Hatching under brownification: DOC-mediated changes in physical, but not chemical properties of water affect hatching patterns of Cladocera resting eggs.

Dissolved organic carbon (DOC) is often related to the brownification of water in continental aquatic systems and to changes in the physiology of zooplankton organisms. Zooplankton resting eggs are particularly sensitive to changes in light and chemical characteristics of water, but the physical and chemical effects associated to DOC on dormant stages have never been tested before. Herein, we tested how DOC affects hatching rates and time to hatching of Cladocera (Diaphanosoma birgei) resting eggs. In order to analyze the chemical (i.e., toxic) and physical (i.e., light attenuation) effects of DOC on hatching patterns, resting eggs were exposed to different concentrations of DOC (0, 50, and 100 mg L-1) in an experimental design which isolated chemical from physical effects. When evaluating the physical effects of DOC, hatching was more than 150% less in 100 than 50 mg L-1 DOC and time to hatching was 66% lesser in 50 mg L-1 DOC than control. Hatchling numbers and time to hatching were not affected by DOC chemical effects. We conclude DOC effects on hatching mainly relied on light attenuation, while chemical effects were likely of minor importance. DOC may change Cladocera emergence patterns mainly through light attenuation in the water column.

New insights on the relation between zooplankton and humic substances in tropical freshwater ecosystems.

Aquatic humic substances (HS) represent about 60-95% of the dissolved organic carbon (DOC) present in coastal environments of northern Rio de Janeiro state, Brazil. Although they are important regulators of processes involving aquatic communities, the response of the zooplankton community to their presence remains poorly understood, especially in natural tropical environments. Therefore, our objective was to elucidate zooplankton community responses along a natural gradient of HS. Such natural humic gradient was obtained in coastal freshwater environments with distinct DOC concentrations (20-200 mg L-1). Results show a decrease in zooplankton density and biomass along the HS gradient. However, microphages organisms (e.g. non-predatory rotifers and smaller testate amoeba, such as Difflugia) were most present in environments with higher concentration of HS, probably due to a stronger importance of the microbial-loop in these environments. Some species - such as Scapholeberis armata (Cladocera) and Lecane boettgeri (Rotifera) were only accounted for environments with high HS concentration, illustrating their potential as bioindicators for HS presence. Nevertheless, we were able to observe the effects of HS on the structure and composition of primary consumers and how these substances might indirectly affect species dynamics. We point out to new findings in highly humic tropical environments, which are still poorly studied and understood.

Effects of the introduction of an omnivorous fish on the biodiversity and functioning of an upland Amazonian lake / Efeitos da introdução de um peixe onívoro sobre a biodiversidade e funcionamento de um lago de altitude amazônico

The introduction of nonnative species is one of the main threats to freshwater ecosystems. Although omnivory and intraguild predation are common in those systems, little is known about the effects of introduced omnivorous fish on pelagic and littoral communities. This study tested predictions of food-web theory regarding the effects of omnivorous fish introduction on previously fishless lakes in the Amazonian uplands of Serra dos Carajás, Pará, Brazil. The trophic structure of two similar lakes, one with and the other without the introduced omnivorous fish Astyanax bimaculatus, was compared using a data series of biotic variables collected from both lakes twice a year from 2010 to 2013. Zooplankton was more abundant in the lake with fish, and the zooplankton composition differed between lakes. Phytoplankton richness and chlorophyll-a were higher in the lake with the introduced fish than in the fishless lake regardless of phosphorus limitation. For the benthic macroinvertebrate communities, species richness and biomass were higher in the fishless lake. Our results also indicate that A. bimaculatus has the potential to link pelagic and littoral habitats through nutrient cycling. The differences observed between the studied lakes are consistent with predictions from food-web theory regarding the effects of multichain omnivorous fish on trophic dynamics. Despite limitations regarding replication at the ecosystem level, it is possible to infer from our findings that the introduction of an omnivorous fish might have changed lake overall functioning.(AU)

A introdução de espécies não nativas é uma ameaça aos ecossistemas de água doce. Embora a onivoria e a predação intraguilda sejam comuns nesses sistemas, os efeitos da introdução de peixes onívoros nas comunidades pelágicas e litorâneas é pouco conhecido. Nós testamos as previsões da teoria da teia trófica considerando os efeitos da introdução de um peixe onívoro em um lago previamente desprovido de peixes localizado na Serra dos Carajás, Pará, Brasil. A estrutura trófica em dois lagos similares, um com a presença do peixe onívoro introduzido Astyanax bimaculatus, e outro sem peixes, foi comparada através de uma série de dados bióticos amostrados entre 2010 e 2013. A comunidade zooplanctônica foi mais abundante no lago com peixe e sua composição diferiu entre os dois lagos. Apesar da limitação por fósforo no lago com peixe, a riqueza do fitoplâncton e a concentração de clorofila-a foram maiores nesse lago. A comunidade de macroinvertebrados bentônicos apresentou maior riqueza e biomassa no lago sem peixe. Nossos resultados também indicam que A. bimaculatus tem o potencial de acoplar as comunidades litorâneas e pelágicas através da reciclagem de nutrientes. Apesar das limitações relacionadas à ausência de replicação no nível de ecossistemas, nós argumentamos que o nosso estudo mostra que a introdução do peixe onívoro pode ter causado mudanças no funcionamento do lago.(AU)

Effects of the introduction of an omnivorous fish on the biodiversity and functioning of an upland Amazonian lake

The introduction of nonnative species is one of the main threats to freshwater ecosystems. Although omnivory and intraguild predation are common in those systems, little is known about the effects of introduced omnivorous fish on pelagic and littoral communities. This study tested predictions of food-web theory regarding the effects of omnivorous fish introduction on previously fishless lakes in the Amazonian uplands of Serra dos Carajás, Pará, Brazil. The trophic structure of two similar lakes, one with and the other without the introduced omnivorous fish Astyanax bimaculatus, was compared using a data series of biotic variables collected from both lakes twice a year from 2010 to 2013. Zooplankton was more abundant in the lake with fish, and the zooplankton composition differed between lakes. Phytoplankton richness and chlorophyll-a were higher in the lake with the introduced fish than in the fishless lake regardless of phosphorus limitation. For the benthic macroinvertebrate communities, species richness and biomass were higher in the fishless lake. Our results also indicate that A. bimaculatus has the potential to link pelagic and littoral habitats through nutrient cycling. The differences observed between the studied lakes are consistent with predictions from food-web theory regarding the effects of multichain omnivorous fish on trophic dynamics. Despite limitations regarding replication at the ecosystem level, it is possible to infer from our findings that the introduction of an omnivorous fish might have changed lake overall functioning. (AU)

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.

Biodiversity effects of ecosystem engineers are stronger on more complex ecosystem processes.

The relative importance of species richness and identity for the diversity-function relationship remains controversial. We mechanistically explored the potential contribution of ecosystem processes complexity (EPC; i.e., the number of pathways and mechanisms through which an ecosystem process can be directly and/or indirectly affected by species and/or their interactions) to the resolution of this controversy. We hypothesized that the complementarity effects of biodiversity will be stronger and that the diversity-function relationship will be more dependent on species richness as the EPC increases. Using a benthic bioturbator community as a model system we tested these predictions across ecosystem processes that could be ordered according to their complexity (suspended material flux < PO4-P flux < NH4-N flux < bacterioplankton production). Consistent with our predictions, species richness explained an increasing proportion of data variation as EPC increased, whereas the contrary was observed for species composition. Nontransgressive overyielding was not affected by EPC, but the magnitude of transgressive overyielding increased significantly with EPC, indicating that complementarity may be stronger as EPC increases. Our results highlight the importance of considering the interactive role of the characteristics of ecosystem processes in our theoretical understanding of the diversity-function relationship and its underlying mechanisms.

Community biomass and bottom up multivariate nutrient complementarity mediate the effects of bioturbator diversity on pelagic production.

Tests of the biodiversity and ecosystem functioning (BEF) relationship have focused little attention on the importance of interactions between species diversity and other attributes of ecological communities such as community biomass. Moreover, BEF research has been mainly derived from studies measuring a single ecosystem process that often represents resource consumption within a given habitat. Focus on single processes has prevented us from exploring the characteristics of ecosystem processes that can be critical in helping us to identify how novel pathways throughout BEF mechanisms may operate. Here, we investigated whether and how the effects of biodiversity mediated by non-trophic interactions among benthic bioturbator species vary according to community biomass and ecosystem processes. We hypothesized that (1) bioturbator biomass and species richness interact to affect the rates of benthic nutrient regeneration [dissolved inorganic nitrogen (DIN) and total dissolved phosphorus (TDP)] and consequently bacterioplankton production (BP) and that (2) the complementarity effects of diversity will be stronger on BP than on nutrient regeneration because the former represents a more integrative process that can be mediated by multivariate nutrient complementarity. We show that the effects of bioturbator diversity on nutrient regeneration increased BP via multivariate nutrient complementarity. Consistent with our prediction, the complementarity effects were significantly stronger on BP than on DIN and TDP. The effects of the biomass-species richness interaction on complementarity varied among the individual processes, but the aggregated measures of complementarity over all ecosystem processes were significantly higher at the highest community biomass level. Our results suggest that the complementarity effects of biodiversity can be stronger on more integrative ecosystem processes, which integrate subsidiary "simpler" processes, via multivariate complementarity. In addition, reductions in community biomass may decrease the strength of interspecific interactions so that the enhanced effects of biodiversity on ecosystem processes can disappear well before species become extinct.

Ecological determinism increases with organism size.

After much debate, there is an emerging consensus that the composition of many ecological communities is determined both by species traits, as proposed by niche theory, as well as by chance events. A critical question for ecology is, therefore, which attributes of species predict the dominance of deterministic or stochastic processes. We outline two hypotheses by which organism size could determine which processes structure ecological communities, and we test these hypotheses by comparing the community structure in bromeliad phytotelmata of three groups of organisms (bacteria, zooplankton, and macroinvertebrates) that encompass a 10 000-fold gradient in body size, but live in the same habitat. Bacteria had no habitat associations, as would be expected from trait-neutral stochastic processes, but still showed exclusion among species pairs, as would be expected from niche-based processes. Macroinvertebrates had strong habitat and species associations, indicating niche-based processes. Zooplankton, with body size between bacteria and macroinvertebrates, showed intermediate habitat associations. We concluded that a key niche process, habitat filtering, strengthened with organism size, possibly because larger organisms are both less plastic in their fundamental niches and more able to be selective in dispersal. These results suggest that the relative importance of deterministic and stochastic processes may be predictable from organism size.

Food web architecture and basal resources interact to determine biomass and stoichiometric cascades along a benthic food web.

Understanding the effects of predators and resources on primary producers has been a major focus of interest in ecology. Within this context, the trophic cascade concept especially concerning the pelagic zone of lakes has been the focus of the majority of these studies. However, littoral food webs could be especially interesting because base trophic levels may be strongly regulated by consumers and prone to be light limited. In this study, the availability of nutrients and light and the presence of an omnivorous fish (Hyphessobrycon bifasciatus) were manipulated in enclosures placed in a humic coastal lagoon (Cabiúnas Lagoon, Macaé - RJ) to evaluate the individual and interactive effects of resource availability (nutrients and light) and food web configuration on the biomass and stoichiometry of periphyton and benthic grazers. Our findings suggest that light and nutrients interact to determine periphyton biomass and stoichiometry, which propagates to the consumer level. We observed a positive effect of the availability of nutrients on periphytic biomass and grazers' biomass, as well as a reduction of periphytic C∶N∶P ratios and an increase of grazers' N and P content. Low light availability constrained the propagation of nutrient effects on periphyton biomass and induced higher periphytic C∶N∶P ratios. The effects of fish presence strongly interacted with resource availability. In general, a positive effect of fish presence was observed for the total biomass of periphyton and grazer's biomass, especially with high resource availability, but the opposite was found for periphytic autotrophic biomass. Fish also had a significant effect on periphyton stoichiometry, but no effect was observed on grazers' stoichiometric ratios. In summary, we observed that the indirect effect of fish predation on periphyton biomass might be dependent on multiple resources and periphyton nutrient stoichiometric variation can affect consumers' stoichiometry.

Natural dissolved humic substances increase the lifespan and promote transgenerational resistance to salt stress in the cladoceran Moina macrocopa.

PURPOSE: Evidence has accumulated that humic substances (HS) are not inert biogeochemicals. Rather, they cause stress symptoms and may modulate the life history of aquatic organisms. Nevertheless, it is still not clear how HS interact with additional stressors and if their effects are transgenerational. We tested the interactive effects of HS and salt to cladocerans, discussing their consequences for the persistence in fluctuating environments, such as coastal lagoons. METHODS: We used life-table experiments to test the effects of natural HS from a polyhumic coastal lagoon (0, 5, 10, 20, 50, and 100 mg dissolved organic carbon (DOC) L(-1)) on the life-history of the cladoceran Moina macrocopa. We further tested the effects of HS (10 mg DOC L(-1)), within and across generations, on the resistance of M. macrocopa to salt stress (5.5 g L(-1)). RESULTS: HS at 5-20 mg DOC L(-1) extended the mean lifespan of M. macrocopa by ~30%. HS also increased body length at maturity by ~4% at 5-50 mg DOC L(-1) and stimulated male offspring production at all tested concentrations. Exposure to HS (even maternal only) alleviated the salt-induced reduction of somatic growth. Co-exposure to HS increased body volume by 12-22% relative to salt-only treatments, while pre-exposure to HS increased body volume by 40-56% in treatments with salt presence, when compared to non-pre-exposed animals. CONCLUSIONS: HS at environmentally realistic concentrations, by acting as mild chemical stressors, modify crucial life-history traits of M. macrocopa, favoring its persistence in fluctuating environments. Some of the effects of HS are even transgenerational.

Synergy of fresh and accumulated organic matter to bacterial growth.

The main goal of this research was to evaluate whether the mixture of fresh labile dissolved organic matter (DOM) and accumulated refractory DOM influences bacterial production, respiration, and growth efficiency (BGE) in aquatic ecosystems. Bacterial batch cultures were set up using DOM leached from aquatic macrophytes as the fresh DOM pool and DOM accumulated from a tropical humic lagoon. Two sets of experiments were performed and bacterial growth was followed in cultures composed of each carbon substrate (first experiment) and by carbon substrates combined (second experiment), with and without the addition of nitrogen and phosphorus. In both experiments, bacterial production, respiration, and BGE were always higher in cultures with N and P additions, indicating a consistent inorganic nutrient limitation. Bacterial production, respiration, and BGE were higher in cultures set up with leachate DOM than in cultures set up with humic DOM, indicating that the quality of the organic matter pool influenced the bacterial growth. Bacterial production and respiration were higher in the mixture of substrates (second experiment) than expected by bacterial production and respiration in single substrate cultures (first experiment). We suggest that the differences in the concentration of some compounds between DOM sources, the co-metabolism on carbon compound decomposition, and the higher diversity of molecules possibly support a greater bacterial diversity which might explain the higher bacterial growth observed. Finally, our results indicate that the mixture of fresh labile and accumulated refractory DOM that naturally occurs in aquatic ecosystems could accelerate the bacterial growth and bacterial DOM removal.

Influence of hydrological pulse on bacterial growth and DOC uptake in a clear-water Amazonian lake.

This study was conducted to evaluate: (1) the bacterial growth and the dissolved organic carbon (DOC) uptake in an Amazonian lake (Lake Batata) at high-water and low-water periods of the flood pulse; (2) the influence of nitrogen and phosphorus (NP) additions on bacterial growth and DOC uptake in Lake Batata at two flood pulse periods; and (3) the bioavailability of the main DOC sources in Lake Batata. Lake Batata is a typical clear-water Amazonian lake, located in the watershed of Trombetas River, Central Amazon, Brazil. Bacterial batch cultures were set up with 90% 0.2-microm filtered water and 10% inoculum from Lake Batata. N-NH(4)NO(3) and P-KH(2)PO(4), with final concentrations of 50 and 5 microM, respectively, were added to the cultures, except for controls. Extra sources of DOC (e.g., algal lysate, plant leachates) were added to constitute six distinct treatments. Bacterial response was measured by maximum bacterial abundance and rates of bacterial production, respiration, DOC uptake, and bacterial growth efficiency (BGE). Bacterial growth and DOC uptake were higher in NP treatments than in controls, indicating a consistent nutrient limitation in Lake Batata. The composition of DOC also seems to be an important regulating factor of bacterial growth in Lake Batata. Seasonally, bacterial growth and DOC bioavailability were higher at low-water period, when the phytoplankton is a significant extra source of DOC, than at high-water period, when the forest is the main source of DOC. DOC bioavailability was better estimated based on the diversity and the diagenetic stage of carbon compounds than on single classes of labile compounds. Changes in BGE were better related to CNP stoichiometry in the water, and the "excess" of organic substrates was oxidized in catabolism, despite the quality of these compounds for bacterial growth. Finally, we conclude that bacterial growth and DOC uptake vary throughout the flood pulse in clear-water Amazonian ecosystems as a result of changes in nutrient concentration and in DOC composition.

Complementary pathways of dissolved organic carbon removal pathways in clear-water Amazonian ecosystems: photochemical degradation and bacterial uptake.

Dissolved organic carbon (DOC) photochemical reactions establish important links between DOC and planktonic bacteria. We hypothesize that seasonal changes in DOC quality, related to the flood pulse, drive the effects of light-DOC interactions on uptake by planktonic bacteria uptake in clear-water Amazonian ecosystems. Water samples from two ecosystems (one lake and one stream) were incubated in sunlight during different hydrological periods and were then exposed to bacterial degradation. Photochemical and bacterial degradation were driven by seasonal DOC inputs. Bacterial mineralization was the main degradation pathway of autochthonous DOC in the lake, while allochthonous DOC was more available for photochemical oxidation. We suggest that sunlight enhances the bacterial uptake of refractory DOC but does not alter uptake of labile forms. We also observed a positive relationship between sunlight and bacterial degradation of DOC, instead of competition. We conclude that photochemical reactions and bacteria complementarily degrade the different sources of DOC during the flood pulse in Amazonian clear-water aquatic ecosystems.