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Human activities and climate change cause abiotic factors to fluctuate through time, sometimes passing thresholds for organismal reproduction and survival. Multiple stressors can independently or interactively impact organisms; however, few studies have examined how they interact when they overlap spatially but occur asynchronously. Fluctuations in salinity have been found in freshwater habitats worldwide. Meanwhile, heatwaves have become more frequent and extreme. High salinity pulses and heatwaves are often decoupled in time but can still collectively impact freshwater zooplankton. The time intervals between them, during which population growth and community recovery could happen, can influence combined effects, but no one has examined these effects. We conducted a mesocosm experiment to examine how different recovery times (0-, 3-, 6-week) between salt treatment and heatwave exposure influence their combined effects. We hypothesized that antagonistic effects would appear when having short recovery time, because previous study found that similar species were affected by the two stressors, but effects would become additive with longer recovery time since fully recovered communities would respond to heatwave similar to undisturbed communities. Our findings showed that, when combined, the two-stressor joint impacts changed from antagonistic to additive with increased recovery time between stressors. Surprisingly, full compositional recovery was not achieved despite a recovery period that was long enough for population growth, suggesting legacy effects from earlier treatment. The recovery was mainly driven by small organisms, such as rotifers and small cladocerans. As a result, communities recovering from previous salt exposure responded differently to heatwaves than undisturbed communities, leading to similar zooplankton communities regardless of the recovery time between stressors. Our research bolsters the understanding and management of multiple-stressor issues by revealing that prior exposure to one stressor has long-lasting impacts on community recovery that can lead to unexpected joint effects of multiple stressors., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Stressors can interact to affect animal fitness, but we have limited knowledge about how temporal variation in stressors may impact their combined effect. This limits our ability to predict the outcomes of pollutants and future dynamic environmental changes. Elevated salinity in freshwater ecosystems has been observed worldwide. Meanwhile, heatwaves have become more frequent and intensified as an outcome of climate change. These two stressors can jointly affect organisms; however, their interaction has rarely been explored in the context of freshwater ecosystems. We conducted lab experiments using Daphnia pulicaria, a key species in lakes, to investigate how elevated salinity and heatwave conditions collectively affect freshwater organisms. We also monitored the impacts of various recovery times between the two stressors. Daphnia physiological conditions (metabolic rate, Na + -K + -ATPase (NKA) activity, and lipid peroxidation level) and life history traits (survival, fecundity, and growth) in response to salt stress as well as mortality in heat treatment were examined. We found that Daphnia responded to elevated salinity by upregulating NKA activity and increasing metabolic rate, causing a high lipid peroxidation level. Survival, fecundity, and growth were all negatively affected by this stressor. These impacts on physiological conditions and life history traits persisted for a few days after the end of the exposure. Heat treatments caused mortality in Daphnia, which increased with rising temperature. Results also showed that individuals that experienced salt exposure were more susceptible to subsequent heat stress, but this effect decreased with increasing recovery time between stressors. Findings from this work suggest that the legacy effects from a previous stressor can reduce individual resistance to a subsequent stressor, adding great difficulties to the prediction of outcomes of multiple stressors. Our work also demonstrates that cross-tolerance/susceptibility and the associated mechanisms remain unclear, necessitating further investigation., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Recreational boating activities enable aquatic invasive species (AIS) dispersal among disconnected lakes, as invertebrates and plants caught on or contained within watercraft and equipment used in invaded waterbodies can survive overland transport. Besides simple preventive measures such as "clean, drain, dry", resource management agencies recommend decontaminating watercraft and equipment using high water pressure, rinsing with hot water, or air-drying to inhibit this mode of secondary spread. There is a lack of studies assessing the efficacy of these methods under realistic conditions and their feasibility for recreational boaters. Hence, we addressed this knowledge gap via experiments on six invertebrate and plant AIS present in Ontario. Washing at high pressures of 900-1200 psi removed the most biological material (90%) from surfaces. Brief (< 10 s) exposure to water at ≥ 60 °C caused nearly 100% mortality among all species tested, except banded mystery snails. Acclimation to temperatures from 15 to 30 °C before hot water exposure had little effect on the minimum temperature required for no survival. Air-drying durations producing complete mortality were ≥ 60 h for zebra mussels and spiny waterfleas, and ≥ 6 days among plants, whereas survival remained high among snails after a week of air-drying. Hot water exposure followed by air-drying was more effective than either method separately against all species tested., (© 2023. The Author(s).)

The widespread salinisation of freshwater ecosystems poses a major threat to the biodiversity, functioning, and services that they provide. Human activities promote freshwater salinisation through multiple drivers (e.g., agriculture, resource extraction, urbanisation) that are amplified by climate change. Due to its complexity, we are still far from fully understanding the ecological and evolutionary consequences of freshwater salinisation. Here, we assess current research gaps and present a research agenda to guide future studies. We identified different gaps in taxonomic groups, levels of biological organisation, and geographic regions. We suggest focusing on global- and landscape-scale processes, functional approaches, genetic and molecular levels, and eco-evolutionary dynamics as key future avenues to predict the consequences of freshwater salinisation for ecosystems and human societies., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Throughout much of the 20th century, unprecedented industrial emissions have led to widespread acidification of regions in North America and Europe and, as lake water pH dropped, aquatic ecosystems have experienced dramatic declines in biodiversity. International emission-control agreements have led to sweeping increases in lake pH, however acid-structured zooplankton communities still persist in many lakes. Concomitantly, calcium concentrations have been declining as a legacy of acidification and are approaching or have reached concentrations that could represent a barrier to the re-establishment of zooplankton communities similar to those in non-acidified or circumneutral reference lakes. To understand how declining calcium may influence the re-establishment of zooplankton in acid-damaged lakes we manipulated calcium and pH using a factorial in-lake mesocosm experiment and assessed their individual and combined effects on a regionally diverse zooplankton assemblage. We found that the impacts of low calcium on zooplankton species were similar to those of acidification and, consequently, may prevent the recovery of acid-structured communities. Abundance of the larger bodied and acid-sensitive Daphnia pulex/pulicaria increased in high pH treatments, albeit nonsignificantly yet, by the end of our experiment, only two individuals were sampled among our 10 low calcium enclosures. In contrast, small acid-tolerant cladocerans, such as Daphnia catawba, Daphnia ambigua, and eubosminids maintained significantly higher abundances in low calcium treatments relative to all other treatment combinations. Although we did not detect an effect of calcium on mean body size, the disproportionately high abundance of small cladocerans in low calcium treatments resulted in low calcium communities with higher overall abundance and lower cladoceran evenness. Our results, along with a landscape comparison demonstrating parallel changes in zooplankton relative abundance from 34 historically acidified lakes, suggests that declining calcium will be an important, on-going factor that may limit the recovery of zooplankton, despite regional improvements in lake pH., (© 2021 The Ecological Society of America.)

Human-induced salinization caused by the use of road deicing salts, agricultural practices, mining operations, and climate change is a major threat to the biodiversity and functioning of freshwater ecosystems. Yet, it is unclear if freshwater ecosystems are protected from salinization by current water quality guidelines. Leveraging an experimental network of land-based and in-lake mesocosms across North America and Europe, we tested how salinization-indicated as elevated chloride (Cl - ) concentration-will affect lake food webs and if two of the lowest Cl - thresholds found globally are sufficient to protect these food webs. Our results indicated that salinization will cause substantial zooplankton mortality at the lowest Cl - thresholds established in Canada (120 mg Cl - /L) and the United States (230 mg Cl - /L) and throughout Europe where Cl - thresholds are generally higher. For instance, at 73% of our study sites, Cl - concentrations that caused a ≥50% reduction in cladoceran abundance were at or below Cl - thresholds in Canada, in the United States, and throughout Europe. Similar trends occurred for copepod and rotifer zooplankton. The loss of zooplankton triggered a cascading effect causing an increase in phytoplankton biomass at 47% of study sites. Such changes in lake food webs could alter nutrient cycling and water clarity and trigger declines in fish production. Current Cl - thresholds across North America and Europe clearly do not adequately protect lake food webs. Water quality guidelines should be developed where they do not exist, and there is an urgent need to reassess existing guidelines to protect lake ecosystems from human-induced salinization., Competing Interests: The authors declare no competing interest.

Salinization of freshwater ecosystems is a growing hazard for organisms and ecosystem functioning worldwide. In northern latitudes, road salt that is being transported into water bodies can cause year-round increases in lake salinity levels. Exploring the environmental factors driving the susceptibility of freshwater zooplankton to road salt is crucial for assessing the impact of salinization on food web processes. We studied the role of essential lipids, i.e., sterols and long-chain polyunsaturated fatty acids (PUFAs), in mediating salt tolerance of the freshwater keystone herbivore Daphnia. Sterols and PUFAs are involved in regulating ion permeability of biological membranes and thus we hypothesized that the susceptibility to salt is affected by the dietary sterol and PUFA supply. Life history experiments revealed opposing effects of sterol and PUFA supplementation on salt tolerance, i.e., tolerance increased upon sterol supplementation but decreased upon PUFA supplementation, which is consistent with their proposed impact on membrane permeability. Our results suggest that the susceptibility of freshwater zooplankton to salinization strongly depends on the dietary lipid supply and thus the phytoplankton community composition. Hence, trophic state related differences in the phytoplankton community composition need to be considered when assessing the consequences of salinization for freshwater ecosystem functioning., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Widespread use of NaCl for road deicing has caused increased chloride concentrations in lakes near urban centers and areas of high road density. Chloride can be toxic, and water quality guidelines have been created to regulate it and protect aquatic life. However, these guidelines may not adequately protect organisms in low-nutrient, soft water lakes such as those underlain by the Precambrian Shield. We tested this hypothesis by conducting laboratory experiments on six Daphnia species using a soft water culture medium. We also examined temporal changes in cladoceran assemblages in the sediments of two small lakes on the Canadian Shield: one near a highway and the other >3 km from roads where salt is applied in the winter. Our results showed that Daphnia were sensitive to low chloride concentrations with decreased reproduction and increased mortality occurring between 5 and 40 mg Cl - /L. Analysis of cladoceran remains in lake sediments revealed changes in assemblage composition that coincided with the initial application of road salt in this region. In contrast, there were no changes detected in the remote lake. We found that 22.7% of recreational lakes in Ontario have chloride concentrations between 5 and 40 mg/L suggesting that cladoceran zooplankton in these lakes may already be experiencing negative effects of chloride.

The invasive species of the genus Bythotrephes introduced in the North American lakes in 1970s-early 1980s was named variously either B. cederstroemi or B. longimanus. The investigation of individuals of the genus from 15 Canadian lakes has allowed us to identify all of them as B. cederströmii Schödler, based on the detailed taxonomic redescription of the Eurasian representatives of the species. The available documented data (figures and photographs) on Bythotrephes from other North American lakes, made it possible to recognize their identity with this species as well. The possible introduction of interspecific hybrids of the genus Bythotrephes in the North American lakes has not been confirmed.

Researchers have long viewed patterns of species association as key to understanding the processes that structure communities. Community-level tests of species association have received the most attention; however, pairwise species associations may offer greater opportunity for linking patterns to specific mechanisms. Although several tests of pairwise association have been developed, there remain gaps in our understanding of their performance. Consequently, it is unclear whether these methods reliably detect patterns of association, or if any one method is superior. We maximized association patterns for single species pairs in synthetic community matrices and examined how accurately five pairwise association tests found that pair, while not finding others (i.e., type I and II error rates). All tests are more likely to miss patterns of association than to falsely detect them. When we maximized association for a species pair that included one or more rare or common species, tests were frequently unable to identify that pair as significantly associated. Consequently, these tests are best suited for identifying significant associations between pairs of species that occur in an intermediate number of samples; for such pairs, three of the five tests considered here detected 100% of the pairs for which we maximized associations., (© 2019 by the Ecological Society of America.)

Calcium levels are declining in eastern North American and western European lakes. This widespread issue is affecting the composition of crustacean zooplankton communities, as the presence and abundance of several calcium-rich daphniid species are declining, while two other daphniids, D. catawba and D. ambigua, that apparently tolerate low calcium environments, are prospering. The physiological basis for low calcium tolerance of these daphniids is unknown. In this study the presence of one Ca-rich (D. pulicaria) and one Ca-poor (D. ambigua) daphniid species in Canadian Shield lakes is assessed in relation to lake water Ca levels. The occurrence of D. ambigua was independent of Ca levels in Ontario lakes, whereas D. pulicaria was more likely to occur in lakes with relatively more Ca. In the laboratory, D. ambigua maintained lower levels of hemolymph Ca 2+ across a range of low Ca levels (0.7 to 7 mg l -1 ) compared with D. pulicaria. The hemolymph pH remained steady across this Ca gradient in D. ambigua while it was significantly more acidic in D. pulicaria in the two lowest Ca treatments. While Ca 2+ uptake was observed adjacent to the surface of D. ambigua individuals, Ca 2+ loss was observed for D. pulicaria assayed under moderately high Ca levels. Based on these observations we propose that D. ambigua is able to survive in low Ca lakes by maintaining low free ionic Ca 2+ levels in the hemolymph which minimizes the Ca gradient across the body wall in low Ca water thus limiting overall Ca loss and facilitating Ca 2+ uptake., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Assessing biological recovery in damaged aquatic environments requires the consideration of multiple spatial and temporal scales. Past research has focused on assessing lake recovery from atmospheric or catchment disturbance at regional or catchment levels. Studies have also rarely considered the influences of adjacent terrestrial characteristics on within-lake habitats, such as subcatchment delta confluences. We used Hyalella azteca, a ubiquitous freshwater amphipod, as a sensitive indicator to assess the importance of local subcatchment scale factors in the context of multiscale lake recovery within the metal mining region of Sudbury, Canada following a period of major reductions in atmospheric pollution. At the regional scale, data from repeated surveys of 40 lakes showed higher probabilities of H. azteca occurrence with higher lake water conductivity, alkalinity, and pH and lower metal concentrations. The importance of metals decreased through time and the importance of higher conductivity, alkalinity, and pH increased. At the subcatchment scale, a subset of six lakes sampled across a colonization gradient revealed higher H. azteca abundances at subcatchment delta sites than non-delta sites in early colonization stages, and that abundance at delta sites was correlated with both within-lake habitat and terrestrial subcatchment characteristics. For example, wetland cover reduced the strength of positive associations between H. azteca abundance and macrophyte density. A single lake from this subset also revealed higher abundances at delta sites associated with higher concentrations of terrestrial organic matter and larger subcatchments. Our results demonstrate that factors affecting recovery can change with the scale of study, and that managing terrestrial-aquatic linkages is important for facilitating recovery processes within damaged lake ecosystems., (© 2017 by the Ecological Society of America.)

Colonist quantity, quality, and arrival frequency can all individually drive the dynamics and extinction of new populations. However, we do not understand which has the strongest influence, nor the circumstances under which their relative importance may change. We conducted a field mesocosm experiment that manipulated colonist quantity, quality, and arrival frequency in two zooplankton species (Daphnia pulicaria and Skistodiaptomus oregonensis). Individuals of each species were cultured under either high or low food concentrations to produce, respectively, 'good' and 'poor' quality colonists. Each species was then introduced at either small (2 individuals) or large introduction quantities (8 individuals) divided over single or multiple introduction events. We found that the extinction of Daphnia pulicaria was not particularly affected by any of our treatments. Introductions of just two individuals performed as well as larger or more frequent introductions, regardless of quality. Conversely, Skistodiaptomus oregonensis extinction was strongly driven by arrival frequency. Populations that arrived in a single event exhibited high rates of extinction (75-83%), with this probability declining dramatically when colonists were introduced over multiple events (33% extinction). Our results show that other less studied aspects of the colonist pool, such as arrival frequency, could be as important to population persistence as the initial quantity of arriving colonists. Additionally, there are potentially numerous species that are well suited to succeeding with a small number of founders, and whose success is therefore not necessarily dependent upon colonist quantity, quality, or arrival frequency.

Aquatic ecosystems depend on terrestrial organic matter (tOM) to regulate many functions, such as food web production and water quality, but an increasing frequency and intensity of drought across northern ecosystems is threatening to disrupt this important connection. Dry conditions reduce tOM export and can also oxidize wetland soils and release stored contaminants into stream flow after rainfall. Here, we test whether these disruptions to terrestrial-aquatic linkages occur during mild summer drought and whether this affects biota across 43 littoral zone sites in 11 lakes. We use copper (Cu) and nickel (Ni) as representative contaminants, and measure abundances of Hyalella azteca, a widespread indicator of ecosystem condition and food web production. We found that tOM concentrations were reduced but correlations with organic soils (wetlands and riparian forests) persisted during mild drought and were sufficient to suppress labile Cu concentrations. Wetlands, however, also became a source of labile Ni to littoral zones, which was linked to reduced abundances of the amphipod H. azteca, on average by up to 70 times across the range of observed Ni concentrations. This reveals a duality in the functional linkage of organic soils to aquatic ecosystems whereby they can help buffer the effects of hydrologic disconnection between catchments and lakes but at the cost of biogeochemical changes that release stored contaminants. As evidence of the toxicity of trace contaminant concentrations and their global dispersion grows, sustaining links among forests, organic soils and aquatic ecosystems in a changing climate will become increasingly important., (© 2016 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

Negative co-occurrence patterns are intriguing because they may reflect the outcome of interspecific interactions and therefore signal how competition shapes communities. However, other factors also contribute to these patterns. For example, theoretical studies as well as two survey-based studies have all suggested that dispersal may also impact these patterns. While natural communities commonly have nonrandom patterns of negative co-occurrence, understanding how different processes drive these patterns requires further research. We tested the influence of dispersal on co-occurrence patterns using a zooplankton mesocosm experiment with four different dispersal treatments varying in the number of dispersers delivered into mesocosms on regular intervals. Our dispersal treatments were intended to adjust the relative importance of dispersal and competition experienced within mesocosms (i.e., high dispersal results in a relatively low influence of competition on species composition and vice versa). Higher dispersal translated into increased zooplankton species richness and inter-mesocosm compositional similarity, and also changed species occupancy patterns such that species occurrences were more even across mesocosms in higher-dispersal treatments. Dispersal treatments also differed markedly in species co-occurrence patterns. Negative co-occurrence patterns were significant for all but the lowest-dispersal treatment, peaked in the intermediate-dispersal treatments, and declined in the highest-dispersal treatment. Stability analyses illustrate that co-occurrence differences are robust to the exclusion of any single mesocosm in null model analyses. Dispersal treatments did not significantly differ with respect to abiotic variation, which has been recognized as a potential driver of negative co-occurrence patterns. These results suggest that not only can dispersal influence patterns of negative co-occurrence via changes to species richness and distribution (occupancy patterns among mesocosms), but the degree to which they do so varies nonlinearly with the strength of dispersal. Critically, because negative co-occurrence patterns were nonsignificant when the contribution of dispersal was lowest, it is possible that dispersal contributes strongly to many observed patterns of negative co-occurrence. Consequently, great care should be taken prior to interpreting significant co-occurrence tests as a product of species interactions.

Theory suggests that communities should be more open to the establishment of regional species following disturbance because disturbance may make more resources available to dispersers. However, after an initial period of high invasibility, growth of the resident community may lead to the monopolization of local resources and decreased probability of successful colonist establishment. During press disturbances (i.e., directional environmental change), it remains unclear what effect regional dispersal will have on local community structure if the establishment of later arriving species is affected by early arriving species (i.e., if priority effects are important). To determine the relationship between time-since-disturbance and invasibility, we conducted a fully factorial field mesocosm experiment that exposed tundra zooplankton communities to two emerging stressors - nutrient and salt addition, and manipulated the arrival timing of regional dispersers. Our results demonstrate that invasibility decreases with increasing time-since-disturbance as abundance (nutrient treatments) or species richness (salt treatments) increases in the resident community. Results suggest that the relative timing of dispersal and environmental change will modify the importance of priority effects in determining species composition after a press disturbance.

Changing environmental conditions are affecting diversity and ecosystem function globally. Theory suggests that dispersal from a regional species pool may buffer against changes in local community diversity and ecosystem function after a disturbance through the establishment of functionally redundant tolerant species. The spatial insurance provided by dispersal may decrease through time after environmental change as the local community monopolizes resources and reduces community invasibility. To test for evidence of the spatial insurance hypothesis and to determine the role dispersal timing plays in this response we conducted a field experiment using crustacean zooplankton communities in a subarctic region that is expected to be highly impacted by climate change - Churchill, Canada. Three experiments were conducted where nutrients, salt, and dispersal were manipulated. The three experiments differed in time-since-disturbance that the dispersers were added. We found that coarse measures of diversity (i.e. species richness, evenness, and Shannon-Weiner diversity) were generally resistant to large magnitude disturbances, and that dispersal had the most impact on diversity when dispersers were added shortly after disturbance. Ecosystem functioning (chl-a) was degraded in disturbed communities, but dispersal recovered ecosystem function to undisturbed levels. This spatial insurance for ecosystem function was mediated through changes in community composition and the relative abundance of functional groups. Results suggest that regional diversity and habitat connectivity will be important in the future to maintain ecosystem function by introducing functionally redundant species to promote compensatory dynamics., (© 2012 Blackwell Publishing Ltd.)

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.

The acidification and ongoing pH recovery of lakes in Killarney Provincial Park, Canada, provide a unique opportunity to increase our understanding of the role of dispersal as communities respond to environmental change. Time lags in community recovery following pH increases in acidified lakes have typically been attributed to local factors; however, no studies have been conducted to determine if colonist availability could also play a role. Moreover, the rates and mechanisms of dispersal to recovering lakes are poorly understood. In this study, we sought to determine if dispersal limitation could impede the recovery of zooplankton communities affected by a regional stressor. To achieve this objective, we used a combination of empirical data collection along with spatial modeling and variation partitioning techniques. Data were collected by measuring dispersal to four recovering lakes in Killarney Park. Dispersal traps were placed next to lakes to measure immigration overland, drift nets were used to measure immigration via streams, and in situ emergence traps were used to quantify immigration from historically deposited resting eggs. Documented dispersal levels were then compared with the theoretical critical density required for reproduction (N(c)) to determine if adequate numbers were dispersing to establish populations of acid-sensitive species in recovering lakes. Spatial modeling and variation partitioning were conducted using community and physical/chemical data for 45 park lakes that were collected in 1972-1973, 1990, and 2005. Field data demonstrated that a variety of zooplankton species were dispersing to recovering lakes through streams and the egg bank, but few individuals were collected dispersing overland. Although we identified 24 species of zooplankton dispersing, only six species absent from the communities of our study lakes were identified from our traps, and two of these species did not disperse in high enough numbers to surpass N(c). Local environmental variables explained the largest proportion of the variation in zooplankton communities (18-37%); however, spatial variables were also important (7-18%). The significant spatial patterns we found in the park's zooplankton communities, combined with the low overland dispersal levels we documented, suggest that dispersal limitation may be a more important impediment to recovery than was previously thought.

Habitat connectivity and regional heterogeneity represent two factors likely to affect biodiversity across different spatial scales. We performed a 3 × 2 factorial design experiment to investigate the effects of connectivity, heterogeneity, and their interaction on artificial pond communities of freshwater invertebrates at the local (α), among-community (β), and regional (γ) scales. Despite expectations that the effects of connectivity would depend on levels of regional heterogeneity, no significant interactions were found for any diversity index investigated at any spatial scale. While observed responses of biodiversity to connectivity and heterogeneity depended to some extent on the diversity index and spatial partitioning formula used, the general pattern shows that these factors largely act at the β scale, as opposed to the α or γ scales. We conclude that the major role of connectivity in aquatic invertebrate communities is to act as a homogenizing force with relatively little effect on diversity at the α or γ levels. Conversely, heterogeneity acts as a force maintaining differences between communities.

In the event of an environmental disturbance, dispersal of native taxa may provide species and genetic diversity to ecosystems, increasing the likelihood that there will be species and genotypes present that are less vulnerable to the disturbance. This may allow communities to maintain functioning during a disturbance and may be particularly important when the perturbation is novel to the system, such as the establishment of an invasive species. We examined how dispersal of native species may influence crustacean zooplankton communities in freshwater lakes invaded by the invertebrate predator, Bythotrephes longimanus. Using large enclosures, we experimentally tested the effect of dispersal on zooplankton community abundance, richness, and composition in (1) a community invaded by Bythotrephes, (2) the same community with the invader removed, and (3) a community that was never invaded. Dispersal increased zooplankton community abundance and richness; however, these effects were usually only significant in the invader-removed treatment. Dispersal tended to make the invader-removed communities more similar to never-invaded communities in abundance, richness, and composition. Dispersal had little effect on zooplankton abundance in the invaded community; however, richness significantly increased, and the community composition changed to resemble a never-invaded community by the end of the experiment. Our results have implications for understanding the role of dispersal during transitory states in communities. Dispersal of native taxa may be particularly important during the period between the arrival and broad-scale establishment of Bythotrephes, as dispersal through space or time (i.e., from resting eggs) may rapidly increase zooplankton abundance when the invader is absent or in low abundances. Overall, our results suggest that communities with strong local predatory and competitive interactions may be closed to immigration from colonists, but that invasive species may alter the conditions under which species can establish. These results have implications for the interaction of native and invasive species across broad spatial scales, as regional dispersal of native taxa may forestall the local extirpation of native species. In particular, transient phases that result from variable persistence of invaders within habitats or across a region may permit native colonists to successfully establish, and thus increase local and regional resistance to future disturbance.

Ecological linkages between species diversity in communities and genetic diversity in populations have potential to influence the assembly of communities in habitats recovering from human disturbance, but few studies have attempted to synthesize relationships between these levels of biological organization, especially for locally adapted species. No such studies have been done in freshwater ecosystems despite the plethora of environmental stressors plaguing aquatic communities around the world. We present the first study to test (1) whether diversity and dissimilarity among communities and populations of a locally adapted species are correlated and (2) whether communities and population haplotypes respond differently to environmental selection and spatial structure of habitats. We used a fragment of mitochondrial DNA (mtDNA) belonging to the gene cytochrome oxidase subunit I (COI) as a neutral tag to discriminate among different population haplotype variants. In boreal lakes with different histories of exposure to anthropogenic acidification, diversity and dissimilarity metrics for crustacean zooplankton communities and locally adapted populations of an abundant and broadly distributed calanoid copepod species, Leptodiaptomus minutus, did not correlate. This discord was likely because zooplankton communities responded more strongly to acidity and acidity-related environmental variables than spatial structure of lakes, whereas the distribution of L. minutus haplotypes was more strongly governed by spatial structure of lakes than environmental selection. Although spatial structure was the dominant driver of haplotype structure among L. minutus lake populations, there were similarities in the types of environmental variables that influenced the distributions of species in communities and haplotypes in populations. How haplotype diversity among populations relates to community diversity depends on the relative influence of spatial structure of habitats and selection at each of these scales of biological organization.

Anthropogenic habitat disturbance can often lead to rapid evolution of environmental tolerances in taxa that are able to withstand the stressor. What we do not understand, however, is how species respond when the stressor no longer exists, especially across landscapes and over a considerable length of time. Once anthropogenic disturbance is removed and if there is an ecological trade-off associated with local adaptation to such an historical stressor, then evolutionary theory would predict evolutionary reversals. On the Boreal Shield, tens of thousands of lakes acidified as a result of SO2 emissions, but many of these lakes are undergoing chemical recovery as a consequence of reduced emissions. We investigated the adaptive consequences of disturbance and recovery to zooplankton living in these lakes by asking (1) if contemporary evolution of acid tolerance had arisen among Leptodiaptomus minutus copepod populations in multiple circum-neutral lakes with and without historical acidification, (2) if L. minutus populations were adaptively responding to reversals in selection in historically acidified lakes that had recovered to pH 6.0 for at least 6-8 years, and (3) if there was a fitness trade-off for L. minutus individuals with high acid tolerance at circum-neutral pH. L. minutus populations had higher acid tolerances in circum-neutral lakes with a history of acidification than in local and distant lakes that were never acidified. However, copepods in circum-neutral acid-recovering lakes were less acid-tolerant than were copepods in lakes with longer recovery time. This adaptive reversal in acid tolerance of L. minutus populations following lake recovery was supported by the results of a laboratory experiment that indicated a fitness trade-off in copepods with high acid tolerances at circum-neutral pH. These responses appear to have a genetic basis and suggest that L. minutus is highly adaptive to changes in environmental conditions. Therefore, restoration managers should focus on removing environmental stressors, and adaptable species will be able to reverse evolutionary responses to environmental disturbance in the years following recovery.

Analyses of temporal patterns of diversity across a wide range of taxa have found that more diverse communities often show smaller compositional changes over time. This generality indicates that high diversity is associated with greater temporal stability in species composition. We examined patterns of diversity and community stability in zooplankton time series data from 36 lakes sampled over a combined 483 years. The species-time relationship was flatter in more species-rich lakes in the temperate zone. However, high-latitude lakes had both low richness and low turnover. These patterns were consistent for turnover both within and among years. Daily, annual and long-term richness were all higher in large lakes while turnover was unaffected by the surface area. Richness on all time scales, as well as turnover within and among years, all declined at high latitude. Species-area relations and latitudinal gradients in richness therefore reflect different temporal components of diversity. Our results suggest that diversity shows strong associations with compositional stability that vary qualitatively across biogeographical provinces. Community stability increases with diversity among lakes in the temperate zone; however, the two are negatively correlated across latitudinal gradients. These patterns indicate that either the direct effects of diversity on stability or their covariance with environmental fluctuations vary with latitude.

Dispersal can be an important determinant of local diversity and species composition, but evidence for effects of the regional species pool on local zooplankton communities has been mixed. Theory and experiments suggest that immigration will be necessary for maintenance of community diversity and functioning during periods of environmental change; conversely, fluctuating resource levels may increase the likelihood of invasion success. We conducted a factorial-design mesocosm experiment to test the effects of a nutrient pulse and weekly immigration from other lakes on the diversity and composition of a pelagic zooplankton community. Contrary to expectations, there were no interactive effects of nutrient enrichment and immigration on any measure of diversity, and the initial shift in community composition in response to the nutrient pulse did not depend on the introduction of new species or genotypes from more productive lakes. Although immigration increased species richness in enclosures, success of most colonising species was poor. However, the dispersal treatment appears to have enabled a stronger predator response to increased herbivore numbers in nutrient-pulsed enclosures, leading to an eventual decline in the abundance of some herbivorous species in response to immigration. We conclude that community invasibility was not influenced by productivity, and that dispersal limitation did not strongly constrain the response of the zooplankton community to our applied disturbance. This indicates an unexpected resistance to change in species composition and diversity in spite of disturbance, and suggests that, in our study system, changes in the abundance of resident species are more important than introductions of new species in the community response to short-term environmental change.

Understanding the factors that affect biological recovery from environmental stressors such as acidification is an important challenge in ecology. Here we report on zooplankton community recovery following the experimental acidification of Little Rock Lake, Wisconsin, USA. One decade following cessation of acid additions to the northern basin of Little Rock Lake (LRL), recovery of the zooplankton community was complete. Approximately 40% of zooplankton species in the lake exhibited a recovery lag in which biological recovery to reference basin levels was delayed by 1-6 yr after pH recovered to the level at which the species originally responded. Delays in recovery such as those we observed in LRL may be attributable to "biological resistance" wherein establishment of viable populations of key acid-sensitive species following water quality improvements is prevented by other components of the community that thrived during acidification. Indeed, we observed that the recovery of species that thrived during acidification tended to precede recovery of species that declined during acidification. In addition, correspondence analysis indicated that the zooplankton community followed different pathways during acidification and recovery, suggesting that there is substantial hysteresis in zooplankton recovery from acidification. By providing an example of a relatively rapid recovery from short-term acidification, zooplankton community recovery from experimental acidification in LRL generally reinforces the positive outlook for recovery reported for other acidified lakes.

We identified some of the sources and sinks of monomethyl mercury (MMHg) and inorganic mercury (HgII) on Ellesmere Island in the Canadian High Arctic. Atmospheric Hg depletion events resulted in the deposition of Hg(II) into the upper layers of snowpacks, where concentrations of total Hg (all forms of Hg) reached over 20 ng/L. However, our data suggest that much of this deposited Hg(II) was rapidly photoreduced to Hg(0) which then evaded back to the atmosphere. As a result, we estimate that net wet and dry deposition of Hg(II) during winter was lower at our sites (0.4-5.9 mg/ha) than wet deposition in more southerly locations in Canada and the United States. We also found quite high concentrations of monomethyl Hg (MMHg) in snowpacks (up to 0.28 ng/L), and at times, most of the Hg in snowpacks was present as MMHg. On the Prince of Wales Icefield nearthe North Water Polynya, we observed a significant correlation between concentrations of Cl and MMHg in snow deposited in the spring, suggesting a marine source of MMHg. We hypothesize that dimethyl Hg fluxes from the ocean to the atmosphere through polynyas and open leads in ice, and is rapidly photolyzed to MMHgCl. We also found that concentrations of MMHg in initial snowmelt on John Evans Glacier (up to 0.24 ng/L) were higher than concentrations of MMHg in the snowpack (up to 0.11 ng/L), likely due to either sublimation of snow or preferential leaching of MMHg from snow during the initial melt phase. This springtime pulse of MMHg to the High Arctic, in conjunction with climate warming and the thinning and melting of sea ice, may be partially responsible for the increase in concentrations of Hg observed in certain Arctic marine mammals in recent decades. Concentrations of MMHg in warm and shallow freshwater ponds on Ellesmere Island were also quite high (up to 3.0 ng/L), leading us to conclude that there are very active regions of microbial Hg(II) methylation in freshwater systems during the short summer season in the High Arctic.

Quantifying chemical variability in different lake types is important for the assessment of both chemical and biological responses to environmental change. For monitoring programs that emphasize a large number of lakes at the expense of frequent samples, high variability may influence how representative single samples are of the average conditions of individual lakes. Intensive temporal data from long-term research sites provide a unique opportunity to assess chemical variability in lakes with different characteristics. We compared the intra- and inter-annual variability of four acidification related variables (Gran alkalinity, pH, sulphate concentration, and total base cation concentration) in four lakes with different flushing rates and acid deposition histories. Variability was highest in lakes with high flushing rates and was not influenced by historic acid deposition in our study lakes. This has implications for the amount of effort required in monitoring programs. Lakes with high flushing rates will require more frequent sampling intervals than lakes with low flushing rates. Consideration of specific lake types should be included in the design of monitoring programs.

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 degrees 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.

Surface water acidity is decreasing in large areas of Europe and North America in response to reductions in atmospheric S deposition, but the ecological responses to these water-quality improvements are uncertain. Biota are recovering in some lakes and rivers, as water quality improves, but they are not yet recovering in others. To make sense of these different responses, and to foster effective management of the acid rain problem, we need to understand 2 things: i) the sequence of ecological steps needed for biotic communities to recover; and ii) where and how to intervene in this process should recovery stall. Here our purpose is to develop conceptual frameworks to serve these 2 needs. In the first framework, the primarily ecological one, a decision tree highlights the sequence of processes necessary for ecological recovery, linking them with management tools and responses to bottlenecks in the process. These bottlenecks are inadequate water quality, an inadequate supply of colonists to permit establishment, and community-level impediments to recovery dynamics. A second, more management-oriented framework identifies where we can intervene to overcome these bottlenecks, and what research is needed to build the models to operationalize the framework. Our ability to assess the benefits of S emission reduction would be simplified if we had models to predict the rate and extent of ecological recovery from acidification. To build such models we must identify the ecological steps in the recovery process. The frameworks we present will advance us towards this goal.