Changes in the feeding behavior of the deposit-feeding polychaete<Emphasis Type="Italic"> Laeonereis acuta</Emphasis> on soft sediments inhabited by burrowing crabs

In this paper, we evaluate the influence of the burrowing crab Chasmagnathus granulatus on the feeding behavior, body condition, burrowing activity, and fecal production of the deposit-feeding polychaete Laeonereis acuta. Previous results and ours show that, due to crab activity, sediment organic matter decreases with depth outside a crab bed, but there were no differences inside. Also, particle sizes were smaller inside the crab bed. Polychaetes showed better body condition inside the crab bed, suggesting higher energy gain. They fed on the surface more frequently outside the crab bed than inside. However, feeding rate was higher inside the crab bed, which is consistent with the larger weight and volume of fecal pellets found inside crab bed. In both areas, the bedload sediment transport was not similar to the sediment ingested by this polychaete. A 1-month exclusion/inclusion experiment performed outside and inside the crab bed showed that the body condition of polychaetes changed between areas but that this was not directly due to crab manipulation. Thus, our results suggest that the higher sediment quality inside a crab bed can positively affect the feeding behavior of the deposit-feeding polychaete L. acuta, increasing its feeding rate and consequently enhancing its body condition.Communicated by P.W. Sammarco, Chauvin     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

Density affects female and male mate searching in the fiddler crab, <Emphasis Type="Italic">Uca beebei</Emphasis>

In most species, only one sex searches for mates while the other waits. Models of sex-specific mate-searching behavior predict single-sex searching, but the factors that determine which sex searches are not understood. In this study, we examine the effects of density and predation risk on mate-searching behavior in the fiddler crab Uca beebei. U. beebei is one of the few fiddler-crab species in which both sexes search for mates. In a field experiment conducted in Panama, we manipulated crab density and perceived predation risk in replicate plots. Females searched more and males searched less at high densities. At high levels of perceived predation risk, both sexes similarly reduced their search rates. Observations of plots that naturally varied in crab density show that females were more likely to search for mates in areas of higher density, where there were more males. Females may preferentially search for mates in high-density areas because the abundance of nearby burrows, into which they can run to escape predators, decreases their costs of searching and because the abundance of males and male burrows facilitates comparisons and thus may increase their benefits from searching. Males at high densities decrease their mate-searching rate perhaps in response to the increase in female searching and to the corresponding increase in the intensity of their competitors' mate-attraction signals.     

Abundance,population structure and claw morphology of the semi-terrestrial crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis> (Fabricius, 1787) on shores of differing wave exposure

Wave action is known to influence the abundance and distribution of intertidal organisms. Wave action will also determine the duration and suitability of various foraging windows (high-tide and low-tide, day and night) for predation and can also affect predator behaviour, both directly by impeding prey handling and indirectly by influencing prey abundance. It remains uncertain whether semi-terrestrial mobile predators such as crabs which can access intertidal prey during emersion when the effects of wave action are minimal, are influenced by exposure. Here, we assessed the effect of wave action on the abundance and population structure (size and gender) of the semi-terrestrial intertidal crab Pachygrapsus marmoratus on rocky shores in Portugal. The activity of P. marmoratus with the tidal cycle on sheltered and exposed shores was established using baited pots at high-tide to examine whether there was activity during intertidal immersion and by low-tide searches. Because prey abundance varies along a wave exposure gradient on most Portuguese shores and because morphology of crab chelipeds are known to be related to diet composition, we further tested the hypothesis that predator stomach contents reflected differences in prey abundance along the horizontal gradient in wave exposure and that this would be correlated with the crab cheliped morphology. Thus, we examined phenotypic variation in P. marmoratus chelipeds across shores of differing exposure to wave action. P. marmoratus was only active during low-tide. Patterns of abundance and population structure of crabs did not vary with exposure to wave action. Stomach contents, however, varied significantly between shores of differing exposure with a higher consumption of hard-shelled prey (mussels) on exposed locations, where this type of prey is more abundant, and a higher consumption of barnacles on sheltered shores. Multivariate geometric analysis of crab claws showed that claws were significantly larger on exposed shores. There was a significant correlation between animals with larger claws and the abundance of mussels in their stomach. Variation in cheliped size may have resulted from differing food availability on sheltered and exposed shores.     

Production of glycine betaine and dimethylsulfoniopropionate in marine phytoplankton. II. N-limited chemostat cultures

The influence of salinity on the time elapsed between two successive molts and the size reached after each molt were studied at 30, 21, 12 and 3‰S in juveniles of two co-occurring grapsid species, Cyrtograpsus angulatus and C. altimanus, cultured under identical conditions of temperature, photoperiod and food. Juvenile growth patterns were compared between these species (which differ in size-at-maturity and maximum size). C. angulatus grew faster than C. altimanus, reflecting a higher increment per molt and a shorter intermolt period. A significant difference existed between the number of instars preceding the size of maturity in both species: >11 in C. angulatus, 6 in C. altimanus. There was evidence of a differential effect of low salinity on growth. By the end of the experiment, individuals of both species were smaller at the lowest salinity (3‰) tested; the largest crabs developed at 21‰ (C. angulatus) and 30‰ (C. altimanus). The size difference between the “optimal” and the less suitable salinities in the sixth crab instar was 12.4% in C. angulatus and 35% in C. altimanus. During early juvenile development (Crab Instars 1 to 4), there were slight differences in intermolt period among salinities in C. angulatus, but large differences in C. altimanus. The longest intermolt period of C. altimanus was at 3‰S and the shortest at 30‰S. In the following instars (5 to 10 in C. angulatus and 5 to 6 in C. altimanus), the longest intermolt period occurred at 21‰S, the shortest at 3‰S, in both species. Interspecific differences in response to low salinities may explain why C. angulatus occurs throughout a whole temperate coastal lagoon, whereas C. altimanus is restricted to its mouth. Received: 12 July 1998 / Accepted: 24 February 1999     

Environmental stress,competition and dominance of Crassostrea virginica near Beaufort,North Carolina,USA

Observations and experiments were made at 2 intertidal areas near Beaufort, North Carolina, USA from May 1977 to July 1978 to determine why the oyster Crassostrea virginica dominated the community in areas protected from wave action but not in areas directly exposed to waves. Barnacles, oysters, the green alga Enteromorpha sp. and the mussel Brachiodontes exustus were the main occupiers of primary space at the mid and low intertidal levels of exposed areas. The intertidal community at the protected site consisted of a mid intertidal occupied by the barnacles Balanus amphitrite and Chthamalus fragilis, and the oyster C. virginica, and a low intertidal dominated by C. virginica. The exposed area was highly variable with high colonization and mortality for all species producing large seasonal changes in structure. The protected site remained constant throughout the year; there was no evidence of further colonization of either barnacles or oysters and mortality was very low. Recolonization experiments, selective removal of species, and growth and survival data demonstrated that C. virginica does not become dominant at exposed locations because (1) the constant wave shock at the ocean site reduces growth and increases mortality of young and adult oysters and (2) oysters are outcompeted by the mussel B. exustus. The monopolization of space by C. virginica at protected sites contrasts with studies north of Cape Hatteras where the abundance of predators produces a more diverse and heterogeneous community. Predation was unimportant in Beaufort because predators were absent at the exposed areas and the oyster drill Urosalpinx cinerea was restricted to the subtidal zone at the protected site. This absence of predators indicates a higher level of environmental stress near Beaufort compared to areas farther north.     

Variations in benthic community structure in a sub-tropical estuary

Abundances of macroinfaunal invertebrates in sand dollar (Dendraster excentricus) beds and in adjacent areas free of sand dollars were studied at 10 intertidal sites in the Pacific Northwest region of the USA and Canada. Each site was sampled once in late summer of 1977. There were no significant differences in overall diversity of species between the beds and adjacent areas; nor did cluster analysis indicate a sand dollar bed community. However, a polychaete (Armandia brevis) was significantly more abundant inside the beds than just outside them. Conversely, in comparison to the beds, adjacent regions contained significantly more specimens of the following species: a bivalve (Transennella tantilla), several tubicolous crustaceans (Corophium spp., Ampelisca agassizi, and Leptochelia savignyi) and two polychaetes (Glycinde polygnatha and Malacoceros arenicola). Sediment characteristics at each study site were not correlated with the presence or absence of sand dollars; moreover, a comparison among the 10 sites revealed no sediment characteristics typical of sand dollar beds. At most of the sites, significantly fewer tube-building invertebrates occurred inside sand dollar beds than outside. The discussion considers the possibility that mechanical disturbance by the sand dollars tends to exclude some macroinvertebrates (especially tubicolous ones) from sand dollar beds.     

Food web analysis in intertidal <Emphasis Type="Italic">Zostera marina</Emphasis> and <Emphasis Type="Italic">Zostera noltii</Emphasis> communities in winter and summer

The food web of two intertidal seagrass (Zostera marina and Zostera noltii) beds that may be influenced by the seasonal variation in food source abundance was studied in winter and in summer with δ¹³C and δ¹⁵N analysis. In spite of high relative variation of abundance of main primary producers at the two sites, the food web did not vary between winter and summer. The δ¹³C range of primary producers was wide. Zostera leaves, the most ¹³C-enriched source, were not consumed directly by grazers. Deposit and filter feeders have a similar δ¹³C and could use a mix of suspended and sedimented organic particulate matter, largely composed of detritus from macroalgae to seagrass. This trophic pathway allows the local incorporation of the high biomass produced by seagrasses. The wide δ¹⁵N range of predators was linked either to a large variety from omnivore to carnivore predators or to the also wide ranges of δ¹⁵N of primary consumers.     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

Influence of predation by a crab on the distribution of the size-groups of three intertidal gastropods in South Australia

Two South Australian rocky intertidal platforms were sampled in April 1981 and December 1982 (Marino Rocks) and in December 1982 (Lady Bay), respectively. Three snail species, Nerita atramentosa Reeve, 1855, Bembicium nanum (Lamarck, 1822) and Austrocochlea concamerata (Wood, 1828), exhibited shore-level size-gradients, with smaller individuals occupying the higher intertidal levels. The reef crab (Ozius truncatus M.-Edwards, 1834), whose distribution overlaps that of the gastropods, is an important predator of these gastropods. The shore-level distribution of shell sizes can be explained by differential selection against smaller individuals by reef crabs. The decreased density of small snails at lower intertidal levels may be the result of either their consumption by reef crabs and/or their movement to higher shore levels where crab predation is less intense.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Biotic resistance to green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, in California bays

The distribution of the introduced European green crab, Carcinus maenas, was investigated in the central California embayments of Bodega Bay Harbor (BBH), Tomales Bay, and Bolinas Lagoon using baited traps and snorkel surveys. Adult green crabs were very spatially limited in all three embayments and occurred primarily in warm, shallow areas that lacked large native Cancer spp. crabs. The green crabs that were found in closest proximity to populations of Cancer spp. exhibited high levels of limb damage and loss; damage was strongly correlated with low ratios of intertidal area: edge, indicative of narrow areas of intertidal that are more easily accessed by large Cancer spp. moving up to forage during periods of tidal inundation. Up to 70% of the green crabs tethered in areas of BBH that are utilized by Cancer spp. experienced limb loss, while those tethered in the marsh, where there are no Cancer spp., were undamaged. The results suggest that the potential distribution of green crabs in the northeastern Pacific will be far less than has been predicted, and that their impacts may be largely attenuated through predation by and competition with native crab species.     

Factors limiting distribution and activity patterns of the soldier crab Dotilla myctiroides in Phuket, South Thailand

This study was carried out between January and March 1995 on the intertidal sand flats of Tang Khen Bay, Phuket, South Thailand, where the soldier crab Dotilla myctiroides (H. Milne-Edwards) occurs in densities of up to 120 m⁻². In this bay, long, ribbon-like sand waves (wavelength 40 m, height 0.4 m) are interspersed with shallow pools, running approximately parallel to the shore. During daylight low-tides, exposure of the sand waves is followed 15 to 20 min later by the emergence of the crabs which have been buried under the sediment surface during high tide. Their subsequent burrowing and feeding activity results in the production of large numbers of sand pellets on the sediment surface. Most crabs retreat down their burrows, and some also plug the burrow entrance, prior to being covered by the incoming tide. The crab burrows have a distinct distribution on the sand waves. Burrows are most dense at the top of each sand wave, and a band of unburrowed sediment adjoins the adjacent tidal pools. Crabs are most abundant between mean high-water neap-tide level and mean low-water neap-tide level, where the median particle size of the surface sediment is ∼2 . Measurements of water-table depth below the sand waves and the exposure time of the sediment indicated that, where sediment size is suitable, the main factor controlling crab distribution is the duration of daytime exposure. This observation is in contrast to those of many previous studies, which have suggested that water-table height and sediment drainage are the main factors controlling the distribution of D. myctiroides. Received: 14 January 1998 / Accepted: 6 May 1999     

How a subsocial intertidal beetle,Bledius spectabilis,prevents flooding and anoxia in its burrow

Summary All stages of Bledius spectabilis Kratz (Staphylinidae) dig wine-bottle shaped burrows in the intertidal Salicornia zone. The adult female lays her eggs around her burrow and, by remaining with them, prevents both flooding by the tide and anoxia. Flooding is prevented by an ever-ready burrow which exploits a surface tension effect and by blocking the burrow once the tide comes in. The burrow has a narrow neck (about 2 mm in diameter) leading to a living chamber 5 mm in diameter. Tides over artificial tubes in agar showed that a critical minimum neck diameter of 2–3 mm prevented sudden flooding, giving time to block the neck with mud. Blocking took about four minutes. Ensuring the burrow is reopened at each low tide is a vital role of the brooding female in anaerobic or impermeable soils. Field models of burrows became anoxic in 4 days, much less than the 4 week long egg stage. A lab model system with anaerobic agar and calculations of oxygen uptake and diffusion supported this conclusion. Mortality of orphaned larvae may be much lower, however, in burrows within large Bledius aggregations because of mitigating good drainage and soil aeration: larval mortality from physical causes in these burrows did not increase over 3 weeks but without the mother, 14% of the eggs were attacked by mould, and two burrows were taken over by a predatory carabid, Dichierotrichus gustavi, and all larvae eaten. The surface tension effects of small air-filled openings may be used by small air-breathing animals in many intertidal habitats. Like Bledius, other intertidal animals, including large ones, may block their burrows at high tide to keep them full of air. Maternal care, in particular the combination of behaviours which protect the brood from the tide and anoxia, enables this airbreathing insect to colonise the inhospitable habitat of the intertidal saltmarsh.     

Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Ultrastructural changes in the gill epithelium of the crab Chasmagnathus granulatus (Decapoda: Grapsidae) in diluted and concentrated seawater

Chasmagnathus granulatus Dana, 1851 is an intertidal estuarine crab that experiences acute salinity changes ranging from <1‰ to full-strength seawater and even hypersaline waters in tide pools concentrated by evaporation. Ultrastructural changes induced by salinity in the posterior gills were examined in crabs collected from the Rio de la Plata estuary Argentina during March 1999. The posterior gills of C. granulatus are involved both in ion uptake and ion secretion depending on the acclimation medium. These organs are mostly lined with a thick tissue, which presents the characteristics of a typical salt-transporting epithelium. Electron microscopy analysis of gill tissue from crabs acclimated to dilute, full, and concentrated seawater (12‰, 34‰, and 45‰ salinity) showed significant development of basolateral membrane interdigitations, with numerous mitochondria and conspicuous apical membrane infoldings. Morphometrical analysis indicated that the subcuticular space delimited by the infolding of the apical membrane was significantly increased in the gills of high-salinity acclimated crabs. Septate junctions, which are thought to define the paracellular permeability, were significantly shorter in high-salinity acclimated crabs, suggesting a possible role of the paracellular pathway in salt secretion.     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Feeding ecology of the mangrove crab <Emphasis Type="Italic">Ucides cordatus</Emphasis> (Ocypodidae): food choice,food quality and assimilation efficiency

Food preferences, consumption rates and dietary assimilation related to food quality were investigated for the large semi-terrestrial and litter-consuming mangrove crab Ucides cordatus cordatus (Ocypodidae, L. 1763) in northern Brazil. Stomach contents were composed of mangrove leaves (61.2%), unidentified plant material and detritus (28.0%), roots (4.9%), sediment (3.3%), bark (2.5%), and animal material (0.1%). U. cordatus prefers Rhizophora mangle over Avicennia germinans leaves despite a higher nitrogen content, lower carbon to nitrogen (C/N) ratio and lower tannin content of the latter. Also, assimilation rates for senescent R. mangle leaves (C: 79.3%, N: 45.4%) were higher than for A. germinans leaves (C: 40.6%, N: 9.1%). Faeces composition indicates that A. germinans leaves were more difficult to masticate and digest mechanically. The leaf-ageing hypothesis, according to which crabs let leaves age in burrows to gain a more palatable and nutritive food, was rejected for U. cordatus since N content, C/N ratio and the abundance of microorganisms did not differ significantly between senescent leaves and leaves taken from burrows. The low microbial biomass on leaf surfaces and in the sediment indicates its minor importance for the nutrition of U. cordatus. It is concluded that high ingestion and assimilation rates of a R. mangle diet together with the consumption of algae allow for a high intake of C, N, and energy. The data suggest that the digestibility of mangrove leaves by U. cordatus is not hampered by tannins. This may have provided a competitive advantage over other leaf-consuming invertebrates unable to digest mangrove litter with high tannin concentrations. Due to the large stock biomass of U. cordatus in the study area, a great amount of finely fragmented faeces is produced (about 7.1 ton dry matter ha⁻¹ year⁻¹ in a R. mangle forest) which is enriched in C, N and bacterial biomass compared to the sediment. The decomposition of mangrove litter, and thus nutrient and energy transfer into the sediment, is greatly enhanced due to litter processing by U. cordatus.     

Microbial reaction rates and bacterial communities in sediment surrounding burrows of two nereidid polychaetes (Nereis diversicolor and N. virens)

The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO₂ and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.