The diatom Thalassiosira rotula affects reproductive success in the copepod Acartia clausi

Fecundity, egg viability and fecal pellet production are reported for Acartia clausi females collected in the Bay of Naples, Italy, from April to October 1992 and fed either with a diatom (Thalassiosira rotula) or dinoflagellate (Prorocentrum minimum) diet, at food saturated conditions. The diatom diet significantly reduced both egg and fecal pellet production as well as hatching success. Blockage of egg development occurred with both axenic and non-axenic cultures of T. rotula, suggesting that inhibitors were provided by the diatoms and not by the bacteria associated with diatom cultures. Low hatching success was also artificially induced by exposing newly spawned A. clausi eggs to high concentrations of diatom extracts, indicating the presence of deleterious, inhibitory compounds blocking copepod embryogenesis. Fecundity and hatching success diminished significantly with female age. In contrast, female longevity was not significantly modified by food type. The presence of males did not significantly alter fecundity or egg viability. Females continued to produce viable eggs throughout the period of incubation, with and without males, in both food conditions, indicating that remating is infrequent and not necessary to sustain viable egg production in this species. The succession in low and high population densities may therefore be the outcome of variations in survival rates of eggs, rather than reproductive protential perse; such variations may strongly depend on the adult copepod diet.     

Scanning electron microscopy investigation of bacterial colonization of the marine copepod Acartia clausi

Association between copepods and bacteria was observed in many scanning electron micrographs. Particular sites on the copepods were selectively colonized by bacteria; the joints of segments and legs, swimming legs and depressed parts of the body surface were found to be densely covered with bacteria. In comparison, bacterial attachment to copepod skeletons in fecal pellets excreted by chaetognaths was not selective; bacteria were sparsely found all over the copepod. Between 9 to 30% of copepods in Tokyo Bay waters had attached bacteria in January and April 1983.     

Relationship between growth rate and egg production in the copepod Acartia clausi hudsonica

Egg production by Acartia clausi hudsonica ceases at low concentrations of Isochrysis galbana as food and at high levels reaches a maximum that increases with temperatures in the natural range. This increase parallels the rate of production of fecal pellets. Females without males can produce about 400 eggs before entering a generally short postreproductive period. Weight increments between copepodite stages are exponential and, assuming isochronal durations of stages and that development time of older stages is the same mutiple of embryonic duration at all temperatures, a temperature-dependent rate is estimated for pre-adult growth. We demonstrate that this growth rate also predicts the observed maximal rate of egg production by the nongrowing adult females. Published data for other copepod species (except Pseudocalanus) are inadequate for wider testing of this hypothesis, but available data do suggest that no such simple rule governs total output of eggs by females of different species.     

Discriminate feeding of the calanoid copepod Acartia clausi in mixtures of phytoplankton and inert particles

Adult female Acartia clausi were allowed to feed in the laboratory on each of three algae, the small green alga Dunaliella tertiolecta (5.6 m ESD), the diatoms Thalassiosira decipiens (13.4 m ESD) and T. nordenskioldii (17.7 m ESD), singly or in mixtures with polystyrene beads (15.7 m ESD). The ingestion rate on beads was much lower than that on cells, even when the supply of beads was one order of magnitude higher than that of cells. Beads offered singly were not ingested. In the experiments with T. nordenskioldii, the more cells adult females ingested, the higher the ingestion rate on beads. In addition, a linear regression equation describing the relationship between bead concentration and ratio of ingested beads to ingested cells was highly significant. These results suggest that beads are ingested by chance in feeding bouts on T. nordenskioldii. The bead interference on the ingestion rate on cells was observed in the experiments with D. tertiolecta and T. decipiens, but not with T. nordenskioldii. In a mixture of beads and T. decipiens, juvenile A. clausi discriminated beads from cells more successfully than adults. The presence of beads did not affect the ingestion rate of juveniles on cells. Starved adult females tended to ingest more beads than well fed individuals. The possible effects of body size and hunger on the discriminate feeding of copepods are discussed.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

Long-term food modification by Acartia clausi: A preliminary view

After acclimation, the copepod Acartia clausi was allowed to graze for 5 days in a mixed suspension of two discrete size classes (species) of the chaining diatoms Thalassiosira spp. derived from continuous culture. Total particle numbers and particle size distributions of Thalassiosira spp. were stable throughout the 5 days, indicating that the effects of algal removal and modification due to grazing were balanced by algal growth. Grazer ingestion is the predominant process affecting all size classes of the smaller diatom population (T. nordenskioldii); however, both ingestion and chain modification are observed with the larger diatom (T. gravida). Although the greatest percentage removal of algal volume occurs in the largest algal size classes for each algal species, the greatest volume removal occurs at the modal peak (T. nordenskioldii) or just to the right of the modal peak (T. gravida). Flask-to-flask replicability of experiments was tested with the single-celled T. fluviatilis, and these tests were compared to the long-term experiments with T. nordenskioldii and T. gravida. Net particle removal occurs on both the large and small algal species in mixed suspension, not just on the larger-cell-side of the T. gravida distribution. Although 80% of the particles ingested are from the T. nordenskioldii distribution, 80% of the volume ingested is from the T. gravida distribution. The apparent multi-peak selection observed in our data is discussed in reference to two separate hypotheses and in light of other recent work pertaining to selective grazing by copepods.     

The effect of low concentrations of phenol and ammonia on egg production rates,fecal pellet production and egg viability of the calanoid copepod Acartia clausi

Estimates of daily fecundity, hatching success and fecal pellet production are reported for Acartia clausi females exposed for 10 d to low levels of phenol and ammonia. Copepods were collected in 1991 and 1992 from a southern coastal area of the Mar Grande of Taranto (southern Italy). A reduction in egg numbers and fecal pellet production was observed for females after 8 d of exposure to 500 g l^-1 phenol concentration. Ammonia (120 g l^-1) produced a significant increment in egg production, but hatching success was reduced by about 50% after nine exposure days. A. clausi was more sensitive to ammonia than phenol at high concentrations (24-h LC₅₀ phenol-32.26 mg l^-1; 24-h LC₅₀ ammonia=0.91 mg l^-1). At low concentrations, only long-term exposure to phenol determined a reduction in fecundity.     

Effects of suspended sediments on egg production of the calanoid copepod Acartia tonsa

The estuarine copepod Acartia tonsa was collected on several occasions between 4 April and 14 August 1985 from Terrebonne Bay, Louisiana (29°08N; 90°36W) and the effects in its diet of suspended sediments, collected from the same area, were measured at five different concentrations of sediment (100 to 1 000 ppm) and six phytoplankton concentrations (500 to 13 000 cells ml^-1 Thalassiosira weissflogii). Egg production rate was used as an index of diet quality. At low phytoplankton concentrations (500 cells ml^-1), and at intermediate phytoplankton concentrations (2 000 cells ml^-1) for previously starved copepods, egg production was reduced by up to 40% at a sediment concentration of 250 ppm and further reduced at higher sediment concentrations. At higher food concentrations (4 000 to 13 000 cells ml^-1), suspended sediment had no effect on egg production rates at sediment concentrations up to 500 ppm. Rates were reduced only at the highest sediment concentration of 1 000 ppm. Under most natural conditions, suspended sediment would not significantly affect egg production rates in A. tonsa.     

Egg production and hatching success in North Adriatic Sea populations of the copepod acartia clausi

Egg production, fecal pellet production and hatching success are reported for Acartia clausi females sampled during three cruises in February 1997, 1998 and June 1997 at 20-24 stations along 4 transects in the North Adriatic Sea. Dramatically low hatching rates were recorded during both diatom bloom events in February as opposed to much higher rates during post-bloom conditions in June, even though A. clausi productivity during the bloom was apparently high. These results are discussed in the light of recent findings on the negative impact of diatoms on copepod reproductive potential.     

EGG PRODUCTION AND HATCHING SUCCESS IN NORTH ADRIATIC SEA POPULATIONS OF THE COPEPOD Acartia clausi

Egg production, fecal pellet production and hatching success are reported for Acartia clausi females sampled during three cruises in February 1997, 1998 and June 1997 at 20-24 stations along 4 transects in the North Adriatic Sea. Dramatically low hatching rates were recorded during both diatom bloom events in February as opposed to much higher rates during post-bloom conditions in June, even though A. clausi productivity during the bloom was apparently high. These results are discussed in the light of recent findings on the negative impact of diatoms on copepod reproductive potential.     

Critical thermal maxima and upper lethal temperatures for the calanoid copepods Acartia tonsa and A. clausi

Determination of the critical thermal maxima and upper lethal temperatures for Acartia tonsa Dana and A. clausi Giesbrecht revealed the relative temperature tolerance of the two species. A. tonsa exhibited a wider range of acclimation temperatures compared to A. clausi. The response of A. tonsa is related to its ubiquity in Atlantic Ocean estuaries (temperate and tropical), and its capacity to live in temperate waters throughout most of the year. Differences between upper lethal temperatures and critical thermal maxima were very narrow for A. tonsa from tropical and subtropical areas. This suggests that, in warm climates, this copepod lives close to its upper lethal temperature limits.Based in part on a dissertation submitted in partial fulfillment for the Ph.D. degree at the University of Rhode Island.     

Diel changes in gut-cell morphology and digestive activity of the marine copepod Acartia tonsa

Morphometric measurements of the gut cells of the marine calanoid copepod Acartia tonsa were made over a 24 h diel cycle to test the hypothesis that feeding rhythms in this copepod are limited by the cycles of blishter-like cells (B-cells) in the gut. Copepods were collected from Long Island Sound, New York, USA, in June 1991. Experimental treatments included low (1 g chlorophyll al^-1) and high (10 g chlorophyll al^-1) food concentrations. Copepods were fixed and embedded in Epon at six time intervals over the 24 h period, and semi-thin sections through the length of Midgut Zone II were analyzed for gut-cell area and vacuole area, with area measurements integrated to yield estimates of gut-cell volume and vacuole volume. Concurrent measurements of gut fluorescence, a measure of gut fullness, and digestive enzyme activities also were made. A diel cycle in gut fluorescence was observed, most notably in the low-food treatment, which exhibited a 3-fold increase in gut pigments. There was little consistent change in digestive enzyme activities over this time span. Gaps in the gut wall indicative of spent B-cells were observed in 17 of 39 copepods, with no trend over time or food treatment. Generally, only a single gap was seen in any one copepod. B-cells were vacuolated throughout the 24 h period. Morphometric analyses revealed a correlation between gut fluorescence and both maximum vacuole area and vacuole volume, as well as percent vacuolated cell volume, in the low-food treatment. The high-food treatment, which had a relatively smaller increase in gut fluorescence (1.5-fold) during the night cycle, showed no significant increase in vacuole size. B-cells appear to have a life-cycle greater than the diel feeding period, and while B-cell vacuoles respond to the ingested food during the diel cycle, production of B-cells does not appear to be a limiting factor in A. tonsa's feeding cycle.     

Carbon requirements of a population of the estuarine copepod Eurytemora affinis

The production of algae in the Patuxent River estuary (Maryland, USA) was much less than the carbon requirements of a population of the calanoid copepod Eurytemora affinis. Detrital carbon was present in quantities that suggest turnover times of 8 to 83 days for this pool, assuming that all algal production was consumed by E. affinis. We hypothesize that E. affinis must consume detritus to meet a part of its requirements for energy.     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Role of the free amino acid pool of the copepod Acartia tonsa in adjustment to salinity change

The free amino acid pool of the calanoid copepod Acartia tonsa was reduced in proportion to the decrease in external salinity within 24 h, with a corresponding increase in ammonia excretion and a transient rise in oxygen consumption. The free amino acid pool was not increased in response to increased salinity. Catabolism of free amino acids is important in the reduction of cellular osmotic pressure in reduced salinities. Antagonistic demands of osmotic preservation and nutritional metabolism on the free amino acid pool may limit the production of the species in waters of higher salinity.     

Impact of epibenthic predation on estuarine intertidal harpacticoid copepod populations

The extent to which energy is transferred directly from benthic meiofauna to epibenthic predators was investigated on an intertidal sand-flat in the Exe estuary, southwest England, during 1981–1982 and compared with data obtained from an intertidal mud-flat in the Lyhner estuary, also in south-west England, between 1978 and 1981. Two species of flatfish (Pleuronectes platessa L. and Platichthys flesus L.), two species of goby [Pomatoschistus microps (Krøyer) and P. minutus (Pallas)], brown shrimp (Crangon crangon L.) and shore crabs (Carcinus maenas L.) are the most common epibenthic predators feeding on the benthic invertebrates in these locatites. Harpacticoid copepods are the only component of the meiofauna to form a significant part of the diet of early juvenile stages of these predators, particularly the invertebrates. Harpacticoids are a more important source of food for predators feeding over the sand-flat than for those feeding on the mud-flat because in the sand-flat alternative prey of suitable size, such as small annelids, are absent. Moreover, the impact of predation on the mud-flat is spread over the whole harpacticoid species spectrum whereas on the sandflat it is confined almost entirely to a single species, Asellopsis intermedia (T. Scott). Flatfish, gobies and shrimp consume daily an estimated 0.01 to 0.1% of the standing stock of A. intermedia and account for between 12 and 22% of the observed reduction in the population of this species between July and October. Therefore, only a very small proportion of total meiofauna biomass is transferred directly to higher trophic levels.     

Evaluation of the effect of antimicrobials in marine cultures,using the copepod Acartia tonsa as a bioindicator

Cultures of the copepod Acartia tonsa are used both in aquaculture and ecotoxicology studies. However, the cultivation of these crustaceans at high densities results in the proliferation of microorganisms that can affect the organisms of interest, leading to illness or death. Antimicrobials inhibit microbial growth and may favour the cultivated species, aiding the development of ecological studies. This study investigated the potential of antimicrobials (antibiotic + antifungal) to inhibit bacteria and fungi when applied to marine zooplankton cultures, using the copepod A. tonsa as a bioindicator of acute toxicity. Treatment with 0.025?g?L^?1 of penicillin G potassium + 0.08?g?L^?1 of streptomycin sulphate + 0.04?g?L^?1 of neomycin sulphate + 0.005?g?L^?1 of nystatin resulted in 95% bacterial inhibition (after 12?h of exposure); however, after this time, the inhibitory effect was lost. The antimicrobial combination tested in this study prevented colonisation by fungi until 168?h after exposure, without causing acute toxicity to A. tonsa. Thus, it has potential for use in marine cultures of less sensitive organisms.     

Feeding of the copepod Acartia tonsa on the diatom Nitzschia closterium and brown algae (Fucus vesiculosus) detritus

Integumental sensilla were examined in 39 species of meso- and bathypelagic shrimps and 6 species of epibenthic shrimps. A tuft organ, consisting of a group of openended, tubular setae and so probably functioning as a chemosensor, is described. A single tuft with ancillary smaller tufts occur on the dorsal median surface of the fourth abdominal segment and two pairs of tufts occur on the proximal dorsal region of the telson of all oplophorid and pandalid shrimps examined. An additional tuft is present on the fifth abdominal segment of the rare bathypelagic Physetocaris microphthalma Chace. No tuft organs were identified in Processa canaliculata Leach, suggesting that they may not occur in all families of caridean shrimps. Acanthephyrid, systellaspid and pandalid shrimps have the integument completely covered by scales. These are extremely delicate and are normally removed during the process of catching the shrimp through abrasion of the integument by the net. The scales are peculiarly orientated, pointing anteriorly in the anterior half and posteriorly in the posterior half of the body. They probably have a sensory function as distance receptors, monitoring water currents and disturbances in the environment surrounding the shrimp.     

Analysis of the biological cycle of Acartia clausi (Copepoda) in a meso-oligotrophic coastal area of the eastern Mediterranean Sea using time-series analysis

We evaluated the duration of Copepodite Stages C1 to C6, the biological cycle and the number of annual generations of the planktonic copepod Acartia clausi in a meso-oligotrophic area of the eastern Mediterranean Sea (Saronikos Gulf, Greece). The results were based on 95 zooplankton samples collected during the period November 1988 through June 1990, at intervals of 1, 2, 7 and 15 d, the sampling intervals being dependent on the abundance of A. clausi. Time-series analysis (cross-correlation) of fluctuations in the comparative abundance (percentages) of the copepodite stages present was used to determine the duration of the development stages and generation length. This methodology could significantly contribute to the identification of cohorts, and hence to the estimation of stage duration, from field data for a given copepod species. The development of A. clausi stages was not isochronal; duration of the first copepodite stage was shorter than that of the last three stages. The mean generation length estimated (28.6 d) is among the highest recorded in the literature for A. clausi at the range of temperatures prevailing in the area (13 to 25°C). Throughout the year there were four or five generations. The possible limiting role of food availability on the duration of each stage and hence on generation length is also discussed.     

Diet and egg production of the copepod Acartia tonsa in Florida Bay. II. Role of the nutritional environment

 As part of an ongoing study of changes in the trophic pathways of Florida Bay's pelagic ecosystem, the nutritional environment (seston protein, lipid and carbohydrate levels), diet (taxon-specific microplankton ingestion rates) and egg production rate of the important planktonic copepod Acartia tonsa were measured off Rankin and Duck Keys in July and September 1997 and in January, March and May 1998. Rankin Key has been the site of extensive sea grass mortality and persistent ultraplankton blooms since 1987. Duck Key has experienced neither of these perturbations. Protist (auto-plus heterotroph) biomass was approximately twice as high off Rankin as off Duck Key. Diatoms, dinoflagellates and heterotrophic protists dominated the food environment off Rankin Key, while cells <5 μm diam often predominated off Duck Key. Protein and carbohydrate concentrations were higher off Rankin Key than Duck Key, while average lipid levels were usually low at both stations. Ingestion rates at both stations frequently approached temperature- and food-dependent maxima for the species, exceeding 100% of estimated body C d⁻¹ on 3 of 5 occasions off Rankin Key. Egg production rates, however, were consistently low (Rankin: 3 to 16 eggs copepod⁻¹ d⁻¹; Duck: 1 to 12 eggs copepod⁻¹ d⁻¹), and gross egg production efficiencies (100% × egg production C/ingested C) averaged <10%. At Duck Key, egg production rate varied with temperature and food concentration, while off Rankin Key, egg production was strongly correlated with seston protein content. The efficiency with which lipids (which were scarce in the seston) were transferred from the diet to the eggs increased exponentially with decreasing seston lipid content. Egg production efficiencies based on protein, however, were independent of seston protein content and never exceeded 10%. Received: 23 December 1998 / Accepted: 23 March 2000