Liselotte S. Johansson

Zooplankton as indicators in lakes: a scientific-based plea for including zooplankton in the ecological quality assessment of lakes according to the European Water Framework Directive (WFD)

With the implementation of the EU Water Framework Directive (WFD), the member states have to classify the ecological status of surface waters following standardised procedures. It was a matter of some surprise to lake ecologists that zooplankton were not included as a biological quality element (BQE) despite their being considered to be an important and integrated component of the pelagic food web. To the best of our knowledge, the decision of omitting zooplankton is not wise, and it has resulted in the withdrawal of zooplankton from many so-far-solid monitoring programmes. Using examples from particularly Danish, Estonian, and the UK lakes, we show that zooplankton (sampled from the water and the sediment) have a strong indicator value, which cannot be covered by sampling fish and phytoplankton without a very comprehensive and costly effort. When selecting the right metrics, zooplankton are cost-efficient indicators of the trophic state and ecological quality of lakes. Moreover, they are important indicators of the success/failure of measures taken to bring the lakes to at least good ecological status. Therefore, we strongly recommend the EU to include zooplankton as a central BQE in the WFD assessments, and undertake similar regional calibration exercises to obtain relevant and robust metrics also for zooplankton as is being done at present in the cases of fish, phytoplankton, macrophytes and benthic invertebrates.

Biomanipulation as a Restoration Tool to Combat Eutrophication: Recent Advances and Future Challenges

Eutrophication resulting from high nutrient loading has been the paramount environmental problem for lakes world-wide for the past four decades. Efforts are being made in many parts of the world to reduce external nutrient loading via improved wastewater treatment or diversion of nutrient-rich inflows. However, even after a reduction of the external phosphorus loading, the effects obtained may be unsatisfactory. This may reflect an insufficient reduction in the external nutrient loading to effectively limit phytoplankton growth. However, the lack of success may also be due to chemical or biological within-lake inertia preventing or delaying improvements. To overcome the resilience and thereby reinforce recovery, a number of physico-chemical and biological restoration methods have been developed. In this chapter, we describe recent developments of biological restoration methods related to eutrophication, their short-term and long-term effects, and discuss the possibility of using combined physico-chemical and biological methods to improve the long-term stability of restoration and to reduce restoration costs. As comprehensive reviews of the effect of fish manipulation in cold temperate lakes are numerous, for these waterbodies, we highlight recent results, including effects on biodiversity and metabolism, and present new approaches of biomanipulation. Our particular focus is, however, directed at biomanipulation in warm lakes and on combined treatments which are far less well described in the literature.

Climate Versus In-Lake Processes as Controls on the Development of Community Structure in a Low-Arctic Lake (South-West Greenland)

The dominant processes determining biological structure in lakes at millennial timescales are complex. In this study, we used a multi-proxy approach to determine the relative importance of in-lake versus indirect processes on the Holocene development of an oligotrophic lake in SW Greenland (66.99°N, 50.97°W). A 14C and 210Pb-dated sediment core covering approximately 8500 years BP was analyzed for organic–inorganic carbon content, pigments, diatoms, chironomids, cladocerans, and stable isotopes (δ13C, δ18O). Relationships among the different proxies and a number of independent controlling variables (Holocene temperature, an isotope-inferred cooling period, and immigration of Betula nana into the catchment) were explored using redundancy analysis (RDA) independent of time. The main ecological trajectories in the lake biota were captured by ordination first axis sample scores (18–32% variance explained). The importance of the arrival of Betula (ca. 6500 years BP) into the catchment was indicated by a series of partial-constrained ordinations, uniquely explaining 12–17% of the variance in chironomids and up to 9% in pigments. Climate influences on lake biota were strongest during a short-lived cooling period (identified by altered stable isotopes) early in the development of the lake when all proxies changed rapidly, although only chironomids had a unique component (8% in a partial-RDA) explained by the cooling event. Holocene climate explained less variance than either catchment changes or biotic relationships. The sediment record at this site indicates the importance of catchment factors for lake development, the complexity of community trends even in relatively simple systems (invertebrates are the top predators in the lake) and the challenges of deriving palaeoclimate inferences from sediment records in low-Arctic freshwater lakes.

Size-based diel migration of zooplankton in Mediterranean shallow lakes assessed from in situ experiments with artificial plants

In warm lakes, fish aggregate within macrophytes, thereby weakening the role of these as a daytime refuge for zooplankton and altering the zooplankton size structure, predation pressure, and water clarity. To elucidate the role of macrophytes as a refuge for zooplankton and their effect on zooplankton size distribution, we established three sets of strandardized artificial plant beds in 11 lakes in Turkey with contrasting fish predation risk and turbidity. Zooplankton were sampled within and outside of each plant beds during day and night. Fish, collected overnight in multimesh-sized gillnets, were abundant both inside and outside the artificial plant beds, impoverishing the usefulness of plants as a daytime refuge for particularly large-bodied zooplankton. Zooplankton size diversity was negatively related to fish abundance. Diel vertical migration was the frequent anti-predator avoidance behavior, but reverse migration was also observed when Chaoborus was present. In contrast to the small-bodied taxa, large- and medium-sized taxa showed intraspecific size-based migration (i.e., individuals of different sizes had different migration patterns). Predators influenced the size structure and diel movement of zooplankton, but the response changed with the size of zooplankton and water clarity.

Temperature effects on fish production across a natural thermal gradient

Abstract Global warming is widely predicted to reduce the biomass production of top predators, or even result in species loss. Several exceptions to this expectation have been identified, however, and it is vital that we understand the underlying mechanisms if we are to improve our ability to predict future trends. Here, we used a natural warming experiment in Iceland and quantitative theoretical predictions to investigate the success of brown trout as top predators across a stream temperature gradient (4–25 °C). Brown trout are at the northern limit of their geographic distribution in this system, with ambient stream temperatures below their optimum for maximal growth, and above it in the warmest streams. A five‐month mark‐recapture study revealed that population abundance, biomass, growth rate, and production of trout all increased with stream temperature. We identified two mechanisms that contributed to these responses: (1) trout became more selective in their diet as stream temperature increased, feeding higher in the food web and increasing in trophic position; and (2) trophic transfer through the food web was more efficient in the warmer streams. We found little evidence to support a third potential mechanism: that external subsidies would play a more important role in the diet of trout with increasing stream temperature. Resource availability was also amplified through the trophic levels with warming, as predicted by metabolic theory in nutrient‐replete systems. These results highlight circumstances in which top predators can thrive in warmer environments and contribute to our knowledge of warming impacts on natural communities and ecosystem functioning.

Cross‐taxon congruence in lake plankton largely independent of environmental gradients

Groups of organisms often have congruent patterns of diversity or community structure due to similar environmental requirements. However, ecological interactions across trophic levels may also promote congruence independent of environmental drivers through selective predation, niche partitioning, or facilitation. We examined congruence between phytoplankton and zooplankton communities using 20 years of monitoring data from 17 Danish lakes, most of which were subject to external nutrient loading reduction after a period of eutrophication. Linear mixed effect models and partial Mantel tests were used to elucidate the extent to which congruence in genus richness and composition was driven by environmental factors. Congruence not explained by environmental controls might indicate ecological interactions across trophic levels of lake plankton. Genus richness and composition of phyto- and zooplankton were significantly congruent. Environmental factors had limited power to explain the genus richness of phyto- and zooplankton (R2: 6% and 12%, respectively). Including richness of the reciprocal group among the predictors markedly improved each respective model and explanatory power (R2: 11% and 17%, respectively). Similarly, a large proportion of the congruence in genus composition was independent of environmental dissimilarity (Mantel r: 0.29 and partial Mantel r after accounting for environmental control: 0.17). The strength of the congruence varied among different groups of phyto- and zooplankton, likely reflecting that the different zooplankton groups differ in their grazing size spectra and thus in the strength of their coupling with the phytoplankton assemblages. Remarkably, congruence with phytoplankton was strongest for the smallest group of zooplankton, rotifers, which graze on a limited size spectrum of phytoplankton. Moreover, congruence was stronger in low-nutrient lakes, suggesting that the strength of the interactions between the two groups weakens with eutrophication. This is likely due to changes in trophic dynamics, where enhanced fish predation on large-bodied zooplankton with increasing nutrient levels results in reduced zooplankton grazing control of phytoplankton assemblages.

Stable isotope analysis confirms substantial differences between subtropical and temperate shallow lake food webs

Differences in trophic web structure in otherwise similar ecosystems as a consequence of direct or indirect effects of ambient temperature differences can lead to changes in ecosystem functioning. Based on nitrogen and carbon stable isotope analysis, we compared the food-web structure in a series of subtropical (Uruguay, 30–35°S) and temperate (Denmark, 55–57°N) shallow lakes. The food-web length was on average one trophic position shorter in the subtropical shallow lakes compared with their temperate counterparts. This may reflect the fact that the large majority of subtropical fish species are omnivores (i.e., feed on more than one trophic level) and have a strong degree of feeding niche overlap. The shapes of the food webs of the subtropical lakes (truncated and trapezoidal) suggest that they are fuelled by a combination of different energy pathways. In contrast, temperate lake food webs tended to be more triangular, likely as a result of more simple pathways with a top predator integrating different carbon sources. The effects of such differences on ecosystem functioning and stability, and the connection with ambient temperature as a major underlying factor, are, however, still incipiently known.

Short-and long term niche segregation and individual specialization of brown trout (Salmo trutta) in species poor Faroese lakes

Trophic niche divergence is considered to be a major process by which species coexistence is facilitated. When studying niche segregation in lake ecosystems, we tend to view the niche on a one-dimensional pelagic-littoral axis. In reality, however, the niche use may be more complex and individual fidelity to a niche may be variable both between and within populations. In order to study this complexity, relative simple systems with few species are needed. In this paper, we study how competitor presence affects the resource use of brown trout (Salmo trutta) in 11 species-poor Faroese lakes by comparing relative abundance, stable isotope ratios and diet in multiple habitats. In the presence of three-spined sticklebacks (Gasterosteus aculeatus), a higher proportion of the trout population was found in the pelagic habitat, and trout in general relied on a more pelagic diet base as compared to trout living in allopatry or in sympatry with Arctic charr (Salvelinus alpinus). Diet analyses revealed, however, that niche-segregation may be more complex than described on a one-dimensional pelagic-littoral axis. Trout from both littoral and offshore benthic habitats had in the presence of sticklebacks a less benthic diet as compared to trout living in allopatry or in sympatry with charr. Furthermore, we found individual habitat specialization between littoral/benthic and pelagic trout in deep lakes. Hence, our findings indicate that for trout populations interspecific competition can drive shifts in both habitat and niche use, but at the same time they illustrate the complexity of the ecological niche in freshwater ecosystems.

Planktonic ciliate community structure in shallow lakes of lowland Western Europe

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although β-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.

Response of northern temperate shallow lakes to reduced nutrient loading, with special emphasis on Danish lakes

Cultural eutrophication has considerably impaired lake ecosystems worldwide (HUTCHINSON 1973, SCHINDLER 1977, SAS 1989, WETZEL 1990). Jnnorthern temperate lakes, total phosphorus (TP) is regarded as the key factor of eutrophication (SCHINDLER I 977). To improve water quality, attempts have been made during the last two decades to reduce the external nutrient Ioading of lakes by improving wastewater treatrnent, by increasing catchment retention capacity and by reducing the phosphorus content of fertilizers and detergents (PHILLIPS et al. 1999, VAN DER MOELEN & PORTIELJE 1999). However, many lakes have subsequently suffered from high interna! loading, delaying recovery (RYDING 198 I, MARSDEN 1989, SAS 1989). While studies o f the response o f nutrient and phytoplankton to nutrient loading reduction are plentiful (SAs I 989), Iittle emphasis has been placed on the higher trophic levels and changes in seasonality. For the Iast I 5 years, eight shallow Danish Iakes have been monitored intensively following nutrient Ioading reduction without the undertaking of additional fish manipulation to precipitate recovery (JEPPESEN et al. 2002 and 2005a, S0NDERGAARD et al. 2002 and 2005). In the present paper we highlight the major results of these studies and include more recent monitoring data from the Iakes. We further refer briefly to a recent comparative study of35 north temperate lakes (JEPPESEN et al. 2005a).