David L. Findlay

Eutrophication of lakes cannot be controlled by reducing nitrogen input: Results of a 37-year whole-ecosystem experiment

Lake 227, a small lake in the Precambrian Shield at the Experimental Lakes Area (ELA), has been fertilized for 37 years with constant annual inputs of phosphorus and decreasing inputs of nitrogen to test the theory that controlling nitrogen inputs can control eutrophication. For the final 16 years (1990–2005), the lake was fertilized with phosphorus alone. Reducing nitrogen inputs increasingly favored nitrogen-fixing cyanobacteria as a response by the phytoplankton community to extreme seasonal nitrogen limitation. Nitrogen fixation was sufficient to allow biomass to continue to be produced in proportion to phosphorus, and the lake remained highly eutrophic, despite showing indications of extreme nitrogen limitation seasonally. To reduce eutrophication, the focus of management must be on decreasing inputs of phosphorus.

EFFECTS OF CLIMATIC WARMING ON LAKES OF THE CENTRAL BOREAL FOREST

Twenty years of climatic, hydrologic, and ecological records for the Experimental Lakes Area of northwestern Ontario show that air and lake temperatures have increased by 2�C and the length of the ice-free season has increased by 3 weeks. Higher than normal evaporation and lower than average precipitation have decreased rates of water renewal in lakes. Concentrations of most chemicals have increased in both lakes and streams because of decreased water renewal and forest fires in the catchments. In Lake 239, populations and diversity of phytoplankton also increased, but primary production showed no consistent trend. Increased wind velocities, increased transparency, and increased exposure to wind of lakes in burned catchments caused thermoclines to deepen. As a result, summer habitats for cold stenothermic organisms like lake trout and opposum shrimp decreased. Our observations may provide a preview of the effects of increased greenhouse warming on boreal lakes.

Long-Term Ecosystem Stress: The Effects of Years of Experimental Acidification on a Small Lake

Experimental acidification of a small lake from an original pH value of 6.8 to 5.0 over an 8-year period caused a number of dramatic changes in the lakes food web. Changes in phytoplankton species, cessation of fish reproduction, disappearance of the benthic crustaceans, and appearance of filamentous algae in the littoral zone were consistent with deductions from synoptic surveys of lakes in regions of high acid deposition. Contrary to what had been expected from synoptic surveys, acidification of Lake 223 did not cause decreases in primary production, rates of decomposition, or nutrient concentrations. Key organisms in the food web leading to lake trout, including Mysis relicta and Pimephales promelas, were eliminated from the lake at pH values as high as 5.8, an indication that irreversible stresses on aquatic ecosystems occur earlier in the acidification process than was heretofore believed. These changes are caused by hydrogen ion alone, and not by the secondary effect of aluminum toxicity. Since no species of fish reproduced at pH values below 5.4, the lake would become fishless within about a decade on the basis of the natural mortalities of the most long-lived species.

Pelagic C : N : P stoichiometry in a eutrophied lake : responses to a whole-lake food-web manipulation

Changes in the ecological stoichiometry of C, N, and P in the pelagic zone are reported from a whole-lake manipulation of the food web of Lake 227, an experimentally eutrophied lake at the Experimental Lakes Area, Canada. Addition of northern pike eliminated populations of planktivorous minnows by the third year (1995) after pike introduction, and in the fourth year after pike addition (1996), a massive increase in the abundance of the large-bodied cladoceran Daphnia pulicaria occurred. Accompanying this increase in Daphnia abundance, zooplankton community N:P declined, seston concentration and C:P ratio decreased, and dissolved N and P pools increased. During peak abundance, zooplankton biomass comprised a significant proportion of total epilimnetic phosphorus (greater than 30%). During the period of increased Daphnia abundance, concentrations of dissolved inorganic nitrogen (TIN) increased more strongly than dissolved phosphorus (TDP), and thus TIN:TDP ratios were elevated. Sedimentation data indicated that increased grazing led to greatly reduced residence times of C, N, and especially P in the water column during 1996. Finally, previously dominant N-fixing cyanobacteria were absent during 1996. Our results show that strong effects of food-web structure can occur in eutrophic lakes and that stoichiometric mechanisms play a potentially important role in generating these effects.

Trophic Dependence of Ecosystem Resistance and Species Compensation in Experimentally Acidified Lake 302S (Canada)

AbstractEcosystem resistance to the impacts of diverse human insults depends on the replacement of sensitive species by ones more tolerant of the stressor. Here we present evidence from a whole-lake acidification experiment (Lake 302S, Experimental Lakes Area, Canada) that resistance and species compensation decline with increasing trophic level. Diverse and fast-growing algal and rotifer assemblages with high dispersal potentials showed significant compensatory species dynamics, resulting in the maintenance of total biomass despite 30%–80% declines in species richness. Canonical correspondence analysis showed that significant compensatory algal and rotifer dynamics were best explained by differential species tolerances of acidified chemical conditions coupled with release from resource limitation and predation. However, less diverse cladoceran, copepod, and fish assemblages showed significant declines in total biomass and weak species compensation with loss of species during acidification. In comparison, algal and zooplankton species dynamics remained relatively synchronized in a nearby unperturbed reference lake (Lake 239) during the experiment. As a result, Lake 302S showed limited ecosystem resistance to anthropogenic acidification. Therefore, we hypothesize that lost species will increase the susceptibility of acidified lakes to the adverse impacts of other environmental stressors (for example, climate warming, stratospheric ozone depletion, invasive species). Consequently, the ecosystem stability of boreal lakes is expected to decline as global change proceeds.

Response of Phytoplankton Communities to Acidification and Recovery in Killarney Park and the Experimental Lakes Area, Ontario

Abstract It has been widely speculated that controls of SO2 emissions would stimulate recovery of acidified freshwater lakes in Canada, the United States and Europe. Phytoplankton communities from 22 lakes near Killarney Park Ontario, covering a pH range from 4.5–7.7, were studied from 1998–2000 and compared to data from experimentally acidified (pH decreased 6.7 to 4.5) and recovered (pH increased to 6.0) Lake 302 South at the Experimental Lakes Area (ELA), northwestern Ontario to assess recovery from acidification. Based on historical data, pH levels have rebounded to above 6.0 in several lakes in the Killarney area that were previously acidified to pH 5.0–5.5. Phytoplankton biomass was not correlated to pH, but there was a highly significant relationship between species richness and pH. Recovery trajectories were observed in a subset of 6 lakes, combining species diversity data from the present study with historical data. Correspondence analysis indicated that several of the lakes that experienced increased pH have shifted towards phytoplankton assemblages typical of circumneutral environments.

Using temporal coherence to determine the response to climate change in Boreal Shield lakes

Climate change is expected to have important impacts on aquatic ecosystems. On the Boreal Shield, mean annual air temperatures are expected to increase 2 to 4 °C over the next 50 years. An important challenge is to predict how changes in climate and climate variability will impact natural systems so that sustainable management policies can be implemented. To predict responses to complex ecosystem changes associated with climate change, we used long-term biotic databases to evaluate how important elements of the biota in Boreal Shield lakes have responded to past fluctuations in climate. Our long-term records span a two decade period where there have been unusually cold years and unusually warm years. We used coherence analyses to test for regionally operating controls on climate, water temperature, pH, and plankton richness and abundance in three regions across Ontario: the Experimental Lakes Area, Sudbury, and Dorset. Inter-annual variation in air temperature was similar among regions, but there was a weak relationship among regions for precipitation. While air temperature was closely related to lake surface temperatures in each of the regions, there were weak relationships between lake surface temperature and richness or abundance of the plankton. However, inter-annual changes in lake chemistry (i.e., pH) were correlated with some biotic variables. In some lakes in Sudbury and Dorset, pH was dependent on extreme events. For example, El Nino related droughts resulted in acidification pulses in some lakes that influenced phytoplankton and zooplankton richness. These results suggest that there can be strong heterogeneity in lake ecosystem responses within and across regions.

Resilience of Epilithic Algal Assemblages in Atmospherically and Experimentally Acidified Boreal Lakes

Abstract Algal assemblages can be highly responsive to environmental changes in recovering acidified lakes. We compared epilithic algal assemblages in boreal lakes during chemical recovery from atmospheric (Killarney Park, Ontario) and experimental (Lake 302S, Experimental Lakes Area, Ontario) acidification to assess the impact of spatial and temporal scale of severe acidification on taxonomic resilience (i.e. recovery rate). Resilience was measured as the distance traveled by lakes in ordination space during pH recovery based on canonical correspondence analysis. Resilience was relatively negligible in the Killarney lakes, suggesting that eight years of experimental acidification in Lake 302S had less impact on biological recovery than did decades of regional acidification. Increases in dissolved organic carbon, dissolved inorganic carbon, and calcium best explained temporal variance of epilithic species abundances in the recovering acidified lakes. In Lake 302S, contrasting trajectories of taxonomic resilience and resistance, i.e. displacement from reference conditions following a perturbation, indicated that ecological factors affecting epilithon differed at corresponding pH levels during recovery and acidification. Our findings reveal that modeling of ecosystem recovery from severe acidification must account for the spatial and temporal scale of the perturbation, and biological delay responses that result in differences between recovery and acidification trajectories.

CARBON AVAILABILITY AND THE PATTERN OF CYANOBACTERIAL DOMINANCE IN ENRICHED LOW-CARBON LAKES1

We hypothesize that the pattern of cyanobacterial dominance in experimentally enriched, low‐carbon lakes is related not only to the resultant N:P ratio but also to the availability of carbon for gas‐vesicle synthesis. We tested this hypothesis by determining the buoyancy responses of a highly gas‐vacuolate, N2‐fixing cyanobacterium to P enrichment with and without induced C limitation. Enrichment of samples of Aphanizomenon schindleri (Kling et al. 1994) from blooms in Lake 227 with combinations of C, N, and P produced rapid buoyancy reductions in P treatments, reductions that were reversed within a generation time in treatments that included C or C and N as well as P. These responses are the first of their kind to be observed in experiments with lake populations of cyano‐bacteria. The rapid buoyancy reductions were associated with polyphosphate accumulations in P‐treated A. schindleri. Differences in buoyancy status after one generation time were linked to differences in relative gas vacuolation between samples treated with P only and samples treated with C and N as well as P. These results may explain the relative success of different types of cyanobacteria in newly enriched, low‐carbon lakes. The availability of C for gasvesicle synthesis may determine whether a low N:P ratio induces N2 fixation by benthic or by planktonic cyanobacteria and whether a high NP ratio leads to dominance by non‐gas‐vacuolate or by highly gas‐vacuolate, non‐N2‐fixers.

Seasonal and decadal patterns in Discostella (Bacillariophyceae) species from bi‐weekly records of two boreal lakes (Experimental Lakes Area, Ontario, Canada)

A recent rise in the relative abundance of Discostella species (D. stelligera and D. pseudostelligera) has been well documented from sedimentary diatom assemblages across the Northern Hemisphere. This unprecedented change over the last ~150 years has been linked to rises in atmospheric temperatures, changes in ice cover, and/or increases in thermal stability, among other factors. The bi‐weekly monitoring data from two boreal lakes at the Experimental Lakes Area (ELA) in northwestern Ontario were analyzed across seasons (spring, summer, and fall) and decades (1970s–2000s). We found that Discostella species are primarily spring/early summer bloomers (i.e., late April to June) in these lakes and changes in concentrations of Discostella over time were most pronounced in the spring or early summer months. Increases in Discostella abundance over time may be linked to earlier ice‐off and a longer period of spring turnover, resulting from increased winter and spring temperatures. It is also possible that a trophic mismatch between the spring diatom bloom and zooplankton is occurring, thus reducing diatom loss rates, and resulting in greater overall abundance. Moreover, the spring dominance of Discostella in our study lakes occurred at a time of the year when nutrient concentrations were at their highest seasonally, suggesting that these taxa are neither limited directly by nutrients, nor responding to enhanced stratification during the summer months in these lakes.