Reproduction ofCalanus glacialis under ice in spring in southeastern Hudson Bay,Canada

In situ egg production rates ofCalanus glacialis Jaschnov females caught under the ice cover of southeastern Hudson Bay, Canada, between early April and mid-June 1986 were measured by direct observations and by an egg-ratio method in order to investigate the role of ice microalgae and under-ice phytoplankton production in copepod reproduction. Egg production commenced, although at low rates, in early May, about 3 wk after the onset of measurable concentrations of ice microalgae at the ice-water interface. Egg production increased to about 12 eggs female^–1 d^–1 in early June, after the ice cover began to melt. The time taken by immature adult females to reach maturity in the field was estimated to be ca. 3 wk (between late April and mid-May) by visual evaluation of gonad maturity in preserved females. In the laboratory, females caught on 10 April and maintained in saturating concentrations of the diatomThalassiosira weissflogii took only a few days to produce eggs at high rates, indicatingC. glacialis was food-limited during the ice algal bloom at the interface. We suggest that ice algal grazing changes the timing, relative to high food concentrations in the water column, of reproduction inC. glacialis by promoting oogenesis and oocyte maturation.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)Please send all reprint requests to Dr. Runge at his current address: Institut Maurice-Lamontagne, Ministère des Pêches et Océans, C. P. 1000, Mont-Joli, Québec G5H 3Z4, Canada     

Early reproduction and development of dominant calanoid copepods in the sea ice zone of the Barents Sea—need for a change of paradigms?

Reproduction and growth of the dominant copepods Calanus finmarchicus, C. glacialis, C. hyperboreus and Pseudocalanus minutus were studied on transects across the sea ice zone of the northern Barents Sea in May and June 1997. C. glacialis and C. finmarchicus were numerically dominant and also the largest component of the biomass. C. hyperboreus was rather rare. Moderate levels of phytoplankton and eventually high concentrations of ice algae supported maximum egg production rates of 53.6 and 48.5 eggs female^–1 day^–1 of C. glacialis in May and June, respectively. Results of incubation experiments were supported by a tremendous abundance of C. glacialis eggs in the water column ranging from 7×10³ to 4.4×10⁴ m^–2 in May and from 9.8×10³ to a maximum of 9.7×10⁴ m^–2 in June. In contrast, C. finmarchicus spawned only in the vicinity of the ice edge, at a maximum rate of 30 eggs female^–1 day^–1. Egg sacs of P. minutus were often observed in the preserved samples, but contained only few eggs, which may be due to loss during sampling. The presence of considerable concentrations of young stages in May and June indicated successful recruitment of C. glacialis and P. minutus. Back calculation using published stage duration estimates indicates March/April as the begin of the reproductive and growth period for these species under the first-year ice of the Barents Sea. Hence, secondary production in the study area starts at the same time as in open water regions and polynyas in the northern North Atlantic. Although the role of ice algae in the nutrition of copepods was not clarified here, the significant relationship between phytoplankton chlorophyll and egg production of C. glacialis suggests that high reproductive activity has already been achieved at moderate food concentrations.Communicated by O. Kinne, Oldendorf/Luhe     

A new source of dimethylsulfide (DMS) for the arctic atmosphere: ice diatoms

We report the first evidence that pennate diatoms growing within the bottom layer of first-year ice in the Arctic produce significant amounts of particulate dimethylsulfoniopropionate (DMSP_p) and dissolved DMSP+DMS. In 1992 in Resolute Passage, a tributary of Barrow Strait, DMSP_p concentrations within the bottom layer of ice reached 1055 mg S m^-3 at the end of the vernal bloom, a value one order of magnitude higher than the maximum value reported in antarctic ice. Bottom-ice concentrations in DMSP_p and DMSP_d+DMS were significantly correlated with the abundance of the dominant pennate diatom Nitzschia frigida. Intracellular concentration in DMSP of ice algae was very low (0.001 pg cell^-1) at the end of April when algae were light-limited and reached 1.17 pg cell^-1 in mid-May following an increase in light and algal growth. We calculate that the rapid release of the dissolved DMSP+DMS from the ice into surface waters following the ice break-up will generate a sea-to-air DMS flux of 0.7 mg S m^-2 d^-1, a pulse ten times higher than the mean arctic summer flux. We estimate that this 1-d pulse represents up to 5% of the annual DMS emission in the Arctic.     

Diel variations in the nutritional physiology of Calanus glacialis from Lat. 78°N in the summer

Calanus glacialis (Jaschnov) from Isfjord, western Spitzbergen, showed no tendency to pronounced diurnal vertical migration or cyclic variations in feeding activity. The samples, taken regularly over a 24-h period and at three different depths in the beginning of August, did however show considerable individual variations in feeding activity. These variations, reflected in the results of gut analyses, were not related to variations in the concentrations of particulate protein, carbohydrate or algal pigments in the water. Analyses of the activities of two digestive enzymes and the electron-transport system (ETS) indicated that those few individuals always present below the photic zone were there temporarily, and showed no signs of slowing down their metabolism. The results suggest that the population of C. glacialis, during the Arctic summer, feeds continuously in the upper water layer, but that intermittent, non-synchronous individual feeding rhythms are the rule.     

In situ feeding rhythms of herbivorous copepods,and the effect of starvation

In situ diel variations in gut pigment contents of neritic (Acartia omorii andPseudocalanus minutus) and oceanic copepods (Calanus plumchrus andC. cristatus) were analyzed.A. omorii andP. minutus were sampled in Onagawa Bay on the east coast of Japan in May and August 1987, andC. plumchrus andC. cristatus were sampled in the Bering Sea in June 1986. Gut pigments were generally high at night, and bimodal feeding rhythms were observed in all species. The first peak of gut pigments occurred between sunset and midnight and was followed by a midnight decrease in gut pigment levels, resulting in eventual evacuation of the gut. The second peak was observed a few hours after sunrise. Incubation experiments indicated that ingestion rates of starved copepods were higher than those of acclimated copepods. This phenomenon was most notable at high food concentrations. Gut pigments of starved copepods rapidly increased after exposure to high concentrations of culturedThalassiosira decipiens. These findings suggest that in situ feeding behavior of herbivorous copepods includes periods of cessation or reduction in feeding during the night, and consequently, feeding activity is periodically enhanced with starvation. Starvation enhanced feeding behavior is most obvious in the large oceanic species,C. plumchrus andC. cristatus and is not distinct in small coastal species such asA. omorii.     

Vertical fine structure of the biomass and composition of algal communities in Arctic pack ice

The biomass and composition of algal communities in sea ice were studied during two summer expeditions to the central Arctic Ocean and the Greenland Sea. Based on algal pigment determination and cell counts, high biomass accumulations were found at the surface, in the interior and in the bottom layer of the ice floes. Pennate diatoms dominated in the bottom layer, while phototrophic flagellates and cysts of unknown origin were the most abundant taxa in the upper parts. The lowermost 20 to 40 cm contained between 4 and 62% of the entire algal biomass. Consequently, ice biological studies, which deal only with the bottom few centimetres of the ice floes, will underestimate algal biomass and production by factors of up to 25. Differences between the results of this study and published data from coastal locations point towards different biological regimes in Arctic sea ice. The algal biomass in coastal ice is about two orders of magnitude higher and composed mainly of diatoms, probably supported by nutrient influx from the water column. In the pack ice of the central Arctic, nutrient supply is probably reduced, and flagellates contribute substantially to total algal biomass. However, methodological problems might partially be responsible for the observed differences. Received: 12 June 1998 / Accepted: 11 December 1998     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Growth and collapse of a spring phytoplankton bloom in the Firth of Clyde,Scotland

A range of physical and chemical parameters were measured through the water column at two previously unstudied shallow-water sites in the Firth of Clyde, Scotland, in the spring of 1991. Sediment traps were also deployed at one of the sites. The results of this survey are presented and discussed. A bloom of Skeletonema costatum was observed at one site through March and into April, when it collapsed completely, the phytoplanktonic cells sedimenting out of the water column. No significant bloom or sedimentation was observed at the second study site.     

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.     

Impact of Land-Based Sources On Marine Environment: Phytoplankton Blooms Within an Annual Cycle

The discharge water into the western region of Alexandria (Egypt), creates suitable conditions for the acceleration of the phytoplankton growth; daily replenishment of nutrients and development of density stratified water column. Water transparency and ammonia concentrations seem to be affecting the variations of the numerical standing crop. Eight phytoplankton bloom pulses of different causative species were observed. Asterionella glacialis represents a newly recorded red tide species in the neritic waters of Alexandria.     

Algal blooms decrease care but increase egg survival in a fish with paternal care

Human-induced eutrophication, resulting in algal blooms and increased water turbidity, is an alarming problem in aquatic systems. Here, we experimentally tested the impact of algal turbidity on parental care, egg fanning, and time in the nest, in the sand goby, Pomatoschistus minutus, a fish with uniparental male care. We allowed males to care for their eggs in either clear water or water made turbid by planktonic algae. In the early brood cycle, males fanned their eggs less in turbid than in clear water, but this difference disappeared later. Despite decreased care, egg survival was higher in turbid conditions, indicating that early fanning may partly be redundant for egg survival and perhaps used more as courtship. Males also spent more time out of their nest in turbid water, perhaps as a means to encounter additional females under conditions of low visibility.     

Resource utilization for decorating in three intertidal majid crabs (Brachyura: Majidae)

 For three spider crabs (Tiarinia cornigera, Micippaplatipes and Pugettia quadridens quadridens), patterns of algal utilization for decorating were compared with the dynamics of algae on an intertidal rocky shore reef where the crabs co-occurred. T. cornigera and P. quadridens quadridens were most abundant from autumn to spring when the dominant algae (Boodleacoacta, Sargassum hemiphyllum, S. thunbergii and Corallina pilulifera) occurred in high coverage, while M. platipes was most abundant from spring to autumn. Monthly change of algae used for decorating was not correlated with algae growing in the crab habitat for T. cornigera, but for M. platipes, it was positively correlated for two algal species, and for P. quadridens quadridens, negatively correlated for one algal species. Each species of the spider crabs used some algal species preferentially for decoration. Decorating preference experiments conducted in the laboratory showed that M. platipes and P. quadridens quadridens exhibited similar preference to their algal utilization in the field, whereas for T. cornigera, algal preference in the experiment differed from utilization in the field. Comparisons between materials used for decoration and gut contents revealed that T. cornigera and M. platipes used algal species differently for decorating and feeding, while P. quadridens quadridens used the same algal species for both decorating and feeding. Different tactics for camouflage are discussed in terms of algal utilizations by the three majid species. Received: 25 September 1999 / Accepted: 22 June 2000     

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.     

Recruitment and juvenile survival in the sea hare Aplysia juliana (Gastropoda: Opisthobranchia)

The specificity in settling response of larval Aplysia juliana (Quoy and Gaimard, 1832) on species of its food algae, Ulva spp., provided a rare opportunity for the quantitative study of recruitment and early survival rates in a non-sessile marine invertebrate. Post-metamorphic juveniles spend about 3 weeks feeding and growing on the algae before moving to the rocky habitat of the adults. Recruitment and survival were studied by collecting weekly samples of algae and holding them in the laboratory until the recruits were large enough to be seen and reliably counted. Recruitment occurred throughout the year, although the rates were relatively low in late winter and spring. Recruitment was monitored relative to algal weight and bottom area. Variation in larval abundance and algal standing crop influenced recruitment to the study area. Mortality during the juvenile stage was very high in all cases, although the rates and shapes of the survivorship curves varied between algal species and location.     

Growth,under laboratory conditions,of turbot,Scophthalmus maximus,from the Ría de Vigo (north-west Spain)

Egg production rates of Calanus glacialis (Jaschnov) were measured in the laboratory (in 1985) and in the field (in 1983–1984). In the laboratory, daily egg production was studied over one month at alternating feeding and fasting conditions. Spawning ceased after 3 d of starvation and started as soon as females were reintroduced to food. Egg production increased stepwise at 3-d intervals. Females survived more than nine months in captivity. In the field, egg production was measured during PREMIZEX 1983 and MIZEX 1984 at 5 and 16 stations, respectively. High egg production was found in polynyas on the East Greenland Shelf, where melt water induced stratification which supported a spring bloom. Highest egg production was converted into 6.1% body carbon female^-1 d^-1. Under thick pack-ice no eggs were spawned. Spawning was induced in females from a station with low food abundance by feeding them on board ship. These results from both experiments and field studies show that egg production in C. glacialis is closely related to food availability. Thus, C. glacialis exhibits a reproductive behavior similar to that of C. finmarchicus, but not C. hyperboreus, the other two dominant species in this region.     

Changes in photosynthetic pigment concentration in seaweeds as a function of water depth

We conducted a study of the relationship between changes in photosynthetic pigment content and water depth in Great Harbor near Woods Hole, Massachusetts, USA, on the green algae Ulva lactuca and Codium fragile and the red algae Porphyra umbilicalis and Chondrus crispus. A calibrated underwater photometer equipped with spectral band filters measured light attenuation by the water column. The depth required for a 10-fold diminution of photon flux was 3.6, 5.3, 6.0 and 6.0 m for red, blue, yellow and green light, respectively. Seaweeds were attached to vertically buoyed lines and left to adapt for 7 days; then, with their positions reversed, they were allowed to readapt for 7 days. All species showed greater photosynthetic pigment content with increased depth. Further, the ratio of phycobiliproteins and chlorophyll b to chlorophyll a increased with depth. Changes in pigment content were reversible and occurred in the absence of cell division. There was a net loss of pigments near the surface (high irradiance), and subsequent synthesis when seaweeds were transferred to a position deep in the water column (low irradiance). In contrast, seaweeds which were found in intertidal habitats changed only their pigment concentration, and not pigment ratio, a phenomena analogous to higher plant sun and shade adaptation. Therefore, seaweeds modify their photon-gathering photosynthetic antennae to ambient light fields in the water column by both intensity adaptation and complementary chromatic adaptation.     

Effects of seasonal pack ice on the distribution of macrozooplankton and micronekton in the northwestern Weddell Sea

The presence of mesopelagic organisms in the guts of surface-foraging seabirds feeding in open areas within seasonal pack ice in the Antarctic has given rise to questions regarding the effects of pack ice on the underlying mesopelagic community. Bottom-moored free-vehicle acoustic instruments were used in concert with midwater trawls and baited traps to examine the abundance, size distribution and vertical distribution of pelagic organisms in the uppermost 100 m of the water column during the austral spring of 1992 in two areas of the northwestern Weddell Sea, one covered by seasonal pack icc and the other free of ice cover. Acoustic largets were more abundant and significantly larger at the open-water station than beneath pack ice. However, targets at the ice-covered site exhibited a pronounced diel pattern, with the largest targets detected only at night. Samples from night trawls at the icecovered site contained several species of large, vertically-migrating mesopelagic fishes, whereas these species were absent from trawls taken during the day. In addition, baited traps deployed in pack ice just beneath the ice-water interface collected large numbers of scavenging lysianassoid amphipods, while deeper traps beneath the ice and traps at the open-water station were empty, indicating the presence of a scavenging community associated with the undersurface of the ice. These results sapport the idea that mesopelagic organisms migrate closer to the surface beneath pack ice than in open water, exposing them to possible predation by surface-foraging seabirds.     

Primary production studies during a Gyrodinium cf. aureolum (Dinophyceae) bloom in the western English Channel

Water column production measurements were made during a bloom of the marine dinoflagellate Gyrodinium cf. aureolum in the Western English Channel in July 1987. The bloom was sampled over a period of 5 d, and primary production was measured using the oxygen light and dark bottle method and the ¹⁴C technique. The photosynthetic quotient (PQ) varied between 1.2 and 3.7. During the period sampled, the proportion of phaeopigments significantly increased (from <5 to >50%) and mixed layer dissolved oxygen saturation decreased from 133 to 114%. A model was used to determine gross water column productivity in the surface mixed layer on two days, and net production values were shown to be close to zero or slightly negative. Comparisons of production values are made with other published G. cf. aureolum bloom data.     

Winter studies on zooplankton in Arctic seas: the Storfjord (Svalbard) and adjacent ice-covered Barents Sea

Zooplankton was sampled in the Storfjord and ice-covered Barents Sea during March 2003. Environmental conditions represented a typical winter situation with low air temperatures, close pack ice, and extremely low chlorophyll concentrations. Polar water dominated the hydrographic regime in the upper layers. Zooplankton distribution reflected spatial variability of hydrography. The copepods Pseudocalanus spp., Oithona similis, Microsetella norvegica together with gastropod larvae were most numerous. Biomass averaged for the entire water column varied from 3.3 to 14.3 mg dry mass m⁻³, Calanus glacialis and Parasagitta elegans contributed most, followed by C. finmarchicus, Oithona similis, and Pseudocalanus spp. Various holoplankters showed reproductive activity, especially cyclopoid copepods and chaetognaths. A few C. glacialis females laid eggs in situ, but when fed diatom cultures rapidly increased their egg production. Meroplankton including larvae of nudibranchia, bivalvia, ophiuroida, polychaeta, and bryozoa were also present. Our data demonstrate that the pelagic community of the seasonally ice-covered Barents Sea was not in a “sleeping” state at the end of the winter, but in addition to dormant stages, a portion of mainly omnivorous and several carnivorous species was reproducing.     

Metabolic responses of the sympagic amphipodsGammarus wilkitzkii andOnisimus glacialis to acute temperature variations

Acutely elevated seawater temperatures had pronounced metabolic effect on the Arctic under-ice amphipodsGammarus wilkitzkii andOnisimus glacialis, collected in May 1986 and 1988 in the Barents Sea. An increased rate of oxygen consumption vs temperature was observed for both species. In the range from 0° to 10 °CG. wilkitzkii andO. glacialis exhibit Q₁₀ values of 3.4 and 3.6, respectively. The results also indicate increased ammonia excretion rates forG. wilkitzkii andO. glacialis by an elevation of temperature from 0° to 10°C, with an overall Q₁₀ of 1.9 and 2.3, respectively. The present study demonstrates an increased O:N ratio with ambient temperature elevation from 0° to 10°C forG. wilkitzkii andO. glacialis, with overall Q₁₀ values of 2.0 and 1.6, respectively. This indicates a temperature-induced change in the metabolic substrate towards lipids.