Effects of mud sedimentation on lugworm ecosystem engineering

Benthic ecosystem engineering organisms attenuate hydrodynamic or biogeochemical stress to ameliorate living conditions. Bioturbating infauna, like the lugworm Arenicola marina, determine intertidal process dynamics by maintaining the sediment oxygenated and sandy. Maintaining the permeability of the surrounding sediment enables them to pump water through the interstitial spaces, greatly increasing the oxygen availability. In a field experiment, both lugworm presence and siltation regime were manipulated to investigate to what extent lugworms are able to cope with sedimentation of increasing mud percentage and how this would affect its ecosystem engineering. Fluorescent tracers were added to experimentally deposited mud to visualise bioturbation effects on fine sediment fractions. Lugworm densities were not affected by an increasing mud percentage in experimentally deposited sediment. Negative effects are expected to occur under deposition with significantly higher mud percentages. Surface chlorophyll a content was a function of experimental mud percentage, with no effect of lugworm bioturbation. Surface roughness and sediment permeability clearly increased by lugworm presence, whereas sediment erosion threshold was not significantly affected by lugworms. The general idea that A. marina removes fine sediment fractions from the bed could not be confirmed. Rather, the main ecosystem engineering effect of A. marina is hydraulic destabilisation of the sediment matrix.     

Distribution and abundance of Abarenicola affinis (Arenicolidae,Polychaeta) in tidal inlets of Otago,New Zealand

Lugworms are important members of the macrobenthos of sheltered intertidal sediments, and this study assessed the abundance and distribution of Abarenicola affinis in Otago, southern New Zealand. Using a faecal cast:lugworm ratio (1.00:1.09) revealed a patchy distribution across inlets, with an overall mean abundance of 11.1 individuals/m² (range: 3.8 to 21.3 individuals/m²). Distribution patterns varied among inlets, with lugworms mostly confined to the high intertidal zone, but spread across all intertidal zones in one inlet. The lugworm habitat was characterised by muddy to medium sand (55–233 µm mean grain size) with low organic matter (< 2%) and chlorophyll a (1–12 µg/g dry sediment) contents. Depending on the inlet, lugworm abundance was related to sediment parameters, indicating that their distribution across Otago inlets is habitat dependent. The present findings also indicate that factors other than those measured contribute to A. affinis distribution and abundance patterns in this region.     

Factors affecting PAH residues in the lugworm Arenicola marina, a sediment feeding polychaete

Being a sediment feeder, the lugworm Arenicola marina could be a suitable candidate organism for the biomonitoring of sediment-bound contaminants in intertidal areas. Since lugworms are only rarely used in environmental monitoring, little is known about the natural variation in their body residue levels. In this study, we assessed the importance of seasonal fluctuations, spatial variation and sexual development to polycyclic aromatic hydrocarbon (PAH) residues in lugworms. Over a period of 15 months, lugworms were sampled along a contamination gradient in the Western Scheldt estuary in the Netherlands. A clear seasonal pattern in the body residue levels of PAH was observed, with lowest levels in March and highest levels just prior to the spawning season in September. During the spawning season the body residue levels showed a marked decrease. Although this seasonal pattern is apparently related to the reproductive cycle of the lugworm, no clear differences in body residue levels were found between animals with or without developed gonads. The contamination gradient, present in the estuary, was clearly reflected in the body residue levels of PAH. The expected gradient of internal concentrations was, however, absent in October, when the spawning period was not yet finished. The variation in lugworm body residue levels was smaller than the fluctuations in sediment PAH levels. It can be concluded that the lugworm is a suitable organism for the biomonitoring of sediment-bound contaminants, provided that attention is paid to its reproductive cycle.     

Small is beautiful: An inverted view of habitat fragmentation in seagrass beds

An intertidal Zostera marina landscape in Torbay, Devon, UK, was sampled to investigate the relationship between patch size, diversity and infaunal assemblage composition with the intention of defining a minimum Zostera patch size where the infaunal seagrass assemblage becomes distinct from the bare sand assemblage. All Zostera patches were found to support a higher level of biodiversity than the surrounding bare sand. However, the size of the Zostera patch had no impact on the level of diversity; it was just the presence or absence of seagrass that made a difference. The sediment and seagrass variables were not significantly different across the range of Zostera patch sizes, indicating that the environment characteristics were homogeneous within the Zostera patches at the patch scale. Multivariate analysis revealed that assemblage composition did vary between the patch types, although the opportunistic polychaete Capitella capitata was present in all patch types and was the most abundant species overall. The presence of opportunistic species and the homogeneity of the Zostera patch variables may be due to the location of this intertidal seagrass bed, which is relatively exposed compared to the locations of other seagrass beds along the south coast of Devon, resulting in a more dynamic and disturbed environment. Nevertheless, our results demonstrate that even small patches of seagrass comprising a few plants support a higher abundance and diversity of infaunal invertebrates than bare sand, indicating that Zostera patches have conservation value whatever their size.     

Sediment segregation by biodiffusing bivalves

The selective processing of sediment fractions (sand and mud; >63 μm and ≤63 μm median grain size) by macrofauna was assessed using two size classes of inert, UV-fluorescent sediment fraction tracers (luminophores). The luminophores were applied to the sediment surface in 16 m² replicated plots, defaunated and control, and left to be reworked by infauna for 32 days. As the macrofaunal assemblage in the ambient sediment and the control plots was dominated by the common cockle Cerastoderma edule, this species was used in an additional mesocosm experiment. The diversity, abundance and biomass of the defaunated macrobenthic assemblage did not return to control values within the experimental period. Both erosion threshold and bed elevation increased in the defaunated plots as a response to the absence of macrofauna and an increase in microphytobenthos growth. In the absence of macrobenthos, we observed an accretion of 7 mm sediment, containing ca. 60% mud. Image analysis of the vertical distribution of the different luminophore size classes showed that the cockles preferentially mobilised fine material from the sediment, thereby rendering it less muddy and effectively increasing the sand:mud ratio. Luminophore profiles and budgets of the mesocosm experiment under “no waves–no current” conditions support the field data very well.     

Effect of macrofaunal bioturbation on bacterial distribution in marine sandy sediments, with special reference to sulphur-oxidising bacteria

We have studied the impact of the bioturbating macrofauna, in particular the lugworm Arenicola marina and the bivalve Cerastoderma edule, on abundances and distribution patterns of total bacteria and of bacteria of selected functional groups in sandy intertidal sediments. The selected groups comprised the colourless sulphur-oxidising bacteria and the anoxygenic phototrophic bacteria, which are expected to occupy small zones at the oxygen–sulphide interface in stable (non-bioturbated) sediments. The presence of a wooden wreck buried in the sediment at 10 cm depth within a large area of intertidal sand flat colonised by lugworms provided a unique opportunity to confront field observations with laboratory simulations. The site with the wooden wreck, which was used as control site, was devoid of both A. marina and C. edule, while the composition of the rest of the zoobenthic community was rather similar to that of the surrounding area. In the field, the density of total bacteria was approximately one order of magnitude higher in the control site than in the natural (bioturbated) site. This can be explained by the higher contents of silt and clay particles (higher surface-area/volume ratio) and higher total organic-carbon contents found at the control site. It appears that the presence of macrofauna affects sedimentation processes, which indirectly influence bacterial dynamics. Samples from the control site have been incubated in the laboratory with A. marina and C. edule added (bioturbated core), while an unamended core served as a control. The laboratory experiments contrasted with the field observations, because it was found that total bacteria were actually higher in the deeper layers of the bioturbated core. Moreover, the populations were more homogeneous (less stratified) and colourless sulphur bacteria were on average less numerous in the bioturbated core. In general, laboratory incubations resulted in a decrease of total bacteria with a concomitant increase of colourless and phototrophic sulphur-oxidising bacteria and thus in modifications of the bacterial community structure. Hence, our results demonstrate that care must be taken in extrapolating results from laboratory experiments (e.g. mesocosm research) to field situations.     

Eiders Somateria mollissima scavenging behind a lugworm boat

The eider is one of the most important molluscivorous birds in the Wadden Sea, where it feeds mainly on blue mussels Mytilus edulis and edible cockles Cerastoderma edule. These prey species are within reach of the birds at all times. Other potential prey of suitable size that are abundantly present, such as several polychaete worms, or the clam Mya arenaria, are taken to a much lesser extent, possibly because they live buried in the sediment and digging them out would take too much effort. Mya may pose another problem because they grow to sizes that prevent eiders from swallowing them. Large Mya also live too deep down in the sediment, but young (small) specimens should be available to eiders. Yet, even these have only rarely been found as prey in eiders in the Wadden Sea. However, diet studies in relation to food abundance have been few, and may have missed prey that do not leave large shell fragments (i.e. in faeces studies). This paper describes observations on eiders taking both Mya and polychaete worms. The eiders fed on these prey in a fashion reminiscent of gulls that scavenge behind fishing vessels: some eiders have learnt to follow professional worm-digging boats that supply a bycatch of molluscs (mainly Mya arenaria) and polychaete worms (mainly Arenicola marina and Nephtys hombergii). Mya and worms were also the main targets of the eiders that fed in a dense flock close to the boat's stern. Faeces found on the flats at low tide comprised mainly cockle shell fragments, a prey rarely taken by the eiders behind the boat. Faeces studies may thus give a highly biased impression of local eider diet.     

Influence of Naineris laevigata (Polychaeta: Orbiniidae) on vertical grain size distribution,and dinoflagellate resting stages in the sediment

Short-term experiments were performed with the polychaete Naineris laevigata Grube, a conveyor-belt deposit feeder, to evaluate: the daily sediment reworking rate, the influence of the activity of this burrowing worm on the vertical redistribution of the sediment and dinoflagellate cysts. We also tested the germination success of cysts after their passage through the gut of the polychaete. Vertical particle distribution was studied in small aquaria with and without worms and with an initially layered sediment. After 30 days, sediment cores were collected in controls and treatments at 3 different depths (upper, intermediate and deep) and the diameter of 100 particles from each depth was measured to obtain the size-frequency distribution. The burrowing activity of 3 worms may completely mix a pre-existent layering within 30 days. The translocation of a mud layer from 4 cm depth towards the surface was observed. The mud layer deposited on the surface by the end of the experiment was at least 1 cm thick.Daily average production of faecal pellets was 0.443±0.032 g per worm. From the first day of the experiment, the particle size distribution of the faecal pellets released at the surface revealed the presence of sediment and cysts originating from underlying layers. Tests showed that the germination percentage of the cysts significantly diminished after passage through the guts of the worms.Our results indicate that N. laevigata is an effective vertical mixer of sediment and plays a role in the vertical transport and germination success of resting stages.     

Effects of the invasive polychaete Marenzelleria spp. on benthic processes and meiobenthos of a species-poor brackish system

The effects of the invasive polychaete Marenzelleria spp. on sediment processes and meiobenthos with an emphasis on free-living nematodes of the Vistula Lagoon (southern Baltic Sea) were investigated in a laboratory microcosm experiment. Marenzelleria occupies an open niche and its deep burying behaviour and feeding strategy represent a new function in the study area. Halos of oxidized sediment along Marenzelleria burrow walls indicated oxygen penetration into the burrows but the polychaete had no significant effect on porewater nutrient concentrations. The results showed, however, the density dependent effects of Marenzelleria on ammonium transport. An enhanced ammonium efflux was recorded at high polychaete densities (2000 ind. m^− 2) but not at low polychaete densities (300 ind. m^− 2).There was no observable impact of the polychaete on total meiobenthic numbers. There was, therefore, no indication that Marenzelleria caused meiofauna mortality. On the contrary, the polychaete significantly affected vertical distribution of meiofauna facilitating the colonization of deeper sediment depths and thus extending the habitat to be used by meiobenthos. In addition, Marenzelleria had a positive impact on the survival of turbellarians.Nevertheless, there was no effect of Marenzelleria on nematode assemblage structure and diversity, indicating that neither the physical presence nor the biological activity of the worm affected the nematode community. This suggests either 1. the limited impact of Marenzelleria on nematodes, resulting from the creation of simple, narrow and un-branched burrows, 2. poor response of nematode community resulting from their low abundance and diversity in the study area, or 3. the overriding role of the harsh chemical environment typical of sediments of the Vistula Lagoon, masking the effect of the bioturbator.     

Effects of laughing gull and shorebird predation on the intertidal fauna at Cape May,New Jersey

The intertidal flats of the Cape May, New Jersey shore of Delaware Bay are populated by large numbers of laughing gulls and migrating shorebirds during the spring and early summer. Exclusion of birds from a shallow slough and a sand bar had only minor effects on the infaunal benthic invertebrate assemblage at either site. The Cape May beaches provide a rich source of food in the form of horseshoe crab (Limulus polyphemus) eggs; foraging on this item may be more profitable than probing the sediment for infauna. Gemma gemma, a small, thick-shelled bivalve, composed over 98% of the benthic infauna at both sites in 1980, and this species may be resistant to predation by certain shorebirds, as suggested by Schneider (1978).     

Spatial intertidal distribution of bivalves and polychaetes in relation to environmental conditions in the Natori River estuary,Japan

This paper aims to reveal spatial variation in the abundance of infaunal bivalves and polychaetes at different spatial scales (station: 200–800 m intervals; plot: 5–20 m), and to reveal environmental variables affecting the spatial distribution of animals in the Natori River estuary, Japan. We found six bivalve species and eight polychaete species from 52 plots at 12 stations. Nuttallia olivacea and Heteromastus sp. were found to be the most abundant species of bivalves and polychaetes, respectively. Assemblage patterns of bivalves and polychaetes were classified into five distinct groups. Substrata (silt-clay contents), salinity, and relative elevation were the variables found to affect the infaunal assemblage patterns. Chlorophyll a was not a significant variable, but benthic animals were absent at sites with extremely low chlorophyll a conditions. Macrobenthic assemblage patterns were different not only between stations but often differed between plots at the same station, reflecting the complex assemblage structure of benthic invertebrates. Detailing such animal–environment relationship is essential in understanding the potential food supply for estuarine fishes.     

Impact of lugworms (Arenicola marina) on mobilization and transport of fine particles and organic matter in marine sediments

We investigated the impact of sediment reworking fauna and hydrodynamics on mobilization and transport of organic matter and fine particles in marine sediments. Experiments were conducted in an annular flume using lugworms (Arenicola marina) as model organisms. The impact of lugworms on sediment characteristics and particle transport was followed through time in sediments experimentally enriched with fine particles (< 63 μm) and organic matter. Parallel experiments were run at low and high water current velocity (11 and 25 cm s^− 1) to evaluate the importance of sediment erosion at the sediment–water interface. There was no impact of fauna on sediment composition and particle transport at current velocity below the sediment erosion threshold. At current velocity above the erosion threshold, sediment reworking by lugworms resulted in dramatic particle transport (12 kg dry matter m^− 2) to an adjacent particle trap within 56 days. The transported matter was enriched 6–8 times in fine particles and organic matter when compared to the initial sediment. This study suggests that sediment reworking fauna is an important controlling factor for the particle composition of marine sediments. A. marina mediated sediment reworking greatly increases the sediment volume exposed to hydrodynamic forcing at the sediment–water interface, and through sediment resuspension control the content of fine particles and organic matter in the entire reworked sediment layer (> 20 cm depth).     

Sublethal effects of oiled sediment on the sand worm, Nereis (neanthes) virens: Induced changes in burrowing and emergence

The burrowing and emergence behavior of sand worms, Nereis (Neanthes) virens Sars, in sediment contaminated with sublethal concentrations of Prudhoe Bay crude oil ranging from 74 to 5222 ppm, was studied in the laboratory. Initial burrowing in oiled sediment did not differ from that observed in unoiled sediment. Emergence of exposed worms was related to the oil concentration and the extent of weathering of oiled sediment. Sand worms buried in unoiled sediment did not emerge. Worms dug from oiled sediment after 12 hours' exposure and placed on unoiled sediment were impaired, resulting in a significant increase in time to burrow. However, after 12h, they recovered and burrowed at normal rates. Worms held in oiled sediment for 96 h also recovered. Possible causes for recovery are discussed. Results suggest that oil-induced aberrations may increase vulnerability to predation.     

Actinian dominated intertidal mudflats: A new case of an extraordinary rare phenomenon from Southern Chile

Generally, estuarine intertidal mudflats constitute important nurseries for fish and foraging grounds for coastal birds by providing a plenitude of mollusks, worms, and crustaceans as prey, which in turn mostly feed on suspended and benthic microalgae, bacteria, and detritus. Despite the high productivity of such habitats, pronounced variability in both salinity and temperature results typically in low diversity. The only sea anemone reported from estuarine mud is the edwardsiid Nematostella vectensis Stephenson, 1935. It occurs widely in the northern hemisphere, and occasionally in extremely high density. Here we document another sea anemone from estuarine mud and muddy sand found in Southern Chile which has similar ecological attributes. Taxonomic confusion has impeded the reporting on this small but prominent member in a macrozoobenthic assemblage, the brooding Anthopleura hermaphroditica (Carlgren, 1899; Anthozoa: Actiniidae). It differs from N. vectensis by the presence of symbiotic algae. Average density under poly- to euhaline conditions in mud and muddy sand at around mid tide level was about 3 actinians per cm². An average abundance of 11,000 m^? 2, a biovolume of 487 cm³ m^? 2, and a biomass of 35.5 g dry organic weight m^? 2 were found in mud and muddy sand in two surveys 20 years apart. The mean fishing area of fully expanded individuals covers 42 ± 25 mm², corresponding to a circular area with a diameter of 7.3 ± 5.7 mm. Preliminary experiments indicate that associated benthos may be relegated to life below surface by the net of tentacles above the sediment. As no predators on A. hermaphroditica could be found on the mudflat, the success of this mixotrophic sea anemone may entail a trophic dead end.     

Effects of seed predators and light level on the distribution of Avicennia marina (Forsk.) Vierh. in tropical,tidal forests

Avicennia marina has an unusual distributional pattern across the intertidal region in mangrove forests of north-eastern Australia. It is often abundant in the low intertidal and high intertidal but is very rare in mid-intertidal areas. The influences of shade intolerance and seed predators on the distribution of Avicennia were investigated. Field observations of 391 light gaps indicated that while seedlings were equally abundant in gaps versus under the neighboring forest canopy, saplings were much more abundant in gaps. Subsequent field studies showed that seedlings survived and grew best in light gaps as opposed to under the forest canopy. Further field experiments revealed that 96·0±3·4% of the post-dispersal propagules of Avicennia are consumed by seed predators, primarily grapsid crabs. Predation on propagules was less in low and high intertidal regions where conspecific adults were common and was highest in the mid intertidal where Avicennia is rarest. Predator exclusion experiments indicated that Avicennia was capable of growing in mid-intertidal areas. It is concluded that the combination of shade intolerance and extensive predation on its' propagules effectively limits the distribution of Avicennia marina across the intertidal and may account for the dominance by members of the Rhizophoraceae in the mangrove forests of northern Australia.     

Distinctly variable mudscapes: Distribution gradients of intertidal macrofauna across the Dutch Wadden Sea

The Wadden Sea is a shallow coastal region, with a large area of sedimentary tidal flats that extends from The Netherlands to Denmark and has been declared a site of international importance in the Dutch and German parts (Ramsar status and UNESCO World Heritage Site). Benthic macrofauna are central to the ecosystem functioning of this area, as they recycle nutrients, decompose organic matter and are an important food source for many secondary consumers, like fish and waterbirds. Due to the environmental gradients characteristic of estuarine systems, it is expected that changes in assemblage composition will be observed across the physical and environmental gradients of the Wadden Sea. First, we explored the spatial variation in assemblage composition of benthic macrofauna across the intertidal part of the Dutch Wadden Sea using 3 years of biomass data. Then, we identified the relative importance of six environmental variables for explaining and predicting changes in assemblage composition across the intertidal areas of the Wadden Sea using generalised dissimilarity modelling (GDM). In accordance with the environmental gradients across this system, the biomass contributed by a few common species differed from west to east and were distinct in the Dollard. In the west, bivalves Mya arenaria, Cerastoderma edule and Ensis directus contributed a relatively large and equal share of the total biomass, whereas C. edule contributed the sole largest share of the total biomass towards the east. The polychaete Alitta succinea became a large share of the total biomass in the upper Ems and in the Dollard estuary, but contributed little elsewhere. Similar to the observed differences in species composition, the spatial patterns in assemblage composition, as predicted by the GDM models, identified the Dollard as distinct and that the prevalence of assemblage types in the west differed to the east. Median grain size, followed by microphytobenthic biomass, and exposure time were the most important variables describing differences in assemblage composition. That the Wadden Sea forms a heterogeneous landscape where assemblage composition varies across multiple gradients has repercussions for management and monitoring.     

Macrofaunal sediment selectivity considerations for beach nourishment programmes

Nowadays, beach nourishment is widely considered as a better alternative compared to the construction of hard structures to protect a sandy coast against detrimental erosive effects, both from an ecological and an engineering perspective. The rare studies conducted on the ecological impact of beach nourishment are short-term, post hoc monitoring investigations of the benthic macrofauna. Little is known of the biological processes during and after nourishment. To allow swift recolonization after nourishment, the characteristics of the nourished beach have to match the habitat demands of the benthic macrofauna. The sediment preference of the key intertidal species Scolelepis squamata, Eurydice pulchra, Bathyporeia pilosa and Bathyporeia sarsi, which dominate many West European sandy beaches, was investigated through laboratory experiments, both in single-species as well as combined-species treatments. While the former aimed at developing guidelines for impact mitigation of beach nourishment, the latter aimed at elucidating the role of biotic interactions in sediment preference. Results of the experiments indicated that B. pilosa and E. pulchra prefer the finest sediment, while B. sarsi had a broader preference and also occurred in medium-coarse sediments. However, the sediment preference of E. pulchra for fine sediments was not confirmed by other field and experimental studies. The polychaete S. squamata had the broadest preference and even showed a high occurrence in coarse sediments that are not naturally occurring on the sandy beaches where the animals were caught for this experiment. However, this polychaete is a cosmopolitan species, not only occurring on fine-grained beaches, but also on coarse-grained beaches worldwide. The preferences imply that beach nourishment with coarse sediment will have a major effect on B. pilosa while effects of coarse sediments on S. squamata will be minor. Finally, interspecific competition with the sympatrically occurring amphipod B. sarsi was found to change the sediment selection of the amphipod B. pilosa towards the coarser sediments where B. sarsi occurred in lower frequencies.     

The effect of the dominant polychaete Scolelepis squamata on nematode colonisation in sandy beach sediments: An experimental approach

The effect of an abundant sandy beach polychaete, Scolelepis squamata, on the colonisation of defaunated sediments by marine nematodes indicates that sandy beach fauna can be partially controlled by biological interactions within and across size groups. Experimental cores, equipped with windows allowing infaunal colonisation, were filled with defaunated sandy beach sediment containing two different treatments with and without S. squamata. These cores were inserted into microcosms filled with sediment with indigenous meiofauna collected from the field. The treatments were incubated in the laboratory at ambient temperature and salinity for 2, 7, 14 and 21 days, in order to follow the colonisation process of the defaunated sediments by the indigenous nematode fauna over time. Nematodes initially colonised both treatments, with abundances of up to 10% of the densities in the control; after 2 weeks, nematode densities in the cores without S. squamata surpassed the control densities. Nematode assemblages in both treatments were not species rich, and also differed in composition from the natural assemblages. The most successful colonising species, Enoplolaimus litoralis, was rare in the surrounding sediment, suggesting that colonisation was determined by species-specific characteristics such as body size, motility and feeding strategy. Initially the presence of macrofauna did not affect the nematode community composition, but after 2 weeks of the experiment, the presence of the polychaete seemed to facilitate the earlier establishment of non-opportunistic species.     

Spatial Overlap and Coexistence in a Mussel-Associated Polychaete Assemblage on a South American Rocky Shore

Abstract. Mussel beds in the intertidal of subtropical South America are known to harbour a large number of invertebrates, particularly polychaetes. This paper presents a quantitative analysis of the patterns of spatial overlap and coexistence in an assemblage of polychaetes associated with mussel beds in a rocky intertidal habitat on the Pacific coast of Peru. Mussel beds are made up of two zones in the intertidal: the lower, more extensive zone formed by Semimytilus algosus and the upper zone formed by Perumytilus purpuratus . Null models were employed to assess the magnitude of spatial overlap between pairs of taxa encompassing a total of seven taxonomic groups. Two taxa belonging to the same functional group tended to be spatially segregated in the Semimytilus bed: Pseudonereis vs. Halosydna (large free-ranging predators) and Scoloplos vs. Mediomastus (deposit feeders), with further segregating tendencies between Pseudonereis and Mediomastus , and between Halosydna and Scoloplos . Small-sized, free-ranging ' Typosyllis ' and the bottom-dwelling scavenging predator Lumbrineris were also spatially associated. Thus, this polychaete assemblage appears to be loosely organised around groupings of taxa which are somewhat different in ecological traits and form mosaic patterns of distribution with reduced overlap within the Semimytilus bed. These relationships were weak to non-existent in the upper, drier Perumytilus habitat where polychaete density was low.     

Tube-forming polychaetes enhance invertebrate diversity and abundance in sandy sediments of Mozambique,Africa

In marine soft-bottom systems, polychaetes can increase habitat complexity by constructing rigid tubes (e.g. several onuphid species) that contrast with surrounding topographically flat sediments. These structures can provide predation refuges and increase larval settlement and thereby increase the richness and abundance of fauna. We collected invertebrate samples from an intertidal flat with low onuphid tube density (2.7 m^–2) in Mozambique and document that more organisms (70 times higher mollusc abundances) and more species (15 times more mollusc species) were found associated with solitary tubes of an onuphid polychaete compared with surrounding sand habitats. These results are in agreement with tube versus sand comparisons from soft-bottom systems in the North Atlantic where polychaete tube densities are often much higher.