Algal “gardening” behavior by nereid polychaetes: Effects on soft-bottom community structure

Nereid polychaetes (Nereis vexillosa and Platynereis bicanaliculata) attach pieces of drift algae to their tube surfaces. The presence of permanent algal cover increases the predictability of the food supply for at least the herbivores (including the nereids) and the deposit feeders, and modulates the temperature and salinity stresses of the marine intertidal soft-bottom environment. However, it may affect the access of organisms to the oxygenated water layers above the sediment surface. This is true for polychaetes that live head downwards in vertical tubes. It is suggested that the attachment behavior of the nereid polychaetes increases the ability of the plant to colonize habitats both temporally and spatially.     

Influence of predation on infaunal abundance in Upper Chesapeake Bay,USA

The importance of predators in controlling the densities of infaunal (>0.5 mm) organisms was investigated in the mesohaline region of the Upper Chesapeake Bay (USA) using field experiments. The role of predators in controlling infaunal density and community characteristics varied with habitat type, season (i.e., predator abundance) and developmental or successional stage of the community. Few infaunal species were adversely affected by predator exclusion. Species that increased greatly in abundance in the absence of predators (e.g. Eteone heteropoda, Streblospio benedicti, Nereis succinea, and juvenile Macoma balthica and Mya arenaria) lived near the sediment-water interface and had major population pulses from fall through spring. Species whose abundances increased moderately or were not affected by predator exclusion were deeper burrowing organisms (e.g. Heteromastus filiformis and adult Mya arenaria), or were relatively small organisms (e.g. Paraprionospio pinnata, Scolecolepides viridis and Peloscolex gabriellae) whose principal predators could be other members of the infauna. Competition did not appear to be an important factor controlling infaunal density in these experiments.This work is Contribution No. 973 of the Center for Environmental and Estuarine Studies of the University of Maryland     

High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Effects of Polinices duplicatus (Gastropoda: Naticidae) on infaunal community structure at Barnstable Harbor,Massachusetts, USA

Effects of the predatory naticid snail Polinices duplicatus (Say) on species composition, diversity and density, of the infauna of intertidal sand-flats at Barnstable Harbor, Massachusetts, USA, were studied using field experiments. Responses of molluscs (prey of P. duplicatus) and nonmolluscs (nonprey) were considered separately to distinguish between the effects of feeding and sediment disturbance during foraging. The fauna of 0.25 m² predator-exclusion cages, coarse-mesh cages, sieved areas, and controls was followed for 1 yr. Species associations within cages from which predators were excluded were denser, more diverse, and richer in molluscs than those in other treatments. Larger areas (3×3 m) of natural bottom were fenced and maintained as snail exclosures and enclosures during two feeding seasons. For both molluscs and nonmolluscs, diversity (H'), number of species, evenness (SD), and density all decreased with increasing snail density. Intense predation pressure on molluscs and selective feeding on thin-shelled bivalves, which were rare, removed individuals and species of molluscs from the community. Comparison of samples taken inside and outside trails made by snails showed that disturbance of the surface sediment layers by snails decreased the abundance of spionid polychaetes and total nonmolluscs. The sipunculan Phascolopsis gouldi and the bivalve Gemma gemma dominated the community in all experimental treatments and were little affected by the activity of Polinices duplicatus. Predation and disturbance by snails lowered community diversity by removing individuals of the less abundant species, and generally maintained population densities below the level where strong competition would occur.     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

A description of a highly diverse soft-bottom community in Kingston Harbour,Jamaica

A highly diverse soft-bottom community in Kingston Harbour, Jamaica is described to provide additional information on the structure of tropical benthic communities. It is shown that the community has a level of diversity unique to stable tropical environments, that dominance by a few species is greatly reduced,and that there are no true parallels to this community in similar environments. The community described seems to fit Sanders' stability-time hypothesis. The density and standing crop of the community are low, and possible explanations for this are discussed. Continuous year-round breeding, short life-spans, and fast turnover rates are suggested as the most important factors regulating biomass in the community described.     

Single-species bacteria in sediments induce larval settlement of the infaunal polychaetes Polydora cornuta and Streblospio benedicti

Larval settlement of the infaunal spionid polychaetes Polydora cornuta and Streblospio benedicti is mediated by sediment-associated microorganisms. To investigate if larval preference for certain sediment is guided by individual sediment-associated bacteria, 13 bacterial isolates (5 phyla) obtained from the natural habitat of adult polychaetes (Wadden Sea, Germany) in 2008 were screened in still-water, no-choice settlement assays. Two isolates (α-Proteobacterium-Strain DF11 and Flavobacterium-Strain 54) significantly triggered larval settlement in comparison with sterile sediment. In still-water, multiple-choice settlement assays comprising natural and sterile sediment and sediment re-inoculated with isolates DF11 and 54, significant preferences for natural sediment and sediment containing bacterial isolates at 10⁸ cells g^?1 were observed. Larval settlement was influenced by bacterial abundance in sediment but the correlation was not strictly positive; thus, maximum larval settlement in response to single bacterial species may occur at certain optimum densities. Non-viable or suspended bacteria and water-soluble bacterial products did not induce larval settlement, suggesting that sediment-associated bacterial settlement cues for P. cornuta and S. benedicti were either produced in situ and/or consisted of heat-labile bacterial products.     

Immunological methods for food web analysis in a soft-bottom benthic community

Basic to nearly any prediction concerning the behavior and structure of entire communities or of their components is knowledge of trophic connections among species. A major impediment to such understanding of soft-bottom benthos is methodological. Because none of the routinely available methods of food web analysis (e.g. visual gut content analysis, direct observation of feeding, tracer techniques) is generally suitable for examining all trophic interactions of benthic infauna, we rought to evaluate the potential of immunological methods for identifying predatorprey relationships in one typical, estuarine, intertidal sand flat. Whole-organism extracts of individual macro- and meiofaunal taxa were injected into rabbits to produce antisera of varying specificity. Double immunodiffusion precipitin tests of antiserum specificity revealed both phyletic and trophic relationships among 20 taxa. Using relatively unspecific antisera, preliminary analysis of the stomach contents of a few surface deposit-feeders and particle browsers was successful, giving positive identification of several trophic links which would otherwise have gone undetected. The production of taxon-specific antisera is expected to provide the methodological tool necessary to document the breadth of trophic connections in a marine benthic food web.     

Effects of an unusual natural temperature increase on a baltic soft-bottom community

The effects of a sudden, extraordinary temperature rise upon a soft-bottom community in the northern Baltic Sea were studied continuously for more than a month, during the summer of 1968. The mean bundance of the total macrofauna on the muddy bottom amounted to ca 6×10³ individuals · m^-2, on the sandy bottom to 2×10⁴ individuals· m^-2. The meiofauna, only sampled in the mud, averaged 1×10⁶ individuals · m^-2, with nematodes, oligochaetes and ostracods as dominating groups. During this period, the redoxpotential-discontinuity (RPD)-layer successively moved upwards in the muddy bottom, being relatively stable in the sand. The mud surface showed a succession of microbial communities. The initially oxidized, yellow surface film, with patches of diatoms and red Spirula, changed to a more grayish layer, with purple areas here interpreted as Chromatium and white spots of Beggiatoa. The latter dominated during the last part of the period, covering vast areas of the bottom like a gigantic cobweb. The endofauna was forced upwards by the emerging RPD-layer, the more sensitive components dying off, facultative anaerobes and nematodes flourishing. The described temperature increase of about 10 C° simulates the effects of a 3000 MW nuclear plant.     

Influence of an artificial reef on the surrounding infaunal community

Artificial reefs have been constructed throughout the world, but their effects on adjacent soft-bottom communities are largely unknown. In December 1986, we investigated the influence of Pendleton Artificial Reef (PAR) in Southern California on the abundance of infauna in the surrounding sand bottom. PAR was constructed in 1980 of quarry rock placed in eight piles, or modules. The artificial reef altered the grain-size distribution of sediments around the reef; sediments close to the modules were coarser than those 10 or 20 m away from the modules. Densities of one of the two most common species, the polychaetePrionospio pygmaeus, were lower near the reef, perhaps due to foraging by reef-associated predators or because the habitat near the reef was less suitable. We found no evidence that foraging by reef-associated fishes caused a widespread reduction in infaunal densities near the reef, and in fact the other most common taxon,Spiophanes spp., had higher densities near the reef. The most conspicuous effect of the artificial reef concerned the tube-dwelling wormDiopatra ornata, which only occurred in close association with the modules. In addition, total infaunal density and the densities of decapods, echinoderms and sipunculids were higher withinD. ornata beds than outside the beds. These results indicate that the densities of some species were enhanced, and others depressed, around the reef, but that the overall effect of the artificial reef on the surrounding infauna was limited to a small area near the modules.     

Human-driven spatial and temporal shift in trophodynamics in the Gulf of Maine,USA

Utilizing marine protected areas (MPAs) to isolate the ecological effects of human influence can help us understand our effect on systems and foster ecosystem-based approaches to management. Specifically, examining invertebrate prey community dynamics inside and outside an MPA may provide a measure of how altering human influence (i.e., certain fishing pressures) affects ecosystem interactions. We measured trophic interactions inside and outside a deep-water temperate MPA over 2 years. Predation rates on tethered, preferred groundfish prey (crabs) were initially identical inside and outside the MPA, but decreased outside the MPA after the commercial groundfish fishing season commenced. Predation trials using a ubiquitous prey species (brittle stars) and a less preferred prey species (urchins) served as controls, showing no MPA effect. Our experiments quantify differential predatory activity resulting from differences in human activity driven by an MPA, demonstrating important effects of fishing and regulations on the strength of trophic interactions.     

Feeding and nutritional ecology of the sea urchin strongylocentrotus drobachiensis in Maine,USA

The sea urchin strongylocentrotus drobachiensis exhibited a high degree of food selectivity, whether foods were presented singly or in combination. Foods ranked from most to least preferred were, in summer, Laminaria longicruris, Chondrus crispus, Corallina officinalis, Ascophyllum nodosum, and Agarum cribrosum, whereas in winter A. nodosum and A. cribrosum exchanged ranks. Food preference was not correlated with caloric content but, because of higher feeding rates on preferred foods, caloric intake was positively correlated with preference. Similarly, food absorption rankings were not correlated with food preference, with the exception of gravimetric efficiencies in winter. However, the absorption of L. longicruris, the most preferred alga, was highest in all measurements. Growth and reproductive development of S. drobachiensis on single species diets were positively correlated with food preference. Highest values occurred with animals fed L. longicruris. Urchins transferred from non-preferred to preferred diets showed increased somatic and reproductive growth compared to control animals on the original, non-preferred diets. Conversely, when transferred from preferred to non-preferred diets, urchins showed reduced growth compared to controls. No combination diet tested supported significantly better gonadal growth than L. longicruris: 25% C. crispus supported slightly better somatic growth than L. longicruris alone. These data support the view that S. drobachiensis has evolved a feeding strategy resulting in the maximization of growth and reproduction.     

Response of a soft-bottom harpacticoid community to stingray (Dasyatis sabina) disturbance

The recolonization by harpacticoid copepods of patches disturbed by stingrays [Dasyatis sabina (LeSeur)] feeding in a subtidal (2–3 m depth) site off the Florida gulf coast (Lat. 29°54.55N, Long. 84°31.45W) was followed. Samples from disturbed patches could no longer be distinguished from control samples in terms of numbers of harpacticoids 29 h after disturbance. No species became significantly disproportionately abundant during the recovery suggesting that small-scale natural disturbances are not an important organizing force in this community.     

Intertidal community structure and oceanographic patterns around Santa Cruz Island, CA, USA

Recent studies suggest that nearshore oceanographic conditions can have important effects on the structure of benthic communities. On Santa Cruz Island (SCI), CA, USA there is a persistent difference in mean annual sea surface temperature (SST) around the island due to its location at the confluence of opposing cold and warm ocean current systems. Over the course of a 4-year study (1997–2001) seawater nutrient and chl-a concentrations, algal tissue C:N ratios, recruitment and growth of filter-feeders (barnacles and mussels), and intertidal community structure were measured at six intertidal sites around the island. There were strong associations between remotely sensed SST and patterns of community structure. Macrophyte abundance was highest at sites with persistently low SST, while recruitment, abundance, and growth of filter-feeding invertebrates were strongly, positively correlated with SST. The cold-water sites were associated with higher nutrient concentrations and lower algal C:N ratios, particularly in the winter months. Values of chl-a were generally low and variable among sites, and were not correlated with the predominant SST gradient. Recruitment of barnacles and mussels was positively correlated with adult abundance across all sites. While detailed experimental studies are needed to further evaluate the mechanisms underlying community dynamics, these results indicate that the confluence of cold- and warm-water masses around SCI may determine the contrasting patterns of intertidal community structure.     

Resource-partitioning and predation impact of a low-latitude myctophid community

This study, based on data collected during summer 1985 in the eastern Gulf of Mexico, examined the degree of overlap in two prime niche parameters, space and food, in 17 of the most abundant myctophid species which inhabit the epipelagic zone at night. Cluster-analyses of vertical distribution information and diet characteristics revealed that while large groups of species overlapped (>60%) in either vertical distributions or diet, when both niche parameters were considered together, little interspecific or intraspecific (size class) overlap occurred. Our data suggest that for myctophids trophic competition is reduced through resource-partitioning, although with considerable overlap at niche boundaries. Niche separation presumably is the result of competition during the evolution of the ecosystem and is maintained presently as diffuse competition: the effect on a species of the combined competition from all other species at that trophic level. We suggest that the large degree of niche overlap enables the packing of over 50 myctophid species in the epipelagic zone at night. Our calculations indicate that myctophid predation nightly removes 2% of the zooplankton biomass. Myctophid predation is selective in that greatest pressure is on certain size classes and types of prey (copepods, ostracods and euphausiids). In the case of copepods, impact is greatest on the larger, more mature stages and hence on the breeding population. It is estimated that myctophids account for at least one-third of the daily production of zooplankton removed from the epipelagic zone by micronekton in the eastern Gulf.     

Factors controlling lead bioavailability in the Butte mining district,Montana, USA

Microprobe analyses of 38 soil and 5 mine-waste samples from Butte, Montana, demonstrated that the samples contain predominantly sulphide/sulphate and oxide/phosphates of lead (Pb)-bearing phases associated with mine waste. The sulphide/sulphate assemblage consists primarily of galena altering to anglesite and plumbojarosite, with secondary jarosite precipitating and rinding the Pb-bearing minerals. In addition, galena was encapsulated within pyrite or quartz grains. The oxide/phosphate assemblage consists of pH-neutral soils in which a plausible paragenetic sequence of PbO to Pb phosphates, PbMnO, or PbFeO is proposed, dependent on the activity of P, Mn, Fe, and Cl in the soil. In addition, Pb-bearing grains are occasionally armoured by the presence of a 1- to 3-(m rind of authigenic silicate. The low solubility of the Pb-bearing minerals resulting from encapsulation in non-Pb-bearing reaction rinds may provide an explanation for the limited Pb bioavailability observed when Butte soils were fed to rats (Freemanet al., 1992). Further evidence of the lack of absorption of lead from these soils is provided by the results of a blood-Pb study indicating very low blood-Pb levels in Butte children. The lower bioavailability of Pb from mining sites, compared to smelting and urban environments, is also due to kinetic limitations that control dissolution rates of Pb-bearing solids relative to the residence time of soil in the gastrointestinal (Gl) tract. When the test soil was fed to New Zealand White rabbits, only 9% of the total Pb was solubilised in the stomach, and therefore available for absorption. Anin vitro assay, developed to estimate maximum available Pb from soil, demonstrates that ingestion of mine-waste-bearing soil results in limited Pb dissolution, and produces results similar to thein vivo testing.     

Multiple determinants of community structure in shallow marsh habitats,Cape Fear River estuary,North Carolina,USA

The nekton of tidal creeks was studied at 17 sampling localities from September 1977 through August 1978, in the Cape Fear River estuary, North Carolina, USA. Prior to these dates, collections were made at 9 stations beginning in January 1977; these data were used to supplement conclusions drawn from the larger effort. Species recruited from the ocean utilized marsh habitats only temporarily and dominated the catches with over 70% of the total abundance. Their distribution was influenced by salinity gradients and to a lesser extent by substrate characteristics. In addition, temporal habitat partitioning with associated size differences of related species played an important role in structuring marsh nekton communities. A clearly defined ecotone was associated with the mesohaline-polyhaline transition zone, in slainities between 14 and 21 S. Numerous marine stenohaline forms were restricted to salinities above 16 S, thus increasing species richness in high salinity marshes. Despite differences in freshwater flows in 1977 and 1978, major features of the various marsh communities (species associations and relative abundances) exhibited little change throughout the Cape Fear estuary, indicating that these communities were relatively persistent in time. Standing crops for ocean-spawned species at the end of the growing season indicated that considerable annual export in the form of living biomass of fish and shellfish takes place from the marshes. Since most individuals of these species return to the ocean in the fall, an important energy link between the marshes and nearshore marine environment is demonstrated.     

Measurement of filtration rates by infaunal bivalves in a recirculating flume

A flume system and protocol for measuring the filtration rate of infaunal bivalves is described. Assemblages of multi-sized clams, at natural densities and in normal filter-feeding positions, removed phytoplankton suspended in a unidirectional flow of water. The free-stream velocity and friction velocity of the flow, and bottom roughness height were similar to those in natural estuarine waters. Continuous variations in phytoplankton (Chroomonas salinay) cell density were used to measure the filtration rate of the suspension-feeding clam Potamocorbula amurensis for periods of 2 to 28 h. Filtration rates of P. amurensis varied from 100 to 580 liters (gd)^-1 over a free-stream velocity range of 9 to 25 cm s^-1. Phytoplankton loss rates were usually constant throughout the experiments. Our results suggest that suspension-feeding by infaunal bivalves is sensitive to flow velocity.     

Seasonal fluctuations of benthic macrofauna in the apalachicola estuary,Florida, USA: The role of predation

Long-term monitoring studies revealed a negative correlation between the abundance of bottom-feeding fishes (Leiostomus xanthurus and Micropogonias undulatus) and benthic macrofaunal density, suggesting a causal relationship. To test this hypothesis, large (3x3 m) topless predator-exclusion cages (mesh opening=6 mm) of a design that produced minimal physical artifacts were constructed at two shallow mud-bottom oligohaline stations. Two-sided control cages and uncaged control areas were also sampled Experiments coincided with maximum abundances of bottom-feeding fishes in the spring and motile macroinvertebrates (penaeid shrimp) in the fall. During short-term (two-month) exclusion experiments in the spring and fall of 1980, large predators (those excluded by 6-mm mesh) were not a major factor in regulating the densities of most macrofaunal species. Predation may be important during other seasons, although this possibility was not tested. Densities of Leiostomus xanthurus and Penaeus setiferus during 1980 were lower than in previous years. Despite the reduced abundance of bottom-feeding fishes, macrofaunal densities still declined during late winter/early spring. It is postulated that increased river flow and lowered salinities during the late winter/early spring may contribute to the decline of macrofaunal densities by reducing larval recruitment.     

An estimate of carbonate production by macrobenthos in some tropical,soft-bottom communities

Productivity figures are available for most of the species comprising a few soft-bottom communities in the Miami area, USA. These are in the form of the ratio of annual somatic production to standing crop, both being measured as drytissue weight. These values have been converted to carbonate productivity ratios, and from these the carbonate production per square metre per year calculated. The values range from less than 1 g to nearly 400 g for sublittoral communities, and about 1 kg for one intertidal area.Contribution No. 1559 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.