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.     

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.     

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.     

Final thermal conditions override the effects of temperature history and dispersal in experimental communities

Predicting the effect of climate change on biodiversity is a multifactorial problem that is complicated by potentially interactive effects with habitat properties and altered species interactions. In a microcosm experiment with communities of microalgae, we analysed whether the effect of rising temperature on diversity depended on the initial or the final temperature of the habitat, on the rate of change, on dispersal and on landscape heterogeneity. We also tested whether the response of species to temperature measured in monoculture allowed prediction of the composition of communities under rising temperature. We found that the final temperature of the habitat was the primary driver of diversity in our experimental communities. Species richness declined faster at higher temperatures. The negative effect of warming was not alleviated by a slower rate of warming or by dispersal among habitats and did not depend on the initial temperature. The response of evenness, however, did depend on the rate of change and on the initial temperature. Community composition was not predictable from monoculture assays, but higher fitness inequality (as seen by larger variance in growth rate among species in monoculture at higher temperatures) explained the faster loss of biodiversity with rising temperature.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

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

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The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

Changes in trophic state and aquatic communities in high Arctic ponds in response to increasing goose populations

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Relations between planktivorous fish abundance,zooplankton and phytoplankton in three lakes of differing productivity

Hydrobiologia - Water chemistry, phytoplankton, zooplankton and fish populations werestudied over several years in three shallow, non-stratified lakes withdiffering nutrient loadings and fish...     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

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

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Temperature Responses to Infrared-Loading and Water Table Manipulations in Peatland Mesocosms

We initiated a multi-factor global change experiment to explore the effects of infrared heat loading (HT) and water table level (WL) treatment on soil temperature (T) in bog and fen peatland mesocosms. We found that the temperature varied highly by year, month, peatland type, soil depth, HT and WL manipulations. The highest effect of HT on the temperature at 25 cm depth was found in June for the bog mesocosms (3.34-4.27℃) but in May for the fen mesocosms (2.32-4.33℃) over the 2-year study period. The effects of WL in the bog mesocosms were only found between August and January, with the wet mesocosms warmer than the dry mesocosms by 0.48-2.03 ℃ over the 2-year study period. In contrast, wetter fen mesocoams were generally cooler by 0.16-3.87℃. Seasonal changes of temperatures elevated by the HT also varied by depth and ecosystem type, with temperature differences at 5 cm and 10 cm depth showing smaller seasonal fluctuations than those at 25 cm and 40 cm in the bog mesocosrns. However, increased HT did not always lead to warmer soil, especially in the fen mesocosms. Both HT and WL manipulations have also changed the length of the non-frozen season.     

Direct and transgenerational effects of an experimental heatwave on early life stages in a freshwater snail

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Spatial variability in the response of lower trophic levels after carp exclusion from a freshwater marsh

Large common carp (Cyprinus carpio >30 cm) wereexcluded from a turbid, eutrophic coastal marsh of Lake Ontario with theconstruction of a fishway at the outlet. The marsh was sampledintensively for 2 seasons prior to (1993, 1994) and following (1997,1998) carp exclusion to study changes in water quality and shifts incommunity structure of phytoplankton and zooplankton. Samples werecollected from May to September in three habitats: open water, vegetated(cattail beds) and sewage lagoon. In the first year after carpexclusion, mean seasonal water turbidity decreased at all sites by49–80%; this was accompanied by growth of submergentplants in shallow, sheltered areas including the vicinity of cattails atthe vegetated site. This drop in turbidity was not significant in thesecond year after exclusion at the open water and lagoon sites, withturbidity levels declining by only 26–54% of1993–1994 values; only the vegetated site showed a sustaineddecrease in turbidity and persistent growth of submergent plants. At thevegetated site, increased clarity was concurrent with a significantreduction in edible algal biomass and an increased representation oflarge zooplankton grazers and substrate-associated cladocerans. At theopen water site, a spring clear-water phase was evident during the firstyear of exclusion and this coincided with the unusual appearance of alarge population of Daphnia. Compared to the other sites, thelagoon remained relatively turbid throughout the study. Results of thisstudy indicate that the response of lower trophic levels tobiomanipulation was variable from site-to-site and contributed to theco-existence of two alternative states in the marsh. In vegetated areas,water clarity was maintained by a positive feedback system betweenzooplankton and submergent macrophytes in the first 2 years followingexclusion. We suggest that both benthivore removal (to reducebioturbation) and planktivore reductions (to produce top down effects)were required to produce clear water and allow submersed macrophytegrowth. Although carp removal likely contributed to a 45%reduction in turbidity, an unusual climactic event in 1997, resulting indelayed fish spawning in the marsh, temporarily reduced zooplanktivoryand favoured zooplankton grazing-induced water clarity improvements.     

Impacts of soil temperature,phenology and plant community composition on invertebrate herbivory in a natural warming experiment

Species and community-level responses to warming are well documented, with plants and invertebrates known to alter their range, phenology or composition as temperature increases. The effects of warming on biotic interactions are less clearly understood, but can have consequences that cascade through ecological networks. Here, we used a natural soil temperature gradient of 5–35°C in the Hengill geothermal valley, Iceland, to investigate the effects of temperature on plant community composition and plant–invertebrate interactions. We quantified the level of invertebrate herbivory on the plant community across the temperature gradient and the interactive effects of temperature, plant phenology (i.e. development stage) and vegetation community composition on the probability of herbivory for three ubiquitous plant species, Cardamine pratensis, Cerastium fontanum and Viola palustris. We found that the percentage cover of graminoids and forbs increased, while the amount of litter decreased, with increasing soil temperature. Invertebrate herbivory also increased with soil temperature at the plant community level, but this was underpinned by different effects of temperature on herbivory for individual plant species, mediated by the seasonal development of plants and the composition of the surrounding vegetation. This illustrates the importance of considering the development stage of organisms in climate change research given the variable effects of temperature on susceptibility to herbivory at different ontogenetic stages.     

Changes in the lower trophic levels as a consequence of the level of fish manipulation in the ponds

Cyprinid fish of different mature age classes (3+ -4+) and stocks (100, 300 and 500 kg/ha) were introduced into each of three experimental ponds with area of 0.3 ha (average depth ca 1.7 m) while the fourth pond was left free of fish. Bream (Abramis brama L.), white bream (Blicca bjoerkna L.) and roach (Rutilus rutilus L.) made up 75% of the total cyprinid biomass, with wild carp (Cyprinus carpio L.) as the remaining 25%. The introduced fish spawned successfully. The high (above 300 kg/ha) planktivorous and benthivorous fish stocks resulted in several qualitative and quantitative alterations of the food chain structure in our simulation pond experiments. These alterations must primarily be assigned to changes caused by both the zooplanktivory and benthivory nature of the stocked fish populations. At the higher levels of fish biomass, Secchi depth was influenced significantly by chlorophyll-a concentration. Most of the variance in suspended solids concentration could be explained by the biomass ratio of the mature benthivorous fish. There was a clear shift in algal cell size in the ponds with the higher fish stocks: ponds with more fish had larger cells later in the summer. The relative influence of young cyprinid fish on crustaceans species composition and biomass, and mature populations on benthic fauna abundance and biomass, was sufficiently greater at higher (300–500 kg/ha) fish stock rates.     

Testing a biological mechanism of the insurance hypothesis in experimental aquatic communities

1.  The insurance hypothesis predicts a stabilizing effect of increasing species richness on community and ecosystem properties. Difference among species' responses to environmental fluctuations provides a general mechanism for the hypothesis. Previous experimental investigations of the insurance hypothesis have not examined this mechanism directly.
2.  First, responses to temperature of four protist species were measured in laboratory microcosms. For each species, we measured the response of intrinsic rate of increase ( r ) and carrying capacity ( K ) to temperature.
3.  Next, communities containing pairs of species were exposed to temperature fluctuations. Community biomass varied less when correlation in K between species (but not r ) was more negative, and this resulted from more negative covariances in population sizes, as predicted. Results were contingent on species identity, with findings differing between analyses including or not including communities containing one particular species.
4.  These findings provide the clearest support to date for this mechanism of the insurance hypothesis. Biodiversity, in terms of differences in species' responses to environmental fluctuations (i.e. functional response diversity) stabilizes community dynamics.     

Multi‐scale responses to warming in an experimental insect metacommunity

In metacommunities, diversity is the product of species interactions at the local scale and dispersal between habitat patches at the regional scale. Although warming can alter both species interactions and dispersal, the combined effects of warming on these two processes remains uncertain. To determine the independent and interactive effects of warming‐induced changes to local species interactions and dispersal, we constructed experimental metacommunities consisting of enclosed milkweed patches seeded with five herbivorous milkweed specialist insect species. We treated metacommunities with two levels of warming (unwarmed and warmed) and three levels of connectivity (isolated, low connectivity, high connectivity). Based on metabolic theory, we predicted that if plant resources were limited, warming would accelerate resource drawdown, causing local insect declines and increasing both insect dispersal and the importance of connectivity to neighboring patches for insect persistence. Conversely, given abundant resources, warming could have positive local effects on insects, and the risk of traversing a corridor to reach a neighboring patch could outweigh the benefits of additional resources. We found support for the latter scenario. Neither resource drawdown nor the weak insect‐insect associations in our system were affected by warming, and most insect species did better locally in warmed conditions and had dispersal responses that were unchanged or indirectly affected by warming. Dispersal across the matrix posed a species‐specific risk that led to declines in two species in connected metacommunities. Combined, this scaled up to cause an interactive effect of warming and connectivity on diversity, with unwarmed metacommunities with low connectivity incurring the most rapid declines in diversity. Overall, this study demonstrates the importance of integrating the complex outcomes of species interactions and spatial structure in understanding community response to climate change.