Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

Regime shifts in a shallow lake over 12 years: Consequences for taxonomic and functional diversities, and ecosystem multifunctionality.

Under increasing nutrient loading, shallow lakes may shift from a state of clear water dominated by submerged macrophytes to a turbid state dominated by phytoplankton or a shaded state dominated by floating macrophytes. How such regime shifts mediate the relationship between taxonomic and functional diversities (FD) and lake multifunctionality is poorly understood. We employed a detailed database describing a shallow lake over a 12-year period during which the lake has displayed all the three states (clear, turbid and shaded) to investigate how species richness, FD of fish and zooplankton, ecosystem multifunctionality and five individual ecosystem functions (nitrogen and phosphorus concentrations, standing fish biomass, algae production and light availability) differ among states. We also evaluated how the relationship between biodiversity (species richness and FD) and multifunctionality is affected by regime shifts. We showed that species richness and the FD of fish and zooplankton were highest during the clear state. The clear state also maintained the highest values of multifunctionality as well as standing fish biomass production, algae biomass and light availability, whereas the turbid and shaded states had higher nutrient concentrations. Functional diversity was the best predictor of multifunctionality. The relationship between FD and multifunctionality was strongly positive during the clear state, but such relationship became flatter after the shift to the turbid or shaded state. Our findings illustrate that focusing on functional traits may provide a more mechanistic understanding of how regime shifts affect biodiversity and the consequences for ecosystem functioning. Regime shifts towards a turbid or shaded state negatively affect the taxonomic diversity and FD of fish and zooplankton, which in turn impairs the multifunctionality of shallow lakes.

Predation by an omnivorous fish and food availability alter zooplankton functional diversity: a microcosm approach.

Studies considering the functional traits of organisms, populations, and communities functional indices increase the understanding of many factors on ecosystem functioning. Here, we analyze the predation effect (by fish) on zooplankton functional diversity and the effects of biomass and density of periphytic algae on zooplankton feeding type trait and body size. We expect that intense predation by fish on zooplankton leads to higher values of zooplankton functional diversity and that food resource will be positively related to the abundance of zooplankton trait and body size. For that, microcosms were established (T1- fish-absence, and T2- fish-presence, both with periphytic algae as food). We observed that fish presence decreased zooplankton functional diversity through modifications in the availability of nutrients and algae, through the middle-out effect. We also observed that body size had a negative relationship with the food resource, reaffirming that high food availability in subtropical lakes is linked to small-bodied zooplankton. The raptorial copepods covariate positively with the periphytic algae, which was an alternative food resource and, in this case, the main form of carbon input into the system. In this study, omnivorous fish reduced zooplankton functional traits, which can alter the energy stock and energy flow in aquatic ecosystems.

Influence of the chlorophyll-a gradient on the community structure of plankton microcrustaceans (Cladocera and Copepoda) in a Neotropical reservoir.

Fish farming in net cages is considered as an alternative to food production in response to elevated population growth, and zooplankton is an important resource to the development of this economic activity. We (i) compared microcrustacean composition in eutrophic and oligotrophic habitats under net tank influence, (ii) investigated changes in species distributions in these habitats, and (iii) indicated which chlorophyll-a concentrations presented thresholds that alter community structure. We expected different responses of species to changes in chlorophyll-a concentration due to net cage management, as chlorophyll-a represents an estimate of food availability. Microcrustacean samplings and chlorophyll-a estimation were made upstream, downstream and close to the net cages, during 120 days, in the Rosana Reservoir (Brazil). Species composition differed significantly (p<0.05) among habitats where in the eutrophic environment was found the largest number of species. However, only in the eutrophic habitats did frequency of occurrence and relative abundance of some species change with chlorophyll-a variation. Thus, net cage management influenced species distribution only in the most productive habitat. These responses can affect ecosystem processes related to trophic dynamics as secondary productivity and nutrient cycling.

Non-predatory mortality of planktonic microcrustaceans (Cladocera and Copepoda) in neotropical semiarid reservoirs.

The accuracy of traditional methods to sample planktonic microcrustaceans depends on two assumptions: that organisms are alive during sampling and that all carcasses can be identified despite their degradation state, but fresh carcasses are not easy to distinguish by traditional methods. Previous studies about mortality have shown that neglecting dead organisms can provide biased ecological information. Thus, our objective was to determine the mortality rate and the proportion of dead microcrustacean in three tropical reservoirs. Sampling was carried out in 12 stations during two periods. The proportion of dead organisms was verified using aniline blue and it varied between 0.6% and 90.6%. The carcass decomposition period varied between 3 to 16 days and microcrustaceans mortality rate varied between 0.005 and 0.314 d-1. Traditional preservation techniques with formalin do not significantly overestimate species abundance. However, these values should not be disregarded, because corrected (disregarding organisms that were dead) and formalin-preserved abundances were correlated with distinct limnological descriptors. Therefore, the traditional formalin preservation technique could provide misleading ecological interpretations. Other studies over larger temporal scales in addition to experiments to evaluate the effects of viruses, parasitism and the toxic effects of cyanobacteria on zooplankton would enlighten mortality rate patterns in freshwater ecosystems.

The influence of heavy metals on toxicogenetic damage in a Brazilian tropical river.

Anthropogenic activities in tropical rivers favor the eutrophication process, which causes increased concentration of heavy metals. The presence and bioaccumulation of metals are directly related to the presence of genotoxic damage in aquatic organisms. Thus, we evaluated the presence of heavy metals (Fe, Zn, Cr, Cu and Al) and performed toxicogenetic tests in surface (S) and bottom (B) of water samples of the Poti river (Piaui/Brazil). Cytotoxicity and genotoxicity tests were performed in Allium cepa, and micronucleus (MN) and comet assay were performed in Oreochromis niloticus. The chemical analysis showed concentrations above the limit for Cu, Cr, Fe and Al according to Brazilian laws, characterizing anthropogenic disturbance in this aquatic environment. Toxicogenetic analysis presented significant cytotoxic, mutagenic and genotoxic effects in different exposure times and water layers (S and B), especially alterations in mitotic spindle defects, MN formations, nuclear bud and DNA strand breaks. Correlations between Fe and cytotoxicity, and Al and mutagenicity were statistically significant and point out to the participation of heavy metals in genotoxic damage. Therefore, Poti river water samples presented toxicogenetic effects on all bioindicators analyzed, which are most likely related to heavy metals pollution.

Correlates of zooplankton beta diversity in tropical lake systems.

The changes in species composition between habitat patches (beta diversity) are likely related to a number of factors, including environmental heterogeneity, connectivity, disturbance and productivity. Here, we used data from aquatic environments in five Brazilian regions over two years and two seasons (rainy and dry seasons or high and low water level periods in floodplain lakes) in each year to test hypotheses underlying zooplankton beta diversity variation. The regions present different levels of hydrological connectivity, where three regions present lakes that are permanent and connected with the main river, while the water bodies of the other two regions consist of permanent lakes and temporary ponds, with no hydrological connections between them. We tested for relationships between zooplankton beta diversity and environmental heterogeneity, spatial extent, hydrological connectivity, seasonality, disturbance and productivity. Negative relationships were detected between zooplankton beta diversity and both hydrological connectivity and disturbance (periodic dry-outs). Hydrological connectivity is likely to affect beta diversity by facilitating dispersal between habitats. In addition, the harsh environmental filter imposed by disturbance selected for only a small portion of the species from the regional pool that were able to cope with periodic dry-outs (e.g., those with a high production of resting eggs). In summary, this study suggests that faunal exchange and disturbance play important roles in structuring local zooplankton communities.

Dispersal ability determines the role of environmental, spatial and temporal drivers of metacommunity structure.

Recently, community ecologists are focusing on the relative importance of local environmental factors and proxies to dispersal limitation to explain spatial variation in community structure. Albeit less explored, temporal processes may also be important in explaining species composition variation in metacommunities occupying dynamic systems. We aimed to evaluate the relative role of environmental, spatial and temporal variables on the metacommunity structure of different organism groups in the Upper Paraná River floodplain (Brazil). We used data on macrophytes, fish, benthic macroinvertebrates, zooplankton, periphyton, and phytoplankton collected in up to 36 habitats during a total of eight sampling campaigns over two years. According to variation partitioning results, the importance of predictors varied among biological groups. Spatial predictors were particularly important for organisms with comparatively lower dispersal ability, such as aquatic macrophytes and fish. On the other hand, environmental predictors were particularly important for organisms with high dispersal ability, such as microalgae, indicating the importance of species sorting processes in shaping the community structure of these organisms. The importance of watercourse distances increased when spatial variables were the main predictors of metacommunity structure. The contribution of temporal predictors was low. Our results emphasize the strength of a trait-based analysis and of better defining spatial variables. More importantly, they supported the view that "all-or-nothing" interpretations on the mechanisms structuring metacommunities are rather the exception than the rule.

Intra and inter-annual structure of zooplankton communities in floodplain lakes: a long-term ecological research study

Water flow management has significantly changed the natural dynamic of floods, which are responsible for the structure and dynamic of aquatic communities in river-floodplain systems. With the aim to elaborate a conceptual framework that describes the main ecological factors associated with zooplankton community structure in the Upper Paraná River, we investigated the mechanisms that regulate the communities structure and their response to inter-annual and hydro-sedimentological variations in the floodplain and the biological factors associated with species abundance in those communities. For this we conducted samplings every six months (potamophase in March and limnophase in September) to characterize intra and inter-annual variations in community structure between 2000 and 2008. The intra-annual differences on the species richness, abundance, Shannon diversity index, and evenness, were conducted using Bayesian procedures to show probabilistic predictions of the data fit to main variation sources. Non-metric multi-dimensional scaling (NMDS), multi-response permutation procedure (MRPP), and indicator species analysis (IndVal) were run to assess and characterize the seasonality of the community structure. During high water (potamophase), hydrologic connectivity favoured exchange and dispersal of species in some lakes, increasing local diversity; during low water (limnophase), higher local productivity favoured opportunistic taxa, increasing species dominance and decreasing local diversity. Food resources and density of small-size fish were biological factors associated with the seasonal dynamic of the zooplankton community; these factors were dependent on hydrosedimentological phase (potamophase or limnophase). Water levels and limnological modifications related to water flow management have promoted replacement and impoverishment of aquatic biota in affected lakes and have indicated the ecological importance of a natural dynamic flood, which displays regular flood pulses. The conceptual model presented encompassed interactions between diverse environmental variables to more understandable mechanisms of the main sources of community variation.

El manejo del régimen de inundación ha cambiado de manera significativa la dinámica natural de las inundaciones, que son responsables de la estructura y dinámica de las comunidades acuáticas en sistemas río-planicie de inundación. En este sentido, investigamos cómo la estructura de las comunidades zooplanctónicas responde a variaciones estacionales e interanuales en los sistemas de llanura de inundación, y los factores biológicos asociados con la abundancia de especies de las comunidades zooplanctonicas. Elaboramos también, un marco conceptual que describe los principales factores ecológicos asociados con la estructura de las comunidades para los sistemas del Alto río Paraná. Para ello se realizaron muestreos cada seis meses (potamophase en marzo y en septiembre limnophase) para caracterizar las variaciones intra e interanuales en la estructura de la comunidad entre 2000 y 2008. Las diferencias estacionales de la riqueza de especies, abundancia, índice de diversidad de Shannon y equitatividad, se llevaron a cabo utilizando procedimientos Bayesianos para mostrar predicciones probabilísticas de los datos ajustados para las principales fuentes variación. Análisis de ordenamiento no-métrico multi-dimensional (NDMS); procedimientos de permutación de respuestas múltiples (MRPP) y análisis de especies indicadoras (IndVal) fueron utilizados para evaluar y clasificar la estacionalidad de la estructura de la comunidad. Durante aguas altas (potamofase), la conectividad hidrológica favoreció el intercambio y dispersión de las especies en algunas lagunas, incrementando la diversidad local; durante aguas bajas (limnofase), la alta productividad local favoreció los táxones oportunistas, incrementando las especies dominante y disminuyendo la diversidad local. La disponibilidad de recursos alimenticios y la densidad de pequeños peces fueron los factores biológicos asociados con la dinámica estacional de la comunidad zooplanctonica; los cuales dependen de la fase hidrosedimentológica (potamofase o limnofase). Modificaciones del régimen hidrológico y limnológico relacionados con el manejo de inundación han promovido el reemplazo y empobrecimiento de la biota acuática, en las lagunas sin conexión afectadas, enfatizando así la importancia ecológica de la dinámica natural de inundaciones, que presenta pulsos regulares de inundación. El modelo conceptual que se presenta abarca desde interacciones entre diversas variables ambientales hasta mecanismos más comprensibles de las principales fuentes de variación de la comunidad.

Intra and inter-annual structure of zooplankton communities in floodplain lakes: a long-term ecological research study.

Water flow management has significantly changed the natural dynamic of floods, which are responsible for the structure and dynamic of aquatic communities in river-floodplain systems. With the aim to elaborate a conceptual framework that describes the main ecological factors associated with zooplankton community structure in the Upper Paraná River, we investigated the mechanisms that regulate the communities structure and their response to inter-annual and hydro-sedimentological variations in the floodplain and the biological factors associated with species abundance in those communities. For this we conducted samplings every six months (potamophase in March and limnophase in September) to characterize intra and inter-annual variations in community structure between 2000 and 2008. The intra-annual differences on the species richness, abundance, Shannon diversity index, and evenness, were conducted using Bayesian procedures to show probabilistic predictions of the data fit to main variation sources. Non-metric multi-dimensional scaling (NMDS), multi-response permutation procedure (MRPP), and indicator species analysis (IndVal) were run to assess and characterize the seasonality of the community structure. During high water (potamophase), hydrologic connectivity favoured exchange and dispersal of species in some lakes, increasing local diversity; during low water (limnophase), higher local productivity favoured opportunistic taxa, increasing species dominance and decreasing local diversity. Food resources and density of small-size fish were biological factors associated with the seasonal dynamic of the zooplankton community; these factors were dependent on hydrosedimentological phase (potamophase or limnophase). Water levels and limnological modifications related to water flow management have promoted replacement and impoverishment of aquatic biota in affected lakes and have indicated the ecological importance of a natural dynamic flood, which displays regular flood pulses. The conceptual model presented encompassed interactions between diverse environmental variables to more understandable mechanisms of the main sources of community variation.