Michelle E. Palmer

The Widespread Threat of Calcium Decline in Fresh Waters

Calcium concentrations are now commonly declining in softwater boreal lakes. Although the mechanisms leading to these declines are generally well known, the consequences for the aquatic biota have not yet been reported. By examining crustacean zooplankton remains preserved in lake sediment cores, we document near extirpations of calcium-rich Daphnia species, which are keystone herbivores in pelagic food webs, concurrent with declining lake-water calcium. A large proportion (62%, 47 to 81% by region) of the Canadian Shield lakes we examined has a calcium concentration approaching or below the threshold at which laboratory Daphnia populations suffer reduced survival and fecundity. The ecological impacts of environmental calcium loss are likely to be both widespread and pronounced.

Should Biological Invasions Be Managed as Natural Disasters

Biological invasions and natural disasters are similar phenomena: Their causes are well understood, hut their occurrences are generally unpredictable and uncontrollable. “Both invasions and natural disasters can generate enormous environmental damage, and the frequency of damaging events is inversely proportional to their magnitude. Many nations invest in personnel training, disaster preparedness, and emergency response plans for extreme natural hazards (e.g., earthquakes), despite the rarity of such events. Similar precautions for invasive species (apart from infectious diseases) are not comprehensively applied by any nation, even though the impacts of invasions are less predictable and often irrevocable. Furthermore, the annual combined economic cost of invasions worldwide exceeds that of natural disasters. Preventative management of invasions—like that of natural disasters—requires international coordination of early-warning systems, immediate access to critical information, specialized training of personnel, and rapid-response strategies.

The state of Lake Simcoe (Ontario, Canada): the effects of multiple stressors on phosphorus and oxygen dynamics

Abstract Lake Simcoe, the largest lake in southern Ontario outside of the Laurentian Great Lakes, is affected by numerous stressors including eutrophication resulting from total phosphorus (TP) loading, climate change, and invasions of exotic species. We synthesized the long-term responses of Lake Simcoe to these stressors by assessing trends in water quality and biological composition over multiple trophic levels. Evidence for climate change included increasing thermal stability of the lake and changes in subfossil diatom communities over time. Although the deep water dissolved oxygen (O2) minimum has increased significantly since TP load reductions, it is still below estimated historical values and the Lake Simcoe Protection Plan end-of-summer target level of 7 mg O2 L-1. Low deep water O2 concentrations corresponded with a decline in coldwater fish abundance. Since 1980, some nutrient concentrations have decreased (spring TP) while others have increased (silica), but many show no obvious changes (ice-free TP, nitrate, ammonium). Increases in water clarity, combined with declines in chlorophyll a and phytoplankton biovolumes in Cook’s Bay, were temporally consistent with declines in TP loading and the lake-wide establishment of dreissenid mussels as a major component of the Lake Simcoe ecosystem. Using an investigative tool, we identified 2 periods when abrupt shifts potentially occurred in multiple parameters: 1986 and 1995-1997. Additional ecosystem level changes such as declines in zooplankton, declines in offshore benthic invertebrate abundance, and increased nearshore invertebrate abundance likely reflect the effects of invasive species. The interaction of these multiple stressors have significantly altered the Lake Simcoe ecosystem.

The jellification of north temperate lakes

Calcium (Ca) concentrations are decreasing in softwater lakes across eastern North America and western Europe. Using long-term contemporary and palaeo-environmental field data, we show that this is precipitating a dramatic change in Canadian lakes: the replacement of previously dominant pelagic herbivores (Ca-rich Daphnia species) by Holopedium glacialis, a jelly-clad, Ca-poor competitor. In some lakes, this transformation is being facilitated by increases in macro-invertebrate predation, both from native (Chaoborus spp.) and introduced (Bythotrephes longimanus) zooplanktivores, to which Holopedium, with its jelly coat, is relatively invulnerable. Greater representation by Holopedium within cladoceran zooplankton communities will reduce nutrient transfer through food webs, given their lower phosphorus content relative to daphniids, and greater absolute abundances may pose long-term problems to water users. The dominance of jelly-clad zooplankton will likely persist while lakewater Ca levels remain low.

Climate change drives coherent trends in physics and oxygen content in North American lakes

Using a 25-year record of monitoring data, we show that recent climate change has affected the thermal properties and oxygen content of seven lakes in south-central Ontario, Canada, and five lakes in north-central Wisconsin, USA. Coherent patterns in autumnal lake warming were driven by increased autumn air temperature in both lake districts. Temperature increases were restricted to the epilimnion and metalimnion of the lakes, resulting in increased thermal stability of the water column. Mixing depths also decreased over the study period. Shallower mixing depths in the Ontario lakes were due to climate-driven increases in lake-water dissolved organic carbon concentrations. Collectively, changes in the thermal regime of the lakes suggest autumn mixing of the water column may be delayed. Metalimnetic oxygen also increased in the Wisconsin lakes, perhaps in response to increased algal production as lake thermal regimes changed. The response of individual lakes to climate change was modified by lake chemistry in the Ontario lake district and by lake chemistry and morphometry in the Wisconsin lake district. Our results demonstrate coherent lake response to climate change and highlight the importance of both regional and local factors in regulating individual lake response to global climate change.

Continuous Bayesian network for studying the causal links between phosphorus loading and plankton patterns in Lake Simcoe, Ontario, Canada.

An ecosystem perspective to restoring beneficial uses in Areas of Concern can be interpreted as a shift from the traditional elucidation of simple cause-effect relationships to a multicausal way of thinking that more effectively accommodates ecosystem complexity. This holistic management paradigm has also pervaded the contemporary ecological modeling practice, making compelling the adoption of more sophisticated ecosystem modeling tools. In this study, our primary objective is to develop a Bayesian hierarchical network of simple ecological models for Lake Simcoe, Ontario, Canada, aiming to establish a realistic representation of the causal connections among exogenous nutrient loading, ambient nutrient conditions, and epilimnetic plankton dynamics. In particular, we used a spatially explicit simple mass-balance model forced with idealized sinusoidal loading to predict total phosphorus concentrations. A structural equation model was then used to delineate the interplay among nutrients, ambient light conditions, phytoplankton, and herbivorous biomass. Our analysis highlights the strength of the causal linkages between total phosphorus and water clarity with phytoplankton as well as the capacity of zooplankton grazing to modulate the algal standing crop. Our Bayesian network is also used to examine the exceedance frequency of threshold values for total phosphorus (15 μg/L) and chlorophyll a (4 μg/L) concentrations under scenarios of phosphorus loading reduction. Our study suggests that a 15% phosphorus loading decrease will still result in >25% violations of the 4 μg chla/L value in the two embayments of Lake Simcoe (Cooks Bay and Kempenfelt Bay). The TP levels will decrease in response to the exogenous loading reductions and this improvement will be primarily manifested in the northcentral segments of the system.

Ice cover extent drives phytoplankton and bacterial community structure in a large north‐temperate lake: implications for a warming climate

Mid-winter limnological surveys of Lake Erie captured extremes in ice extent ranging from expansive ice cover in 2010 and 2011 to nearly ice-free waters in 2012. Consistent with a warming climate, ice cover on the Great Lakes is in decline, thus the ice-free condition encountered may foreshadow the lakes future winter state. Here, we show that pronounced changes in annual ice cover are accompanied by equally important shifts in phytoplankton and bacterial community structure. Expansive ice cover supported phytoplankton blooms of filamentous diatoms. By comparison, ice free conditions promoted the growth of smaller sized cells that attained lower total biomass. We propose that isothermal mixing and elevated turbidity in the absence of ice cover resulted in light limitation of the phytoplankton during winter. Additional insights into microbial community dynamics were gleaned from short 16S rRNA tag (Itag) Illumina sequencing. UniFrac analysis of Itag sequences showed clear separation of microbial communities related to presence or absence of ice cover. Whereas the ecological implications of the changing bacterial community are unclear at this time, it is likely that the observed shift from a phytoplankton community dominated by filamentous diatoms to smaller cells will have far reaching ecosystem effects including food web disruptions.

Bythotrephes invasion elevates trophic position of zooplankton and fish: implications for contaminant biomagnification

We estimated the effects of Bythotrephes longimanus invasion on the trophic position (TP) of zooplankton communities and lake herring, Coregonus artedi. Temporal changes in lacustrine zooplankton communities following Bythotrephes invasion were contrasted with non-invaded reference lakes, and along with published information on zooplankton and herring diets, formed the basis of estimated changes in TP. The TP of zooplankton communities and lake herring increased significantly following the invasion of Bythotrephes, whereas TP in reference lakes decreased (zooplankton) or did not change significantly (lake herring) over a similar time frame. Elevated TP following Bythotrephes invasion was most prominent in lakes that also supported the glacial relict, Mysis diluvania, suggesting a possible synergistic interaction between these two species on zooplankton community composition. Our analysis indicated that elevated TPs of zooplankton communities and lake herring are not simply due to the presence of Bythotrephes, but rather reflect changes in the zooplankton community induced by Bythotrephes; namely, a major reduction in the proportion of herbivorous cladoceran biomass and a concomitant increase in the proportion of omnivorous and/or predatory copepod biomass in invaded lakes. We demonstrated that increases in TP of the magnitude reported here can lead to substantial increases in fish contaminant concentrations. In light of these results, we discuss potential mechanisms that may be responsible for the disconnect between empirical and theoretical evidence that mid-trophic level species invasions (e.g., Bythotrephes) elevate contaminant burdens of consumer species, and provide testable hypotheses to evaluate these mechanisms.

Gauging recovery of zooplankton from historical acid and metal contamination: the influence of temporal changes in restoration targets

Summary Clearly defined restoration targets are necessary to judge the effectiveness of management actions in restoring damaged ecosystems. However, the identification of appropriate targets is difficult in a rapidly changing world. Historical reference conditions commonly provide recovery targets, but they may not be appropriate if present-day environments have shifted in response to regional or global drivers. Such shifts may need to be incorporated into restoration targets to avoid erroneous conclusions about the recovery of ecosystems damaged by localized stressors. No previous study has examined whether the selection of historical vs. present-day reference conditions alters judgments of the recovery of historically damaged ecosystems. We examined 35-year trends in the zooplankton communities of four lakes polluted by smelter emissions in Sudbury, Ontario, Canada. Recovery was assessed by comparing the Sudbury lakes to both historical (1983–1984) and present-day (2004–2006) conditions in a set of minimally impacted reference lakes in south-central Ontario. Sudbury zooplankton communities improved substantially over time when compared with both the historical and present-day recovery targets. However, recovery occurred later, and improvements differed quantitatively when judged against the present-day vs. historical targets. These differences were attributable to regional shifts in zooplankton communities that happened after the historical sampling period but were reflected in the present-day data. Species richness in two Sudbury lakes met recovery targets and communities in all four lakes became more similar to those in the reference lakes. However, the continued absence of many daphniids, cyclopoids and large calanoids indicated that the lakes had not fully recovered and further monitoring is needed. Synthesis and applications. Our results show that the choice of reference condition can alter recovery assessments. This finding emphasizes the importance of establishing clearly defined restoration goals to ensure appropriate choice of reference conditions. Restoration is unlikely to be judged as successful if an historical reference point is used to guide management actions meant to restore an ecosystem to present-day regional conditions.

Forecasting cyanobacteria dominance in Canadian temperate lakes

Predictive models based on broad scale, spatial surveys typically identify nutrients and climate as the most important predictors of cyanobacteria abundance; however these models generally have low predictive power because at smaller geographic scales numerous other factors may be equally or more important. At the lake level, for example, the ability to forecast cyanobacteria dominance is of tremendous value to lake managers as they can use such models to communicate exposure risks associated with recreational and drinking water use, and possible exposure to algal toxins, in advance of bloom occurrence. We used detailed algal, limnological and meteorological data from two temperate lakes in south-central Ontario, Canada to determine the factors that are closely linked to cyanobacteria dominance, and to develop easy to use models to forecast cyanobacteria biovolume. For Brandy Lake (BL), the strongest and most parsimonious model for forecasting % cyanobacteria biovolume (% CB) included water column stability, hypolimnetic TP, and % cyanobacteria biovolume two weeks prior. For Three Mile Lake (TML), the best model for forecasting % CB included water column stability, hypolimnetic TP concentration, and 7-d mean wind speed. The models for forecasting % CB in BL and TML are fundamentally different in their lag periods (BL = lag 1 model and TML = lag 2 model) and in some predictor variables despite the close proximity of the study lakes. We speculate that three main factors (nutrient concentrations, water transparency and lake morphometry) may have contributed to differences in the models developed, and may account for variation observed in models derived from large spatial surveys. Our results illustrate that while forecast models can be developed to determine when cyanobacteria will dominate within two temperate lakes, the models require detailed, lake-specific calibration to be effective as risk-management tools.