Contrasts between dystrophic and clearwater lakes in the long‐term effects of acidification on cladoceran assemblages

1. Most studies on zooplankton responses to acidification have focused on clearwater lakes with a dramatic acidification history. The role of dissolved organic carbon (DOC) in moderating zooplankton responses to acidification in naturally acidic, dystrophic lakes is less well understood and is partially impeded by a lack of baseline data. 2. Cladocera leave identifiable remains preserved in lake sediments that can be used to provide information on pre‐industrial species assemblages and their responses to environmental stressors such as acidification. Therefore, we used palaeolimnological approaches to track cladoceran assemblage responses to acidification since c.1850 (inferred from sedimentary diatom assemblages) in three acidified lakes in Kejimkujik National Park (Nova Scotia, Canada) that differ markedly in DOC content. These include two highly dystrophic lakes (Kejimkujik and Pebbleogittch lakes), and one clearwater lake (Beaverskin Lake). 3. In dystrophic Pebbleogittch Lake, an increase in the acid‐tolerant, jelly‐clad, pelagic taxon Holopedium glacialis occurred coincident with diatom‐inferred pH (DI‐pH) declines, but no other notable cladoceran assemblage shifts occurred. Similarly, Cladocera assemblages did not appear to respond to lakewater acidification in dystrophic Kejimkujik Lake. 4. In contrast, in the clearwater Beaverskin Lake, several observed shifts in cladoceran assemblage corresponded to DI‐pH declines, including an increase in the proportion of littoral taxa and an increase in Hill’s N2 species diversity. This may indicate increased water clarity as a result of acidification‐related decreases in DOC, which may have enhanced growth of emergent aquatic macrophytes and improved visibility for planktivorous fish, leading to increased predation on pelagic taxa. Species shifts within the littoral assemblage of Beaverskin Lake may reflect the differing tolerances of littoral taxa to low pH and aluminium toxicity. 5. Overall, our results suggest that cladoceran assemblages in naturally acidic, dystrophic lakes may be resilient against additional pH declines related to industrial emissions of acidifying agents, as dystrophic lakes are less vulnerable to increased aluminium toxicity and acidification‐induced increases in water clarity and often have a pre‐industrial cladoceran assemblage already adapted to acidic conditions.     

Are cladoceran diversity and community structure linked to spatial heterogeneity in urban landscapes and pond environments?

Biodiversity patterns in cladoceran communities were investigated in urban waterbodies in relation with residential land use, pond management, and waterbody environments. We evaluated species richness in the pelagic and littoral zones of eighteen waterbodies of a large Canadian city. Gamma diversity (26 species) observed at a small scale in the urban survey was important comparatively to large-scale surveys of lakes. Beta diversity ranged from 1 to 8 species among waterbodies. We tested if littoral species greatly contributed to regional diversity in urban waterbodies. Littoral species (Chydoridae, Ilyocryptidae, Macrothricidae, Polyphemidae) accounted for 58% of the total species pool. We distinguished five cladoceran assemblages associated to different waterbodies (temporary ponds, permanent lakes, and wetlands). Cladoceran communities were more diverse and variable in permanent lakes than in temporary ponds. Changes in cladoceran species assemblages among waterbodies were driven by variations in waterbody size and phosphorus enrichment, macrophyte and algal biomass, urban density, pond management practices, and the presence of potential predators as fish and macroinvertebrates. Our study indicates that both artificial ponds and lakes and natural wetlands are valuable habitats for the conservation of cladoceran biodiversity and rare endemic species in urban regions. Further research on pond management strategies promoting urban aquatic biodiversity should be undertaken.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

Patterns in the Distribution of Cladocerans (Crustacea: Branchiopoda) in Lakes Across a North–south Transect in Alaska, USA

The remains of cladocerans were examined from the surface sediments of 51 freshwater sites along a north–south transect spanning Alaska. We identified 27 cladoceran taxa from the sediments, consisting primarily of littoral chydorid species. Variations in cladoceran assemblages were related to measured physical and chemical variables using multivariate techniques. Redundancy analysis (RDA) indicated that lake depth, total phosphorus (TP), and altitude all had a significant influence in determining the composition of cladoceran assemblages. Cladoceran communities in tundra and forest-tundra lakes, which were relatively shallow and nutrient-poor, had relatively low abundances of pelagic Cladocera, and were primarily composed of several littoral chydorid species. Among pelagic cladoceran species, there was a distinct shift in dominance from the Bosminidae in lakes in the southern boreal forest region to Daphniidae in lakes in the northern boreal forest. Daphnia dominated lakes had significantly higher total phosphorus, specific conductivity, and calcium concentrations than lakes dominated by Eubosmina. Overall, the relative importance of physical and chemical factors in structuring cladocerans is similar to other previously studied regions, and suggests the Cladocera may be useful as ecological and paleoenvironmental indicators in this region.     

Seasonal fluctuations in macrophyte cover and water transparency of four brown-water lakes: implications for crustacean zooplankton in littoral and pelagic habitats

The dynamics of crustacean zooplankton in the littoral and pelagic zones of four forest lakes having variable water qualities (colour range 130–340 mg Pt l⁻¹, Secchi depth 70–160 cm) were studied. The biomass of zooplankton was higher in the littoral zone than in the pelagic zone only in the lake having the highest transparency. In the three other lakes, biomass was significantly higher in the pelagic zone than in the littoral zone. In the two lakes with highest transparency, the littoral biomass of cladocerans significantly followed the development of macrophyte vegetation, and cladoceran biomass reached the maximum value at the time of highest macrophyte coverage. In lakes with lowest transparency, littoral zooplankton biomass developed independently of macrophyte density and decreased when macrophyte beds were densest. The seasonal development of the littoral copepod biomass did not follow the development of macrophytes in any of the lakes. The mean size of cladocerans in the pelagic zone decreased with increasing Secchi depth of the lake, whereas in the littoral zone no such phenomenon was detected. Seasonally, when water transparency increased temporarily in two of the lakes, the mean size of cladocerans in the pelagic zone decreased steeply. For copepods, no relationship between water transparency and body size was observed. The results suggested that in humic lakes the importance of the littoral zone as a refuge decreases with decreasing transparency of the water and that low water transparency protects cladocerans from fish predation. All the observed between-lake differences could not be explained by fish predation, but were probably attributed to the presence of chaoborid larvae with variable densities. Feeding efficiency of chaoborids is not affected by visibility and thus they can obscure the relationship between water quality, fish density, and the structure of crustacean zooplankton assemblages. Handling editor: S. I. Dodson     

Species richness of pelagic Cladocera of large lakes in the eastern hemisphere

The composition of the pelagic cladoceran species assemblages in some large, well-studied, lakes of Europe, Asia and Africa is reviewed based on original and literature data. In general, pelagic cladocerans are taxonomically less well studied than some littoral groups. Only the limnetic members of the family Sididae and some Daphnia have recently been revised, whereas knowledge of species diversity of other Daphniidae (including moinas) and Bosminidae is missing. Future detailed taxonomic studies may lead to considerable changes in understanding of limnetic zooplankton composition.     

Using invertebrate remains and pigments in the sediment to infer changes in trophic structure after fish introduction in Lake Fogo: a crater lake in the Azores

Fish introduction may have marked effects on the trophic dynamics and ecological state of former fishless lakes, but due to scarcity of historical data this can seldom be documented. We used remains of cladoceran, chironomid and pigment assemblages in the sediment archive to unravel the effect of introduction of carp (Cyprinus carpio), rainbow trout (Oncorhynchus mykiss) and a cyprinid (Chondrostoma oligolepis) in Lake Fogo, the Azores (Portugal). The stratigraphical record showed two major shifts in community assemblage coinciding with the time of introduction of carp (AD ca. 1890) and trout (AD 1941), respectively. Carp introduction was followed by an abrupt and major decline in the abundance of chironomids, a shift in the cladoceran community from a benthic to a more pelagic dominated community, and Daphnia size was significantly reduced. Pigment assemblages also indicated a shift from a benthic to a pelagic dominated ecosystem, as cryptophytes became markedly more abundant at the expense of benthic diatoms. Trout introduction was followed by a return to a more benthic cladoceran and benthic algae (pigments) dominated state, which we attribute to trout predation on carp leading to improved water clarity. A steady increase in the abundance of pigments and cladoceran remains followed, suggesting enhanced productivity, which may be attributed to enhanced atmospheric nitrogen deposition and introduction of C. oligolepis. We conclude that fish introduction has profoundly altered the trophic dynamics and the relative importance of benthic and pelagic production in this species poor and natural fishless lake in the Azores, and likely in most others lakes at the archipelago islands as fish stocking has been a widespread practice.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Reconstructing the past density of planktivorous fish and trophic structure from sedimentary zooplankton fossils: a surface sediment calibration data set from shallow lakes

1. As quantitative information on historical changes in fish community structure is difficult to obtain directly from fish remains in lake sediments, transfer function for planktivorous fish abundance has been developed based on zooplankton remains in surface sediment (upper 1 cm). The transfer function was derived using weighted average regression and calibration against contemporary data on planktivorous fish catch per unit effort (PF-CPUE) in multiple mesh size gill nets. Zooplankton remains were chosen because zooplankton community structure in lakes is highly sensitive to changes in fish predation pressure. The calibration data set consisted of thirty lakes differing in PF-CPUE (range 18–369 fish net^–1), epilimnion total phosphorus (range 0.025–1.28 mg P l^–1) and submerged macrophyte coverage (0–57%). 2. Correlation of log-transformed PF-CPUE, total phosphorus and submerged macrophyte coverage v the percentage abundance in the sediment of the dominant cladocerans and rotifers revealed that the typical pelagic species correlated most highly to PF-CPUE, while the littoral species correlated most highly to submerged macrophyte coverage. Consequently, only pelagic species were taken into consideration when establishing the fish transfer function. 3. Canonical correspondence analysis (CCA) revealed that the pelagic zooplankton assemblage was highly significantly related to PF-CPUE (axis 1), whereas the influence of total phosphorus and submerged macrophyte coverage was insignificant. Predicted PF-CPUE based on weighted average regression without (WA) and with (WA(tol)) downweighting of zooplankton species tolerance correlated significantly with the observed values (r² = 0.64 and 0.60 and RMSE = 0.54 and 0.56, respectively). A marginally better relationship (r² = 0.67) was obtained using WA maximum likelihood estimated optima and tolerance. 4. It is now possible, quantitatively, to reconstruct the historical development in planktivorous fish abundance based on zooplankton fossil records. As good relationships exist between contemporary PF-CPUE data and indicators such as the zooplankton/phytoplankton biomass ratio, Secchi depth and the maximum depth distribution of submerged macrophytes, it is now also possible to derive information on past changes in lake water quality and trophic structure. It will probably prove possible further to improve the transfer function by including other invertebrate remains, e.g. chironomids, Chaoborus, snails, etc., and its scope could be widened by including deeper lakes, more oligotrophic lakes, more acidic lakes and lakes with extensive submerged macrophyte coverage (in the latter case to enable use of the information in the fossil record on plant-associated cladocerans).     

Cladocera assemblages from the surface sediments of south-central Ontario (Canada) lakes and their relationships to measured environmental variables

Fossil cladoceran remains preserved in surface sediment samples from 44 oligotrophic lakes in south-central Ontario were examined to evaluate the relationships between species assemblages and measured environmental variables. Differences in cladoceran assemblages were related to physical and chemical variables using multivariate techniques. Redundancy Analysis (RDA) identified five environmental variables as significantly influencing assemblage composition: sulphate (SO₄²⁻), calcium (Ca²⁺), pH, maximum lake depth (Z _max) and dissolved organic carbon (DOC). There was a distinct separation of lakes and taxa along the ion gradient based on SO₄, Ca and pH. Additionally, cladoceran communities in coloured, shallow lakes had relatively higher abundances of littoral chydorid species and the pelagic taxa Holopedium spp., and the Daphnia pulex complex. Deep, clear lakes had relatively higher abundances of other pelagic taxa. Predation by fish (measured as presence–absence) and Chaoborus (measured as density) were less significant than some of the physico-chemical variables in influencing cladoceran assemblage structure. However, this could be due to the limited resolution of the predation data that was available at the time of this study. The distribution of cladocerans in the surface sediment, and their relation to these important environmental variables, suggests that there is considerable potential for the use of sedimentary cladoceran remains as environmental indicators in south-central Ontario lakes. Handling editor: J. Saros     

Trophic structure in the pelagial of 25 shallow New Zealand lakes: changes along nutrient and fish gradients

To gain better insight into the importance of predator and resourcecontrol in New Zealand lakes we surveyed the late summer trophicstructure of 25 shallow South Island lakes with contrastingnutrient levels (6–603 µg TP l^–1) and fishdensities. Total catch of fish per net (CPUE) in multi-meshgillnets placed in the open water and the littoral zones waspositively related with the nutrient level. Trout CPUE was negativelycorrelated with total phosphorus (TP) and total nitrogen (TN).Zooplankton seemed largely influenced by fish, as high fishCPUE coincided with low zooplankton and Daphnia biomass, lowaverage weight of cladocerans, low contribution of Daphnia tototal cladoceran biomass, low ratio of calanoids to total copepodbiomass and low ratio of zooplankton biomass to phytoplanktonbiomass. However, chlorophyll a was only slightly negativelyrelated to Daphnia biomass and not to zooplankton biomass ina multiple regression that included TN and TP. Ciliate abundancewas positively related to chlorophyll a and negatively to Daphniabiomass, but not to total zooplankton biomass, while no relationshipswere found between heterotrophic nanoflagellates and zooplankton.The relationships between fish abundance and nutrients and fishabundance and zooplankton:phytoplankton ratio and between chlorophylla and TP largely followed the pattern obtained for 42 northtemperate Danish lakes. We conclude that fish, including trout,have a major effect on the zooplankton community structure andbiomass in the pelagial of the shallow oligotrophic to slightlyeutrophic New Zealand lakes, but that the cascading effectson phytoplankton and protist are apparently modest.     

Distribution patterns of Cladocera in subarctic Fennoscandian lakes and their potential in environmental reconstruction

Multivariate statistical techniques were used to examine the relationships between surface-sediment cladoceran assemblages and 28 physical and chemical variables in 53 small subarctic lakes from northern Fennoscandia. The lakes were distributed along a steep eco-climatic gradient, spanning boreal corniferous forest to treeless tundra. In general, the sites were small, oligotrophic, and bathymetrically simple, with little or no disturbance in their catchments. From the initial 53 localities, only 36 contained a sufficient number of cladoceran remains for reasonable quantification. From these, a total of 29 cladoceran taxa representing 19 genera were identified, comprising predominantly littoral chydorid species. A constrained redundancy analysis (RDA) and associated Monte Carlo permutation tests indicated that maximum lake depth, sediment organic content, epilimnetic summer temperature, lake perimeter, and lake catchment area made statistically significant (p le; 0.05) contributions to explaining the variance in the cladoceran taxon data. These five variables together accounted for 67.7% of the explained variance, and made a unique contribution of 26.8% to the total variance: all physical determinants independently captured 33.2% of the total variance. The significance of the most powerful explanatory variables is discussed in the paper in detail, and autecological information regarding the most common cladoceran taxa is given. To assess the potential of cladoceran assemblages in environmental reconstruction, quantitative inference models for mean July water and air temperatures were developed for the cladoceran assemblage using partial least squares (PLS) regression. The final prediction model yielded a root mean squared error of prediction (RMSEP). as assessed by jackknifing, of 1.19°C for Cladocera-water temperature data-set, whereas the cladoceran assemblages showed only very weak relationships to mean July air temperature. The overall results emphasize the role of physical factors in regulating species abundance and distributions in these environmentally sensitive ecotonal lakes.     

The role of cladocerans in tracking long-term change in shallow lake trophic status

Shallow lakes have been affected by a variety of human activities profoundly altering their ecological structure and function. Cladocerans have been used to track change resulting from a variety of drivers at a number of time scales. Aquatic macrophytes are well recognised as reflecting the ecological condition of a lake. Here, we compare the plant macrofossils with the sub-fossil cladoceran assemblages from 20 dated sediment cores. Co-correspondence analysis was used to determine the degree of commonality of change in community composition of the two biological groups through time. This analysis revealed very high levels of agreement in the nature and timing of change at all the sites examined with very high correlation coefficients between the axis 1 scores for macrofossils and cladocerans. Furthermore, at all sites a high proportion of the variance (min 20%, max 54%) in the macrofossil data was explained by the change in the cladoceran assemblage. Sub-fossil macrofossil and cladoceran assemblages, from at least from 1700 AD onwards, were examined in more detail at three sites: Ormesby Great Broad, Felbrigg Lake and Lake Søbygaard. There was very good accord in the main shifts of the cladoceran and macrofossil assemblages at all three sites. This may reflect the long-term shift in the principal focus of primary production from the benthic to the pelagic habitat. We suggest that the combination of their central position in the food-web and the presence of both pelagic and benthic taxa make cladocerans a strong candidate as the single best indicator of (palaeo) ecological condition related to changing trophic status and alteration in food-web structure in shallow lakes.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Steady state phytoplankton assemblages during thermal stratification in deep alpine lakes. Do they occur?

Phytoplankton seasonal and long-term succession can be described and functionally classified by associations similar as in terrestrial vegetation studies. Such a concept has to include 'climax' into pelagic succession which in turn leads to the question to what extent steady state assemblages occur and if during periods of dynamic equilibrium can be identified. Here we explore the situation with respect to the above question for deep, nutrient poor, alpine lakes in Austria. We first track the long-term development of phytoplankton biomass, their taxonomic structure and their relation to total phosphorus and chlorophyll-a as predictors of trophic state over the past 35 years. We then analyse this data set for coherent algal associations which can be ascribed to trait separated functional groups according to Reynolds et al. (2002). A three year period of stable environmental conditions has then be extracted from the progression of trophic state indices, having similar dominating species each year. These years were finally analysed for steady state conditions according to definitions given in Sommer et al. (1993). During thermal stratification, achievement of an equilibrium could be ruled out although coexistence of several dominating species lasted for several weeks. Habitat templates were constructed from environmental variables prior to biomass peaks for two species important in summer assemblages, the dinoflagellate Ceratium hirundinella and the diatom Fragilaria crotonensis. In summary, functional groups proved to be a valid and useful concept to describe species succession of phytoplankton in deep alpine lakes while pelagic climax is much less clear and steady state conditions were never met.     

Functional shifts in lake zooplankton communities with hypereutrophication

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Relating planktonic microbial food web structure in lentic freshwater ecosystems to water quality and land use

Relationships among picoplankton, protozoa, phytoplankton, plantnutrients, lake type, drainage basin morphology and land coverwere studied in 45 water bodies in South Island, New Zealandthat ranged from large, deep, ultra-oligotrophic lakes to shallow,macrophyte-dominated ponds and swamps. The biomasses of mostheterotrophic components of the pelagic microbial food webswere positively related to phytoplankton and features of thedrainage basin that enhanced nutrient input, and imply strongresource-driven structuring of pelagic microbial food webs.Prokaryotic picophytoplankton biomass was negatively relatedto indices of eutrophication, and the picoautotroph contributionto total microbial food web biomass declined with increasingtotal phosphorus concentration from 16.5% in deep lakes to <0.02%in swamps and ponds. Biomass ratios of (picoplankton plus protozoa):phytoplanktonranged from 40:60 in swamps and ponds to >70:30 in deep lakes,and indicate the potential importance of microbial food websin carbon transfer to higher trophic levels in deep, less productivelakes. Strong relationships exist between land use in the catchmentand pelagic microbial food web structure and biomass acrossa wide range in size and trophic state of water bodies in heterogeneouslandscapes.     

Sub-fossils of cladocerans in the surface sediment of 135 lakes as proxies for community structure of zooplankton,fish abundance and lake temperature

To elucidate the possibilities of using zooplankton remains in the surface sediment to describe present-days community structure and population dynamics of zooplankton, fish abundance and temperature, we compared contemporary data sampled in the pelagial during summer with the sediment record from the upper 1 cm of the sediment in 135 lakes covering a latitude gradient from Greenland in the north to New Zealand in the south. The abundance of three genera Bosmina, Daphnia and Ceriodaphnia of the total pool of ephippia was significantly related to the total abundance of the same taxa in the pelagic zone. However, in most lakes the abundance of Ceriodaphnia was higher in the sediment than in the water, which may be attributed to the overall preference by this genus for the littoral habitat. Using contemporary data from 27 Danish lakes sampled fortnightly during summer for 10 years, we found substantial inter-annual variations in the abundance of Daphnia spp., Ceriodaphnia spp., B. longirostris and B. coregoni. Yet, the sediment record mimicked the medium level well for most of the lakes, which suggests that the sediment record provides an integrated picture of the pelagic cladoceran community, which otherwise can be obtained only by long-term frequent contemporary sampling for several years. The contribution of Daphnia to the sum of Daphnia and Bosmina ephippia was negatively correlated with the abundance of fish expressed as catch per night in multi-mesh sized gill nets (CPUE). Yet, region-specific differences occurred, which partly could be eliminated by including nutrient state expressed as total phosphorus (TP) in a multiple regression. The average ratio of ephippia to the sum of ephippia and carapaces of Bosmina varied 40-fold between the sampling regions and was significantly negatively related to summer mean air temperature, and for Danish lakes also, albeit weakly, to fish CPUE but not to chlorophyll a. Apparently, temperature is the most important factor determining the ratio of parthenogenetic to ephippia producing specimens of Bosmina. We conclude that the sediment record of cladocerans is a useful indicator of community structure of pelagic cladocerans and the abundance of fish and temperature.     

The paradox of re‐oligotrophication: the role of bottom–up versus top–down controls on the phytoplankton community

Increases in phytoplankton biomass have been widely observed over the past decades, even in lakes experiencing nutrient reduction. However, the mechanisms giving rise to this trend remain unclear. Here, we unveil the potential mechanisms through quantifying the relative contribution of bottom–up versus top–down control in determining biomasses of phytoplankton assemblages in Lake Geneva. Specifically, we apply nonlinear time series analysis, convergent cross mapping (CCM), to decipher the degree of bottom–up versus top–down control among phytoplankton assemblages via quantifying 1) causal links between environmental factors and various phytoplankton assemblages and 2) the relative importance of bottom–up, top–down, and environmental effects. We show that the recent increase in total phytoplankton biomass, albeit with phosphorus reduction, was mainly caused by a particular phytoplankton assemblage which was better adapted to the re‐oligotrophicated environment characterized by relatively low phosphorus concentrations and warm water temperature, and poorly controlled by zooplankton grazing. Our findings suggest that zooplankton act as a critical driver of phytoplankton biomass and strongly impact the dynamics of recovery from eutrophication. Therefore, our phytoplankton assemblage approach in combination with causal identification of top–down versus bottom–up controls provides insights into the reason why phytoplankton biomass may increase in lakes undergoing phosphorus reduction.