Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

Food supply impacts sediment reworking by <Emphasis Type="Italic">Nereis diversicolor</Emphasis>

This study assessed the effects of food supply on sediment reworking by Nereis diversicolor. We hypothesized that food supply would enhance sediment reworking and that the frequency of food supply would affect the intensity of bioturbation. Mesocosm experiments consisted of four treatments: (1) without worms (control cores), (2) with worms and no food supply, (3) with worms and daily food supply, (4) with worms and weekly food supply. Fluorescent particles, used as tracers, were spread over the sediment surface. Sediment reworking was quantified after 28 days based on the tracer distribution profiles. Results showed that sediment reworking by N. diversicolor was exclusively due to non-local transport processes. Food supply greatly increased non-local transport coefficients (more than 3 times) in comparison with those measured in the absence of a food supply. However, the intensity of sediment bioturbation by these worms was unaffected by the frequency of food supply. This study showed that environmental conditions affecting the quantity of food supply at the water-sediment interface could strongly influence bioturbation process. Handling editor: P. Viaroli     

Impact of Oil on Bacterial Community Structure in Bioturbated Sediments

Oil spills threaten coastlines where biological processes supply essential ecosystem services. Therefore, it is crucial to understand how oil influences the microbial communities in sediments that play key roles in ecosystem functioning. Ecosystems such as sediments are characterized by intensive bioturbation due to burrowing macrofauna that may modify the microbial metabolisms. It is thus essential to consider the bioturbation when determining the impact of oil on microbial communities. In this study, an experimental laboratory device maintaining pristine collected mudflat sediments in microcosms closer to true environmental conditions – with tidal cycles and natural seawater – was used to simulate an oil spill under bioturbation conditions. Different conditions were applied to the microcosms including an addition of: standardized oil (Blend Arabian Light crude oil, 25.6 mg.g⁻¹ wet sediment), the common burrowing organism Hediste (Nereis) diversicolor and both the oil and H. diversicolor. The addition of H. diversicolor and its associated bioturbation did not affect the removal of petroleum hydrocarbons. After 270 days, 60% of hydrocarbons had been removed in all microcosms irrespective of the H. diversicolor addition. However, 16S-rRNA gene and 16S-cDNA T-RFLP and RT-PCR-amplicon libraries analysis showed an effect of the condition on the bacterial community structure, composition, and dynamics, supported by PerMANOVA analysis. The 16S-cDNA libraries from microcosms where H. diversicolor was added (oiled and un-oiled) showed a marked dominance of sequences related to Gammaproteobacteria. However, in the oiled-library sequences associated to Deltaproteobacteria and Bacteroidetes were also highly represented. The 16S-cDNA libraries from oiled-microcosms (with and without H. diversicolor addition) revealed two distinct microbial communities characterized by different phylotypes associated to known hydrocarbonoclastic bacteria and dominated by Gammaproteobacteria and Deltaproteobacteria. In the oiled-microcosms, the addition of H. diversicolor reduced the phylotype-richness, sequences associated to Actinobacteria, Firmicutes and Plantomycetes were not detected. These observations highlight the influence of the bioturbation on the bacterial community structure without affecting the biodegradation capacities.     

Rapid reworking of subtidal sediments by burrowing spatangoid urchins

The mixing and displacement of sediment by benthic macrofauna (bioturbation) has major biogeochemical implications, and can control rates of organic matter degradation and carbon burial. Large, abundant, mobile macrofauna often dominate sediment bioturbation, and heart urchins of the genus Echinocardium are regarded as key sediment bioturbators in marine systems throughout the world. To better understand the bioturbation potential and functional role of Echinocardium, we developed a mathematical model and parameterized it with field data from six locations in northern New Zealand in order to estimate bioturbation rates in these places. Although urchin sizes and densities were measured in consecutive years at all six locations, we obtained a third model parameter, urchin movement rate, from one time and place only (Site OB5). Because confidence in model output was greatest at OB5, and since OB5 had the highest sediment reworking rate of all sites, our model yielded a good upper bound estimate for the bioturbation potential of Echinocardium in the areas examined. The volume of sediment displaced by Echinocardium populations reached 20,000 cm³ m⁻² d⁻¹ at OB5, suggesting that surface sediment is reworked about every 3 days at sites where Echinocardium is abundant. Experimental work with a fluorescent tracer at OB5 suggested limited downward particle movement as a result of Echinocardium bioturbation, though vertical profiles of chlorophyll a and organic matter content indicated well mixed sediment. The loss of Echinocardium because of broad-scale anthropogenic disturbance to the seabed could have major consequences on marine ecosystem functioning.     

Nereis diversicolor effect on the stability of cohesive intertidal sediments

The effect of the polychaete Nereis diversicolor on the stability of natural cohesive sediments was investigated in the laboratory. Three densities (450, 600 and 1200 ind m⁻²) of N. diversicolor were used. Sediment shear strength was measured using a cone penetrometer. Sediment erodability was assessed using an annular flume (current velocities from 5 to 55 cm s⁻¹) in which flow velocity was increased incrementally, and water sampled to quantify suspended material in order to derive critical erosion velocity and erosion rates. At low current velocities ( <25 cm s⁻¹), we found N. diversicolor to have a stabilising effect, reflected by an increase of up to 20% in the critical erosion velocity. This is related to an enhancement of ~50% in shear strength, due probably to gallery building activities, responsible for the promotion of lateral compaction, an increase in the area of the sediment–water interface, and enhanced microphytobenthos production. Once the sediment began to erode, the stabilising effect of N. diversicolor reverses, leading to an increase of up to 40% in eroded matter due to compaction, which resulted in the erosion of larger aggregates. The balance between the effect of N. diversicolor on herbivory and microphytobenthos production due to the presence of galleries is discussed. Our results indicate that neither chlorophyll a, nor shear strength nor critical erosion velocity are good indicators of erodability. This underlines the need to include biogeochemical processes in any realistic sediment transport model.     

Macrofauna distribution and bioturbation on tidal confluences of the dutch Wadden Sea

Macrofauna density and bioturbation intensity (measured with X-ray radiography), were studied in the Dutch Wadden Sea near Ameland and in Mok Bay, Texel, in September 1988. The bivalveMacoma balthica and the polychaeteHeteromastus filiformis were widely distributed in the areas studied. The cockleCerastoderma edule and the polychaeteScoloplos armiger did not occur in areas with high clay content, while the molluscsHydrobia ulvae andMya arenaria preferred fine sediments. The polychaeteNereis diversicolor preferred the higher parts of the intertidal. In the Ameland area disturbance of primary sediment structures was highest near the shore and near the middle of the tidal confluence where physical reworking was low. The sheltered Mok Bay sediments were completely reworked by deposit-feeders. Bioturbation intensity and deposit-feeder (bioturbator) density were, however, not highly correlated. A number of stations showed higher bioturbation towards the surface and this may be related to the high abundance of near surface dwelling macrofauna, due to eutrophication and organic enrichment of the sediments in recent years.     

Bioturbation of Burrowing Crabs Promotes Sediment Turnover and Carbon and Nitrogen Movements in an Estuarine Salt Marsh

Ecological functions of bioturbation in ecosystems have received increasing attention over the recent decades, and crab burrowing has been considered as one of the major bioturbations affecting the physical and chemical processes in salt marshes. This study assessed the integrated effects of crab excavating and burrow mimic trapping on sediment turnover and vertical C and N distributions in a Chinese salt marsh in the Yangtze River estuary. Crab burrowing increased soil water content and the turnover of carbon and nitrogen and decreased bulk soil density. Vertical movement of materials, nutrient cycling and reuse driven by crab burrowing might be obstructed by vegetation (Phragmites australis and Spartina alterniflora communities). The amount of soil excavated by crab burrowing was higher than that deposited into burrow mimics. In Phragmites marshes, Spartina marshes and unvegetated mudflats, net transport of soil to the marsh surface was 171.73, 109.54, and 374.95 g m⁻² d⁻¹, respectively; and the corresponding estimated soil turnover time was 2.89, 4.07 and 1.83 years, respectively. Crab burrowing in salt marshes can mix surface and deeper soil over a period of years, accelerating litter decomposition and promoting the efficient reuse of nutrients by plants. Therefore, bioturbation affects soil physical processes and functioning of ecosystems, and needs to be addressed in ecosystem management.     

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.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

Nitrate Reduction Functional Genes and Nitrate Reduction Potentials Persist in Deeper Estuarine Sediments. Why?

Denitrification and dissimilatory nitrate reduction to ammonium (DNRA) are processes occurring simultaneously under oxygen-limited or anaerobic conditions, where both compete for nitrate and organic carbon. Despite their ecological importance, there has been little investigation of how denitrification and DNRA potentials and related functional genes vary vertically with sediment depth. Nitrate reduction potentials measured in sediment depth profiles along the Colne estuary were in the upper range of nitrate reduction rates reported from other sediments and showed the existence of strong decreasing trends both with increasing depth and along the estuary. Denitrification potential decreased along the estuary, decreasing more rapidly with depth towards the estuary mouth. In contrast, DNRA potential increased along the estuary. Significant decreases in copy numbers of 16S rRNA and nitrate reducing genes were observed along the estuary and from surface to deeper sediments. Both metabolic potentials and functional genes persisted at sediment depths where porewater nitrate was absent. Transport of nitrate by bioturbation, based on macrofauna distributions, could only account for the upper 10 cm depth of sediment. A several fold higher combined freeze-lysable KCl-extractable nitrate pool compared to porewater nitrate was detected. We hypothesised that his could be attributed to intracellular nitrate pools from nitrate accumulating microorganisms like Thioploca or Beggiatoa. However, pyrosequencing analysis did not detect any such organisms, leaving other bacteria, microbenthic algae, or foraminiferans which have also been shown to accumulate nitrate, as possible candidates. The importance and bioavailability of a KCl-extractable nitrate sediment pool remains to be tested. The significant variation in the vertical pattern and abundance of the various nitrate reducing genes phylotypes reasonably suggests differences in their activity throughout the sediment column. This raises interesting questions as to what the alternative metabolic roles for the various nitrate reductases could be, analogous to the alternative metabolic roles found for nitrite reductases.     

The effects of chemical pollution on the bioturbation potential of estuarine intertidal mudflats

Bioturbation by benthic infauna has important implications for the fate of contaminants as well as for changes to the sediment structure, chemistry and transport characteristics. There is an extensive literature dealing with the influence of sedimentary variables on the structure and function of infaunal marine and estuarine organisms but less is known of the converse, the influence of biota on sedimentary structure. Although some work has been carried out regarding spatial and temporal patterns of bioturbation, little attention has been given to the effects of pollution. The paper gives a framework of animal sediment relationships in an intertidal environment and discusses the general role of macrofauna in structuring and modifying sedimentary features. A brief outline of the various techniques used for quantifying the degree of bioturbation is given and some of these techniques have then been used to demonstrate the effect of a petrochemical discharge on the bioturbation potential of intertidal communities in the Humber estuary, eastern England. These studies indicate an increase in bioturbation with increasing distance from the source of pollution, not only because of differences in abundance, animal size and depth of activity but also because of the difference in species composition between the communities. As a means of interpreting the responses, the species present have been broadly classified in terms of their feeding strategy and sediment modification potential. The paper concludes by discussing the potential impact, in terms of effect on sediment transport, of selectively removing the different guilds (by pollution). Received: 8 February 1999 / Received in revised form: 10 May 1999 / Accepted: 14 May 1999     

Filter-feeding in the polychaeteNereis diversicolor: a review

An overview of recent findings concerning filter-feeding inNereis diversicolor is given. It has been discovered that the facultative filter-feederN. diversicolor may meet its metabolic requirements on a pure diet of phytoplankton, just as a typical obligate filter-feeder. Consequently, this worm is likely to be a hitherto undervalued key organism in the control of phytoplankton production in many shallow brackish water areas.     

Bacterial abundance and aerobic microbial activity across natural and oyster aquaculture habitats during summer conditions in a northeastern Pacific estuary

We measured sediment properties and the abundance and aerobic metabolism of microbes in Willapa Bay, Washington, USA, to test the response of sediment microbes to oyster aquaculture. Sites spanned the estuary gradient (practical salinity units ranged from 24 to 30 under seasonally low river flows) and six different low-intertidal habitat types: eelgrass (Zostera marina), unstructured tideflat, oyster hummocks (reefs of Crassostrea gigas), longline oyster aquaculture, hand-picked on-bottom oyster aquaculture, and dredged on-bottom oyster aquaculture. Aerobic metabolism was assessed by sole-source carbon use (SSCU) of 31 carbon sources on Biolog plates. Sediments generally became siltier and more organically enriched into the estuary, but no consistent differences in sediment properties occurred across habitat types. Bacterial cell density tracked organic content. Across the estuary gradient, overall aerobic SSCU increased less steeply than bacterial cell density, possibly as anaerobic metabolism became more important. Across habitats, aerobic SSCU differed significantly in both overall metabolism and diversity of carbon sources. Aerobic metabolism was generally lower for sediment microbes from intertidal on-bottom oyster aquaculture than from eelgrass. Humans indirectly alter microbial activity through biogenic habitats created during aquaculture, but, as has been shown for bivalves more generally, these changes were relatively small, particularly in comparison to sediment changes along estuarine gradients.     

Stress in estuarine communities: Lessons from the highly-impacted Bilbao estuary (Spain)

The highly-impacted Bilbao estuary was studied toquantify the response of estuarine communities tostress. Total heterotrophic biomass, a measure ofsediment infauna, declined sharply from the mouth tothe upper reaches of the estuary. Differentstatistical methods selected the depletion ofdissolved oxygen (DO) at the bottom of the watercolumn as the main correlate with the observedpattern. Biomass size-spectra (BSS) of estuarinecommunities suggested that `unimpacted' sites at themouth of the estuary had a high proportion of biomassin the larger size classes under regular andpredictable environmental conditions. In contrast, inthe impacted middle and inner parts of the estuarybiomass was concentrated in the smaller size classes,indicating a clear effect of stressful environmentalconditions on larger bodied organisms.A progressive increase of DO values has beenforecasted by the local Water Authority after thecompletion of a sewerage plan for the whole estuary,and predictions of water quality criteria were used toestimate community biomass figures following thismanagement. In order to predict the recovery at thelevel of individual species, several bioassays withthe native species Nereis diversicolor andScrobicularia plana were performed. Results obtainedwith the latter, more-sensitive species, suggest thatchemical contamination in sediments from the Bilbaoestuary has the potential to cause deleterious effectson indigenous benthic organisms, which could delayfaunal recovery in some parts of the estuary oncewater-oxic conditions improve.     

USING QUANTITATIVE PCR TO DETERMINE THE DISTRIBUTION OF A SEMICRYPTIC BENTHIC DIATOM,NAVICULA PHYLLEPTA (BACILLARIOPHYCEAE)1

Evidence has accumulated during the last decade showing that many established diatom morpho‐species actually consist of several semicryptic or truly cryptic species. As these species are difficult or even impossible to differentiate by microscopic analysis, there is virtually no information on how they behave in natural environments. In this study, we developed a quantitative real‐time PCR (qPCR) assay using TaqMan probes^® targeted to the internal transcribed spacer 1 (ITS1) to assess the spatial distribution and seasonal dynamics of an important component of the microphytobenthos of intertidal sediments. Navicula phyllepta Kützing is a brackish‐marine morpho‐species with a cosmopolitan distribution. Axenic clones of this species were isolated from natural assemblages of benthic diatoms at different intertidal stations in the Westerschelde estuary (The Netherlands). At least two distinct semicryptic species of N. phyllepta were present, as shown by differences in the quantity of DNA per cell, the ITS1 sequences and the copy number of ITS per cell. DNA and chl a concentrations extracted from sediment surface samples were closely correlated, showing that the DNA used for subsequent analysis mostly belonged to the microalgal community. The results of real‐time qPCR from sites throughout the estuary and over several seasons agreed well with microscopic counts. Additionally, the seasonal pattern of the two forms of N. phyllepta showed an overlapping, but unique distribution along the estuary.     

The hydrologic environment of Pool 19 of the Mississippi River

Invertebrate populations and biomass were studied over four years (1978–1981) in a coastal, saline lagoon in eastern England. Both temperature and salinities fluctuated sharply, with salinities as high as 72.4 gl⁻¹. Of 14 taxa recorded, only three occurred in significant numbers. Nereis diversicolor exhibited summer peaks over all four years, but there was a gradual trend of decline. Corophium volutator showed peaks in 1978 and 1979, but was largely absent thereafter. Chironomus salinarius showed a peak in numbers in 1978, but was rare in 1979 and largely absent in the following two years. The peak biomass was more than 4 000 mg m-2 in 1978, but was less than half this in subsequent years, when it consisted mainly of N. diversicolor. Laboratory experiments tended to confirm that high summer salinities were the cause of poor invertebrate success. The controlled ingress of brackish water into the lagoon from the adjacent estuary should result in a greater and more predictable production of invertebrates, which form the food supply of the chicks of an important colony of avocets Recurvirostra avocetta.     

Interpretation of the fossil record of Acanthaster planci from the Great Barrier Reef: a reply to criticism

The nature and interpretation of the fossil record of Acanthaster planci from the GBR is reviewed in the light of comments from Keesing et al. (1992) and Pandolfi (1992). Skeletal remains of A. planci in reef-top sediment of many reefs has been derived from very large numbers of individuals, indicating substantial, long-term mortality at reef-top locations. The fossil record provides useful perspective on mortality patterns in the absence of substantive ecological data. The incidence of skeletal elements on reefs where they are abundant cannot be adequately accounted for by the mortality of non-outbreak populations as estimated by recent surveys. Analysis of all available data reaffirms a relationship between the incidence of skeletal elements in surface sediment and observed outbreak history. There is no presently identified taphonomic mechanism by which the accumulation of A. planci skeletal elements released on death might be systematically biased relative to other skeletal components of reefal sediment. Because of skeletal degradation, physical transport and extensive bioturbation that applies in shallow-water reefal sediment, reconstructive taphonomic analysis of A. planci skeletal remains is not achievable. Core sediment, on which interpretation of the longterm fossil record of A. planci is based, is homogeneous, unstratified, and has experienced substantial time averaging due to pervasive bioturbation. Extensive bulk sediment dating has shown that the cores have retained a general age structure but fine-scale stratigraphic detail, required for the recognition of outbreak events from the fossil record available in reefal sediment is unlikely. As required by the principle of simplicity, the proposition that abundant A. planci skeletal elements found in sediment from Green Island, John Brewer and other reefs of the GBR represent the time-averaged product of outbreaking populations should be adopted as the favoured working hypothesis. Other alternative explantions have been advanced but all require patterns or processes that have yet to be substantiated.     

Selectivity of Perinereis aibuhitensis (Polychaeta,Nereididae) feeding on sediment

Deposit feeding polychaetes play an important role in the acceleration of the biogeochemical processes of the sediment through bioturbation. Feeding is one of the important factors of bioturbation. However, knowledge of the feeding biology of polychaetes, especially the subsurface deposit feeder, is limited. The objective of this study is to characterize the feeding selection of Perinereis aibuhitensis with different body weights. The animals were fed with natural sediment from their original habitat in the lab. The feed intake and particle size of sediment were measured to find any evidence of feeding selection. A two-way ANOVA showed that the particle size class significantly affected the particle size frequency of the ingested and the remaining sediment (P?P?>?0.05). Bivariate correlation analysis showed that the particle size frequency of sediment, ingested and remaining sediment in different size classes were linearly related (P?P?P?P. aibuhitensis preferred smaller particles in the sediment, which was limited by the particle size distribution of the sediment in which they live. The nitrogen and organic carbon contents in the faeces might be the thresholds when P. aibuhitensis selects sediment particles as food. These results demonstrated the particle selectivity of P. aibuhitensis, and may be applicable to other subsurface deposit feeding polychaetes.     

Lugworm exclusion experiment: Responses by deposit feeding worms to biogenic habitat transformations

On six 400 m² plots over 3 years, we excluded the sessile subduction and conveyer-belt feeding polychaete Arenicola marina which generates a pit-and-mound topography at the sediment surface from intertidal sands near the island of Sylt, Germany. This experiment was used to test whether other abundant deposit feeding polychaetes (the discretely motile and surface feeding ragworm Nereis diversicolor and the subsurface-feeding, motile orbiniid polychaete Scoloplos cf. armiger) benefit from competitive release. Ragworms took advantage from the absence of lugworms. Presumably they responded to a more stable and nutritious surface layer at lugworm exclusion plots (relief from inhibitive bioturbation). Contrary to this, S. cf. armiger was negatively affected by the exclusion of A. marina. It may have suffered from higher sulfide concentrations in the less irrigated and less permeable sediment where lugworms were absent. For adult worms of both species these results were consistent in 2 out of 3 years examined. Recruitment by N. diversicolor was highly variable between years and occurred either irrespective of experimental treatments or the response was inconsistent. Juveniles of S. cf. armiger benefited from the presence of A. marina and aggregated near lugworm tail shafts where inflow of oxygen rich water was high and sulfide concentrations were low. Biogenic habitat mediated effects of lugworms on both deposit feeders were in the same order of magnitude as abundance variation in space and time. Thus, A. marina was one of the key factors structuring the deposit feeding community. It is suggested that arenicolids modify the composition of the associated polychaete assemblage primarily through habitat transformation.