Antipredator responses of three Daphnia species within the D. longispina species complex to two invertebrate predators.

Prey communities in natural environments face a diverse array of predators with distinct hunting techniques. However, most studies have focused only on the interactions between a single prey species and one or more predators and typically only one of many induced defense traits, which limits our understanding of the broader effects of predators on prey communities. In this study, we conducted a common garden experiment using five clones each of three Daphnia species (D. cucullata, D. galeata, and D. longispina) from the D. longispina species complex to investigate the plasticity of predator-induced defenses in response to two predators in a community ecology setting. Five clones from each species were subjected to predator kairomones from two closely related invertebrate predators that are common in several European lakes, Bythotrephes longimanus or Leptodora kindtii for a duration of 10 days, and the morphological traits of body size, head size, spina size, and the presence of spinules on the ventral and dorsal carapace margins were measured. We show that among the species within this species complex there are different antipredator reactions to the invertebrate predators. The induced responses exhibited were species, trait, and predator-specific. Notably, D. galeata and D. cucullata developed distinctive helmets as defensive mechanisms, while microdefenses were induced in D. galeata and D. longispina, but not in D. cucullata. This demonstrates that the expression of micro- and macrodefenses across species was unrelated, highlighting the possible independent evolution of microstructures as defensive modules in Daphnia's antipredator strategies. This study is the first to document both micro- and macrodefensive phenotypic plasticity in three co-occurring Daphnia species within the D. longispina species complex. The differences in inducible defenses may have a substantial impact on how these three species cohabit with Bythotrephes and Leptodora.

Impact of climate warming on phenological asynchrony of plankton dynamics across Europe.

Climate warming alters the seasonal timing of biological events. This raises concerns that species-specific responses to warming may de-synchronize co-evolved consumer-resource phenologies, resulting in trophic mismatch and altered ecosystem dynamics. We explored the effects of warming on the synchrony of two events: the onset of the phytoplankton spring bloom and the spring/summer maximum of the grazer Daphnia. Simulation of 16 lake types over 31 years at 1907 North African and European locations under 5 climate scenarios revealed that the current median phenological delay between the two events varies greatly (20-190 days) across lake types and geographic locations. Warming moves both events forward in time and can lengthen or shorten the delay between them by up to ±60 days. Our simulations suggest large geographic and lake-specific variations in phenological synchrony, provide quantitative predictions of its dependence on physical lake properties and geographic location and highlight research needs concerning its ecological consequences.

Can young-of-the-year invasive fish keep up with young-of-the-year native fish? A comparison of feeding rates between invasive sticklebacks and whitefish.

Invasion of non-native species might alter food web structure and the strength of top-down control within lake ecosystems. As top-down control exerted by fish populations is often dominated by young of the year fish, the impact of new fish species might depend on the feeding rates of the juvenile fish. Here we provide comparative analyses of feeding rates of juvenile whitefish (Coregonus wartmanni) - a native and specialised planktivore and an invasive generalist (sticklebacks, Gasterosteus aculeatus). We studied feedings rates of whitefish and sticklebacks in aquaria experiments using 2 cm to 8 cm fish feeding on seven zooplankton species common to Lake Constance. As whitefish hatch several months earlier than sticklebacks, 0+ whitefish are larger than 0+ sticklebacks throughout the year and hence are predicted to have higher feeding rates on especially large zooplankton species. We show that sticklebacks as small as 2 cm were able to feed on the largest zooplankton species of Lake Constance. Further, stickleback feeding rates were similar to both the same size 0+ whitefish and the larger 0+ whitefish co-occurring with smaller 0+ sticklebacks. Hence, 0+ sticklebacks will compete with 0+ whitefish for the same zooplankton species, therefore the invasion of sticklebacks is unlikely to change the relative feeding pressure by individual 0+ fish on zooplankton species.

Reversed evolution of grazer resistance to cyanobacteria.

Exploring the capability of organisms to cope with human-caused environmental change is crucial for assessing the risk of extinction and biodiversity loss. We study the consequences of changing nutrient pollution for the freshwater keystone grazer, Daphnia, in a large lake with a well-documented history of eutrophication and oligotrophication. Experiments using decades-old genotypes resurrected from the sediment egg bank revealed that nutrient enrichment in the middle of the 20th century, resulting in the proliferation of harmful cyanobacteria, led to the rapid evolution of grazer resistance to cyanobacteria. We show here that the subsequent reduction in nutrient input, accompanied by a decrease in cyanobacteria, resulted in the re-emergence of highly susceptible Daphnia genotypes. Expression and subsequent loss of grazer resistance occurred at high evolutionary rates, suggesting opposing selection and that maintaining resistance was costly. We provide a rare example of reversed evolution of a fitness-relevant trait in response to relaxed selection.

Local and continental-scale controls of the onset of spring phytoplankton blooms: Conclusions from a proxy-based model.

A key phenological event in the annual cycle of many pelagic ecosystems is the onset of the spring algal bloom (OAB). Descriptions of the factors controlling the OAB in temperate to polar lakes have been limited to isolated studies of single systems and conceptual models. Here we present a validated modelling approach that, for the first time, enables a quantitative prediction of the OAB and a systematic assessment of the processes controlling its timing on a continental scale. We used a weather-driven, one-dimensional lake model to simulate the seasonal dynamics of the underwater light climate in 16 lake types characterized by the factorial combination of four lake depths with four levels of water transparency. We did so at 1962 locations across Western Europe and over 31 years (1979-2009). Assuming that phytoplankton production is light-limited in winter, we identified four patterns of OAB control across lake types and climate zones. OAB timing is controlled by (i) the timing of ice-off in ice-covered clear or shallow lakes, (ii) the onset of thermal stratification in sufficiently deep and turbid lakes and (iii) the seasonal increase in incident radiation in all other lakes, except for (iv) ice-free, shallow and clear lakes in the south, where phytoplankton is not light-limited. The model predicts that OAB timing should respond to two pervasive environmental changes, global warming and browning, in opposite ways. OAB timing should be highly sensitive to warming in lakes where it is controlled by either ice-off or the onset of stratification, but resilient to warming in lakes where it is controlled by incident radiation. Conversely, OAB timing should be most sensitive to browning where it is controlled by incident radiation, but resilient to browning where it is controlled by ice-off or the onset of stratification. Available lake data are consistent with our findings.

Increased winter drownings in ice-covered regions with warmer winters.

Winter activities on ice are culturally important for many countries, yet they constitute a high safety risk depending upon the stability of the ice. Because consistently cold periods are required to form stable and thick ice, warmer winters could degrade ice conditions and increase the likelihood of falling through the ice. This study provides the first large-scale assessment of winter drowning from 10 Northern Hemisphere countries. We documented over 4000 winter drowning events. Winter drownings increased exponentially in regions with warmer winters when air temperatures neared 0°C. The largest number of drownings occurred when winter air temperatures were between -5°C and 0°C, when ice is less stable, and also in regions where indigenous traditions and livelihood require extended time on ice. Rates of drowning were greatest late in the winter season when ice stability declines. Children and adults up to the age of 39 were at the highest risk of winter drownings. Beyond temperature, differences in cultures, regulations, and human behaviours can be important additional risk factors. Our findings indicate the potential for increased human mortality with warmer winter air temperatures. Incorporating drowning prevention plans would improve adaptation strategies to a changing climate.

Storm impacts on phytoplankton community dynamics in lakes.

In many regions across the globe, extreme weather events such as storms have increased in frequency, intensity, and duration due to climate change. Ecological theory predicts that such extreme events should have large impacts on ecosystem structure and function. High winds and precipitation associated with storms can affect lakes via short-term runoff events from watersheds and physical mixing of the water column. In addition, lakes connected to rivers and streams will also experience flushing due to high flow rates. Although we have a well-developed understanding of how wind and precipitation events can alter lake physical processes and some aspects of biogeochemical cycling, our mechanistic understanding of the emergent responses of phytoplankton communities is poor. Here we provide a comprehensive synthesis that identifies how storms interact with lake and watershed attributes and their antecedent conditions to generate changes in lake physical and chemical environments. Such changes can restructure phytoplankton communities and their dynamics, as well as result in altered ecological function (e.g., carbon, nutrient and energy cycling) in the short- and long-term. We summarize the current understanding of storm-induced phytoplankton dynamics, identify knowledge gaps with a systematic review of the literature, and suggest future research directions across a gradient of lake types and environmental conditions.

A single Thaumarchaeon drives nitrification in deep oligotrophic Lake Constance.

Ammonia released during organic matter mineralization is converted during nitrification to nitrate. We followed spatiotemporal dynamics of the nitrifying microbial community in deep oligotrophic Lake Constance. Depth-dependent decrease of total ammonium (0.01-0.84 µM) indicated the hypolimnion as the major place of nitrification with 15 N-isotope dilution measurements indicating a threefold daily turnover of hypolimnetic total ammonium. This was mirrored by a strong increase of ammonia-oxidizing Thaumarchaeota towards the hypolimnion (13%-21% of bacterioplankton) throughout spring to autumn as revealed by amplicon sequencing and quantitative polymerase chain reaction. Ammonia-oxidizing bacteria were typically two orders of magnitude less abundant and completely ammonia-oxidizing (comammox) bacteria were not detected. Both, 16S rRNA gene and amoA (encoding ammonia monooxygenase subunit B) analyses identified only one major species-level operational taxonomic unit (OTU) of Thaumarchaeota (99% of all ammonia oxidizers in the hypolimnion), which was affiliated to Nitrosopumilus spp. The relative abundance distribution of the single Thaumarchaeon strongly correlated to an equally abundant Chloroflexi clade CL500-11 OTU and a Nitrospira OTU that was one order of magnitude less abundant. The latter dominated among recognized nitrite oxidizers. This extremely low diversity of nitrifiers shows how vulnerable the ecosystem process of nitrification may be in Lake Constance as Central Europe's third largest lake.

Resilience to changes in lake trophic state: Nutrient allocation into Daphnia resting eggs.

During past decades, many lakes underwent drastic human-caused changes in trophic state with strong implications for population dynamics and food web processes. We investigated the influence of trophic state on nutrient allocation into Daphnia resting eggs. The production of resting eggs is an important survival strategy, allowing Daphnia to cope with unfavorable environmental conditions. Allocation of essential nutrients into resting eggs may crucially influence embryonic development and offspring survival and thus is of great ecological and evolutionary interest. The capacity of Daphnia to adjust the allocation of nutrients into resting eggs may depend on the dietary nutrient supply, which may vary with trophic state-related changes in the phytoplankton community composition. Resting eggs were isolated from sediment cores taken from Lake Constance, a large prealpine lake with a distinct eutrophication and reoligotrophication history, and analyzed for elemental (carbon, nitrogen, and phosphorus) and biochemical (sterols and fatty acids) nutrients. Carbon allocation into Daphnia resting eggs continuously decreased over time, irrespective of changes in trophic state. The allocation of nitrogen into Daphnia resting eggs followed the changes in trophic state, that is, nitrogen concentrations in resting eggs increased with eutrophication and decreased again with reoligotrophication. The allocation of phosphorus, sterols and long-chain polyunsaturated fatty acids, such as eicosapentaenoic acid, into Daphnia resting eggs did not change significantly over time. Changes in trophic state strikingly influenced all trophic levels in Lake Constance. However, nutrient allocation into Daphnia resting eggs was mostly resilient to changes in lake trophic state.

Calanoid copepod zooplankton density is positively associated with water residence time across the continental United States.

Inherent differences between naturally-formed lakes and human-made reservoirs may play an important role in shaping zooplankton community structure. For example, because many reservoirs are created by impounding and managing lotic systems for specific human purposes, zooplankton communities may be affected by factors that are unique to reservoirs, such as shorter water residence times and a reservoir's management regime, compared to natural lakes. However, the environmental factors that structure zooplankton communities in natural lakes vs. reservoirs may vary at the continental scale and remain largely unknown. We analyzed data from the 2007 U.S. Environmental Protection Agency's National Lakes Assessment and the U.S. Army Corps of Engineers' National Inventory of Dams to compare large-bodied crustacean zooplankton communities (defined here as individuals retained by 0.243 mm mesh size) in natural lakes and reservoirs across the continental U.S. using multiple linear regressions and regression tree analyses. We found that large-bodied crustacean zooplankton density was overall higher in natural lakes compared to reservoirs when the effect of latitude was controlled. The difference between waterbody types was driven by calanoid copepods, which were also more likely to be dominant in the >0.243 mm zooplankton community in natural lakes than in reservoirs. Regression tree analyses revealed that water residence time was not a major driver of calanoid copepod density in natural lakes but was one of the most important drivers of calanoid copepod density in reservoirs, which had on average 0.5-year shorter water residence times than natural lakes. Reservoirs managed for purposes that resulted in shorter residence times (e.g., hydroelectric power) had lower zooplankton densities than reservoirs managed for purposes that resulted in longer residence times (e.g., irrigation). Consequently, our results indicate that water residence time may be an important characteristic driving differing large-bodied zooplankton dynamics between reservoirs and natural lakes.

A European Multi Lake Survey dataset of environmental variables, phytoplankton pigments and cyanotoxins.

Under ongoing climate change and increasing anthropogenic activity, which continuously challenge ecosystem resilience, an in-depth understanding of ecological processes is urgently needed. Lakes, as providers of numerous ecosystem services, face multiple stressors that threaten their functioning. Harmful cyanobacterial blooms are a persistent problem resulting from nutrient pollution and climate-change induced stressors, like poor transparency, increased water temperature and enhanced stratification. Consistency in data collection and analysis methods is necessary to achieve fully comparable datasets and for statistical validity, avoiding issues linked to disparate data sources. The European Multi Lake Survey (EMLS) in summer 2015 was an initiative among scientists from 27 countries to collect and analyse lake physical, chemical and biological variables in a fully standardized manner. This database includes in-situ lake variables along with nutrient, pigment and cyanotoxin data of 369 lakes in Europe, which were centrally analysed in dedicated laboratories. Publishing the EMLS methods and dataset might inspire similar initiatives to study across large geographic areas that will contribute to better understanding lake responses in a changing environment.

Trait selection and co-existence of phytoplankton in partially mixed systems: Trait based modelling and potential of an aggregated approach.

Trait selection and co-existence in phytoplankton communities in partially mixed water columns is investigated using trait based modelling. In the models employed, trait selection results from phytoplankton competition for two limiting resources, light and nutrients. The study employs spatially resolved models, in which the phytoplankton community is represented as a large number of trait-groups characterized by fixed trait combinations (trade-offs). Results from the trait-group resolving model (RM) are compared to results from an aggregated trait based model with adaptive traits (AM). Differences in specific production resulting from a trade-off between the half saturation constants of light and nutrients are sufficient to support evolutionary stable co-existence confirming that co-existence does not require differences in resource consumption. If abiotic conditions lead to the selection of a single trait group in RM, AM provides excellent approximations of the development of total biomass, average community trait and trait variance in the phytoplankton community. However, if selection leads to bimodal trait distributions, e.g. to co-existence of two trait groups (or species), functionally important properties of the phytoplankton community cannot be adequately represented by the aggregated information provided by AM. Because the increase in variance due to the development of bimodal trait distributions cannot be distinguished from an increase in variance due to an increase in trait diversity, the development of trait variance in AM models is not a reliable measure of trait diversity. Furthermore, AM may not provide reliable simulations of trophic interactions if the performance of the consumers depends on the traits of their resources. However, AM may support exploration of the consequences of environmental conditions and of the parameterization of species for co-existence within communities.

Dynamics and drivers of phytoplankton richness and composition along productivity gradient.

The shape of the productivity-richness relationship (PRR) for phytoplankton has been suggested to be unimodal, that is, the richness peaks at intermediate productivity levels. However, the mechanistic understanding for this pattern is still widely debated. In this study, we observed a unimodal PRR within 71 lakes along the Yangtze River encompassing an altitude range of 0-2700m, and an over 2200km distance from the upper reaches to the lower reaches. At low productivity, the competition for resources and regulatory processes jointly affected phytoplankton richness and composition, and their explanatory power depend on the gradient scale of driving factors. The variation of temperature attributing to altitudinal difference explained the majority of the variations of phytoplankton. If the altitude variation in temperature was eliminated, the explanatory power of temperature decreased from 31.7 to 7.6, and the independent effect of each resource and regulatory variable were limited and not decisive. At high productivity, the negative feedback of increased productivity (light limitation) affected the phytoplankton species richness and composition. The light-sensitive species disappeared, low-light-adapted species was retained and the phytoplankton composition gradually became similar with an increase in productivity. The findings contribute to an increased understanding of the mechanisms resulting in a hump-shaped PRR for phytoplankton.

Cross-ecosystem fluxes: Export of polyunsaturated fatty acids from aquatic to terrestrial ecosystems via emerging insects.

Cross-ecosystem fluxes can crucially influence the productivity of adjacent habitats. Emerging aquatic insects represent one important pathway through which freshwater-derived organic matter can enter terrestrial food webs. Aquatic insects may be of superior food quality for terrestrial consumers because they contain high concentrations of essential polyunsaturated fatty acids (PUFA). We quantified the export of PUFA via emerging insects from a midsize, mesotrophic lake. Insects were collected using emergence traps installed above different water depths and subjected to fatty acid analyses. Insect emergence from different depth zones and seasonal mean fatty acid concentrations in different insect groups were used to estimate PUFA fluxes. In total, 80.5mg PUFA m-2yr-1 were exported, of which 32.8mgm-2yr-1 were eicosapentaenoic acid (EPA), 7.8mgm-2yr-1 were arachidonic acid (ARA), and 2.6mgm-2yr-1 were docosahexaenoic acid (DHA). While Chironomidae contributed most to insect biomass and total PUFA export, Chaoborus flavicans contributed most to the export of EPA, ARA, and especially DHA. The export of total insect biomass from one square meter declined with depth and the timing at which 50% of total insect biomass emerged was correlated with the water depths over which the traps were installed, suggesting that insect-mediated PUFA fluxes are strongly affected by lake morphometry. Applying a conceptual model developed to assess insect deposition rates on land to our insect-mediated PUFA export data revealed an average total PUFA deposition rate of 150mgm-2yr-1 within 100m inland from the shore. We propose that PUFA export can be reliably estimated using taxon-specific information on emergent insect biomass and seasonal mean body PUFA concentrations of adult insects provided here. Our data indicate that insect-mediated PUFA fluxes from lakes are substantial, implying that freshwater-derived PUFA can crucially influence food web processes in adjacent terrestrial habitats.

Global impacts of the 1980s regime shift.

Despite evidence from a number of Earth systems that abrupt temporal changes known as regime shifts are important, their nature, scale and mechanisms remain poorly documented and understood. Applying principal component analysis, change-point analysis and a sequential t-test analysis of regime shifts to 72 time series, we confirm that the 1980s regime shift represented a major change in the Earth's biophysical systems from the upper atmosphere to the depths of the ocean and from the Arctic to the Antarctic, and occurred at slightly different times around the world. Using historical climate model simulations from the Coupled Model Intercomparison Project Phase 5 (CMIP5) and statistical modelling of historical temperatures, we then demonstrate that this event was triggered by rapid global warming from anthropogenic plus natural forcing, the latter associated with the recovery from the El Chichón volcanic eruption. The shift in temperature that occurred at this time is hypothesized as the main forcing for a cascade of abrupt environmental changes. Within the context of the last century or more, the 1980s event was unique in terms of its global scope and scale; our observed consequences imply that if unavoidable natural events such as major volcanic eruptions interact with anthropogenic warming unforeseen multiplier effects may occur.

Trophic mismatch requires seasonal heterogeneity of warming.

Climate warming has been shown to advance the phenology of species. Asynchronous changes in phenology between interacting species may disrupt feeding interactions (phenological mismatch), which could have tremendous consequences for ecosystem functioning. Long-term field observations have suggested asynchronous shifts in phenology with warming, whereas experimental studies have not been conclusive. Using proxy-based modeling of three trophic levels (algae, herbivores, and fish), we .show that asynchronous changes in phenology only occur if warming is seasonally heterogeneous, but not if warming is constant throughout the year. If warming is seasonally heterogeneous, the degree and even direction of asynchrony depends on the specific seasonality of the warming. Conclusions about phenological mismatches in food web interactions may therefore produce controversial results if the analyses do not distinguish between seasonally constant and seasonal specific warming. Furthermore, our results suggest that predicting asynchrony between interacting species requires reliable warming predictions that resolve sub-seasonal time scales.

Testing the stress gradient hypothesis in herbivore communities facilitation peaks at intermediate nutrient levels.

The role of positive interactions in structuring plant and animal communities is increasingly recognized, but the generality of current theoretical models has remained practically unexplored in animal communities. The stress gradient hypothesis predicts a linear increase in the intensity of facilitation as environmental conditions become increasingly stressful, whereas other theoretical models predict a maximum at intermediate environmental stress. We tested how competition and facilitation between herbivores change over a manipulated gradient of nutrient availability. We studied the effect of grazing by pond snails (Lymnaea stagnalis L.) as bulk grazers on aquatic caterpillars (Acentria ephemerella Denis and Schiffermüller) as small specialist grazers along an experimental gradient of environmental nutrient concentration. Higher nutrient levels increased overall total plant biomass but induced a shift toward dominance of filamentous algae at the expense of macrophytes. Facilitation of caterpillars by snail presence peaked at intermediate nutrient levels. Both caterpillar biomass and caterpillar grazing on macrophytes were highest at intermediate nutrient levels. Snails facilitated caterpillars possibly by removing filamentous algae and increasing access to the macrophyte resource, whereas they did not affect macrophyte biomass or C: nutrient ratios, a measure of food quality. We conclude that competition and facilitation in herbivore communities change along nutrient availability gradients that affect plant biomass and community composition. Understanding how interspecific interactions may change in strength and direction along environmental gradients is important to predict how the diversity and structure of communities may respond to the introduction or removal of herbivore species in ecosystems.

Compensatory dynamics and the stability of phytoplankton biomass during four decades of eutrophication and oligotrophication.

The link between compensatory dynamics and regime shifts is not well understood. We analyse a regime shift in phytoplankton in a large lake with respect to: (1) environmental forcing and (2) the type of dynamics (compensatory or synchronous) between phytoplankton groups. The regime shift in phytoplankton was related to gradual changes in nutrient levels, but unrelated to an almost concurrent shift in climatic conditions. The relationship between total phytoplankton biomass and phosphorus concentrations was sigmoid. Trajectories of phytoplankton biomass and community dynamics suggest that eutrophication effects can successfully be reversed when management efforts decrease nutrient loading to a level sufficiently low to overcome community resilience. The regime shift was associated with a loss of biomass compensation and compensatory dynamics among the phytoplankton groups. This suggests that the type of interactions is important for a better understanding of the existence and shape of nonlinear responses of phytoplankton biomass to environmental change.

Uniform temperature dependency in the phenology of a keystone herbivore in lakes of the Northern Hemisphere.

Spring phenologies are advancing in many ecosystems associated with climate warming causing unpredictable changes in ecosystem functioning. Here we establish a phenological model for Daphnia, an aquatic keystone herbivore based on decadal data on water temperatures and the timing of Daphnia population maxima from Lake Constance, a large European lake. We tested this model with long-term time-series data from two lakes (Müggelsee, Germany; Lake Washington, USA), and with observations from a diverse set of 49 lakes/sites distributed widely across the Northern Hemisphere (NH). The model successfully captured the observed temporal variation of Daphnia phenology in the two case study sites (r(2) = 0.25 and 0.39 for Müggelsee and Lake Washington, respectively) and large-scale spatial variation in the NH (R(2) = 0.57). These results suggest that Daphnia phenology follows a uniform temperature dependency in NH lakes. Our approach--based on temperature phenologies--has large potential to study and predict phenologies of animal and plant populations across large latitudinal gradients in other ecosystems.

Role of phytoplankton cell size on the competition for nutrients and light in incompletely mixed systems.

We investigate the effects of algal cell size on the competition for nutrients and light in an incompletely mixed water column, employing a spatially explicit variable internal stores approach and previously published allometric scaling relationships for modeling phytoplankton growth. We analyze the interplay between the size-dependent vertical assimilation and uptake profiles and the role of environmental settings such as mixing intensity, nutrient loading and background turbidity for the outcome of competition. Our results suggest that a potentially beneficial factor for resource competition in spatially heterogeneous systems is a low ratio of subsistence nutrient quota to the maximal quota, q(min)/q(max), which is a decreasing function of cell size according to allometric relationships. Environmental parameters such as mixing intensity and nutrient availability are shown to modulate the relevance of the q(min)/q(max) ratio for the competitive outcome and thereby have non-monotonic impacts on the algal size selection. The outcome of competition further depends on the temporal and spatial variability of mixing. In particular, the presence of a metalimnion with low diffusivity and periodic perturbation of the depth of the metalimnion strongly influences the relative success of differently sized algae. This suggests that the anticipated reduction in wind induced mixing events due to climate warming will have context-dependent consequences for algal size selection.