Differential effects of native and invasive algal wrack on macrofaunal assemblages inhabiting exposed sandy beaches

Many sandy beaches worldwide receive large amounts of drift seaweed, known as wrack, from offshore algal beds and closer rocky intertidal shores. Despite the important influence of algal wrack on macrofaunal assemblages from different coastal systems, relatively little attention has been paid to the macrofaunal responses in sandy beaches to macrophyte wrack supplies. Algal wrack is a key resource, i.e. for food and/or refuge, for beach invertebrates while its availability can affect diversity and abundance of intertidal animals including shorebirds, but the role of certain types of wrack and its location on the shore has not been examined experimentally to date. In this paper, we use experimental manipulation of two species of brown seaweeds, i.e. artificial wrack patches made up of the native macroalgae Saccorhiza polyschides and the invasive species Sargassum muticum, to test hypotheses about influences on macrofaunal assemblages inhabiting the drift line and supratidal levels of exposed beaches. Results pointed out that different types of wrack deposits were not used uniformly by invertebrates. Nutritional value differed between the two species of wrack. In most cases, the carbohydrates, lipids and organic carbon content were greater in patches of S. muticum than in patches of S. polyschides. Data also provided evidences that nutritional content and microclimatic conditions of wrack deposits, i.e. temperature and humidity, might affect macrofaunal assemblages.     

Small-scale burial of macroalgal detritus in marine sediments: Effects of Ulva spp. on the spatial distribution of macrofauna assemblages

Patches of dead seaweeds can deposit, bury, and age into the sediment. Decomposition and release of algal-derived nutrients can influence patterns of distribution of benthic organisms. Here, I investigated how small-scale burial of Ulva spp. affected spatial variation of macrofauna in intertidal sediment. I deliberately buried Ulva detritus under the surface of 50 × 50 cm² patches of sediment in three intertidal flats of the Oosterschelde estuary (The Netherlands). Results showed that there was no accumulation of particulate organic carbon and nitrogen in the sediment at the scales examined. The biomass of microphytobenthos did not show any change and there was evidence that grazing was important all over the study area. Burial did not alter composition and diversity of macrofauna, but some animals (Corophium volutator, Eteone spp. and Scoloplos armiger) had less numbers in the plots where detritus was buried than in the controls. These findings showed that burial of macroalgal detritus does not represent a major source of variation at the scales examined. It is suggested that in these sediments, recycling of detritus is fast and it buffers the effects of excess organic matter in the system.     

Effects of sandy beach cusps on wrack accumulation,sediment characteristics and macrofaunal assemblages

Beach‐cast marine macroalgae and seagrasses, collectively termed wrack, provide shelter and habitat for beach fauna and can often provide a large input of energy and nutrients to sandy beaches. Wrack deposition on sandy beaches varies spatially and could be affected by morphological features on the beach face such as cusps. This study tested a series of hypotheses regarding the differences in wrack deposits, sediments and macrofaunal assemblages between cusp bays and horns on two beaches in South Australia. Bays had greater cover and larger pieces of wrack than horns. Sediment organic matter content was greater on horns than in bays but mean particle size did not differ consistently between bays and horns. Macrofaunal diversity was higher in bays and this pattern was probably driven by differences in the cover of wrack between bays and horns. Cusp morphology thus influences the distribution of wrack on the beach face, which in turn influences the distribution of macrofauna. Studies of sandy beaches with cusps should therefore be explicitly designed to sample cusp features and associated wrack deposits.     

Response of deposit-feeders to exclusion of epibenthic predators in a Mediterranean intertidal flat

Deposit-feeders are common components of macrofaunal assemblages in intertidal soft sediments. Predation has been considered to have a central role in affecting their distribution and population dynamics. This study investigates the effect of epibenthic predators on deposit-feeders, inhabiting the shallow layers of the sediment (surface and subsurface) and also the deepest layers (e.g., burrowing shrimp Upogebia pusilla; Petagna). The experiment was conducted in summer 2000 (August-September) at three different sites on an intertidal flat in Mediterranean Sea. In the field, predators were excluded using cages, placed on the surface of the sediment. It was predicted that under the cages, (i) abundances of animals would increase and (ii) species composition of assemblages would change as an effect of the exclusion of predators. Potential artefacts caused by the cages were controlled using partial cages (PC). Composition of organic matter and porosity were also analysed under PC and in natural controls to test whether the presence of cages would change sediment characteristics on the surface. At the end of the experiment, there were no significant differences in the composition of organic matter between PC and the undisturbed sediment (UC). After 8 weeks from the beginning of the experiment, exclusion of predators enhanced the abundance of U. pusilla. There were, however, no clear-cut changes in the species composition of macrofaunal assemblages and densities of animals did not increase under the cages. Indeed, some animals (Oligochaeta, Brania oculata, and Tanais dulongii) were less abundant under the cages (EC) than outside (PC and UC). We propose that predation might play a role in regulating interspecific relationships among some surface deposit-feeders and the burrowing shrimp U. pusilla.     

Long-Term Field Study Reveals Subtle Effects of the Invasive Alga Sargassum muticum upon the Epibiota of Zostera marina

Invasive species can alter coastal ecosystems both directly, e.g. through competition for substratum and nutrients, and indirectly. Indirect effects may be mediated by creation of dissimilar or inimical habitats, changes in predator and/or prey assemblages, alterations in associated biota, and perturbations of water movement and thermal regimes. Previous studies have shown that invasive algae can modify native habitat architecture, disrupt intricately linked food webs and alter epibiotic assemblages. In the UK, the seagrass Zostera marina supports a diverse epibiotic assemblage, influencing key factors such as sediment dynamics, depositional regime and trophic linkages. Increasing encroachment of the invasive alga Sargassum muticum into seagrass meadows changes the physical and chemical characteristics of the local environment and creates the potential for changes in the epibionts associated with the seagrass blades, threatening the integrity of the seagrass ecosystem. We investigated the effects of S. muticum invasion upon the epibiota of Z. marina in a drowned river valley in SW England seasonally from spring to autumn over four years in an in-situ manipulative experiment, comparing permanent quadrats with and without artificially introduced S. muticum. Epibiota were weighed, identified to the most detailed operational taxonomic unit (OTU) possible, and unitary organisms were enumerated. Multivariate PERMANOVA+ analysis revealed significant differences in epibiont assemblages between Sargassum treatments. Linear mixed effects models indicated that differences in epibiota assemblage composition were not reflected as significant differences in mean biomass per sample, or number of epibiont OTUs per sample. We conclude that S. muticum invasion into Z. marina meadows may significantly alter the species composition and abundance distribution of epibiotic assemblages found on the blades of the seagrass. Thus S. muticum invasion could have more wide-reaching effects on processes within coastal ecosystems than predicted purely by direct effects.     

Spatial changes in sublittoral soft-bottom polychaete assemblages due to river inputs and sewage discharges

Spatial changes in species composition and structure of sublittoral soft-bottom polychaete assemblages along a transect parallel to the shoreline off the Barcelonès and Maresme regions on the Spanish Catalan coast (NW Mediterranean sea) are described. The transect (20 m mean depth) covers 30 km through an area affected by both water sewage discharges and river outflow. These effluents generate a spatial gradient of both silt and organic enrichment, according to the prevailing southerly currents in the zone. South of the effluents, eutrophized sediments supported a macrofaunal assemblage which was typical of polluted bottoms, characterized by the massive presence of Capitella capitata and Malacoceros fuliginosus, high abundance and biomass, low values of species richness, and a simplified trophic structure dominated by surface and subsurface-deposit feeders. North of the effluents, both the mud and organic matter content in sediment decreased drastically and the polychaete assemblages described were progressively less affected with distance. Thus, a clear change in species composition, as well as low values of abundance and biomass, increases in species richness, and a more complex trophic structure on the polychaete assemblages was observed north rather than south of the effluents.     

The role of microorganisms in mobilization and fixation of phosphorus in sediments

Cycling of phosphorus (P) at the sediment/water interface is generally considered to be an abiotic process. Sediment bacteria are assumed to play only an indirect role by accelerating the transfer of electron from electron donors to electron acceptors, thus providing the necessary conditions for redox-and pH-dependent, abiotic sorption/desorption or precipitation/dissolution reactions. Results summarized in this review suggest that

1. in eutrophic lakes, sediment bacteria contain as much P as settles with organic detritus during one year
2. in oligotrophic lakes, P incorporated in benthic bacterial biomass may exceed the yearly deposition of bioavailable P several times
3. storage and release of P by sediment bacteria are redox-dependent processes
4. an appreciable amount of P buried in the sediment is associated with the organic fraction
5. sediment bacteria not only regenerate PO₄, they also contribute to the production of refractory, organic P compounds, and
6. in oligotrophic lakes, a larger fraction of the P settled with organic detritus is converted to refractory organic compounds by benthic microorganisms than in eutrophic lakes.

From this we conclude that benthic bacteria do more than just mineralize organic P compounds. Especially in oligotrophic lakes, they also may regulate the flux of P across the sediment/water interface and contribute to its terminal burial by the production of refractory organic P compounds.     

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.     

Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Weak diurnal changes in the biochemical properties and benthic macrofauna of urbanised mangrove forests and mudflats

Diurnal changes in the biochemical properties and the benthic macrofaunal assemblage of sediments in urbanised mangrove forests and their adjacent mudflats in Sydney Harbour were investigated. Behavioural and physiological changes in the microphytobenthos between day and night were predicted to cause diurnal changes in the micro-scale depth distribution of chlorophylls a and b and colloidal carbohydrate. In addition, because macrofauna can alter sediment properties, diurnal changes in the macrofaunal assemblages were investigated. The microphytobenthos at the study sites were predominantly filamentous green algae, although diatoms were present. Samples for biochemical analysis were collected from the top 2 mm of sediment using mini-cryolanders, during low tide in the day and at night. Three biochemical properties of the sediments were measured spectrophotometrically: chlorophylls a and b (surrogate for microphytobenthos biomass) and colloidal carbohydrate. The amount of chlorophylls tended to be less at night than during the day, but site to site variability was large and these differences were generally small and not significant. Depth profiles indicated that there was some redistribution of pigments in the surface 2 mm between day and night, possibly due to migration of microphytobenthos or physiological changes. There was no significant difference in chlorophylls between the mangrove forest and adjacent mudflat, with the exception of chlorophyll b at one sampling time, which was larger in the mangrove forest than on the mudflat. Colloidal carbohydrate was significantly larger in the mangrove forest and significantly less on the mudflat during the day at one site at one time, but otherwise showed no significant differences between day and night or between the mangrove forest and mudflat. Whilst there were some differences in the benthic macrofaunal assemblages between day and night, these differences were only significant for spionids and polychaetes at one time. There were, however, significant differences in assemblages of benthic macrofauna between the mangrove forest and mudflat, probably due to structural differences between these habitats such as the presence of pneumatophores, shade and leaf litter. In summary, there was some minor diurnal variation in the measured biochemical properties of the sediment, but not in the macrofaunal assemblage. Diurnal changes should, therefore, be considered when investigating biochemical properties in these habitats, but they are not a major influence. These findings contrast to previous studies on diatom dominated mudflats in Europe, where stronger diurnal changes in biochemical properties were found. Diurnal changes in the macrofauna assemblages were largely insignificant and therefore could not explain the changes in the biochemical properties. Diurnal effects on the macrofauna in these habitats are more likely to be via altered behaviours and this requires further investigation.     

The effect of natural populations of the burrowing and grazing soldier crab (Mictyris longicarpus) on sediment irrigation, benthic metabolism and nitrogen fluxes

The aim of this study was to determine the effect of sediment grazing and burrowing activities of natural populations of Mictyris longicarpus on benthic metabolism, nitrogen flux and irrigation rates by comparing sediments taken from minimum disturbance exclusion cages and adjacent sediments subject to M. longicarpus activities. M. longicarpus reduced sediment surface chlorophyll a (approximately 77%), organic carbon (approximately 95%) and total nitrogen concentrations (approximately 99%) in comparison to ungrazed sediments. Consequently, they significantly reduced gross benthic O₂ production (about 71%) and sediment O₂ consumption (approximately 46%). Mean N₂ fluxes showed net effluxes (276-430 μmol m⁻² day⁻¹) in the presences of M. longicarpus and net uptakes (194.09-449.21 μmol m⁻² day⁻¹) where they were excluded. The net uptake of N₂ was most likely due to cyanobacteria fixing of N₂, as dense microbial mats became established over the sediment surface in the absence of M. longicarpus grazing activity. Sediment irrigation/transport rates calculated from CsCl tracer dilution indicated greater irrigation rates in the exclusions (12.12-16.22 l m⁻² h⁻¹) compared to inhabited sediments (6.33-11.73 l m⁻² h⁻¹) and this was again was most likely due to the lack of grazing pressure which allowed large populations of small burrowing polychaetes to inhabit the organic matter rich exclusion sediments. As such, the main influence of M. longicarpus was the interception and consumption of transported organic material, benthic microalgae and other small infaunal organisms resulting in the removal of approximately 0.06 g m⁻² day⁻¹ of nitrogen and 12.12 g m⁻² day⁻¹ of organic carbon. This “cleansing” of the sediments reduced sediment metabolism and the flux of solutes across the sediment water interface and ultimately the heavy predation of M. longicarpus by transient species such as stingrays, results in a net loss of carbon and nitrogen from the system.     

The effect of artificial defaunation on bacterial assemblages of intertidal sediments

The use of the artificial defaunation of sediments is widespread among studies examining the disturbance and recovery of benthic macrofaunal communities. Standard methods of defaunation include driving the sediment to anoxia, freezing and sieving. In this study we performed a field experiment to test the assumption that the bacterial assemblages are unaffected by these methods of defaunation. Same-sized patches of sediment were defaunated by covering sediment with plastic sheeting (weighted by concrete blocks), freezing or sieving (1-mm mesh). Macrofaunal counts of sediment cores, taken to determine the effectiveness of each defaunation method, indicated that although none of the treatments removed 100% of macrofauna, all resulted in reduced macrofaunal presence, with the sieved treatment being the most effective. Bacterial samples were taken over the course of a month to determine both the initial and long-term effects of defaunation on bacterial community structure. Numerical effects were determined via epifluorescence microscopy, whereas differences in community composition were followed using PCR and denaturing gradient gel electrophoresis (DGGE). The anoxic treatments resulted in significant numerical changes in both active and total cell counts over time, while the frozen and sieved treatments caused less apparent changes. All of the treatments initially changed the composition of the community; however, anoxic and sieved treatments resulted in subtle changes while the frozen treatment produced more notable and variable changes within the community. The composition of the bacterial community in all of the treatment plots trended towards recovery, or convergence towards that of ambient sediments, by the t = 25-day sampling period.     

Impacts of the invasive alga Sargassum muticum on ecosystem functioning and food web structure

Biological invasions have the potential to cause severe alterations to the biodiversity of natural ecosystems. At the same time, variation in the diversity and composition of native communities may have an important influence on the impact of invasions. Here, effects of the invasive Japanese wireweed, Sargassum muticum, were tested across a range of native marine algal assemblages using a combined additive and substitutive design. The invasive alga significantly reduced primary production, an important component of ecosystem functioning, and increased connectance, a key property of the food webs associated with the algal resources. These impacts were mediated by changes in the proportions of intermediate and top species, as well as apparent reductions in faunal species richness and diversity. Some key alterations to faunal species composition (including the arrival of generalist species associated with S. muticum) may have been important in determining these patterns. Overall results suggest that S. muticum not only directly impeded the native algal community, but that these effects extended indirectly to the native fauna and therefore caused major changes throughout the ecosystem.     

Glacial Holocene environment of the southeastern Okhotsk Sea: evidence from geochemical and palaeontological data

Environmental conditions and productivity changes in the southeastern Okhotsk Sea have been reconstructed for the last 20 ka using planktonic and benthic foraminiferal oxygen isotope records and calcium carbonate, organic carbon and opal content data from two sediment cores. Species variability in benthic foraminiferal and diatom assemblages provides additional palaeoceanographic evidence. AMS radiocarbon dating of the sediments and oxygen isotope stratigraphy serve as the basis for the age models of the cores for the last 20 ¹⁴C kyr and for correlation between environmental variations in the Okhotsk Sea, and regional and global climate changes. Benthic foraminiferal assemblages in the two cores (depth 1590 and 1175 m) varied with time, so that we could recognise seven zones with different species composition. Changes in the benthic foraminiferal assemblages parallel major environmental and productivity variations. During the last glaciation, fluxes of organic matter to the sea floor showed strong seasonal variations, indicated by the presence of abundant A. weddellensis and infaunal Uvigerina spp. Benthic foraminiferal assemblages changed with warming at 12.5–11 and 10–8 ¹⁴C kyr BP, when productivity blooms and high organic fluxes were coeval with global meltwater pulses 1A and 1B. Younger Dryas cooling caused a decline in productivity (11–10 kyr BP) affecting the benthic faunal community. Subsequent warming triggered intensive diatom production, opal accumulation and a strong oxygen deficiency, causing significant changes in benthic fauna assemblages from 5.26–4.4 kyr BP to present time.     

Effect of organic carbon flux and dissolved oxygen on the benthic foraminiferal oxygen index (BFOI)

Variations in oceanic primary productivity, flux of organic carbon to the sediments, and dissolved-oxygen levels in the water column are thought to be important in the control of benthic foraminiferal test size, wall thickness, morphology, and species composition of assemblages by many foraminiferal paleontologists. Aspects of these processes should be reflected by the benthic foraminiferal oxygen index (BFOI) based on these foraminiferal characteristics. However, analyses indicate that the BFOI correlates most strongly with dissolved-oxygen levels in overlying water (R²=0.81), weakly with oceanic primary productivity (R²=0.55), and weakly with organic carbon flux to the sediments (R²=0.51). Although both dissolved oxygen and organic carbon flux are main controlling factors for benthic foraminiferal assemblages, the BFOI is a useful indicator extracted from benthic foraminiferal assemblages for estimating the condition of dissolved oxygen in Cretaceous and Cenozoic oceans.     

Biomass dynamics of exotic Sargassum muticum and native Halidrys siliquosa in Limfjorden,Denmark—Implications of species replacements on turnover rates

The expansion of Sargassum muticum in the Danish estuary Limfjorden between 1984 and 1997 was followed by a decrease in abundance of native perennial macroalgae such as Halidrys siliquosa. Although commonly associated with the expansion of exotic species, it is unknown whether such structural changes affect ecosystem properties such as the production and turnover of organic matter and associated nutrients. We hypothesized that S. muticum possesses ‘ephemeral’ traits relative to the species it has replaced, potentially leading to faster and more variable turnover of organic matter. The biomass dynamics of S. muticum and H. siliquosa was therefore compared in order to assess the potential effects of the expansion of Sargassum. The biomass of Sargassum was highly variable among seasons while that of Halidrys remained almost constant over the year. Sargassum grew faster than Halidrys and other perennial algae and the annual productivity was therefore high (P/B = 12 year⁻¹) and exceeded that of Halidrys (P/B = 5 year⁻¹) and most probably also that of other perennial algae in the system. The major grazer on macroalgae in Limfjorden, the sea urchin Psammechinus miliaris, preferred Sargassum to Halidrys, but estimated losses due to grazing were negligible for both species and most of the production may therefore enter the detritus pool. Detritus from Sargassum decomposed faster and more completely than detritus from Halidrys and other slow-growing perennial macrophytes. High productivity and fast decomposition suggest that the increasing dominance of S. muticum have increased turnover of organic matter and associated nutrients in Limfjorden and we suggest that the ecological effects of the invasion to some extent resemble those imposed by increasing dominance of ephemeral algae following eutrophication.     

Early stages of leaf decomposition are mediated by aquatic fungi in the hyporheic zone of woodland streams

1. Leaf litter constitutes the major source of organic matter and energy in woodland stream ecosystems. A substantial part of leaf litter entering running waters may be buried in the streambed as a consequence of flooding and sediment movement. While decomposition of leaf litter in surface waters is relatively well understood, its fate when incorporated into river sediments, as well as the involvement of invertebrate and fungal decomposers in such conditions, remain poorly documented. 2. We tested experimentally the hypotheses that the small interstices of the sediment restrict the access of the largest shredders to buried organic matter without compromising that of aquatic hyphomycetes and that fungal decomposers in the hyporheic zone, at least partly, compensate for the role of invertebrate detritivores in the benthic zone. 3. Alder leaves were introduced in a stream either buried in the sediment (hyporheic), buried after 2 weeks of exposure at the sediment surface (benthic‐hyporheic), or exposed at the sediment surface for the entire experiment (benthic). Leaf decomposition was markedly faster on the streambed surface than in the two other treatments (2.1‐ and 2.8‐fold faster than in the benthic‐hyporheic and hyporheic treatments, respectively). 4. Fungal assemblages were generally less diverse in the hyporheic habitat with a few species tending to be relatively favoured by such conditions. Both fungal biomass and sporulation rates were reduced in the hyporheic treatment, with the leaves subject to the benthic‐hyporheic treatment exhibiting an intermediate pattern. The initial 2‐week stage in the benthic habitat shaped the fungal assemblages, even for leaves later subjected to the hyporheic conditions. 5. The abundance and biomass of shredders drastically decreased with burial, except for Leuctra spp., which increased and was by far the most common leaf‐associated taxon in the hyporheic zone. Leuctra spp. was one of the rare shredder taxa displaying morphological characteristics that increased performance within the limited space of sediment interstices. 6. The carbon budgets indicated that the relative contributions of the two main decomposers, shredders and fungi, varied considerably depending on the location within the streambed. While the shredder biomass represented almost 50% of the initial carbon transformed after 80 days in the benthic treatment, its contribution was <0.3% in the hyporheic one and 2.0% in the combined benthic‐hyporheic treatment. In contrast, mycelial and conidial production in the permanently hyporheic environment accounted for 12% of leaf mass loss, i.e. 2–3 times more than in the two other conditions. These results suggest that the role of fungi is particularly important in the hyporheic zone. 7. Our findings indicate that burial within the substratum reduces the litter breakdown rate by limiting the access of both invertebrate and fungal decomposers to leaves. As a consequence, the hyporheic zone may be an important region of organic matter storage in woodland streams and serve as a fungal inoculum reservoir contributing to further dispersal. Through the temporary retention of litter by burial, the hyporheic zone must play a significant role in the carbon metabolism and overall functioning of headwater stream ecosystems.     

The impact of infauna (Nereis diversicolor and Saduria entomon) on the redistribution and biomass of macroalgae on marine soft bottoms

Mass occurrence of macroalgae is a phenomenon attributed to eutrophication, and can lead to drastic changes in the benthic communities on soft bottoms. While the negative effects of macroalgal blooms on the macrozoobenthos have been studied extensively, the effects of the infauna on the macroalgal material have not previously been studied in the northern Baltic Sea. The impact of the infaunal species Nereis diversicolor and Saduria entomon on the burial and biomass of Enteromorpha spp., Cladophora glomerata and Fucus vesiculosus, was assessed through a series of microcosm experiments. Results show that S. entomon did not significantly affect the biomass of the algae, nor actively relocate them. N. diversicolor redistributed the filamentous green algae into the sediment, down to 4 cm at most, and decreased the biomass of the filamentous algae by 140-360%. Furthermore, the loss of biomass promoted in presence of polychaetes proved to be a density dependent process. The effect on the perennial macroalgal species, F. vesiculosus, was less clear, as no redistribution or significant change in macroalgal biomass was observed. Our findings show that infauna can contribute to a loss in macroalgal biomass through feeding and burrowing activities leading to the redistribution and incorporation of the detritus into bioturbated sediment.     

Structuring factors in a marine soft bottom community during eutrophication—an experiment with radio-labelled phytodetritus

The quantitative development and uptake of radio-labelled phytodetritus in benthic macro- and meiofauna was studied in a 5-month experiment in two mesocosms, one of which received a single large nutrient (N and P) addition, while the other served as control. In reponse to the 12-fold increase in phytoplankton biomass noted after 2 weeks and the resulting enhanced accumulation of fresh phytodetritus, the abundance and biomass of the polychaetes Mediomastus ambiseta and Polydora ligni and the mud anemone Cerianthiopsis americanus increased significantly in the enriched tank. The abundances of P. ligni and M. ambiseta increased 37-fold and 12-fold, respectively, within the first two months of the experiment. No other macrofaunal or meiofaunal taxa showed any consistent positive quantitative response to the increased input of phytodetritus. In the control tank no considerable change in the benthic community structure was noted. The measurements of radio-label uptake within the benthic fauna showed that the quantitatively most successful species utilized fresh phytodetritus highly. However, a high degree of utilization of fresh detritus was also shown by taxa that did not respond quantitatively within the 5 month of the experiment, and almost all taxa showed a preference for fresh detritus over older organic material. Within the benthic meiofauna, kinorhynchs and especially foraminiferans showed a remarkably low preference for fresh detritus. A budget calculation comparing the total amounts of labelled organic carbon bound in animal tissue and in the sediment indicated that at any time at least 75% of this carbon was available for assimilation by deposit feeders. These results suggest that factors other than the availability of food, such as competition for space by a few opportunistic macrofauna species, limited the response of other species within this benthic community to the increased input of phytodetritus.