorganisms in the WAP in winter. Moreover, because of regional warming, sea-ice retreat is happening earlier in spring, at the onset of the production season. In this context, this study describes the dynamics of the marine microbial community in the Melchior Archipelago (WAP) from fall to spring 2006. Samples were collected monthly to biweekly at four depths from the surface to the aphotic layer. The abundance and carbon content of bacteria, phytoplankton and microzooplankton were analyzed using flow cytometry and inverted microscopy, and bacterial richness was examined by PCR–DGGE. As expected, due to the extreme environmental conditions, the microbial community abundance and biomass were low in fall and winter. Bacterial abundance ranged from 1.2 to 2.8 × 10⁵ cells ml^?1 showing a slight increase in spring. Phytoplankton biomass was low and dominated by small cells (<2 μm) in fall and winter (average chlorophyll a concentration, Chl-a, of, respectively, 0.3 and 0.13 μg l^?1). Phytoplankton biomass increased in spring (Chl-a up to 1.13 μg l^?1), and, despite potentially adequate growth conditions, this rise was small and phytoplankton was still dominated by small cells (2–20 μm). In addition, the early disappearing of sea-ice in spring 2006 let the surface water exposed to ultraviolet B radiations (UVBR, 280–320 nm), which seemed to have a negative impact on the microbial community in surface waters.     

Farm pond restoration using Chara vulgaris vegetation

Four aged Madison County, New York farm ponds were selected to see if various treatments could be used to restore the water quality. One pond was untreated and used as a control; another pond was partially drained and exposed to the drying and oxidizing effects of the air over the fall and winter; the other two ponds were drained and the accumulated sediment removed by bulldozing. In these latter two ponds, Chara vulgaris vegetation was inoculated following the restoration process. C. vulgaris growth rapidly became the dominant producer where this inoculation was accomplished in the fall of 1976, and it is expected that the other pond will also become a C. vulgaris pond in 1978 — after its oogonia have undergone the requisite winter dormancy period.Early C. vulgaris growth was found to be associated with clear water conditions and lessened phytoplankton growth; short, bushy, light-inhibited growth by the algae stabilized the bottom against wind-caused turbidity because of its rhizoidal growth within the substrate. Pioneer C. vulgaris growth was also found to be highly productive, significantly lowering the pond's CO₂ readings.Investigators of aquatic systems are cautioned to be cognizant of the effect of epiphytic growth on successional events in such environments. Such epiphytes are surely important, if not prime, causes of the demise of various aquatic macrophytes.The partial draining and exposing of a pond over the fall and winter did not yield significantly improved water conditions.     

BIOMASS,PRODUCTION, AND LITTERFALL IN THE COASTAL SAGE SCRUB OF SOUTHERN CALIFORNIA

A 22-year-old stand of coastal sage scrub in the coastal mountains of southern California had a peak standing aboveground biomass of 1,417 g/m², determined by dimension analysis. Annual aboveground net primary production was 255 g/m²/yr, determined by monthly twig harvests of dominant species and the clipping production of subordinate species. The stand was codominated by two drought-deciduous species, Salvia leucophylla and Artemisia californica, which together comprised 81% of the biomass. Annual litterfall was measured at 194 g/m²/yr. These biomass, production, and litterfall values are less than those measured in most evergreen chaparral communities in California. Seasonally, the two dominant shrubs began aboveground production in the winter soon after the first rains and continued growth for six months until early summer. A massive leaf fall occurred in May–June as the summer drought began, but twig and inflorescence production for both species continued at a high rate into the summer months. Salvia leucophylla had two shoot types: 1) an early spring canopy shoot that elongated rapidly, produced the inflorescence, and died in mid-summer; and 2) a short side shoot produced in late spring with small dense leaves that were retained during the summer drought and early winter. Artemisia californica produced a single cohort of twigs in the early spring, most of which carried inflorescences by late summer.     

Impact of fish predation on cladoceran body weight distribution and zooplankton grazing in lakes during winter

1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter. 2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia). 3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte‐rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish‐biomanipulated lakes during the winter season. 4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio. 5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.     

Limnological studies of the lakes and streams of the upper Qu'Appelle River System,Saskatchewan, Canada

High phytoplankton productivity characterizes the eutrophic lakes of the upper Qu'Appelle River system. Annual primary production varied from 187 to 561 g C m^? while daily areal production varied from 290 to 8 575 mg C m^?2. The A_max range was 164 to 315 mg C m^?3h^?1 with the highest rates usually occurring in August or September. A_max values on any given day normally occurred in the top 0.5 m. Blue-green algae (Aphanizomenon flos-aquae, Microcystis aeruginosa, Oscillatoria prolifica) dominated the phytoplankton communities during the summer and fall. Diatoms (Asterionella formosa, Fragilaria capucina, Stephanodiscus niagarae) often dominated the spring communities but sometimes persisted as dominants through the summer. Fragilaria on occasion was present at very high concentrations in the late fall. Ceratium hirundinella was sometimes dominant but was usually an important part of the biomass. Green algae, although always present, rarely formed an important part of the biomass with Pediastrum duplex the only exception. A species list is appended.     

Primary production of aquatic macrophytes and their epiphytes in two shallow lakes (Peipsi and Võrtsjärv) in Estonia

In shallow lakes with large littoral zones, epiphytes and submerged macrophytes can make an important contribution to the total annual primary production. We investigated the primary production (PP) of phytoplankton, submerged macrophytes, and their epiphytes, from June to August 2005, in two large shallow lakes. The production of pelagic and littoral phytoplankton and of the dominant submerged macrophytes in the littoral zone (Potamogeton perfoliatus in Lake Peipsi and P. perfoliatus and Myriopyllum spicatum in Lake Võrtsjärv) and of their epiphytes was measured using a modified ¹⁴C method. The total PP of the submerged macrophyte area was similar in both lakes: 12.4 g C m^?2 day^?1 in Peipsi and 12.0 g C m^?2 day^?1 in Võrtsjärv. In Peipsi, 84.2% of this production was accounted for by macrophytes, while the shares of phytoplankton and epiphytes were low (15.6 and 0.16%, respectively). In Võrtsjärv, macrophytes contributed 58%, phytoplankton 41.9% and epiphytes 0.1% of the PP in the submerged macrophyte area. Epiphyte production in both lakes was very low in comparison with that of phytoplankton and macrophytes: 0.01, 5.04, and 6.97 g C m^?2 day^?1, respectively, in Võrtsjärv, and 0.02, 1.93, and 10.5 g C m^?2 day^?1, respectively, in Peipsi. The PP of the littoral area contributed 10% of the total summer PP of Lake Peipsi sensu stricto and 35.5% of the total summer PP of Lake Võrtsjärv.     

Zooplankton allochthony is spatially heterogeneous in a boreal lake

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Seasonal Development of Phytoplankton in Fish Ponds

At the fish ponds under study the authors defined several types of plankton which have been frequently found during the season (April – October). These types are: Early-spring maximum of phytoplankton, Depression of phytoplankton, Bloom of Aphanizomenon, Maximum of Chlorococcales. The periods of “depression” seem to be typical for the managed carp ponds in the spring. They are characterized by the low density of rapidly reproducing algal populations (e. g. Cryptomonas) and by the dense populations of large cladocerans of the genus Daphnia. Chlorophyll in phytoplankton is less than 5μg/l, transparency is higher than 2 meters. Periods of the spring depression may be followed by the maxima of either Aphanizomenon or Chlorococcales, with concentrations of chlorophyll increasing to 100 μg/l and more. The change from the phase of “depression” to the “maximum of Chlorococcales” is accompanied by decrease in numbers of Daphnia and increase in numbers of the small cladoceran species, but all the mechanisms responsible for the transition are not yet fully understood.     

Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

Remote sensing of phytoplankton-macrophyte coexistence in shallow hypereutrophic fluvial lakes

We investigated with remote sensing (APEX images) the coexistence of phytoplankton and macrophytes in three interconnected shallow and hypereutrophic fluvial lakes (Mantua Lakes, Northern Italy). High concentrations of chlorophyll-a, up to 60 mg m^?3, were determined in the open water between well-developed stands of floating-leaved, submerged, and emergent macrophytes. Our data suggest a general inhibition of phytoplankton by macrophytes, evidenced by decreasing chlorophyll-a concentrations in proximity of macrophyte stands. Chlorophyll-a concentrations halved in the proximity of emergent stands (~6 mg m^?3 within 21 m from the stand border) when compared to the outer zones (~13 mg m^?3). Contrasting trends were observed for submerged stands, where concentrations decreased inwards from ~8 to ~3 mg m^?3. Floating leaved stands had a neutral effect, chlorophyll-a being nearly constant in both inner and outer zones. Overall, remotely-sensed data allow evaluation of quantitative and spatially defined interactions of macrophytes and phytoplankton at the whole ecosystem scale.