Egg production of Calanus finmarchicus : effect of temperature, food and season

The use of the egg production rate of herbivorous copepods as an important parameter for understanding population dynamics and as an index of secondary production requires knowledge of the regulatory mechanisms involved and of the response to changes in food concentrations and temperature. Furthermore, the effects of season and generation on egg production have to be studied. In this context data are presented for Calanus finmarchicus from the northern North Atlantic. Prefed and prestarved females were exposed to different concentrations of the diatom Thalassiosira antarctica over 1 to 2 wk at 0 or 5 °C, and egg deposition was controlled daily. Egg production increased with higher food concentrations, but much less when prestarved. The effect of temperatures between −1.5 and 8 °C on egg production was studied in females maintained at optimum feeding conditions. Egg production rate increased exponentially over the whole temperature range by a factor of 5.2, from 14.2 to 73.4 eggs female⁻¹ d⁻¹, and carbon-specific egg production by 4, from 2.1 to 8.5% body C d⁻¹. The response to starvation was also temperature dependent. In both the temperature and feeding experiments egg production rate was regulated mainly by changes of the spawning interval, while changes of clutch size were independent of experimental conditions. Different responses to optimum feeding conditions were observed in females collected in monthly intervals on three occasions between March and May. The March females deposited more clutches than the April and May females. In May, >50% of the females did not spawn at all. Maximum egg production rates were never >25% of the rate expected at 5 °C, indicating endogenous control of egg production in addition to food and temperature effects. Received: 4 August 1996 / Accepted: 11 September 1996     

Abundance,distribution and prey composition of scyphomedusae in the southern North Sea

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita, Cyanea lamarckii, C. capillata and Chrysaora hysoscella were studied in the southern North Sea in 2004 and 2005. Three different patterns of seasonal occurrence of medusae were distinguished: (1) the early occurring C. lamarckii (February–August), (2) C. capillata and A. aurita (April–August) and (3) the late appearing C. hysoscella (July/August–September). Cyanea lamarckii was the most frequently encountered species in this study; its highest mean abundance was 1.8 ± 2.7 ind. 100 m⁻³. The prey spectra of C. lamarckii, C. capillata and C. hysoscella contained several copepod and other crustacean species and thus make them potential competitors with fish larvae. Medusae in this study also consumed fish eggs and larvae, including clupeids, in all months analysed. Although peak spawning of sprat (Sprattus sprattus) coincides with the maximum abundance of medusae (May–June) the relative low abundance of all medusae species in this study makes jellyfish predation unlikely to be a factor controlling sprat recruitment in the time frame investigated.     

Life cycle of the copepod Calanus hyperboreus in the Greenland Sea

The seasonal ontogenetic migration of the Arctic copepod Calanus hyperboreus was described from surface-to-bottom hauls in the central Greenland Sea Gyre (GSG) and in the Westspitsbergen Current (WSC). All stages except females spent the winter below 500 m in the GSG and below 1000 m in the WSC. Seasonal ascent begins in April, and descent in July. For the C.␣hyperboreus population an active downward transport of 8.1 g m⁻² dry weight during 8 months of overwintering was estimated, similar to flux rates of particulate matter in sediment traps. Seasonal distribution of biomass was determined from weight measurements of single stages. Annual means varied from 4.0 to 9.2 g m⁻² in two different years in the GSG and were 1.1 in 1 year in the WSC. The life cycle in the Greenland Sea was reconstructed from field data on stage composition, vertical distribution, reproduction, and moult cycle phase from tooth development of CV. Laboratory experiments were conducted on the moulting of CIV and CV in fall. A 3-year (males) and 3- to 4-year (females) life cycle is proposed for the GSG and 2 to 3 years for the WSC. However, the small number of young larvae and the incomplete spring ascent by older copepodites observed in the WSC cast doubt on the reproductive success in the WSC. A suite of physiological strategies and adaptations performed by the developmental stages support survival of this species in harsh environments. Received: 25 January 1997 / Accepted: 11 February 1997     

Reproductive biology of deep-water calanoid copepods from the Arctic Ocean

Reproductive modes and egg production were studied in 15 species of meso- and bathypelagic copepods from nine Calanoida families in the Arctic Ocean. During shipboard incubation, females of seven species released eggs freely into the water and females of three species produced membrane-bound egg sacs. One species, Aetideopsis rostrata, produced a mass of eggs looking like an egg sac, but the “sac” lacked a membrane and disintegrated within 2 h. Females of four additional species were encountered with membrane-bound egg sacs in the preserved samples. In most families, only one reproductive mode, either egg-carrying (the Euchaetidae, Augaptilidae), or broadcast spawning (the Heterorhabdidae, Spinocalanidae, Scolecitrichidae, Tharybidae, Bathypontiidae) was observed. In contrast, different genera of the Aetideidae family demonstrated different reproductive modes, with broadcast spawning predominant in the benthopelagic species, and both broadcast spawning and egg-brooding in the planktonic species. Clutch size and egg diameter varied widely between species in both broadcast spawners and egg-brooders. In broadcast spawners, the clutch size varied from 1 to 95 eggs female⁻¹, while the average egg diameter ranged from 152 to 440 μm. The clutch size for egg brooders varied between 3 and 82, while average egg diameter varied from 258 to 732 μm. Deep-water broadcast spawners produced much larger eggs compared to surface-dwelling broadcast spawning species. This larger egg size may result in a reduction, or elimination, of feeding during naupliar stages, thereby improving the survival potential of deep-water species.     

Feeding ecology of Spinocalanus antarcticus, a mesopelagic copepod with a looped gut

Spinocalanus antarcticus, an abundant mesopelagic copepod in polar seas, has a greatly elongated and looped midgut, contrary to most other copepod species. The total gut length is 1.77, 1.86 and 1.90 times the total body length in adult females, CV and CIV, respectively. Gross morphology of the midgut is similar in all copepodite stages and adults. It is described here from specimens collected in the Arctic Ocean. In stratified samples from the deep Amundsen and Makarov Basins S. antarcticus showed a clear preference for the depth layer between 100 and 500 m. Generally, the guts were packed with material, but most of it was impossible to identify. In most specimens the digestive tract was filled with undefined detritus particles ("detritus balls"). They were almost spherical, heterogeneous organic aggregates of 40-100 µm diameter, with small clay-sized mineral flakes imbedded. Mineral particles in the size range of 1-10 µm were found in large quantities in the guts of many specimens. Cysts of Chrysophycea and dinoflagellates and fragments of dinoflagellates, diatoms, tintinnids and radiolarians, as well as skeletons of silicoflagellates, were rather rare; some animal remnants were also found. A high carbon/nitrogen ratio (8.9) and very high lipid content (54% of dry weight) indicated a very good nutritional state. The adaptive significance and possible feeding strategy of this deep-water copepod is discussed.     

Seasonal development of scyphozoan medusae and the predatory impact of Aurelia aurita on the zooplankton community in the Bornholm Basin (central Baltic Sea)

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita and Cyanea capillata was studied in the Bornholm Basin (central Baltic Sea) in 2002. Seasonal changes in prey composition and predatory impact were investigated by analyzing stomach contents. A. aurita occurred from July to November, with a maximum mean abundance of 2.3 ind. per 100 m³ in August, whereas C. capillata was caught in much smaller numbers from July to September. No ephyrae of either species were found; therefore, advection of medusae from the western Baltic Sea is assumed. From July to October, ~80% of A. aurita medusae was distributed in the upper 20 m above the thermocline, whereas C. capillata occurred only in the halocline below 45 m. A. aurita did not migrate vertically and fed mainly on the most abundant cladoceran species Bosmina coregoni maritima. Further prey organisms were the cladocerans Evadne nordmanni and Podon spp., mollusk larvae and copepods. Copepod nauplii and copepodite stages I–III were not eaten by the medusae, neither were fish eggs and larvae used as prey. Based on mean medusa and zooplankton abundance from the upper 20 m, the predatory impact was very low. In August, when mean abundance of A. aurita was highest, only 0.1% of the copepod and 0.5% of the cladoceran standing stock were eaten per day. However, in regions with higher medusa or lower zooplankton abundance, up to 7.9% of the cladoceran standing stock was consumed per day. Hence, A. aurita did not regulate the zooplankton community in the Bornholm Basin, and fish larvae did not suffer from competition with and predation by the medusae.Communicated by O. Kinne, Oldendorf/Luhe     

Egg production of the Arctic copepod Calanus glacialis: laboratory experiments

Egg production of single female Calanus glacialis Jaschnov fed with Thalassiosira antarctica at super-abundant concentrations (>300 g C l^-1) was determined over several weeks. Experiments were performed directly after collection from the East Greeland Current in June 1987 and 1988, and during resumed feeding after long-term starvation over 4 (October 1988), 4.5 (October 1987) and 6.5 (January 1988) mo. In addition, in June 1987, short-term starvation experiments of 5 and 15 d were conducted. Egg production was closely related to feeding in all experiments. While directly after collection eggs were produced within a few days, it took 2 wk (October 1987 and 1988) and 10 d (January 1988), respectively, to resume egg production after long-term starvation. During long-term starvation periods eggs were not laid. The decrease in total egg production with duration of starvation was due to decreasing clutch size and increasing spawning interval. In contrast, short-term starvation experiments only affected spawning interval. Interannual variability in egg production was high, with much higher clutch sizes in 1988. Average production rates in June 1988 correponded to 5% body C female^-1 d^-1, the maximum was 7.4% (1 274 eggs in 23 d). Carbon content of eggs was 0.40 g egg^-1. C. glacialis is well adapted to pulsed food events in the Arctic by its longevity; its ability to preserve its reproductive potential over several months; its rapid mobilization of ovaries; and by its high egg production rates. The implication of prolonged spawning capacity on life cycle studies is discussed.     

Egg production ofCalanus finmarchicus at low temperature

Reproduction ofCalanus finmarchicus Gunnerus collected in June 1988 in Polar water and in April 1989 in Atlantic water was studied. Single females were kept at 0°C in the laboratory for 22 d (Polar) and 77 d (Atlantic) with superabundant food concentration (> 400µg Cl^–1) of the diatomThalassiosira antarctica. There was no significant difference between the two populations, although more spent females were found in Polar water, probably due to the different dates of collection. The hypothesis of low temperature determining the geographic range ofC. finmarchicus via reproductive failure is not supported. Mean daily egg production rate of all females from Atlantic water over a 60 d period was 24.4, corresponding to 5.5% body C female^–1 d^–1, when an egg carbon content of 0.23µg is assumed. Coefficient of variation was 25%. Maximum values were 53.2 eggs female^–1 d^–1, corresponding to 12.1% body C d^–1. The highest number of eggs spawned by a single female was 3101, corresponding to a seven-fold turnover of body C during the investigation period; >20% of females produced > 2000 eggs. Body carbon content did not change significantly during the experiment; the C:N ratio increased slightly, indicating lipid accumulation. Delay of response to starvation periods of 2, 4 and 7 d duration was always 2 d: egg production ceased 2 d after the onset of starvation and continued 2 d after onset of feeding.     

Life cycle of <Emphasis Type="Italic">Pseudocalanus acuspes</Emphasis> Giesbrecht (Copepoda,Calanoida) in the Central Baltic Sea: II. Reproduction,growth and secondary production

The population dynamics of Pseudocalanus acuspes in the Central Baltic Sea were studied from March 2002 to May 2003 on a monthly basis. All stages were present year round with a stage shift from nauplii to older copepodite stages over the course of the year. Biomass, estimated from prosome length, peaked between May and September with maximum recorded values of 594 and 855 mg C m⁻² in May 2002 and 2003, respectively. Differences in biomass between stations up to a factor of 20 were observed especially in April/May and October. Mean egg production rate (EPR) showed a seasonal course and was highest in April 2002 and 2003 with 3.6 and 2.1 eggs f⁻¹ day⁻¹, respectively, corresponding to a mean weight-specific egg production rate (SEPR) of 0.13 and 0.04. Egg production seems to be limited by food from May on. Stage durations determined from moulting experiments turned out to be extremely long. Maximum growth rates based on stage durations of 15–25 days at 4°C in May and July 2003 amounted for 0.03–0.05 day⁻¹ in CI-CIV. Comparing these rates with rates derived from temperature–development relationships for P. acuspes from the literature resulted in five times higher growth rates for the latter case. Secondary production reached values up to 9.1 mg C m⁻² day⁻¹ (method for continuously reproducing populations) and 10.5 mg C m⁻² day⁻¹ (increment summation).     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.