Marine calanoid resting eggs in Norway: Abundance and distribution of two copepod species in the sediment of an enclosed marine basin

Abundance and distribution of calanoid resting eggs in sediment were examined in 1988–1989 in an enclosed marine basin (Svartatjønn; western Norway). The basin undergoes rotenone treatment and draining every winter. Egg densities were highest in autumn, when up to 2 × 10⁶ eggs m^–2 were found, and lowest in summer. Egg densities were also highest in the deeper parts of the basin and in the upper 1 cm of sediment. Hatching was initiated in late January by pumping warmer deep water from the outside area into the basin. Evidence is given for resting-egg formation inEurytemora affinis Poppe, 1880 andAcartia clausi Gurney, 1931 (present phenotype reclassified asA. teclae by Bradford 1976: N. Z. Jl mar. Freshwat. Res. 10: 159–202). Seasonal differences in inhibition of hatching, together with the longevity and tolerance of these eggs, strongly suggest that they are diapause eggs.     

Calanoid resting eggs in the Baltic Sea: implications for the population dynamics of Acartia bifilosa (Copepoda)

Resting eggs of Acartia spp. (presumably A. bifilosa Giesbr.) at densities between 65 and 125 eggs cm^-2 were found in sediment off the southern coast of Finland and represent the first observations of dormant calanoid eggs in the Baltic Sea. The highest egg densities were found at sediment depths between 4 and 8 cm. The hatching percentage varied between 0 and 90% in the different sediment layers and at different experimental temperatures (13, 16 and 19°C). Some nauplii emerged from the eggs extracted from sediment layers 9 to 10 cm beneath the surface layer and other nauplii emerged from eggs after 82 d of dark storage at 3°C. Light was not needed to trigger hatching; the nauplii were able to hatch in darkness when the eggs were resuspended in filtered seawater. Indirect evidence suggests that the marked spring peak of Acartia spp. nauplii seen in a monitoring data from the years 1973 to 1984 was derived from benthic eggs and not from spawning. The possible mechanisms for induction of hatching and the adaptive benefits of resting egg production for A. bifilosa are discussed.     

Mesozooplankton resting eggs in the Baltic Sea: identification and vertical distribution in laminated and mixed sediments

Benthic resting eggs of rotifers, cladocerans and calanoid copepods were studied at two sites with different hydrographical and sediment properties off the SW coast of Finland, the northern Baltic Sea between 1991 and 1993. Vertical distribution of the resting eggs in the sediment was studied down to 10 cm depth, and hatching of the eggs extracted from different 1-cm thick sediment layers was experimentally tested. The larvae of eight mesozooplankton taxa emerged from the eggs: Calanoida (mostlyAcartia bifilosa), cladoceransBosmina spp.,Daphnia spp.,Evadne nordmanni, andPodon spp. (mostlyP. polyphemoides), and rotifersSynchaeta spp.,Keratella spp. andAsplanchna priodonta. Some calanoid and cladoceran eggs hatched after 7 mo storage of the sediment at 3°C. A few calanoid and cladoceran eggs hatched from the 8 to 10 cm deep sediment layer. Their age, estimated by means of¹³⁷Cs analysis, was ca. 8 to 10 yr. Vertical distribution of the benthic eggs reflected the different sediment structures of the study sites. The eggs of all zooplankton taxa were more stratified in the laminated sediment than in the disturbed sediment. We suggest that the intensity of deep water mixing is the key factor for regulating the vertical distribution and hatching probability of mesozooplankton resting eggs, since it largely determines the stratification of the sedimenting material, distribution of oxygen within the sediment, and the living conditions of benthic animals causing bioturbation.     

The effect of temperature on the development and hatching of resting eggs of non-indigenous predatory cladoceran Cercopagis pengoi in the Gulf of Finland, Baltic Sea

The effect of temperature on the development and hatching of resting eggs of the Ponto-Caspian Cercopagis pengoi was studied experimentally in the Gulf of Finland, Baltic Sea. Morphological changes were monitored as the development of the resting eggs proceeded. Sexual reproduction of the C. pengoi population in the Gulf of Finland was evaluated by combining the data from hatching experiments and resting egg abundances in the sediment. Development time of resting eggs was dependent on temperature: increase in the temperature shortened the time needed until hatching. Hatching success was also dependent on incubation temperature. Almost sixfold increase in hatching success was detected when temperature increased two degrees above the storage temperature. Average resting egg abundances varied between 0.16 and 0.49 eggs cm⁻³ in the 0–6 cm sediment layer.     

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

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

Calanoid copepod eggs in sea-bottom muds. III. Effects of temperature,salinity and other factors on the hatching of resting eggs of Tortanus forcipatus

The “resting” eggs of a marine neritic copepod, Tortanus forcipatus Giesbrecht, recovered from sea-bottom sediment were hatched in the laboratory. Hatching occurred at temperatures of 13° to 30°C, no eggs hatched at 10°C. Temperatures around 25°C were found to be optimal for hatching, although the range of optimal temperatures for hatching was approximately 5°C lower in eggs stored for 14 to 15 months than in those stored for 1 to 2 months. A wide range of salinity, from 18 to 54%S, was favourable for hachting. Eggs failed to hatch within the sediment mud, which suggests that they are in a state of dormancy in the mud. Hatching was successful under both light and dark conditions.     

Longevity of subitaneous and diapause eggs of Centropages hamatus (Copepoda: Calanoida) from the northern Gulf of Mexico

Copepod resting eggs are abundant in the seabed of many bays and estuaries where they provide a potential source of recruits for growth of planktonic populations. In the northeastern Gulf of Mexico the copepod Centropages hamatus (Lillejeborg) occurs in the water column only during the late fall, winter and early spring. The species produces subitaneous and diapause eggs, and both egg types have been found in the seabed. We determined the longevity of these two egg types to ascertain their potential for contributing to the growth of the planktonic population and for sustaining a persistent egg bank. Eggs were collected from females and incubated in the laboratory under temperature and oxygen conditions chosen to simulate field conditions. The diapause eggs were also exposed to sulfide. The total hatching success of subitaneous eggs in two experiments declined from highs of 78 and 97% to zero after 60 and 90 d of exposure to anoxia. The total hatching success of diapause eggs that were exposed to anoxia for 90 d however was typically greater than 80%. Some diapause eggs hatched after being incubated under anoxia for 437 d. Diapause eggs survived longer at ambient field temperatures when incubated under anoxia (437 d) compared to normoxia (118 d). Exposure to sulfide did not result in greater mortality of diapause eggs compared to anoxia alone. Diapause eggs that were incubated at ambient field temperatures did not hatch when exposed to normoxia until the temperature dropped to <20 °C. The results of this study suggest that C.␣hamatus sustain a short-term reserve of subitaneous eggs in the seabed that provides recruits for the current year's population. The greater longevity of diapause eggs suggests that they sustain the seasonal reappearance of the species year after year in the northeastern Gulf of Mexico. However, the contribution of diapause eggs of C. hamatus from the Gulf of Mexico to a persistent egg bank is questionable since hatching ceased after 437 d. Received: 30 July 1997 / Accepted: 18 January 1998     

Biological effects of oil on early development of the Baltic herring Clupea harengus membras

The effects of petroleum hydrocarbons from two crude oils and one fuel oil (No. 1) were studied on the ontogenic development of the Baltic herring Clupea harengus membras L. Herring eggs exposed to water-soluble fractions of the oils at the time of fertilization showed no decrease in numbers of fertilized eggs compared to eggs exposed 6 or 72 h after fertilization. During embryongenesis, treatment with 3.1 to 8.9 ppm or 3.3 to 11.9 ppm total oil hydrocarbons from light fuel oil and the two crude oils respectively, gave rise to alterations in embryonic activity, decreased heart rate, and premature or delayed hatching. Although many larvae hatched from eggs exposed to contaminated water (3.1 to 11.9 ppm total oil hydrocarbons), the majority of the (70 to 100%) were malformed or dead 1 day after hatching. Exposure of eggs to 5.4–5.8 ppm total oil hydrocarbons resulted in significantly (P<0.001) decreased lengths of the larvae. Increased temperature (from 9° to 14°C) aggravated the effects of the oils. The results are discussed in relation to the potential effects of oil spills and chronic oil pollution on fish eggs and larvae in the Baltic Sea.     

Calanoid copepod eggs in sea-bottom muds. V. Seasonal changes in hatching of subitaneous and diapause eggs of Tortanus forcipatus

A warm-water neritic calanoid copepod, Tortanus forcipatus, has been found to lay two physiologically different types of eggs (subitaneous and diapause eggs), which are separately spawned in response to different seasonal environmental conditions. Subitaneous eggs are produced when the planktonic populations are at their numerical maxima, and are only shed until late summer. After peak spawning, true resting eggs are spawned which undergo diapause for 1 to 3 months. The eggs recover from their diapause around mid-winter, but hatching does not take place until the water temperature at the sea bottom exceeds 15°C in early summer.     

Effects of oil pollution on the freezing tolerance and solute concentration of the blue mussel Mytilus edulis

Exposure of blue mussels (Mytilus edulis L.) to oil-polluted sea water for 7 d in the laboratory did not affect their body fluid concentrations of inorganic ions and free amino acids. Mussels exposed to-4°C for 12 h did not freeze, whereas freezing occurred invariably in mussels exposed to-7°C or lower temperatures for the same period. Following freezing at-10°C, oil polluted mussels resumed normal activity considerably more slowly than unpolluted ones, but oil-polluted as well as unpolluted mussels showed normal activity 3 h after thawing. Freezing at-15°C was invariably lethal to individuals of both groups. One possible explanation of the delayed recovery of oil-polluted mussels frozen at-10°C may be that oil components had become concentrated to toxic levels as the amount of solvent water diminished during freezing.     

Interactions between temperature and salinity during brooding on subsequent zoeal development of the mud crab Rhithropanopeus harrisii

Female mud crabs, Rhithropanopeus harrisii, carrying newly extruded eggs, were collected from the Petaluma River (San Francisco Bay Estuarine System, California, USA) in summer 1985, and exposed to factorial combinations of temperature (20°, 25° or 30°C) and salinity (2, 5, 15, 25, or 32%.). Upon hatching, dry weights of 12 to 15 h-old zoeae were determined. Subgroups of the remaining zoeae were transferred from hatching salinities to the salinities listed above and raised until metamorphosis to megalopa. Low salinities reduced zoeal dry weights by as much as 25%. Temperature played a secondary role in reduction of hatching weight of zoeae. Survival of larvae through zoeal development was best when hatching and rearing salinities were the same; in this case, overall survival increased with temperature. Both duration of zoeal development and megalopal dry weights were strongly influenced by temperature and rearing salinity, with only a small contribution from hatching salinity. The influence of hatching salinity was most obvious at extremes of the range tested. These studies indicate that physical conditions during embryogenesis profoundly influence subsequent larval development. Interpretation of experimental approaches to study ecophysiological adaptations of larval stages should not neglect the role of physical conditions during embryogenesis.     

The effects of temperature and salinity on egg production and hatching success of Baltic Acartia tonsa (Copepoda: Calanoida): a laboratory investigation

The functional response of the aspects of reproductive success of a southwestern Baltic population of Acartia tonsa (Copepoda: Calanoida) was quantified in the laboratory using wide ranges in temperatures and salinities. Specifically, daily egg production (EP, # female⁻¹ day⁻¹) was determined for 4 or 5 days at 18 different temperatures between 5 and 34°C and the time course and success of hatching were evaluated at 10 different temperatures between 5 and 23°C. The effect of salinity (0 to 34 psu) on egg hatching success was also examined. The highest mean rates of EP were observed between 22 and 23°C (46.8–50.9 eggs female⁻¹ day⁻¹). When studied at 18 psu, hatching success of eggs increased with increasing temperature and was highest (92.2%) at 23°C. No hatching was observed for eggs incubated at low temperatures (≤12°C) that were produced by females acclimated to temperatures ≤10°C indicating a possible thermal threshold between 10.0 and 13.0°C below which only the production of diapause (or low quality) eggs exists in this population. When tested at 18°C, the hatching success of eggs incubated at 15 different salinities increased asymptotically with increasing salinity and was maximal (81.4–84.5%) between 17 and 25 psu. The high reproductive success observed over wide ranges in temperatures and salinities in this Baltic population demonstrates one of the mechanisms responsible for the cosmopolitan distribution of this species within productive, estuarine and marine habitats.     

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

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

Effects of anoxia and hypoxia on the dormancy and survival of subitaneous eggs of<Emphasis Type="Italic"> Acartia bifilosa</Emphasis> (Copepoda: Calanoida)

Subitaneous eggs of the copepod Acartia bifilosa were incubated at low oxygen concentrations and in anoxic conditions to study the induction of dormancy and the longevity of the resting eggs. Eggs were incubated in hypoxic conditions for 8 or 10 days, and also in anoxic conditions for 8–441 days. After the anoxic and hypoxic incubations, the unhatched eggs were subsequently incubated in normal oxygen conditions to reveal the viability of the eggs. Low oxygen concentrations (< 0.17 ml l^–1) induced dormancy in the eggs: they did not hatch at these concentrations in the hypox experiments but did hatch afterwards. Eggs survived well in anoxic conditions: after 10 months at 4°C, approximately 40% of the eggs still hatched, while some eggs survived in anoxic conditions for a year at 13°C, which is longer than hitherto reported for subitaneous eggs of any copepod species. Also, sediment that contained eggs of copepods was stored in the laboratory to reveal the long-term survival of the eggs. When incubated after 1.8 years storage, more than 40% of A. bifilosa eggs hatched. The study shows that subitaneous eggs may also be used as a survival strategy on seasonal time scales. This may partly explain why A. bifilosa does not produce diapause eggs in the present study area in the northern Baltic Sea.Communicated by M. Kühl, Helsingør     

Tisbe pori n. sp. (Copepoda: Harpacticoida) from the Mediterranean coast of Israel and its cultivation in the laboratory

Tisbe pori n. sp., from the Mediterranean coast of Mikhmoret, Israel, was cultivated in the laboratory and is described. The relative food values of 9 species of marine algae were evaluated according to the copepods' development time (length of period from hatching to reproductive maturity). Individuals fed on Dunaliella tertiolecta or D. primolecta developed the fastest. The rate of feeding of the adults on Dunaliella primolacta was determined. In the temperature range 17° to 31.5°C, the shortest development time (7 days) was at 26° to 28°C. The range of salinity tolerated by adults was 26 to 48.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

Respiratory physiology in summer diapause embryos of the neustonic copepodAnomalocera patersoni

The respiratory physiology of summer diapausing eggs of the neustonic copepodAnomalocera patersoni, maintained under constant temperature (13 °C) and light (12 h light:12 h dark) conditions, was characterized by a bell-shaped curve, with low O₂ uptake levels at the beginning of dormancy. This was followed by a steady rise in O₂ consumption with maximum levels of 0.002 l O₂ embryo^–1 h^–1 70 d after spawning. A slow diminution in O₂ uptake then occurred until Day 150 when minimum values of 0.0003 l O₂ embryo^–1 h^–1 were recorded, coinciding with the hatching of the first embryos. Embryos continued to hatch asynchronously up to 360 d from the moment of egg laying. When eggs were subjected to 20 °C, the respiratory activity was almost three times higher than at 13 °C, even though both respiratory curves were similar. The elevated metabolism in eggs kept at 20 °C led to death of the embryos possibly due to a total depletion of metabolic reserves. ATP content also differed at the two temperatures. Diapause eggs kept at 20 °C showed no rapid rise in ATP content as opposed to those kept at 13 °C. The results of temperature shock experiments, in which eggs were first kept at winter temperatures for several weeks, after which the temperature was raised to 20 °C for another number of weeks prior to a second period of chilling at 13 °C, showed that as long as embryos were kept at 20 °C no hatching occurred. By contrast, hatching was observed after 10 d following the resumption of winter temperatures, suggesting that low environmental temperatures are an essential prerequisite for hatching of these eggs. The type of diapause inA. patersoni differs considerably from the one described in insects and in another neustonic copepod,Pontella mediterrana. In this case, there is a U-shaped respiratory curve with greatest O₂ consumption prior to the onset or upon breaking of diapause. Differences in the two types of diapause seem to involve not only differences in O₂ consumption levels but also in the sequence of metabolic changes with time and the metabolic requirements during sommer and winter dormancy.     

Hatching and viability of copepod eggs at two stages of embryological development: anoxic/hypoxic effect

The effects of low oxygen concentrations on the hatching and viability of copepod eggs at two stages of embryological development were investigated. Fully developed eggs from Acartia tonsa (Dana) and Labidocera aestiva (Wheeler) collected between July and September 1991 at Turkey Point, Florida, USA, hatched at lower oxygen concentrations than newly spawned eggs given the same incubation periods. Since many of the newly spawned eggs subsequently hatched when exposed to normoxic conditions, it is likely that the exposure to low oxygen delayed embryonic development. At oxygen concentrations where no initial hatching occurred, the subsequent hatching success of fully developed eggs of A. tonsa was higher than that of newly spawned eggs, indicating that newly spawned eggs were more sensitive to low oxygen concentrations. No such difference was observed for the eggs of L. aestiva.     

Brooding behaviour and reproductive success in two species of free-living simultaneous hermaphrodites

Although polyclads are amongst the most structurally simple of the triploblastic metazoans, they adopt a wide range of reproductive strategies. Parental care behaviour in this group is yet to be quantified for any species. We assessed the significance of brooding behaviour to the reproductive success of two free-living marine flatworms. Echinoplana celerrima and Stylochus pygmaeus were collected from the field and placed in pairs in containers of filtered seawater where they laid batches of eggs. Both parents were then removed from half of the containers and the brooding behaviour and hatching success of eggs were quantified. There were interspecific differences in brooding behaviour. Egg masses were covered by one E. celerrima parent for 12 ± 2% of time, whereas egg masses of S. pygmaeus were covered by one or both parents simultaneously for 85 ± 8% of time. Egg batches were abandoned by both species immediately prior to the onset of hatching (10–12 days). Hatching success was generally high (~90%) and brooding did not enhance the hatching success of eggs. We assessed the significance of parental care to hatching success of E. celerrima egg masses in the presence of three potential egg predators; in the presence of other organisms. E. celerrima devoted less time to brooding; however, hatching success was not affected. The amount of time spent brooding eggs differed greatly between the two polyclad species but was not essential to their reproductive success under benign conditions. Parental care may be of adaptive value under more stressful environmental conditions commonly experienced in estuarine environments such as lowered salinity, increased hypoxia or turbidity. Covering egg batches may play an additional role of advertising sexual status and a willingness to care for eggs.