The first nauplius and the copepodite stages ofThalestris longimana Claus, 1863 (Copepoda,Harpacticoida, Thalestridae) and their bearing on the reconstruction of phylogenetic relationships

The first nauplius and 5 copepodite stages ofThalestris longimana Claus, 1863 are described. Sexual dimorphism makes its first appearance in the copepodid IV. Brief remarks are given on cultivation and on ecology of the species. The significance of both the naupliar and the copepodite development for the reconstruction of phylogenetic relationships of Thalestridae is discussed.     

Postembryonic development of Drescheriella glacialis Dahms & Dieckmann (Copepoda,Harpacticoida) reared in the laboratory

Summary All 6 naupliar and 5 copepodite stages of Drescheriella glacialis, a tisbid from Antarctic fast ice, are described. A key for the identification of the naupliar stages is given. Sexual dimorphism makes its first appearance in the copepodid IV. The setae configuration of the exopod of the second antenna in copepodid I is considered to be an ancestral state, and subsequent ontogenetic reductions are proposed to be apomorphic processes.     

Description of the developmental stages in Acartia bifilosa Giesbrecht (Copepoda: Calanoida)

The developmental stages and increase in length of Acartia bifilosa,a dominant planktonic copepod in Kyeonggi Bay, Yellow Sea, Korea,were investigated in a laboratory. Adults laid eggs at a rateof 2.5 eggs day^–1 female^–1 and up to 97% of eggshatched. The eggs were 92 µm in diameter and larger thanthe eggs of other Acartia species. The body segmentation wasrecognizable at the fifth nauplius stage and sex distinctionwas possible at the fourth copepodite stage. Thesix naupliusstages took -7.5 days and the five copepodite stages -8.5 days.Sexually mature males and females appeared at the 16th day,and by the 20th day the adults laid eggs that hatched and grewsuccessfully into the nauplius stages. The detailed morphologicalcharacteristics of each developmental stage are presented anddiscussed in relation to A.dausi.     

The year cycle of Eudiaptomus vulgaris (Schmeil, 1896) (Copepoda,Calanoida) in a small,acid water body during 1973. Development in the natural habitat and relationships between temperature and duration of development stages

The development of a population of Eudiaptomus vulgaris (Schmeil, 1896) in the Meeuwenven, a shallow acid guanotrofic moorland pool, has been described during one year. The population hibernates as copepodite 5 stages, adults and, to a small extent as naupliar stages N1, N2 and N3 (which could not develop further at low temperatures in autumn). In spring the population development starts at temperatures above 10°C and shows 3 or 4 pulses a year. An attempt has been made to explain seasonal changes in the size of adult males and females and in the sex ratio.In order to establish the duration of the various development stages, the animals have been cultured at different temperatures under illumination with 2000 Lux at a daylength of 14 hours. An adequate quantity of food from the natural habitat was available.Total egg development and total naupliar and copepodite development have been compared with the results of other workers, especially with those from Eckstein (1964), who studied Eudiaptomus vulgaris in the deep Schluchsee. The duration curves do not differ markedly with those of Eckstein and are strongly temperature dependent.The relation between the development times of the various stages with temperature can be generally expressed as parabolic regressions of the type D = a + b₁T + b₂T², the C₅ and adult stages being the only exception at higher temperatures. Comparison of the relative duration of the stages at different temperatures did show that younger stages can take a larger share of the total development time at lower temperatures, stage N6 being the most temperature-sensitive.     

Lipid composition of copepodite stages and adult females of Calanus glacialis in Arctic waters of the Barents Sea

Summary The composition of lipid classes and their component fatty acid are described for copepodite stages III, IV, V, and adult females of Calanus glacialis sampled from Arctic waters of the Barents Sea during summer. Was esters were the principal component of the lipid in all copepodite stages examined, averaging 73% over all the stages. The proportion of triacylglycerols varied from 1.5% to 10.5% of total lipid among copepodite stages. The lipids of adult females contained lower levels of wax esters and higher levels of triacylglycerols than copepodite stages III, IV and V. Fatty alcohols of wax esters from copepods sampled in June and July were dominated by 20:1 (n-9) and 22:1 (n-11) alcohols with the proportion of 20:1 (n-9) increasing from stages III to adult female. 14:0 and 16:0 alcohols were the principal fatty alcohols of wax esters of a sample comprising mainly of copepodites stage III taken in August. 16:1 (n-7), 20:1 (n-9) and 20:5 (n-3) were the major fatty acids in the was esters of animals captured in June and July whereas 18:1 (n-9) predominated in the August sample. The polar lipids of the copepodite stage III from August also contained lower levels of polyunsaturated fatty acids than from all stages of copepods from June and July. The data are discussed in relation to life cycle strategies and trophic aspects of Calanus glacialis in the Arctic pelagic community of the Barents Sea.     

Variation nycthémérale de la répartition verticale et de l'efficacité de collecte du zooplancton en lagune Ebrié (Côte d'Ivoire)

Vertical distribution and night and day collecting efficiencies of both nets and a Schindler trap, were studied in two shallow (4 m) unstratified bays of the Ebrié Lagoon. Some of the organisms migrate (Mysidacae, copepodite and adult stages of Acartia clausi and Pseudodiaptomus hessei) and others do not (all the nauplii, copepodite and adult stages of Oithona brevicornis, rotifers, mollusca larvae). The first are found more or less near the bottom at noon and are evenly distributed throughout the water at the beginning of the night. Mysidaceae and P. hessei strongly exhibit this pattern, none (Mysidacea) or very few specimens being collected in the water during the day. For A. clausi, depth concentration increases from C1 to adult. Net and trap estimated night densities are higher than the day ones for migrating organisms, except for C1 or A. clausi. The Night/Day ratios are equal to 28 (net) and 25 (trap) for P. hessei, 5 and 13 for the adults of A. clausi and lies between 2 and 5 for the different copepodite stages of this last species. The possible causes and consequences of the vertical distribution are discussed, the interference of turbulence on vertical distribution being outlined.     

The effects of temperature on metabolic rates of different life stages of Calanus glacialis in the Barents Sea

Summary The effects of temperature on rates of respiration, excretion and gut evacuation were examined for copepodite stages and adult female Calanus glacialis collected in areas close to the ice-edge during the arctic summer in the Barents Sea. The various life history stages responded differently to acute temperature changes above the in situ temperature (ca.-1.7°C). Respiration rates of early copepodite stages (C I to C IV) were very variable whereas excretion rates declined with increasing temperature in the range from-1.7°C to +5°C. Rate of oxygen consumption of adult females were independent of temperature between-1.7° and +5°C, but increased as temperature increased from +5° to 10°C. Rates of excretion of copepodite stage V and adult females were independent of temperature in the range from-1.7° to +2°C, whereas excretion of copepodite stages III and IV was negatively related to temperature in the range from-1.7° to +5°C. In C IV, C V and adult females the instantaneous rate of gut evacuation increased with increasing temperature. The different response patterns of metabolic rates of small copepodite stages, copepodite stage V and adult females C. glacialis to acute temperature changes suggest that the capacity for adjustment of ammonia excretion is better developed in C V's and adult females than in the younger life stages.     

Ecological investigations of the zooplankton community of Balsfjorden,northern Norway: Generation cycle,seasonal vertical distribution,and seasonal variations in body weight and carbon and nitrogen content of the copepod Metridia longa (Lubbock)

Metridia longa (Lubbock) has an annual life cycle in Balsfjorden (69°21'N:19°06'E) with spawning occurring from early to mid-May. Development through copepodite stages I-V takes place during summer. At the end of August copepodite stage V accounted for >80% of the population. Overwintering (October-March) was mainly performed by adults. At noon throughout the year copepodite stage V and adults were found within the bottom 30 m of the fjord. The majority of stage IV occurred within the bottom 80 m, while copepodite stages I-III were mainly concentrated within the surface 50 m. Body length, weight, carbon and nitrogen content, and $\text{C}\text{N}$ ratio were determined for copepodite stage V, adult males and females throughout the year. While body length was constant, weight and C and N content varied with season. Lowest $\text{C}\text{N}$ ratios (4–6) were found in copepodite stage V in July and in adults in late winter. Highest $\text{C}\text{N}$ ratios (16–20) were measured in adults from October through early winter. These results are discussed in relation to the life cycle of M. longa and the primary production cycle of the fjord. It is argued that M. longa overwinters in an active state compared with Calanus finmarchicus (Gunnerus) from the same locality.     

Diapause in the life cycle of Calanoides carinatus (Kroyer), (Copepoda, Calanoida)

The distribution and stage composition of Calanoidescarinatus (Kroyer) are described together with changes in protein and lipid content, respiration rate and gonad development in copepodite stages IV and V (CV) and adults in the Northern Benguela current (17°S, 23°S and 25°S).During active upwelling the population consisted of two parts: the surface part over the shelf was represented by all development stages, while the deep part offshore was dominated (90–95%) by diapausal CVs. In the surface CVs the surplus assimilated energy was allocated to structural growth and maturation or to synthesis of reserve lipids. CVs with large oil sacs and high lipid content descended into deeper layers and formed diapausal stock; they were characterized by a dramatic decrease of respiration rate. Increase of gonad size in association with decrease of oil sac volume in diapausal CV suggests that reserve lipids were expended not only for respiration but also for gonad development. The moulting of dispausal CVs into adults took place in deep water.These results are discussed in relation to the life cycle of the C. carinatus population and the factors causing the formation and termination of the diapausal phase.     

Ecological investigations on the zooplankton community of Balsfjorden,northern Norway: Generation cycles,and variations in body weight and body content of carbon and nitrogen related to overwintering and reproduction in the copepod Calanus finmarchicus (Gunnerus)

The generation cycles of Calanus finmarchicus (Gunnerus) are described together with the seasonal variations in length, wet wt, dry wt, carbon content, nitrogen content and $\text{C}\text{N}$ ratio in copepodite stage IV, V and stage VI males and females from Balsfjorden (69°21′N: 19°06′E), a subarctic fjord in northern Norway. C. finmarchicus overwinters in copepodite stage IV (≈ 20%) or V (≈80%) and produces one generation a year. Variations in body weight and body content of carbon and nitrogen in the different copepodite stages showed a pronounced seasonal pattern. For instance, the $\text{C}\text{N}$ ratio was lowest (4.9) in adult females during the spawning period. Copepodite stage IV and V had higher $\text{C}\text{N}$ values in summer and autumn (12 to 14) than in spring (8 to 10). Variations in length, weight and chemical composition revealed that the overwintering stock of C. finmarchicus went through two growth phases during this period. From September to January no significant changes in the measured variables were detected. During the second phase of the overwintering period, January to April, the different stages showed a profound decrease in weight and change in chemical composition. This seemed to be connected with the onset of sexual differentiation in stage V starting in January, subsequent moulting into adults and gonad maturation in these adults. These results are further discussed in relation to the different prevailing hypotheses concerning overwintering strategy in Calanus species.     

The distribution and diel vertical migration of Pseudodiaptomus hessei (Mrázek) (Calanoida: Copepoda) in a subtropical lake in southern Africa

The vertical distribution and diel migratory behaviour of Pseudodiaptomus hessei in a freshwater lake is described. All stages showed a pronounced diel migration. During daylight naupliar and copepodite instars were almost exclusively benthic in shallow areas (<10m). Copepodite V and adult stages were predominantly benthic even at 40 m. The diurnal vertical distribution pattern implies age-related differences in photosensitivity and a possible depth-regulatory mechanism, based on the existence of differential photosensitivity, is offered to account for day-depth control. Nocturnal vertical distribution, studied approximately fortnightly during 1970–71, showed no clear seasonal variation. Variations in pattern, most obvious in adult instars, correlated with prevalent lunar conditions. Nocturnal or midnight sinking was evident, particularly in post-naupliar stages. Dusk rise and dawn descent were performed at very low light intensities and paralleled changes in light penetration. Dawn descent was less rapid than the dusk rise. An ephemeral dawn rise was sometimes observed but was not a consistent feature of all stages. An endogenous rhythm of locomotor activity was recorded under laboratory conditions. Activity was low during daylight, increased sharply at dusk and then decreased through the night to reach daytime levels at dawn. The stimulus provided by changes in light intensity and/or endogenous activity changes can account for the dusk and dawn movements of P. hessei but a definitive identification of the direct migratory stimulus is not possible. Attention is called to a closer examination of the role of endogenous activity rhythms in vertical migratory behaviour. The possible adaptive value of vertical migration to P. hessei is examined briefly.     

First observations of predation by New Zealand Greenshell mussels (Perna canaliculus) on zooplankton

Observations made overseas of predation by blue mussels and zebra mussels on mesozooplankton (>200 μm) have raised concern within New Zealand that the Greenshell mussel, Perna canaliculus, which is cultured in large tonnages throughout hundreds of marine farms within the New Zealand coastal zone, could exert ecologically detrimental effects by preying on zooplankton. We conducted experiments at Clova Bay, Pelorus Sound in May 2002 to determine the rates that P. canaliculus ingests prey, up to and including the mesozooplankton size range. Single mussels from farms were incubated with seawater enriched with zooplankton (>60 μm) in gently circulated 15-l pails. Depletion of chlorophyll-a (chl-a), ciliate microzooplankton, and nauplii, copepodites, and adults of copepods was determined over 5 h, relative to controls with no mussels. Two experiments were made over consecutive days. Gut contents of these experimental mussels, and of mussels examined soon after collection from a farm, were described.Gut contents of experimental and of freshly collected mussels (standard shell length ∼90 mm) had numerous copepod parts, whole copepods and larval bivalves present. Experimental mussels cleared chl-a and ciliates from 59- to 137-l individual⁻¹ day⁻¹, respectively, averaged across the two experiments. Faster ciliate than chl-a clearance was probably caused by the high proportion (56%) of phytoplankton below the retention size for P. canaliculus (ca. 5 μm) and by faster ciliate grazing in controls than treatments. The average clearance rates of adult, copepodite, and naupliar copepod stages by mussels were 20, 31, and 49 l individual⁻¹ day⁻¹, respectively. The clearance rates of each copepod stage were not significantly different between the two experiments. Clearance of nauplii was significantly greater than of adults and copepodites, while adult and copepodite clearance rates were nearly significantly different. The mean lengths of the adult, copepodite, and naupliar copepods were 430, 265, and 165 μm, respectively. The decreasing clearance rates with increasing size and development of prey (from ciliates, through naupliar, copepodite to adult copepods), suggested that prey escape ability, related to body size and/or morphology, affected capture rates. Mussel faecal samples indicated complete digestion of the gut contents. Pseudofaecal samples showed very low rejection rates of mesozooplankton by mussels. The results are considered in context of current biophysical modelling studies of impacts of large mussel farms in New Zealand. Designs of future experiments to improve accuracy of estimates of mesozooplankton clearance rates by P. canaliculus are considered.     

The prediction of development rate and the effect of temperature for the meiobenthic copepod,Microarthridion littorale (Poppe)

Specimens of the meiobenthic copepod, Microarthridion littorale (Poppe ), were cultured in the laboratory on substrata consisting of sediment, benthic diatoms, and detrital filtrate. No significant differences were found in egg development times of M. littorale cultured in winter vs. summer, and we suggest several possible interpretations. Egg development time was independent of the number of clutches a female carried in the laboratory, suggesting confidence in estimates of egg development time without knowledge of the number of broods a female has produced. Egg, naupliar, and copepodite development rates were determined over a range of temperatures. Regression analyses indicated that both Bělehrádek's temperature function and a curvilinear model adequately described the relationship between development and temperature for all three developmental stages. A linear model was also adequate to describe copepodite development (Lack of Fit Test, P = 0.05). Eight of the reared females produced egg sacs, allowing us to estimate generation time (egg to egg) as 29.6 days at 25°C and 21.6 days at 33°C. Generation times were compared to those for a closely related benthic copepod, Tachidius discipes. Life history parameters between the two differed, and we advise caution when using predictions of development time derived from a species other than the one being studied.     

Feeding rhythmicity in a migratory copepod (Pseudodiaptomus hessei (Mrázek))

SUMMARY. This paper describes a day/night feeding rhythm in the copepod Pseudodiaptomus hessei in subtropical Lake Sibaya (27°20' S, 32°40' E) and examines the origin of this rhythmicity on the basis of field and laboratory observations.
Under natural conditions, adult and late copepodite stages (C_IV-C_V) of this copepod were predominantly benthic during daylight and migrated at dusk into pelagic waters in which they remained until dawn. Pronounced changes in feeding activity (based on the proportion of animals which contained food in the gut) were associated with this transition from a benthic to pelagic distribution; feeding activity was low by day and high at night. The younger copepodite stages (C_I-C_III) were less strongly benthic during daylight and showed correspondingly less pronounced diel changes in feeding activity.
Laboratory experiments involving concurrent light/dark and continuous dark regimes showed that batches of adult and C_IV—C_V stages fed throughout 24 h, although faecal pellet production increased with time under both light regimes. The laboratory observations are interpreted as negating the possibility that diel differences observed in the field arose through an endogenous feeding rhythm. The possibility that food resources in near-bottom waters was inadequate to support reasonable feeding activity is rejected on several counts. It is concluded that the feeding rhythm in P. hessei arises from an inferred infaunal distribution during daylight which would preclude filter-feeding activity. The circumstantial evidence supporting this suggestion is given and the suggestion is evaluated.     

Post-embryonic developmental plasticity of Arctodiaptomus salinus (Copepoda: Calanoida) at different temperatures

Ephemeral and semi-permanent saline wetlands undergo frequentunpredictable disturbances such as episodes of high salinityor desiccation. Organisms living in these systems are adaptedto survive and develop in very fluctuating and unpredictableconditions and, therefore, could help us to understand lifecycle strategies and plasticity in the context of global warming.Arctodiaptomus salinus is a common species occurring in suchenvironments and is often the main constituent of their zooplanktoncommunity, which is characterized by low diversity and shorttrophic webs. Despite the qualitative and quantitative importanceof A. salinus in these systems, no attention has been paid tothe effect of temperature on its population dynamics. For thisreason, variability in post-embryonic development times andstage-specific mortality of A. salinus were studied for thetemperature-range over which this species occurs. Mortalitywas higher in earlier naupliar and first copepodite stages.The highest developmental variability and mortality occurredat the highest temperature. Developmental instability couldexplain the high variability at high temperatures and the fluctuatingspeed of development throughout the successive stages. The interactionbetween mortality and developmental variability might be animportant mechanism ensuring persistence of A. salinus populationsin these fluctuating and stressing environments.     

A new culture system for in situ observation of the growth and development of Eucyclops serrulatus (Copepoda: Cyclopoida)

A practical and convenient method of rearing Eucyclops serrulatus in a microculture environment is described. A complete life cycle of E. serrulatus was maintained in a narrow space on a microscope slide glass on which a cover glass of 22 x 40 mm in size was mounted at a height of 0.8 mm. The culture medium was constituted by bottled mineral water boiled with grains of Glycine max (soybean). Chilomonas paramecium, a free-living protozoan organism, was provided as live food. Growth of nauplii hatched from eggs to the first stage of copepodite took an average of 7.7 days, and the growth of copepodite 1 to the egg-bearing adult female took an average of 20.1 days in the microculture cell with an average life time of 44.7 days. Continuous passage of copepods was successfully maintained as long as sufficient medium and food were provided. The microculture method enables an in situ microscopic observation on the growth and developmental process of helminth larvae experimentally infected to copepods as well as of copepod itself. Furthermore, it does not require anesthetization and, therefore, minimize the amount of stress exposed to copepods during the handling process.     

Errors in the estimation of recruitment of early stages of mesocyclops leuckarti (claus) caused by the diurnal periodicity of egg-production

The seasonal and bathymetrical distribution of Mesocyclops leuckarti (Claus) was investigated during 1969–1975 Development time of egg, naupliar and copepodite stages were found experimentally at three temperatures (15°, 22° and 27°C). The calculations of population dynamics generated some unexplained results.Experimental study of the influence of light-dark conditions and temperature on the process of laying and hatching, together with the field data, helped us to describe an artifact when sampling is done at early morning.Contribution no. 123. Israel Oceanographic & Limnological Research Ltd.     

Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.     

On the trophic fate of Phaeocystis pouchetii (Harlot). III. Functional responses in grazing demonstrated on juvenile stages of Calanus finmarchicus (Copepoda) fed diatoms and Phaeocystis

The objective of this study was to quantify the functional responsein feeding rate in the various developmental stages of Calanusfinmarchicus to different concentrations of the diatoms Thalassiosiranordenskioeldii and Porosira glacialis, and the haptophyseanPhaeocystis pouchetii. Grazing of copepodite stage I–VC.finmarchicus was measured using two different approaches.Feeding rates were obtained from either incubation experiments,estimating the rate of removal of particles from suspension,or by quantifying the turnover rate of the plant pigments inthe gut. Clearance as a function of algal concentration (1–30µg plant pigment 1^–1) was described in juvenilestages of C.finmarchicus fed the diatoms T.nordenskioeldii [20µm equivalent spherical diameter (ESD)], P.glacialis (40µm ESD), and two size categories (30–100 µmand >100 µm ESD) of the gelatinous alga P.pouchetii.When the copepodite stages were fed T.nordenskioeldii, the gutcontent of plant pigments was in general higher than when fedP.glacialis. Rates obtained were variable when the same copepoditestages were offered the two size categories of P.pouchetii,but within the same order of magnitude as those obtained forthe larger diatom. At unialgal diets, diatoms were more readilyconsumed than the larger size fraction among colonies of P.pouchetiiby copepodite stage I–III C.finmarchicus. But given anappropriate prey size, C.finmarchicus grazed both diatoms andcolonies of gelatinous algae at equal rates. A linear relationshipbetween gut content and food concentrations <10 µgchlorophyll 1^–1 was found. This indicates that the ingestionrate in C.finmarchicus is directly proportional to the ambientfood concentration during the most productive period in Mayand June in high latitudes irrespective of algal species present. ¹Present address: Marine Biological Laboratory, University ofCopenhagen, Strandpromenaden 5, DK-3000 Helsingør, Denmark ²Present address: Greater Copenhagen Council, Gl. KøgeLandevej 1–3, DK-2550 Valby, Denmark