Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Ecological Memory of Historical Contamination Influences the Response of Phytoplankton Communities

Ecological memory (EM) recognizes the importance of previous stress encounters in promoting community tolerance and thereby enhances ecosystem stability, provided that gained tolerances are preserved during non-stress periods. Drawing from this concept, we hypothesized that the recruitment of tolerant species can be facilitated by imposing an initial sorting process (conditioning) during the early stages of community assembly, which should result in higher production (biomass development and photosynthetic efficiency) and stable community composition. To test this, phytoplankton resting stages were germinated from lake sediments originating from two catchments that differed in contamination history: one impacted by long-term herbicides and pesticides exposures (historically contaminated lake) from an agricultural catchment compared to a low-impacted one (near-pristine lake) from a forested catchment. Conditioning was achieved by adding an herbicide (Isoproturon, which was commonly used in the catchment of the historically contaminated lake) during germination. Afterward, the communities obtained from germination were exposed to an increasing gradient of Isoproturon. As hypothesized, upon conditioning, the phytoplankton assemblages from the historically contaminated lake were able to rapidly restore photosynthetic efficiency (p?>?0.01) and became structurally (community composition) more resistant to Isoproturon. The communities of the near-pristine lake did not yield these positive effects regardless of conditioning, supporting that EM was a unique attribute of the historically stressed ecosystem. Moreover, assemblages that displayed higher structural resistance concurrently yielded lower biomass, indicating that benefits of EM in increasing structural stability may trade-off with production. Our results clearly indicate that EM can foster ecosystem stability to a recurring stressor.

     

Dispersal-mediated trophic interactions can generate apparent patterns of dispersal limitation in aquatic metacommunities

Dispersal is a major organising force in metacommunities, which may facilitate compositional responses of local communities to environmental change and affect ecosystem function. Organism groups differ widely in their dispersal abilities and their communities are therefore expected to have different adaptive abilities. In mesocosms, we studied the simultaneous compositional response of three plankton communities (zoo-, phyto- and bacterioplankton) to a primary productivity gradient and evaluated how this response was mediated by dispersal intensity. Dispersal enhanced responses in all three planktonic groups, which also affected ecosystem functioning. Yet, variation partitioning analyses indicated that responses in phytoplankton and bacterial communities were not only controlled by dispersal directly but also indirectly through complex trophic interactions. Our results indicate that metacommunity patterns emerging from dispersal can cascade through the food web and generate patterns of apparent dispersal limitation in organisms at other trophic levels.     

Community composition has greater impact on the functioning of marine phytoplankton communities than ocean acidification

Ecosystem functioning is simultaneously affected by changes in community composition and environmental change such as increasing atmospheric carbon dioxide (CO₂) and subsequent ocean acidification. However, it largely remains uncertain how the effects of these factors compare to each other. Addressing this question, we experimentally tested the hypothesis that initial community composition and elevated CO₂ are equally important to the regulation of phytoplankton biomass. We full‐factorially exposed three compositionally different marine phytoplankton communities to two different CO₂ levels and examined the effects and relative importance (ω²) of the two factors and their interaction on phytoplankton biomass at bloom peak. The results showed that initial community composition had a significantly greater impact than elevated CO₂ on phytoplankton biomass, which varied largely among communities. We suggest that the different initial ratios between cyanobacteria, diatoms, and dinoflagellates might be the key for the varying competitive and thus functional outcome among communities. Furthermore, the results showed that depending on initial community composition elevated CO₂ selected for larger sized diatoms, which led to increased total phytoplankton biomass. This study highlights the relevance of initial community composition, which strongly drives the functional outcome, when assessing impacts of climate change on ecosystem functioning. In particular, the increase in phytoplankton biomass driven by the gain of larger sized diatoms in response to elevated CO₂ potentially has strong implications for nutrient cycling and carbon export in future oceans.     

Grazer diversity affects resistance to multiple stressors in an experimental seagrass ecosystem

When multiple stressors act simultaneously, their effects on ecosystems become more difficult to predict. In the face of multiple stressors, diverse ecosystems may be more stable if species respond differently to stressors or if functionally similar species can compensate for stressor effects on focal species. Many habitats around the globe are threatened by multiple stressors, including highly productive seagrass habitats. For example, in Chesapeake Bay, USA, regional climate change predictions suggest that elevated temperature and freshwater inputs are likely to be increasingly important stressors. Using seagrass mesocosms as a model system, we tested whether species richness of crustacean grazers buffers ecosystem properties against the impacts of elevated temperature and freshwater pulse stressors in a fully factorial experiment. Grazer species responded to pulsed salinity changes differently; abundance of Elasmopus levis responded negatively to freshwater pulses, whereas abundance of Gammarus mucronatus and Erichsonella attenuata responded positively or neutrally. Consistent with the hypothesis that biodiversity provides resistance stability, biomass of epiphytic algae that form the base of the food web was less affected by stressors in species‐rich grazer treatments than in single‐species grazer treatments. Stochastic (among‐replicate) variation of sessile invertebrate biomass within treatments was also reduced in more diverse grazer treatments. Therefore, grazer species richness tended to increase the resistance stability of both major components of the seagrass fouling community, algae and invertebrates, in the face of environmental stressors. Finally, in our model system, multi‐stressor impacts suggested a pattern of antagonism contrary to previous assumptions of synergistic stressor effects. Overall, our results confirm that invertebrate grazer species are functionally diverse in their response to environmental stressors, but are largely functionally redundant in their grazing effects leading to greater resistance stability of certain ecosystem properties in diverse grazer assemblages even when influenced by multiple environmental stressors.     

Consistent Richness-Biomass Relationship across Environmental Gradients in a Marine Macroalgal-Dominated Subtidal Community on the Western Antarctic Peninsula

Biodiversity loss has spurred the biodiversity-ecosystem functioning research over a range of ecosystems. In Antarctica, however, the relationship of taxonomic and functional diversity with ecosystem properties (e.g., community biomass) has received less attention, despite the presence of sharp and dynamic environmental stress gradients that might modulate these properties. Here, we investigated whether the richness-biomass relationship in macrobenthic subtidal communities is still apparent after accounting for environmental stress gradients in Fildes Bay, King George Island, Antarctica. Measurements of biomass of mobile and sessile macrobenthic taxa were conducted in the austral summer 2013/4 across two environmental stress gradients: distance from nearest glaciers and subtidal depth (from 5 to 30 m). In general, community biomass increased with distance from glaciers and water depth. However, generalised additive models showed that distance from glaciers and depth accounted for negligible proportions of variation in the number of functional groups (i.e., functional richness) and community biomass when compared to taxonomic richness. Functional richness and community biomass were positive and saturating functions of taxonomic richness. Large endemic, canopy-forming brown algae of the order Desmarestiales dominated the community biomass across both gradients. Accordingly, differences in the composition of taxa accounted for a significant and large proportion (51%) of variation in community biomass in comparison with functional richness (10%). Our results suggest that the environmental factors here analysed may be less important than biodiversity in shaping mesoscale (several km) biomass patterns in this Antarctic system. We suggest that further manipulative, hypothesis-driven research should address the role of biodiversity and species’ functional traits in the responses of Antarctic subtidal communities to environmental variation.     

Functional richness outperforms taxonomic richness in predicting ecosystem functioning in natural phytoplankton communities

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Trophic Dependence of Ecosystem Resistance and Species Compensation in Experimentally Acidified Lake 302S (Canada)

Ecosystem resistance to the impacts of diverse human insults depends on the replacement of sensitive species by ones more tolerant of the stressor. Here we present evidence from a whole-lake acidification experiment (Lake 302S, Experimental Lakes Area, Canada) that resistance and species compensation decline with increasing trophic level. Diverse and fast-growing algal and rotifer assemblages with high dispersal potentials showed significant compensatory species dynamics, resulting in the maintenance of total biomass despite 30%–80% declines in species richness. Canonical correspondence analysis showed that significant compensatory algal and rotifer dynamics were best explained by differential species tolerances of acidified chemical conditions coupled with release from resource limitation and predation. However, less diverse cladoceran, copepod, and fish assemblages showed significant declines in total biomass and weak species compensation with loss of species during acidification. In comparison, algal and zooplankton species dynamics remained relatively synchronized in a nearby unperturbed reference lake (Lake 239) during the experiment. As a result, Lake 302S showed limited ecosystem resistance to anthropogenic acidification. Therefore, we hypothesize that lost species will increase the susceptibility of acidified lakes to the adverse impacts of other environmental stressors (for example, climate warming, stratospheric ozone depletion, invasive species). Consequently, the ecosystem stability of boreal lakes is expected to decline as global change proceeds. Received 2 January 2001; accepted 12 July 2002.     

Testing a ‘genes‐to‐ecosystems’ approach to understanding aquatic–terrestrial linkages

A ‘genes‐to‐ecosystems’ approach has been proposed as a novel avenue for integrating the consequences of intraspecific genetic variation with the underlying genetic architecture of a species to shed light on the relationships among hierarchies of ecological organization (genes → individuals → communities → ecosystems). However, attempts to identify genes with major effect on the structure of communities and/or ecosystem processes have been limited and a comprehensive test of this approach has yet to emerge. Here, we present an interdisciplinary field study that integrated a common garden containing different genotypes of a dominant, riparian tree, Populus trichocarpa, and aquatic mesocosms to determine how intraspecific variation in leaf litter alters both terrestrial and aquatic communities and ecosystem functioning. Moreover, we incorporate data from extensive trait screening and genome‐wide association studies estimating the heritability and genes associated with litter characteristics. We found that tree genotypes varied considerably in the quality and production of leaf litter, which contributed to variation in phytoplankton abundances, as well as nutrient dynamics and light availability in aquatic mesocosms. These ‘after‐life’ effects of litter from different genotypes were comparable to the responses of terrestrial communities associated with the living foliage. We found that multiple litter traits corresponding with aquatic community and ecosystem responses differed in their heritability. Moreover, the underlying genetic architecture of these traits was complex, and many genes contributed only a small proportion to phenotypic variation. Our results provide further evidence that genetic variation is a key component of aquatic–terrestrial linkages, but challenge the ability to predict community or ecosystem responses based on the actions of one or a few genes.     

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.     

Stressor‐induced biodiversity gradients: revisiting biodiversity–ecosystem functioning relationships

Biodiversity–ecosystem functioning experiments typically inspect functioning in randomly composed communities, representing broad gradients of taxonomic richness. We tested if the resulting evenness gradients and evenness–functioning relationships reflect those found in communities facing evenness loss caused by anthropogenic stressors. To this end, we exposed marine benthic diatom communities to a series of treatments with the herbicide atrazine, and analysed the relationship between the resulting gradients of evenness and ecosystem functioning (primary production, energy content and sediment stabilization). Atrazine exposure resulted in narrower evenness gradients and steeper evenness–functioning relations than produced by the design of random community assembly. The disproportionately large decrease in functioning following atrazine treatment was related to selective atrazine effects on the species that contributed most to the ecosystem functions considered. Our findings demonstrate that the sensitivity to stress and the contribution to ecosystem functioning at the species level should be both considered to understand biodiversity and ecosystem functioning under anthropogenic stress. Synthesis Biodiversity loss affects ecosystem functioning, yet biodiversity–ecosystem functioning relations have mainly been investigated using communities with random species loss. In nature however, species are lost according to their sensitivity to environmental stress. In the present study, biodiversity loss and biodiversity–ecosystem functioning relations in randomly composed diatom communities were compared to those induced by the pesticide atrazine. Stress exposure resulted in smaller biodiversity loss but steeper decrease in functioning than in randomly composed communities, due to selective atrazine effects on the best performing species. Therefore, species‐specific sensitivity and contribution to ecosystem functioning need to be considered to predict biodiversity and ecosystem functioning under anthropogenic stress.     

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.     

Biodiversity effects on ecosystem functioning change along environmental stress gradients

Positive relationship between biodiversity and ecosystem functioning has been observed in many studies, but how this relationship is affected by environmental stress is largely unknown. To explore this influence, we measured the biomass of microalgae grown in microcosms along two stress gradients, heat and salinity, and compared our results with 13 published case studies that measured biodiversity–ecosystem functioning relationships under varying environmental conditions. We found that positive effects of biodiversity on ecosystem functioning decreased with increasing stress intensity in absolute terms. However, in relative terms, increasing stress had a stronger negative effect on low‐diversity communities. This shows that more diverse biotic communities are functionally less susceptible to environmental stress, emphasises the need to maintain high levels of biodiversity as an insurance against impacts of changing environmental conditions and sets the stage for exploring the mechanisms underlying biodiversity effects in stressed ecosystems.     

Response of cyanobacteria and phytoplankton abundance to warming,extreme rainfall events and nutrient enrichment

Cyanobacterial blooms are an increasing threat to water quality and global water security caused by the nutrient enrichment of freshwaters. There is also a broad consensus that blooms are increasing with global warming, but the impacts of other concomitant environmental changes, such as an increase in extreme rainfall events, may affect this response. One of the potential effects of high rainfall events on phytoplankton communities is greater loss of biomass through hydraulic flushing. Here we used a shallow lake mesocosm experiment to test the combined effects of: warming (ambient vs. +4°C increase), high rainfall (flushing) events (no events vs. seasonal events) and nutrient loading (eutrophic vs. hypertrophic) on total phytoplankton chlorophyll‐a and cyanobacterial abundance and composition. Our hypotheses were that: (a) total phytoplankton and cyanobacterial abundance would be higher in heated mesocosms; (b) the stimulatory effects of warming on cyanobacterial abundance would be enhanced in higher nutrient mesocosms, resulting in a synergistic interaction; (c) the recovery of biomass from flushing induced losses would be quicker in heated and nutrient‐enriched treatments, and during the growing season. The results supported the first and, in part, the third hypotheses: total phytoplankton and cyanobacterial abundance increased in heated mesocosms with an increase in common bloom‐forming taxa—Microcystis spp. and Dolichospermum spp. Recovery from flushing was slowest in the winter, but unaffected by warming or higher nutrient loading. Contrary to the second hypothesis, an antagonistic interaction between warming and nutrient enrichment was detected for both cyanobacteria and chlorophyll‐a demonstrating that ecological surprises can occur, dependent on the environmental context. While this study highlights the clear need to mitigate against global warming, oversimplification of global change effects on cyanobacteria should be avoided; stressor gradients and seasonal effects should be considered as important factors shaping the response.     

Spatial insurance against a heatwave differs between trophic levels in experimental aquatic communities

Climate change-related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better-adapted taxa. Yet, experimental evidence for such predictions from multi-trophic communities and pulse-type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat-induced increase in metabolic costs, resulting in weaker top-down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer-lasting changes in ecosystem functioning.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Regional zooplankton biodiversity provides limited buffering of pond ecosystems against climate change

1. Climate change and other human-driven environmental perturbations are causing reductions in biodiversity and impacting the functioning of ecosystems on a global scale. Metacommunity theory suggests that ecosystem connectivity may reduce the magnitude of these impacts if the regional species pool contains functionally redundant species that differ in their environmental tolerances. Dispersal may increase the resistance of local ecosystems to environmental stress by providing regional species with traits adapted to novel conditions. 2. We tested this theory by subjecting freshwater zooplankton communities in mesocosms that were either connected to or isolated from the larger regional species pool to a factorial manipulation of experimental warming and increased salinity. 3. Compensation by regional taxa depended on the source of stress. Warming tolerant regional taxa partially compensated for reductions in heat sensitive local taxa but similar compensation did not occur under increased salinity. 4. Dispersal-mediated species invasions dampened the effects of warming on summer net ecosystem productivity. However, this buffering effect did not occur in the fall or for periphyton growth, the only other ecosystem function affected by the stress treatments. 5. The results indicate that regional biodiversity can provide insurance in a dynamic environment but that the buffering capacity is limited to some ecosystem processes and sources of stress. Maintaining regional biodiversity and habitat connectivity may therefore provide some limited insurance for local ecosystems in changing environments, but is unable to impart resistance against all sources of environmental stress.     

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.     

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.     

Long‐term allelopathic control of phytoplankton by the submerged macrophyte Elodea nuttallii

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