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.     

Planktonic dispersal dampens temporal trophic cascades in pond metacommunities

Trophic cascades, in which changes in predation affect the biomass of lower trophic levels, vary substantially in strength and incidence. Most work to explain this variation has focused on local factors and has ignored larger regional effects. To study how metacommunity dynamics can alter trophic cascades, we constructed mesocosm metacommunities consisting of three pond communities with heterogeneous levels of fish predation and examined how planktonic dispersal rate (5–140% per week) affected biomass partitioning. Two of the three communities differed continually in the occurrence of fish and supported different but constant environments in a 'spatial trophic cascade,' while the third community supported temporally variable fish occurrence in a 'temporal trophic cascade.' We find that the presence, but the not the magnitude, of dispersal dampens temporal trophic cascades through an increase in grazer biomass. In contrast, dispersal has no effect on the strength of spatial cascades due to strong sorting pressures in the communities with constant presence or absence of fish as top predators.     

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.     

An experimental demonstration of trophic interactions affecting water quality of Rice Lake, Ontario (Canada)

Eight cylindrical enclosures (3 m diameter, 2.7 m long, V = 20m³) were installed in eutrophic Rice Lake (Ontario, Canada) in late spring of 1987. Fish (yearling yellow perch (Perca flavescens) and macrophytes (Potamogeton crispus) presence and absence were set at the beginning of the experiment to yield four combinations of duplicate treatments. The purpose of the experiment was to determine if the phytoplankton, zooplankton, macrophytes and fish species resident in the lake interact to influence water quality (major ions, phosphorus, algal densities and water clarity).The presence of fish was associated with: (1) decreased biomass of total zooplankton, (2) decreased number of species in the zooplankton, (3) decreased average size of several zooplankton taxa, (4) higher total phosphorus concentrations, (5) higher phytoplankton and chlorophyll a concentrations, (6) lower water clarity, (7) lower potassium levels during macrophyte die-back, (8) lower pH and higher conductivity in the presence of macrophytes. Biomass of large Daphnia species (but not total zooplankton) was highly correlated with the algal response (r ² = 0.995) and was associated with reduced biomass of several algal taxa including some large forms (Mougeotia, Oedogonium) and several colonial blue-green algae. However, no significant control of late summer growth of the bloom-forming blue-green alga Anabaena planctonica Brun. was achieved by the Daphnia presence-fish absence treatment. Release of phosphorus to the water column during the die-back of P. crispus was not an important phenomenon.     

Predator‐induced changes in dissolved organic carbon dynamics

The fate of dissolved organic carbon (DOC) is partly determined by its availability to microbial degradation. Organisms at upper trophic levels could influence the bioavailability of DOC via cascading effects on primary producers and bacteria. Here we experimentally tested whether the presence of fish in aquatic food webs can indirectly affect the composition of the DOC pool. We found that fish had strong positive effects on phytoplankton biomass that affected the dynamics of DOC composition. Specifically, fish increased protein‐like, algae‐derived DOC mid‐experiment, concurrent with the strongest fish‐induced increase in phytoplankton biomass. Fish also increased bacterial abundance, altered the community composition and diversity of bacteria, and temporarily increased DOC compounds with fluorescence properties indicative of microbially‐reprocessed organic matter. Overall, our experiment revealed that fish can positively influence the substrate (algae‐produced DOC) and the key players (bacteria) of the microbial carbon pump. Consequently, fish could contribute to carbon sequestration by stimulating both the production of bioavailable DOC and the microbial degradation of bioavailable to persistent DOC. We propose this as a novel mechanism whereby the loss of predators from global ecosystems could alter carbon cycling.     

On the taxonomy of the genus Megadiaptomus Kiefer,including the description of a new species (Copepoda,Calanoida) from India

The insecticide permethrin (a synthetic pyrethroid) was applied into enclosures (1 m diameter and 3.5 m deep) placed in a pond. The chemical was rapidly removed from the water to the sediments. Daphnia rosea and its predator, Chaoborus flavicans were seriously affected by this application and disappeared from the enclosure. Acanthodiaptomus pacificus increased as Chaoborus decreased. Two species of Cladocera which had not been observed before the treatment, established their populations after 10 days, when Chaoborus had not recovered. Whereas Tropocyclops pracinus declined when permethrin was applied at 10 µg 1^–1, the number of rotifer Keratella valga increased, suggesting a close relationship between these two species.Photosynthesis and phytoplankton were not significantly affected by permethrin, except for Ceratium hirundinella. The dominance of Dinobryon divergens continued in the treated enclosures, whereas other flagellate species, Scenedesmus and Nitzschia occurred during the second half of the experiments in the control enclosure and pond.     

Towards a multi‐trophic extension of metacommunity ecology

Metacommunity theory provides an understanding of how spatial processes determine the structure and function of communities at local and regional scales. Although metacommunity theory has considered trophic dynamics in the past, it has been performed idiosyncratically with a wide selection of possible dynamics. Trophic metacommunity theory needs a synthesis of a few influential axis to simplify future predictions and tests. We propose an extension of metacommunity ecology that addresses these shortcomings by incorporating variability among trophic levels in ‘spatial use properties’. We define ‘spatial use properties’ as a set of traits (dispersal, migration, foraging and spatial information processing) that set the spatial and temporal scales of organismal movement, and thus scales of interspecific interactions. Progress towards a synthetic predictive framework can be made by (1) documenting patterns of spatial use properties in natural food webs and (2) using theory and experiments to test how trophic structure in spatial use properties affects metacommunity dynamics.     

Size‐based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn,Iceland

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HSS revisited: multi‐channel processes mediate trophic control across a productivity gradient

Classical food web theory holds that energy channels are regulated by top‐down control with increasing productivity, arising from within‐channel processes. However, these hypotheses do not consider the existence of parallel energy channels linked by shared resource pools and which can fuel generalist predators, imposing trophic control arising from multi‐channel processes. Using 23 large marine food webs, we show that food web responses to increasing productivity are consistent with the apparent trophic cascade hypothesis (ATCH) – with rising productivity predators derive an increasing fraction of their diet from increasingly productive bottom‐up controlled detritus channels, thereby subsidising predator biomass, and in turn strengthening top‐down control in parallel grazing channels. These results testify to a fundamental role of detritus channels specifically and multi‐channel processes in general in mediating food web response to productivity and demonstrate that the ATCH provides an alternative explanation for classical predictions of top‐down control.     

δ15N of zooplankton species in subarctic lakes in northern Sweden: effects of diet and trophic fractionation

1. To assess the use of stable nitrogen isotopes (δ¹⁵N) for reconstructing trophic relationships in planktonic food webs, crustacean zooplankton species and pelagic dissolved and particulate matter were analysed in 14 subarctic lakes in northern Sweden. The lakes are situated along an altitudinal gradient and show a substantial variation in nutrient content and energy mobilization by bacterioplankton and phytoplankton. 2. The δ¹⁵N of dissolved and particulate matter was comparatively low, suggesting efficient N recycling and low losses of depleted N from the pelagic zone of these unproductive lakes. 3. Copepods had a systematically higher δ¹⁵N than cladocerans, with an average difference of 3.1–4.9‰ within lakes, implying different trophic positions of the two groups. Comparisons of nitrogen pools and energy fluxes suggest that the low cladoceran δ¹⁵N was a result of feeding on bacteria. 4. The difference in δ¹⁵N between copepods and cladocerans declined with decreasing bacterioplankton production among lakes, due either to increasing trophic isotope fractionation or decreasing relative importance of bacteria in the diet of cladocerans.     

Spatial heterogeneity of biomass turnover has contrasting effects on synchrony and stability in trophic metacommunities

Spatial heterogeneity is a fundamental feature of ecosystems, and ecologists have identified it as a factor promoting the stability of population dynamics. In particular, differences in interaction strengths and resource supply between patches generate an asymmetry of biomass turnover with a fast and a slow patch coupled by a mobile predator. Here, we demonstrate that asymmetry leads to opposite stability patterns in metacommunities receiving localized perturbations depending on the characteristics of the perturbed patch. Perturbing prey in the fast patch synchronizes the dynamics of prey biomass between the two patches and destabilizes predator dynamics by increasing the predator's temporal variability. Conversely, perturbing prey in the slow patch decreases the synchrony of the prey's dynamics and stabilizes predator dynamics. Our results have implications for conservation ecology and suggest reinforcing protection policies in fast patches to dampen the effects of perturbations and promote the stability of population dynamics at the regional scale.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Plankton metacommunities in floodplain wetlands under contrasting hydrological conditions

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Omnivory does not prevent trophic cascades in pelagic food webs

1. Strong trophic cascades have been well documented in pelagic food webs of temperate lakes. In contrast, the limited available evidence suggests that strong cascades are less typical in tropical lakes.
2. To measure the effects of omnivorous tilapia on planktonic communities and water transparency of a small man-made tropical lake, we performed a 5-week in situ enclosure experiment with five densities of fish randomly allocated to 20 enclosures. Zooplankton and Phytoplankton biomasses as well as water transparency were measured weekly.
3. Results show that omnivorous tilapia significantly decreased the abundance of large Cladocerans, increased the abundance of small algae (greatest axial linear dimension <50  μ m) and decreased water transparency as predicted by trophic cascade theory.
4. Therefore, omnivory was not a sufficient factor to prevent a trophic cascade in this pelagic community, although the cascade effect was weaker than reported from many north temperate, nutrient-rich lakes.     

Tri‐trophic interactions: bridging species,communities and ecosystems

A vast body of research demonstrates that many ecological and evolutionary processes can only be understood from a tri‐trophic viewpoint, that is, one that moves beyond the pairwise interactions of neighbouring trophic levels to consider the emergent features of interactions among multiple trophic levels. Despite its unifying potential, tri‐trophic research has been fragmented, following two distinct paths. One has focused on the population biology and evolutionary ecology of simple food chains of interacting species. The other has focused on bottom‐up and top‐down controls over the distribution of biomass across trophic levels and other ecosystem‐level variables. Here, we propose pathways to bridge these two long‐standing perspectives. We argue that an expanded theory of tri‐trophic interactions (TTIs) can unify our understanding of biological processes across scales and levels of organisation, ranging from species evolution and pairwise interactions to community structure and ecosystem function. To do so requires addressing how community structure and ecosystem function arise as emergent properties of component TTIs, and, in turn, how species traits and TTIs are shaped by the ecosystem processes and the abiotic environment in which they are embedded. We conclude that novel insights will come from applying tri‐trophic theory systematically across all levels of biological organisation.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

Trophic transfer and trophic modification of fatty acids in high Arctic lakes

1. A comparative study of fatty acid (FA) profiles in particulate matter (seston) and the key grazer Daphnia was performed in six high Arctic ponds (79°N, Svalbard). The ponds were all small and shallow, but followed a strong gradient with respect to nutrient content and optical properties. 2. A distinct locality‐specific pattern was detected by principal component analysis of FA profiles, where samples from each locality clustered both with regard to seston and Daphnia. Linear discriminant analysis using nine sestonic fatty acids as discriminant functions gave on average a correct prediction of the Daphnia lake membership in 47% of cases, with very high predictability in some lakes but poor predictability in others. 3. No significant correlation was detected between FA and nutrient concentration or levels of dissolved organic carbon. The major determinant of FA profiles as judged from a redundancy analysis was the taxonomic composition of phytoplankton communities, notably the biomass of Chlorophyceae. 4. The FA profiles of Daphnia were for some FAs strongly enriched relative to the seston, while diluted for others. Among the polyunsaturated fatty acids (PUFAs), a pronounced magnification of eicosapentaenoic acid (EPA, 20 : 5 n‐3), and to some extent 18 : 3 n‐3 and 20 : 4 n‐6 was found, while docosahexaenoic acid (DHA, 22 : 6 n‐3) contributed in general less to FAs in Daphnia than in seston and was hardly detectable in Daphnia from most localities. 5. The overall content of PUFAs in Daphnia was consistently high, close to 40% of total FA in all investigated localities, despite major differences in seston PUFA content. Daphnia thus acts as a regulator with regard to overall PUFAs, reflecting its physiological constraints and relatively fixed demands for PUFAs in general. The distinct locality‐specific profiles in Daphnia strongly suggest a kind of FA‐fingerprint, but our data do not allow strict statements on the use of specific FAs as trophic markers.