Complex Effects of Ecosystem Engineer Loss on Benthic Ecosystem Response to Detrital Macroalgae

Ecosystem engineers change abiotic conditions, community assembly and ecosystem functioning. Consequently, their loss may modify thresholds of ecosystem response to disturbance and undermine ecosystem stability. This study investigates how loss of the bioturbating lugworm Arenicola marina modifies the response to macroalgal detrital enrichment of sediment biogeochemical properties, microphytobenthos and macrofauna assemblages. A field manipulative experiment was done on an intertidal sandflat (Oosterschelde estuary, The Netherlands). Lugworms were deliberately excluded from 1× m sediment plots and different amounts of detrital Ulva (0, 200 or 600 g Wet Weight) were added twice. Sediment biogeochemistry changes were evaluated through benthic respiration, sediment organic carbon content and porewater inorganic carbon as well as detrital macroalgae remaining in the sediment one month after enrichment. Microalgal biomass and macrofauna composition were measured at the same time. Macroalgal carbon mineralization and transfer to the benthic consumers were also investigated during decomposition at low enrichment level (200 g WW). The interaction between lugworm exclusion and detrital enrichment did not modify sediment organic carbon or benthic respiration. Weak but significant changes were instead found for porewater inorganic carbon and microalgal biomass. Lugworm exclusion caused an increase of porewater carbon and a decrease of microalgal biomass, while detrital enrichment drove these values back to values typical of lugworm-dominated sediments. Lugworm exclusion also decreased the amount of macroalgae remaining into the sediment and accelerated detrital carbon mineralization and CO₂ release to the water column. Eventually, the interaction between lugworm exclusion and detrital enrichment affected macrofauna abundance and diversity, which collapsed at high level of enrichment only when the lugworms were present. This study reveals that in nature the role of this ecosystem engineer may be variable and sometimes have no or even negative effects on stability, conversely to what it should be expected based on current research knowledge.     

Composition of inorganic and organic nutrient sources influences phytoplankton community structure in the New River Estuary, North Carolina

The New River Estuary, NC, is a nutrient-sensitive, eutrophic water body that is prone to harmful algal blooms. High annual loading from the watershed of varying nutrient forms, including inorganic phosphorus and inorganic and organic nitrogen, may be linked to the persistence of algal blooms in the estuary. In order to evaluate phytoplankton response to nutrient inputs, a series of in situ nutrient addition experiments were carried out during June 2010 to July 2011 on water from an estuarine site known to support algal blooms. Estuarine water was enriched with nutrients consisting of individual and combined sources of dissolved inorganic nitrogen, orthophosphate, urea, and a natural dissolved organic nitrogen (DON) addition derived from upstream New River water. The combined inorganic N and P addition most frequently stimulated phytoplankton biomass production as total chlorophyll a. The responses of diagnostic (of major algal groups) photopigments were also evaluated. Significant increases in peridinin (dinoflagellates), chlorophyll b (chlorophytes), and myxoxanthophyll (cyanobacteria) were most frequently promoted by additions containing riverine DON. Significant increases in zeaxanthin (cyanobacteria) were more frequently promoted by inorganic nitrogen additions, while increases in fucoxanthin (diatoms) and alloxanthin (cryptophytes) were not promoted consistently by any one nutrient treatment. Evaluating the impact of varying nutrient forms on phytoplankton community dynamics is necessary in order to develop strategies to avoid long-term changes in community structure and larger-scale changes in ecosystem condition.     

Benthic microalgae in coral reef sediments of the southern Great Barrier Reef,Australia

The abundance and productivity of benthic microalgae in coral reef sediments are poorly known compared with other, more conspicuous (e.g. coral zooxanthellae, macroalgae) primary producers of coral reef habitats. A survey of the distribution, biomass, and productivity of benthic microalgae on a platform reef flat and in a cross-shelf transect in the southern Great Barrier Reef indicated that benthic microalgae are ubiquitous, abundant (up to 995.0 mg chlorophyll (chl) a m^–2), and productive (up to 110 mg O₂ m^–2 h^–1) components of the reef ecosystem. Concentrations of benthic microalgae, expressed as chlorophyll a per surface area, were approximately 100-fold greater than the integrated water column concentrations of microalgae throughout the region. Benthic microalgal biomass was greater on the shallow water platform reef than in the deeper waters of the cross-shelf transect. In both areas the benthic microalgal communities had a similar composition, dominated by pennate diatoms, dinoflagellates, and cyanobacteria. Benthic microalgal populations were potentially nutrient-limited, based on responses to nitrogen and phosphorus enrichments in short-term (7-day) microcosm experiments. Benthic microalgal productivity, measured by O₂ evolution, indicated productive communities responsive to light and nutrient availability. The benthic microalgal concentrations observed (92–995 mg chl a m^–2) were high relative to other reports, particularly compared with temperate regions. This abundance of productive plants in both reef and shelf sediments in the southern Great Barrier Reef suggests that benthic microalgae are key components of coral reef ecosystems.Communicated by Environmental Editor, B.C. Hatcher     

Benthic community responses to nutrient enrichment and predator exclusion: Influence of background nutrient concentrations and interactive effects

Potential community effects of nutrient enhancement are a topic of theoretical interest and increasing management concern in coastal marine systems. While increased nutrient levels may lead to increased microalgal production and biomass, studies have provided variable evidence regarding the existence of upward cascade effects on macrofauna. In benthic marine communities, limitation by predation or factors preventing recruitment response may contribute to weak coupling between resource availability and macrobenthos abundances. We conducted blocked nutrient addition and predator exclusion experiments in the intertidal of two estuaries that varied in background nutrient concentrations (Cape Fear and White Oak, southeastern North Carolina). Benthic community comparisons were also made among these and two other North Carolina estuaries to examine correlations in distribution patterns. Cape Fear, which had the highest background nitrogen and phosphorus concentrations, also had highest ambient benthic microalgal biomass. There was no significant response of microalgal biomass to local nutrient additions in Cape Fear and only one macrofaunal taxon during one season exhibited abundance responses to nutrient additions. White Oak, with lower background nutrient levels, was characterized by significant microalgal responses to nutrient additions and significant macrofauna abundance responses for 50% of the species examined during summer experiments. However, all of these macrofauna declined in abundance with nutrient enhancement while biomass remained constant or significantly increased with nutrient additions. This suggests a complex response of macrofauna to nutrient additions in this estuary with greater biomass per individual but a corresponding decline in abundances. Top-down/bottom-up interactive effects were observed for haustoriid amphipods, which were uncommon or absent when predators had access, but exhibited strong biomass responses to nutrient enhancement when predators were excluded. These results support a growing body of literature that indicates the importance of background conditions in regulating benthic community responses to nutrient enhancement. However, responses may be complex with biomass per individual rather than densities being the primary response variable for some taxa and predator moderation of responses occurring for some taxa but not others.     

Sediment-water fluxes of nutrients in an Antarctic coastal environment: influence of bioturbation

Rates of exchanges of nitrate and ammonium across the sediment-water interface were measured in an inshore marine environment at Signy Island, South Orkney Islands, Antarctica, over 6 months from August 1991 to February 1992. The sediment was a source of ammonium to the water column but a sink of nitrate, although nitrate exchange rates were very variable. Concentration profiles of nitrate and ammonium in the sediment porewater corroborated the measured vertical exchanges. Bioturbation, by a largely amphipod benthic infauna which was confined to the top 2 cm of sediment, was investigated experimentally. Removal of bioturbation depressed sedimentary O₂ uptake by 33% and sedimentary release of NH₄ ⁺ by 50%. In contrast, in the absence of bioturbation, the removal of NO₃ ^– from the water column by the sediment increased in rate. The measured fluxes of ammonium and nitrate from the sediment did not match with the amount of nitrogen mineralised within the sediment, and urea may account for the difference. It is suggested that the export of nitrogen from the bottom sediment may be significant in sustaining primary production in the Antarctic inshore environment. Ammonium and urea are preferred to nitrate as a nitrogen source by phytoplankton. The nitrate concentrations in the sediment porewater were low, but a large pool of nitrate was identified in the top 0–2 cm layer, which was released by KCl extraction or by freezing of the sediment. This extractable pool of nitrate did not equilibrate with the soluble nitrate pool in the sediment, but seemed to be released from components of the benthic infauna, which were also largely confined to the top 0–2 cm. The physiological role of this nitrate is unknown.     

Nitrogen cycling in coastal marine ecosystems

It is generally considered that nitrogen availability is one of the major factors regulating primary production in temperate coastal marine environments. Coastal regions often receive large anthropogenic inputs of nitrogen that cause eutrophication. The impact of these nitrogen additions has a profound effect in estuaries and coastal lagoons where water exchange is limited. Such increased nutrient loading promotes the growth of phytoplankton and fast growing pelagic macroalgae while rooted plants (sea-grasses) and benthic are suppressed due to reduced light availability. This shift from benthic to pelagic primary production introduces large diurnal variations in oxygen concentrations in the water column. In addition oxygen consumption in the surface sediments increases due to the deposition of readily degradable biomass. In this review the physico-chemical and biological factors regulating nitrogen cycling in coastal marine ecosystems are considered in relation to developing effective management programmes to rehabilitate seagrass communities in lagoons currently dominated by pelagic macroalgae and/or cyanobacteria.     

Enrichment experiments and primary production at Sagres (SW Portugal)

Water was collected from the Sagres station (SW Portugal) in September 2002, at a site adjacent to the upwelling centre of Cabo São Vicente, during relaxation of upwelling conditions. Surface and depth samples were enriched with inorganic nutrients in order to evaluate their relative influence on the microalgal assemblage. Small-scale, short-term bioassays involved separate in vitro additions of nitrogen and phosphorus. Enrichments with nitrogen led to a general increase of primary production, suggesting nitrogen as the primary potential nutrient limiting microalgal growth during this period, as well as altering the relative microplanktonic composition in favour of diatoms.     

Elevated CO2 affects porewater chemistry in a brackish marsh

As atmospheric CO₂ concentrations continue to rise and impact plant communities, concomitant shifts in belowground microbial processes are likely, but poorly understood. We measured monthly porewater concentrations of sulfate, sulfide, methane (CH₄), dissolved inorganic carbon and dissolved organic carbon over a 5-year period in a brackish marsh. Samples were collected using porewater wells (i.e., sippers) in a Schoenoplectus americanus-dominated (C₃ sedge) community, a Spartina patens-dominated (C₄ grass) community and a mixed (C₃ and C₄) community within the marsh. Plant communities were exposed to ambient and elevated (ambient + 340 ppm) CO₂ levels for 15 years prior to porewater sampling, and the treatments continued over the course of our sampling. Sulfate reduction was stimulated by elevated CO₂ in the C₃-dominated community, but not in the C₄-dominated community. Elevated CO₂ also resulted in higher porewater concentrations of CH₄ and dissolved organic carbon in the C₃-dominated system, though inhibition of CH₄ production by sulfate reduction appears to temper the porewater CH₄ response. These patterns mirror the typical divergent responses of C₃ and C₄ plants to elevated CO₂ seen in this ecosystem. Porewater concentrations of nitrogen (as ammonium) and phosphorus did not decrease despite increased plant biomass in the C₃-dominated community, suggesting nutrients do not strongly limit the sustained vegetation response to elevated CO₂. Overall, our data demonstrate that elevated CO₂ drives changes in porewater chemistry and suggest that increased plant productivity likely stimulates microbial decomposition through increases in dissolved organic carbon availability.     

Differential effects of chronic and acute simulated seawater intrusion on tidal freshwater marsh carbon cycling

Tidal freshwater ecosystems experience acute seawater intrusion associated with periodic droughts, but are expected to become chronically salinized as sea level rises. Here we report the results from an experimental manipulation in a tidal freshwater Zizaniopsis miliacea marsh on the Altamaha River, GA where diluted seawater was added to replicate marsh plots on either a press (constant) or pulse (2 months per year) basis. We measured changes in porewater chemistry (SO₄^2?, Cl^?, organic C, inorganic nitrogen and phosphorus), ecosystem CO₂ and CH₄ exchange, and microbial extracellular enzyme activity. We found that press (chronic) seawater additions increased porewater chloride and sulfate almost immediately, and ammonium and phosphate after 2–4 months. Chronic increases in salinity also decreased net ecosystem exchange, resulting in reduced CO₂ and CH₄ emissions from press plots. Our pulse treatment, designed to mimic natural salinity incursion in the Altamaha River (September and October), temporarily increased porewater ammonium concentrations but had few lasting effects on porewater chemistry or ecosystem carbon balance. Our findings suggest that long-term, chronic saltwater intrusion will lead to reduced C fixation and the potential for increased nutrient (N, P) export while acute pulses of saltwater will have temporary effects.     

Partitioning soil respiration: quantifying the artifacts of the trenching method

Sediment porewater nutrients often occur at concentrations that are orders of magnitude higher than nutrients in overlying waters, and accordingly may subsidise growth of benthic macroalgal mats in estuarine ecosystems. The relative contribution of porewater nutrients is expected to be particularly important for macroalgae entrained in intertidal mudflat sediments, where access to water column nutrients is tidally constrained. In this study, filamentous Gracilaria chilensis thalli were simultaneously exposed to sediment and overlying water nutrient sources, labelled using ¹⁵N tracers (¹⁵NH₄⁺ or ¹⁵NO₃^?) during a 5-day experiment. Dissolved inorganic N (DIN) uptake from porewater and overlying water accounted for 33 and 52%, respectively, of the N estimated as necessary to support the growth of G. chilensis, despite the two-fold lower DIN concentration of the overlying water and its periodic availability (8 h day^?1). Of the total N assimilated by the plants,?~?15% could not be accounted for, supporting the acquisition of other N forms in order to meet demand. We also found that regardless of background NH₄⁺:NO₃^? ratios (i.e. 1:3 in overlying water and 12:1 in porewater), plants accumulated ¹⁵NH₄⁺ significantly more readily than ¹⁵NO₃^?, indicating a preference for NH₄⁺. This ability to utilise multiple sources and species of N relatively rapidly may partly explain the competitive success of entrained macroalgae relative to non-entrained species and historically abundant seagrass beds in these environments. These results underscore the significance of both internal nutrient loading and external inputs as important in sustaining opportunistic macroalgal blooms in shallow estuaries.     

Detrital diversity influences estuarine ecosystem performance

Global losses of seagrasses and mangroves, eutrophication‐driven increases in ephemeral algae, and macrophyte invasions have impacted estuarine detrital resources. To understand the implications of these changes on benthic ecosystem processes, we tested the hypotheses that detrital source richness, mix identity, and biomass influence benthic primary production, metabolism, and nutrient fluxes. On an estuarine muddy sandflat, we manipulated the availability of eight detrital sources, including mangrove, seagrass, and invasive and native algal species that have undergone substantial changes in distribution. Mixes of these detrital sources were randomly assigned to one of 12 treatments and dried detrital material was added to seventy‐two 0.25 m² plots (n = 6 plots). The treatments included combinations of either two or four detrital sources and high (60 g) or low (40 g) levels of enrichments. After 2 months, the dark, light, and net uptake of NH₄⁺, dissolved inorganic nitrogen, and the dark efflux of dissolved organic nitrogen were each significantly influenced by the identity of detrital mixes, rather than detrital source richness or biomass. However, gross and net primary productivity, average oxygen flux, and net NO_X and dissolved inorganic phosphorous fluxes were significantly greater in treatments with low than with high detrital source richness. These results demonstrate that changes in detrital source richness and mix identity may be important drivers of estuarine ecosystem performance. Continued impacts to estuarine macrophytes may, therefore, further alter detritus‐fueled productivity and processes in estuaries. Specific tests that address predicted future changes to detrital resources are required to determine the consequences of this significant environmental problem.     

Contributions of benthic and planktonic primary producers to nitrate and ammonium uptake fluxes in a nutrient-poor shallow coastal area (Corsica, NW Mediterranean)

By using the stable isotope ¹⁵N, we have measured in situ the uptake of nitrate and ammonium by the seagrass Posidonia oceanica, its leaf epiphyte community, the brown macroalgae Halopteris scoparia and the suspended particulate organic matter (SPOM). In Revellata Bay (Gulf of Calvi, Western Corsica), which is a very nutrient-poor region, the specific uptake rates (V) (μg N g N⁻¹ h⁻¹) of SPOM measured at ambient concentrations are 10-1000 higher than those of benthic primary producers. Macroalgae have intermediary V, between the seagrass leaf and leaf epiphytes. V are quite variable and the reasons for this variability remain unclear.Despite the difference of specific uptake rates found between benthic and pelagic primary producers, when integrating the uptake fluxes for a water column of 10 m depth, the contribution of benthic primary producers to N uptake fluxes (g N m⁻² h⁻¹) is significant, corresponding on average to 40% of total uptake flux. This results from the dominance in terms of N biomass of benthic primary producers in this shallow nutrient-poor area. When reported for the entire volume of the Revellata Bay, the contribution of benthic primary producers is reduced to 5-10% of total N uptake flux.Although this contribution could appear relatively low, it results in a significant direct transfer of inorganic nitrogen from the water column to the benthic compartment. By this transfer, the benthic plants act as a biological pump incorporating the pelagic N into the benthic compartment for a time longer than the characteristic time of phytoplankton dynamics (month-years vs. day-week).     

Nutrient pulsing as a regulator of phytoplankton abundance and community composition in Galveston Bay, Texas

Galveston Bay, Texas, is a large shallow estuary with a watershed that includes 60% of the major industrial facilities of Texas. However, the system exhibits low to moderate (2-20 μg l⁻¹) microalgal biomass with sporadic phytoplankton blooms. Both nitrogen (N) and phosphate (P) limitation of phytoplankton growth have been proposed for the estuary. However, shifts between N and P limitation of algae growth may occur due to annual fluctuations in nutrient concentrations. The primary goal of this work was to determine the primary limiting nutrient for phytoplankton in Galveston Bay. Nutrient addition bioassays were used to assess short-term (1-2 days) phytoplankton responses (both biomass and community composition) to potentially limiting nutrients. The experimental bioassays were conducted over an annual cycle using natural water collected from the center to lower part of the estuary. Total phytoplankton biomass increased in the nitrate (10 μM) additions in 11 of the 13 bioassays, but no significant increases were detected in the phosphate (3 μM)-only additions. Bioassay results suggest that the phytoplankton community was usually not phosphate limited. All major groups increased in biomass following nitrate additions but diatoms increased in biomass at a faster rate than other groups, shifting the community composition toward higher relative abundance of diatoms. The results of this study suggest that pulsed N input events preferentially favor increases in diatom biomass in this estuary. The broader implications of this study are that N pulsing events, primarily due to river discharge, play an important role in structuring the phytoplankton community in the Galveston Bay estuary.     

Microphytobenthic Primary Production and Sedimentary Carbohydrates Along Salinity Gradients in the Lagoons of Grado and Marano (Northern Adriatic Sea)

Primary production of the microphytobenthic community and carbohydrates concentrations were studied in the lagoonal system of Grado and Marano, located in the Northern Adriatic coast. Sediment samples were collected along a salinity gradient. Abundance and species composition of the microphytobenthic communities were analysed and the benthic microalgal biomass was estimated as Chlorophyll a (Chl a). Primary production of benthic diatoms was estimated using ¹⁴C-tracer. Extracellular carbohydrates were extracted from the sediment and separated in two operationally defined fractions (colloidal and EDTA-extractable). Salinity was higher in the Grado lagoon, where the benthic microalgal community was mainly composed of marine diatoms. In the Marano lagoon, which has a lower salinity, freshwater species were also found. In both lagoons, photosynthetic efficiency showed an inverse relationship with salinity and a direct relationship with the main biological variables. Photosynthetic activity was directly related to Chl a and abundance of benthic microalgae, suggesting that in the benthic system microalgal community is responsible for primary production. Overall, salinity was also influent on the microphytobenthic primary production, which was greater in the more saline Grado lagoon.     

Mediation of benthic–pelagic coupling by microphytobenthos: an energy- and material-based model for initiation of blooms of Aureococcus anophagefferens

We present a conceptual model for initiation of blooms of the estuarine brown-tide pelagophyte Aureococcus anophagefferens. The model is based on the observation that in addition to its well-documented stimulation by organic nutrients, Aureococcus is pre-adapted to low light levels. Its relatively low maximum (light-saturated) growth rate makes it a poor competitor with other estuarine species at high light under acclimated conditions. Its large photosynthetic antenna and relatively low quota of photoprotective pigments make it more susceptible to photoinhibition than other species to which it is compared. These same characteristics give it a competitive advantage at low light levels. In its shallow habitat, both the light level and the rate of nutrient supply from groundwater and benthic porewater are determined by the degree of benthic coupling. Experimental manipulations in a microcosm and a survey of the literature demonstrate the ability of the sediment-associated microphytobenthos (MPB) to regulate both the light- and nutrient-environment in the overlying water column. The model predicts that the growth dynamics of the MPB are such that the benthic/water column interactions tend towards one of two stable states. In one, a well-developed population of MPB restricts resuspension of particulate material and efflux of dissolved nutrients, resulting in clear and nutrient-poor overlying waters. This condition does not favor growth of Aureococcus. In the alternative state, erosion of the MPB results in turbid, nutrient-rich waters that do favor bloom initiation. Alternation between the states is caused by external physical forcing, through wind-driven mixing of the water column. Field data from Quantuck Bay, New York (USA), failed to document the transition from non-bloom to bloom conditions. Even so, they are consistent with the model’s predictions.     

Coral reefs modify their seawater carbon chemistry – case study from a barrier reef (Moorea,French Polynesia)

Changes in the carbonate chemistry of coral reef waters are driven by carbon fluxes from two sources: concentrations of CO₂ in the atmospheric and source water, and the primary production/respiration and calcification/dissolution of the benthic community. Recent model analyses have shown that, depending on the composition of the reef community, the air‐sea flux of CO₂ driven by benthic community processes can exceed that due to increases in atmospheric CO₂ (ocean acidification). We field test this model and examine the role of three key members of benthic reef communities in modifying the chemistry of the ocean source water: corals, macroalgae, and sand. Building on data from previous carbon flux studies along a reef‐flat transect in Moorea (French Polynesia), we illustrate that the drawdown of total dissolved inorganic carbon (C_T) due to photosynthesis and calcification of reef communities can exceed the draw down of total alkalinity (A_T) due to calcification of corals and calcifying algae, leading to a net increase in aragonite saturation state (Ω_a). We use the model to test how changes in atmospheric CO₂ forcing and benthic community structure affect the overall calcification rates on the reef flat. Results show that between the preindustrial period and 1992, ocean acidification caused reef flat calcification rates to decline by an estimated 15%, but loss of coral cover caused calcification rates to decline by at least three times that amount. The results also show that the upstream–downstream patterns of carbonate chemistry were affected by the spatial patterns of benthic community structure. Changes in the ratio of photosynthesis to calcification can thus partially compensate for ocean acidification, at least on shallow reef flats. With no change in benthic community structure, however, ocean acidification depressed net calcification of the reef flat consistent with findings of previous studies.     

Seasonal changes in eelgrass functions: current velocity reduction,prevention of sediment resuspension,and control of sediment–water column nutrient flux in relation to eelgrass dynamics

We assessed seasonal changes in eelgrass (Zostera marina) functions, i.e., reduction of current velocity, buffering of sediment resuspension, and control of dissolved inorganic nitrogen flux between the sediments and the water column, using field observations and experiments in the Akkeshi-ko estuary, Hokkaido, Japan. We also analyzed the relationships between eelgrass traits and functions. The efficiency of the reduction in current velocity increased with the development of the eelgrass canopy. Sediment resuspension was inhibited from May to August, during which time the eelgrass canopy developed. Eelgrass controlled the NH₄ ⁺ concentration of sediment porewater through root nutrient uptake, affecting NH₄ ⁺ flux between the sediments and the water column. Fluctuations in eelgrass functions and coincident changes in dynamics resulted in seasonal changes in the eelgrass environment, which may in turn affect the dynamics of organisms inhabiting eelgrass beds, e.g., mysids and epiphytic algae. Moreover, the developed eelgrass canopy trapped a large amount of material during spring and summer, which was resuspended into the surrounding ecosystem in autumn when the canopy and its functions (i.e., reduction of current and sediment resuspension) diminished. These results suggest that seasonal changes in eelgrass functions also affect communities within marginal coastal ecosystems through the control of allochthonous resources. Handling editor: P. Viaroli     

Quantifying the trophic base for benthic secondary production in the Nakdong River estuary of Korea using stable C and N isotopes

The overall dependence of benthic secondary production on the main primary producers at three different habitats in the Nakdong River estuarine system, Korea, was estimated. Inventories of macrobenthic invertebrate biomass were combined with multiple-isotope-mixing models to evaluate the trophic base, comparing Scirpus triqueter-dominated and Phragmites australis-dominated marshes and bare intertidal flat. The feasible contributions of four main food sources, marsh macrophytes, the microphytobenthos, and riverine and marine suspended particulate organic matter (RPOM and MPOM), to the consumer biomasses were calculated using the isotopic mixing model. After weighting the feasible contributions of food sources to each taxon by the consumer biomass, the resultant values were summed for all the consumers at each habitat to quantify the trophic base of the benthic invertebrate community. Dual-isotope-mixing model calculations verified the varying dependence on those potential food sources among the functional feeding groups. In addition, the dependence on each source of the same functional group varied between bare intertidal and salt-marsh habitats, shifting from a dominance of benthic and pelagic microalgal sources on the former habitat to a mixed food source at the latter habitat. The biomasses of the species comprising each functional group differed among habitats and sampling dates, so that each functional group made a different contribution to the whole benthic community and its basal food source. Given the calculation of the overall dependence of macrozoobenthic community on each food source, our results indicate that the microphytobenthic source dominates (nearly half) the trophic base in all the intertidal habitats of different vegetational compositions. Marsh-macrophyte-derived organic matter and RPOM served as considerable subsidies only to salt-marsh food webs, reflecting the use of the mixed food source by salt-marsh-bed consumers. Conversely, the dominance of MPOM in the total food base was equal to that of the microphytobenthos in the bare intertidal ecosystem but increased during spring−summer in the salt-marsh systems. Our results also suggest that the river discharge concentrated during the summer monsoon does not lead to any shift in trophic base for estuarine secondary production.     

Predation regulation of sedimentary faunal structure: potential effects of a fishery-induced switch in predators in a Newfoundland sub-Arctic fjord

The collapse of the cod fishery in Newfoundland has coincided with marked increases in abundances of snow crab, pandalid shrimp, and other crustaceans that prey on sedimentary infauna. A 3-year sampling program in Bonne Bay, Newfoundland indicates differences in composition and number of these predators in the two main arms of the fjord that coincide with strong differences in benthic community structure. To test whether predation pressure contributes to the observed patterns in sedimentary fauna, exclusion field experiments with full and partial cages were deployed in both arms at 30-m depth and sampled along with ambient sediments at 0-, 4-, and 8-week periods. Predation significantly influenced species composition, abundance and, in some cases, diversity. The most striking changes included increases in the polychaetes Phöloe tecta and Ophelina cylindricaudata in exclusions relative to controls, and concurrent declines in the polychaete Paradoneis lyra and the cumacean Lamphros fuscata. In laboratory experiments, fresh non-disturbed sediment cores from each experimental area were either protected or exposed to snow crab, the most abundant predator in the bay. A snow crab inclusion experiment was also carried out in the field, using cages similar to those used for exclusions. Despite differences in sedimentary faunas in the two arms, both types of experiments detected a predator effect that was very similar to that documented in exclusion experiments. Thus, despite differences in the scales associated with each type of manipulation, our results suggest that crab predation is a significant structuring force in Newfoundland sedimentary communities. Given the historical changes that have occurred in predator composition as a result of cod over-fishing, we hypothesize that broad-scale community changes may be taking place in North Atlantic benthic ecosystems.     

Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.