Regional zooplankton dispersal provides spatial insurance for ecosystem function

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.     

Spatial insurance in multi‐trophic metacommunities

Metacommunity theory suggests that dispersal is a key driver of diversity and ecosystem functioning in changing environments. The capacity of dispersal to mitigate effects of environmental change might vary among trophic groups, potentially resulting in changes in trophic interactions and food web structure. In a mesocosm experiment, we compared the compositional response of bacteria, phyto‐ and zooplankton to a factorial manipulation of acidification and dispersal. We found that the buffering capacity of dispersal varied among trophic groups: dispersal alleviated the negative effect of acidification on phytoplankton diversity mid‐experiment, but had no effect on the diversity of zooplankton and bacteria. Likewise, trophic groups differed in whether dispersal facilitated compositional change. Dispersal accelerated changes in phytoplankton composition under acidification, possibly mediated by changes in trophic interactions, but had no effect on the composition of zooplankton and bacteria. Overall, our results suggest that the potential for spatial insurance can vary among trophic groups.     

Ecosystem Functions across Trophic Levels Are Linked to Functional and Phylogenetic Diversity

In experimental systems, it has been shown that biodiversity indices based on traits or phylogeny can outperform species richness as predictors of plant ecosystem function. However, it is unclear whether this pattern extends to the function of food webs in natural ecosystems. Here we tested whether zooplankton functional and phylogenetic diversity explains the functioning of 23 natural pond communities. We used two measures of ecosystem function: (1) zooplankton community biomass and (2) phytoplankton abundance (Chl a). We tested for diversity-ecosystem function relationships within and across trophic levels. We found a strong correlation between zooplankton diversity and ecosystem function, whereas local environmental conditions were less important. Further, the positive diversity-ecosystem function relationships were more pronounced for measures of functional and phylogenetic diversity than for species richness. Zooplankton and phytoplankton biomass were best predicted by different indices, suggesting that the two functions are dependent upon different aspects of diversity. Zooplankton community biomass was best predicted by zooplankton trait-based functional richness, while phytoplankton abundance was best predicted by zooplankton phylogenetic diversity. Our results suggest that the positive relationship between diversity and ecosystem function can extend across trophic levels in natural environments, and that greater insight into variation in ecosystem function can be gained by combining functional and phylogenetic diversity measures.     

Evenness effects mask richness effects on ecosystem functioning at macro‐scales in lakes

Biodiversity–ecosystem functioning (BEF) theory has largely focused on species richness, although studies have demonstrated that evenness may have stronger effects. While theory and numerous small‐scale studies support positive BEF relationships, regional studies have documented negative effects of evenness on ecosystem functioning. We analysed a lake dataset spanning the continental US to evaluate whether strong evenness effects are common at broad spatial scales and if BEF relationships are similar across diverse regions and trophic levels. At the continental scale, phytoplankton evenness explained more variance in phytoplankton and zooplankton resource use efficiency (RUE; ratio of biomass to resources) than richness. For individual regions, slopes of phytoplankton evenness–RUE relationships were consistently negative and positive for phytoplankton and zooplankton RUE, respectively, and most slopes did not significantly differ among regions. Findings suggest that negative evenness effects may be more common than previously documented and are not exceptions restricted to highly disturbed systems.     

Congruence,but no cascade—Pelagic biodiversity across three trophic levels in Nordic lakes

Covariation in species richness and community structure across taxonomical groups (cross‐taxon congruence) has practical consequences for the identification of biodiversity surrogates and proxies, as well as theoretical ramifications for understanding the mechanisms maintaining and sustaining biodiversity. We found there to exist a high cross‐taxon congruence between phytoplankton, zooplankton, and fish in 73 large Scandinavian lakes across a 750 km longitudinal transect. The fraction of the total diversity variation explained by local environment alone was small for all trophic levels while a substantial fraction could be explained by spatial gradient variables. Almost half of the explained variation could not be resolved between local and spatial factors, possibly due to confounding issues between longitude and landscape productivity. There is strong consensus that the longitudinal gradient found in the regional fish community results from postglacial dispersal limitations, while there is much less evidence for the species richness and community structure gradients at lower trophic levels being directly affected by dispersal limitation over the same time scale. We found strong support for bidirectional interactions between fish and zooplankton species richness, while corresponding interactions between phytoplankton and zooplankton richness were much weaker. Both the weakening of the linkage at lower trophic levels and the bidirectional nature of the interaction indicates that the underlying mechanism must be qualitatively different from a trophic cascade.     

Loss of Rare Fish Species from Tropical Floodplain Food Webs Affects Community Structure and Ecosystem Multifunctionality in a Mesocosm Experiment

Experiments with realistic scenarios of species loss from multitrophic ecosystems may improve insight into how biodiversity affects ecosystem functioning. Using 1000 L mesocoms, we examined effects of nonrandom species loss on community structure and ecosystem functioning of experimental food webs based on multitrophic tropical floodplain lagoon ecosystems. Realistic biodiversity scenarios were developed based on long-term field surveys, and experimental assemblages replicated sequential loss of rare species which occurred across all trophic levels of these complex food webs. Response variables represented multiple components of ecosystem functioning, including nutrient cycling, primary and secondary production, organic matter accumulation and whole ecosystem metabolism. Species richness significantly affected ecosystem function, even after statistically controlling for potentially confounding factors such as total biomass and direct trophic interactions. Overall, loss of rare species was generally associated with lower nutrient concentrations, phytoplankton and zooplankton densities, and whole ecosystem metabolism when compared with more diverse assemblages. This pattern was also observed for overall ecosystem multifunctionality, a combined metric representing the ability of an ecosystem to simultaneously maintain multiple functions. One key exception was attributed to time-dependent effects of intraguild predation, which initially increased values for most ecosystem response variables, but resulted in decreases over time likely due to reduced nutrient remineralization by surviving predators. At the same time, loss of species did not result in strong trophic cascades, possibly a result of compensation and complexity of these multitrophic ecosystems along with a dominance of bottom-up effects. Our results indicate that although rare species may comprise minor components of communities, their loss can have profound ecosystem consequences across multiple trophic levels due to a combination of direct and indirect effects in diverse multitrophic ecosystems.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

Predators, resources, and trophic chains in the regulation of plankton population and biomass in oligothrophic lakes

The relative strength of "top-down" versus "bottom-up" control of plankton community structure and biomass in two small oligotrophic lakes (with and without fish), located near the Polar circle (Russia), has been investigated for two years, 1996 and 1997. The comparative analyses of zooplankton biomass and species abundance showed strong negative effect of fish, stickeback (Pungitius pungitius L.), on the zooplankton community species, size structure and biomass of particular prey species but no effect on the biomass of the whole trophic level. An intensive predation in Verkhneye lake has lead to: 1) sixfold decline in biomass of large cladoceran Holopedium gibberum comparing to the lake lacking predator, 2) shift in the size mode in zooplankton community and the replacement of the typical large grazers by small species--Bosmina longirostris and rotifers. Their abundance and biomass even increased, demonstrating the stimulating effect of fish on the "inefficient" and unprofitable prey organisms. The analysis of contributions of different factors into the cladoceran's birth rate changes was applied to demonstrate the relative impact of predators and resources on zooplankton abundance. An occasional introduction of the stickleback to Vodoprovodnoye lake (the reference lake in 1996) in summer 1997 lead to drastic canges in this ecosystem: devastating decrease of zooplankton biomass and complete elimination of five previously dominant grazer species. The abundance of edible phytoplankton was slightly higher in the lake with fish in 1996 and considerably higher in the lake where fish has appeared in 1997 showing the prevailing "top-down" control of phytoplankton in oligotrophic ecosystem. The reasons of trophic cascade appearance in oligotrophic lakes are also discussed.     

Five Years of Experimental Warming Increases the Biodiversity and Productivity of Phytoplankton

Phytoplankton are key components of aquatic ecosystems, fixing CO₂ from the atmosphere through photosynthesis and supporting secondary production, yet relatively little is known about how future global warming might alter their biodiversity and associated ecosystem functioning. Here, we explore how the structure, function, and biodiversity of a planktonic metacommunity was altered after five years of experimental warming. Our outdoor mesocosm experiment was open to natural dispersal from the regional species pool, allowing us to explore the effects of experimental warming in the context of metacommunity dynamics. Warming of 4°C led to a 67% increase in the species richness of the phytoplankton, more evenly-distributed abundance, and higher rates of gross primary productivity. Warming elevated productivity indirectly, by increasing the biodiversity and biomass of the local phytoplankton communities. Warming also systematically shifted the taxonomic and functional trait composition of the phytoplankton, favoring large, colonial, inedible phytoplankton taxa, suggesting stronger top-down control, mediated by zooplankton grazing played an important role. Overall, our findings suggest that temperature can modulate species coexistence, and through such mechanisms, global warming could, in some cases, increase the species richness and productivity of phytoplankton communities.     

Ecosystem consequences of the regional species pool

In this study we analyse with an experiment how the spatial area from which the species pool is sampled, i.e. local vs regional, can have functional consequences for ecosystem processes and structure. Particularly, we analysed how major stocks and processes respond to a change in basal productivity and how this depends on the source of the species pool. The experimental microcosms used in this experiment were developed by inoculating nutrient solution with samples of rockpool sediment containing resting stages. Here we show that communities developed from sediment samples of single origin (local communities) exhibit important differences in ecosystem structure and their response to a productivity gradient compared to systems developed from mixed samples of multiple and spatially dispersed origin (regional communities). The difference between local and regional communities was most pronounced for zooplankton which had much lower abundances in the local communities compared to the regional communities resulting in cascading effects on phytoplankton. We demonstrate that the spatial extend from which the species pool is sampled can affect ecosystem structure and functioning comparable in magnitude to changes in the basal productivity of the system. We discuss with a conceptual model how the regional species pool is important for sustaining diversity in functional groups, and how higher trophic levels are more sensitive to changes in the regional species pool.     

A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014–2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.     

Warming induces synchrony and destabilizes experimental pond zooplankton metacommunities

The spatial insurance hypothesis predicts that intermediate rates of dispersal between patches in a metacommunity allow species to track favourable conditions, preserving diversity and stabilizing biomass at local and regional scales. However, theory is unclear as to whether dispersal will provide spatial insurance when environmental conditions are changing directionally. In particular, increased temperatures as a result of climate change are expected to cause synchronous growth or decline across species and communities, and this has the potential to erode the stabilizing compensatory dynamics facilitated by dispersal. Here we report on an experimental test of how dispersal affects the diversity and stability of metacommunities under warming using replicate two‐patch pond zooplankton metacommunities. Initial differences in local community composition and abiotic conditions were established by seeding each patch in the metacommunities with plankton and sediment from one of two natural ponds that differed in water chemistry and species composition. We exposed metacommunities to a 2°C increase in average ambient temperature, crossed with three rates of dispersal (none, intermediate, high). In ambient conditions, intermediate dispersal rates preserved diversity and stabilized metacommunities by promoting spatially asynchronous fluctuations in biomass, especially between local populations of the dominant genus, Ceriodaphnia. However, warming synchronized their populations so that these effects of dispersal were lost. Furthermore, because the stabilizing effect of dispersal was primarily due to asynchronous fluctuations between populations of a single genus, metacommunity biomass was stabilized, but dispersal did not stabilize local community biomass. Our results show that dispersal can preserve diversity and provide stability to metacommunities, but also show that this benefit can be eroded when warming is directional and synchronous across patches of a metacommunity, as is expected with climate warming.     

Effect of (a)synchronous light fluctuation on diversity,functional and structural stability of a marine phytoplankton metacommunity

Disentangling the mechanisms that maintain the stability of communities and ecosystem properties has become a major research focus in ecology in the face of anthropogenic environmental change. Dispersal plays a pivotal role in maintaining diversity in spatially subdivided communities, but only a few experiments have simultaneously investigated how dispersal and environmental fluctuation affect community dynamics and ecosystem stability. We performed an experimental study using marine phytoplankton species as model organisms to test these mechanisms in a metacommunity context. We established three levels of dispersal and exposed the phytoplankton to fluctuating light levels, where fluctuations were either spatially asynchronous or synchronous across patches of the metacommunity. Dispersal had no effect on diversity and ecosystem function (biomass), while light fluctuations affected both evenness and community biomass. The temporal variability of community biomass was reduced by fluctuating light and temporal beta diversity was influenced interactively by dispersal and fluctuation, whereas spatial variability in community biomass and beta diversity were barely affected by treatments. Along the establishing gradient of species richness and dominance, community biomass increased but temporal variability of biomass decreased, thus highest stability was associated with species-rich but highly uneven communities and less influenced by compensatory dynamics. In conclusion, both specific traits (dominance) and diversity (richness) affected the stability of metacommunities under fluctuating conditions.     

Influence of Pulsed Inflows and Nutrient Loading on Zooplankton and Phytoplankton Community Structure and Biomass in Microcosm Experiments Using Estuarine Assemblages

Productivity and community structure of phytoplankton and zooplankton are influenced by hydrologic disturbances in many ways. In a recent modeling study it was suggested that pulsed inflows might enhance zooplankton performance, curb accumulation of phytoplankton accumulated biomass, and promote phytoplankton species diversity. We tested these predictions by performing microcosm experiments on natural plankton assemblages from the Nueces Delta, TX, USA. On three occasions (March, June, and September 2001), experiments of semi-continuous and flow-through design were conducted using natural plankton assemblages. We investigated the effect of two different inflow and nutrient loading regimes on zooplankton biomass, and phytoplankton biomass and diversity, i.e., continuous and pulsed inflows of 3 day frequency. Despite differences in initial community structure on these three occasions, as well as the very different communities that arose between experimental designs, our findings showed that pulsed inflows altered plankton dynamics. In all cases, pulsed inflows resulted in greater zooplankton biomass. In most cases, pulsed inflows resulted in lower phytoplankton biomass and higher diversity. We speculate that greater phytoplankton diversity in the pulsed flow treatments favored selectively feeding zooplankton, whose better performance prevented higher accumulation of phytoplankton biomass.     

Intraspecific variation in a predator drives cascading variation in primary producer community composition

Predation has important cascading impacts on primary producer biomass and community composition in many ecosystems. While most studies have focused on the consequences of interspecific or density differences in predators, it is recognized that phenotypic variation within species can have strong and cascading community and ecosystem consequences at lower trophic levels. In coastal New England lakes, both the presence and life history form of the zooplanktivorous fish alewife, Alosa pseudoharengus, have strong influence on the biomass, size structure and community composition of crustacean zooplankton communities. Here we test the hypothesis that alewife presence and life history will have cascading impacts on phytoplankton biomass and community composition in a mesocosm experiment that previously reported strong biomass and compositional differences of crustacean zooplankton communities among alewife treatments. We show that alewife life history led to small but statistically significant differences in phytoplankton community composition among treatments. This compositional difference was driven primarily by an increase in the density of two edible phytoplankton genera associated with lower zooplankton biomass in the anadromous alewife treatment. Our results show that intraspecific variation in a predator can have cascading effects on primary producer communities. However we did not observe significant differences in total algal biomass.     

Cyanobacteria dominance influences resource use efficiency and community turnover in phytoplankton and zooplankton communities

Freshwater biodiversity loss potentially disrupts ecosystem services related to water quality and may negatively impact ecosystem functioning and temporal community turnover. We analysed a data set containing phytoplankton and zooplankton community data from 131 lakes through 9 years in an agricultural region to test predictions that plankton communities with low biodiversity are less efficient in their use of limiting resources and display greater community turnover (measured as community dissimilarity). Phytoplankton resource use efficiency (RUE = biomass per unit resource) was negatively related to phytoplankton evenness (measured as Pielou's evenness), whereas zooplankton RUE was positively related to phytoplankton evenness. Phytoplankton and zooplankton RUE were high and low, respectively, when Cyanobacteria, especially Microcystis sp., dominated. Phytoplankton communities displayed slower community turnover rates when dominated by few genera. Our findings, which counter findings of many terrestrial studies, suggest that Cyanobacteria dominance may play important roles in ecosystem functioning and community turnover in nutrient‐enriched lakes.     

Biogeographical patterns of freshwater micro‐ and macroorganisms: a comparison between phytoplankton,zooplankton and fish in the eastern Mediterranean

Aim We investigated the biogeographical patterns of phytoplankton, zooplankton and fish in freshwater ecosystems. We tested whether spatial distance or environmental heterogeneity act as potential factors controlling community composition. Location Northern and central Greece, eastern Mediterranean. Method Data on 310 phytoplankton, 72 zooplankton and 37 fish species were collected from seven freshwater systems. Species occurrence data were used to generate similarity matrices describing community composition. We performed Mantel tests to compare spatial patterns in community composition of phytoplankton, zooplankton and fish. Next, we examined the correlation between geographical distance and the degree of similarity in community composition. The analysis was repeated for different taxonomic, trophic and size‐based groups of the microorganisms studied. We assessed different environmental variables (topographic and limnological) as predictors of community composition. Results Phytoplankton community composition showed a strong positive correlation with environmental heterogeneity but was not correlated with the geographical distance between systems. Zooplankton community composition was unrelated to geographical distance and was only weakly correlated with environmental variables. In contrast, fish community similarity decayed significantly with distance. We found no relationship along all pairwise comparisons of the compositional matrices of the three groups. The pairwise comparisons of the different taxonomic, trophic and size‐based groups of the microorganism communities studied were in accordance with the results for the entire microorganism community. Main conclusions Our results support the proposition that the biogeography of microorganisms does not demonstrate a distance–decay pattern and further suggest that, in reality, the drivers of distribution depend on the specific community examined. In contrast, the biogeography of macroorganisms was affected by geographical distance. These differences reflect the dispersal abilities of the different organisms. The microorganisms exhibit passive dispersal through the air, with local environmental conditions structuring their community composition. On the other hand, for macroorganisms such as fish, the terrestrial environment could pose barriers to their dispersal; with fish structuring distinctive communities over greater distances. Overall, we suggest that the biogeography of freshwater phytoplankton and zooplankton reflects contemporary environmental conditions, while the biogeographical patterns for fish inhabiting the same systems are related to factors affecting their dispersal ability.     

Understanding the Spatio-Temporal Response of Coral Reef Fish Communities to Natural Disturbances: Insights from Beta-Diversity Decomposition

Understanding how communities respond to natural disturbances is fundamental to assess the mechanisms of ecosystem resistance and resilience. However, ecosystem responses to natural disturbances are rarely monitored both through space and time, while the factors promoting ecosystem stability act at various temporal and spatial scales. Hence, assessing both the spatial and temporal variations in species composition is important to comprehensively explore the effects of natural disturbances. Here, we suggest a framework to better scrutinize the mechanisms underlying community responses to disturbances through both time and space. Our analytical approach is based on beta diversity decomposition into two components, replacement and biomass difference. We illustrate this approach using a 9-year monitoring of coral reef fish communities off Moorea Island (French Polynesia), which encompassed two severe natural disturbances: a crown-of-thorns starfish outbreak and a hurricane. These disturbances triggered a fast logistic decline in coral cover, which suffered a 90% decrease on all reefs. However, we found that the coral reef fish composition remained largely stable through time and space whereas compensatory changes in biomass among species were responsible for most of the temporal fluctuations, as outlined by the overall high contribution of the replacement component to total beta diversity. This suggests that, despite the severity of the two disturbances, fish communities exhibited high resistance and the ability to reorganize their compositions to maintain the same level of total community biomass as before the disturbances. We further investigated the spatial congruence of this pattern and showed that temporal dynamics involved different species across sites; yet, herbivores controlling the proliferation of algae that compete with coral communities were consistently favored. These results suggest that compensatory changes in biomass among species and spatial heterogeneity in species responses can provide further insurance against natural disturbances in coral reef ecosystems by promoting high levels of key species (herbivores). They can also allow the ecosystem to recover more quickly.     

Food web de‐synchronization in England's largest lake: an assessment based on multiple phenological metrics

Phenological changes have been observed globally for marine, freshwater and terrestrial species, and are an important element of the global biological ‘fingerprint’ of climate change. Differences in rates of change could desynchronize seasonal species interactions within a food web, threatening ecosystem functioning. Quantification of this risk is hampered by the rarity of long‐term data for multiple interacting species from the same ecosystem and by the diversity of possible phenological metrics, which vary in their ecological relevance to food web interactions. We compare phenological change for phytoplankton (chlorophyll a), zooplankton (Daphnia) and fish (perch, Perca fluviatilis) in two basins of Windermere over 40 years and determine whether change has differed among trophic levels, while explicitly accounting for among‐metric differences in rates of change. Though rates of change differed markedly among the nine metrics used, seasonal events shifted earlier for all metrics and trophic levels: zooplankton advanced most, and fish least, rapidly. Evidence of altered synchrony was found in both lake basins, when combining information from all phenological metrics. However, comparisons based on single metrics did not consistently detect this signal. A multimetric approach showed that across trophic levels, earlier phenological events have been associated with increasing water temperature. However, for phytoplankton and zooplankton, phenological change was also associated with changes in resource availability. Lower silicate, and higher phosphorus, concentrations were associated with earlier phytoplankton growth, and earlier phytoplankton growth was associated with earlier zooplankton growth. The developing trophic mismatch detected between the dominant fish species in Windermere and important zooplankton food resources may ultimately affect fish survival and portend significant impacts upon ecosystem functioning. We advocate that future studies on phenological synchrony combine data from multiple phenological metrics, to increase confidence in assessments of change and likely ecological consequences.     

Productivity-diversity relationships in lake plankton communities

One of the most intriguing environmental gradients connected with variation in diversity is ecosystem productivity. The role of diversity in ecosystems is pivotal, because species richness can be both a cause and a consequence of primary production. However, the mechanisms behind the varying productivity-diversity relationships (PDR) remain poorly understood. Moreover, large-scale studies on PDR across taxa are urgently needed. Here, we examined the relationships between resource supply and phyto-, bacterio-, and zooplankton richness in 100 small boreal lakes. We studied the PDR locally within the drainage systems and regionally across the systems. Second, we studied the relationships between resource availability, species richness, biomass and resource ratio (N:P) in phytoplankton communities using Structural Equation Modeling (SEM) for testing the multivariate hypothesis of PDR. At the local scale, the PDR showed variable patterns ranging from positive linear and unimodal to negative linear relationships for all planktonic groups. At the regional scale, PDRs were significantly linear and positive for phyto- and zooplankton. Phytoplankton richness and the amount of chlorophyll a showed a positive linear relationship indicating that communities consisting of higher number of species were able to produce higher levels of biomass. According to the SEM, phytoplankton biomass was largely related to resource availability, yet there was a pathway via community richness. Finally, we found that species richness at all trophic levels was correlated with several environmental factors, and was also related to richness at the other trophic levels. This study showed that the PDRs in freshwaters show scale-dependency. We also documented that the PDR complies with the multivariate model showing that plant biomass is not mirroring merely the resource availability, but is also influenced by richness. This highlights the need for conserving diversity in order to maintain ecosystem processes in freshwaters.