Impact of a burrowing deposit-feeder, Monoporeia affinis, on viable zooplankton resting eggs in the northern Baltic Sea

We investigated the effect of different densities of the burrowing deposit-feeding amphipod Monoporeia affinis on the potential for recruitment of zooplankton from benthic resting eggs. Hatching of resting eggs was induced in the laboratory on sliced and resuspended 1-cm depth-sections of sediment cores, collected at six stations in an archipelago area of the Gulf of Bothnia, Baltic Sea. The uppermost 5 cm of the sediment was studied. The most common species that hatched was Eurytemora affinis (Copepoda). Individuals from another copepod genus, Acartia, hatched in significant numbers only in the cores from two stations with low amphipod abundance. Cores from stations with high amphipod densities showed a deeper distribution of emerging E. affinis nauplii compared with stations with few amphipods; the oxidised sediment layer was also deeper at high M. affinis densities than at low. Total (0 to 5 cm strata pooled) number of hatched E. affinis nauplii was independent of amphipod density. This indicates that the effect of M. affinis on E. affinis eggs involves deeper burial due to bioturbation, rather than predation. Decreased benthic recruitment of zooplankton at localities with high M. affinis density is suggested, since more deeply positioned eggs are less likely to hatch. When hatching was induced in intact, non-sliced cores from one station, the number of E. affinis nauplii that hatched was on average 43% of the number that hatched in the upper centimetre of the sliced cores from the same station. This fraction (43%), if applied to the other stations, implied a potential for benthic recruitment of up to 80 000 ind m⁻² for E. affinis. Due to its high abundance, M. affinis is likely to greatly reduce benthic recruitment of zooplankton in this system. Received: 17 September 1999 / Accepted: 23 February 2000     

Deposit-feeding amphipods (Monoporeia affinis) reduce the recruitment of copepod nauplii from benthic resting eggs in the northern Baltic Sea

We experimentally investigated the effect of different densities of the burrowing, deposit-feeding amphipod Monoporeia affinis on the recruitment of zooplankton from benthic resting eggs. Intact sediment cores with in situ density and species composition of zooplankton resting eggs and benthic fauna were collected in the northern Bothnian Sea, part of the Baltic Sea. We removed as many M. affinis as possible from the cores and then added different numbers of M. affinis to the cores to generate a range of densities. The cores were exposed to different densities of M. affinis for either 3 or 40 days, after which the hatched zooplankton was registered. One subset of the cores were initially incubated under low temperature (2–3 °C, to prevent hatching) for 37 days (the resting phase), to allow for effects of M. affinis on unhatched resting eggs. These cores were then incubated under higher temperature (13 °C) for 3 days (the hatching phase), to induce hatching and allow for effects on hatching or hatched specimens. In a second subset of cores with the same time and temperature schedule, the M. affinis density was experimentally reduced at the start of the hatching phase, to evaluate the effect of M. affinis during the hatching phase. To a third subset of cores, we immediately initiated the hatching phase, without an experimental resting phase, to evaluate the effects induced during the resting phase. The most common zooplankton species that hatched was Eurytemora affinis (Copepoda), followed by Bosmina longispina maritima (Cladocera). In all cores that were subjected to a resting phase, the numbers of hatched E. affinis were log-linearly negatively related to density of M. affinis. An increase of M. affinis density from 1,000 to 5,000 individuals m⁻², normal field densities, reduced the hatching by 60–70%. The negative impact was mainly exerted during the hatching phase, suggesting predation on, burial of or physical injury of hatching nauplii or eggs in a late development stage as likely mechanisms. Also, the number of B. longispina maritima that hatched was reduced by M. affinis during the hatching phase, but no clear relation to density of M. affinis could be identified. The results show that M. affinis can reduce recruitment to zooplankton from benthic resting eggs. Such impact by the benthos on resting stages of zooplankton is therefore a potentially significant link between the benthic and pelagic systems. Received: 10 August 2000 / Accepted: 13 November 2000     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Survival, growth and reproduction of the salt-marsh amphipod Uhlorchestia spartinophila reared on natural diets of senescent and dead Spartina alterniflora leaves

The talitrid amphipod Uhlorchestia spartinophila lives in close association with standing-dead leaves of the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the Atlantic coast of North America. This study probed the strength of the trophic link between the amphipod population and the decomposition process in this detrital-based ecosystem. We measured survival, growth and reproductive output in groups of amphipods reared for 6 wk on five diets derived from sheath and blade portions of S. alterniflora leaves just prior to (senescent) and during (dead) decomposition. In unfed treatments, the daily specific mortality rate was 0.391 and starved amphipods survived no longer than 11 d. Among the fed treatments, a diet of senescent sheaths resulted in the lowest survival (20%) and yielded no offspring. Groups fed senescent blades, dead sheaths, dead blades and unwashed dead sheaths had survival rates of 56 to 84% and produced 5.0 to 12.5 offspring replicate⁻¹. Sex ratio usually favored females, but approached unity in treatments with high overall survival, suggesting that quality of available food resources may influence sex ratio in this species. Mean specific growth rates (mm mm⁻¹ d⁻¹) ranged from 0.013 to 0.016, and matched previous estimates of growth from field populations. Overall ecological performance (survival + growth + reproduction) was similar for all food treatments, except senescent sheaths, which yielded a final mean (±SD) dry biomass (0.4 ± 0.42 mg replicate⁻¹) of amphipods significantly lower than that of other diets (1.7 ± 0.81 to 2.6 ± 0.69 mg replicate⁻¹). Natural diets derived from decomposing cordgrass leaves can fulfill the nutritional requirements of U. spartinophila populations, but variation in initial amounts of living fungal biomass among the five experimental diets only partially explained the responses of amphipods in our experiment. Structural characteristics and variation in rates of fungal occupation within different portions of cordgrass leaves may affect the amphipod's ability to access plant production made available by decomposers. Received: 12 December 1996 / Accepted: 18 December 1996     

Life under Arctic sea ice – reproduction strategies of two sympagic (ice-associated) amphipod species, Gammarus wilkitzkii and Apherusa glacialis

The reproductive strategies of two gammaridean amphipod species, Gammarus wilkitzkii and Apherusa glacialis, that permanently inhabit the Arctic sea ice were investigated. G. wilkitzkii reaches sexual maturity at an age of 2 years and produces 128 ± 54 eggs fem.⁻¹ yr⁻¹. Mating takes place during fall and winter, and the development of the large eggs (0.60 to 0.80 mm diam.) lasts 6 to 7 months. The sex ratio of G. wilkitzkii was dominated by males in a proportion of 1.5:1. In vivo studies showed that juveniles are released in batches from the brood pouches of the females during April and May. A. glacialis reaches sexual maturity at the age of 1 year and produces 555 ± 151 eggs fem.⁻¹ yr⁻¹. The eggs are between 0.18 and 0.23 mm in diameter, and are the smallest known for gammaridean amphipods. Eggs are kept in packages of two to eight in the brood pouches of females. The sex ratio of A. glacialis was dominated by females in a proportion of 3:1. The high fecundity of both amphipod species, the release of juveniles in batches over a period of time, a high proportion of females (A. glacialis), and an elongated life-span with multiple spawnings (G. wilkitzkii) are discussed as possible adaptations to the specific and highly variable conditions under Arctic sea ice. Received: 29 December 1999 / Accepted: 8 March 2000     

Population and reproductive biology of two sibling amphipod species from ascidians and sponges

 The population dynamics and reproductive biology of an ascidian- and a sponge-dwelling amphipod were examined. The two undescribed amphipod species, Leucothoe“ascidicola” and L.“spongicola”, are closely related to each other, and occur in ascidians and sponges, respectively, along the Florida Atlantic coast. L. “ascidicola” was abundant in solitary ascidians during fall 1997, disappeared during spring/early summer, and became abundant again in September 1998. During the time when L. “ascidicola” were absent from their hosts, a copepod became a frequent inhabitant of the ascidians but disappeared again when L.“ascidicola” returned to the ascidians in September 1998. The numbers of L.“spongicola” in sponges increased substantially during spring, when high reproductive activity was observed. Following this reproductive peak, both adult and juvenile amphipods apparently left the sponges, and during the summer amphipod numbers in the sponges were very low. Another small amphipod species, which often co-occurred with L.“spongicola”, showed less seasonal variation and was found in sponges throughout the whole study period. The percentage of ovigerous females per host unit was usually lower in the ascidian-dwelling than in the sponge-dwelling amphipods. In solitary ascidians, L.“ascidicola” amphipods usually occurred in groups of several adults, yet there never was more than one ovigerous female per ascidian. In contrast, several ovigerous L.“spongicola” females were found to cohabit in the same spongocoel. This suggests that intrasexual aggression may be stronger among reproductive amphipod females in the ascidians than in sponges. The size distributions of juvenile cohorts indicate that juvenile L.“ascidicola” remain for relatively long time periods in the parental ascidian, where they may reach sexual maturity. In contrast, in L.“spongicola”, only cohorts of very small juveniles could be identified, indicating that juveniles disperse shortly after emerging from the female's brood pouch. It is concluded that extended parental care is of very short duration or does not occur in the sponge-dwelling amphipod L. “spongicola”, possibly because fast-growing sponges with a highly branched spongocoel system do not allow long-lasting coexistence of parent-offspring groups. In contrast, the discrete character of the solitary ascidians may enhance the potential for exclusion of other species, resource monopolization by reproductive females, and furthermore for long-lasting extended parental care in the ascidian-dwelling amphipod. Groups of single parents together with cohorts of large juveniles are reported in the literature for amphipods and isopods from brachiopods, bivalves and ascidians, suggesting that these discrete biotic microhabitats may favor the evolution of extended parental care in peracarid crustaceans. Received: 30 July 1999 / Accepted: 8 May 2000     

Seasonal variation of membrane fluidity of the naturally acclimatized Baltic Sea amphipods Gammarus spp. and Monoporeia affinis

The membrane fluidity of the Baltic Sea crustaceans Gammarus spp. and Monoporeia affinis was studied in different seasons. Gammarus spp. were collected at a location with stable salinity and with temperature fluctuations from about 0 to 20 °C, and M. affinis at a deep location with stable salinity and a constant low temperature of about 1.5 to 4.5 °C. The membrane fluidity was measured from preparations enriched with plasma and mitochondrial membranes employing a fluorescence polarization technique using 1,6-diphenyl-1,3,5-hexatriene (DPH) as a probe. The measurements were carried out without any preceding acclimation of the animals. In Gammarus spp., the membrane fluidity increased in the order: summer animals < autumn animals < spring animals. The differences between seasons were significant, with pronounced homeoviscous adaptation, and correlated well with the water temperature. It is suggested that in Gammarus spp., temperature is the determining factor for membrane fluidity. In M. affinis, the membranes of summer animals were the most fluid, and there was a significant difference only between summer and spring animals. In M. affinis the difference was not connected to the water temperature and no homeoviscous adaptation was noticed. The deep-water M. affinis experience a long period of fresh food deficiency, which probably affects the membrane fluidity. Although there are seasonal differences in fluidities between Gammarus spp. and M. affinis, it is interesting to note that all data obtained from M. affinis settled between the polarization lines of spring and summer Gammarus spp., regardless of the prevailing temperatures. Received: 30 June 1999 / Accepted: 7 June 2000     

Effects of DDT sediment-contamination on macrofaunal community structure and composition in San Francisco Bay

The objectives of this study were to determine the effects of sediment contamination on the benthic macrofauna and to predict macrofaunal changes following remediation at a Superfund (uncontrolled hazardous waste) site in San Francisco Bay, California, USA. DDT and its metabolites (ΣDDT) were the contaminants of concern. With few small-scale exceptions, all (>100) other sediment contaminants ever measured at the site were present at background or non-toxic levels. In hierarchical regressions [Y=f(X ₁, X ₂, X ₃), where X ₁=sediment %silt + clay, X ₂=sediment total organic carbon (OC), and X ₃=log ₁₀ (ΣDDT μg g⁻¹ OC)] with data from samples collected at the study site, log₁₀(ΣDDT μg g⁻¹ OC) explained a highly significant amount of the variance in the infaunal index (II ) and log₁₀(number of Amphipoda excluding Grandidierella japonica + 1) after statistically controlling for the potential effects of sediment %silt + clay and OC. The ratios of change of II and log₁₀(number of Amphipoda excluding G. japonica + 1) with respect to log₁₀(ΣDDT μg g⁻¹ OC) were about −9:1 and −0.4:1, respectively. Most of the 92 species collected were present at low to moderate densities over the entire range of ΣDDT sediment concentrations. The bivalve Theora lubrica, tubificids, most polychaetes, a tanaid (Zeuxo normani), and an amphipod (G. japonica), were common, while four other amphipods (Ampelisca abdita, Corophium heteroceratum, Photis brevipes, Dulichia rhabdoplastis), a phoronid (Phoronis cf. pallida), a bivalve (Cryptomya californica), and a cumacean (Eudorella pacifica), were rare or absent from sites with high ΣDDT sediment-concentrations. Received: 1 August 1997 / Accepted: 13 August 1997     

Field distribution and sulphide tolerance of Capitella capitata (Annelida: Polychaeta) around shallow water hydrothermal vents off Milos (Aegean Sea). A new sibling species?

The cosmopolitan polychaete Capitella capitata, known as a complex of opportunistic sibling species, usually dominates the macrobenthos of polluted or unpredictable environments. A population of C. capitata, termed Capitella sp. M, was found in a shallow water hydrothermal vent area south of Milos (Greece). Here, this population occurs close to vent outlets (termed the “transition zone”), an environment with steep gradients of temperature, salinity and pH and increased sulphide concentrations of up to 710 μM. The field distribution of C. capitata in relation to sulphide concentrations around the vent outlets was investigated and sulphide tolerance experiments were conducted on laboratory-cultured worms to elucidate possible adaptations of Capitella sp. M to these extreme environmental conditions. In order to investigate whether the population from the Milos hydrothermal vent area can be considered a distinct sibling species within the C. capitata complex, crossbreeding experiments and analysis of general protein patterns were conducted with Capitella sp. M and three other C. capitata populations of different ecological ranges. Capitella sp. M showed high resistance (median survival time: 107 ± 38 h) to anoxia plus high sulphide concentrations of 740 μM. It seems that the ability to survive high-sulphide conditions in combination with reduced interspecific competition enables the polychaete to maintain a continuous population in this rigorous habitat. From the extremely high tolerance to anoxia and sulphide, shown in both the crossbreeding experiments and the analysis of total proteins, it can be concluded that Capitella sp. M from the Milos hydrothermal vent area represents a separate sibling species within the C. capitata complex. Received: 3 March 1997 / Accepted: 12 September 1997     

Effects of cadmium on Pontoporeia affinis (Crustacea: Amphipoda) in laboratory soft-bottom microcosms

Laboratory soft bottom microcosms, containing sediment with natural populations of meio-and microfauna and the amphipod Pontoporeia affinis Lindström, were used to investigate chronic effects of cadmium on a typical Baltic Sea benthic community. Temperature, salinity, organic content of the sediment and the life cycles of the amphipods in laboratory control microcosms were in good agreement with conditions in the Baltic Sea. In Experiment 1, P. affinis was exposed to four assayed cadmium concentrations, 0 (control), 6.3, 41 and 127 g Cd·1^-1, for 265 d. At the lowest cadmium concentration (6.3 g Cd·1^-1), the sediment contained about 700 times, and P. affinis about 1 700 times, the water concentration at the end of the experiments. In microcosms without P. affinis, the sediments accumulated less cadmium. Effects on P. affinis embryogenesis were detected after 105 d of exposure in all tested cadmium concentrations. Mortality of juveniles was significantly increased after 265 d of exposure in mean assayed concentrations of 41 and 127 g Cd·1^-1. In Experiment 2, P. affinis was exposed to cadmium in assayed concentrations, 0, 6.5 and 46 g Cd·1^-1 for 460 d. At the lowest water concentration (6.5 g Cd·1^-1), the prolonged exposure did not increase the sediment concentration, which indicates a steady state, while concentrations in P. affinis reached about 3 500 times the lowest water concentration after 460 d. Juvenile P. affinis matured sexually and reproduced normally in all microcosms, but their offspring died in all microcosms with added Cd. This occurred at water concentrations not uncommon in polluted marine areas. The effects are serious in comparison to those found in conventional short-term laboratory experiments with the same concentrations and demonstrate the importance of long-term experiments in ecotoxicology.     

Nitrogen- versus phosphorus-limited growth and sources of nutrients for coral reef macroalgae

Recent investigations of nutrient-limited productivity in coral reef macroalgae have led to the conclusion that phosphorus, rather than nitrogen, is the primary limiting nutrient. In this study, comparison of the dissolved inorganic nitrogen:phosphorus ratio in the water column of Kaneohe Bay, Hawaii, with tissue nitrogen:phosphorus ratios in macroalgae from Kaneohe Bay suggested that nitrogen, rather than phosphorus, generally limits productivity in this system. Results of nutrient-enrichment experiments in a flow-through culture system indicated that inorganic nitrogen limited the growth rates of 8 out of 9 macroalgae species tested. In 6 of the species tested, specific growth rates of thalli cultured in unenriched seawater from the Kaneohe Bay water column were zero or negative after 12 d. These results suggest that, in order to persist in low-nutrient coral reef systems, some macroalgae require high rates of nutrient advection or access to benthic nutrient sources in addition to nutrients in the overlying water column. Nutrient concentrations in water samples collected from the microenvironments inhabited or created by macroalgae were compared to nutrient concentrations in the overlying water column. On protected reef flats, inorganic nitrogen concentrations within dense mats of Gracilaria salicornia and Kappaphycus alvarezii, and inorganic nitrogen and phosphate concentrations in sediment porewater near the rhizophytic algae Caulerpa racemosa and C. sertularioides were significantly higher than in the water column. The sediments associated with these mat-forming and rhizophytic species appear to function as localized nutrient sources, making sustained growth possible despite the oligotrophic water column. In wave-exposed habitats such as the Kaneohe Bay Barrier Reef flat, water motion is higher than at protected sites, sediment nutrient concentrations are low, and zones of high nutrient concentrations do not develop near or beneath macroalgae, including dense Sargassum echinocarpum canopies. Under these conditions, macroalgae evidently depend on rapid advection of low-nutrient water from the water column, rather than benthic nutrient sources, to sustain growth. Received: 1 December 1997 / Accepted: 9 July 1998     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Production and biomass accumulation of periphytic diatoms growing on glass slides during a 1-year cycle in a subtropical estuarine environment (Bay of Paranaguá, southern Brazil)

Production rates, chlorophyll concentrations and general composition of periphytic diatom communities growing on glass slides were studied in relation to environmental parameters during one seasonal cycle in the Bay of Paranaguá, southern Brazil. Slides were routinely submersed at 1, 2 and 3 m depth and recovered weekly for microscopic examinations, analyses of chlorophyll, cell counts and in situ photosynthetic incubations using the Winkler titration method. Water samples were also collected at surface and bottom layers for determinations of temperature, salinity, nutrients and chlorophyll in the water. The periphytic community was mainly formed by epipelic and epipsammic species, dominated by Navicula phyllepta, Cylindrotheca closterium, Navicula spp. and Amphora sp. Weekly chlorophyll a and cell accumulations on slides varied from <1–32 mg m⁻² and up to 31 × 10⁸cells m⁻², respectively. Photosynthetic rates varied from <1 to 35 mg oxygen mg chlorophyll a ⁻¹ h⁻¹, with higher values in summer. Daily production varied from 5 to 3,600 mg oxygen m⁻² day⁻¹ (<0.01–1.4 g carbon m⁻² day⁻¹). Multiple regression analysis revealed that vertical differences in light conditions and grazing pressure jointly affected the influence of temperature on the seasonal patterns of cell densities and chlorophyll concentrations according to depth. Received: 27 April 2000 / Accepted: 16 August 2000     

Behavioural response to gradually declining oxygen concentration by Baltic Sea macrobenthic crustaceans

Behavioural responses to decreasing oxygen concentrations were studied in the amphipods Monoporeia affinis (Lindström) and Pontoporeia femorata Krøyer and the isopod Saduria entomon (L.), all benthic macrofauna species from the Baltic Sea. Infrared time-lapse video recording was used. Seven oxygen concentrations were studied, starting near saturation (about 90%), followed by about 65, 49, 33, 16, 8 and 5% oxygen saturation, with each oxygen level maintained for 24?h. M. affinis was more active than P. femorata. The duration of swimming activity of M. affinis was shorter at ≤33% oxygen saturation compared with at the two highest levels. The duration of swimming by P. femorata was longest at 90, 65 and 5% oxygen saturation. For both amphipods the frequency of swimming activity was highest at 5% oxygen saturation. Above 7% oxygen saturation, S. entomon spent most of its time in the sediment, whereas at levels ≤7% it was most often found on the sediment surface. Both amphipods were predominantly night active (nocturnal), whereas no diurnal variation in behaviour was detected for S. entomon. It is suggested that the amphipods minimise the costs associated with obtaining oxygen as well as their risk of predation by moving little on or above the sediment until a lower critical oxygen level is reached. Below this level, they show an oxygen-seeking behaviour. S. entomon also hid in the sediment for as long as possible, suddenly coming to the surface once the oxygen concentration became intolerably low. Its behaviour is also indicative of a trade-off between predation risk and obtaining sufficient oxygen. It is suggested that the change in behaviour shown at about 33% oxygen saturation could lead to a long-term alteration in community structure in the deeper parts of the northern Baltic proper, whereas the change at about 8% oxygen saturation reflects a more acute, short-term response.     

Impact of the polychaete Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon fluoranthene

Polychaetes belonging to the genus Capitella are often present in high numbers in organic-rich sediments polluted with, e.g., oil components, and Capitella spp. may have a great impact on the biogeochemistry of these sediments. We examined the influence of Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon, fluoranthene. Capitella sp. I were added to microcosms (10 000 ind m⁻²) and the impact of a pulse-sedimentation of fluoranthene-contaminated sediment (3 mm layer) was studied for a period of 12 d after sedimentation. The sediment oxygen uptake and total sediment metabolism (TCO₂ production) increased in cores with worms (71 to 131%), whereas the anaerobic activity, measured as sulfate reduction rate 12 d after sedimentation, was lower compared to cores without worms. The effect of fluoranthene on sulfate reduction was most pronounced in the presence of worms, with a 34% reduction versus 16% in cores without worms. The reduced sulfur pools in cores with worms were smaller than in cores without worms, suggesting that the reduced anaerobic activity was caused by increased oxidation of the sediment, which may favor O₂ and other electron-acceptors (e.g. NO₃ ⁻, Fe³⁺, Mn⁴⁺) in organic matter decomposition. The sediment oxygen uptake and TCO₂ production did not show significant changes due to fluoranthene treatment, indicating that these parameters were either less sensitive to fluoranthene stress or recovered more rapidly (i.e. within 48 h) than sulfate reduction rates. Bioturbation by Capitella sp. I altered the depth profile of fluoranthene such that fluoranthene was found in deeper sediment layers (down to 2 cm) where diffusional loss and microbial breakdown probably are reduced relative to surface layers. In cores without worms, fluoranthene was found down to 1 cm, with 75% remaining in the upper 5 mm. Received: 5 December 1996 / Accepted: 11 February 1997     

Physiological and behavioral aspects of Calanus euxinus females (Copepoda: Calanoida) during vertical migration across temperature and oxygen gradients

 In the Black Sea, during summer stratification, Calanus euxinus (Hulsemann) undertakes diel vertical migrations with an amplitude of about 117 m from oxygenated, warm (18 °C) surface layers to hypoxic (∼0.8 mg O₂ l⁻¹) zones with lower temperature (7.9 °C). When such changes in temperature and oxygen concentration are reproduced in the laboratory, total metabolism, basal metabolism and scope of activity of copepods decrease 7.2, 7.8 and 6.7 times, respectively, while the frequency of locomotory acts and mechanical power decline 3.4- and 9.5-fold, respectively. These changes allowed the copepods to conserve a significant portion of food consumed near the surface for transformation to lipid reserves. Diel respiratory oxygen consumption of migrating individuals, calculated so as to include actual duration of residence in layers with different temperature and oxygen concentrations, is estimated at 17.87 μg O₂ ind⁻¹. The net energy cost of vertical migration made up only 11.6% of the total. Copepods expend 78.6% of diel energy losses during approximately 10 h in the surface layers, while about 5.4% is required during about 9 h at depth. Hypoxia is shown to have a significant metabolic advantage during diel vertical migrations of C. euxinus in the Black Sea. Received: 1 October 1999 / Accepted: 11 July 2000     

Dimethylsulfoniopropionate and dimethylsulfide in the marine flatworm Convoluta roscoffensis and its algal symbiont

 A distinct smell of dimethylsulfide (DMS) was noted at the edge of the intertidal mudflat of Marennes-Oléron Bay, at the French Atlantic coast, where dense populations of the marine flatworm Convoluta roscoffensis Graff (Platyhelminthes: Turbellaria) were present. DMS is the cleavage product of dimethylsulfoniopropionate (DMSP). DMSP was shown to be present in high amounts in sediment containing the flatworm as well as in axenic cultures of the symbiotic alga Tetraselmis sp. that was isolated from the flatworm. In untreated sediment samples containing C. roscoffensis the concentration of DMS was as high as ∼55 μmol l⁻¹ sediment, and in samples that were fixed with glutaraldehyde the concentration of DMS was even three orders of magnitude higher (∼66 mmol l⁻¹ sediment). This rapid cleavage of DMSP to DMS in fixed samples was unexpected. Pure DMSP was stable in glutaraldehyde, and it was therefore concluded that a DMSP-lyase was responsible for cleavage in the field samples. The isolated symbiotic alga, Tetraselmis sp., did not show DMSP-lyase activity, indicating that DMSP-lyase may have been present in the flatworm, although the role of bacteria could not be excluded. The Chl a-specific DMSP content of C. roscoffensis (∼200 mmol g⁻¹) was much higher than that of Tetraselmis sp. (∼30 mmol g⁻¹). Possibly, DMSP was not only present in the symbiotic alga, but was also incorporated in the body tissue of the flatworm. It remains unclear what the function of DMSP is in C. roscoffensis. In Tetraselmis sp., but not in C. roscoffensis, DMSP increased with increasing salinity. It was concluded that salinity probably does not play an important role in the dynamics of DMSP and DMS in sediment containing C. roscoffensis. Received: 21 January 2000 / Accepted: 29 August 2000     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Sediment processing in Capitella spp. (Polychaeta: Capitellidae): strain-specific differences and effects of the organic toxicant fluoranthene

The effects of the widespread polycyclic aromatic hydrocarbon (PAH) fluoranthene on sediment-processing rates in the infaunal polychaete Capitella spp. were investigated by comparing five populations of this deposit-feeding species complex: Capitella sp. I from New York, USA; Capitella sp. M from Milos, Greece; Capitella sp. S from Sylt, Germany; and two unidentified Capitella populations from salmon farm sediments –Capitella population K from Kilmelford, Scotland and Capitella population C from Cranford, Ireland. Replicate worms from each strain were exposed to 0, 10, and 95 μg (g dry wt sediment)⁻¹ fluoranthene (=μg/g fluoranthene) for a period of 16 days. Initial and final wet and dry weights (mg) of worms and worm-specific growth rates (WSGRs) were calculated. Sediment processing was measured as the sum of the total dry weight of pellets produced during the experiment, and we estimated size-specific processing rates (SSPRs) as a measure of sediment processed per mg worm dry weight per day. The five populations of Capitella spp. differed significantly in body size, WSGR, and sediment-processing rates. Capitella sp. I grew faster than all of the other populations. Capitella population C and Capitella population K from fish farm sediments, with the largest body lengths (up to 52.0 ± 27.2 mm), had the highest processing rates, whereas the small Capitella sp. S (up to 17.1 ± 5.6 mm) had the lowest. There were also significant differences in SSPR among populations with Capitella sp. I having a higher SSPR (about 12 × body wt/day) than Capitella population C (about 5 × body wt/day) and Capitella population K (3 × body wt/day). The fluoranthene concentrations used in the present study, while representing moderately to highly contaminated conditions, had only marginal effects on sediment-processing and growth rates of all of the Capitella populations examined. Processing of contaminated sediment by Capitella spp. may be important in the remediation of PAH-contaminated sediment. Received: 16 January 2000 / Accepted: 28 August 2000     

Development, maturation, brood size and generation length of the mesopelagic amphipod Cyphocaris challengeri (Gammaridea: Lysianassidae) off southwest Hokkaido, Japan

 Using the number of segments of pleopod rami as a marker of instar number, the population structure (instar composition) of the mesopelagic gammarid amphipod Cyphocaris challengeri was investigated by monthly samplings from May 1997 to April 1999 at a station off southwest Hokkaido, Japan. Laboratory-rearing experiments were also conducted to establish the relationship between the number of segments of pleopod rami and instar number, and to estimate the growth pattern of this gammarid based on the intermolt period and molt-increment data. Stratified sampling in the field (0 to 200 and 200 to 400 m depth strata) showed this species occurred mainly at 200 to 400 m depth during the day. Instar analysis indicated that C. challengeri has 12 instars in females and 11 instars in males. Based on observations of secondary sexual characters, Instars 1 to 6 were designated juveniles (Instars 1 to 3 occurred in the marsupia of gravid females); in males, 7 to 9 were immature and 10 and 11 were mature, while in females 7 and 8 were immature and 9 to 12 were mature. Off southwest Hokkaido, Instar 4 (just released from a female's marsupium) was found throughout the year, with a peak abundance occurring in April to July of each year. A sequential development of Instar 4 to 9 (youngest adult instar) through the year was observed. Generation length (i.e. the time required to grow from Instar 4 to 10) was estimated from a laboratory-obtained growth curve to be 216 to 584 d at the in situ temperature range (2 to 5 °C), which is consistent with observations on field populations. Specimens older than Instar 9 were rare in the field and could not be used in laboratory-rearing experiments, so longevity could not be estimated. Eggs were oval and measured 0.6 mm (large diameter). Brood size ranged from 20 to 65. Comparing the present results with those of epipelagic hyperiid amphipods, the nearly identical growth rates together with the production of fewer but larger eggs seen in C. challengeri appear to reflect to the typical life mode of deep-living pelagic crustaceans. Received: 14 February 2000 / Accepted: 6 July 2000