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.     

Impact of irrigation on oxygen flux into the sediment: intermittent pumping by Callianassa subterranea and “piston-pumping” by Lanice conchilega

O₂-flux into sediments attributed to the pumping behaviour of two macrofauna species, Callianassa subterranea (Decapoda) and Lanice conchilega (Polychaeta) was investigated. Samples were obtained from the North Sea near Helgoland in 1989 and 1990. The two species were found to transport roughly similar amounts (3 mmolm^-2d^-1) of oxygen into the sediment although they displayed markedly different pumping behaviours. Irrigation by C. subterranea was intermittent and characterized by regularly recurring breathing currents which lasted 2.6 min and were separated by 40-min pauses. In addition to this regular intermittent irrigation, an irregular mode was observed. C. subterranea constructed a complex burrow system. At least half of the burrow wall was not in contact with oxygenated water, however, and thus not effective as additional interface for O₂-exchange. Sediment expelled from the burrow increased the total oxygen uptake (TOU) relative to the surrounding sediment surface. L. conchilega moved water much more frequently (every 4 min) than C. subterranea. We suggest that L. conchilega acted as a piston when moving in its tube, exchanging burrow water with the overlying water. This mechanism, termed piston-pumping, is also potentially important in other smaller tube dwelling organisms. At a shallow water station in the southern North Sea 21 ind of C. subterranea constructed 1.6 m² burrow surface per m². L. conchilega (300 ind m^-2) created only 0.37 m²m^-2 tube surface. On the basis of the abundance and oxygen transport associated with pumping activity, it is calculated that the two species increase TOU by 85% compared to O₂-flux across the sediment-water interface.     

Molecular association of Cu,Zn, Cd and 210Po in the digestive gland of the squid Nototodarus gouldi

In July 1985, very fine burrows extending at least to a sediment depth of 50 cm were found in sediments of the Vöring-Plateau, Norwegian continental slope, at depths of 1 261 to 1 969 m. Such burrows have been described from other areas of the oceans, but until now the producers have been unknown. In the box-cores from Vöring-Plateau sipunculans of the genus Golfingia, subgenus Nephasoma, were identified as inhabitants and producers of these Lebensspuren, which correspond with the fossil ichnogenus Trichichnus.     

Ecological implications of the presence of the tube-building polychaete <Emphasis Type="Italic">Lanice conchilega</Emphasis> on soft-bottom benthic ecosystems

The common tube-building polychaete Lanice conchilega is known as a habitat structuring species and can form dense aggregations. The effects of L. conchilega on the surrounding benthic community have received little attention, especially in subtidal areas. Therefore, the presence of L. conchilega in different habitats in the North Sea and its effect on the abundance, species richness, diversity and community structure in these habitats are evaluated in the present paper, based on data from the ICES North Sea Benthos Survey of 2000. Lanice conchilega has a wide geographical distribution and a low habitat specialization, but optimally occurs in shallow fine sands. In the present study, the presence of L. conchilega resulted in a density increase and a significant (positive) correlation of the benthos density with the density of L. conchilega. Furthermore, the species richness (number of species) increased with increasing density of L. conchilega. This trend was, however, not consistent: the number of species reached more or less an asymptotic value or even decreased after reaching a critical density of L. conchilega (>500–1,000 ind/m2), as observed in shallow fine sands. The same overall pattern was detected concerning the expected number of species. The N ₁-diversity index showed similar or slightly higher values in L. conchilega patches compared to patches without L. conchilega. From the results of the community analysis, it can be concluded that the species, which were responsible for the increase of the diversity, belonged to the overall species-pool of that habitat. The effects on density and diversity differed between the four discerned habitats (shallow muddy sand, shallow fine sand, shallow medium sand and deep fine sand), and were most pronounced in shallow fine sands. These patterns can be attributed to the habitat structuring capacity of L. conchilega. The mechanisms responsible for the increase of the habitat quality in patches of L. conchilega can be summarized as (1) changes in the hydrodynamics, (2) increases of the habitat stability and oxygen supply, and (3) a creation of habitat heterogeneity in a uniform environment. In this way, L. conchilega alters the habitat characteristics and affects other organisms, and can therefore even be considered as an ecosystem engineer. In other words, L. conchilega patches are responsible for an increased habitat quality in an otherwise uniform habitat, which results in a higher survival of the surrounding benthic species.     

Mating system of the burrowing crab Neohelice granulata (Brachyura: Varunidae) in two contrasting environments: effect of burrow architecture

Few studies conducted in crustaceans have demonstrated how habitat features could shape the mating systems. Here, the burrow of Neohelice granulata was considered as a resource used for mating, and its architecture was characterized in two contrasting study sites: Mar Chiquita Lagoon (MCL), an estuary composed of muddy sediments, and San Antonio Oeste (SAO), a marine bay composed of gravel sediment. Burrow features differed between study sites and occupant gender. Large males constructed burrows with a chamber in MCL and with a widened entrance in SAO, while small males constructed the same narrow burrows as females at both study sites. Field experiments demonstrated that burrows with chambers or widened entries are places used for copulation, although successful post-copulatory guarding was displayed only in those with chambers. The intensity of the agonistic encounters and the success of males in winning resources (burrows/females) also depend on the habitat characteristics. N. granulata shows a resource defense mating system where males employ different mating strategies according to the burrow architecture to ensure mating success.     

Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Negative interactions between two SW Atlantic intertidal crabs in soft-bottom habitats

The intertidal crabs Chasmagnathus granulatus and Cyrtograpsus angulatus coexist across the SW Atlantic intertidal. Previous studies in this region suggest that C. granulatus displace C. angulatus in soft sediment, where C. granulatus build burrows (“burrowing beds”). We examined variation in abundance, size-frequency distribution, sex ratio, incidence of autotomies, and diet of both species in C. granulatus crab beds and adjacent areas without burrows. We also experimentally tested the hypothesis that in the absence of C. granulatus, C. angulatus will build burrows. Only large sized individuals of C. angulatus venture into C. granulatus crab beds. The sex ratio of C. angulatus was always biased towards females, with higher bias outside crab beds (1:8 outside, 1:2 inside). Although the items consumed in the crab beds did not differ from those consumed outside, the males of C. angulatus had a higher frequency of empty stomachs in crab beds. The incidence of limb autotomies of C. angulatus was higher outside C. granulatus crab bed areas. After a long rainy period in which C. granulatus was absent from these areas, the pattern of habitat use of C. angulatus changed. During this period C. angulatus showed higher abundance in the areas, where C. granulatus previously constructed burrows, and there were no differences between areas in any of the measured parameters. In the absence of C. granulatus, C. angulatus built their own burrows and never used C. granulatus burrows. The interaction between C. granulatus and C. angulatus may be a good example of competitive exclusion, when the shared resource is the access to surface soft-sediment.     

Burrow structure, burrowing and feeding behaviour of Corallianassa longiventris and Pestarella tyrrhena (Crustacea, Thalassinidea, Callianassidae)

The behaviour of the Caribbean Corallianassa longiventris and the Mediterranean Pestarella tyrrhena, two burrowing thalassinideans, was studied in situ and in laboratory aquaria. Burrows of C. longiventris were closed most of the time; they consist of a deep U (down to 1.5 m) with upper and deeper chambers, some of them filled with macrophyte debris. The burrows of P. tyrrhena reached down to a maximum depth of 54 cm and consisted of a shallow U with a mound and a funnel, and a spiral shaft from which several, often debris-filled chambers branched off. The appearance of C. longiventris at the sediment surface to collect debris is strongly triggered by wave swell or odours from plant and animal juices; its burrows are opened within 10 min. The surface activity of P. tyrrhena was relatively less frequent and less predictable. Inside the burrows, both species exhibited different patterns of time allocated to 25 defined behavioural states. After being offered seagrass debris, P. tyrrhena spent relatively less time manipulating this debris, but it handled sediment more often than C. longiventris. During frequent mining events, both species showed sediment-sorting behaviour, which brought a parcel of sediment in close contact with the mouthparts; some of this sediment may be ingested because the fecal rods produced by both shrimps contain very fine sediment particles. Seagrass debris is irregularly tended by P. tyrrhena after its introduction into the chambers. Such material ultimately becomes buried. Corallianassa longiventris frequently returns to its debris chambers to pick up pieces of seagrass, which are subsequently cut with the chelae or ripped with the third maxilliped and then transported to another empty chamber nearby. Pieces become smaller with time and show curved cutting edges and bite marks. After 100 to 140 days, 2 to 6 g(dw) seagrass debris are consumed in this manner by individuals of this species. The debris-related behaviour of P. tyrrhena probably enriches the sediment around the burrow for stochastic encounters during later mining events. Such an indirect benefit may also be effective on a population level because other individuals may also encounter this buried nutrient source. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Toxiwasp∗

TOXIWASP combines most of the kinetic structure of EXAMS 2 with the transport capabilities of WASP (Water Analysis Simulation Program). TOXIWASP uses variable chemical degradation rates from chemical properties and the environmental conditions of the aquatic ecosystem. These rates are reduced from pseudo first‐order rates to first‐order rates including the processes hydrolysis, biotransfor‐mation, phototransformation, oxidation, and volatilisation. Assuming ultimate local equilibrium, and using a chemical dependent partition coefficient as well as spatially varying environmental carbon fractions, sorption onto sediments and biomass is calculated. Environmental alternations could be specified in any time scale by providing monitoring data.

TOXIWASP generates total sediment and chemical concentrations every time step in every segment, including surface water, subsurface water, surface bed and subsurface bed. Advection, dispersion, mass loading, sedimentation, and scour affect sediment concentration in the water column and in the bed sediment concentrations depend on burial and erosion. In addition chemical concentrations are influenced by degradation, sediment‐water dispersion, and percolation. Lateral transport of chemical within the bed is neglected and transport data are not calculated in the program. TOXIWASP is developed to model stratified lakes, reservoirs, large rivers, estuaries, and coastal waters. As for EXAMS 2 (Burns et al.²) the TOXIWASP user has to accept the model's inability to connect the water body to a chemically contaminated atmosphere.     

Mesocosm and in situ observations of the burrowing shrimp Calocaris templemani (Decapoda: Thalassinidea) and its bioturbation activities in soft sediments of the Laurentian Trough

In vivo observations in laboratory mesocosms and aquaria, accompanied with in situ photographic surveys, have shown that the burrowing shrimp Calocaris templemani has a significant impact on bottom sediment dynamics and geochemistry in the St. Lawrence Estuary. This burrowing shrimp establishes and maintains complex semi-permanent burrows made up of several interconnected, ‘U-shaped’ galleries with generally four or more openings to the sediment surface. In the Estuary, at 345 m depth, Calocaris average density was estimated at 3.4 individuals m^?2. Observed individual burrows reached a maximum volume of 0.54 L. C. templemani displaces this volume of mostly anoxic sediments from the subsurface layers (down to 15 cm) to the sediment surface, thereby obscuring some of the natural stratification patterns. With an estimated turnover rate of about 8 L m^?2 year^?1 of sediment, our calculations suggest that over a period of about 18.75 years, all the sediment to a depth of 15 cm will have been reworked by C. templemani alone.     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.     

Fiddler crab burrowing affects growth and production of the white mangrove (<Emphasis Type="Italic">Laguncularia racemosa</Emphasis>) in a restored Florida coastal marsh

Positive plant–animal interactions are important in community ecology, but relatively little attention has been paid to their effect on the production of mangroves, dominant halophytic trees in tropical coastal marshes. Here, the role of fiddler crab (Uca spp.) burrowing on the growth and production of the white mangrove, Laguncularia racemosa (<2 years old), was examined in a restored marsh in Tampa Bay, Florida (27°41.65 N, 82°30.34 W) with manipulative experiments from June 2006 to May 2007. Fiddler crab burrowing significantly increased mangrove height by 27%, trunk diameter by 25%, and leaf production by 15%, compared to mangroves in crab exclusion enclosures. Additionally, the exclusion of fiddler crabs significantly increased interstitial water salinity from 32.4 to 44.2, and decreased the oxidation–reduction potential of the low organic sediments, but did not affect soil pH or sulfide concentration. Mangrove height, trunk diameter, and leaf production along a transect that varied in crab burrow density were positively associated with the number of crab burrows. Further, the density of sympatric Spartina alterniflora shoots was positively correlated with crab burrow density along the transect. As in temperate marshes, fiddler crabs can have significant ecological effects on mangrove communities, serving as ecological engineers by modulating the amount of resources available to marsh plants, and by altering the physical, chemical, and biological state of these soft sediment communities. In restored coastal systems that typically have very poor sediment quality, techniques such as soil amendment could be used to facilitate a more natural interaction between crabs and mangroves in ecosystem development.     

Active swimming in meiobenthic copepods of seagrass beds: geographic comparisons of abundances and reproductive characteristics

The entry of meiobenthic copepods from sediments or seagrass blades into the water column and reproductive characteristics of actively migrating fauna were investigated from 1981–1986 in a temperate intertidal Zostera capricorni seagrass bed in Pautahanui Inlet, New Zealand and in a subtidal Thalassia testudinum bed in Tampa Bay, Florida, USA. Emergence of copepods in New Zealand varied over a tidalcycle, while in Florida a distinct diel periodicity was displayed. Selected copepod species in New Zealand had similar numbers emerging from sediments and/or blades over a 6 h period as the common copepods actively migrating from sediments in Florida. Daily abundances of emerging copepods (24 h) in Tampa Bay, Florida, were substantially greater than those in New Zealand, where migration is linked to tidal cover. In Z. capricorni meadows in New Zealand, sex ratios of copepods in sediments and on blades were dominated by females; males dominated water-column samples. In T. testudinum meadows in Tampa Bay, sex ratios of males to females, although of a lowermagnitude than in Z. capricorni beds, were higher in trap than in sediment samples. Differences in sex ratios, the availability in emergence traps of females of appropriate stage for mating, and observations on clasping in live samples from traps suggest that swimming behavior in copepods may be partly linked to prenuptial courtship. Meiobenthic copepods may use the water column as an important habitat for reproductive behavior.     

Reproductive biology of a filter-feeding amphipod, Leptocheirus pinguis , with extended parental care

Leptocheirus pinguis (Stimpson, 1853) is a widely distributed, abundant, endobenthic amphipod that engages in extended parental care, i.e. females host their juveniles in their burrows for extended time periods. I examined reproduction and population biology of L. pinguis at mean low water (MLW) in muddy sediment in Lowes Cove, Maine, USA. Cores around individual burrows were taken monthly in 1994 and 1995, and four seasonal samples were taken at different tidal heights. During the major reproductive periods in spring/early summer and in the fall, females produced several consecutive broods and hosted growing offspring in their burrows. Juveniles remained in their mothers' burrows until they reached a length of 5 mm (approximately one-third adult size) or more. At the study site, the majority of amphipods in individual burrows were adult females. Following the main reproductive periods, subadult individuals were found in their own burrows, but densities did not increase following the reproductive period in spring/early summer 1994, probably because large numbers of L. pinguis emigrated via the water column between June and December. L. pinguis is an annual species. Many members of the cohort born in spring/early summer start reproducing in the fall, and survive until the following spring when they produce several broods. Members of the cohort born in the fall start reproducing the following spring and also produce several consecutive broods. Both the spring/early summer and fall cohorts die off after the major reproductive period in the following spring/early summer. High standing stocks of microphytobenthos occur in soft-bottoms at MLW, and I conclude that L. pinguis can engage in extended parental care there because its food is abundant year-round. The limited expandability and low stability of burrows in soft-bottoms at MLW do not permit long persistence of parent–offspring groups in L.␣pinguis. Received: 27 March 1997 / Accepted: 30 July 1997     

Behavior and growth ofMercenaria mercenaria during simulated storm events

Experiments were conducted in April–August 1989 on juvenileMercenaria mercenaria (L.) in an oscillatory water tunnel to simulate resuspension of bottom sediments by waves and to determine the effects of shortterm storm events on particle ingestion, pseudofeces production, and shell growth. Juveniles (mean length = 19.2 mm) were subjected to identical concentrations of algae in both low-flow, gentle waves (maximum velocity = 7 cm s^–1) and high-velocity storm waves (maximum velocity = 22 cm s^–1). Suspended sediment levels reached 193 mg 1^–1 at 1 cm above the bed during storms. Shell growth decreased by a maximum of 38% during the storm when levels of phytoplankton were high (average cell concentration = 43 × 10⁶ cells 1^–1), and by 18% when phytoplankton levels were low (av cell conc = 6 × 10⁶ cells 1^–1). Orientation of clam siphons was not related to flow direction. Significantly more pseudofeces were produced when the clams were subjected to increased sediment resuspension under waves, and in troughs of sand ripples. The size of sediment grains ingested did not vary significantly among the flow treatments. The decrease in shell growth during storms may be due to a reduction in filtration rate coupled with a decrease in net energy gained from filtration due to costs of pseudofeces production. The magnitude of the decrease seems to be related to concentration of algae, water temperature, age of clams and sediment transport mode (bed load or suspended load). Thus, the interpretation of growth increments must be made in the context of these environmental variables.     

Ammonium regeneration and assimilation in eelgrass (Zostera marina) beds

Regeneration and assimilation of ammonium in the water column and in sediments of eelgrass (Zostera marina L.) beds of Izembek Lagoon and Crane Cove, Alaska, USA and Mangoku-Ura, northeastern Japan, were investigated by using a ¹⁵N isotope dilution technique. In the water column of Mangoku-Ura, ammonium was regenerated at a rate of 12 nmol l^-1 h^-1 and assimilated at a rate of 74 nmol l^-1 h^-1. The ammonium regeneration rate in sediments ranged from 2 to 150 nmol g^-1 h^-1, and with one exception, exceeded ammonium assimilation in sediments (0.3 to 77 nmol g^-1 h^-1). The ammonium regeneration in the water column was of little significance for the nitrogen supply to the eelgrass bed ecosystem. Net ammonium production (regeneration minus assimilation) in the sediment of Izembek Laggon met nitrogen demand for eelgrass growth, suggesting that ammonium regeneration in the sediments was very important for the nitrogen cycle in the eelgrass bed ecosystem.     

Observation of bioturbation and hyporheic flux in streambeds

In the Elkhorn River, burrows, tubes, and sediment mounds created by invertebrate bioturbation were observed in the exposed streambed and commonly concentrated on the fine-sediment patches, which consist of silt, clay, and organic matter. These invertebrate activities could loosen the thin layer of clogging sediments and result in an increase of pore size in the sediments, leading to greater vertical hydraulic conductivity of the streambed (K _v ). The measurements of the vertical hydraulic gradient across the submerged streambed show that vertical flux in the hyporheic zone can alter directions (upward versus downward) for two locations only a few meters apart. In situ permeameter tests show that streambed K _v in the upper sediment layer is much higher than that in the lower sediment layer, and the calculated K _v in the submerged streambed is consistently greater than that in the clogged sediments around the shorelines of the sand bars. Moreover, a phenomenon of gas bubble release at the water-sediment interface from the subsurface sediments was observed in the groundwater seepage zone where flow velocity is extremely small. The bursting of gas bubbles can potentially break the thin clogging layer of sediments and enhance the vertical hydraulic conductivity of the streambed.     

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.     

Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3–28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2–1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

Reproductive biology of a deposit-feeding amphipod, Casco bigelowi, with extended parental care

The reproductive traits of a deposit-feeding amphipod that engages in extended parental care were examined. At the study site in Lowes Cove, Maine, USA, Casco bigelowi (Blake, 1929) occurred in highest densities in soft sediments just below mean low water (MLW). During most months, the sex ratio was ≃1. Many females hosted males in their burrows throughout the summer, but after fertilization of females in September, all adult males disappeared from the study area. In October almost 80% of the females were ovigerous, and in November about the same percentage was parental, i.e. caring for juveniles in their burrows. The females produced only one brood each in late fall which they accommodated in their burrows for 2 mo or longer. The average number of juveniles per female was ∼20 in November, and continuously decreased until January. Juveniles reached sizes >10 mm length in the maternal burrows. In early December the first juveniles were found in their own burrows, but major recruitment took place in late December and January. It is concluded that for C. bigelowi, the delay of recruitment into the winter months with low predation pressure and the large offspring size at this time are major advantages gained by extended parental care. C. bigelowi is host to the peritrich ciliate Cothurnia sp. on its gills, and during the summer months >70% of all amphipods had ciliates on their gills. Juveniles still living in their mother's burrows showed infestation rates similar to that of the parent; those of highly infested mothers were more heavily infested than those of “clean” mothers. Facilitated epibiont transmission during intimate and long-lasting (2␣mo) parent–offspring associations may be a consequence of extended parental care. Received: 25 November 1997 / Accepted: 14 April 1998