Can we predict nutrient limitation in streams and rivers?

1. Anthropogenic impacts on the biogeochemical cycles of nitrogen (N) and phosphorus (P) affect natural ecosystems worldwide. Modelling is required to predict where and when these key nutrients limit primary production in freshwaters. 2. We reviewed 382 nutrient‐enrichment experiments to examine which factors promote limitation of microphytobenthos biomass by N or P in streams and rivers. Using regression models, we examined whether the response of microphytobenthos biomass to N and P additions could be predicted by the absolute N and P concentrations in the water, the water N:P ratio or a combination of the two. 3. The absolute N concentration in the water was the best predictor of the magnitude of the response of microphytobenthos biomass to N additions. In comparison, the N:P ratio was the best predictor of whether or not N was limiting. However, predictions were uncertain except at extreme N:P ratios <1 : 1 and >100 : 1. 4. The absolute P concentration in the water was the best predictor of the magnitude of the response of microphytobenthos biomass to P additions. Neither the absolute nor the relative N and P concentrations predicted whether or not P was limiting. 5. The absolute and the relative N and P water concentrations contribute significant and complementary insights into the responses of microphytobenthos biomass to nutrient enrichment in running waters. However, ability to predict nutrient limitation from these concentrations is constrained by substantial error in the models. In the future, the prediction ability of models of nutrient limitation might be improved by focussing on regional scales and accounting for additional factors such as light and disturbance.     

Seasonal variation in primary production of microphytobenthos at the Isshiki intertidal flat in Mikawa Bay

Seasonal variations in photosynthetic rates by microphytobenthos and phytoplankton at the Isshiki tidal flat in Mikawa Bay were measured with a ¹⁴C combustion method. In addition, diurnal variations in the photosynthetic rate and photosynthesis versus irradiance (P-I) curves were obtained through in situ incubation. The photosynthetic rate of microphytobenthos (annual average, 13.9 ± 6.4 mg C m⁻² h⁻¹) did not show a remarkable change, and they maintained a higher production rate than phytoplankton (annual average 9.0 ± 5.1 mg C m⁻² h⁻¹) throughout the year. The P-I curves from in situ experiments showed that the photosynthetic activity of microphytobenthos at the laboratory irradiance (250 μE m⁻² s⁻¹) was 56% of that at the maximum irradiance (1200 μE m⁻² s⁻¹) in situ. In the in situ experiments, the chlorophyll a concentration, photosynthetic rate, and activity of microphytobenthos varied greatly throughout the day, influenced by tidal submersion/emersion and daylight. From an analysis of these results, it is considered that microphytobenthos contributed greatly to primary production in this ecosystem throughout the year by adapting suitably to intertidal environments. Received: July 28, 1999 / Accepted: October 10, 1999     

Estimating stable carbon isotope values of microphytobenthos in the Arctic for application to food web studies

Most studies on Arctic food webs have neglected microphytobenthos as a potential food source because we currently lack robust measurements of δ¹³C values for microphytobenthos from this environment. As a result, the role of microphytobenthos in high latitude marine food webs is not well understood. We combined field measurements of the concentration of aqueous carbon dioxide and the stable carbon isotopic composition of dissolved inorganic carbon (δ¹³C_DIC) from bottom water in the Beaufort and Chukchi seas with a set of stable carbon isotopic fractionation factors reflecting differences in algal taxonomy and physiology to estimate the stable carbon isotope composition of microphytobenthos-derived total organic carbon (δ¹³C_p). The δ¹³C_p for Phaeodactylum tricornutum, a pennate diatom likely to be a dominant microphytobenthos taxon, was estimated to be ?23.9 ± 0.4 ‰ as compared to a centric diatom (Porosira glacialis, δ¹³C_p = ?20.0 ± 1.6 ‰) and a marine haptophyte (Emiliana huxleyi, δ¹³C_p = ?22.7 ± 0.5 ‰) at a growth rate (µ) of 0.1 divisions per day (d^?1). δ¹³C_p values increased by ~2.5 ‰ when µ increased from 0.1 to a maximum growth rate of 1.4 d^?1. We compared our estimates of δ¹³C_p values for microphytobenthos with published measurements for other carbon sources in the Arctic and sub-Arctic. We found that microphytobenthos values overlapped with pelagic sources, yet differed from riverine and ice-derived carbon sources. These model results provide valuable insight into the range of possible isotopic values for microphytobenthos from this region, but we remain cautious in regard to the conclusiveness of these findings given the paucity of field measurements currently available for model validation.     

Benthic Diatoms of the Russian Waters of the Sea of Japan and Adjacent Sea Areas

Russian Journal of Marine Biology - This review analyzes the current state of knowledge of the microphytobenthos in the Russian waters of the Sea of Japan and adjacent sea areas. A retrospective...     

Reevaluation of the nutrient mineralization process by infaunal bivalves (Ruditapes philippinarum) in a shallow lagoon in Hokkaido, Japan

Previous estimations of nutrient mineralization in the water column by infaunal bivalves might have been overestimated because of underestimation of the uptake process by microphytobenthos in the field. We conducted field surveys of environmental conditions and quantitative sampling of Ruditapes philippinarum in a shallow lagoon system (Hichirippu Lagoon, eastern Hokkaido, Japan) in August 2006. We recorded the spatial distribution pattern and the molar ratio of dissolved inorganic nutrients to determine the limiting nutrients for microphytobenthos, to evaluate the input and output of nutrients at the entrance of the lagoon station, and to estimate potential nutrient mineralization by R. philippinarum. Our aim was to reevaluate the nutrient mineralization process by infaunal bivalve species. In this study, the mean standing stock of microphytobenthos inhabiting surface sediment (5 mm thick) on the tidal flats was 100 times higher than that of phytoplankton (1 m depth). Low N/P and high Si/N ratios (mean = 2.6 and 17.6, respectively) near the entrance of the lagoon compared to those of microphytobenthos (N:P:Si = 10.1:1:18) clearly suggest N deficiency. The flux of NH₄-N coming into the lagoon was 3.4 kmolN d^− 1, and the flux out was − 3.7 kmolN d^− 1. Thus, assuming that there would have been no phytoplankton and microphytobenthos uptake during the day, 0.3 kmolN d^− 1 of NH₄-N was produced within the lagoon. However, the NH₄-N mineralization rate of the clams has been estimated to be approximately 7.7 ± 6.8 kmolN d^− 1. Thus, 96% (7.4 kmolN d^− 1, i.e., 7.7 kmolN d^− 1 minus 0.3 kmolN d^− 1) of the NH₄-N mineralized by the clam was consumed by microphytobenthos. In contrast, if all the NH₄-N inflow (3.1 kmolN d^− 1) was consumed by the microalgae before outflow, 52% (4.0 kmolN d^− 1, i.e., 7.7 kmolN d^− 1 minus 3.7 kmolN d^− 1) of the NH₄-N mineralized by the clams should have been consumed by microphytobenthos. Microphytobenthos on the tidal flats (11.3 ± 11.8 kmolN) used all of the surplus nutrients (between 4.0 and 7.4 kmolN d^− 1), and the temporal division rate [=(NH₄-N uptake)/(standing stock of microphytobenthos)] of microphytobenthos would have to be between 0.35 and 0.65 d^− 1. Residual NH₄-N (0.3 - 3.7 kmolN d^− 1) was the water-column source and accounted for 12-148% of NH₄-N in the water column near the entrance of the lagoon (2.5 ± 1.4 kmolN) per day. This is the first field-based observation with a quantitative evaluation of nutrient mineralization by infaunal bivalves and nutrient uptake by microphytobenthos.     

Biodegradation of photosynthetically produced extracellular organic carbon from intertidal benthic algae

14C-labeled extracellular products of a natural microphytobenthic community and two species of benthic diatoms (Nitzschia hybridaeformis and Amphora coffeaeformis) were fractionated into extracellular dissolved organic carbon (14C-EDOC), organic carbon extracted with EDTA (14C-EDTA-extractable OC) and extracellular polymeric substances (14C-EPS). The biodegradation of this labeled extracellular organic carbon by bacteria in sediments was examined to determine the processes of enzymatic degradation of photosynthetically-produced extracellular organic carbon from microphytobenthos in an intertidal flat ecosystem. In addition, primary production as well as extracellular enzyme activities (beta- and alpha-glucosidase) were measured to evaluate the possible relationship between organic carbon production and microbiological degradation at the Isshiki intertidal flat in Mikawa Bay, Japan. With all three 14C-fractions extracted from a natural microphytobenthic assemblage and two species of benthic diatoms, more than 50% of the added substrates were mineralized within 24 h by the bacterial community in sediments. At that time, the percentage of high-molecular-weight compounds (>5 K MW) to total MW compounds of 14C-EDTA-extractable OC and 14C-EPS fractions decreased within 24 h from 50.9 to 6.6% and 74.5 to 11.1%, respectively. In situ, beta- and alpha-glucosidase activity in sediment was higher than in the seawater column (at a depth of 1 m), though the photosynthetic production of microphytobenthos was equal to that of phytoplankton. Based on our previous studies that microphytobenthos produced much more extracellular products than phytoplankton, it is assumed from these results that carbon flowing into the microbial loop through the mediation of enzymatic degradation of extracellular products in a benthic system exceeds that in the overlying water column.     

Are epiphytes a significant component of intertidal Zostera noltii beds?

The role of epiphytes in an intertidal Zostera noltii seagrass bed in Marennes-Oléron Bay was assessed in comparison with the other main benthic primary producers (Z. noltii, microphytobenthos) at two bathymetric levels and on a seasonal basis. Assemblage and biomass of epiphytes were studied using scanning electron microscopy (SEM). Z. noltii and its detrital matter followed a typical seasonal pattern: microphytobenthos was present in large quantities throughout the year representing 21% of the total biomass while detrital matter, above-ground parts and below-ground parts accounted for 65, 9 and 5%, respectively. Only two species of epiphytic diatoms, Cocconeis scutellum and Cocconeis placentula, were observed on seagrass leaves. Epiphyte biomass was very low, representing on average less than 0.001% of that of microphytobenthos or leaves. This low epiphyte biomass is linked with the absence of macroalgae and also with the low biovolume of Cocconeis, which formed a monolayer of cells on leaves. This can be explained by the severe conditions of the intertidal and the high leaf turn-over of Z. noltii leaves.     

The response of planktonic and microbenthic algal assemblages to nutrient enrichment in shallow coastal waters,southwest Sweden

Field and laboratory nutrient (nitrogen and phosphorus) enrichment experiments were performed using natural phytoplankton and microphytobenthic assemblages from the brackish water Öresund, S.W. Sweden. The response of algae from a low-nutrient area (Falsterbo Canal) was compared to that of algae from a polluted, nutrient-rich area (Lomma Bay).The biomass (measured as chlorophyll a) of both phytoplankton and microphytobenthos from the Falsterbo Canal increased after the addition of nitrogen. Phytoplankton growth was stimulated by the addition of phosphorus to the nitrogen-rich water of the polluted Lomma Bay. Sediment chlorophyll a showed no significant increase after the addition of nutrients in the Lomma Bay. In containers without sediment, phytoplankton uptake was calculated to account for ≈ 90% of the disappearance of inorganic fixed nitrogen from the water. In the sediment containers the microphytobenthos was estimated to account for ≈20% of the nitrogen uptake. The rest was presumably lost mainly through denitrification.When containers with microphytobenthos from Lomma Bay were kept in the dark, phosphorus was released at a rate of up to ≈ 180 μM · m^?2 · day^?1. We suggest that by producing oxygen microbenthic algae keep the sediment surface oxygenated thereby decreasing phosphorus transport from the sediment to the overlying water.     

Species identity, diversity and microbial carbon flow in reassembling macrobenthic communities

Intense disturbance may locally destroy patches of habitat and shape the landscape into a mosaic of reassembling communities. The development of ecosystem properties during such community reassembly is poorly understood. In intertidal bare sediments, trophic relations between microphytobenthos or heterotrophic bacteria and macrofauna invertebrates may guarantee fundamental ecosystem properties such as carbon flow through the food web. We studied the dynamic relation between reassembling macrofauna communities and such microbial carbon flow during recovery after severe disturbance. We deliberately induced prolonged hypoxia in winter and early summer and allowed recolonisation for periods of two and five months. Carbon flow was quantified from basal resources (microphytobenthos and bacteria) to intermediate consumers using ¹³C as a tracer. Within the period of study (5 months), microbial carbon flow fully recovered, although macrofauna diversity was still very low compared to the natural communities (ranging from 6 to 17 species). More than 90% of microbial carbon flow to macrofauna was due to the consumers that recolonised within two months. Two of these species were dominant contributors to microphytobenthos carbon transfer to fauna. Furthermore, at an early stage of reassembly, this ecosystem property was remarkably similar when disturbance took place at different times of the year (winter or early summer), although there were differences in assemblage composition and functional diversity. We conclude that species assemblages and ecosystem function developed relatively independently in this benthic system. We discuss which ecological factors may have caused such non-parallel development of macrofaunal communities and carbon flow.     

Trophic cul-de-sac, Pyrazus ebeninus, limits trophic transfer through an estuarine detritus-based food web

The importance to food‐webs of trophic cul‐de‐sacs, species that channel energy flow away from higher trophic levels, is seldom considered outside of the pelagic systems in which they were first identified. On intertidal mudflats, inputs of detritus from saltmarshes, macroalgae or microphytobenthos are generally regarded as a major structuring force underpinning food‐webs and there has been no consideration of trophic cul‐de‐sacs to date. A fully orthogonal three‐factor experiment manipulating the density of the abundant gastropod, Pyrazus ebeninus, detritus and macrobenthic predators on a Sydney mudflat revealed large deleterious effects of the gastropod, irrespective of detrital loading or the presence of predators. Two months after experimental manipulation, the standing‐stock of microphytobenthos in plots with high (44 per m²) densities of P. ebeninus was 20% less than in plots with low (4 per m²) densities. Increasing densities of P. ebeninus from low to high halved the abundance of macroinvertebrates and the average number of species. In contrast, the addition of detritus had differing effects on microphytobenthos (positively affected) and macroinvertebrates (negatively affected). Over the two‐months of our experiment, no predatory mortality of P. ebeninus was observed and high densities of P. ebeninus decreased impacts of predators on macroinvertebrate abundances. Given that the dynamics of southeast Australian mudflats are driven more by disturbance than seasonality in predators and their interactions with prey, it is likely that Pyrazus would be similarly resistant to predation and have negative effects on benthic assemblages at other times of the year, outside of our study period. Thus, in reducing microphytobenthos and the abundance and species richness of macrofauna, high abundances of the detritivore P. ebeninus may severely limit the flow of energy up the food chain to commercially‐important species. This study therefore suggests that trophic cul‐de‐sacs are not limited to the eutrophied pelagic systems in which they were first identified, but may exist in other systems as well.     

Benthic Reef Primary Production in Response to Large Amplitude Internal Waves at the Similan Islands (Andaman Sea,Thailand)

Coral reefs are facing rapidly changing environments, but implications for reef ecosystem functioning and important services, such as productivity, are difficult to predict. Comparative investigations on coral reefs that are naturally exposed to differing environmental settings can provide essential information in this context. One prevalent phenomenon regularly introducing alterations in water chemistry into coral reefs are internal waves. This study therefore investigates the effect of large amplitude internal waves (LAIW) on primary productivity in coral reefs at the Similan Islands (Andaman Sea, Thailand). The LAIW-exposed west sides of the islands are subjected to sudden drops in water temperature accompanied by enhanced inorganic nutrient concentrations compared to the sheltered east. At the central island, Ko Miang, east and west reefs are only few hundred meters apart, but feature pronounced differences. On the west lower live coral cover (-38 %) coincides with higher turf algae cover (+64 %) and growth (+54 %) compared to the east side. Turf algae and the reef sand-associated microphytobenthos displayed similar chlorophyll a contents on both island sides, but under LAIW exposure, turf algae exhibited higher net photosynthesis (+23 %), whereas the microphytobenthos displayed reduced net and gross photosynthesis (-19 % and -26 %, respectively) accompanied by lower respiration (-42 %). In contrast, the predominant coral Porites lutea showed higher chlorophyll a tissues contents (+42 %) on the LAIW-exposed west in response to lower light availability and higher inorganic nutrient concentrations, but net photosynthesis was comparable for both sides. Turf algae were the major primary producers on the west side, whereas microphytobenthos dominated on the east. The overall primary production rate (comprising all main benthic primary producers) was similar on both island sides, which indicates high primary production variability under different environmental conditions.     

Microphytobenthic dynamics in a Wadden Sea intertidal flat – Part II: Seasonal and spatial variability of non-diatom community components in relation to abiotic parameters

The microphytobenthos colonizing the intertidal flats forms an important component of the Wadden Sea. Ten sampling points along a 1-km transect were studied in a fringe area of the Solthörn tidal flat, southern North Sea, in order to determine seasonal differences in the microphytobenthos. An accompanying paper deals with the major component of the flora, the diatoms; here we, focus on the minor taxonomic groups. From May 2008 to May 2009 surface sediments were collected during low tide. Variation of environmental factors as well as microphytobenthic density (abundance and chlorophyll a) were monitored. The area investigated was a mixed-sediment mudflat, with a gradient from coarse to fine. Highest biomass was obtained in summer 2008 with 215.9?±?12.6?mg chlorophyll a m^–2. In late autumn the chlorophyll a concentration decreased continuously at all investigated stations. Lowest values were detected in December 2008. Species abundances varied considerably both along the transect and seasonally, depending on species-specific requirements as well as hydrodynamic conditions (tidal currents). Higher densities of benthic pro- and eukaryotic microalgae were observed in sites characterized by fine sediments. Apart from the diatoms, the most abundant microphytobenthic group was the cyanophytes. Coccoid cyanophytes, mainly Merismopedia sp., were most abundant during summer, with cell numbers up to 5.72?×?10⁶ cells cm^?2, while diatoms dominated in winter, spring and autumn. Filamentous cyanophytes, particularly Microcoleus chthonoplastes, were most abundant during autumn, while coccoid chlorophytes (spring: Chlorococcum submarinum, Crucigenia tetrapedia, Tetraselmis suecica), euglenophytes (summer: Euglena obtusa), dinophytes (autumn: Amphidinium operculatum, A. herdmanii) and cryptophytes (autumn: Hillea marina, Hemiselmis virescens) contributed to the microphytobenthos during warmer seasons. The statistical analysis confirmed that the composition of the microphytobenthos was related to sediment features and to characteristics of particular seasons.     

Coupling between microphytobenthic biomass and fiddler crab feeding

Among the organic matter ingested by fiddler crabs, microphytobenthos is of fundamental importance because it is their main N source. Microphytobenthos abundance generally develop semilunar changes as the dynamics of tidal exposures and day-night cycle are not held constant across days, modifying the balance between growth and mortality. In this study we explored the coupling between temporal dynamics in microphytobenthos abundance and crab feeding activity. We measured the Chlorophyll a content in the 2 mm surficial sediment surrounding the burrows and the crab feeding activity over two semilunar cycles. Chlorophyll a and crab feeding activity showed biweekly cyclic dynamics. Crabs did not concentrate feeding activity around days with maximum abundance of microhytobenthos. This phase difference between both dynamics could be the result of the crab feeding impact, but a crab experimental exclusion showed that the temporal dynamics of Chlorophyll a content stayed unchanged when feeding activity was removed. Comparisons between fed and unfed sediment suggest that the feeding efficiency changes with tidal dynamic. Crabs achieved more than 50% of Chlorophyll a extraction during days of highest feeding activity, and less than 30% during days of low feeding activity or low microhytobenthos abundance. Furthermore, comparisons of fed sediment between consecutive days indicated that Chlorophyll a was completely replenished during days with high flooding tides, but partially replenished during days near neap tides. Environmental conditions affecting feeding efficiency may select crabs to concentrate feeding activity before days with the highest microhytobenthos abundance. The low feeding impact on microphytobenthos dynamics suggests that fiddler crabs would not control microhytobenthos abundance and thus unable to absorb the increasing eutrophication of studied estuarine areas.     

The circatidal rhythm of the estuarine gastropod Hydrobia ulvae (Gastropoda: Hydrobiidae)

Intertidal animals display a suite of cyclic behaviours that evolved as adaptations to the predictable cycle of inundation and exposure. In estuarine habitats, mud snails from the genus Hydrobia are among the most abundant grazers, and have received considerable attention with respect to the behavioural mechanisms mediating locomotion, dispersal, and feeding, although the nature of the control of these processes has remained elusive. In particular, it is not clear whether endogenous activity patterns are related to periodic changes of microphytobenthos biomass at the sediment surface, or whether they are timed to the tidal cycle at all. In the present study, we address the crawling activity of Hydrobia ulvae under constant conditions, as well as the effects of individual size and previous short‐term exposure to tides of different range, by recording immersed individual snails under constant dark conditions. We show that the species displays an overt circatidal pattern of crawling, with activity peaks around high water, and that the start of inundation may act as an entrainment agent of the rhythm. Moreover, the results obtained indicate that smaller snails display higher levels of activity, although neither the size nor previous in situ influence of tidal range has an effect on the period and on the amplitude of the rhythm. These findings suggest that fluctuations of microphytobenthos biomass are not a sufficiently strong selective pressure to have shaped locomotor activity in H. ulvae. Moreover, feeding of H. ulvae should take place mostly during high water and be independent of periodic fluctuations of microphytobenthos biomass at the surface of the sediment. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 439–450.     

PHOTOSYNTHETIC RESPONSE OF NATURAL ASSEMBLAGES OF MARINE BENTHIC MICROALGAE TO SHORT-AND LONG-TERM VARIATIONS OF INCIDENT IRRADIANCE IN BAFFIN BAY,TEXAS1

This study was designed to understand the high variability characterizing primary production rates of microphytobenthos. The photosynthetic efficiency (α^B) and photosynthetic capacity (P^B_max) of the microphytobenthos were measured at different times of the day on two different dates (8 May and 7 July 1990). In July, unusually low light conditions were caused by the development of a brown tide (chrysophytes). Both light-limited and light-saturated photosynthesis changed at hourly and monthly scales. There was a linear relationship between α^B and P^B_max, suggesting a common response to environmental factors [α^B= 0.0075(±0.00063)·P^B_max+ 0.00097(±0.0071), R²= 0.94]. Incident irradiance at the sediment-water interface was the primary physical factor that explained variability of both α^B (84%) and P^B_max (92%). Temperature had a negative but minor effect that explained an extra 8% and 2% of the variance, respectively. There was no diel rhythm of α^B and P^B_max and incident irradiance was regulated by wind-induced currents. Therefore, microphytobenthos photosynthesis seemed to be primarily controlled by wind events in Baffin Bay.     

The response of meiofauna and microphytobenthos to engineering effects of fiddler crabs on a subtropical intertidal sandflat

Fiddler crabs are key bioturbators on tidal flats. During their intense bioturbation process, they manipulate large amounts of sediment, altering the physical state of existing materials. We investigated whether different types of sediment bioturbation produced by fiddler crabs modulate meiofaunal assemblages and microphytobenthic content. We hypothesized that sedimentary structures produced by burrowing (the burrow itself and the excavation pellets) and feeding (feeding pellets) generate different microenvironments compared with areas without apparent signs of fiddler crab disturbance, affecting both meiofauna and microphytobenthos, independent of the sampling period. Our results indicate that the engineering effects of burrow construction and maintenance and the engineering effects of fiddler crab foraging modulate meiofaunal assemblages in different ways. Overall, meiofauna from burrows and excavation pellets was more abundant and diverse than at control sites, whereas feeding pellets contained poor meiofaunal assemblages. By contrast, only foraging effects were detected on microphytobenthos; independent of the sampling period, Chl a and phaeopigment content were higher in the feeding pellets, but similar among burrows, excavation pellets and control sites. The present study demonstrates that the different engineering effects of fiddler crabs are an important source of habitat heterogeneity and a structuring agent of meiofaunal assemblages on subtropical tidal flats.     

Effects of desiccation on the photosynthetic activity of intertidal microphytobenthos biofilms as studied by optical methods

Due to the periodic exposure to air during periods of low tide, desiccation can be expected to cause important limiting effects on the photosynthetic activity of intertidal microphytobenthos biofilms. This work addresses the study of the short-term effects of desiccation on microphytobenthos using a new, simple methodological approach to non-destructively estimate the water content of muddy intertidal sediments. The method is based on the non-destructive measurement of the specular reflectance in the visible spectral region, shown to be linearly related to the water content of the uppermost 200 µm of the sediment. During air exposure, water loss by the surface sediment layers was shown to induce marked decreases in both the photosynthetic activity, as measured by the maximum quantum yield of photosystem II, F_v/F_m, and the surface microalgal biomass, as estimated from the diffusive reflectance biomass index NDVI. The effects of desiccation were largely dependent on the rate of sediment de-watering. For a same level of desiccation, samples under fast desiccation (exposed to wind of 4.2 m s^− 1) showed much larger effects on F_v/F_m and NDVI comparatively to samples under slow desiccation (maintained under still air). By showing the rapid and significant effects of desiccation on microphytobenthos biofilm functioning, the results of this study have potentially important implications for the modelling of primary productivity of estuarine intertidal areas, as desiccation and factors inducing it may result in previously unaccounted effects on photosynthetic performance and productive biomass.     

Effects of the physical structure of mangrove vegetation on a benthic faunal community

Clarification of the role of the physical structure of mangrove for benthic faunal communities was sought by investigating the impacts of canopy shade and root structure on (1) the physical environment, including temperature, moisture and grain size of the substrate sediment, (2) benthic faunal distribution, and (3) food resource availability, using a field manipulated experiment at an intertidal mangrove forest around Sikao Creek, Trang Province, Thailand. Five treatments were established, including artificial shade cover, root simulated structure, mix (shade and structure) and control of the canopy gap (which had no mangrove vegetation), in addition to forest control under the mangrove canopy. Following 18 months of observation, species' richness and abundance of epifauna were found to have increased in shaded treatments, which had low temperature and high moisture substrate. Food resource conditions had also altered from abundant microphytobenthos (relatively high nutritional values) to enriched mangrove detritus due to shading. These results indicated that the physical structure of mangrove vegetation facilitates the habitation of intertidal epifauna, canopy shade having an important function in providing cooler wetter surface substrate, despite also inducing a reduction of favorable food resources (i.e. microphytobenthos).     

Community structure and food web based on stable isotopes (δN and δC) analysis of a North Eastern Atlantic maerl bed

Maerl beds are highly biodiverse biogenic substrata that have been receiving increasing attention in the last decade. Although maerl beds represent important nursery areas for commercial fishes and molluscs, little is known on the trophic web of their communities. Community structure parameters of maerl bed of the Bay of Brest (species richness, abundance, biomass and dominating species) were studied in parallel with the carbon and nitrogen isotopic composition of their main benthic species (macrofaunal, and megafaunal organisms) in order to assess the trophic levels and differences in the potential food sources of maerl inhabitants. The major potential sources of energy were identified to originate either from epiphytic macroalgae and microphytobenthos both growing on maerl thalli, together with sedimenting (sedimentary) particulate organic matter (POM) originating from the water column. The majority of the macro- and megafaunal organisms investigated were filter feeders, selective-deposit feeders and predators/scavengers. Filter feeders fall into three different groups representing different trophic pathways (i) sponges feeding directly on POM (water column filter feeders I), (ii) ascidians and holothurians feeding on POM and probably captured pelagic preys (water column filter feeders II), and (iii) filter feeding molluscs and crustaceans were hypothesised to feed on microphytobenthos or on decaying sedimented POM (Interface filter feeders). Selective deposit feeders were also divided into two subgroups. Carnivores were also distinguished between those with scavenging habits and true predators. Coupling of the trophic levels observed with the community biomass structure revealed that most of the benthic biomass derives its food from detritic sedimented POM and/or microphytobenthos, with interface filter feeders (23% of the biomass), selective deposit feeders (12%). Carnivores made up to 14% of the total biomass. Generally stable isotopes ratio mean values overlap and cover a large range within feeding types, indicating a strong overlap in food sources and a high degree of complexity of the food web presumably due to the diversity of the potential food sources.