Current water quality guidelines across North America and Europe do not protect lakes from salinization.

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.

Habitat isolation reduces intra- and interspecific biodiversity and stability.

Fragmentation is predicted to reduce biodiversity and stability by increasing habitat isolation and impeding dispersal among patches. These effects may manifest at both the interspecific and intraspecific levels, yet few studies have simultaneously explored dispersal effects across levels of organization. We used field mesocosm experiments to examine how habitat isolation (in the form of dispersal rate) alters inter- and intraspecific stability and diversity in local zooplankton communities. We observed effects of increasing dispersal rate at both the intra- and interspecific levels. Increasing dispersal increased local species diversity and reduced mean temporal variability of populations. At the intraspecific level, Daphnia pulex clonal diversity was enhanced by dispersal and mean temporal variability of clone abundances through time was reduced.

Eco-Evolutionary Dynamics in the Wild: Clonal Turnover and Stability in Daphnia Populations.

There is increasing recognition of the importance of rapid adaptation in the dynamics of populations and communities. While the effects of rapid adaptation on the stability of populations have been shown in experimental systems, demonstration of their impacts in natural populations are rare. We examined the relationship between clonal dynamics and population stability of natural Daphnia pulex populations experiencing seasonal environmental variation. We show that the degree of asynchrony in a population's clonal dynamics is tightly linked to its population-level stability. Populations whose clonal abundances were more asynchronous were more stable temporally. Variation in asynchrony was related to variability in primary productivity, and experiments using clones from the study populations revealed significant genotype by environment interactions in response to food level. This suggests that clonal turnover was not due to neutral dynamics alone but may be linked to variation in functional traits associated with resource acquisition and conversion.

Impacts of dispersal on rapid adaptation and dynamic stability of Daphnia in fluctuating environments.

Prior ecological research has shown that spatial processes can enhance the temporal stability of populations in fluctuating environments. Less explored is the effect of dispersal on rapid adaptation and its concomitant impact on population dynamics. For asexually reproducing populations, theory predicts that dispersal in fluctuating environments can facilitate asynchrony among clones and enhance stability by reducing temporal variability of total population abundance. This effect is predicted when clones exhibit heritable variation in environmental optima and when fluctuations occur asynchronously among patches. We tested this in the field using artificial ponds and metapopulations composed of a diverse assemblage of Daphnia pulex clones. We directly manipulated dispersal presence/absence and environmental fluctuations in the form of nutrient pulses. Consistent with predictions, dispersal enhanced temporal asynchrony among clones in the presence of nutrient pulses; this in turn stabilized population dynamics. This effect only emerged when patches experienced spatially asynchronous nutrient pulses (dispersal had no effect when patches were synchronously pulsed). Clonal asynchrony was driven by strong positive selection for a single clone that exhibited a performance advantage under conditions of low resource availability. Our work highlights the importance of dispersal as a driver of eco-evolutionary dynamics and population stability in variable environments.

Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystems.

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities.

The stabilizing effects of genetic diversity on predator-prey dynamics.

Heterogeneity among prey in their susceptibility to predation is a potentially important stabilizer of predator-prey interactions, reducing the magnitude of population oscillations and enhancing total prey population abundance. When microevolutionary responses of prey populations occur at time scales comparable to population dynamics, adaptive responses in prey defense can, in theory, stabilize predator-prey dynamics and reduce top-down effects on prey abundance. While experiments have tested these predictions, less explored are the consequences of the evolution of prey phenotypes that can persist in both vulnerable and invulnerable classes. We tested this experimentally using a laboratory aquatic system composed of the rotifer Brachionus calyciflorus as a predator and the prey Synura petersenii, a colony-forming alga that exhibits genetic variation in its propensity to form colonies and colony size (larger colonies are a defense against predators). Prey populations of either low initial genetic diversity and low adaptive capacity or high initial genetic diversity and high adaptive capacity were crossed with predator presence and absence. Dynamics measured over the last 127 days of the 167-day experiment revealed no effects of initial prey genetic diversity on the average abundance or temporal variability of predator populations. However, genetic diversity and predator presence/absence interactively affected prey population abundance and stability; diversity of prey had no effects in the absence of predators but stabilized dynamics and increased total prey abundance in the presence of predators. The size structure of the genetically diverse prey populations diverged from single strain populations in the presence of predators, showing increases in colony size and in the relative abundance of cells found in colonies. Our work sheds light on the adaptive value of colony formation and supports the general view that genetic diversity and intraspecific trait variation of prey can play a vital role in the short-term dynamics and stability of planktonic predator-prey systems.

Transient dynamics and the destabilizing effects of prey heterogeneity.

The presence of prey heterogeneity and weakly interacting prey species is frequently viewed as a stabilizer of predator-prey dynamics, countering the destabilizing effects of enrichment and reducing the amplitude of population cycles. However, prior model explorations have largely focused on long-term, dynamic attractors rather than transient dynamics. Recent theoretical work shows that the presence of prey that are defended from predation can have strongly divergent effects on dynamics depending on time scale: prey heterogeneity can counteract the destabilizing effects of enrichment on predator-prey dynamics at long time scales but strongly destabilize systems during transient phases by creating long periods of low predator/prey abundance and increasing extinction probability (an effect that is amplified with increasing enrichment). We tested these general predictions using a planktonic system composed of a zooplankton predator and multiple algal prey. We first parameterized a model of our system to generate predictions and tested these experimentally. Our results qualitatively supported several model predictions. During transient phases, presence of defended algal prey increased predator extinctions at low and high enrichment levels compared to systems with only a single edible prey. This destabilizing effect was moderated at higher dilution rates, as predicted by our model. When examining dynamics beyond initial oscillations, presence of the defended prey increased predator-prey temporal variability at high nutrient enrichment but had no effect at low nutrient levels. Our results highlight the importance of considering transient dynamics when assessing the role of stabilizing factors on the dynamics of food webs.

Environmental noise, genetic diversity and the evolution of evolvability and robustness in model gene networks.

The ability of organisms to adapt and persist in the face of environmental change is accepted as a fundamental feature of natural systems. More contentious is whether the capacity of organisms to adapt (or "evolvability") can itself evolve and the mechanisms underlying such responses. Using model gene networks, I provide evidence that evolvability emerges more readily when populations experience positively autocorrelated environmental noise (red noise) compared to populations in stable or randomly varying (white noise) environments. Evolvability was correlated with increasing genetic robustness to effects on network viability and decreasing robustness to effects on phenotypic expression; populations whose networks displayed greater viability robustness and lower phenotypic robustness produced more additive genetic variation and adapted more rapidly in novel environments. Patterns of selection for robustness varied antagonistically with epistatic effects of mutations on viability and phenotypic expression, suggesting that trade-offs between these properties may constrain their evolutionary responses. Evolution of evolvability and robustness was stronger in sexual populations compared to asexual populations indicating that enhanced genetic variation under fluctuating selection combined with recombination load is a primary driver of the emergence of evolvability. These results provide insight into the mechanisms potentially underlying rapid adaptation as well as the environmental conditions that drive the evolution of genetic interactions.

Dispersal promotes compensatory dynamics and stability in forced metacommunities.

Understanding the factors that govern the stability of populations and communities has gained increasing importance as habitat fragmentation and environmental perturbations continue to escalate due to human activities. Dispersal is commonly viewed as essential to the maintenance of diversity in spatially subdivided communities, but few experiments have explored how dispersal interacts with the spatiotemporal components of environmental perturbations to determine community-level stability. We examined these processes using an experimental planktonic system composed of three competing species of zooplankton. We subjected zooplankton metacommunities to varying levels of dispersal and pH perturbations that varied in their degree of spatial synchrony. We show that dispersal can reverse the destabilizing effects of environmental forcing when perturbations are spatially asynchronous. Asynchrony in pH perturbations generated spatially and temporally varying species refugia that promoted source-sink dynamics and allowed prolonged persistence of zooplankton species that were otherwise extirpated in synchronously varying metacommunities. This, in turn, increased local species diversity, promoted compensatory population dynamics, and enhanced local community-level stability. Our results indicate that patterns of spatial covariation in environmental variability are critical to predicting the effects of dispersal on the dynamics and persistence of communities.

Periodically forced food-chain dynamics: model predictions and experimental validation.

Despite the recognition of the importance of seasonal forcing in nature, remarkably few studies have theoretically explored periodically forced community dynamics. Here we employ a novel approach called "successional state dynamics" (SSD) to model a seasonally forced predator-prey system. We first generated analytical predictions of the effects of altered seasonality on species persistence and the timing of community state transitions. We then parameterized the model using a zooplankton-phytoplankton system and tested quantitative predictions using controlled experiments. In the majority of cases, timing of zooplankton and algal population peaks matched model predictions. Decreases in growing-period length delayed algal blooms, consequently delaying peaks in zooplankton abundance. Predictions of increased probability of predator extinction at low growing-period lengths were also upheld experimentally. Our results highlight the utility of the SSD modeling approach as a framework for predicting the effects of altered seasonality on the structure and dynamics of multitrophic communities.

Resurrecting the ghost of competition past with dormant zooplankton eggs.

A common prediction of evolutionary theory is that the strength of interspecific competition should decline over time among sympatric populations of competing species. Here we provide experimental evidence of historical declines in competition effects among competing zooplankton populations. Using diapausing eggs, we resurrected clones of three species of zooplankton obtained from different periods of community assembly in a single lake. We show that clones of Daphnia ambigua obtained from early in assembly when D. ambigua was dominant became extinct in competition with clones of Daphnia pulicaria and Daphnia dentifera (the current lake dominants). In contrast, D. ambigua clones obtained from later in the lake's history experienced weaker competition effects and persisted with D. dentifera. While we cannot rule out the role of intraspecific competition within D. ambigua, our results are in line with the view that natural selection favors reduced interaction strength among co-occurring species, facilitating coexistence and population persistence.

Species richness and allometric scaling jointly determine biomass in model aquatic food webs.

1. Allometric theory makes specific predictions about how density, and consequently biomass, scale with organism size within trophic levels, across trophic levels and across food webs. 2. Diversity-yield relationships suggest that more diverse food webs can sometimes support more biomass through mechanisms involving niche complementarity or selection effects that are sometimes attributed to organism size. 3. We combine the above two approaches and show that, generally, density and biomass scale with organism size within and between trophic levels as predicted by allometric theory. Further, food webs converged in total biomass despite persistent differences in the composition and size of the organisms among food webs; species richness explained deviations from the constant yield of biomass expected from size-abundance relationships. 4. Our results suggest that organism size plays only a transient role in controlling community biomass because population increases or decreases lead to rapid convergence in biomass. Species richness affects community biomass independently by effectively increasing the mass of organisms that can be supported in a given productivity regime.

Population and community resilience in multitrophic communities.

Diversity-stability relationships have long been a topic of controversy in ecology, but one whose importance has been re-highlighted by increasing large-scale threats to global biodiversity. The ability of a community to recover from a perturbation (or resilience) is a common measure of stability that has received a large amount of theoretical attention. Yet, general expectations regarding diversity-resilience relations remain elusive. Moreover, the effects of productivity and its interaction with diversity on resilience are equally unclear. We examined the effects of species diversity, species composition, and productivity on population-and community-level resilience in experimental aquatic food webs composed of bacteria, algae, heterotrophic protozoa, and rotifers. Productivity manipulations were crossed with manipulations of the number of species and species compositions within trophic groups. Resilience was measured by perturbing communities with a nonselective, density-independent, mortality event and comparing responses over time between perturbed communities and controls. We found evidence that species diversity can enhance resilience at the community level (i.e., total community biomass), though this effect was more strongly expressed in low-productivity treatments. Diversity effects on resilience were driven by a sampling/selection effect, with resilient communities showing rapid response and dominance by a minority of species (primarily unicellular algae). In contrast, diversity had no effect on mean population-level resilience. Instead, the ability of a community's populations to recover from perturbations was dependent on species composition. We found no evidence of an effect of productivity, either positive or negative, on community- or population-level resilience. Our results indicate that the role of diversity as an insurer of stability may depend on the level of biological organization at which stability is measured, with effects emerging only when focusing on aggregate community properties.

Context-dependent effects of Daphnia pulex on pond ecosystem function: observational and experimental evidence.

Large-bodied zooplankton of the genus Daphnia are thought to be keystone species in freshwater pelagic habitats, potentially able to exert strong grazing effects and enhance phosphorus limitation of algae. I examined the degree to which Daphnia pulex differ from small-bodied zooplankton in their effects on algal biomass, seston C:P and N:P, total nitrogen and total phosphorus. This was done with both survey data from natural ponds and an in situ experiment in which D. pulex was compared to a small zooplankton assemblage under low and high nutrient conditions and in two different ponds. D. pulex effects on algae were only evident under high nutrient conditions. In natural ponds, D. pulex dominance resulted in a significantly weaker chlorophyll-total phosphorus relationship, with the divergence between D. pulex and small zooplankton-dominated systems being greatest in highly enriched ponds. In the experiment, D. pulex exerted stronger top-down control in enriched treatments only and tended to graze algae to lower levels in the more productive pond. Dynamics of C:P over the course of the experiment did not reveal strong effects of zooplankton composition. However, data on the final date of the experiment provided some evidence that D. pulex can enhance phosphorus limitation of algae; total phosphorus was lower and C:P higher in D. pulex treatments. Survey results revealed no effects of D. pulex on seston C:P or N:P, suggesting that this species may not be an important factor governing phosphorus limitation of algae in natural ponds.