Age-structure, recruitment, and adult mortality in populations of the Atlantic surfclam, Spisula solidissima, from 1978 to 1997

The Atlantic surfclam, Spisula solidissima (Dillwyn, 1817), is a dominant member of the benthic community on the continental shelf from Georges Bank to North Carolina, USA. This bivalve has supported a major fishery, primarily off New Jersey and the Delmarva Peninsula, since the 1960s. Early papers documented that these populations were at historical lows in the mid-1970s owing to commercial harvesting and a hypoxic event off New Jersey. It was also shown that major recruitment took place off New Jersey in 1976 and off the Delmarva Peninsula in 1977. Because the size frequencies of surfclams from federal surveys do not show distinct year classes, there has been uncertainty about the number of year classes in these populations throughout the 1980s and 1990s. The present study describes changes in population age- and size-structure from 1978 to 1997 in federal waters (≥5.5 km from shore) of the USA. Given the 30 to 35 year life span of S. solidissima, these populations could be composed of many year classes. Yet, these populations were composed of only two to three year classes in 1978. Through annual recruitment, the number of year classes increased over time, and populations off New Jersey and the Delmarva Peninsula contained at least 19 year classes in 1997. This major change in population structure over time was not evident from examination of available size-frequency data, and could only be inferred from data on age-composition. Throughout the 1980s and 1990s, the surfclam fishery was supported by multiple year classes. The mean and variance of recruitment to the New Jersey region, as indicated by the abundance of 4-year-olds over time, was greater than that off Delmarva, particularly between 1980 and 1986. The instantaneous rate of adult mortality, which includes the effect of harvesting, was approximately 0.26 yr⁻¹ in each region. Received: 11 September 1998 / Accepted: 1 February 1999     

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.     

Growth of the Atlantic surfclam,Spisula solidissima,from Georges Bank to the Delmarva Peninsula,USA

Age/shell length data for offshore surfclam, Spisula solidissima (Dillwyn, 1817), populations were used to estimate the parameters of the von Bertalanffy growth model by time period and region, from Georges Bank in the north to the Delmarva Peninsula in the south. Randomization tests were used to compare curves. We tested the a priori hypothesis that growth curves would change over time in the south (i.e., New Jersey and Delmarva) but remain constant in the north (i.e., Long Island and South New England). This hypothesis was proposed because surfclam population structure in the south had been altered by the hypoxic event of 1976, and possibly by intense, long-term commercial harvesting. Northern regions, unaffected by these factors, served as natural controls. Based on a comparison of data collected in 1980 with pooled data from 1989 and 1992, the hypothesis was supported. Both the growth coefficient (k) and maximum shell length (L ) declined between two time periods in the two southern regions, while during the same time interval, no change occurred in the two northern regions. Differences in growth between regions were often statistically significant. For example, compared with the southern regions, the growth coefficient on Georges Bank was larger, and those clams attained a smaller maximum length. In a comparison of adjacent regions from Delmarva to S. New England, k increased from south to north. This could imply faster growth in cooler water, as well as no relationship between growth and primary productivity. Alternatively, size-selective mortality, imposed by the commercial fishery, was discussed as a mechanism that might account for this unexpected pattern.     

Phylogeography of surfclams,<Emphasis Type="Italic"> Spisula solidissima</Emphasis>, in the western North Atlantic based on mitochondrial and nuclear DNA sequences

The Atlantic surfclam, Spisula solidissima (Dillwyn), is broadly distributed in sandy sediments of the western North Atlantic between the Gulf of St. Lawrence and the Gulf of Mexico. In the United States, a substantial commercial fishery between Long Island and Cape Hatteras harvests offshore populations of one subspecies, S. s. solidissima. A smaller coastal form, S. s. similis Say (also known as S. s. raveneli Conrad), has a partially sympatric geographic distribution, but differs in several life-history characteristics. DNA sequence variation in mitochondrial cytochrome oxidase I (COI) and in introns at two nuclear calmodulin loci was examined to measure genetic divergence between the two subspecies and to test for population structure among populations of S. s. solidissima. Surfclams were collected from seven localities between 1994 and 2001. Based on both mitochondrial and nuclear DNA variation, the two subspecies of S. solidissima are reciprocally monophyletic, with a net COI divergence of 13.9%, indicating long-term reproductive isolation. The only significant differentiation among populations of S. s. solidissima (based on an AMOVA analysis of COI sequences) was between the Gulf of St. Lawrence and more southerly populations. A long internal branch in the S. s. solidissima genealogy coupled with low haplotype diversity in the northern-most population suggests that populations north and south of Nova Scotia have been isolated from each other in the past, with gene exchange more recently. Populations of S. s. similis from Atlantic and Gulf of Mexico coasts had a net COI divergence of 9.2%. Thus, diversification of Spisula spp. clams in the western North Atlantic involved an early adaptive divergence between coastal and offshore forms, with later barriers to dispersal emerging in the offshore form from north to south and in the coastal form between Atlantic and Gulf of Mexico populations.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).     

Age and growth rate determinations for the Atlantic surf clam Spisula solidissima (Bivalvia: Mactracea), based on internal growth lines in shell cross-sections

Marked and recovered surf clams, Spisula solidissima Dillwyn, from Virginia (USA) deposited one internal growth line during a period of 11/2 years, probably in response to spawning during late summer. We have used these annual growth lines to make growth curves for two samples of New Jersey (USA) clams, one from inshore (1.8 km from shore, 15 m deep), the other from offshore (17.5 km from shore, 28 m deep) waters. The offshore and inshore clams grow at approximately the same rate until Age 3 years, after which time the growth rates differ, as does the ultimate lifespan; the offshore clams grow more rapidly and attain a greater age — up to 31 years. Comparison of our growth curves with other published curves revealed a close correspondence to a curve which was based on a study of growth over a 5-year period. Curves based on external growth lines probably underestimate growth rate in early life and overestimate it in later years.     

Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Vertical distribution of bivalve larvae along a cross-shelf transect during summer upwelling and downwelling

Previous time-series studies of meroplankton abundances in the LEO-15 research area off Tuckerton, New Jersey, USA (39°28′N, 74°15′W) indicated short-lived (6–12 h) pulses in larval surfclam (Spisula solidissima Dillwyn) concentration often associated with the initiation of downwelling. To examine possible larval surfclam (and other bivalve) concentrating mechanisms during upwelling and downwelling, six sets of adaptive mobile zooplankton pump samples were taken in July 1998 at different depths at five to six stations along a 25-km transect perpendicular to the coastline and crossing Beach Haven Ridge at LEO-15. Sampling was guided by near real-time, satellite imagery of sea surface temperature overlain by sea surface currents from a shore-based ocean surface current radar (OSCR) unit. A Seabird CTD on the mobile pump frame near the intake provided information on thermocline depth, and sampling depths were adjusted according to the temperature profiles. Near shore, the thermocline was tilted down during downwelling, and up during upwelling. The highest concentrations of surfclam larvae occurred near the bottom at a station near Beach Haven Ridge during downwelling, and just above the thermocline 3 km further offshore during well-developed upwelling. For other bivalve taxa, the larvae were concentrated near the thermocline (Anomia simplex Orbigny and Pholadidae spp.) or concentrated upslope near the bottom (Mytilidae spp.) during upwelling, and the larvae were concentrated near the bottom or were moved downslope during downwelling. Donax fossor Say larvae were found near the surface or above the thermocline during upwelling and downwelling. The general patterns of larval bivalve distribution appear to be influenced by water mass movement during upwelling and downwelling. The larval concentration patterns of individual species are likely a consequence of advection due to upwelling and downwelling circulation, vertical shear in the front region, species-specific larval behaviors, and larval sources.     

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.     

A functional interpretation of cilia and mucocyte distributions on the abfrontal surface of bivalve gills

The lack of fundamental data on the abfrontal surface of bivalve gills has prompted a comparative study of cilia and mucocytes on this surface, using scanning electron microscopy and histology on eight species of bivalves, representing seven families and the four major gill types: Mytilus edulis, Modiolus modiolus, Arca zebra, Placopecten magellanicus, Crassostrea virginica, Spisula solidissima, Mercenaria mercenaria, and Mya arenaria. Abfrontal cilia and mucocytes were found in all species studied, with types and densities differing within and between gill types. The three species of homorhabdic filibranchs presented different densities of abfrontal cilia and mucocytes, from very dense in M. edulis to sparse in A. zebra. The heterorhabdic gills had intermediate cilia and mucocyte densities, with highest concentrations of both abfrontal cilia and mucocytes on the principal filaments. The eulamellibranchs showed low ciliary densities together with high mucocyte densities, especially in S. solidissima, where the abfrontal mucocytes were glandular. These results indicate that: (1) the abfrontal surface is a vestigial mucociliary surface; (2) the abfrontal surface cannot participate in water pumping in most species, due to low ciliary densities; and (3) species with high densities of abfrontal mucocytes could utilize abfrontal mucus to reduce drag, especially in the highly fused gills, such as those of the eulamellibranchs. The differing distributions of abfrontal cilia and mucocytes may reflect different selective pressures acting on the gills within the various taxa. Received: 12 February 2000 / Accepted: 10 September 2000     

Comparative study of the carbon cycle in Venus verrucosa fed on bacteria and phytoplankton

Following a previous study, this paper deals with the utilisation of a phytoplankton suspension (Pavlova lutheri) by the clam Venus verrucosa Linné, 1767 as a function of time. Experimental clams filtered and ingested 100% of the suspension after 2 h. The radioactivity recovered in the whole soft body of the clams was 83.8% after 2 h, 66.0% after 10 h and 46.0% after 41 h. The faeces contained 13.7% after 3 h and 34.3% after 41 h. Carbon-dioxide radioactivity (gas plus dissolved) increased slowly to 12.4% after 41 h. When the water was not changed after 17 h, the clams reabsorbed a great part of their dissolved products and faeces. Under these latter conditions the radioactivity recovered from the whole soft body of the clams was roughly the same as that recovered after the start of the experiment. The aim of this study was the comparison of the consumption of two kinds of food: bacteria and phytoplankton. The work was carried out over three years, from 1977 to 1980.     

Evolutionary relationships of deep-sea hydrothermal vent and cold-water seep clams (Bivalvia: Vesicomyidae): results from the mitochondrial cytochrome oxidase subunit I

Phylogenetic relationships among vesicomyid clams (Bivalvia: Vesicomyidae) and their placement within the order Heterodonta were examined using mitochondrial encoded cytochrome oxidase subunit I (COI) DNA sequences. The presently analyzed vesicomyids represent a recent monophyletic radiation that probably occurred within the Cenozoic. Nucleotide phylogenetic analyses resolved discrete clades that were consistent with currently recognized species: Calyptogena magnifica, C. ponderosa, Ectenagena extenta, C. phaseoliformis, Vesicomya cordata, Calyptogena n. sp. (Gulf of Mexico), C. kaikoi, C. nautilei, C. solidissima and C. soyoae (Type-A). However, specimens variously identified as: V. gigas, C. kilmeri, C. pacifica, and V. lepta comprised two “species complexes”, each composed of multiple evolutionary lineages. Most taxa are limited to hydrothermal-vent or cold-seep habitats, but the “vent” versus “seep” clams do not constitute separate monophyletic groups. Current applications of the generic names Calyptogena, Ectenagena, and Vesicomya are not consistent with phylogenetic inferences. Received: 24 July 1997 / Accepted: 22 August 1997     

Growth and sexual maturity of the northern propellerclam (<Emphasis Type="Italic">Cyrtodaria siliqua</Emphasis>) in Eastern Canada,with bomb radiocarbon age validation

The northern propellerclam Cyrtodaria siliqua is a common bycatch in the Arctic surfclam, Mactromeris polynyma fishery on Banquereau Bank in Eastern Canada. Samples of the propellerclam from this exploited fishery were used to determine the life history characteristics of the population. The age structure of the population is dominated by old animals to ages exceeding 100 years. We validated the age estimates for the propellerclam through analysis of bomb-produced radiocarbon in the shell growth increments deposited before, during and after the atmospheric atomic bomb testing periods of the 1950s and 1960s. Radiocarbon from shells with presumed birth dates between the late 1950s and 1970s clearly reflected the sharp increase in oceanic radiocarbon attributable to previous nuclear testing, indicating that age estimates based on shell increment counts are accurate. Estimates of von Bertalanffy growth parameters revealed that the growth rate of the population was relatively rapid for the first 20 years of life, slowing down to very low growth rates thereafter. Sexual maturity was estimated as being reached at 28.6 mm in length and 4.7 years in age. Size–weight morphometric relationships were also calculated.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Protracted estuarine phase in the life cycle of the marine pufferfish Torquigener pleurogramma

The present study was undertaken to elucidate the way in which the Swan Estuary in south-western Australia is used by the common blowfish Torquigener pleurogramma, a representative of the abundant and widely distributed family Tetraodontidae. T. pleurogramm were collected by beach seine and otter trawl from the Swan Estuary between February 1977 and December 1980 and between May 1984 and February 1986. While T. pleurogramma feeds on a wide variety of organisms in the estuary, the main components of its diet are polychaetes and amphipods for fish <130 mm and bivalve molluscs for larger fish. Numbers of blowfish were inversely correlated with water depth, with densities on the banks (water depth <1.5 m) sometimes reaching 5 fish m^-2, and tended to be greater at night than during the day. The density of T. pleurogramma in the shallows was positively correlated with salinity and inversely correlated with distance from the estuary mouth. Numbers increased greatly in the latter half of 1980 and 1985 as a result of the recruitment of large numbers of the 0+ age class (i.e., fish in their first year of life). Blowfish were represented by seven age classes in the estuary and attained a maximum size of 230 mm (220 g). By the end of their first and second years of life, fish had reached approximately 90 mm (14 g) and 125 mm (39 g), respectively. Sexual maturity was generally not reached until the end of the second year of life. The presence of higher gonadosomatic indices and more mature gonads in fish collected just outside than within the estuary indicate that T. pleurogramma leaves the estuary before spawning. Comparisons between lengthfrequency data, allied with information on the prevalence and intensity of gill parasites, indicate that assemblages in estuarine and neighbouring inshore waters remain distinct for many months and that growth within the estuary is faster than in inshore marine environments.     

Analysis of growth rate in Mya arenaria using the Von Bertalanffy equation

Field studies were conducted in Gloucester, Massachusetts, USA, to determine linear shell growth rates for Mya arenaria. These rates were then compared with those reported for the same species from other locations. Most shell deposition occurred from March through November of each year. Winter interruptions in growth were not as marked in the small clams as in the larger ones (>60.0 mm). Annual variations in growth were slight during the period 1973–1974. Growth of mature clams (>35.0 mm) slowed during the spawning season. No significant sexual dimorphism in mean annual growth rates was detected. Winter rings were shown to be a reliable method for determining age in clams from Gloucester. Age-size relationships, based on two independent measures of annual growth, winter rings and tagging experiments, were computed using the Von Bertalanffy growth equation. No well-defined latitudinal patcerns in growth could be established for M. arenaria.     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.     

Sclerochronological records of temperature and growth from shells of Mercenaria mercenaria from Narragansett Bay,Rhode Island

Annual internal growth increments in shells of hard clams (Mercenaria mercenaria) provide an accurate record of both growth history and some types of environmental change. These increments were used to determine age and growth rate of hard clams collected in 1984–1985 from ten sites in Narragansett Bay, Rhode Island, USA, and to assess geographic variation in growth within the bay. The regional comparisons were facilitated by modeling the clam growth using the von Bertalanffy equation and the parameter of Gallucci and Quinn. The optimum region for hard clam growth was near the head of the bay, with areas further north (Providence River) and south (lower bay) not as productive. The relationship between size and age was similar to that reported for hard clams from northern New Jersey. Using dendrochronological techniques, variations in growth were analyzed on a temporal (year-to-year) basis. The comparative longevity of the hard clams (individuals with 40 annual bands were encountered) demonstrated that sclerochronologies of several decades were possible. A 26 yr growth record based upon 100 individuals was established, covering the years 1959–1984. Significant temporal variations occurred in bivalve growth, and a broad trend of increasing growth indices over the last two decades of the record was noted. The yearly standardized growth index values were highly and positively correlated with mean annual water temperatures in Narragansett Bay for the same time interval. The incremental shell records of these bivalves provide a quantitative indication of marine climatic (temperature) variability and growth which can be used in hindcasting the effects of natural or anthropogenic environmental perturbations.     

UV-absorbing substances in the tunic of a colonial ascidian protect its symbiont, Prochloron sp., from damage by UV-B radiation

The infaunal bivalve Ruditapes decussatus L. was collected from Ria Formosa, Faro, southern Portugal, and subjected to a range of hypoxic conditions and anoxia. Physiological measurements, clearance rates, respiration rates and absorption efficiency were undertaken at slightly different oxygen partial pressures (11, 6, 3 and 1.2 kPa for clearance rates and absorption efficiency and 12, 7, 5, 1.9 and 0.9 kPa for respiration rates). Metabolic rates under hypoxia were measured as oxygen consumption and anoxic metabolism was measured using direct calorimetry. Increasing hypoxia resulted in lower clearance rates, leading to lower ingestion rates and reduced faeces production. Clearance and ingestion rates declined below ˜6 kPa, reflecting decreasing ventilation and feeding activity, although complete cessation was not observed even at 1.2 kPa. Under extreme hypoxia (< 2 kPa) clams showed an irregular behaviour, with valves either closed or only slightly open, and siphons compressed or retracted. Clearance rate was 12% and respiration rate was 35% of normoxic rates. R. decussatus responded to increasing hypoxia by lowering its metabolic rate. Regulation of respiration was absent through moderate hypoxia (˜␣7␣kPa), but was observed in the lower hypoxia range (7 to 0.9 kPa). Under anoxia, rates of heat dissipation were 3.6% of normoxic rates. The low anoxic metabolic rate is indicative of a reduced energy expenditure, and this energy-saving mechanism is common in bivalves. Scope for growth was always pos itive, and even at low oxygen levels clams did not have to utilize their energy reserves. The ability to reduce metabolic costs but still meet the maintenance costs by aerobic catabolism enables R. decussatus to tolerate hypoxia. Such conditions can occur, particularly in the summer, in southern Portugal. Received: 19 July 1996 / Accepted: 17 September 1996     

Intraspecific competition in<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> F</Emphasis>.<Emphasis Type="Italic"> evanescens</Emphasis> (Phaeophyceae: Fucales) germlings: effects of settlement density,nutrient concentration,and temperature

Survival and growth of early post-settlement stages are critical for the development of seaweed populations. Fucoid germlings commonly settle in dense monospecific aggregates, where intraspecific competition and environmental variables (e.g. nutrient concentration and temperature) may affect survival and growth. Using factorial experiments, we determined the effects of settlement density (~10, ~50 and ~250 germlings cm^–2), nutrient enrichment (from ~10 to ~40 µM N and from ~0.5 to ~2.5 µM P), and temperature (7°C and 17°C) on Fucus serratus and F. evanescens germlings in laboratory cultures over 3 months. Settlement density, nutrient concentration and temperature interactively affected growth of germlings, and the magnitude of this interaction varied between the two species. This represents the first record of such factorial interactions in Fucus spp. germlings. Intraspecific competition, estimated as the relative reduction in germling growth and survival from low to high densities, increased with decreasing nutrient concentration and increasing temperature in both species. While temperature and nutrient concentration had little effect on germling size distributions, size inequality and skewness generally increased with germling density, indicating that a few large individuals gained dominance and suppressed many smaller ones at high density. Self-thinning increased with settlement density and depended on nutrient concentration and species at high density. At high density, self-thinning increased with decreasing nutrient levels in F. evanescens, but not in F. serratus. At low density, nutrient enrichment increased germling growth in F. evanescens, but not in F. serratus, whereas growth in both species was stimulated by nutrient enrichment at higher densities. These results suggest that germling growth and self-thinning are more sensitive to variation in nutrient concentration in F. evanescens than in F. serratus. The potential implications of our findings for the understanding of eutrophication-related abundance changes in both species in southern Norway are discussed.Communicated by L. Hagerman, Helsingør