Kimmo T. Tolonen

History of heavy metal pollution in Finland as recorded by lake sediments

Abstract Profiles of heavy metals and major metals were determined in sediment cores from 18 small, headwater drainage and seepage lakes in Finland. More than half of the lakes have recently been acidified. The results show an increase of Pb deposition starting in the early 1800s. Most of the Pb profiles show a levelling off or slight decrease in the deposition rate in the post-1980 sediments. The first signs of increases of Zn, Cd and Hg typically appear during the 1800s; a prominent increase in the sedimentation of these elements is typically confined to this century. Clear subsurface (2–5 cm) concentration maxima of Zn and Cd were observed in cores from many lakes in south Finland. The maxima are not related to changes in emission and atmospheric deposition. Subtraction of the background fluxes from the total fluxes of the 1980s gives the following percentages for atmospheric deposition: Pb, 74–97% of the total; Cd, 62–92%; Zn, 39–91%; and Hg, 70–89%. These figures apply to southern and central Finland, areas that also have the highest deposition of acidic compounds. In northern Finland the increase in deposition starts later and has remained smaller than in the south. About half of the profiles of Cu and V and a third of the Ni profiles showed increased accumulation, typically after 1950. Recent increases in the concentration and accumulation rates of Na, K and Mg, observed in most cores in southernmost Finland, may also be of atmospheric origin. This is suggested by the chemistry of bulk deposition.

Littoral species diversity and biomass: concordance among organismal groups and the effects of environmental variables

Abstract.Biomass and species diversity (richness and evenness) of littoral organisms were explored in 27 sites in three basins of the large Lake Saimaa system in eastern Finland. The basins differed in degree of nutrient loading and trophic status. Six organismal groups, i.e., phytoplankton, periphyton, macrophytes, crustacean zooplankton, benthic macroinvertebrates and fish were studied. Factors affecting the biomass and diversity of these groups were explored by multiple stepwise regression analysis. The biomass of different groups was explained by the same variables, mainly nutrients, while diversity was associated with different environmental factors among the studied groups. The biomass of periphyton, phytoplankton, zooplankton, and fish correlated significantly with each other. There was also an apparent association between the biomass of macrophytes and that of benthic invertebrates. However, no significant correlations were found among the diversity of the studied groups. In accordance with previous studies, our results did not support the existence of species-rich hotspots or the possibility of using any surrogate taxon to reveal overall biodiversity. Thus, for conservation planning, biological surveys should include extensive collection of taxonomic groups and organisms at all trophic levels.

Indicator groups and congruence of assemblage similarity, species richness and environmental relationships in littoral macroinvertebrates

The rapid decrease of biodiversity and limited resources for surveying it have forced researchers to devise short-cuts for biodiversity surveys and conservation planning. These short-cuts include environmental surrogates, higher taxon surrogates, indicator species and indicator groups. We considered indicator groups as surrogates for wholesale biodiversity and cross-taxon congruence in biodiversity patterns in littoral macroinvertebrates of boreal lakes. Despite the fact that we considered indicator groups amongst a wide variety of taxa, such as two-winged flies, mayflies, caddisflies, beetles, bugs and molluscs, none of the proposed groups possessed all of the qualities of a good indicator taxon for biodiversity surveys and conservation planning. We found generally weak, yet typically significant, relationships between the proposed indicator groups and remaining taxa in both species richness and assemblage similarity. Low congruence was paralleled by somewhat differing relationships of the taxonomic groups to various environmental features of lakes. Furthermore, the relationships of most indicator groups to the environmental features of lakes were not particularly strong. The present findings are unfortunate, because indicator groups did not perform well in predicting the wholesale biodiversity of littoral macroinvertebrates. Thus, there appears to be no short-cut for considering all groups of macroinvertebrates in biodiversity surveys, conservation planning and characterisation of environmental relationships of lake littoral assemblages.

Natural variation of profundal macroinvertebrate communities in boreal lakes is related to lake morphometry: implications for bioassessment

Modern biological assessment of aquatic systems is often based on the reference condition approach, which requires characterization of biota in undisturbed conditions. For this approach, it is essential to recognize the origins and degree of natural variation in communities. We used data from 55 minimally disturbed Finnish lake basins to investigate the natural variation of profundal macroinvertebrate community composition in relation to environmental factors. Partial canonical correspondence analysis showed that most (68% combined, 39% uniquely) of the total explained compositional variation (29%) was correlated with environmental variables insensitive to human activities, especially lake morphometry. In contrast, the unique contributions of geographical location and variables influenced by human activities (dissolved oxygen and total phosphorus) were substantially smaller (5.5% and 22%, respectively). All of the explained variation (38%–64%) of three widely used community metrics was also correlated wit...

Comparison of sampling methods and habitat types for detecting impacts on lake littoral macroinvertebrate assemblages along a gradient of human disturbance.

We explored environmental variables structuring littoral macroinvertebrate communities in a large lake basin along a gradient of nutrient enrichment. Furthermore, we evaluated sensitivity and cost-effi ciency of different sampling schemes (i.e. combinations of three habitat types and a number of standard sampling methods) to detect changes in macroinvertebrate communities along this anthropogenic disturbance gradient. Partial canonical ordination analysis showed that habitat characteristics accounted for a major part (56 % uniquely) of the explained variation in the species composition of invertebrate communities. When different mesohabitats were examined separately, assemblage variation of stony bottoms was most strongly associated with human-induced changes in water chemistry (51 % of the explained variation). In sandy and vegetated habitats, shore morphometry and habitat characteristics were more infl uential factors. The most time-consuming phases of sample processing were sorting and identifi cation, whilst the time-costs of fi eld sampling were relatively low. The duration of sample treatment was positively correlated with the numbers of individuals counted and taxa identifi ed. Additionally, processing time was associated with habitat type, samples from stony bottoms being the least laborious. Kick-net sampling on stony shores was the most cost-effi cient sampling scheme with low costs and highest sensitivity to detect anthropogenic impacts.

The role of benthivorous and planktivorous fish in a mesotrophic lake ecosystem

The effectiveness of fish removal as a tool for biomanipulation and restoration was studied from 1993 to 1997 in shallow Lake Pohjalampi (North Karelia, eastern Finland). The external nutrient loading into this lake was low and nearly stable throughout the study period. During a period of 5 years, a total of > 200 kg ha−1 of fish, mainly roach and bream, were caught and thus the total fish biomass was reduced by nearly 80%. However, in 1996 a very dense year-class of perch hatched and the number of planktivorous fish increased again. These changes in the fish community resulted in a decreased fish predation on benthic invertebrates, whereas the predation of fish on zooplankton increased. The responses in the prey communities were consistent with these changes: the biomass of benthic invertebrates increased and that of zooplankton decreased. At the same time, nutrient and chlorophyll a : concentrations declined. The decreasing chlorophyll a:P ratio indicated also that the nutrient cycling in the lake was reduced and despite the increased predation on herbivorous zooplankton the lake was shifted to a more oligotrophic state.

Phantom midge-based models for inferring past fish abundances

We sampled living and subfossil phantom midge (Diptera: Chaoboridae) larvae from surface sediments of 21 small lakes in Southern Sweden to examine the influence of fish and selected abiotic variables on the abundance and species composition of chaoborid assemblages. We expected total Chaoborus abundance to be inversely correlated with fish abundance and Chaoborus species most sensitive to fish predation to be found only in fishless lakes. We aimed to use the observed relationships to develop models to reconstruct past fish abundances from chaoborid remains and the abiotic environment. C. flavicans occurred in almost every lake, whereas subfossil C. obscuripes were found in the surface sediments of only one fishless lake. The density of living C. flavicans larvae correlated negatively with fish abundance, lake order and size. The concentration of C. flavicans subfossils was negatively associated with pH, lake size, water transparency and fish abundance. Regression models that included lake morphometry and landscape position as additional predictors of fish abundance performed better than models that used only Chaoborus predictors. The explained variance in fish abundance varied from 52 to 86%. Leave-one-out cross-validation indicated moderate performance of the two best models. These models explained 51 and 56% of the observed untransformed fish density and biomass, respectively. In addition, all Chaoborus models were unbiased in closely following the 1:1 reference line in plots of observed versus predicted values. These results are a promising step in developing midge-based paleolimnological reconstructions of past fish abundance, and the approach might be improved by including chironomid remains in the models.

Relative roles of spatial processes, natural factors and anthropogenic stressors in structuring a lake macroinvertebrate metacommunity

Studies of aquatic metacommunities have so far been focused almost entirely on relatively isolated systems, such as a set of streams, lakes or ponds. Here, we aimed to quantify the relative importance of spatial processes, natural factors and anthropogenic stressors in structuring of a macroinvertebrate metacommunity within a large, highly-connected shallow lake system. The roles of different drivers were evaluated for the entire metacommunity, 10 trait-based deconstructed metacommunities and four common species by incorporating extensive sampling and a large number of abiotic explanatory variables. Contrary to our expectations, we found that variation in community structure among sites was mostly correlated to spatial and wind-wave variables rather than anthropogenic disturbance factors even though the lake presented strong environmental gradients associated with long-term human pressures. In addition, the relative importance of the three groups of drivers varied slightly among the deconstructed trait matrices (i.e. based on dispersal ability, feeding mode and degree of occurrence). Importantly, the distributions of the most common species showed significant and strong spatial autocorrelation, indicating the prominent role of high dispersal rate for their distributions. These findings suggest that the influences of high dispersal rates and natural disturbance may even override the roles of anthropogenic stressors in metacommunity organization in highly-connected aquatic systems. Hence, we strongly encourage that spatial processes and natural drivers are taken into account in the development of bioassessment approaches in highly-connected aquatic systems.

Environmental filtering and spatial effects on metacommunity organisation differ among littoral macroinvertebrate groups deconstructed by biological traits

We examined spatial and environmental effects on the deconstructed assemblages of littoral macroinvertebrates within a large lake. We deconstructed assemblages by three biological trait groups: body size, dispersal mode and oviposition behaviour. We expected that spatial effects on assemblage structuring decrease and environmental effects increase with increasing body size. We also expected stronger environmental filtering and weaker spatial effect on the assemblages of flying species compared with assemblages of non-flying species. Stronger effect of environmental filtering was expected on the assemblages with species attaching eggs compared with assemblages of species with free eggs. We used redundancy analysis with variation partitioning to examine spatial and environmental effects on the deconstructed assemblages. As expected, the importance of environmental filtering increased and that of spatial effects decreased with increasing body size. Opposite to our expectations, assemblages of non-flying species were more affected by environmental conditions compared to assemblages of flying species. Concurring with our expectations, the importance of environmental filtering was higher in structuring assemblages of species attaching eggs than in structuring those with freely laid eggs. The amount of unexplained variation was higher for assemblages with small-sized to medium-sized species, flying species and species with free eggs than those with large-sized species, non-flying species and species with attached eggs. Our observations of decreasing spatial and increasing environmental effects with increasing body size of assemblages deviated from the results of previous studies. These results suggest differing metacommunity dynamics between within-lake and among-lake levels and between studies covering contrasting taxonomic groups and body size ranges.

Do different facets of littoral macroinvertebrate diversity show congruent patterns in a large lake system

Modern biodiversity research focuses on multiple diversity facets because different indices may describe different ecological and environmental processes, as well as the effects of varied disturbances of natural and anthropogenic origins. We investigated littoral macroinvertebrate diversity in a large boreal lake system and specifically explored congruence of indices within and between the three diversity facets: species diversity, functional diversity and taxonomic distinctness. First, we found that the indices of taxonomic distinctness were the most sensitive indicators of eutrophication. Second, we observed that most correlations between the indices within the same diversity facet, and between the indices of functional and species diversity, were relatively strong. However, the indices of taxonomic distinctness (Δ+ and Λ+) were weakly associated with other metrics of diversity, emphasising the importance of taxonomic distinctness as a complementary dimension of biodiversity. Therefore, our observations support the importance to examine multiple facets for mapping biodiversity or for assessing the effects of anthropogenic disturbances on biological communities.