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     

Diet composition and feeding intensity of horse mackerel,<Emphasis Type="Italic"> Trachurus trachurus</Emphasis> (Osteichthyes: Carangidae) in the eastern Adriatic

Diet composition and feeding intensity of the horse mackerel, Trachurus trachurus, collected in the eastern Adriatic Sea, were examined. Stomach contents of 1,200 specimens, taken at monthly intervals (January–December 1996), were analyzed. The stomachs contents consisted of five major prey groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda and Teleostei. Euphausiacea constituted the most important food resource by weight, number and frequency occurrence. Teleosts were the second most important food category, while mysids, decapods and cephalopods were occasional food. There was little seasonal variation in diet. Euphausiids were dominant prey during all seasons, and were especially abundant from spring and autumn. Feeding intensity of horse mackerel varied during the year. The lowest intensity of feeding was recorded in winter (February, March) and early spring (April) during the lower sea temperature as well as at the time of intensive spawning. Feeding activity increased upon spawning period (May and June) and was also higher during summer. Feeding intensity and diet composition changed during the diurnal cycle. The highest feeding intensity was recorded at night and during early morning hours. Euphausiids, mysids and a greater part of teleosts dominated night and morning diet, while decapods and cephalopods were most frequently in the daily and evening diet.Communicated by O. Kinne, Oldendorf/Luhe     

Vertical distribution and mortality of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during overwintering in oceanic waters southwest of Iceland

The seasonal vertical distribution and the predatory regime encountered by Calanus finmarchicus were studied along a transect across the Reykjanes Ridge in the oceanic area southwest of Iceland from data collected during four cruises between November 1996 and June 1997. The mortality for the overwintering period was estimated using linear regressions of density estimates from November 1996 to April 1997. In addition, we also estimated the mortality of the oldest population stages (C4 and older) in April and June by applying the vertical table method. During winter (November/December–January/February), the animals mainly resided at a depth of ∼300–1,500 m in the water of Atlantic origin. Ascent to upper layers took place mainly during March and April, and continued until May. During all cruises, continuous deep-scattering layers were observed, mainly within the range of 400–500 m to 700–800 m depth. Based on sampling with a Harstad pelagic trawl in April, the scattering was mainly ascribed to jellyfish (mainly Periphylla periphylla), small mesopelagic oceanic fishes (several species but Benthosema glaciale and Maurolicus muelleri were most abundant), euphausiids (mainly Meganyctiphanes norvegica) and shrimps (mainly Sergestes arcticus). These species may represent a predatory threat to overwintering C. finmarchicus. From November to April, daily per capita mortality rates were estimated to be (mean ± 95% CL) 0.004 ± 0.0028 for the total data set, and 0.004 ± 0.0033 and 0.004 ± 0.0023 (day^-1) for the Iceland Basin and Irminger Basin, respectively. Mortality rates were higher later in life (mean ± 95% CL) for C5/females (0.13 ± 0.044) and C5/males (0.19 ± 0.051) than for C4/C5 (0.00 ± 0.035) when averaged over all samples taken in April and June 1997. We discuss how the observed distribution and mortality rates of overwintering C. finmarchicus might be related to predatory regime.     

Recruitment,growth, and diel vertical migration of Euphausia pacifica in a temperate fjord

The pelagic crustacean Euphausia pacifica Hansen was sampled with a multiple-sample 1.0 m² Tucker trawl and a multiple-sample 1.0 m² vertical net in Dabob Bay, Washington on 17 dates between May 1985 and October 1987. Size (stage) structure and abundance of the population were determined for each date, while vertical distribution and diel migration were determined for 13 dates. Although internannual variability in both timing and magnitude of events occurred, consistent patterns were discernable. The population produced a large pulse of larvae (2 to 5 mm) in late spring of each year, apparently in response to the vernal phytoplankton bloom. Much lower abundances of larvae occurred during summer and autumn of each year, and larvae were completely absent during winter. Recruitment to the juvenile (6 to 9 mm) and adult (10 mm) stages was strongest during the summer, with abundances of these individuals peaking in summer and autumn. Individual growth rates, determined by modal progression analysis, were calculated for E. pacifica. Rates ranged from zero for some adult cohorts during the winter to 0.12 mm d^-1 for larvae during spring. The latter are among the highest ever reported for this species in the field. The vertical distributions and diel vertical migrations (DVM) of E. pacifica varied seasonally and between size (stage) classes. At night, all size classes were distributed in the surface layer (upper 25 m) irrespective of season or year. During the day, the larger/older stages were always distributed at middepths (50 to 125 m). In contrast, the daytime distribution of the larvae was more variable, being concentrated at the surface during spring and early summer of 1985, and at increasing depths later in the summer and autumn of 1985 and again in spring of 1986. This resulted in invariant DVM in the juveniles and adults, but variable DVM in the larvae, the latter of which is hypothesized to be a response to variable abundances of zooplanktivorous fish.     

Seasonality of reproduction in the cushion star Odontaster validus at Signy Island, Antarctica

The reproductive and larval biology of the common Antarctic cushion-star Odontaster validus Koehler (Echinodermata: Asteroidea) were studied in a shallow-water population at Signy Island in the maritime Antarctic. The gonad index peaked in April/May, and injection with 1-methyladenine indicated that competence to spawn peaked in May/June. Gastrulae were first detected in May, and maximal water-column larval densities were measured in July. Reproduction was therefore highly seasonal, although not all individuals appeared to spawn each year. The pyloric caeca index increased in summer (typically November to March, but variable between years) and decreased during winter (June to October), suggesting that feeding activity was seasonal. All aspects of biology investigated showed significant interannual variability. Comparison with previous studies in the high Antarctic at McMurdo Sound indicated a similar if more pronounced seasonality at Signy, but the warmer seawater temperatures in the maritime Antarctic (particularly in late winter/early spring from September onwards) resulted in faster larval development at Signy. Received: 21 April 1997 / Accepted: 28 January 1998     

Trophodynamics and seasonal cycle of the copepod Pseudocalanus acuspes in the Central Baltic Sea (Bornholm Basin): evidence from lipid composition

Seasonal lipid dynamics of the copepod Pseudocalanus acuspes were studied in the Bornholm Basin (Central Baltic Sea) on a monthly basis from March 2002 until March 2003 and were interpreted in light of life cycle strategies and diet selection. The individual total lipid content of females ranged from 0.9 to 1.8 μg, with relative wax ester content reaching a significant maximum in May (44% of total lipids) and minimum (17% of total lipids) in April and November. Significant changes in size, lipid content, lipid classes and fatty acid composition of structural as well as storage lipids suggested five characteristic seasonal phases that were induced by different feeding histories and environmental conditions. Storage lipids were characterized by 18:1(n−9) as major component, which ranged between 44% of total fatty acids in June and 23% in February. The strong coherence between 18:1(n−9) in the seston lipids and the occurrence of ciliates emphasized the importance of ciliates in the diet of P. acuspes. As indicated by changes in the amounts of fatty acid markers, other food sources varied over the year, suggesting an opportunistic feeding behavior. The spring period was characterized by an increase in typical diatom and dinoflagellate markers, whereas other sources, potentially cyanobacteria, became more important during summer. The life cycle strategy is discussed with respect to extant adaptations to high latitudinal habitats.     

Water-soluble vitamins in natural plankton (copepods) during two consecutive spring blooms compared to vitamins in Artemia franciscana nauplii and metanauplii

Weight-specific contents of asorbic acid, thiamin, riboflavin, niacin, pantothenic acid, vitamin B₆, biotin, folate and vitamin B₁₂ were analysed in natural zooplankton collected from a sea water lagoon in a halibut fry production plant. The samples of natural zooplankton were collected from April/May to the end of June during two consecutive spring blooms. For comparison, the same vitamins were measured in nauplii of Artemia franciscana, just after hatching and after enrichment for 24 and 48 h with a commercial diet. The changes in vitamin content in zooplankton, which consisted mainly of Temora longicornis, during the two spring blooms were <2.5-fold for each specific vitamin. The large number of high quality halibut fry produced using the plankton as a dietary source suggests that the plankton contained sufficient amounts of all water-soluble vitamins. With the exception of thiamin and ascorbic acid, Artemia– both newly hatched and enriched for 24 and 48 h – contained higher levels of water-soluble vitamins than natural zooplankton. Enrichment feeding of Artemia for 48 h stabilised or increased the level of the water-soluble vitamins. Received: 8 March 1999 / Accepted: 21 February 2000     

High reproduction of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during a diatom-dominated spring bloom

Feeding, egg production, hatching success and early naupliar development of Calanus finmarchicus were measured in three north Norwegian fjords during a spring bloom dominated by diatoms and the haptophyte Phaeocystis pouchetii. Majority of the copepod diet consisted of diatoms, mainly Thalassiosira spp. and Chaetoceros spp., with clearance rates up to 10 ml ind⁻¹ h⁻¹ for individual algae species/groups. Egg production rates were high, ranging from ca 40 up to 90 eggs f⁻¹ d⁻¹, with a hatching success of 70–85%, and fast naupliar development through the first non-feeding stages. There was no correlation between the egg or nauplii production and diatom abundance, but the hatching success was slightly negatively correlated with diatom biomass. However, the overall high reproductive rates suggested that the main food items were not harmful for C. finmarchicus reproduction in the area, although direct chemical measurements were not conducted. The high population egg production (>1,20,000 eggs m⁻² d⁻¹) indicated that a large part of the annual reproduction took place during the investigation, which stresses the importance of diatom-dominated spring phytoplankton bloom for population recruitment of C. finmarchicus in these northern ecosystems.     

Effects of starvation,season, and diet on the free amino acid and protein content of<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> females

Effects of food availability and season on the free amino acid (FAA) and total protein content of the copepod Calanus finmarchicus females were investigated in two mesocosm experiments on the Norwegian west coast in spring and autumn. Starved C. finmarchicus females showed no change in total FAA content, but the FAA pool composition changed drastically. During the first 10 days of starvation the protein content showed a moderate decline (<2 µg ind ^-1); however, during the following 21 days the total content was drastically reduced, from 63 to 9 µg ind ^-1. This supports the notion of a sequential catabolism of endogenous nutrients during starvation. In females at high food concentrations, the body protein content increased during spring, but decreased during autumn. The FAA pool composition of females differed between spring and autumn in 14 of the 18 FAA investigated. Reduced body protein content and increased proportion of essential free amino acid were observed during starvation. Similar changes were observed in females sampled at the end of the mesocosm experiments in the autumn. The results suggest that mature C. finmarchicus females are in a negative protein balance during autumn, despite high food concentrations, contributing to a lower fitness than in females maturing during the spring.     

Inverse vertical migration and feeding in glacier lanternfish (<Emphasis Type="Italic">Benthosema glaciale</Emphasis>)

A bottom-mounted upward-facing 38-kHz echo sounder was deployed at ~400 m and cabled to shore in Masfjorden (~60^°52′N, ~5^°24′E), Norway. The scattering layers seen during autumn (September–October) 2008 were identified by trawling. Glacier lanternfish (Benthosema glaciale) were mainly distributed below ~200 m and displayed three different diel behavioral strategies: normal diel vertical migration (NDVM), inverse DVM (IDVM) and no DVM (NoDVM). The IDVM group was the focus of this study. It consisted of 2-year and older individuals migrating to ~200–270 m during the daytime, while descending back to deeper than ~270 m during the night. Stomach content analysis revealed increased feeding during the daytime on overwintering Calanus sp. We conclude that visually searching glacier lanternfish performing IDVM benefit from the faint daytime light in mid-waters when preying on overwintering Calanus sp.     

Biology of euphausiids in the subarctic waters north of Iceland

The seasonal abundance, maturity, spawning, and population dynamics of Thysanoessa inermis (Krøyer, 1846), T. longicaudata (Krøyer, 1846), and Meganyctiphanes norvegica (M. Sars, 1857) were studied in the subarctic waters north of Iceland from February 1993 to February 1994. The material was sampled at approximately monthly intervals along a transect of eight stations extending from 66°16′ to 68°00′N at 18°50′W. Information on temperature and chlorophyll a concentrations is also presented. Spring warming of the water began in March to April and maximum temperatures were recorded in August (3.8?°C). The spring bloom of the phytoplankton started in late March and highest chlorophyll a concentrations were measured during middle to late April (7.0?mg chlorophyll a m^?3). T. inermis was the dominant species in the samples, constituting 77% of juvenile, male and female euphausiids present. The greatest abundance of juvenile, male and female T. inermis and M. norvegica was observed during autumn and winter, with lower abundance in spring and summer. T. longicaudata showed only limited changes in seasonal abundance. Male T. inermis had spermatophores in their ejaculatory ducts from February to May, while mature females had spermatophores attached during April and May. T. longicaudata males bore spermatophores from February to July, whereas females only bore spermatophores in April and May. M. norvegica males had spermatophores from February to April, while the single female with spermatophores was caught in February. Euphausiid eggs were first recorded during the latter part of April; the highest numbers of eggs were observed in the samples taken in late May. Maximum numbers of nauplii of both Thysanoessa spp. and M. norvegica were recorded in late May. The main spawning of the euphausiids coincided with the phytoplankton spring bloom. Most male T. inermis took part in breeding at 1 yr of age while most females appeared not to mature until 2 yr of age. T. inermis has a life span of just over 2 yr, T. longicaudata appears to live just over 1 yr. Limited data did not allow the life span of M. norvegica to be determined.     

Biannual spawning periods and resultant divergent patterns of growth in the estuarine goby Pseudogobius olorum: temperature-induced?

Samples of juveniles and adults of the goby Pseudogobius olorum were collected from seven sites in the shallows of the upper Swan Estuary, Western Australia, using a 3 mm-mesh seine net on one or two occasions in each month between September 1983 and April 1985. The mean gonadosomatic index of female fish rose from very low values in winter (June–August) to a sharp peak in mid-spring (October), reflecting the rapid maturation of ovaries over, this period. Ovaries with post-ovulatory follicles and ovaries that were undergoing degeneration were present, in November and December, but were then either rare or absent in those members of the corresponding cohort which survived into January and February. Female fish with advanced oocytes and mature ovaries were not found in December and January, but were present in February to April. The above trends exhibited by ovarian maturity indices, together with the appearance of larvae and small fish in both spring and autumn, demonstrate that P. olorum spawns in both spring and autumn and at best to only a limited extent in summer. Length-frequency and gonadal data show that the progeny of the spring-spawning group frequently spawn in the following autumn, when they are 5 mo old, and that those of the autumn-spawning group frequently spawn in the following spring, when they are 7 mo old. Some representatives of these two spawning groups survive through the winter and summer, respectively, to breed in a second season. Growth of the progeny of the spring-spawning group was relatively rapid between late spring and mid-autumn, whereas that of the autumn-spawning group was negligible during winter, but then inceased markedly in spring. It is proposed that the biannual spawning periods in each year by P. olorum in the Swan Estuary developed as a result of a rise in water temperature over the last few thousand years. Such a rise would have brought forward further into spring and extended later into autumn the periods when the water temperatures lie within the range (20 to 25°C) at which P. olorum typically spawns. However, mid-summer is now characterised by water temperatures >25°C, which are considered less conducive to reproductive success.     

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     

Biological weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. II. Calanus finmarchicus (Copepoda) eggs

The copepod Calanus finmarchicus Gunnerus is a key component of the planktonic food web in the Gulf of St. Lawrence, Canada. In this region, productivity-determining biophysical interactions occur in the upper 0 to 30 m of the water column. The eggs and nauplii of C. finmarchicus are found in this layer. Measurements of the diffuse attenuation coefficients for solar ultraviolet-B radiation (280 to 320 nm, UV-B) at various locations in this region indicated maximum 10% depths (the depth to which 10% of the surface energy penetrates) of 3 to 4 m at a wavelength of 310 nm. This represents a significant percentage of the summer mixed-layer water column: organisms residing in this layer are exposed to UV-B radiation. Laboratory experiments using a Xenon-arc-lamp based solar simulator revealed that C. finmarchicus embryos exposed to UV-B exhibited high wavelength-dependent mortality. The strongest effects occurred under exposures to wavelengths below 312 nm. A significant percentage of nauplii hatched from eggs exposed to these wavelengths exhibited malformations indicative of errors in pattern formation during embryogenesis. At the shorter wavelengths (<305 nm), UV-B-induced mortality was strongly dependent on cumulative exposure. The biological weighting function (BWF) derived for UV-B-induced mortality in C. finmarchicus eggs is similar to that reported for naked DNA. This suggests that the UV-B-induced mortality effect on C. finmarchicus embryos is a direct result of DNA damage. There was no evidence of a detrimental effect of ultraviolet-A radiation (320 to 400 nm). Calculations based upon the BWF indicate that, under current noon surface irradiance, 50% of C. finmarchicus eggs located at or very near (within 10 cm) the ocean surface will be dead after 2.5 h of exposure. Under solar spectral irradiance simulating a 20% decrease in ozone layer thickness, this time drops to 2.2 h. These are first-order estimates based upon irradiance taken at a time of day during which the values would be maximal. Nonetheless, they illustrate the relative changes in UV-B effects that will result from ozone layer depletions expected over the coming decades. It is also important to point out that variability in cloud cover, water quality, and vertical distribution and displacement within the mixed layer, can all have a greater effect on the flux of UV-B radiation to which C. finmarchicus eggs are exposed than will ozone layer depletion at these latitudes. Received: 2 March 1998 / Accepted: 18 December 1998     

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.     

Effects of season and maternal food on<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> reproduction,with emphasis on free amino acids

Effects of maternal food environment and season were examined during spring and autumn on females, eggs and nauplii of Calanus finmarchicus, in different natural prey suspensions or cultures of Rhodomonas baltica. Females sampled in spring were in general larger, had higher protein content, and showed higher egg production and hatching rates, than in autumn. The cumulative egg production was almost double in spring compared to autumn (females fed R. baltica). Females had higher content of free amino acids (FAA) and free essential amino acids (EAA) in autumn than in the spring. Also, the FAA contents in eggs and nauplii were higher in autumn than in spring. In contrast, the composition of EAA in eggs was constant between seasons, indicating maternal regulation. The highest cumulative egg production was correlated with a high similarity in the free pool of EAA in the food suspension and the female copepod. Thus, the data support the hypothesis that similarity in the free pool of EAA of diet and female promotes high fecundity and egg hatching success in C. finmarchicus.     

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     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Fission and its effect on population structure of Holothuria atra (Echinodermata: Holothuroidea) in Taiwan

Populations of Holothuria atra Jaeger occurring at Wanlitung and Nanwan were sampled monthly at spring tide from March 1990 to February 1992 in southern Taiwan. At Wanlitung, small individuals with evidence of fission and regeneration were abundant in shallow tidepools on the wide reef flats (density: 98±40 ( ± SD) 100 m^-2, body weight: 6 to 182 g). At Nanwan, large individuals with no evidence of fission were sparse in deep tidepools and narrow flats [density: 0.24±0.07 ( ± SD) 100 m^{- 2}, body weight: 351 to 1400 g]. At Wanlitung, fission occurred throughout the year with an average monthly fission frequency of 4.5%, peaking at 18% in September 1990 and 16% in August 1991. Fission was accomplished by revolving, twisting, and stretching of the body, resulting in two unequal fragments. The minimal weight of 3007 individuals measured during 1 yr at Wanlitung was 6 g. All ten individuals weighing 6 to 9 g ( ± SD = 6.7 ± 1.1) 100 m^{- 2} showed evidence of regeneration from fission products. Nine large individuals (>500 g) transferred from Nanwan to Wanlitung and 50 small individuals (<100 g) transferred from Wanlitung to Nanwan in February 1991 showed no evidence of fission during the following year. Fission occurs in small individuals living in shallow tidepools, suggesting that fission probably is triggered by a stressful environment resulting from solar radiation at the unusually low water level of spring tide. Fission of H. atra at Wanlitung results in a population consisting of small individuals.