Microbial food web in a large shallow lake (Lake Balaton, Hungary)

Seasonal variations of phyto-, bacterio- and colourless flagellate plankton were followed across a year in the large shallow Lake Balaton (Hungary). Yearly average chlorophyll-a concentration was 11 µg 1^–1, while the corresponding values of bacterioplankton and heterotrophic nanoflagellate (HNF) plankton biomass (fresh weight) were 0.24 mg 1^–1 and 0.35 mg 1^–1, respectively. About half of planktonic primary production was channelled through bacterioplankton on the yearly basis. However, there was no significant correlation between phytoplankton biomass and bacterial abundance. Bacterial specific growth rates were in the range of 0.009 and 0.09 h^–1, and ended to follow the seasonal changes in water temperature. In some periods of the year, predator-prey relationships between the HNF and bacterial abundance were obvious. The estimated HNF grazing on bacteria varied between 3% and 227% of the daily bacterial production. On an annual basis, 87% of bacterial cell production was grazed by HNF plankton.     

An attempt to trace eutrophication in a shallow lake (Balaton,Hungary) using chironomids

Lake Balaton, the largest shallow lake in Central Europe, is about 20 000 years old. An enormous increase in tourism and the disproportionate building development of the last few decades has resulted in the acceleration of eutrophication in the lake. Widespread research to reveal the causes of water-quality deterioration and possible ways of protection against it have recently started. The investigation of the larvae of non-biting midges (Diptera: Chironomidae) in the sediment of the open-water zone has also begun. The contemporary faunal composition strongly correlates with the trophic gradient along the longitudinal axis of the lake. We therefore supposed that the eutrophication process should be identifiable from the analysis of subfossil chironomid head capsules from the upper (15 cm thick) layer of the sediment. We found that quantitative results could only be obtained when fragments as well as relatively intact head capsules are considered. Our data verify that the originally oligo-mesotrophic community has been gradually replaced by eutrophic species in a west to east direction. Large-bodied larvae belonging to the Chironomus plumosus group mix the sediment down to 15 cm as they build their tubes and consequently alter the original proportions of head capsules at the different levels. So the sequence of communities through the sediment-layers is not quite reliable.     

The role of storms in the summer succession of the phytoplankton community in a shallow lake (Lake Balaton, Hungary)

The authors carried out day-to-day studies on the phytoplanktonof the largest shallow lake of Central Europe in the summerof 1976, 1977 and 1978. Data were analysed with diversity andcluster analyses. (i) Wind-induced stirring-up of the sedimentplays important role in the summer phytoplankton successionof the shallow lake. (ii) After storms algae with very smallcell sizes show synchronous development. In long calm periodsalgae with larger cell sizes exhibit organised population dynamics.(iii) In calm periods a shift from r-selection to predominantK-selection can be suspected. After the next storm importanceof r-selectionists again increases. (iv) Effect of changinggrowth- and loss rates on the population dynamics of the mostabundant species is discussed based on measured generation timesand calculated elimination. The phenomena discussed in the papercan appear to different extents in the summer phytoplanktonsuccession of other shallow, polimictic lakes.     

Composition,density and feeding of crustacean zooplankton community in a shallow,temperate lake (Lake Balaton,Hungary)

Composition density and filtering rates of crustacean zooplankton were studied in the open water and among the macrophytes of the oligo-mesotrophic part of Lake Balaton from 1981 to 1983. From the individual filtering rates of the different populations and the densities community grazing rates were derived.Copepoda made up 79–90% of crustacean plankton community in the open lake and 95–97% of it in littoral zone. Among them the nauplii dominated. At the end of summer 1982, when Anabaenopsis was in bloom, the filter-feeding species (Eudiaptomus, Daphnia) practically disappeared, being replaced by cyclopoids. Daphnia had the highest filtering rates followed by those of Eudiaptomus and Diaphanosoma. Among copepods, the filtering rates in decreasing order were: ovigerous > all adults > copepodites > nauplii. > . The filtering rates of the different species varied both seasonally as well as from year to year. In 1983, when the concentration of organic seston decreased, filtering rate increased compared with those in the earlier years. During the water bloom in 1982, the rates decreased by 70% on the average.The community grazing rate was very low (3% per day) in the open lake and among macrophytes, both in 1981 and 1982; also the share of crustacean zooplankton in grazing was very low. In 1983, together with the improving of water quality, the community grazing rate increased 4-fold. In 1981 and 1983 the rates were influenced by water temperature but in 1982 by seston concentration.     

Daily egg production ofTemora longicornis (Copepoda,Calanoida) during winter and early spring in the Marsdiep (Southern North Sea)

Mature females of the copepodTemora longicornis from the Marsdiep tidal inlet were incubated at ambient water temperature during the winter of 1989/1990 to estimate their egg production during 24 hours. Throughout winter this production remained at a level of 0 to 10 eggs per female. In March a steep rise to 60 to 80 eggs coincided with an equally steep rise in chlorophyll-a from 4 to 20 g.l^–1. Excess food given during incubation tended to increase the observed production rate only in October at relatively high temperature. Additive effects of rising food level and temperature on daily egg production were found in spring. In March and April 1988 and 1990 the observed maximum daily egg production per female increased from about 20 at 5°C and 5 g.l^–1 chlorophyll to 70 at almost 10°C and 20 g.l^–1. The egg production in winter gives a prebloom potential to maintain a stock of premature stages ready to utilize the algal spring bloom. Early spawning before April can also contribute to the development of the abundance peak of adults in May–June. Due to growth rates increasing with temperature in the months March and April, most of the adults at their peak density may have hatched from eggs before the period of maximum egg production rate at the end of April. The exceptionally high production rate observed at the end of March may be related to a developingPhaeocystis bloom.     

Life history of Corophium curvispinum G. O. Sars (Crustacea,Amphipoda) living on macrophytes in Lake Balaton

Samples of the amphipod Corophium curvispinum were collected from Myriophyllum spicatum twice a month during the vegetation period. The animals were sexed, their length measured and the eggs of females were counted. Length frequency histograms served to follow the life history of C. curvispinum.Reproduction lasts from May to October. The animals born in May–June reproduce in the same year while the summer-autumn generation reproduces the following year. The life span is about 1 year. The highest fecundity was found in June, the lowest in October.     

Phytoplankton nitrogen demand and the significance of internal and external nitrogen sources in a large shallow lake (Lake Balaton,Hungary)

Since the middle of 1990s the trend of Lake Balaton towards an increasingly trophic status has been reversed, but N₂-fixing cyanobacteria are occasionally dominant, endangering water quality in summer. The sources of nitrogen and its uptake by growing phytoplankton were therefore studied. Experiments were carried out on samples collected from the middle of the Eastern (Siófok) and Western (Keszthely) basins between February and October 2001. Ammonium, urea and nitrate uptake and ammonium regeneration were measured in the upper 5-cm layer of sediment using the ¹⁵N-technique. Ammonium was determined by an improved microdiffusion assay. N₂ fixation rates were measured by the acetylene-reduction method. Ammonium regeneration rates in the sediment were similar in the two basins. They were relatively low in winter (0.13 and 0.16 μg N cm^?3 day^?1 in the Eastern and Western basin, respectively), increased slowly in the spring (0.38 and 0.45 μg N cm^?3 day^?1) and peaked in late summer (0.82 and 1.29 μg N cm^?3 day^?1, respectively). Ammonium uptake was predominant in spring in the Eastern basin and in summer in the Western basin, coincident with the cyanobacterial bloom. The amount of N₂ fixed was less than one third of the internal load during summer when external N loading was insignificant. Potentially, the phytoplankton N demand could be supported entirely by the internal N load via ammonium regeneration in the water column and sediment. However, the quantity of N from ammonium regeneration in the upper layer of sediment combined with that from the water column would limit the standing phytoplankton crop in spring in both basins and in late summer in the Western basin, especially when the algal biomass increases suddenly.     

Stir-up effect of wind on a more-or-less stratified shallow lake phytoplankton community,Lake Balaton,Hungary

Microstratification of phytoplankton in the large shallow Lake Balaton (Hungary) was studied during a 24 h period. Dissolved O₂ showed biological stratification; flagellates exhibited a definite circadian rhythm. In the middle of the investigation a heavy storm broke out which was followed by the disappearance of differences between different layers of water. Storm-induced destratification is described by cluster-analysis. Abundances of dominant species changed differently in connection with the storm. Numbers of Nitzschia sp. increased due to stirring up from the sediment surface. Numbers of single-celled or colony-forming species (Cyclotella comta, Crucigenia quadrata, Coelosphaerium kuetzingianum) practically did not change. Numbers of all the three dominant filamentous species (Aphanizomenon fos-aquae f. klebahnii, Lyngbya limnetica, Planctonema lauterbornii) significantly decreased, which might be attributed to an unknown loss process and was followed by a competitive displacement by algae of small cell size.     

Mesozooplankton assemblage and first record of Paracartia grani Sars G.O., 1904 (Copepoda: Calanoida) in the Western harbour of Genova (Ligurian Sea).

Harbours are characterized by high pollutant charge and by the occurrence of well adapted and resistant species. This paper reports the results of an annual plankton survey (May 1997-June 1998) carried out in the western harbour of Genova (Ligurian Sea) and in its mouth. Plankton samples were collected by horizontal trawls using a WP2 net. Copepods were the bulk of plankton in almost all samples. Eight copepod species were recognized: Paracalanus parvus and Acartia clausi were the most abundant. The first record of Paracartia grani in the harbour of Genova is here reported; this species, which is known to occur in polluted harbour waters of the Mediterranean Sea and was found in semi-confined Mediterranean and Atlantic coastal areas, was dominant during October 1997. Also Clausocalanus spp., Centropages typicus, Oithona helgolandica, Oithona nana, Farranula spp., Eurytemora spp., Isias clavipes and Lucicutia spp. were frequently sampled. Among other zooplankters, cladocerans, ostracods and tunicates occurred frequently, while cnidarians, mysids and chaetognaths showed low densities. These results show the occurrence of a well defined harbour plankton and point out the differences between harbour and neritic plankton of the Gulf of Genova.     

Physiology and population structure of Calanus finmarchicus (Copepoda: Calanoida) during a Lagrangian tracer release experiment in the North Atlantic

A population of Calanus finmarchicus was followed in the opensea for a period of 10 days at the end of June 1996 using atracer release method. Gut fluorescence, egg production, carboncontent and stage specific abundance, together with phytoplanktonconcentration and composition, were measured during this period.Chlorophyll levels were less than 1 µg l^–1 and thephytoplankton population was dominated by coccolithophorids.Gut fluorescence decreased during a period with strong winds,whereas egg production remained constant. Estimated phytoplanktoningestion was too low to cover egg production requirements.However, the microzooplankton concentration seemed to be highenough to complement phytoplankton ingestion, enabling egg productioncarbon requirements to be covered. C/N ratios of adult femalesand stage CV were low, indicating low levels of lipid storage.The population was dominated by stage CIV, but shifted to adominance of younger stages during the study. Mortality calculatedfrom population structure indicates that higher death ratesoccurred at stage CV.     

Parasitic regulation in a population of Boeckella hamata Brehm (Copepoda: Calanoida)

SUMMARY. 1. The impact of parasitism by a fungus Aphanomyces sp. (Phycomycetes: Saprotegniaceae) on a population of Boeckella hamata in Lake Mahinerangi in New Zealand was studied over 20 months. The incidence of infection among adult female copepods exceeded 12% throughout winter (May-October) and was highest in autumn (46%) when population birth rates were depressed by 48.8%. During most of the study, parasitism lowered the rates of population recruitment and growth by more than 10%. These effects on the B. hamata population exceeded those of Aphanomyces on Boeckella dilatata Sars in another study.
2. It is hypothesized that the high incidence of parasitism by Aphanomyces and its importance at times in regulating populations of Boeckella in New Zealand lakes may be an ecological consequence of life in predator-poor ecosystems.     

Diversity patterns of trait-based phytoplankton functional groups in two basins of a large, shallow lake (Lake Balaton, Hungary) with different trophic state

The application of functional approaches in understanding phytoplankton community-level responses to changes in the environment has become increasingly widespread in recent years. Eutrophication is known to cause profound modifications in ecosystem processes; however, the underlying relationships between environmental drivers and phytoplankton diversity and functioning are complex and largely unknown. Therefore, in the present study, we investigated and compared the temporal diversity patterns of phytoplankton functional groups in the mesotrophic eastern and eutrophic western basin of the shallow Lake Balaton situated in Hungary. Diversity data were derived from taxonomic composition and biomass data corresponding to the years 2005–2006 and 2008–2009. With the use of cluster analysis, phytoplankton species were classified into eight distinct groups representing different combinations of functionally relevant traits including greatest axial linear dimension; surface-to-volume ratio; photosynthetic pigment composition; N₂ fixation; phagotrophic potential; growth form/complexity (unicell, filamentous, colony-, or coenobium-forming); and motility. Our results have revealed that there is a significant inverse relationship between the functional group diversity used in our study and trophic state (total phytoplankton biomass) as opposed to species diversity, where no correlation was observed. In addition, group evenness showed an even stronger negative correlation with trophic state, while species evenness yielded only a weak relationship. The observed variability in functional group diversity suggests that such an approach could provide an efficient means of revealing structural changes in phytoplankton communities, establishing new hypotheses and highlighting fundamental points in ecosystem functioning.     

Distribution of submerged macrophytes along environmental gradients in large,shallow Lake Balaton (Hungary)

Distribution of submerged macrophytes was assessed at 15 littoral sites in large, shallow Lake Balaton using the echo sounding technique. Sites represented the conspicuous longitudinal and transverse environmental gradients. Absence/presence of plants, water depth, bottom consistency and distance from the shore have been derived from echo sounding data. Turbidity and meteorological variables were recorded at a single site from 2004 to 2006. Site-specific variability in underwater light conditions (an indicator of stress) and exposure to breaking waves (an indicator of disturbance) were estimated by simple models.     

Long-term changes in the population of Eurytemora affinis (Copepoda,Calanoida) in the Gironde estuary (1978–1992)

Long-term changes in the population of Eurytemora affinis were investigated during 15 years (1978–1992) at a fixed station in the Gironde estuary (South West France). Total numbers, sex-ratio, % ovigerous females, proportion of copepodites and clutch-size were taken into account and their long-term distribution was related to temperature, river flow, salinity, suspended matter and chlorophyll a concentrations after the seasonal effect was removed for all variables.There was a great decrease of the river flow during the period of investigation, due to a general deficit in pluviosity. On the contrary, the general trend for salinity was an increase from 1978 to 1992. The same pattern was observed for chlorophyll, indicating an intrusion of marine phytoplankton. Water temperature increased significantly during the study period, following the general increase in air temperature observed in the area. For the suspended particulate matter (SPM), an increase was observed between 1978 and 1981–1982, then a very sharp decrease occurred from 1984 onwards.Numbers of E. affinis were inversely correlated with temperature and salinity, and positively correlated with the river flow. E. affinis was negatively correlated with chlorophyll concentration because of the covariation with salinity. No clear long-term trend was observed for the sex-ratio. A significant correlation was found between females carrying egg-sacs and SPM concentration, probably due to a decrease of the predation pressure in very turbid waters. The percentages of copepodites tended to decrease with time and were inversely correlated with temperature. Clutch-size significantly decreased during the 15-year period. This trend was mainly explained by temperature and salinity.It is concluded that, in the absence of strong human alteration in the Gironde estuary the long-term distribution of zooplankton can be explained by the natural environmental variability. The dominant factors are the river flow which governs the movement of the populations along the estuary and the temperature which influences the reproductive processes. Any change in climate will have consequences on the pelagic community.     

Gladioferens Henry (Copepoda: Calanoida) discovered in Antarctica: G. antarcticus sp. nov. described from a lake in the Bunger Hills

A new species of Gladioferens is described from a freshwater lake in the Bunger Hills, Greater Antarctica. A key to the six species of Gladioferens is provided for both sexes. Most species of the subgenus containing G. antarcticus sp. nov. are remarkable in possessing asymmetries in the second legs (male) and fourth legs (female). This suggests that the mating behaviour of these species may be radically different from that in most heterarthrandrian calanoids and calls for further study. However, G. antarcticus is more primitive than its consubgeners in having, in the male, symmetrical second legs and 3-segmented endopods in the fifth legs. The genus shows a highly disjunct distribution (Australia, New Zealand and Bunger Hills) and its biogeography is now of outstanding interest.     

Feeding of Boeckella gracilipes (Copepoda, Calanoida) on ciliates and phytoflagellates in an ultraoligotrophic Andean lake

The calanoid copepod, Boeckella gracilipes, is the dominantcrustacean zooplankton in South Andean deep ultra-oligotrophiclakes. Combining field and experimental data we explored thefeeding of the copepod and its access to the mixotrophic ciliate,Ophrydium naumanni, in Lake Moreno Oeste (Patagonia, Argentina).Phytoplankton was dominated by nanoflagellates throughout thewater column. Ophrydium naumanni, which accumulates much ofthe chlorophyll a, as do copepodites and adults of B. gracilipes,has a deep distribution during the day, with maximal abundancesaround 30 m depth. Mouth-part morphology analysis of B. gracilipesindicated that the copepod has an omnivorous diet. Laboratoryexperiments showed that B. gracilipes could access O. naumannionly when it is offered as a single food item. However, whennatural phytoplankton and ciliate assemblages (including O.naumanni)are offered, B. gracilipes did not eat Ophrydium and preyedon the oligotrich, Strombidium viride, and phytoflagellateslike Chrysochromulina parva. The range of ingested sizes wasbroad (3.9–33 µm of equivalent spherical diameter)but all selected particles were motile ones with distinctivemovements, which would enhance the copepod particle detection.     

Meroplankton dynamics in a saline,turbulent, turbid shallow lake (Neusiedlersee,Austria and Hungary)

Phytoplankton species composition, horizontal distribution, seasonal- and long-term dynamics are investigated in relation to some environmental factors, based on about 700 samples taken between 1968 and 1992 in the shallow, turbid, turbulent, saline Neusiedlersee (Austria/Hungary). We deduced:

1. The phytoplankton is relatively poor in species which is attributable to high salinity and turbidity. Blue-green algae of picoalgal size, meroplanktonic diatoms, green algae either with gelée or elongated form are the most important groups in the plankton;
2. No consequent horizontal differences in the distribution of phytoplankton biomass were found; the high contribution of diatoms (69%) and green algae (22%) is characteristic;
3. Despite the low percentage contribution (0.54%) of phytoplankton dry weight in total seston, the two variables correlate closely because of simultaneous resuspension after wind actions. The prevalence of both low phytoplankton/seston ratio and significant correlation between these two variables is indicative to the presence of a meroplankton;
4. Phytoplankton biomass, especially that of diatoms, varies within wide limits in the short term as a combined effect of growth/loss, transport by horizontal water currents and periodic resuspension from the sediment surface. Diatoms like Fragilaria, Surirella and Campylodiscus are involved especially. Parameters of population dynamics (growth rate, annual cycles, length of stationary phases, standing crop) were estimated based on moving averages. Growth rates were smaller than those of ‘normal’ planktonic species; however, the similarities of seasonal pattern suggest that the carrying capacity of the lake is very constant;
5. Picoalgal biomass is very high in the lake; its contribution to total biomass can exceed 75%, especially in spring. The large cellular chlorophyll a content of Neusiedlersee's phytoplankton can most probably be explained by this high picoalgal standing crop, which is not included in the routine biomass estimations;
6. Concerning long-term changes of phytoplankton, periodic appearances and disappearances are very characteristic. Annual average biomass of most species is growing for several (3–5) years, then declining with a similar rate and these periods are recurrent. Climatic drying-out periods, during which water level lowers, conductivity increases, nutrients and their ratios change following trend-like periodicities, and they are supposed to be responsible for the observed longterm periodicity of dominant species. An increased nutrient load in the seventies was superimposed on this cyclicity;
7. Most of the species can be found in smaller or larger amounts in the littoral microhabitats (canals, inner ponds) of the lake in periods when they are absent or very rare in the open water. Thus, these littoral microhabitats play a very important role in the survival and recruitment of planktonic populations to the open water.