The subtidal pump: a mechanism of interstitial water exchange by wave action

Sandy subtidal sediments are part of the earth's largest filter system. Water flow through bottom sediments is driven by wave action on the sea surface. The mechanisms involved are described, including a theoretical deduction and field measurements. As an example, the total water exchange through part of the West Atlantic shelf is computed and the influence of the phenomenon is discussed from a biological point of view and with regard to its importance for the world's oceans.     

Food-particle size and selection by bivalve larvae in a temperate embayment

Epifluorescence microscopy was used to analyze the stomach contents of bivalve larvae collected in the Baie des Chaleurs (western Gulf of St. Lawrence, Canada) in order to document food-particle sizes, compare feeding among taxa, and compare the diet with the in situ phytoplankton community. Stomach contents were mainly composed of small autotrophic flagellates (<5 μm) and cyanobacteria (<2 μm), reflecting the microbial food web which characterizes these waters. More than half (55%) of all veligers examined contained algal cells of 5 to 15 μm, whereas only 3% had cells of 15 to 25 μm. Differences in the size ranges of ingested algal cells among similar-sized larvae of different species suggests that veligers actively selected food particles. Among the smallest veligers (185 to 260 μm), scallops (Placopecten magellicanus) and mussels (Mytilus edulis) ingested more <5 μm and 5 to 15 μm algae than clams (Mya arenaria). Among larger veligers (261 to 405 μm), clams contained significantly more <5 μm cells than mussels, whereas mussels contained significantly more 5 to 15 μm algae than clams. Algal cells of 15 to 25 μm were preferentially ingested by mussel veligers. Feeding also differed between different-sized veligers within taxa, i.e. the smallest clam veligers ingested fewer of 5 to 15 μm algae than the larger size classes. Mussel veligers ingested significantly more 15 to 25 μm and fewer <5 μm cells as their size increased. The dominance of ultraplankton in the nearshore waters of Baie des Chaleurs and in the stomach contents suggests that veliger larvae may be an important export path for carbon produced by small phytoplankton. Received: 17 July 1996 / Accepted: 20 September 1996     

Assessment of space utilisation in a subtidal temperate bryozoan community

On subtidal hard substrata, Bryozoa are often very abundant, both in terms of numbers of colonies and numbers of species. To investigate how so many species are able to co-exist, often in close proximity to each other, subtidal material was dredged from an area of bryozoan-rich coarse shelly sea-bed off the south-west of the Isle of Man (British Isles) in autumn 1984 and 1985. A survey was performed on the distribution of colonies of eight common, encrusting cheilostome bryozoan species across the surfaces of four types of empty, disarticulated bivalve shells. Each species was found to be significantly commoner on certain substratum types such as rough surfaces or inner concave surfaces. Fenestrulina malusii (Audouin) showed a further, more specific, tendency to be signficantly more abundant on the grooves of the corrugated inner surfaces of left valves of Pecten maximus (L.). In general however there was much overlap in distribution and it is proposed that such large numbers of species are able to co-exist because of the evolution of different life-history strategies ensuring that each species uses the available temporally and spatially heterogenous substrata in subtly different ways; some species, for example, occupying substrata that are optimal to bryozoan growth and other species, by reaching maturity earlier, being able to inhabit more ephemeral suboptimal substrata.     

Pelagic-benthic coupling in a subtidal system of the North-Western Mediterranean

Suspended particulate matter, zooplankton, and macrobenthos dynamics were investigated in a shallow area of the Ligurian Sea (north-west Mediterranean) characterized by wide temporal variability over an annual cycle. As indicated by multivariate analyses, the seasonal dynamics can be summarized as follows: (1) a late winter-early spring phytoplankton bloom followed by high zooplankton and macrobenthos densities during the spring months; (2) low-quality particulate suspended matter in summer, and an increase in the importance of zooplankton taxa with a wide range of feeding strategies, a decrease in macrofaunal abundance, and an increase in deposit-feeders and predators; and (3) a second phytoplankton bloom in autumn, followed by an increase in copepod density and a low macrofaunal abundance. In conclusion, pelagic and benthic communities in the coastal area of the Ligurian Sea mainly seem to be controlled bottom-up. Our results suggest that the quality of the particulate organic matter may play an important role in determining the temporal changes of both plankton and benthic assemblages, while the direct influence of other environmental features (such as sediment grain size) is relevant only for some macrobenthic taxa (e.g. crustaceans).     

Plant facilitation of a belowground predator

Interest in facilitative predator plant interactions has focused upon above-ground systems. Underground physical conditions are distinctive, however, and we provide evidence that bush lupine, Lupinus arboreus, facilitates the survival of the predatory nematode Heterorhabditis marelatus. Because H. marelatus is prone to desiccation and lupines maintain a zone of moist soil around their taproots even during dry periods, we hypothesized that dry-season nematode survival under lupines might be higher than in the surrounding grasslands. We performed field surveys and measured nematode survival in lupine and grassland rhizospheres under wet- and dry-season conditions. Nematodes survived the crucial summer period better under lupines than in grasslands; however, this advantage disappeared in wet, winter soils. Modeling the probability of nematode population extinction showed that, while even large nematode cohorts were likely to go extinct in grasslands, even small cohorts in lupine rhizospheres were likely to survive until the arrival of the next prey generation. Because this nematode predator has a strong top-down effect on lupine survival via its effect on root-boring larvae of the ghost moth Hepialus californicus, this facilitative interaction may enable a belowground trophic cascade. Similar cases of predator facilitation in seasonally stressful environments are probably common in nature.     

Long-distance dispersal of a subantarctic brooding bivalve (Gaimardia trapesina) by kelp-rafting

The probability of successful dispersal by sessile benthic invertebrates is thought to strongly influence their geographic distribution and population genetics. Generally, species with long-lived planktonic larvae are expected to exhibit wider distribution patterns than those species which brood their young, due to their presumably greater potential for dispersal. In some cases, however, brooding species exhibit broad distributions and show evidence of genetic exchange with geographically distant populations. One potential factor that has been invoked as an expianation is dispersal by floating and rafting of adults and egg masses. Several studies have shown that it is possible for sessile adults to disperse on the order of several to many thousand kilometers by rafting on debris in ocean currents. With very few exceptions, however, direct evidence of rafting in the open ocean has been lacking. We present evidence of long-distance (1300 to 2000 km) dispersal of a brooding pelecypod,Gaimardia trapesina (Lamarck, 1819), in the Southern Ocean in the vicinity of Cape Horn, the Falkland Islands, and the antarctic island South Georgia (54°S; 37°W). Data on survival and fecundity rates ofG. trapesina and the prevalence of kelp rafts collected during the austral winter of 1993 indicate that dispersal by rafting can occur over ecologically relevant time scales and could potentially serve as a significant means of genetic exchange between populations.     

Digestion in a subtidal population of Mercenaria mercenaria (Bivalvia)

The digestive processes of Mercenaria mercenaria (Linn.) were studied in a natural subtidal population. Hourly sampling of 4 to 5 quahogs was conducted over one 25 h period in August, 1978, at Woods Hole, Ma., USA. Crystalline style length did not vary significantly with time, tide, particulate C, N or suspended material in the water. Changes in secretory activity of the style sac epithelium were not evident from histological sections. Based on cellular morphological differences, digestive tubules were individually classified into one of 4 categories, indicative of the state of intracellular digestion within the digestive cells. All 4 tubule types were present in individuals from each sample hour, signifying that intracellular digestion occurs continually within the digestive gland. A 3 h time lag is evident between times of peak C, N or suspended materials and apparent peak levels of absorptive (Type II) tubules.Contribution No. 76 Marine Science Institute, Northeastern University; Nahant, MA 01908, USA     

Biogenic disturbance determines invasion success in a subtidal soft-sediment system

Theoretically, disturbance and diversity can influence the success of invasive colonists if (1) resource limitation is a prime determinant of invasion success and (2) disturbance and diversity affect the availability of required resources. However, resource limitation is not of overriding importance in all systems, as exemplified by marine soft sediments, one of Earth's most widespread habitat types. Here, we tested the disturbance-invasion hypothesis in a marine soft-sediment system by altering rates of biogenic disturbance and tracking the natural colonization of plots by invasive species. Levels of sediment disturbance were controlled by manipulating densities of burrowing spatangoid urchins, the dominant biogenic sediment mixers in the system. Colonization success by two invasive species (a gobiid fish and a semelid bivalve) was greatest in plots with sediment disturbance rates < 500 cm(3) x m(-2) x d(-1), at the low end of the experimental disturbance gradient (0 to > 9000 cm(3) x m(-2) x d(-1)). Invasive colonization declined with increasing levels of sediment disturbance, counter to the disturbance-invasion hypothesis. Increased sediment disturbance by the urchins also reduced the richness and diversity of native macrofauna (particularly small, sedentary, surface feeders), though there was no evidence of increased availability of resources with increased disturbance that would have facilitated invasive colonization: sediment food resources (chlorophyll a and organic matter content) did not increase, and space and access to overlying water were not limited (low invertebrate abundance). Thus, our study revealed the importance of biogenic disturbance in promoting invasion resistance in a marine soft-sediment community, providing further evidence of the valuable role of bioturbation in soft-sediment systems (bioturbation also affects carbon processing, nutrient recycling, oxygen dynamics, benthic community structure, and so on.). Bioturbation rates are influenced by the presence and abundance of large burrowing species (like spatangoid urchins). Therefore, mass mortalities of large bioturbators could inflate invasion risk and alter other aspects of ecosystem performance in marine soft-sediment habitats.     

Selectivity in feeding by the deposit-feeding bivalve Macoma nasuta

Macoma nasuta Conrad is primarily a deposit-feeding bivalve sucking the top millimeter of the sediment surface. Growth experiments show that surface sediment supports growth better than detritus falling from the water column. Gut clearance time is between 1 and 9 h (12°C). Fecal pellets are ejected in a regular rhythm. However, the total amount of feces per unit time shows considerable individual variation. Due to sorting in the mantle cavity, about 97% (dry weight) of the surface material is ejected again as pseudofeces. Selectivity by the bivalve is estimated by comparison of particle size and organic composition of sandy and muddy sediments and compared with feces produced by clams fed these sediments. Fecal pellets are in all cases richer in organic components than the sediment, indicating a high degree of selectivity. Ingestion and digestion of small animals (meiofauna) occur, but many of the ingested specimens survive. It is not possible to estimate the assimilation of organic matter by simple difference between the ingested sediment and the ejected feces. The difficulties in calculating energy budgets which arise from selective feeding and associated bacteria are discussed.     

Impact of predation-disturbance by large epifauna on sediment-dwelling harpacticoid copepods: Field experiments in a subtidal seagrass bed

The hypothesis was tested that predation-disturbance by epibenthic macrofauna affects the abundance of sediment-dwelling harpacticoid copepods. The copepods inhabited a subtidal seagrass (Zostera marina L.) bed in British Columbia, Canada. The response of the harpacticoid community was observed in controlled field experiments in which epibenthic predators-disturbers were excluded from portions of the seagrass bed. Controlled, exclusion-cage (0.8 m² area, 7-mm mesh) experiments were conducted within the seagrass bed from late March/early April to mid-June in both 1986 and 1987. Sampling was conducted biweekly. Exclusion of large epibenthic predators-disturbers had little effect on sediment-dwelling harpacticoid copepod density. Total harpacticoid numbers and abundances of dominant species generally did not increase in the exclusion treatment relative to the control. Shifts in species composition of the harpacticoid community did not occur. The treatment control was adequate in simulating the exclusion cage structure. It appears that large epibenthic predators-disturbers have little impact on the abundance of harpacticoid copepod populations at this study site.     

Sclerotized protein in the shell matrix of a bivalve mollusc

The epsilon-amino groups of lysine and phenolic groups of tyrosine are most heavily concentrated in the newlydeposited organic matrix of the shell of the bivalve Mercenaria mercenaria. A phenoloxidase enzyme which oxidizes L-dihydroxyphenylalanine is present only in this new area of the shell matrix. Scanning electron micrographs of calcified secretions of the shell show that accretion lines, thought to be layers of organic matrix separating diurnal acceretions of calcium carbonate, are not developed until up to 4 d after deposition of shell material. These results suggest that the shell matrix is hardened by some kind of polymerization, and that lysine and tyrosine residues in the matrix are involved in the process. Accretion lines in polished and etched sections become visible only after complete hardening of the polymer occurs.     

Responses of Asterias rubens to bivalve prey in a Y-maze

Experiments carried out in a Y-maze apparatus using a constant number of mussels, Mytilus edulis L., as prey showed the presence of a food-seeking period in the predatory starfish Asterias rubens L. collected in the Menai Strait, Anglesey, North Wales. The starfish orientated towards the mussels in the experiment conducted from Novemeber-December to May. The food-seeking activity was not significantly different from random expectation between June and October. It is suggested that this seasonal change in the behaviour of A. rubens can be due either to a lowering of chemo-sensitivity and failure to respond by A. rubens, or to a seasonal change in production of attractant(s) by the prey, or to a combination of the above. A. rubens does not move towards a bait of oyster spat (Ostrea edulis L.) in the Y-maze unless it has been previously conditioned to eat the oysters for about a month. In the Menai Strait, oysters would rarely be encountered by A. rubens and, therefore, the experiments confirmed the concept of ingestive conditioning (Wood, 1968).     

Inheritance of a nuclear DNA polymorphism assayed in single bivalve larvae

We examined the inheritance of theMytilus edulis CaM-1 Intron 3 locus, a non-coding DNA locus with two potentially neutral length-variants. The polymerase chain reaction (PCR) was used to determine theCaM-1 genotype for 799 larvae obtained from 11 laboratory crosses. Larvae were typed singly only 8 to 24 h after fertilization. We find evidence that each allele can be inherited from either sex, that there are no barriers to fertilization between gametes of different genotypes, and that most larvae have genotypes compatible with Mendelian inheritance of the locus. Deviations from expected genotype frequencies were found in some crosses; we suggest that a contributing factor is aneuploidy in early larvae, but a major cause is more likely to be selection at a locus linked toCaM-1, occurring under the artificial laboratory culture conditions. This study demonstrates the feasibility of applying molecular genetic techniques to early larval stages of marine bivalves, and presents a new non-destructive biopsy method for DNA analysis from living adult mussels.     

Genetic structure of the subtidal red alga Delisea pulchra

We used random amplified polymorphic DNA (RAPDs) to examine small-scale spatial genetic structure in the red alga Delisea pulchra (Greville) Montagne at two locations near Sydney, Australia. We examined genetic structure among plants at four spatial scales ranging from 2 km apart down to <50 cm apart between locations, among sites within locations, among quadrats within sites, and among plants within quadrats. Haploid stages of D. pulchra were absent from the populations studied, suggesting that they are maintained through asexual reproduction of diploid plants. Consistent with this, we found that 19 RAPD phenotypes scored in this study had multiple individuals, indicating the presence of clones in these populations. However, there were no RAPD phenotypes common to two locations separated by only 2 km. Analysis of molecular variance revealed that strong genetic differences occurred between plants from these two locations, with 46.3% of the total genetic variation occurring at this scale, most probably reflecting limited gene flow. Within each location, <25% of the genetic variation was attributable to differences among sites or quadrats, indicating gene flow at those smaller scales. Most of the variation within each location occurred at the smallest spatial scale, among plants within 0.25 m² quadrats. Nonetheless, some pairwise genetic distances (φST) between sites or quadrats within locations were large, indicating some genetic divergence on smaller scales. Genetic distance was independent of spatial distance within both locations, suggesting that fine-scale differences within locations were most probably caused by variation in fine-scale patterns of water movement or fine-scale natural selection. We assessed the impact of one potential selective agent, grazing sea urchins, on the fine-scale genetic structure of D. pulchra. There was no evidence that grazing by sea urchins affected the genetic structure of D. pulchra. In combination with demographic data, our results indicated that local populations of D. pulchra within locations were relatively open and that fine-scale genetic structure was probably constrained by gene flow. At the larger scale however, strong genetic differentiation indicated little gene flow between locations and restricted dispersal of spores. Received: 22 April 1999 / Accepted: 29 November 1999     

Circatidal swimming activity rhythm in a subtidal cumacean Dimorphostylis asiatica (Crustacea)

Dimorphostylis asiatica, a cumacean crustacea inhabiting the sublittoral line of the Seto Inland Sea, shows a rhythmic pattern of swimming activity coinciding with daily and tidal components in the field. This activity pattern was bimodal at first, i.e., circatidal activity coinciding with high tides at their habitat. The mean free-running period (i.e., bitidal interval) of this endogenous rhythm was 23.1 h at 10°C, which was significantly shorter than the environmental tidal cycle. In most cases (90% of the records) this bimodal activity became unimodal within 10 d. The unimodal period was 24 to 27.5 h, which is markedly longer than the preceding bimodal period. The bimodal pattern observed in the present study was not observed in the field. The difference between field and laboratory activity patterns can be explained in terms of direct response of the bimodal circatidal rhythm to the day-night cycle in the field. Field observations were made and laboratory experiments conducted between 1988 and 1992.     

Life history of the temperate subtidal gastropod Umbonium costatum

The life history and reproductive biology of the trochid sand snail Umbonium costatum (Kiener) were investigated on a subtidal sandy shore in Hakodate Bay, Japan between 1988 and 1991. Female U. costatum mature at 1 yr of age (shell diameter-11 mm), reproduce twice (in June-July and September-October) in successive years, grow to a maximum size (shell diameter=22 mm) at age 8 yr, increase annual fecundity with age from 2000 (age 1 yr) to 37000 (age 8 yr), and show a maximum monthly gonad somatic index of 8% which is constant among ages. In comparison to a previously studied life history of a tropical Umbonium vestiatium, temperate U. costatum shows more sustained growth and a longer life span after maturation. This could be explained by: (1) the optimal size model concerned with resource investment, in gametes (Sebens 1987); and by (2) bet hedging to compensate large variability in larval success at high latitudes. These two hypotheses are not mutually exclusive, but both are based on season-related extremes of environment at high latitudes where the period suitable for reproduction is short.     

Effects of long-term physical disturbance by commercial scallop fishing on subtidal epifaunal assemblages and habitats

 This paper examines spatial differences in the distribution of by-catch assemblages from the scallop [Pecten maximus (L.) and Aequipecten opercularis (L.)] fishing grounds in the North Irish Sea, during 1995. The sites examined have been exposed to differing known levels of fishing disturbance by scallop dredging, based on unusually high-resolution data extracted from fishermens' logbooks. Uni- and multi-variate techniques have been used on a production dataset (a value which incorporates both abundance and biomass figures), as well as abundance and biomass data individually. The original species list was reduced to higher taxonomic groupings in line with the theory that the latter is more appropriate for detecting anthropogenic change. Species diversity and richness, total number of species, and total number of individuals all decrease significantly with increasing fishing effort. Species dominance increases with effort. Total abundance, biomass and production, and the production of most of the major individual taxa investigated decrease significantly with increasing effort. Multivariate analysis reveals a significant relationship between fishing effort and by-catch assemblage structure. The taxa most responsible for the differences are the echinoids and cnidarians, but prosobranch molluscs and crustaceans also contribute to the differences. By-catch assemblage structure is more closely related to fishing effort than any other environmental parameter investigated, including depth and sediment type. We observed an approximately linear decrease in diversity with increasing fishing disturbance, and suggest this is primarily due to selective removal of sensitive species and, more importantly, habitat homogenisation. These results were interpreted in the light of ecological theories relating disturbance to community structure. The argument that invertebrate scavenger populations benefit from prolonged exposure to fishing disturbance was also examined, but no supporting evidence was found. Received: 29 January 2000 / Accepted: 31 May 2000     

A demographic study of a subtidal population of the New Zealand articulate brachiopod Terebratella inconspicua

Size-frequency distributions from subtidal populations of Terebratella inconspicua (Sowerby, 1846) consistently display prominent modes at 18 to 22 mm shell length, representing many years accumulation of non-growing adults. In addition, these distributions contain seasonally variable modes made up of new recruits which are present in very high numbers towards the end of the breeding season, but are reduced to very low numbers by the end of their first year of life. Growth of recruits follows a sigmoid curve with considerable variability between individuals. Some adults survive at least 14 years. Females are 2 years old before they breed. Larvae are brooded internally by the parent and spawned in autumn and spring. The density of recruitment is positively correlated with the density of residents. Larvae commonly settle on the shells of large brachiopods; these recruits can survive to breed, but their survivorship as adults is lower than that of conspecifics attached to rock. Total mortality in the population is size (=age)-dependent; the mean life expectancy of a new recruit is 0.54 years compared with 2.57 years for a 1 year-old. Sources of mortality are considered.     

Social facilitation of selective mortality

Territorial defense by breeders influences access to resources near defended nest sites by intruder species and may have indirect effects on other species within the territory, leading to local patchiness in distribution patterns. The present study demonstrates that adult males of a damselfish, Pomacentrus amboinensis, indirectly facilitate the increased survival of conspecific juveniles through the territorial defense of their nesting site from potential egg predators. Moreover, male territoriality results in a shift in the selectivity of predation on newly settled juveniles. We monitored the fate of pairs of predator-naive, newly settled P. amboinensis placed inside and outside nesting territories. Individuals within a pair differed in size by approximately 1 mm and were tagged for individual identification. Away from male territories larger juveniles had greater survival, while within territories, larger juveniles suffered higher mortality. Behavioral observations indicated that the moonwrasse Thalassoma lunare, a predator of benthic eggs and small fishes, had reduced access to juveniles within male territories, while another predator on small fishes, the dottyback Pseudochromis fuscus, had unobstructed access to male territories. Experimental removal of P. fuscus indicated that the shift in the direction of phenotypic selection on newly settled juveniles was the indirect effect of aggression by nest-guarding male damselfish, which resulted in differential access to male territories by these two predators of small fishes. Evidence suggests that behavioral interactions between the resident community and intruders will influence patchiness in selective pressures imposed on benthic prey by influencing both the composition of predator types that can access the prey resource and their relative abundance. How this spatial and temporal patchiness in predator pressure interacts with spatial patchiness of recruiting prey will have a major influence on the resulting distribution of juveniles and their phenotypic traits.