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.     

Seasonal changes in feeding, gonad development and lipid stores in Calanusfinmarchicus and C. hyperboreus from Malangen, northern Norway

 Seasonal dynamics of feeding activity, oil sac volume, gonad development, sex ratio and spawning periods in the two sibling species Calanusfinmarchicus and C. hyperboreus, the key zooplankton copepod organisms throughout the northern Atlantic waters, were studied simultaneously in Malangen, northern Norway, during 1992. We were also tracing differences in surface- and deep-dwelling components of these populations in terms of oil sac volume and gonad development during the time period when the G₁ is preparing for a subsequent generation (G₂) or hibernation. The main difference in the life cycle strategies of these species is the earlier maturation and spawning of C. hyperboreus. No feeding activity in either of the two species was found in February, but both commenced feeding in March, prior to the spring phytoplankton peak. The larger copepod, C. hyperboreus, had a more intensive energy deposition than C. finmarchicus. The period of active feeding was much shorter for the former species, only from March through July in copepodite stages CIV and CV, and even less in females – March and April. Basically, a similar pattern of seasonal changes in gonad length and lipids was observed in the two species. In June, oil sacs in the surface- and deep-dwelling specimens were about equal, during the rest of the year, lipids in the deep CVs exceeded those in the surface. We propose that as copepods accumulated sufficient lipid reserves, they started to descend, while others, containing less fat, stayed in the upper layers feeding. The mean length of the gonads in the surface-dwelling copepods was consistently less than in their deep counterparts from October to February, so that gonad development at the expense of accumulated reserves during resting stage was confirmed. C. finmarchicus males were found in considerable numbers only in February and March, and were only occasionally found in the upper layers (0–100 m), while adult male C. hyperboreus were present from October to March, but were never found in the surface layers. The differences in life cycle timing among the two species are discussed in relation to tradeoffs with regard to foraging strategies, generation numbers, bioenergetics and predator avoidance. Received: 31 March 1999 / Accepted: 23 November 1999     

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.     

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     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

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.     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Short-term effects of sucralose on Calanus finmarchicus and Calanus glacialis in Disko Bay,Greenland

The potential effects of sucralose on the Arctic copepods Calanus finmarchicus and Calanus glacialis were studied in Disko Bay, Greenland. Sucralose is a non-calorie sweetener and chlorine derivate of sucrose containing three chlorine atoms. Scandinavian screening studies of sucralose in 2007, revealed sucralose in all effluent samples. To investigate whether sucralose is harmful to the Arctic aquatic ecosystems, possible short-term effects were investigated on egg production, hatching rate, food intake and mortality of two species of Arctic copepods. The copepods were exposed to six different concentrations (0–50,000 ng · L^?1) of sucralose, which spans the range of concentrations found in the screening studies. Exposure led to no mortality among the copepods. Food intake by C. glacialis increased with increasing concentrations of sucralose. In C. finmarchicus, food intake did not differ with increasing concentrations. No effect of sucralose was observed on egg production of C. finmarchicus. Despite increased food intake with increasing concentrations of sucralose, C. glacialis did not increase its egg production. The results show that both species responded weakly to sucralose, but with C. glacialis being possibly slightly more sensitive to sucralose than C. finmarchicus.     

Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.     

Large-scale oceanic distribution and population structure of Calanus finmarchicus, in relation to physical environment, food and predators

The investigation was carried out from 62°N to 73°N and from 14°E to 11°W in the Norwegian Sea during 19 June-12 July 1997. Regional differences in the phase of the seasonal development of the plankton community were evident, most pronounced across the Arctic front. In the Coastal and eastern Atlantic domains, post-bloom conditions prevailed, characterised by low chlorophyll a (chl a) levels and a phytoplankton assemblage dominated by coccolithophorids and small flagellates. During the study period, egg production rates of Calanus finmarchicus were low (<10 eggs female^-1 day^-1), older copepodite stages dominated, and the seasonal descent to deeper waters had started. In the Arctic domain, bloom conditions were evident by high chl a levels and a high abundance of large diatoms. Egg production rates were higher (a maximum of 29 eggs female^-1 day^-1), but the dominance of stages CI-CIII indicated that considerable spawning had already occurred prior to the spring bloom. The seasonal descent had barely started. Both invertebrate and fish predators were most abundant in the Coastal and eastern Atlantic domains, with abundance strongly decreasing north-westwards. No tight relationship between total abundance of invertebrate or fish predators and that of C. finmarchicus was apparent. However, a weak, but significant, relationship between abundance of young stages of chaetognaths and Euchaeta spp. versus young stages of C. finmarchicus was found, indicating that these invertebrate predators develop parallel to the development of the new cohort of C. finmarchicus. In early summer, C. finmarchicus had reached overwintering stages, and had started to accumulate in deeper waters in areas with the highest abundance of horizontally migratory planktivorous fish.     

Live discrimination of Calanus glacialis and C. finmarchicus females: can we trust phenological differences?

Two key players in the Arctic and subarctic marine ecosystem are the calanoid copepods, Calanus finmarchicus and C. glacialis. Although morphologically very similar, these sibling species have different life cycles and roles in the Arctic pelagic marine ecosystem. Considering that the distribution of C. glacialis corresponds to Arctic water masses and C. finmarchicus to Atlantic water masses, the species are frequently used as climate indicators. Consequently, correct identification of the two species is essential if we want to understand climate-impacted changes on Calanus-dominated marine ecosystems such as the Arctic. Here, we present a novel morphological character (redness) to distinguish live females of C. glacialis and C. finmarchicus and compare it to morphological (prosome length) and genetic identification. The characters are tested on 300 live females of C. glacialis and C. finmarchicus from Disko Bay, western Greenland. Our analysis confirms that length cannot be used as a stand-alone criterion for separation. The results based on the new morphological character were verified genetically using a single mitochondrial marker (16S) and nuclear loci (six microsatellites and 12 InDels). The pigmentation criterion was also used on individuals (n = 89) from Young Sound fjord, northeast Greenland to determine whether the technique was viable in different geographical locations. Genetic markers based on mitochondrial and nuclear loci were corroborative in their identification of individuals and revealed no hybrids. Molecular identification confirmed that live females of the two species from Greenlandic waters, both East and West, can easily be separated by the red pigmentation of the antenna and somites of C. glacialis in contrast to the pale opaque antenna and somites of C. finmarchicus, confirming that the pigmentation criterion is valid for separation of the two species.     

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

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

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

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

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.     

Reproductive strategy of<Emphasis Type="Italic"> Calanoides carinatus</Emphasis> and<Emphasis Type="Italic"> Calanus helgolandicus</Emphasis> during a summer upwelling event off NW Spain

Reproductive activity and production of the calanoid copepods Calanus helgolandicus and Calanoides carinatus were measured during a summer upwelling event off the coast of NW Spain. The upwelling pattern affected the distribution and fecundity of both species in the study area. The demographic composition of both populations and the stage of gonad maturation (e.g. the high abundance of fertilised females with mature ova) indicated active reproduction. C. carinatus, a highly fecund species associated with the African upwelling zones and considered as an upwelling specialist, showed low production rates (overall means of 15 eggs female^–1 day^–1 and 3% body C day^–1), despite the fact that the food conditions (high phytoplankton biomass dominated by diatoms) seemed to be optimal for this species. By contrast, C. helgolandicus, a temperate species that shows a strong link between spring phytoplankton blooms and reproduction time, seems to be flexible enough to take full advantage of shorter-term, enhanced feeding conditions associated with the pulsed nature of the summer coastal upwelling. Both the egg and carbon-specific production rates attained by this species (overall means of 26 eggs female^–1 day^–1 and 12% body C day^–1) were similar to values reported for a spring bloom situation. This high production would imply a long spring–summer recruitment event of C. helgolandicus in these waters. For both species the stage of gonad maturation was significantly correlated with their egg production rates and likely influenced by the food conditions; a species-specific nutritional requirement for final oogenesis is suggested. The carbon condition factor (carbon weight/prosome volume) of C. carinatus females was higher than that of C. helgolandicus, suggesting differential use of the carbon ingested; C. helgolandicus seems to use all ingested carbon to produce eggs at a high rates, whereas C. carinatus seems to store part of the ingested carbon as lipid reserves to ensure female survival and to support production during subsequent unfavourable food conditions.Communicated by S.A. Poulet, Roscoff     

Egg production and lipid content of Calanus glacialis in spring: indication of a food-dependent and food-independent reproductive mode

Female Calanus glacialis were collected in early May 1989 in the pack ice region of the western Barents Sea and were fed or starved over 11 wk. Both groups laid eggs continuously during this period, however, fed females laid up to six times more eggs. During the first 10 d after collection, both groups spawned at low rates. There-after, fed females strongly increased spawning rates and maintained high egg production levels over 11 wk, while the rates of starved females decreased. During starvation they lost 70% body carbon, 50% body nitrogen and 70% lipids. The wax ester portion decreased from 86 to ca. 60% of total lipids. Three phases of gonad development and lipid metabolism were distinguished: early gonad development; gonad maturation with a rapid decrease in lipids, especially wax esters; and spawning under fed and starved conditions, where in fed females food provided most of the energy, whereas in starved females the lipid content strongly decreased.     

Oil exposure in a warmer Arctic: potential impacts on key zooplankton species

Oil exploration activities are rapidly increasing in Arctic marine areas with potentially higher risks of oil spills to the environment. Water temperatures in Arctic marine areas are simultaneously increasing as a result of global warming. Potential effects of a combination of increased water temperature and exposure to the PAH pyrene were investigated on fecal pellet and, egg production and hatching success of two copepod species, Calanus finmarchicus and Calanus glacialis, sampled in Disko Bay, Greenland on 23–25 April 2008. The two species were exposed daily to nominal pyrene concentrations of 0-0.01-0.1-1-10-100 nM at water temperatures of 0.5, 5 and 8°C for 9 and 7 days, respectively. Daily measurements of faecal pellet production, egg production and hatching showed different responses of the two species to the applied stressors. When temperature increased, low concentrations of pyrene caused a decrease in faecal pellet production by C. finmarchicus, whereas C. glacialis faecal pellet production showed no negative response to pyrene exposure when temperatures increased. Pyrene exposure decreased egg production of C. finmarchicus at all temperatures, but the species was more sensitive at 0.5 and 8°C. A lag period of 1 day before egg production began was prolonged with several days when warmer water was combined with pyrene exposure. Egg production by C. glacialis was only negatively affected by pyrene in a dose-dependent manner at 0.5°C. Hatching success in both species was not affected by pyrene, where increased water temperatures led to a higher hatching success. In conclusion, C. glacialis seemed to be the less sensitive of the two species to the stress combination of increased water temperature and pyrene exposure. As a consequence of the differential responses of the two species, their competition can be impaired with a consequent impact on energy transfer between trophic levels.     

Long-term experiments on lifespan, reproductive activity and timing of reproduction in the Arctic copepod Calanus hyperboreus

Several experiments were conducted with starved and fed females of the Arctic copepod Calanus hyperboreus (1) to investigate their lifespan, reproductive period, egg production and egg viability; (2) to study the effect of origin, i.e., Atlantic and Arctic Waters in the Greenland Sea, on the timing of reproduction; and (3) to study the effect of time of collection on the onset of reproductive activity as a first approach to study control mechanisms of the reproductive cycle. Females collected in October produced up to 1,000 eggs and had a maximum lifespan of 164 days without feeding, whereas fed females produced up to ca. 6,000 eggs and survived up to 806 days. These observations support earlier assumptions that females were multiannual-iteroparous, i.e., capable to spawn in successive years, which would be unique for calanoid copepods. In starved females, clutch size decreased significantly with each spawning event. Viable eggs were produced during most of the life time. There was no difference in the timing of reproductive activity between females from the West Spitsbergen Current and the Greenland Sea Gyre. Fed and starved females collected in May and June began to spawn circa 2 and 4 months after collection, respectively, whereas females collected in August and October started spawning at the same time, in the middle of October. This indicates initiation of reproductive activity in the field in August, coincident with the descent into deep waters. Potential cues for the untimely spawning of females collected in spring and ‘unnatural’ feeding in fall experiments are discussed. Their large size, robustness and combination of different types of diapause in their life cycle make C. hyperboreus a good model organism to study diapause control mechanisms.     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

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

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