Pierluigi Viaroli

Impact of Tapes philippinarum farming on nutrient dynamics and benthic respiration in the Sacca di Goro

AbstractThe introduction of the short-necked clam Tapes philippinarum into the Sacca di Goro has over a short period made this coastal environment one of the top European clam production sites. In recent years, this activity has been seriously impacted due to the appearance in the lagoon of large macroalgal beds and the occurrence of dystrophic events causing anoxia and massive deaths of molluscs in the cultivated areas. Tapes cultivation sites now cover more than one third of the lagoon surface at densities sometimes attaining 2000–2500 adult individuals m−2; such densities and the harvesting methods, based on sediment dredging, probably have a strong impact on the benthic system. Whilst a number of studies have reported water–sediment interface induced modifications due to oyster or mussel farming there have been few attempts to quantify how clam farming affects biogeochemical cycles of oxygen and nutrients, in particular in the Sacca di Goro. Two areas, a farmed and a control one, were compared for benthic fluxes and results were correlated with clam biomass. Oxygen, carbon dioxide, ammonium, reactive silica and phosphorus fluxes were stimulated several fold by the presence of Tapes due to the clams, respiration and excretion activities, but also to the reducing conditions in the surface sediments. On average, the whole lagoon dark sediment O2 demand and CO2 production were stimulated by a factor of, respectively, 1.8 and 3.3, whilst nutrient release was 6.5 times higher for NH4+ and 4.6 times higher for PO43-. Our results indicate that clam farmers should carefully consider sustainable densities of Tapes in order to prevent the risk of sediment and water anoxia. Rapid nutrient recycling (up to 4000 μmol NH4+ m−2 h−1 and 150 μmol PO43- m−2 h−1) stimulated by the high biodegradability of clam faeces and pseudofaeces could in turn favour macroalgal growth.

Growth of the seaweed Ulva rigida C. Agardh in relation to biomass densities, internal nutrient pools and external nutrient supply in the Sacca di Goro lagoon (Northern Italy)

Growth of the seaweed Ulva rigida C. Agardh was investigated in relation to biomass densities, internal nutrient pools and external nutrient supply. Research was carried out from 23 March to 5 July 1994 in the Sacca di Goro (Po Delta, Northern Italy), whose south-eastern part was covered by extensive mats of Ulva rigida. Two types of field experiments were conducted by incubating Ulva thalli inside large cages. In the first experiment, beginning on 23 March, 100 g of wet thalli were placed into the cages, allowed to grow for two weeks, then collected and replaced. This procedure was repeated 8 times over the study period. In the second experiment, Ulva thalli were left inside the cages and collected at selected time intervals (14, 27, 41, 64 and 76 days) in order to simulate the effects of increased density on growth and nutrient storage.We recorded specific growth rates (NGR) ranging from 0.025 to 0.081 d−1 for a period up to two months in the repeated short-term experiments performed at relatively low initial algal densities (300–500 g AFDW m−3). These NGR resulted significantly related to dissolved inorganic nitrogen (DIN) in the water column. Tissue concentrations of total Kjeldahl nitrogen (TN) were almost constant, while extractable nitrate decreased in a similar manner to DIN in the water column. Total phosphorus showed considerable variation, probably linked to pulsed freshwater inflow.In the long-term incubation experiment, NGR of Ulva was inversely related to density. Internal concentrations of both total P and TN reached maximum values after one month; thereafter P concentration remained almost constant, while TN decreased below 2% w/w (by dry weight). The TN decrease was also accompanied by an abrupt decrease in nitrate tissue concentration. The biomass incubated over the two month period suffered a progressive N limitation as shown by a decreasing NY ratio (49.4 to 14.6). The reciprocal control of Ulva against biogeochemical environment and viceversa is a key factor in explaining both resource competition and successional stages in primary producer communities dominated by Ulva. However, when the biomass exceeds a critical threshold level, approximately 1 kg AFDW m−3, the macroalgal community switches from active production to rapid decomposition, probably as a result of selfshading, biomass density and development of anaerobic conditions within the macroalgal beds.

Nitrate uptake and storage in the seaweed Ulva rigida C. Agardh in relation to nitrate availability and thallus nitrate content in a eutrophic coastal lagoon (Sacca di Goro, Po River Delta, Italy)

The seasonal cycle of biomass and tissue composition of Ulva rigida C. Agardh, in relation to nitrogen availability in the water column, was studied in 1991–1992 in the Sacca di Goro, a highly eutrophic lagoon in the Po River Delta (Italy). Nitrate uptake rates and storage capacity were also determined in laboratory experiments. The seasonal growth of U. rigida was related to the seasonal trend of nitrogen concentration in the water column. U. rigida biomass increased exponentially during spring and attained peaks of about 300–400 g dry mass (DM) m−2 in June. As biomass increased, U. rigida depleted nitrate in the water column. Thallus nitrate reserves also declined from 100 μmol N (g DM)−1 to almost undetectable levels, and total thallus nitrogen declined from 4% to 2.5% DM and 1.25% DM in 1991 and 1992, respectively. During summer, U. rigida decomposition increased, and organic nitrogen concentrations in the water column increased. The uptake experiments demonstrated an inverse relationship between thallus nitrate content and nitrate uptake rates. A modified Michaelis–Menten equation that accounts for thallus nitrate fit the uptake data well. U. rigida can accumulate up to about 400–500 μmol nitrate (g DM)−1 in cellular reserves. U. rigida in the Sacca di Goro has higher Km and lower Vmax/Km ratios for nitrate uptake than other chlorophycean species, indicating a low efficiency of uptake at low nitrate concentrations. This low uptake efficiency, and the ability to exploit N availability by storing cellular nitrate pools in excess of immediate growth needs, may represent a physiological response to an eutrophic environment where nitrate is in large supply for most of the year.

Nitrogen cycling networks of coastal ecosystems: influence of trophic status and primary producer form

Abstract We have used ecological network analysis to compare nitrogen cycles from five well-researched coastal ecosystems. These included a representative ricefield and two lagoons (Tancada and Encanysada lagoons) in the Ebro River delta, Spain; a region of the Sacca di Goro, a lagoon at the mouth of the Po River, Italy; and a drowned river estuary in North Carolina, USA, the Neuse River estuary. We constructed networks for the various systems and ranked them by trophic status (i.e., degree of eutrophication) using four indices. We then considered the importance of (1) trophic status, (2) growth form of dominant primary producer and (3) water residence time to the intensity and pattern of recycling and to the manner in which the systems can “filter” N. Three indices of flux (rate of import, primary producivity and total systems throughput) gave similar rankings of trophic status among ecosystems with the Italian and U.S. systems being most eutrophic, ricefields next, and then the two Spanish lagoons. Patterns of N export and of cycling within the systems were most closely related to the growth form of dominant primary producers. Phytoplankton, with their rapid turnover rate, foster rapid recycling within the water column and continuous transfer to sediments and export. Submersed and emergent aquatic vegetation and macroalgae create lags and pulses within systems by sequestering N during growth and releasing it during senescence, death and decomposition. Trends in cycling among systems relative to trophic status or water residence appear largely secondary to primary producer growth form.

Nutrient and iron limitation to Ulva blooms in a eutrophic coastal lagoon (Sacca di Goro, Italy)

Growth patterns and bloom formation of the green seaweed Ulva rigida were analysed in the eutrophic Sacca di Goro lagoon (Po River Delta, Italy). Variations of standing biomasses and elemental composition of Ulva were analysed through an annual cycle with respect to nitrogen, phosphorus and iron. Growth rates, nutrient and iron uptake and nitrate storage by macroalgal thalli were also assessed with field experiments during the formation of a spring bloom. The control of Ulva growth and the bloom formation depended on multiple factors, especially on nitrogen availability and iron deficiency. In the nitrate rich waters of the Sacca di Goro lagoon, nitrate accumulation in Ulva thalli was inversely related with Fe uptake, indicating an influence of Fe limitation on N acquisition. Since length and magnitude of nitrate luxury uptake are inversely related to the size of the intracellular nitrate pools, in nitrate rich waters the fast growing Ulva may face risk of N-limitation not only when exposed to low N concentrations or at high biomass levels, but also when exposed to pulsed dissolved nitrate concentrations at low iron availability. The potential Fe limitation could be affected by processes controlled by geochemical reactions and by macroalgal growth and decomposition. Both Fe oxidation during the active macroalgal growth and the formation of insoluble FeS and FeS2 during bloom collapse can result in a drastic decrease of soluble iron. Thus, a potential limitation of Fe to macroalgae can occur, determining positive feedbacks and potentially controlling the extent of bloom development and persistence.

Microphytobenthos activity and fluxes at the sediment-water interface: interactions and spatial variability

In this study oxygen and nutrient fluxes and denitrification rates across the sediment-water interface were measured via intact core incubations with a twofold aim: show whether microphytobenthos activity affects these processes and analyse the dispersion of replicate measurements. Eighteen intact sediment cores (i.d. 8 cm) were randomly sampled from a shallow microtidal brackish pond at Tjarno, on the west coast of Sweden, and were incubated in light and in darkness simulating in situ conditions. During incubation O2, inorganic N and SiO2 fluxes and denitrification rates (isotope pairing) were measured. Assuming mean values of 18 cores as best estimate of true average (BEA), the accuracy of O2, NH4+, NO3- and SiO2 fluxes calculated for an increasing number of subsamples was tested. At the investigated site, microalgae strongly influenced benthic O2, inorganic N and SiO2 fluxes and coupled (Dn) and uncoupled (Dw) denitrification through their photosynthetic activity. In the shift between dark and light conditions NH4+ and SiO2 effluxes (60 and 110 µmol m-2h-1) and Dn (5 µmol m-2 h-1) dropped to zero, NO3- uptake (70 µmol m-2 h-1) showed a 30% increase, while Dw (20 µmol m-2 h-1) showed an 80% decrease. For O2 and NO3- dark fluxes, 4 core replicates were sufficient to obtain averages within 5-10% of the best estimated mean, while 10-20% accuracy was obtained with 4-12 replicates for SiO2 and >10 replicates for NH4+ dark fluxes. Mean accuracy was considerably lower for all light incubations, probably due to the patchy distribution of the benthic microalgal community.

Animal-sediment relationships: Evaluating the 'Pearson-Rosenberg paradigm' in Mediterranean coastal lagoons

We investigated the applicability of the Pearson-Rosenberg (P-R) conceptual model describing a generalized pattern of response of benthic communities in relation to organic enrichment to Mediterranean Sea coastal lagoons. Consistent with P-R model predictions, benthic diversity and abundance showed two different peaks at low (>2.5-5 mg g(-1)) and high (>25-30 mg g(-1)) total organic carbon (TOC) ranges, respectively. We identified TOC thresholds indicating that risks of reduced benthic diversity should be relatively low at TOC valuesabout 28 mg g(-1), and intermediate at values in-between. Predictive ability within these ranges was high based on results of re-sampling simulation. While not a direct measure of causality, it is anticipated that these TOC thresholds should serve as a general screening-level indicator for evaluating the likelihood of reduced sediment quality and associated bioeffects in such eutrophic systems of the Mediterranean Sea.

Short-term influence of recolonisation by the polycheate worm [2pt] Nereis succinea on oxygen and nitrogen fluxes and [2pt] denitrification: a microcosm simulation

The short-term effects of sediment recolonisation by Nereis succinea on sediment-water column fluxes of oxygen and dissolved inorganic nitrogen, and rates of denitrification, were studied in microcosms of homogenised, sieved sediments. The added worms enhanced oxygen uptake by the sediments, due to the increased surface area provided by the burrow walls and the degree of stimulation was stable with time. Similarly, ammonium fluxes to the water column were stimulated by N. succinea, but declined over the 3 day incubation in all microcosms including the controls. Nitrate fluxes were generally greater in the faunated microcosms, but highly variable with time. Denitrification rates were positively stimulated by N. succinea populations, denitrification of water column nitrate was stimulated 10-fold in comparison to denitrification coupled to nitrification in the sediments. Rates of denitrification of water column nitrate were not significantly different from rates in undisturbed sediment cores with similar densities of N. succinea, whereas rates of coupled nitrification–denitrification were 3-fold lower in the experimental set-up. These results may reflect the relative growth rates of nitrifying and denitrifying bacteria, which allow more rapid colonisation of new burrow surfaces by denitrifier compared to nitrifier populations. The data indicate that recolonisation by burrowing macrofauna of the highly reduced sediments of the Sacca di Goro, Lagoon, Italy, following the annual dystrophic crisis, may play a significant role in the reoxidation and detoxification of the sediments. The increased rates of denitrification associated with the worm burrows, may promote nitrogen losses, but due to the low capacity of nitrifying bacteria to colonise the new burrow structures, these losses would be highly dependent upon water column nitrate concentrations.

Seasonal variations of sulphate reduction rates sulphur pools and iron availability in the sediment of a dystrophic lagoon (Sacca di Goro, Italy)

The aim of this work was to analyse factors which regulate sulphide mobility in the sediments of a dystrophic lagoon (Sacca di Goro, the southern lagoon of the Po river delta, Italy). In 1995–96, sediment oxygen demand and variations in organic matter content, redox potential, iron availability, inorganic sulphur concentrations and sulphate reduction rates were measured in sediment profiles at three stations (G, 4 and 17) representative of the main areas of the lagoon. Stations differed mainly in their salinity range and primary producer communities. High concentrations of reactive iron (110-275 pmol ml-1) and low sulphate reduction rates (0.8–16.1 mmol m−2 d−1) were measured in the sediment. Moreover, high concentrations of reactive ferric iron were detected in winter and spring at the stations closest to the freshwater inputs. Nevertheless, in summer, high concentrations of free sulphides were detected in the porewater, although most of the reactive ferrous iron was not sulphide-bound, indicating that not all of the reactive iron pool was available to buffer against sulphide release and thus measures of reactive iron pools may not be a good measure of the true buffering capacity of the sediment. Furthermore, a considerable production of sulphide may occur in the decayingUlva biomass in the water column, where its concentration will be independent of the potential buffering capacity of iron in the sediment. Therefore when assessing the vulnerability of coastal lagoons to dystrophic events, both the size and availability of the reactive iron pool as well as the site of sulphide production must be taken into consideration.

Physical factors and dissolved reactive silica affect phytoplankton community structure and dynamics in a lowland eutrophic river (Po river, Italy)

We tested the hypothesis that species composition and persistence of phytoplankton communities in nutrient rich lowland rivers depends mainly on physical factors. The study aimed to analyse the effects of water discharge, temperature and chemistry on phytoplankton dynamic and species composition in the lowland reach of the eutrophic Po river (Italy). Both taxonomical and morpho-functional methods were used. True planktonic and tychoplanktic (i.e. detached taxa of benthic origin that remain in suspension) species were found, among which only a few taxa and functional groups prevailed. Diatoms were the most abundant, with a clear dominance of species either sensitive to the onset of water stratification or well adapted to turbid waters. Phytoplankton abundance, biomass and chlorophyll-a followed similar trends, attaining the highest values in summer, at low discharge rates. Correlation and multivariate analysis revealed that the development of a stable phytoplankton community was mainly controlled by water discharge rates. Namely, changes in water flow rates induced major variations in the community structure. The seasonal succession of phytoplankton assemblages was also related to water temperature and dissolved reactive silica availability to some extent overlapping flow effects.