Naupliar growth versus egg production in the calanoid copepod Centropages typicus

Growth rates of the copepod Centropages typicus were assessedduring May 1998 in the Alboran Sea. The goal of the work wasto compare field growth rates of juvenile (artificial cohortmethod) and adult copepods (female egg production rates) underan extensive range of natural conditions. The results showedthat adult and juvenile growth rates were similar in some stations,while in others no such relationship was apparent. In contrastto the common belief that juvenile growth may be saturated undernatural conditions whereas adults may be food-limited, in mostof our study, adult growth rates were greater than juveniles.We discuss these results in the light of food availability atthe surveyed stations. In summary, the assumption that copepodgrowth rates estimated through egg production rates of adultsare equivalent to juvenile growth is not always valid.     

Laboratory study of the spawning rate of the calanoid copepod Centropages typicus: effect of fluctuating food concentration

The spawning rate of laboratory-reared Centropages typicus fedHymenomonas elongala increases with food concentration, up toa value of {small tilde}2800 µg C (16 500 cells) ml^–1.An alternation of a low food (1000 cells ml^–1) and highfood concentration (16 500 cells ml^–1) is not favourableto egg release when its periodicity is 1 or 2 days, whereasit may be of advantage if it is longer (3–6 days). Inthe latter case, Centropages typicus will benefit best fromthe rich food diet if this coincides with (or just follows)the last moult.     

Food type induces different reproductive responses in the copepod Centropages typicus

Egg production and hatching success were determined for individualCentropages typicus fed two diatoms (Thalassiosira rotula andPhaeodactylum tricomutum) and two dinoflagellates (Prorocen-trumminimum and Gonyaulaxpolyedrd). Both reproductive responseswere strongly affected by food type. Females incubated withoutmales produced eggs with all diets, but fecundity was twiceas high with the larger T. rotula and G.polyedra cells. In contrast,hatching success was 2–3 times higher with the dinoflagellatediets. The presence of males did not enhance egg productionrates. Males also did not improve hatching success when thediet consisted of the diatom T.rotula. However, egg viabilitywas higher for couples fed the dinoflagellate G.polyedra, indicatingthat egg viability was possibly being controlled by both rematingand food type. Egg viability was artificially lowered by exposingnewly spawned eggs to high concentrations (10⁴–10⁹ pgC) of extracts from T.rotula, whereas the development of eggsproceeded normally at all concentrations of extracts from P.minimum.Blockage of egg development was not due to anoxia, but to thepresence of intracellular, deleterious chemical compounds indiatoms, suggesting that bottom-up prey control mechanisms underlieseasonal fluctuations in C.typicus populations at sea.     

Seasonal cycles in population abundances and egg production rates in the planktonic copepods Centropages typicus and Acartia clausi

Laboratory studies on the seasonal cycling of egg productionrates for Centropages typicus and Acartia clausi did not reflectseasonal changes in densities for these species at sea. Maximumegg production rates occurred from autumn to spring when populationabundances at sea were low. In late spring, an increment inpopulation numbers was followed by a drastic decline in eggdeposition which continued into summer. The percentage lossdue to mortality, calculated using egg data and total numberof copepodid stages CIII and CIV recorded at sea {small tilde}2weeks later, indicated greatest mortality (80–99%) fromeggs to copepodids during periods of high breeding intensity.These results suggest that high and low density phases in populationnumbers may be the outcome of variations in reproductive potentialand survival rates of eggs and immature stages rather than reproductivepotential per se.     

Seasonal fluctuations in fecundity and hatching success in the planktonic copepod Centropages typicus

A mismatch of {small tilde}2–3 months was observed betweenreproductive rates and population abundances in the planktoniccopepod Centropages typicus, with the highest production atrelatively low female abundance and low production at high abundance,during the course of a 2 year study in 1989 and 1990 in neriticwaters of the Gulf of Naples. During this period, egg mortalitywas at times severe, with values as high as 59% in February1989. The seasonal trend in percentage hatching success didnot match seasonal fluctuations in breeding intensity, and wasnot correlated with variations in environmental variables suchas temperature and chlorophyll a. The results of experimentsusing the fluorescent dye Hoechst 33342 showed that unhatchedeggs had been fertilized and that in most cases developmenthad proceeded to an advanced stage before death of the embryo.The causes of high egg mortality are uncertain, but here wegive evidence, for the first time, that egg mortality may notonly be due to infertility caused by failure to remate.     

The effect of reproductive condition on egg production rates in the planktonic copepod Centropages typicus

Centropages typicus females oscillated between a dark, ripestage of oogenesis and a light, unripe stage of primary oocytes.The number of eggs released after 24 h under different experimentalconditions was strongly dependent on the reproductive statusof females at the time of incubation.     

Is fecundity modified by crowding in the copepod Centropages typicus?

Experiments were conducted to test the effect of overcrowdingon egg production rates in the copepod Centropages typicus.Fertility was not significantly reduced by the presence of otherconspecifics when the volume of water was the same for femalesincubated alone or in the presence of other females and males.There was no evidence of chemical cues affecting egg productionwhen females were kept in large (1 female in 100 ml sea water[SW] or small (1 female in 25 ml SW) volumes of water, withor without partitions, and in the presence or absence of foodor males. Also, spawning rate was not reduced when females wereincubated in flow-through experimental chambers which alloweddiffusion of pheromones between females incubated alone or inhigh densities (100 ind. in 50 ml SW). Only when females wereplaced at unnaturally high densities (1 female ml^–1) wasegg production significantly reduced, probably due to physicaldisturbance and increased number of collisions rather than chemicalcommunication among individuals. The results show that chemicalcues do not affect spawning behaviour when population densitiesare close to field conditions. By contrast, overcrowding inducedby swarming or aggregation may potentially limit reproductivesuccess due to reduced feeding, and production and cannibalismof eggs.     

Laboratory studies on egg and faecal pellet production of Centropages typicus: effect of age, effect of temperature, individual variability

The rates of egg production of individually reared females ofCentropager typicus have been observed from the moulting C5-adultto death under constant conditions of food (10 000 cells ml^–1of Hymenomonas elongara) and temperature (15 and 20°C).Preliminaiy experiments showed that virgin females could produceunfecunded eggs, but that the distinctive spines of the eggsof C.typicus were only obtained after fecundation occurred.High and continuous egg production needs several matings, andthe following experiments were run with a couple. In all experiments,both males and females fed and produced faecal pellets. Thefemales show a high variability in spawning life periods, inspawning rhythms, in daily rates and, consequently, in cumulatednumbers of eggs spawned during their life. The highest dailyrates of egg production were obtained in the first half of thefemale's life, and were as high at 20°C as at 15°C.Nevertheless, the females survived longer at 15°C than at20°C, and finally produced 47.6% more eggs during theirwhole life at 15°C than at 20°C, which can be partiallydue to the difference in size (almost 10% bigger at 15°Cthan at 20°C). The production of faecal pellets occurredduring the whole of life and was more regular than egg production,with a maximum daily rate in the first part of life, and showeda decrease until death. The decline in daily egg productionwith age was parallel to the decrease in daily faecal pelletproduction. The daily production of faecal pellets was higherat 15°C than at 20°C. At 20°C, a good correlationwas observed between the total egg production per female andthe total faecal pellet production per female. The individual-basedegg production rates of our experiments have been compared withrates obtained in different conditions by several authors, andwere lower than in other published studies, probably due tothe food offered. We emphasize the difficulties of comparingresults which have been obtained for the entire life of thefemales with those based on daily egg production. The totalegg production of a female is an important parameter in regardto the population d because it fixes the maximal rate of losses(mortality plus dispersion) for which a population can continueto develop.     

Mediterranean marine copepods: basin-scale trends of the calanoid Centropages typicus

The Mediterranean Sea is located in a crossroad of mid-latitude and subtropical climatic modes that enhance contrasting environmental conditions over both latitudinal and longitudinal ranges. Here, we show that the large-scale environmental forcing is reflected in the basin scale trends of the adult population of the calanoid copepod Centropages typicus. The species is distributed over the whole Mediterranean basin, and maximal abundances were found in the north-western basin associated to oceanic fronts, and in the Adriatic Sea associated to shallow and semi enclosed waters. The peak of main abundances of C. typicus correlates with the latitudinal temperature gradient and the highest seasonal abundances occurred in spring within the 14–18°C temperature window. Such thermal cline may define the latitudinal geographic region where C. typicus seasonally dominates the >200 μm-sized spring copepod community in the Mediterranean Sea. The approach used here is generally applicable to investigate the large-scale spatial patterns of other planktonic organisms and to identify favourable environmental windows for population development.     

Observations on egg production rates and seasonal changes in the internal morphology of Mediterranean populations of Acartia clausi and Centropages typicus

Egg production rates in wild populations of Acartia clausi and Centropages typicus, sampled biweekly in the Gulf of Naples from October 1985 to July 1987, showed marked seasonal fluctuations with maximum values in early spring that proceeded the annual maxima for adult female densities in summer. A positive correlation between chlorophyll a concentrations and egg production was evident only during the early spring phytoplankton bloom. A strong diminution in egg deposition occurred later in spring and continued throughout the summer notwithstanding high chlorophyll concentrations. In winter, when population abundances for adult females were lowest, egg production rates were always higher than in summer. Differences in egg production rates coincided with pronounced morphological changes between summer and winter populations of both species. The most striking of these changes consisted, in winter, in the presence of a dark brown fluid-like mass of granular material that seemed to freely bathe the gonads. The presence of this substance only during periods of elevated egg production suggests that it may enhance egg production rates when the adult population reaches minimum annual levels. Such a mechanism of self-regulation may operate to dampen the effects of environmental variability thereby contributing to maintain a conservative structure in coastal copepod communities.     

Development, growth and egg production of Centropages abdominalis in the eastern subarctic Pacific

Centropages abdominalis is a neritic, omnivorous, temporallyabundant copepod present throughout the subarctic Pacific andits marginal seas. The two main objectives of this study wereto determine how temperature influences the development of C.abdominalis and whether growth rates of in situ populationsmay be limited by available food. At 6.9°C, median developmenttime from eggs laid to 50% adults was 42 days and the averageweight-specific somatic growth rate was 0.17 day^–1. At4.6°C, median development time to adult was 59 days (projected)and growth rate averaged 0.08 day^–1, suggesting that 4.6°Cmay be approaching the lower temperature for development andgrowth in this species. The functional relationship betweendevelopment time and temperature was established over the temperaturerange in which this species occurs. The in situ adult growthrates between 10 and 13°C averaged 0.14 day^–1 andwere generally lower than the laboratory-reared juvenile growthrates, which may indicate that adult C. abdominalis are foodlimited in the field during summer and autumn.     

Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus

Diel variations in vertical distribution, gut pigment content,ammonium excretion and egg production were investigated foradult females of Acartia erythraea and A.pacifica in the verticallymixed Inland Sea of Japan and Centropages furcatus in the stratified,neritic Gulf of Mexico. Gut pigment content and egg productionrate were maximal at night and ammonium excretion was maximalduring the daytime. Neither A.erythraea nor A.pacifica adultfemales showed an apparent diel migration, but the former werehighly concentrated in the surface layer during the afternoon.In contrast, C.furcatus adult females showed a clear diel migration,residing immediately above the bottom during the daytime andbeing concentrated between 10 and 25 m depth during the nighttime.Individual-based data on gut content and excretion and egg productionrates were combined with vertical-distribution data to calculatepopulation values. In the Inland Sea of Japan, the resultantpattern for Acartia spp. reflected the diel variation in physiologicalrates and even distribution of adult females, except for theafternoon, surface aggregation of A.erythraea. In the Gulf ofMexico, the pattern for C.furcatus reflected largely the dielvariation in each rate process and the heterogeneous distributionof adult females in the water column. Elevated nocturnal feedingactivity of these copepods may be due to an endogenous rhythm.The daytime maximum in ammonium excretion and night-time maximumin egg production rate indicated approximate half-day and daytime lags, respectively, after the intake of food until itsconversion into dissolved excreta and released eggs.     

Short communication. Food availability and diel feeding rhythms in the marine copepods Acartia grani and Centropages typicus

The effects of food availability on the daily feeding pattern of planktonic copepods was studied for two species: Acartia grani (cultured and wild specimens) and Centropages typicus (wild specimens). Both species showed clear diel feeding rhythms at high food concentrations, with significantly higher clearance rates during night hours. However, at limiting food conditions, while C.typicus maintained its daily feeding pattern, A.grani fed continuously, with similar day and night clearance rates. These results could be explained in relation to differences in predation risks and probability of experiencing starvation episodes in their respective habitats, as well as their capacity for dampening food fluctuations. On the other hand, the maintenance (although with less intensity) of the diel feeding rhythm in A.grani, when cultured in the laboratory for long periods of time (>10 generations) and in the absence of predators, suggests the importance of the endogenous control of copepod feeding patterns.      

Estimated, in situ, rates of egg production for the copepod centropages typicus (Krøyer) in the new york bight

Centropages typicus (Krøyer) and Pseudocalanus sp. are the two predominant copepods in the Continental Shelf waters immediately south of Long Island, New York. The estimated, in situ, rate of egg production for Centropages typicus ranged from 5 to 230 eggs female^?1 day^?1 during this study. Variability was partly attributed to seasonal variation in water temperature and partly to variations in the physiological condition of individual females. It could not be shown that the ability to produce eggs varied seasonally due to factors related to food. Egg production by Pseudocalamus sp. is probably from 1–10 eggs female^?1 day^?1. The two reproductive behaviours result in average abundances of adults that are approximately the same, although the peaks are at different times of the year, indicating that Centropages typicus has a higher mortality between the egg and adult stages than Pseudocalanus sp.     

Copepod egg production, moulting and growth rates, and secondary production, in the Skagerrak in August 1988

Measurements of hydrography, chlorophyll, moulting rates ofjuvenile copepods and egg production rates of adult female copepodswere made at eight stations along a transect across the Skagerrak.The goals of the study were to determine (i) if there were correlationsbetween spatial variations in hydrography, phytoplankton andcopepod production rates, (ii) if copepod egg production rateswere correlated with juvenile growth rates, and (iii) if therewas evidence of food-niche separation among co-occumng femalecopepods The 200 km wide Skagerrak had a stratified water columnin the center and a mixed water column along the margins. Suchspatial variations should lead to a dominance of small phytoplanktoncells in the center and large cells along the margins; however,during our study blooms of Gyrodinium aureolum and Ceratium(three species) masked any locally driven differences in cellsize: 50% of chla was >11 µm, 5% in the 11–50µm fraction and 45% <50 µm. averaged for allstations. Chlorophyll ranged from 0.2 to 2.5 µg l^–1at most depths and stations. Specific growth rates of copepodsaveraged 0.10 day^–1 for adult females and 0.27 day^–1for juveniles The latter is similar to maximum rates known fromlaboratory studies, thus were probably not food-limited. Eggproduction rates were food-limited with the degree of limitationvarying among species: 75% of maximum for Centropages typicus, 50% for Calanus finmarchicus, 30% for Paracalanus parvus and 15% for Acartia longiremis and Temora longicornis. Thedegree of limitation was unrelated to female body size suggestingfood-niche separation among adults. Copepod production, summedover all species, ranged from 3 to 8 mg carbon m^–3day^–1and averaged 4.6 mg carbon m^–1 day^–1. Egg productionaccounted for 25% of the total.     

Temperature impact on reproduction and development of congener copepod populations

The goal of this study was to relate the temperature response of all developmental stages and reproductive biology of two congener copepod pairs inhabiting different biogeographic regions to their geographic distribution patterns. Survival of adult females and egg production, embryonic development and hatching success of the genera Centropages and Temora from two stations, in the North Sea and the Mediterranean, were studied in laboratory experiments in a temperature range from 2 to 35 °C. Postembryonic development was determined from cohorts raised at temperatures between 10 and 20 °C with surplus food. Tolerance limits and optima of female survival, reproduction and development distinguished the northern species Centropages hamatus and Temora longicornis from the southern T. stylifera, while C. typicus, which is found in both regions, was intermediate. Thus, thermal preferences could in part explain distribution patterns of these species. While C. hamatus and the two Temora species showed distinct temperature ranges, C. typicus was able to tolerate different temperature conditions, resulting in its wide distribution range from the subarctic to the tropics. However, the thermal range of a species did not necessarily correlate with the optimal temperatures in the experiments. Optima of egg production and stage development were surprisingly low in T. stylifera, which has a mere southern distribution.     

Persistent spatial and temporal abundance patterns for the late-stage copepodites of Centropages typicus (Copepoda: Calanoida) in the US Northeast continental shelf ecosystem

The annual cycle of abundance and the monthly distributions of the copepod Centropages typicus are described for US Northeast Continental Shelf waters from samples collected on broadscale plankton surveys during 1977-87. High numbers of the copepod were captured throughout the region during the autumn months in weak south-north and onshore-offshore abundance gradients. The highest individual station densities were found near the mouth of the major estuaries and the heaviest broad-scale concentrations were usually located where bottom depth ranged from 20 to 39 m. Numbers declined throughout the ecosystem after winter arrived, less so in the southern half of the region where C.typicus abundance remained high year round in nearshore and midshelf waters between New York City and Chesapeake Bay. The copepod's abundance fell to much lower levels further north in the Georges Bank and Gulf of Maine subareas, and disappeared entirely from shelf waters in the northernmost offshore region until summer. Interannual abundance variability was substantial, but no long-term trend was detected. Analyses of samples collected from 1988 to 1996 on Georges Bank during early autumn indicate that abundance levels of C.typicus have been high here in the 1990s, completely recovered from low density values measured there in 1986 and 1987. Temperature and food availability were found to be the key factors that determine the copepod's distribution and annual abundance cycle. Mean abundance was high throughout the ecosystem where surface temperature was >9C and in regions where annual mean chlorophyll levels exceeded 1 mg m^-3. The copepod's abundance appeared to be independent from variation in water column salinity.      

Pigment gut contents of copepods and deep phytoplankton maximum in the Western Mediterranean

The feeding activity of Centropages typicus in the Western Mediterraneanwas evaluated during the summer stratification period, whena deep phytoplankton and chlorophyll maximum (DCM) develops.The lack of significant day-night differences in gut contentssuggests continuous phytoplankton ingestion, despite nictemeralmigration of this species. Copepods captured in the DCM hadgreater phytoplankton gut contents, both in nocturnal and diurnalsamples. The observed pattern of phytoplankton ingestion andvertical migration implies copepod-mediated upward transportof nutrients, which might contribute to the maintenance of primaryproduction in the photic layers.     

Seasonal Cycles of Egg Production of Two Planktonic Copepods, Centropages typicus and Temora stylifera, in the North-western Mediterranean Sea

Reproduction of the dominant copepods Centropages typicus andTemora stylifera was studied at a permanent station in the LigurianSea (north-western Mediterranean). Seasonal patterns of eggproduction, clutch size, egg size and female prosome lengthwere followed from January 1998 to December 1999. Female carboncontent and weight-specific egg production were compared inautumn 1998 and spring 1999. Reproductive patterns of C. typicusand T. stylifera were very similar, indicating that reproductionwas affected by the same environmental factors. Reproductiveactivity was highest in autumn in both species and years. Asecond peak of egg production was observed in early summer,which was less intense in 1999 after a bloom of salps. Egg productionrates reached maximal values of 33.5 and 33.3 eggs female^–1day^–1 and annual means of 10.8 and 11.7 eggs female^–1day^–1 in Centropages and Temora, respectively. Maximalweight-specific egg production was 0.21 day^–1 in bothspecies in November 1998, when female carbon contents were 6.7(C. typicus) and 12.0 µg (T. stylifera). No statisticalrelationship between egg production and food availability ortemperature was detected. Reproductive activity did not reflectthe seasonal abundance patterns, with C. typicus dominatingin spring and T. stylifera in autumn.     

Copepod grazing on a phytoplankton community containing the toxic dinoflagellate Dinophysis acuminata

Copepod grazing on the toxic dinofiagellate Dinophysis acuminatafrom the west coast of France (La Rochelle) was studied witha concentrated (40–70 µm) phytoplankton assemblagedominated by Leptocylindrus danicus, D.acuminata, Ceratium fususand Ceratium furca. Copepod nauplii were also present. Threeto five copepods/copepodites (Acartia clausi, Isias clavipesand Centropages typicus) were incubated together with the phytoplankton.Dinophysis acuminata was grazed upon by all copepod species.However, to some extent, I.clavipes and C.typicus avoided itas food. Dinophysis acuminata cells represented for them only5–10% of total ingested carbon during the first 24 h,and almost all individuals survived and thrived well. In contrast,A.clausi did not avoid D.acuminata. which represented 30% ofingested carbon in 1 day. Acartia clausi then had a lower survivalthan the two other copepod species. However, the survival ofA.clausi was high in control incubations, where a plankton communitywithout D.acuminata was used as food. It is concluded that theokadaic acid of D acuminata is potentially toxic to some grazers,and/or might function as an allelopathic grazer repellent.