Morpho-functional defences of Mediterranean sea urchins,<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> and<Emphasis Type="Italic"> Arbacia lixula</Emphasis>, against fish predators

Morpho-functional features potentially involved in defence mechanisms against fish predators (i.e. attachment tenacity, spine length, and test robustness and thickness) have been assessed in two Mediterranean sea urchins, Paracentrotus lividus and Arbacia lixula. All four morpho-functional features were significantly and positively related to individual size for both species of sea urchins. Test robustness (i.e. static load needed to break sea urchin tests) was significantly greater for A. lixula (from 3,450 to 15,000 g depending on size) than for P. lividus (1,180–11,180 g). Attachment tenacity (i.e. force needed to dislodge sea urchins from the rocky substrate) was greater in A. lixula (280–3,300 g) than in P. lividus (110–1,450 g), and the difference tended to decrease in relation to smaller sea urchin size. Spine length was greater in A. lixula (1.5–2.9 cm) than in P. lividus (0.5–2.3 cm), but the difference decreased for larger sea urchin size. Test thickness was slightly greater (but not significantly) in A. lixula (0.35–1.10 mm) than in P. lividus (0.12–0.90 mm). These results provide evidence that morpho-functional features of sea urchins could be involved in affecting predation rates by fishes upon P. lividus and A. lixula, with potential implications for the population structure and distribution patterns of the two sea urchins in shallow rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

Utilization of a fumigated sediment by two benthic deposit-feeders:Abra alba (Mollusca: Bivalvia) andEupolymnia nebulosa (Annelida: Polychaeta)

The present study tested the utilization of dead microbial biomass by two benthic deposit-feeders:Abra alba (Wood) (Mollusca: Bivalvia) andEupolymnia nebulosa (Montagu) (Annelida: Polychaeta). Clams were collected in the Canet lagoon during spring 1989. Worms were collected in the Port-Vendres harbour during spring 1989. The¹⁴C-labelled (glutamic acid, 24 h) sediment used during the study was sterilized with 1% chloroform, washed with sterile seawater, and dried (60°C; 48 h). This sterilisation procedure, called fumigation is the least harmful to the sediment (Novitsky 1986). Both clams and worms were incubated in the presence of the fumigated sediment for 5, 10, 20, and 50 h. At the end of each experiment we recorded the radioactivity in four compartments: (1) sediment, (2) dissolved organic matter (DOM), (3) CO₂, and (4) animals. The radioactivity of the sediment was subdivided into five fractions: (i) soluble in 2N HCl, (ii) soluble in hot 5% trichloroacetic acid (TCA), (iii) soluble in 1N NaOH, (iv) soluble in hot 6N HCl, (v) residual (after combustion in a Leco carbon analyser). In the first set of experiments, after 20 h of incubation, 5.4 and 4.7% of the total radioactivity was taken up by clams and worms, respectively. However, a model revealed that this uptake could have been correlated with the release of radiolabelled DOM (33% of total radioactivity during the first 5 h). In order to test this assumption, we used the same protocol with three additional washes of the fumigated sediment. This resulted in a significantly lower uptake by the clams (1.9% of the total radioactivity byt = 50 h), whereas the worms exhibited an uptake similar to that in the initial experiment (5.1% of total radioactivity byt = 50 h). These results underline the importance of considering interactions with DOM when applying radiotracer techniques to the study of benthic food chains. The average ingestion rates of fumigated sediment byA. alba andE. nebulosa were 5.2 10^–2 mg sediment dry wt mg^–1 clam h^–1 and 3.5 10^–2 mg sediment dry wt mg^–1 worm h^–1, respectively, which is comparable to previous data reported for other deposit-feeding bivalves and polychaetes feeding on natural sediment or detritus. The low radioactivity recorded for CO₂ together with the similarity of the changes in the partitioning of the radioactivity within the sediment between control experiments and experiments carried out in the presence of clams or worms suggest low assimilation efficiencies. Therefore, the present study supports the fact that dead microbial biomass does not constitute an important food source for benthic deposit-feeders.     

Population age structure and rhizome growth of<Emphasis Type="Italic"> Cymodocea nodosa</Emphasis> in the Ria Formosa (southern Portugal)

The population age structure and derived population dynamics (recruitment, growth and mortality), rhizome growth and flowering effort of Cymodocea nodosa in the Ria Formosa (south of Portugal) were examined using reconstruction techniques. The horizontal rhizome elongation rates were low (13.8–30.7 cm year^–1), which is consistent with the low population growth rate (–2.35 to 0.29 year^–1). The vertical elongation rate was proportional to the horizontal growth except for one stand that was subjected to sand movements and had higher elongation rates. The mean leaf production varied from 11.8 to 14 leaves year^–1. The average shoot age varied from 0.70 to 1.04 years, and the oldest shoot found was 7.6 years. The mortality rates were the highest as yet reported for this species (0.99–3.70 year^–1). Observed low growth and high mortality are possibly associated with the development of eutrophic conditions in the Ria Formosa. The flowering probability found in Ria Formosa stands (<0.009–0.90 shoots^–1 year^–1) was comparable to that found in the Mediterranean and off the Canary Islands. Only two stands with flowering events were found inthe Ria Formosa population, which indicates that the stands are highly dependent on clonal growth to maintain themselves inthe Ria Formosa.Communicated by O. Kinne, Oldendorf/Luhe     

Tolerance of Mediterranean seagrasses (<Emphasis Type="Italic">Posidonia oceanica</Emphasis> and <Emphasis Type="Italic">Cymodocea nodosa</Emphasis>) to hypersaline stress: water relations and osmolyte concentrations

The present study examines for the first time the effects of increased salinity on water relations and osmolyte (carbohydrates and amino acids) concentrations in two Mediterranean seagrass species, Posidonia oceanica and Cymodocea nodosa, which are adapted to growth in environments with contrasting salinity and have a known differential sensitivity to alterations in ambient salinity. The specific aim was to obtain insights into their respective capacities to cope with natural or anthropogenically induced (e.g. desalination plants) hypersaline stress and its ecological implications. To this end, large plant fragments of both seagrass species were maintained for 47 days in a laboratory mesocosm system under ambient salinity (37 psu; control) and three chronic hypersaline conditions (39, 41 and 43 psu). Analyses of leaf-tissue osmolality indicated that both species followed a dehydration avoidance strategy, decreasing their leaf water potential (Ψ_w) as the external salinity increased, but using different physiological mechanisms: whereas P. oceanica leaves exhibited a reduction in osmotic potential (Ψ_π), C. nodosa leaves maintained osmotic stability through a decrease in turgor pressure (Ψ_p) probably mediated through cell-hardening processes. Accordingly, the concentrations of soluble sugars and some amino acids (mainly Pro and Gly) suggested the activation of osmoregulatory processes in P. oceanica leaves, but not in C. nodosa leaves. Osmotic adjustments probably interfered with leaf growth and shoot survival of P. oceanica under hypersaline stress, whereas C. nodosa showed a more efficient physiological capacity to maintain plant performance under the same experimental conditions. These results are consistent with the more euryhaline ecological behaviour of C. nodosa and contribute to understanding the high vulnerability shown by P. oceanica to even mild increments in seawater salinity.     

Adaptations of subtropical Venus clams to predation and desiccation: endurance of<Emphasis Type="Italic"> Gafrarium tumidum</Emphasis> and avoidance of<Emphasis Type="Italic"> Ruditapes variegatus</Emphasis>

Intertidal endobenthic bivalves are often dislodged from sediments by hydrodynamic forces. As a result, they encounter the dangers of predation and desiccation, which are generally harsh near the sediment surface. To cope with such dangers, the bivalves possibly possess: (1) a strong body to endure predation and desiccation stress, (2) quick mobility to avoid the stresses, or (3) a high growth rate for attaining a size refuge. The present study examined which of these modes are adopted by the subtropical cobbled-shore Venus clams Gafrarium tumidum (Röding, 1798) and Ruditapes variegatus (Sowerby, 1852), revealing the following interspecific differences. (1) G. tumidum survived better than R. variegatus did in harsh experimental conditions, namely: the experimental cages exposed to predation and desiccation on a cobbled shore; a laboratory aquarium with a predatory crab Scylla serrata; and ovens with high temperatures (27°C and 34°C). (2) R. variegatus was more mobile than G. tumidum was, digging into the sediment on a cobbled shore more rapidly at both high and low tides. (3) The two species with shell lengths 20–30 mm showed similar growth rates (median: –0.2 to 44.5 m day^–1) in seasonal mark–recapture surveys over 2 years. Overall, to cope with the dangers of predation and desiccation G. tumidum appears to have a strong body, while R. variegatus displays rapid mobility, and neither species seems to attain a size refuge through rapid growth. Such species-specific modes are discussed in relation to the interspecific differences found in shell morphology.Communicated by T. Ikeda, Hakodate     

Primary production of Posidonia oceanica in the Mediterranean Basin

Primary production of the marine phanerogam Posidonia oceanica (Linnaeus) Delile was measured by lepidochronological analyses at 22 sites in the Mediterranean Sea (Corsica, France, Italy, Sardinia and Turkey), between 1983 and 1992, to determine spatial and temporal variations. Leaf production (blade and sheath) ranged from 310 to 1 540 mg dry wt shoot^–1 yr^–1, depending on site and depth. Rhizome production ranged from 24 to 120 mg dry wt shoot^–1 yr^–1 (6% of average leaf production). At some sites the results obtained by lepidochronological analysis were consistent with earlier results obtained by classic methods (e.g. leaf-marking). While primary production per shoot (mg dry wt shoot^–1 yr^–1) displayed no significant differences between sites, primary production of the P. oceanica meadow (g dry wt m^–2 yr^–1) decreased with increasing depth at all sites studied. This decrease correlated with reduced density of the meadow (number of shoots per m²) with increasing depth. Past primary production was also extrapolated at three sites at the island of Ischia (Italy) for a period of 5 yr in order to determine interannual variations over a period of several years. While major variations were recorded for the surface stations (5 and 10 m depth), production remained stable at the deepest station (20 m depth). Given the large geographical scale of the study (location, depth range), it would appear that while P. oceanica production remains considerable, the values recorded in the literature on the basis of classical analyses (surface stations) represent maxima, and cannot be generalised for meadows as a whole.     

Habitat associations and diet of young-of-the-year Pacific cod (<Emphasis Type="Italic">Gadus macrocephalus</Emphasis>) near Kodiak,Alaska

The influence of environmental variables and habitat on growth and survival of juvenile gadoid species in the Atlantic has been clearly demonstrated; conversely, in the North Pacific little is known about the habitat and ecology of juvenile Pacific cod (Gadus macrocephalus Tilesius). The hypothesis that density of young-of-the-year (YOY) Pacific cod in nearshore habitats is predicted by shallow depth and the presence of eelgrass and macroalgae was tested in a variety of nearshore habitats adjacent to commercial fishing grounds near Kodiak Island, AK. From 10 to 22 August 2002, a beach seine and small-meshed beam trawl were used to capture YOY Pacific cod (n = 254) ranging from 42 to 110 mm fork length. Depth, water temperature, salinity, sediment grain size, and percent cover by emergent structure (i.e. tube-dwelling polychaetes, sea cucumber mounds, macroalgae) were measured prior to fishing. Density of YOY Pacific cod was highest in areas of moderate depth (15–20 m) and positively and linearly related to percent cover by sea cucumber mounds and to salinity. No previous studies have documented fish utilizing sea cucumber mounds as habitat. Furthermore, eelgrass and macroalgae were inconsequential to cod distribution. Diets consisted mainly of small calanoid copepods, mysids, and gammarid amphipods and were significantly related to cod length and depth. This work provides important information on previously undocumented factors that affect distribution and feeding of YOY Pacific cod, which ultimately influence growth and survival in this species.     

The ecology of cirratulid mudballs on the Oman margin, northwest Arabian Sea

Mudball-building cirratulid polychaetes have been described previously only from the southern California margin. During a study of oxygen minimum-zone benthos in fall 1994, we observed dense aggregations of agglutinated mudballs at 840 to 875 m on the Oman margin in the northwest Arabian Sea. These were inhabited, and probably constructed, by a cirratulid polychaete species in the genus Monticellina. The mudballs were cigar-shaped, 4.5 to 25 mm long, and positioned vertically so as to protrude several millimeters above the sediment–water interface. Total mudball densities were ∼16 000 m⁻². Occupied mudballs occurred at densities of 2 112 m⁻²; 89% were in the uppermost 2 cm of sediment, and no occupied mudballs were found below 10 cm. Organisms other than the cirratulid were present on 1.7% of the mudballs examined, and included epizoic polychaetes, agglutinated and calcareous Foraminifera. Various polychaetes, a nemertean and nematodes were found inside tests. Mudball abundance exhibited positive associations with densities of several paraonid polychaete species, and with densities of burrowing and subsurface-deposit-feeding polychaetes. Negative associations were observed between mudballs and three tube-building taxa (two polychaetes and an amphipod). Mudball-inhabiting cirratulids are abundant in at least two low-oxygen, margin settings. We expect further sampling of bathyal environments to yield additional systems in which cirratulid mudballs are common. Such observations are valuable because mudballs appear to represent a significant source of heterogeneity that can influence macrofaunal community structure in deep-sea sediments. Received: 15 January 1997 / Accepted: 30 January 1997     

Effects of temperature,salinity, and predators on mortality of and colonization by the invasive hydrozoan<Emphasis Type="Italic"> Moerisia lyonsi</Emphasis>

The successful invasion of non-indigeneous species depends on initial colonization as well as establishing a self-maintaining population. The invasive hydrozoan Moerisia lyonsi (Boulenger, 1908), possibly originating from low-salinity waters in the Black Sea and Middle East regions, has become established in low-salinity waters in several estuaries of North America, including Chesapeake Bay. The effects of temperature and salinity on mortality of M. lyonsi polyps were examined in the laboratory in February 2001 in the presence of abundant food. The polyps of M. lyonsi were directly transferred from 20°C and 10 salinity to one of 45 combinations of temperature (10–29°C) and salinity (1–40). Polyp mortality within 7 days occurred only in low-temperature treatments with salinities of 35–40. Surviving polyps reproduced asexually in salinities of 1–40 at 20–29°C, and in salinities of 1–25 at 15°C, but not in any salinities at 10°C. The greatest asexual reproduction rates, an index for population survival potential, occurred at salinities of 5–20. Survival and reproduction of M. lyonsi over such broad temperature and salinity ranges indicate that M. lyonsi may colonize and establish populations throughout the Chesapeake Bay; however, M. lyonsi medusae were reported only at salinities <9.3 there. This discrepancy may be due to the effects of predators. The scyphomedusan Chrysaora quinquecirrha (Desor, 1848), but not the ctenophore Mnemiopsis leidyi (A. Agassiz, 1865) consumed M. lyonsi medusae in laboratory experiments in August–September 2001. Populations of M. lyonsi do not appear to be limited by temperature and salinity conditions; however, their distribution in Chesapeake Bay may be restricted to low salinities not inhabited by predators.Communicated by J.P. Grassle, New Brunswick     

The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds

Benthic filter feeding macrofauna organisms may be an important factor linking sediment and water column. Transport of water and concomitantly of suspended matter is directly related to the size of the benthic filter feeding population. This paper aims to determine the potential for water transport by the bivalve Mya arenaria along a coastal stretch of roughly 100 km length in the southern Baltic Sea. Quantification of population filtration rates specific to the area is based on distribution, abundance and biomass of M. arenaria and calculated according to previously published filtration rate–biomass relations. Calculated rates range up to >8 m³ m^–2 day^–1 (at 5–20 m water depth in sandy sediment) with the potential to locally process a volume of water equivalent to the water column within <1 day. Data from 1991–2002 at one site suggest that the area-specific potential population filtration rate remains remarkably constant in time despite changes in population structure of M. arenaria. The related impact on pore water exchange within the permeable sediment associated with the leakage of water from the gape of M. arenaria valves is discussed.Communicated by L. Hagerman, Helsingør     

Behavior and growth ofMercenaria mercenaria during simulated storm events

Experiments were conducted in April–August 1989 on juvenileMercenaria mercenaria (L.) in an oscillatory water tunnel to simulate resuspension of bottom sediments by waves and to determine the effects of shortterm storm events on particle ingestion, pseudofeces production, and shell growth. Juveniles (mean length = 19.2 mm) were subjected to identical concentrations of algae in both low-flow, gentle waves (maximum velocity = 7 cm s^–1) and high-velocity storm waves (maximum velocity = 22 cm s^–1). Suspended sediment levels reached 193 mg 1^–1 at 1 cm above the bed during storms. Shell growth decreased by a maximum of 38% during the storm when levels of phytoplankton were high (average cell concentration = 43 × 10⁶ cells 1^–1), and by 18% when phytoplankton levels were low (av cell conc = 6 × 10⁶ cells 1^–1). Orientation of clam siphons was not related to flow direction. Significantly more pseudofeces were produced when the clams were subjected to increased sediment resuspension under waves, and in troughs of sand ripples. The size of sediment grains ingested did not vary significantly among the flow treatments. The decrease in shell growth during storms may be due to a reduction in filtration rate coupled with a decrease in net energy gained from filtration due to costs of pseudofeces production. The magnitude of the decrease seems to be related to concentration of algae, water temperature, age of clams and sediment transport mode (bed load or suspended load). Thus, the interpretation of growth increments must be made in the context of these environmental variables.     

Benthic macrophyte communities in the Ghar El Melh lagoon (North Tunisia)

This work deals with the biodiversity and distribution of benthic macrophytes in the Ghar El Melh lagoon, a Mediterranean coastal lagoon located in the North of Tunisia. An inventory was made of the benthic flora and submerged macrophyte communities were mapped during two successive campaigns (the summer of 1999 and the winter of 2000). The following 24 macrophyte species were identified: seven red algae, two brown algae, 11 green algae, and four marine angiosperms. The results were compared with available data from the literature. Ruppia cirrhosa is the most dominant species. It is found in all lagoon parts, except in the west sector.Ruppia beds are usually associated withCladophora forming heterogeneous communities. During summerRuppia cirrhosa shows a large distribution, covering an area of ca. 21.4 km², with dense, extensive beds covering 80–100%. In winter, severalCladophora species have a very large distribution as well, covering nearly an area of 28.5 km² with an average cover of 46%. The green algaeCaulerpa prolifera is confined to the eastern part of the lagoon which is mainly affected by seawater. In comparison with previous situations, many transformations were observed in biodiversity and spatial distribution of the dominant communities. Thus,Cymodocea nodosa andZostera beds, which dominated in the 1970s, were replaced byZostera andCaulerpa prolifera in the 1980s and are currently succeeded byRuppia cirrhosa andCladophora. Restoration of the Ghar El Melh lagoon will enable an increase in the exchange with the open sea and the circulation of water, in particular in the confined zones. This should considerably improve the water quality and would positively influence the phytobenthic communities.     

Rates of primary productivity and growth in<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> measured at different depths,over an annual cycle,at West Island,South Australia

The pattern of growth (biomass accumulation) in Ecklonia radiata throughout the year and across a depth profile was investigated using the traditional hole-punch method, and the information presented in context with concurrently measured in situ net productivity rates. The rate of net daily productivity showed a lack of consistent seasonal variability, remaining constant throughout the year at two of the four depths measured (3 m and 12 m), and becoming higher during winter at another (5 m). Throughout the year, rates of net daily productivity differed significantly across the depth profile. Net daily productivity rates averaged 0.017 g C g^–1 dwt day^–1 and 0.005 g C g^–1 dwt day^–1 at a depth of 3 m (1,394 mol O₂ g^–1 dwt day^–1) and 10 m (382 mol O₂ g^–1 dwt day^–1) respectively. In contrast, the biomass accumulation rate of E. radiata was highly seasonal, with low rates of growth occurring in autumn (0.002 g dwt g^–1 dwt day^–1 at both 3 and 10 m) and summer (0.007 and 0.004 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and higher rates in spring (0.016 and 0.007 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and winter (0.015 and 0.008 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively). The proportion of assimilated carbon used for biomass accumulation varied throughout the year, between 5% and 41% at 3 m and between 28% and 128% at 10 m. The rates of biomass accumulation at all depths represented only a small proportion of the amount of carbon assimilated annually.Communicated by P.W. Sammarco, Chauvin     

Comparative analysis of macrofaunal species richness and composition in Posidonia oceanica, Cymodocea nodosa and leaf litter beds

We investigated macrofaunal species richness and composition in Posidonia oceanica, Cymodocea nodosa and Leaf litter beds within a coastal area of the Gulf of Oristano in proximity of the Cabras lagoon (western Sardinia, Italy). A total of 124 taxa were found, of which 116 were identified at the species level. They were analyzed based on both taxonomic and substrate affinity classification. Presence/absence analysis revealed that P. oceanica, C. nodosa and Leaf litter were all characterized by a conspicuous number of soft-bottom polychaetes (e.g., Prionospio multibranchiata and Ampharete acutifrons) and crustaceans (e.g., Corophium sextonae and Dynamene bidentatus), also known as detritivores. There were also major differences between the three habitats investigated. Consistent with its structural complexity, P. oceanica showed the highest species richness [E(S ₅₀)] and the most diversified macrofaunal assemblages, both in terms of taxonomic groups and taxa associated with different substrates. The two seagrasses, however, showed a similar species composition and differed from Leaf litter for the exclusive presence of hard-bottom species (e.g., the tunicate Phallusia fumigata) and seagrass-associated species (e.g., the polychaete Syllis garciai and the decapod Paguristes syrtensis). In contrast, Leaf litter showed the most differences between the habitats, and was characterized by the bivalves Abra alba and Cerastoderma glaucum, not found in seagrass beds, and by Loripes lacteus and Ruditapes decussatus. Leaf litter also had the highest content of organic matter (26.7% ± 1.4) and total organic carbon (10.3% ± 0.4). Our results confirmed the facilitative role of living seagrasses, in particular P. oceanica, as related to their structural complexity, for numerous species from different substrates (e.g., hard bottom species). This study also showed that leaf litter beds act as a particular environment where sediment instability, leaf breakdown, and organic matter enrichment and decomposition strongly influence animal distribution. Finally, our results highlighted the ecological and trophic importance of seagrass-derived detritus and the associated macroinvertebrate detritivores within seagrass-dominated systems.     

Marine calanoid resting eggs in Norway: Abundance and distribution of two copepod species in the sediment of an enclosed marine basin

Abundance and distribution of calanoid resting eggs in sediment were examined in 1988–1989 in an enclosed marine basin (Svartatjønn; western Norway). The basin undergoes rotenone treatment and draining every winter. Egg densities were highest in autumn, when up to 2 × 10⁶ eggs m^–2 were found, and lowest in summer. Egg densities were also highest in the deeper parts of the basin and in the upper 1 cm of sediment. Hatching was initiated in late January by pumping warmer deep water from the outside area into the basin. Evidence is given for resting-egg formation inEurytemora affinis Poppe, 1880 andAcartia clausi Gurney, 1931 (present phenotype reclassified asA. teclae by Bradford 1976: N. Z. Jl mar. Freshwat. Res. 10: 159–202). Seasonal differences in inhibition of hatching, together with the longevity and tolerance of these eggs, strongly suggest that they are diapause eggs.     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.^–1 day^–1) were calculated for this species from laboratory estimates of energy losses due to routine (R_R), active (R_A), and feeding (R_SDA) oxygen consumption rates (J ind.^–1 h^–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to R_R and R_A increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q₁₀=3.01) than between 10°C and 14°C (Q₁₀=2.85). Rates of R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.^–1 day^–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Respiration rate and cost of swimming for Antarctic krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in large groups in the laboratory

Constructing realistic energy budgets for Antarctic krill, Euphausia superba, is hampered by the lack of data on the metabolic costs associated with swimming. In this study respiration rates and pleopod beating rates were measured at six current speeds. Pleopod beating rates increased linearly with current speed, reaching a maximum of 6 beats s^–1 at 17 cm s^–1. There was a concomitant linear increase in respiration rate, from 1.8 mg O₂ g_D^–1 h^–1 at 3 cm s^–1 to 8.0 mg O₂ g_D^–1 h^–1 at 17 cm s^–1. The size of the group tested (50, 100 and 300 krill) did not have a significant effect on pleopod beating rates or oxygen consumption (ANCOVA, F=0.264; P>0.05). The cost of transport reached a maximum of 75 J g^–1 km^–1 at 5 cm s^–1, and then decreased with increasing current speed to 29 J g^–1 km^–1. When considered in light of energy budgets for E. superba, these data indicate that the cost of swimming could account for up to 73% of total daily metabolic expenditure during early summer.Communicated by G.F. Humphrey, Sydney     

Annual abundance and biomass of aloricate ciliates in tropical neritic waters off Kingston,Jamaica

The daily abundance of aloricate ciliates at Lime Cay, Jamaica, a shallow neritic site, ranged from 29 to 118 × 10⁶ m^–2 (0.97 to 3.93 × 10⁶ m^–3) between November 1985 and November 1986. Biomass was converted to kilojoules (1 kcal=4.1855 kJ) assuming 42% carbon, 20.15 kJ (g dry wt)^–1, and 20% cell shrinkage. Biomass ranged from 0.40 to 3.00 kJ m^–2 (13.3 to 100 J m^–3; 0.28 to 2.08µg C l^–1) with an annual mean of 1.11 kJ m^–2 (36.8 J m^–3; 0.764µg C l^–1). Nanociliates (<20µm equivalent spherical diameter, ESD) dominated abundance, but microciliates (> 20µm ESD) dominated biomass.Strombidium, Strobilidium, Tontonia andLaboea species were conspicuous taxa. Annual production estimates of the aloricate assemblage, based on literature growth rates, ranged from 404 kJ m^–2 yr^–1 (37 J m^–3 d^–1) to 1614 kJ m^–2 yr^–1 (147 J m^–3 d^–1). A compromise estimate of 689 kJ m^–2 yr^–1 (i.e., 63 J m^–3 d^–1) is comparable to other estimates from tropical and subtropical regions. A model of annual energy flow through 11 planktonic compartments suggests the total ciliate assemblage (aloricates and tintinnines) to be as productive as metazoan herbivores and metazoan carnivores.     

Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

Temporary cyst formation of<Emphasis Type="Italic"> Heterocapsa triquetra</Emphasis> (Dinophyceae) in natural populations

Heterocapsa triquetra (Ehrenberg) Stein is a phototrophic marine dinoflagellate with wide coastal distribution. It is known to be capable of mixotrophy and diel vertical migration. The species was particularly abundant in the Gulf of Finland (the Baltic Sea) during the summers of 1996 and 1998, leading to discolouration of water on the south-west coast of Finland. Large-scale (50 m³) coastal mesocosm experiments in the north-west Gulf of Finland (the Baltic Sea) in the summers of 1996 and 1998 with daily mineral nutrient additions provoked a biomass increase of phytoplankton dominated by H. triquetra. From the first days of the experiment temporary cysts of H. triquetra were found in the bottom sediment water of the mesocosms. Maximum temporary cyst production rates reached values up to 20×10⁶ cysts m^–2 day^–1, accounting for <1% of the depth-integrated motile population size. The environmental features favouring temporary cyst production remain uncertain; zooplankton grazing and nutrient stress are potential factors. Temporary cysts of H. triquetra were observed in a unialgal culture (f/2 medium) isolated in summer 1999 from Eel Pond (Woods Hole, Mass., USA).Communicated by M. Kühl, Helsingør