Annual population development and production by Calanus finmarchicus, C. glacialis and C. hyperboreus in Disko Bay, western Greenland

The populations of the copepod species Calanus finmarchicus, C. glacialis and C. hyperboreus were investigated in Disko Bay during a 14-month period in 1996-1997. The three species were predominant in the copepod community. The biomass reached a maximum at the beginning of June (127 mg C m^-3). From the end of July until the end of April the following year, the biomass was <1-6 mg C m^-3. All three species showed seasonal ontogenetic migration. The spring ascent for all three species was just prior to or in association with the break-up of sea ice and the development of the spring bloom, whereas descent occurred over a larger time span during summer. The main overwintering stages were CV for C. finmarchicus, CIV and CV for C. glacialis and C. hyperboreus. Peak abundance of juvenile copepodites, representing the new generation, was in August for C. finmarchicus, in July for C. glacialis and in May/June for C. hyperboreus. From the timing of reproduction and the population development, the life cycles were deduced to be 1 year for C. finmarchicus and at least 2 years for C. glacialis and C. hyperboreus. Secondary production and potential grazing impact of the Calanus community were estimated by two methods based on specific egg-production rates and temperature-dependent production. The Calanus community was not able to control the primary producers during the spring bloom but probably did during post-bloom. The estimates also indicated that grazing on ciliates and heterotrophic dinoflagellates contributes as an essential food source in the post-bloom period.     

Population structure of two Antarctic ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, in winter sea ice

The spatial distribution and population structure of two dominant ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, were studied during winter at nine locations in east Antarctic fast ice. These species accounted for at least 90% of the total metazoan abundance at each location. Abundances were high, reaching 175 individuals l^-1 (190,000 m^-2) for D. glacialis and 660 l^-1 (901,000 m^-2) for P. antarctica. These abundances were probably partly supported by the high biomass of ice-algae (Pearson correlation coefficient, r=0.75), as indicated by chlorophyll-a concentrations (1.7-10.1 µg l^-1). The population structures of each species suggested very different life-history strategies. All developmental stages of D. glacialis were isolated from the ice cores, including females with egg sacs, supporting the hypothesis that this species reproduces in the sea ice during winter. This strategy might assist D. glacialis in leading a continually colonising existence, whereby it responds opportunistically to the availability of favourable habitat patches. The populations of P. antarctica were composed primarily of nauplii (>99%), consistent with past observations of a synchronised life cycle for this species. The strong coupling of the developmental cycle of P. antarctica to the growth and decay of sea ice suggests that local extinctions might occur in areas where ice break-out is unpredictable.     

Population development of the invader ctenophore Mnemiopsis leidyi, in the Black Sea and in other seas of the Mediterranean basin

In the last two decades of the twentieth century, the ctenophore Mnemiopsis leidyi (A. Agassiz) has invaded the Black, Azov, Marmara and Aegean Seas, and, recently, the Caspian Sea. Here, we compare its spatial and temporal distribution, seasonal dynamics and the time and duration of reproduction. We also discuss factors that control its abundance throughout its invasive range and its effect on ecosystems. Observations are based on the long-term field data collected by three research institutes. An analysis of the effects of temperature, salinity, prey (zoo- and ichthyoplankton) availability and predation (by ctenophores of the genus Beroe) on M. leidyi population size, and the effects of M. leidyi on zoo- and ichthyoplankton, and on fish populations in the Black and Azov Seas is also provided. With the Black Sea current, M. leidyi spreads to the upper layers of the Sea of Marmara, where it now occurs around the year. At regular intervals, the Black Sea current also takes it to the northern Aegean Sea. In contrast, it has to re-invade the Sea of Azov every spring or summer, dying out during winter when the temperature drops below 4°C. The warm summer and mild winter temperatures, relatively low salinity and abundance of prey in the Black Sea are close to optimal for M. leidyi, while they are suboptimal in the northern Aegean Sea, where salinity and temperature are often too high. In the Black Sea the absence of gelatinous and other predators led to an enormous ctenophore abundance for a decade, but with the appearance of Beroe ovata in 1999, M. leidyi abundance greatly decreased. Analysis of seasonal dynamics of M. leidyi in the Black Sea and in other seas of the Mediterranean basin indicates similarities in the timing of maximum abundance and biomass, in spite of some differences in the initiation and duration of reproduction. A peak biomass and density occurred in 1989 in the Black and Azov Seas and in 1990 in the other seas. The M. leidyi invasion negatively affected the ecosystems of the Black Sea and the Sea of Azov. The zooplankton, ichthyoplankton and zooplanktivorous fish stocks all underwent profound changes. Similar effects, but less pronounced, were recorded in the Sea of Marmara. Effects on Mediterranean food chains have, so far, remained insignificant. Salinity is probably supraoptimal here, and several predators prevent M. leidyi from reaching outbreak levels.     

Reproductive cycles of three dominant Calanus species in Disko Bay, West Greenland

This study describes the annual reproductive cycles of the three dominant Calanus species, C. finmarchicus, C. glacialis and C. hyperboreus, in Disko Bay (West Greenland) in relation to seasonal phytoplankton development. Relative abundance of females, copepodite stage V (CV) and males, and the developmental stage of the female gonad were examined from plankton samples collected at weekly to monthly intervals from May 1996 to June 1997 with a WP2 net or a pump. During spring and summer, egg production rates were determined. Females of all three species were present year round. Maximum relative abundance was reached by C. hyperboreus females at the beginning of February, by C. glacialis in mid-February, and by C. finmarchicus in April. All three species reproduced successfully in Disko Bay. Their reproductive cycles were considerably different with respect to the timing of final gonad maturation and spawning, and hence in their relation to seasonal phytoplankton development. In all three species, early gonad development took place during winter, before living food became plentiful, suggesting that these processes were largely food independent. Final gonad maturation and spawning in C. finmarchicus was related to the phytoplankton concentration, reflecting that final gonad maturation processes are food dependent in this species. C. glacialis females matured and spawned prior to the spring bloom. Our results indicate that first internal lipid stores and later ice alga grazing supplied final gonad maturation and egg production. Maximum egg production rates of C. glacialis were found in spring and summer, when the chlorophyll a concentration was high. Mature female C. hyperboreus were found from February until mid-April, when the chlorophyll a concentration was still low. In this species, reproductive activity was decoupled from phytoplankton development, and final maturation processes and spawning were solely fuelled by internal energy stores.     

Under-ice amphipods in the Greenland Sea and Fram Strait (Arctic): environmental controls and seasonal patterns below the pack ice

During three "Polarstern" cruises to the ice-covered Greenland Sea (spring 1997, summer 1994, autumn 1995) studies on the under-ice habitat (morphology, hydrography, ice-algal biomass) and on the macrofaunal, autochthonous under-ice amphipods (species diversity, abundance) were carried out in order to describe environmental controls and seasonal patterns in this community. In spring, the ice underside was rather smooth and whitish, while in summer melting structures and sloughed-off ice-algal threads were observed, in autumn detritus clumps accumulated in depressions at the ice underside. Only in summer, a thin layer of warm (up to -0.6°C) and less saline (as low as S=6.3) water was found at the ice-water interface above Polar Water. Integrated ice-algal biomass was highest during autumn (2.6 mg chl a m^-2) and lowest during summer (1.2 mg chl a m^-2). Four species of under-ice amphipods occurred in spring and summer (Apherusa glacialis, Onisimus glacialis, O. nanseni, Gammarus wilkitzkii), but only the last species was observed at the ice underside in autumn. A. glacialis and G. wilkitzkii were equally abundant in spring; A. glacialis dominated in summer. The highest total abundance of amphipods occurred during summer (31.9 ind. m^-2), compared to lower abundances in spring and autumn (5.3 and 1.1 ind. m^-2, respectively). A factor analysis revealed seasonal patterns in the data set, which mainly influenced A. glacialis, and species-specific relations between several environmental factors and the distribution of under-ice amphipods. Abundance of A. glacialis was closely related to the under-ice hydrography and ice-algal biomass, whereas the other amphipod species were more influenced by the under-ice morphology. It is therefore stated that the observed thinning of the Arctic sea ice and the resulting increased meltwater input and change in morphology of floes will have a profoundly adverse effect on the under-ice amphipods.     

Differences in life-cycle traits of<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> originating from 60°N and 69°N,when reared in mesocosms at 69°N

Two different Calanus finmarchicus (Gunnerus) cohorts originating from 60°N (Bergen) and 69°N (Tromsø) were investigated in equal environmental conditions to study their different physiological responses to the same environment. A two-plus-two-bag mesocosm study was carried out between March and July 1998, in Håkøybotn, Tromsø, in order to determine development and mortality rates of the two parallel cohorts of C. finmarchicus. For practical reasons, the cohort from Bergen was incubated 10 days earlier than the Tromsø cohort. Consequently, they were exposed to elevated food conditions as compared to the Tromsø cohort. A high initial mortality among the Bergen cohort could clearly be ascribed, by genetic discrimination, to "contamination" with C. helgolandicus. After this initial mass mortality, the mortality was constantly 0.03-0.04 day^у. In cohorts starting from naupliar stage I, there was no significant difference in development or growth, the median development time (NI-CIV) differing by only 7 days (~6%). The difference in development time can be explained to a large extent (~4 days) by temperature differences. This is substantiated with model simulations using a physiological model developed for C. finmarchicus. There was a time lag in incubation between the two cohorts, resulting in elevated temperature during incubation of the Tromsø cohort. A fraction of both cohorts differentiated sexually at stage CV, with males differentiating before females. Females from both cohorts produced eggs, but specific egg production differed significantly (P>0.001, t-test). This was supported by elevated RNA:DNA ratio in females from the Bergen cohort. Both cohorts demonstrated quite similar development and physiological growth rates and, consequently, are considered to belong to the same genetic population inhabiting the Norwegian Shelf. The study demonstrates that C. finmarchicus is capable of adaptation to changes in environment and, thereby, demonstrates a significant physiological plasticity.     

Mesozooplankton community structure, abundance and biomass in the central Arctic Ocean

During the "International Arctic Ocean Expedition 1991" (20 August-21 September 1991) mesozooplankton was sampled at six stations in the Nansen, Amundsen and Makarov Basins of the central Arctic Ocean from 1,500 m depth to the surface by multiple opening/closing net hauls. Total mesozooplankton abundance decreased from 268 ind. m^-3 in the surface layer (0-50 m) to <25 ind. m^-3 below 200 m depth. The small copepods Oithona similis and Microcalanus pygmaeus, as well as copepod nauplii, were most abundant close to the surface, while Oncaea borealis and Spinocalanus spp. frequently occurred at greater depth. Mesozooplankton dry mass (DM) integrated over the upper 1,500 m of the water column was surprisingly stable throughout the investigation area and measured 2.0ǂ.3 g DM m^-2. Dry mass in the upper 50 m measured 20.9 mg m^-3 and was dominated by Calanus hyperboreus (57.4%) and C. glacialis (21.1%). C. finmarchicus was very abundant only in the Nansen Basin. Below 200 m the calanoid copepods Metridia longa, Microcalanus pygmaeus and Pareuchaeta spp., the decapod Hymenodora glacialis and chaetognaths of the genus Eukrohnia were the principal contributors to biomass values of <1 mg DM m^-3. Hence, vertical changes in abundance, biomass and species composition were much more pronounced than regional differences between the basins. Three different mesozooplankton communities were differentiated according to their faunistic composition and are discussed in context with the major water masses: Polar Surface Water, Atlantic Layer and Arctic Deep Water.     

Diet composition and quality for <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> egg production and hatching success off south-west Iceland

Egg production and hatching success of the copepod Calanus finmarchicus was measured during spring and summer in the waters south-west of Iceland. Egg-production rates varied greatly, both temporally and spatially, with highest average rates found at a station with low chlorophyll-a concentrations (0.4 mg m^-3). Excluding this high production rate from statistical analysis, the remaining egg-production rates were found to be positively correlated with phytoplankton biomass, as well as with parameters representing healthy phytoplankton condition, food quality and diatom-type fatty acids. Hatching success of eggs was negatively correlated with some saturated and monounsaturated fatty acids related to phytoplankton senescence.     

Burrowing behaviour and sediment reworking in the heart urchin Brissopsis lyrifera Forbes (Spatangoida)

Sediment reworking due to burrowing and feeding was studied in the spatangoid Brissopsis lyrifera, at two different temperatures (7°C and 13°C). Spine activity and burrowing behaviour were recorded with a real-time video camera. Reworked sediment volume was calculated from tracks produced by the heart urchin. Ingestion rates were measured by feeding the heart urchins with luminophores. Temperature had a significant effect on the bioturbation activity of B. lyrifera. At 13°C reworked sediment volume due to burrowing was 22 ml sediment h^-1 and at 7°C 14 ml sediment h^-1. The ingested amount of sediment was 0.08 and 0.02 g dry sediment h^-1 in 13 and 7°C, respectively. Reworked sediment volume due to burrowing was 60-150 times higher than the volume ingested. The large reworked volume is a consequence of B. lyrifera moving with a rocking motion through the sediment. The spines were continuously transporting sediment around the test with 5-min metachronal wave cycles.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

Large viruses and infected microeukaryotes in Ross Sea summer pack ice habitats

A variety of Ross Sea summer pack ice habitats between 66 and 75°S were examined for viruses 𔒦 nm capsid diameter. Maximum abundances of these viruses likely to infect eukaryotes were 10⁶-10⁷ ml^-1 brine in surface, interior, and bottom habitats and constituted up to 18% of the total (all sizes) viruses. There is abundant ultrastructural evidence for infection of a variety of microheterotrophs and some autotrophs. One station exhibited the classical characteristics of a lower latitude algal bloom with potential viral control. The blooming alga, Pyramimonas tychotreta Daugbjerg 2000, was infected, as were two abundant heterotrophs, Cryothecomonas spp. and an unidentified flagellate, that fed on P. tychotreta. Infections were observed in only one life history stage (multiflagellate cells) of P. tychotreta, suggesting a relationship among virus-induced lysis, life-history stages, physiology, and environmental factors regulating the life cycle. There is good evidence that diatoms are not a likely source of the large viruses, and viruses in general are not a major food source for ice microheterotrophs in summer.     

Effects of the 1996 and 1998 positive sea-surface temperature anomalies on corals,coral diseases and fish in the Arabian Gulf (Dubai,UAE)

Two positive sea-surface temperature anomalies occurred in the Arabian Gulf in short sequence. Between May and August 1996 and 1998, sea-surface temperatures in the southern Arabian Gulf were elevated by 2°C above average. The consequences for coral fauna, coral diseases and coral regeneration were studied in Dubai (United Arab Emirates) between Jebel Ali and Ras Hasyan. In 1996, coral death was widespread, affecting primarily the genus Acropora. In Acropora-dominated areas, live coral cover was reduced from 90% to about 26% in 1996, while in 1998 only a reduction from 26% to 22% of the remaining coral cover occurred. In the study area, all six Acropora species suffered total mortality in 1996, thus the coral fauna was reduced from 34 species to 27. The nearest areas with surviving Acropora were 30 km to the east (Deira) and 20 km to the west (Al Jazira). Massive coral species suffered negligible mortality, and slowly increased in space cover. The Acropora overkill turned 7.9 km² (19.7% of total coral-covered area) of previously lush coral gardens into a dead framework that was increasingly bioeroded. Acropora recruitment only started in 1998, average recruit size in 1999 was 7Dž cm, and recruits were rare. Prior to the mass mortality event, coral diseases were common and seasonal (14LJ% of corals, mainly Acropora, affected in summer, in winter 7Lj%, mainly massives), after the mortality event seasonality was lost and infection remained below winter levels (6LJ%, only massives infected). In fish, overall species richness decreased from 95 to 64 species in point counts, but frequency only decreased in one species (Pseudochromis persicus). Guild structure changed inasmuch as herbivores and planktivores increased, and invertivores decreased, although differences were not statistically significant. The most abundant family, both prior to and after the coral mass mortality, was Lutjanidae. It appears that even though much of the coral was dead, the maintenance of structural complexity allowed the fish assemblage to avoid a similar catastrophic change to that experienced by the coral assemblage.     

Oxygen and carbon stable isotopic profiles of the fan mussel, Pinna nobilis, and reconstruction of sea surface temperatures in the Mediterranean

Stable oxygen and carbon isotope profiles ('¹⁸O_skeletal and '¹³C_skeletal), taken along the direction of growth from the umbo to the shell margin in shells of the pinnid Pinna nobilis, were used to reconstruct sea surface temperatures (SST) in the south-east Mediterranean and ontogenetic records of metabolic CO₂ incorporation. Comparison of the seasonal cycle of SST, predicted from the '¹⁸O_skeletal record of a small (young) rapidly growing pinnid and temperature measured with a continuous in situ recorder showed that P. nobilis calcifies under isotopic equilibrium with surrounding seawater, thus indicating that P. nobilis shells can be used as a reliable predictor of SST. A 10-year SST record for the south-east Mediterranean was reconstructed from the shell profiles of four pinnid shells of different sizes and ages collected in 1995 and 1996. Reliable resolution of the seasonal SST could only be achieved during the first 4 years of shell growth. As the pinnids grew older, the temperature record was poorly resolved because the shell growth had diminished with age, resulting in time-averaging of the record. The amplitude of the generated seasonal temperature cycle compared favourably (DŽ°C) with a long-term temperature record from northern Mediterranean waters. Clear seasonal cycles in '¹³C_skeletal were observed with an amplitude of ~1.0‰, similar to the calculated seasonal changes in '¹³C of seawater (0.6‰) overlying seagrass meadows. An ontogenetic trend towards less positive '¹³C_skeletal values was too large to be attributed to any decrease in '¹³C in seawater resulting from the invasion of anthropogenic CO₂. It is suggested that the temporal changes of '¹³C_skeletal are due to incorporation of respiratory CO₂ into the extrapallial fluid and reflect changes in the metabolic activity of the pinnid rather than changes in the isotopic composition of dissolved inorganic carbon within the surrounding seawater.     

Size-related trophodynamic changes in three target fish species recovering from intensive trawling

The consequences for food-web structure of protecting marine communities from trawling have been little explored. We investigated the Gulf of Castellammare, northwest Sicily, where a trawl ban initiated in 1989 has led to great increases in the biomass of some target species. Using stable isotopes of carbon and nitrogen, our objectives were to see if mean trophic level and omnivory had increased after the ban in three of the most common fishery-target species, namely, the anglerfish Lophius budegassa, Mediterranean hake Merluccius merluccius, and red mullet Mullus barbatus. We compared size data from before and after the fishery closure, but we also compared recently derived data from outside and inside the closed area. In all three species '¹³C and '¹⁵N were found to differ markedly depending on fish size. '¹³C was found to decrease in L. budegassa and Merluccius merluccius but increase in Mullus barbatus with increasing fish length. '¹⁵N increased in all three species, and this was thought to reflect feeding at increasingly higher trophic levels during the animals' lives. Mean length, and therefore '¹⁵N-derived trophic level, increased after the trawl ban only in the anglerfish L. budegassa. Based on '¹³C and '¹⁵N data, none of the species became more or less omnivorous after the ban. After 9 years of no trawling, increases in numerical abundance were not accompanied by substantial size-related trophodynamic shifts in any of the three species of fish studied.     

Settlement patterns of newly settled plaice (Pleuronectes platessa) in a non-tidal Swedish fjord in relation to larval supply and benthic predators

In demersal fish species with a pelagic larval stage, settlement patterns may be a consequence of variations in larval supply, habitat selection at settlement, and processes acting between the time of settlement and the time of benthic sampling. This study describes temporal (1994-1998) and spatial variation in plaice (Pleuronectes platessa L.) settlement densities in four semi-isolated nursery areas with similar habitat characteristics, in the non-tidal Gullmarsfjord on the west coast of Sweden. Juvenile abundance varied by a factor of ten, both among years and among nursery grounds. For the 3 years when larval sampling was undertaken (1994-1996) and all nursery areas, there was a significant positive relationship between larval supply and juvenile abundance (linear regression: r²=0.45, n=24, P<0.001). On the southern side of the fjord, a significant positive relationship between larval and juvenile abundance was found in one area (r²=0.62, n=6, P<0.05). The absolute mortality rate of plaice after settlement was related to the initial settlement density (r²=0.95, n=20, P<0.001), and to the abundance of predatory shrimps Crangon crangon (r²=0.44, n=20, P<0.01). Plaice otoliths were found in 6% of the shrimp stomachs analysed from an area with high density (13.3 m^-2) of newly settled plaice. The present study suggests that the density of juvenile plaice was limited by larval supply to the nursery grounds. Consistency in the relative abundance of juveniles among nursery grounds between years also suggested that some nursery areas may be in the settlement shadow of others. The irregular nature of the coastline in combination with larval depletion could thereby cause small-scale (10³-10⁴ m) variation in settlement densities of the same order of magnitude as the inter-annual variability in recruitment to individual nursery grounds.     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Combined effect of temperature and food concentration on the grazing rate of the rotifer Brachionus plicatilis

We developed a predictive relationship to determine the grazing rate of Brachionus plicatilis at given temperatures and food concentrations; this function could be applied to experimental culturing and aquaculture practices. Grazing experiments were conducted at temperatures between 5°C and 40°C and at food concentrations, of the flagellate Isochrysis galbana, ranging between ~0 and 10⁶ ml^-1. In total, 136 grazing rates were determined, using the prey depletion method, for rotifers acclimated to treatments for 0.5 or 4 h. The response of grazing rate to temperature and food concentration was described using a model that combined a rectangular hyperbolic function for food concentration and a sigmoidal function for temperature. Using non-linear curve-fitting methods an equation was obtained: G=(452F)/(159000+F)Ǵ.94/(1+2190002T^-4.35) , where G is the grazing rate (flagellates rotifer^-1 min^-1), F is the food concentration (flagellates ml^-1), and T is temperature (°C). The equation indicates a maximum grazing rate of ~35 prey rotifer^-1 min^-1, above ~4᎒⁵ prey ml^-1 and 25°C.     

Temperature acclimation of upper and lower thermal limits and freeze resistance in the nonindigenous brown mussel, Perna perna (L.), from the Gulf of Mexico

Acute and chronic upper and lower thermal limits and freeze resistance were investigated in the nonindigenous brown mussel, Perna perna, from the Texas Gulf of Mexico coast in order to assess its potential distribution in North American coastal waters. This species' long-term, incipient lower and upper thermal limits were 7.5°C and 30°C, congruent with the seasonal ambient water temperature range of 10-30°C reported for other populations worldwide. Effects of temperature acclimation and individual size on survival time were most pronounced on chronic exposures to lethal temperatures approaching incipient lower or upper thermal limits. When exposed to temperature increasing at 0.1°C min^-1, the acute upper lethal limit was 44°C regardless of acclimation temperature or individual size. P. perna had a limited freeze resistance, being intolerant of emersion at -2.5°C. This species' narrow incipient thermal limits, limited capacity for temperature acclimation and poor freeze resistance may account for its restriction to subtidal and lower eulittoral zones of cooler subtropical rocky shores. Near extinction of P. perna from Texas Gulf of Mexico waters occurred in the summer of 1997 when mean surface-water temperatures approached its incipient upper limit of 30°C.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.     

Abundance, growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the Oyashio region, western subarctic Pacific

Abundance and life-cycle features of the mesopelagic hyperiid amphipod Primno abyssalis (formerly P. macropa) in the Oyashio region, western subarctic Pacific, were investigated using samples collected between July 1996 and July 1998. P. abyssalis was collected throughout the entire survey period, with abundance peaks occurring in spring to autumn. While all maturity stages of males and females were observed throughout the study period, the peak reproduction season was in summer. Instar analysis based on the segment number of the pleopod rami indicated that hatched juveniles molted 10 times before becoming adult males and 13 times before becoming adult females. Judging from the dry and ash-free dry weights of each instar, males and females continued to feed throughout the final instar stage. Based on cohort analysis of seasonal samples and laboratory observations on molting frequencies, growth in body length of P. abyssalis was linear with time, and estimated generation lengths were 2.3-3.8 years for females and 1.4-1.9 years for males. Brood size of females ranged from 66 to 337 and increased with increasing female body length. Lifetime fecundity, calculated as the sum of six successive broods, was 1,004. Compared with P. abyssalis in the southern Sea of Japan, those in the Oyashio region have a larger number of adult instars (six versus five for females, three 3 vs one for males), a lower growth rate (0.014 mm day^-1 vs 0.021 mm day^-1), and mature earlier (instar 13 vs instar 15 for females; instar 10 vs instar 11 for males). These characteristics are considered to be advantageous life-history traits to counteract higher niche competition within the mesopelagic community and higher predation pressure by mesopelagic fishes in the Oyashio region than in the Sea of Japan.