Naticid snail predation on early post-settlement surfclams (<Emphasis Type="Italic">Spisula solidissima</Emphasis>) on the inner continental shelf of New Jersey,USA

Naticid gastropod predators leave characteristic boreholes in the shells of their bivalve prey that allow the evaluation of spatial differences in occurrence and intensity of predation. This approach, extensively used in paleobiological research, was used to compare spatial variation in predation by moonsnails (Euspira heros Say) on initial recruits of the Atlantic surfclam (Spisula solidissima Dillwyn). A four-year sampling at two 12-m deep stations on either flank of Beach Haven Ridge on the inner continental shelf detected temporal and spatial differences in surfclam densities following a large settlement pulse in early July. Between-station density differences were large during the first peak in surfclam density, but decreased in August-September, when densities at both stations also decreased. Surfclam seasonal peaks were followed by peaks in density of articulated shells (recent mortality due to non-crustacean predators and/or intrinsic mortality) and surfclams with boreholes (mortality due to naticids). The proportion of surfclams with boreholes was consistently higher (more than twice, on average) at the station where the highest densities of surfclams were also detected. These spatial differences are interpreted as differences in predation intensity resulting from a corresponding higher abundance of moonsnails at the same station. Although naticid predation is not the primary source of surfclam mortality, it consistently contributes to the reduction of spatial differences in density initially created by dissimilar levels of larval surfclam settlement.     

Vertical distribution of fish larvae in the Canaries-African coastal transition zone in summer

This study reports the vertical distribution of fish larvae during the 1999 summer upwelling season in the Canaries-African Coastal Transition Zone (the Canaries-ACTZ). The transition between the African coastal upwelling and the typical subtropical offshore conditions is a region of intense mesoscale activity that supports a larval fish population dominated by African neritic species. During the study, the thermal stratification extended almost to the surface everywhere, and the surface mixed layer was typically shallow or non-existent. Upwelling occurred on the African shelf in a limited coastal sub-area of our sampling. The vertical distributions of the entire larval fish population, as well as of individual species, were independent of the seasonal thermocline. Fish larvae and mesozooplankton were concentrated at intermediate depths regardless of the thermocline position, probably because of its weak signature and spatial and temporal variability. Day/night vertical distributions suggest that some species did not perform diel vertical migration (DVM), whereas others showed either type I DVM or type II DVM. The opposing DVM patterns of different species compensate for each other resulting in no net DVM for the larval fish population as a whole.     

Differential ingestion of zooplankton by four species of bivalves (Mollusca) in the Mali Ston Bay,Croatia

This study provides information about differences in composition of ingested zooplankton amongst bivalve species coexisting in the same area in a period from May 2009 to December 2010. The study was conducted at the Mali Ston Bay (42°51′ N, 17°40′ E)—the most important bivalve aquaculture area in the eastern Adriatic Sea. Stomach content analysis was performed on cultured species—Ostrea edulis and Mytilus galloprovincialis, and commercially important bivalve species from their natural environment—Modiolus barbatus and Arca noae. Results confirmed carnivory in bivalves, both from natural and cultured populations, but cultured species had higher numbers of zooplankters than those living on the seabed. The most abundant taxa were bivalve larvae, followed by tintinnids, copepods, unidentified eggs and gastropod larvae. Recorded numbers of bivalve larvae in M. galloprovincialis stomach were the highest so far reported and show that mussels impact the availability of natural spat.     

Ontogenetic changes in the vertical distribution of giant scallop larvae,Placopecten magellanicus,in 9-m deep mesocosms as a function of light,food, and temperature stratification

To understand how thermal stratification and food abundance affects the vertical distribution of giant scallop larvae Placopecten magellanicus (Gmelin), a mesocosm study was conducted in January and February 1992. The position of larvae was followed over 55 d in replicated 9-m deep tanks in relation to a sharp thermocline and the presence or absence of phytoplankton. Growth and vertical position of larvae were monitored in separate treatments which included phytoplankton added above the thermocline, below the thermocline, throughout the mesocosm, or absent from the mesocosm. Changes in the vertical position of larvae over time were quantified with a new, profiling, video-optical instrument capable of semi-automatically identifying, counting and sizing larvae. The strong diurnal migration of scallop larvae resulted in aggregations at two interfaces: the air/water interface during the night, and at the thermocline during the day. At times, the concentration of larvae within cm of the surface was > 100 times that in the remaining water column. The formation of bioconvective cells of swimming larvae at the air/water interface allowed larval aggregations to persist throughout the period of darkness. Regardless of the distribution of food, larvae remained above the thermocline during most of the experiment. Therefore, only in those treatments where food was also present above the thermocline was larval growth relatively high. Larger larvae penetrated the thermocline only after reaching a shell length of about 200 m; thus larval size, rather than chronological age, was more important in describing their vertical distribution. The rapid increase in kinematic viscosity with decreasing water temperature at the thermocline may retard the movement of larvae and contribute to aggregation at this interface. The influence of larval size on their vertical distribution, and the resulting potential for horizontal transport to settlement sites, points to the importance of persistent hydrographic features as critical factors contributing to settlement variance in scallops.     

High-frequency observations of early-stage larval abundance: do storms trigger synchronous larval release in <Emphasis Type="Italic">Semibalanus balanoides</Emphasis>?

The acorn barnacle, Semibalanus balanoides, is thought to release larvae in response to phytoplankton blooms, but there is evidence that another, unidentified cue for release may exist. We conducted high-frequency sampling in Little Harbor, Massachusetts, USA, to determine whether early-stage larval abundance was related to several environmental variables, and to characterize vertical distributions of the larvae. Larval concentrations peaked at 2.52 and 1.02 individuals l⁻¹ during two storms. Larvae were more abundant near the surface than near the bottom. We suggest the hypothesis that turbid conditions and upward-swimming behavior may protect newly-released larvae from predation and cannibalism. Future studies should test this hypothesis with barnacles and other invertebrates.     

Intra- and interannual variability in the larval fish assemblage off Gran Canaria (Canary Islands) over 2005–2007

The present study provides the longest and most intensive plankton and larval fish seasonal variability analysis in the Canary Islands and forms a basis for understanding life cycle scheduling and interactions among species, as well as the potential variability in transport processes of early life stages. Larval fish assemblages were studied weekly at Gran Canaria Island, Canary Islands, from January 2005 to June 2007, which represented two contrasting hydrological and biological periods. The former year was characterized by lower temperature and salinity that increased through 2006 and 2007. In contrast, chlorophyll concentration and mesozooplankton biomass decreased through the same period, especially when only the late winter bloom period was evaluated. However, ichthyoplankton abundance did not exhibit any clear pattern, as larger values were observed during 2006. The larval fish community of this oceanic island, located near the NW African upwelling, was composed of both neritic and oceanic taxa. Two families accounted for almost half of the collected larvae: Clupeidae (21.9%) and Myctophidae (20.5%). Although total larval concentration did not exhibit any seasonal peak linked to changes in zooplankton, the ichthyoplankton composition gradually changed during the year due to the high diversity and extended spawning periods of the fish species represented in samples. “Winter” and “summer” larval assemblages were identified, corresponding to the mixing and stratification periods of the water column, respectively. These assemblages were characterized by changes in the contribution of the most abundant annual taxa (Sardinella aurita, Cyclothone braueri, Ceratoscopelus spp. and Gobids) and by the presence of larvae of winter (Pagellus bogaraveo, Pomacentridae sp1) or summer spawners (Pomacentridae sp2, Trachinus draco, Arnoglossus thori, Tetraodontidae sp1). Upwelling filaments shed from the NW African coast reached the sampling area three times during this study, but changes in the local larval community were only detected in August 2005.     

<Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> blooms and clonal isolates from the northwest Atlantic coast cause rapid mortality in larvae of multiple bivalve species

Globally, many commercial bivalve populations have declined in recent decades. In addition to overharvesting and habitat loss, the increasing frequency and intensity of harmful algal blooms (HABs) are likely to contribute to bivalve losses, particularly in cases where blooms negatively impact larval stages. This paper reports on the lethal effects of clonal cultures and blooms of Cochlodinium polykrikoides from the US Atlantic coast on the larvae of three species of commercially and ecologically valuable bivalves: the Eastern oyster (Crassostrea virginica), the bay scallop (Argopecten irradians), and the Northern quahog (hard clam; Mercenaria mercenaria). Both cultures and blooms of C. polykrikoides were highly toxic to all three species of bivalve larvae causing 80–100% mortality during 24- to 72-h exposures at concentrations of 1–2 × 10³ cells ml⁻¹. Toxicity was dependent on cell densities, growth stage of C. polykrikoides (i.e. cultures in exponential stage growth were more toxic than later stages), exposure time of larvae to cells (i.e. longer exposure caused higher mortality), the age of larvae (i.e. younger larvae were more sensitive), and the relative abundance of C. polykrikoides (i.e. the presence of other microalgae decreased toxicity). Free radical-scavenging enzymes (peroxidase and catalase) and the removal of C. polykrikoides cells (i.e. culture filtrate) significantly increased larval survival suggesting toxicity is maximized by contact with live cells and may involve labile toxins bound by these compounds including e.g. reactive oxygen species. The toxicity of C. polykrikoides to bivalve larvae was generally more severe than other HAB species (e.g. Karenia brevis, Karlodinium veneficum, Alexandrium tamarense, Prorocentrum minimum). Since the bivalves in this study spawn in the months when C. polykrikoides blooms on the east coast of North America, these results suggest that these blooms may have detrimental effects on efforts to restore these already diminished populations.     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

Where are larvae of the vermetid gastropod <Emphasis Type="Italic">Dendropoma maximum</Emphasis> on the continuum of larval nutritional strategies?

This study evaluated whether larvae of the Indo-Pacific vermetid gastropod Dendropoma maximum are obligate planktotrophs, or whether they exhibit an intermediate feeding strategy. Experiments were conducted in Moorea, French Polynesia (149°50′W, 17°30′S), Sep–Oct 2009, to examine D. maximum larval growth and metamorphic responses to different diets and amounts of food. Dendropoma maximum larvae required particulate food to undergo metamorphosis, but were able to survive and grow in the absence of food for up to 20 days. Larvae in Low and Unfed food treatments exhibited phenotypic plasticity by growing a larger velum (the larval feeding structure) compared with those in high food. Unfed D. maximum larvae had a slower initial growth rate; however, by 11-day post-hatch fed and unfed larvae had converged on the same mean shell height (553 μm), which was only 10% larger than the initial size at hatching. Therefore, although the nutritional strategy of D. maximum larvae is best described as obligate planktotrophy, it appears to approach an intermediate feeding strategy.     

Ingestion of bivalve larvae by<Emphasis Type="Italic"> Mytilus edulis</Emphasis>: experimental and field demonstrations of larviphagy in farmed blue mussels

Ingestion of bivalve larvae by Mytilus edulis was investigated. Laboratory experiments revealed that ~ 90% of bivalve larvae offered to mussels was ingested and apparently fully digested. The shell of the bivalve larvae offered no protection against digestive processes, resulting in high larval mortality once inside the stomach. Stomach content analysis (September 2001–January 2003) showed that bivalve larvae were ingested by farmed mussels year-round, with the exception of March 2002. Numbers of ingested larvae were highest in October 2001 and May 2002, which coincides with known spawning times of farmed mussels in Ireland. Mussels ingested a large size-range of bivalve larvae, suggesting that all stages of the bivalve life cycle are vulnerable to predation. It is suggested that adult bivalves routinely filter larvae from the surrounding water and that, given the high biomass of mussels present in mussel farms, filtration by adult bivalves significantly reduces numbers of bivalve larvae in nearby waters.Communicated by J.P. Thorpe, Port Erin     

Ecological investigations of the late-stage phyllosoma and puerulus larvae of the western rock lobster Panulirus longipes cygnus

The distribution and abundance of the late-stage phyllosoma larvae of Panulirus longipes cygnus George and the distribution and densities of the final larval stage, the puerulus, both in the plankton and at settlement along the coast, were investigated. A total of 3,617 late-stage phyllosoma (Stages VI to IX) and 301 puerulus larvae were caught at 187 plankton stations during the July to November periods 1974, 1975 and 1976 off the west coast of Australia between 29°00 to 32°30S and 113°30 to 115°00E. The depth range sampled was 0 to 35 m on the continental shelf and 0 to 90 m off the shelf. During onshore/offshore cruises with similar sampling effort on and off the shelf, 1,169 late-stage phyllosoma larvae were taken, of which only 9 were caught on the shelf, and these near the outer edge. A series of cruises sampling two areas beyond the shelf near 29°30 and 32°00S yielded 2448 late-stage phyllosoma, with greater densities of larvae in the northern location. The settlement of puerulus-stage larvae along the coast in the same geographical range was also greater in the north than in the south. The data from the onshore/offshore cruises showed a definite effect of moon phase on numbers of puerulus larvae caught on the shelf, with higher catches near new moon. The low numbers of puerulus larvae (usually 0, 1 or 2 individuals) caught at all stations showed that the puerulus stage is sparsely distributed in the plankton. Fewer puerulus larvae were present at the surface than at lower depths, but it was not possible to determine a depth preference for the puerulus between 10 m and the lowest depths sampled because of the low catch numbers. No relationships were found between puerulus larvae density and surface-water temperature, salinity, or plankton biomass at each station. Data on the larval distributions indicate that, near the end of their planktonic existence, the majority of the late-stage phyllosoma larvae of P. longipes cygnus are not carried onto the shelf, where mixing of oceanic and continental shelf waters occurs only on the outer third, but are transported southward by oceanic circulation beyond the shelf. The puerulus moults from the last phyllosoma stage beyond the shelf and completes the larval cycle by swimming across the shelf and settling in the shallow reef areas.     

Vertical structure of very nearshore larval fish assemblages in a temperate rocky coast

Small-scale vertical patterns of larval distribution were studied at a very nearshore larval fish assemblage, during the spring–summer period of several years, at two depth strata (surface and bottom) using sub-surface and bottom trawls. A total of 4,589 larvae (2,016 from surface samples and 2,573 from bottom samples) belonging to 62 taxa included in 22 families were collected. Most larvae belonged to coastal species. Although inter-annual variations in larval density and diversity could be found, total larval abundance was always higher near the bottom whereas diversity was higher at the surface. A marked distinction between the structure of surface and bottom assemblages was found. Sixteen taxa explained 95% of the similarity among surface samples. Larvae which contributed most to this similarity included species like clupeiformes, sparids and serranids, and also blenniids, tripterygiids and some labrids. In the bottom samples, fewer species were present, with only six taxa, almost exclusively from species which lay demersal eggs, contributing to 95% of the similarity between samples. Larvae present at the surface were significantly smaller than at the bottom. For some of the most abundant species found at the bottom, only small larvae occurred at the surface while the whole range of sizes was present at the bottom, indicating that larvae may be completing the entire pelagic phase near the adults’ habitat. These results indicate that larval retention near the reefs probably occurs for these species, although for others dispersal seems to be the prevailing mechanism.     

Lethal effects of Northwest Atlantic Ocean isolates of the dinoflagellate, Scrippsiella trochoidea, on Eastern oyster (Crassostrea virginica) and Northern quahog (Mercenaria mercenaria) larvae

The thecate dinoflagellate Scrippsiella trochoidea is a cosmopolitan, bloom-forming alga that has been generally considered non-toxic. Here, we report that environmentally relevant cell densities (10⁴ cells mL⁻¹) of Scrippsiella trochoidea strains isolated from the Northwest Atlantic Ocean caused 100% mortality in Eastern oyster (Crassostrea virginica) larvae during 3-day exposures while parallel control larvae exhibited 100% survival. S. trochoidea also exhibited lethal effects on Northern quahog (Mercenaria mercenaria) larvae (70% mortality during 3-day exposure) but were non-toxic to juvenile fish (Cyprinodon variegates). The cultures of S. trochoidea were more lethal to Northern quahog larvae than ten other species of harmful algae, including the highly toxic species Cochlodinium polykrikoides. Scrippsiella trochoidea cultures within later stages of growth were more toxic than exponential growth stages to bivalve larvae, and the toxicity was dose dependent. Furthermore, toxicity was maintained in the cultures that were sonicated, boiled, and frozen as well as in resuspended residues of the culture but was significantly lower in cell-free culture media. Collectively, these results suggest that S. trochoidea causes mortality in bivalve larvae through a physicochemical rather than strictly chemical mechanism, such as clogging of larval feeding apparatuses by materials produced by S. trochoidea (e.g., lipids, extracellular polysaccharides, and/or cell debris) which accumulate as cells in culture or blooms age. This is the first report of the lethal effects of Scrippsiella trochoidea on shellfish larvae.     

Life history of a dominant polychaete, <Emphasis Type="Italic">Polygordius jouinae</Emphasis>, in inner continental shelf sands of the Mid-Atlantic Bight,USA

The life history of the polychaete Polygordius jouinae Ramey, Fiege, and Leander 2006 was studied in the sandy sediments of the LEO-15 research area off the coast of Tuckerton, New Jersey (39°27.69′N, 74°15.81′W). Divers collected sediment cores biweekly, from February 2004 to November 2005 to examine the temporal patterns of abundance and size–frequency distributions of P. jouinae. These parameters were also examined (for the first time) for P. jouinae in samples taken in ambient sediments and in three reciprocal 3 to 5-day sediment transplant experiments that were part of a previous, unrelated study at coarse sand and muddy sand sites (∼3 km apart) at LEO-15 in the summers of 1994 and 1995. Analysis of these previous data along with the current data provided a larger set of values for P. jouinae, as well as further insight into the life history, habitat selection, and dispersal of P. jouinae (not considered in previous studies). Polygordius jouinae is gonochoristic and the population was mostly composed of sexually mature individuals in late May 2004 and 2005. The reproductive period usually occurs from May to August and individuals that spawn generally have a 1-year life span. Non-spawning individuals of P. jouinae could potentially live longer than 1 year and therefore the population may sometimes consist of individuals of two generations. Recruitment begins no later than July as recently settled individuals (≤9 mm body length), which were not present during the first week of June 1995, 2004, and 2005, first appeared in July in all 3 years. The smallest individual was 2.01 mm long and 0.07 mm wide and this provides the first estimate of size at initial recruitment. Although some worms in sediment trays likely settled as larvae, most worms were recently settled juveniles (length categories 6 and 9 mm) transported in from the surrounding sediments, presumably as a consequence of re-suspension or bed load transport. As expected, larger individuals were present later in September. Although it is unknown whether P. jouinae were transported actively or passively these data suggest that recently settled juveniles are capable of habitat selection (prefer sand over mud) and show that post-settlement transport and selection influence community structure.     

Endogenous swimming rhythms in estuarine crab megalopae: implications for flood-tide transport

Up-estuary migration of crab larvae to adult habitats is thought to be accomplished by selective tidal transport in which late-stage larvae enter the water column on flood tides and remain on or near the bottom on ebb tides. This study measured endogenous rhythms in swimming by the last larval stage (megalopa) of blue crabs Callinectes sapidus and fiddler crabs Uca spp. Previous field studies found that megalopae of both species were only abundant in the estuarine water column on nocturnal rising tides. Megalopae were collected from the Newport River Estuary, North Carolina (34°41N; 76°40W) during August–September 1992 and swimming activity was recorded for 4.5 to 7 d under constant conditions with a video system. Rhythms exhibited by both genera in the laboratory were not identical to those recorded in the field. Uca spp. displayed a circatidal rhythm, with maximum swimming occurring near the time of high tide in the field. Rhythm amplitude increased when crushed oyster shells were present, which suggested that megalopae bury or cling to the substrate during quiescent periods. In contrast, C. sapidus had a circadian rhythm in which maximum swimming coincided with the day phase in the field. In most trials, the activity of blue crab megalopae was unrelated to the expected tidal cycle. It was concluded that a tidal rhythm in swimming was the behavioral basis of flood-tide transport for fiddler crab larvae. The endogenous rhythm in blue crabs does not participate in transport, which probably results from behavioral responses to environmental cues associated with flood tide.     

Spatial patterns of ichthyoplankton distribution in relation to hydrographic features in the Northern Benguela region

The spatial distribution of the most abundant eggs and larvae of teleost fish species on the continental shelf and slope off the northern Benguela region was studied in April 1986. The horizontal and vertical distribution of eggs and larvae were analysed together with environmental data, in order to determine patterns of ichthyoplankton distribution. Both species composition and relative egg and larval abundance levels exhibited important latitudinal differences during a period of quiescent upwelling with an intense intrusion of Angolan water into the system. Larval diversity was higher in the northern part of the study area, where, because of the intrusion of the warmer Angolan water, the water column was more stratified than in the southern part, where the affect of upwelling of South Atlantic Central Water was continuous and only a few species spawned. The frontal zone appeared to be a nursery ground for the most important pelagic species of the region:Trachurus trachurus capensis, Engraulis capensis, andSardinops ocellatus. Vertical egg and larval distributions showed evidence of stratification, with highest concentrations located in the uppermost 50 m. In comparison, during periods of intense upwelling, longitudinal gradients were responsible for the horizontal distribution of ichthyoplankton, and the vertical distribution of eggs and larvae were much more extensive because of the greater mixing of the water column.     

Settlement-competency periods of larvae of three species of scleractinian corals

The pattern of settlement over time of three broadcast spawning coral species (Cyphastrea serailia, Acanthastrea lordhowensis, and Goniastrea australensis) from the Solitary Islands (30°00′S; 153°20′E) was studied in 1995 and 1996 in order to determine the maximum length of time these larvae could remain in the water column and still retain the ability to settle and metamorphose. Larvae were maintained in aquaria and the number which had settled on biologically-conditioned tile pairs was monitored every 5 to 10 d. While the majority of larvae settled quickly after becoming competent, some larvae survived and settled for extended periods after spawning. Competency periods ranged from 26 d for C. serailia to 56 d for G. australensis and 78 d for A. lordhowensis. These data greatly extend the known competency periods for larvae of broadcast-spawning corals and indicate the potential for transport of broadcast-spawned coral larvae over large distances. Medium to long-distance larval dispersal of the species studied provides a mechanism for their widespread distribution in subtropical regions, on reefs which are often widely spaced and relatively isolated. Received: 27 May 1997 / Accepted: 27 November 1997     

Behaviour of echinoid larvae around sharp haloclines: effects of the salinity gradient and dietary conditioning

We examined larval response to a range of sharp haloclines and determined the effect of dietary conditioning on that response in the sea urchins Echinometra lucunter and Arbacia punctulata. We reared larvae in the laboratory under a high or low concentration of either single (Isochrysis galbana) or mixed (Isochrysis galbana, Dunaliella tertiolecta, Thalassiosira weissflogii) microalgal species. For both species of sea urchins, rate of larval development was faster and age-specific larval length and width were greater in high-ration than low-ration diets. We examined the distribution of two- and four-arm larvae of E. lucunter from each diet treatment and of four-arm larvae of A. punctulata from the high-ration diets in cylinders with experimentally constructed haloclines. In three of the halocline treatments, the salinity of the bottom layer was 33‰ and that of the top layer was 21, 24 or 27‰ (21/33, 24/33 and 27/33) and in a fourth one, the salinities of the bottom and top layer were 30 and 21‰, respectively (21/30). The position of larvae in the cylinders varied with the steepness of the halocline and with dietary conditioning for both sea urchin species and all developmental stages tested. Significantly more larvae crossed the haloclines into water of 24 and 27‰ salinity than into water of 21‰ salinity. We observed an effect of diet on the position of larvae in the cylinders, and that effect varied among halocline treatments for both species. The proportion of larvae of E.lucunter that crossed the halocline was greater in low- than high-ration diets in the 24/33 and 27/33 treatments. Position of four-arm larvae in the cylinders also varied with food quality in high-ration diets: for E.lucunter in the 24/33 treatments, and for A. punctulata in the 21/30 treatments, more larvae from the single- than from the mixed-species diets were present above the halocline. Salinity in the adult habitat during most of the active reproductive period ranged from 15 to 40‰. We showed that larvae can respond to gradients in salinity, and therefore can remain within a water mass of higher salinity overlying the adult habitat. However, survival of poorly fed larvae may be increased if they are introduced into a new water mass and carried away from a nutritionally poor environment. Received: 9 July 1997 / Accepted: 12 January 1998     

Spatial and temporal genetic homogeneity in the Arctic surfclam (<Emphasis Type="Italic">Mactromeris polynyma</Emphasis>)

Commercially harvested marine bivalve populations show a broad range of population-genetic patterns that may be driven by planktonic larval dispersal (gene flow) or by historical (genetic drift) and ecological processes (selection). We characterized microsatellite genetic variation among populations and year classes of the commercially harvested Arctic surfclam, Mactromeris polynyma, in order to test the relative significance of gene flow and drift on three spatial scales: within commercially harvested populations in the northwest Atlantic; among Atlantic populations; and between the Atlantic and Pacific oceans. We found small nonsignificant genetic subdivision among eight populations from the northwest Atlantic (F _ST = 0.002). All of these Atlantic populations were highly significantly differentiated from a northeast Pacific population (F _ST = 0.087); all populations showed high inbreeding coefficients (F _IS = 0.432). We tested one likely source of heterozygote deficits by aging individual clams and exploring genetic variation among age classes within populations (a temporal Wahlund effect). Populations showed strikingly different patterns of age structure, but we found little differentiation among age classes. In one case, we were able to analyze genetic diversity between age classes older or younger than the advent of intensive commercial harvesting. The results generally suggest spatially broad and temporally persistent genetic homogeneity of these bivalves. We discuss the implications of the results for the biology and management of surfclam populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Settlement of <Emphasis Type="Italic">Macoma balthica</Emphasis> larvae in response to benthic diatom films

The role of multi-species benthic diatom films (BDF) in the settlement of late pediveliger larvae of the bivalve Macoma balthica was investigated in still-water bioassays and multiple choice flume experiments. Axenic diatom cultures that were isolated from a tidal mudflat inhabited by M. balthica were selected to develop BDF sediment treatments characterized by a different community structure, biomass, and amount of extracellular polymeric substances (EPS). Control sediments had no added diatoms. Although all larvae settled and initiated burrowing within the first minute after their addition in still water, regardless of treatment, only 48–52% had completely penetrated the high diatom biomass treatments after 5 min, while on average 80 and 69% of the larvae had settled and burrowed into the control sediments and BDF with a low diatom biomass (<3.5 μg Chl a g⁻¹ dry sediment), respectively. The percentage of larvae settling and burrowing into the sediment was negatively correlated with the concentration of Chl a and EPS of the BDF. This suggests higher physical resistance to bivalve penetration by the BDF with higher diatom biomass and more associated sugar and protein compounds. The larval settlement rate in annular flume experiments at flow velocities of 5 and 15 cm s⁻¹ was distinctly lower compared to the still-water assays. Only 4.6–5.8% of the larvae were recovered from BDF and control sediments after 3 h. Nonetheless, a clear settlement preference was observed for BDF in the flume experiments; i.e., larvae settled significantly more in BDF compared to control sediments irrespective of flow speed. Comparison with the settlement of polystyrene mimics and freeze-killed larvae led to the conclusion that active selection, active secondary dispersal and, at low flow velocities (5 cm s⁻¹), passive adhesion to the sediment are important mechanisms determining the settlement of M. balthica larvae in estuarine biofilms.