Tõnu Feldmann

Dynamic carbon budget of a large shallow lake assessed by a mass balance approach

To study the role of large and shallow hemiboreal lakes in carbon processing, we calculated a 3-year carbon mass balance for Lake Võrtsjärv (Estonia) based on in situ measurements. This balance took into account hydrological and biogeochemical processes affecting dissolved inorganic (DIC), dissolved organic (DOC) and particulate organic (POC) carbon species. Accumulation varied greatly on a seasonal and yearly basis. The lake exported carbon during most of the year except during spring floods and in late autumn. In-lake processes were responsible for exporting POC and storing DOC while DIC switched between storage and export. The carbon cycle was alternatively dominated in 2009 by biogeochemical processes and in 2011 by riverine fluxes, whereas in 2010 the two process types were of the same magnitude. These results suggest that the role of large shallow lakes like Võrtsjärv in the global C cycle is equally driven by hydrological factors, in particular seasonal water level changes, and by biogeochemical in-lake reactions.

Benthic foodweb structure in a large shallow lake studied by stable isotope analysis

Abstract: The benthic foodweb structure of Lake Võrtsjärv, a large (270 km2), shallow, and turbid Estonian lake, was evaluated based on C and N stable-isotope signatures (&dgr;13C, &dgr;15N). Variation in &dgr;13C between sampling sites was not related to site proximity to the littoral zone or the more vegetated southern part of the lake, but rather appeared to be influenced by in-situ site peculiarities. &dgr;13C was stable temporally and between functional feeding groups, a result implying that the whole benthic food web of the lake relies largely on the same C source admixture, essentially particulate organic matter (POM). Thus, the foodweb composition of Lake Võrtsjärv is remarkably homogeneous given the lakes large surface area. Apparent trophic-level &dgr;15N fractionation between total collectors and total predators (mean 1.7‰) was lower than the value of 3.4‰ generally adopted in foodweb studies, but the higher value was valid for specific prey—predator links. The low &dgr;13C signature of some chironomid samples indicated probable assimilation of methane-oxidizing bacteria (MOB) by these sediment-dwelling invertebrates. However, the lack of similar 13C depletion in benthic filterers (mussels) indicated that the MOB layer is essentially confined to the sediments and does not reach the water column, which probably constrains transfer of methane-derived C through the food web to fish in this lake. Our study demonstrates that the benthic food web of shallow turbid lakes like Võrtsjärv is simplified and is mostly sustained by phytoplanktonic C sources.

From bacteria to piscivorous fish: estimates of whole-lake and component-specific metabolism with an ecosystem approach.

The influence of functional group specific production and respiration patterns on a lakes metabolic balance remains poorly investigated to date compared to whole-system estimates of metabolism. We employed a summed component ecosystem approach for assessing lake-wide and functional group-specific metabolism (gross primary production (GPP) and respiration (R)) in shallow and eutrophic Lake Võrtsjärv in central Estonia during three years. Eleven functional groups were considered: piscivorous and benthivorous fish; phyto-, bacterio-, proto- and metazooplankton; benthic macroinvertebrates, bacteria and ciliates; macrophytes and their associated epiphytes. Metabolism of these groups was assessed by allometric equations coupled with daily records of temperature and hydrology of the lake and measurements of food web functional groups biomass. Results revealed that heterotrophy dominated most of the year, with a short autotrophic period observed in late spring. Most of the metabolism of the lake could be attributed to planktonic functional groups, with phytoplankton contributing the highest share (90% of GPP and 43% of R). A surge of protozooplankton and bacterioplankton populations forming the microbial loop caused the shift from auto- to heterotrophy in midsummer. Conversely, the benthic functional groups had overall a very small contribution to lake metabolism. We validated our ecosystem approach by comparing the GPP and R with those calculated from O2 measurements in the lake. Our findings are also in line with earlier productivity studies made with 14C or chlorophyll a (chl-a) based equations. Ideally, the ecosystem approach should be combined with diel O2 approach for investigating critical periods of metabolism shifts caused by dynamics in food-web processes.

Factors influencing macrophyte metrics in Estonian coastal lakes in the light of ecological status assessment

Macrophytes and factors affecting their distribution were studied in 19 coastal lakes of Estonia. The aim of the study was to determine the factors influencing the distribution of macrophytes in coastal lakes and to assess the suitability of valid macrophyte metrics. Our hypothesis was that in coastal lakes most of the macrophyte distribution patterns are caused by lake-specific variables. Morphological, physico-chemical and catchment area characteristics of the lakes varied greatly. Lakes were in different development stages—lakes nearest to the Baltic Sea were younger and more influenced by brackish water and the furthest lakes were older with more freshwater. All that variability was reflected in macrophyte parameters. Factor analysis of environmental indices divided them into three groups—catchment area, morphometric and water chemistry factors. The first factor may be considered as a pressure and the other two as lake-type-specific factors. Lake catchment area parameters had an influence on Bolboschoenus maritimus, Chara tomentosa and Typha latifolia abundance. Morphometric parameters had an influence on the depth distribution of macrophytes and water chemistry factors on the abundance of helophytes. Current indicator species showed more variability associated with lake-specific factors than with changes in status or the influence of pressures.

Ciliate communities of a large shallow lake: association with macrophyte beds.

We investigated the influence of macrophyte composition on ciliate community structure in a large, shallow, eutrophic Lake Võrtsjärv. We hypothesized that macrophyte composition must have strong influence on the dispersal of ecologically different ciliate groups in a shallow lake and that more diverse macrophyte stands cause also a greater diversity in the ciliate community. In Võrtsjärv macrophyte distribution is spatially strongly polarized both in east-west and north-south directions in relation to abiotic factors. Phragmites australis and Myriophyllum spicatum were the most widespread species occurring in most parts of the lake. Correlation of environmental, macrophyte and planktonic ciliate variables confirmed the suggested spatial gradients. More diverse macrophyte stands supported a high species richness and abundance of epiplanktonic community but showed negative influence on the number and abundance of euplanktonic ciliate taxa. Opposite trends were found relative to the abundance of P. australis. Benthic ciliates showed a similar distribution pattern to euplanktonic taxa being most abundant in sites were the Shannon-Weaver index for macrophytes was low. Strong polarizing effect of the lakes vegetation on planktonic ciliate diversity was reflected in correlations of the number of ciliate taxa as well as the numbers of eu- and epiplanktonic taxa with geographic co-ordinates.

The influence of zooplankton enrichment on the microbial loop in a shallow, eutrophic lake.

With increasing primary productivity, ciliates may become the most important members of the microbial loop and form a central linkage in the transformation of microbial production to upper trophic levels. How metazooplankters, especially copepods, regulate ciliate community structure in shallow eutrophic waters is not completely clear. We carried out mesocosm experiments with different cyclopoid copepod enrichments in a shallow eutrophic lake to examine the responses of ciliate community structure and abundance to changes in cyclopoid copepod biomass and to detect any cascading effects on bacterioplankton and edible phytoplankton. Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by the decline of predaceous ciliate abundance, which would otherwise graze effectively on the small-sized ciliates. The inverse relationship between crustacean zooplankton and large predaceous ciliates is an important feature adjusting not only the structure of the ciliate community but also the energy transfer between meta- and protozooplankton. Still we could not detect any cascading effects on bacterio- or phytoplankton that would be caused by the structural changes in the ciliate community.

Macrophytes in relation to ecological factors in a lowland river in Estonia

Compared to lakes an d the coastal sea, the macroflora o f rivers in Estonia has lacked investigation; nevertheless, the floristic list ofmacrophytes collected from 396 reaches on 150 Estonian rivers and streams includes 124 taxa ofvascular plants, 24 taxa of mosses, and 34 taxa of macroalgae (JARVEKÜLG 2001). Because vascular plants in rivers are mostly perennial, constant occurrence of the same species in the same river reach would be expected; however, we observed changes in the floristic composition, number of species, coverage, and composition of the dominating species visiting the same sites over years during the monitoring of rivers. Recent studies (PAAL & TREI 2004, 2006, TREI & PALL 2004) and monitoring data have shown that changes in the macroflora often proceed in different directions species disappeared and reappeared on the same site over years. Trends have not been evident, however, and the causes of the changes remain unclear. The aims of this study were (l) to establish the floristic composition and coverage of macrophytes in a lowland river over 3 years; (2) to monitor changes in species composition and coverage along the river from year to year; and (3) to determine the environmental factors governing the distribution o f macrophytes in an Estonian river.