The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

Egg production and hatching success in the calanoid copepods Calanus helgolandicus and Calanus finmarchicus in the North Sea from March to September 2001

Spatial and seasonal egg production rates (E_r) and egg hatchingsuccess in the copepods Calanus finmarchicus and Calanus helgolandicuswere measured in the North Sea from March to September. Foodavailability was monitored by chlorophyll and protist concentrationsand three size fractions of seston fatty acids. Seasonal andspatial distribution and production differed between the species.Calanus finmarchicus was found only offshore of the 50-m isobath,with decreasing E_r (37–28 eggs female^–1 day^–1)from March to July. Calanus helgolandicus had two abundancepeaks, in spring and autumn, with a low in May during whichtime the highest E_r were observed (38 eggs female^–1 day^–1).At other times, E_r in C. helgolandicus remained lower than inC. finmarchicus (     

Egg production and growth rates of Calanus euxinus (Copepoda) in the Black Sea

Egg production rates (EPRs) of Calanus euxinus were measuredin the Black Sea during October 2000 and May 2001. EPRs weregenerally low, on average 1.7 eggs female^–1 day^–1in October 2000 and 3.9 eggs female^–1 day^–1 in May2001. The relationships between EPRs and gonad maturity, depth-integratedchlorophyll a (Chl a) and mean surface layer temperature wereexamined. The EPRs were not related to depth-integrated Chla, but were negatively correlated with temperature. EPRs werestrongly related to the proportion of mature females. Growthrates of C. euxinus were derived from the EPRs. The mean growthrate was 0.011 day^–1 in October 2000 and 0.03 day^–1in May 2001. Growth rates were not significantly correlatedwith Chl a concentrations, but were negatively related to femaleweight and temperature.     

The relative importance of egg production rate, hatching success, hatching duration and egg sinking in population recruitment of two species of marine copepods

The disappearance of spawned copepod eggs can, at times, approach100% day^–1 and may be a bottleneck to population recruitmentof marine copepods. We examined the egg production rate andegg hatching success of Centropages hamatus and Temora longicomis(Copepoda: Calanoida) on natural diets, and the role of delayedhatching combined with high sinking rates in removing theireggs from the water column. Cumulative hatching success withinIS days was consistently high from March to June 1996:     

Seasonal variation of reproduction rates and body size of Calanus sinicus in the Southern Yellow Sea, China

Annual variations of egg production rate (EPR) and clutch sizeof Calanus sinicus, as well as body size of females (prosomelength and dry weight), were investigated at a series of stationsin the Southern Yellow Sea by onboard incubation. Calanus sinicuswas spawning in all the 11 cruises investigated, and the annualvariation of EPR was bimodal. Monthly average EPR was highestfrom May to July, respectively, 5.97, 5.36 and 6.30 eggs female^–1d^–1, then decreased dramatically to only 1.37 eggs female^–1d^–1 in August and attained the lowest 1.07 eggs female^–1d^–1 in October. In November, average EPR increased againto 4.31 eggs female^–1 d^–1. Seasonal variation ofclutch size was similar to EPR, except that it decreased graduallyafter August rather than dramatically as did EPR. Prosome lengthof females was maximum in May and minimum in October, but dryweight was highest in November. Monthly average EPR correlatedbetter with prosome length than dry weight, while clutch sizewas rather determined by dry weight of females. It is suggestedthat egg production of C. sinicus was active during two discontinuousperiods when both surface and bottom temperature fell into itsfavorite range (i.e. 10–23°C), and different reproductivestrategies were adopted in these two reproductive peaks: otherthan the highest EPR, longer prosome length was also achievedby C. sinicus from May to July, while females in November developedshorter bodies but accumulated more energy for reproduction.     

Distribution and species-specific egg production of Pseudocalanus in the Gulf of Alaska

Pseudocalanus species are important contributors to the secondaryproduction of the northern hemisphere mid- to high-latitudeoceans. In the coastal Gulf of Alaska, Pseudocalanus are presentyear round and are represented by three species. In 2001, Pseudocalanusmimus was the dominant Pseudocalanus species on the shelf duringspring and summer, comprising 30–100% of the total, whilePseudocalanus newmani dominated in Prince William Sound (10–90%).Pseudocalanus minutus were only abundant in Prince William Soundduring early spring. Egg production (by number and volume) wasa function of female prosome length and decreased from springto summer; however, significant variability was attributableto regional influences that were independent of size. For thesame sized female, P. newmani produced more eggs per clutchthan P. mimus. Pseudocalanus mimus, however, tended to havea larger mean egg size than P. newmani. Consequently, clutchvolumes of the two species were indistinguishable. Pseudocalanusegg production rates (EPRs) (eggs female^–1 day^–1)were lower in July and August (ca. 2–4) than April andMay (ca. 1–9), but total egg production by the population(eggs day^–1) was nearly equivalent for the two time periodsdue to higher female concentrations in summer.     

Zooplankton feeding ecology: does a diet of Phaeocystis support good copepod grazing, survival, egg production and egg hatching success?

The bloom-forming alga Phaeocystis is ingested by a varietyof zooplankton grazers, but is thought to be a poor source offood. We examined copepod grazing on solitary Phaeocystis cellsby adult females of Temora stylifera, and survival, fecal pelletproduction, egg production and egg hatching success in Calanushelgolandicus and T. stylifera over periods of 15 consecutivedays. Phaeocystis cell concentrations were high (1.2–3.6x 10⁴ cells ml^–1 for C. helgolandicus and 2.5–7.9x 10⁴ cells ml^–1 for T. stylifera), but within the rangeof maxima recorded for natural blooms. Both copepods survivedwell and continuously produced fecal pellets (indicating continuousgrazing) on a diet of Phaeocystis. However, egg production ratesfor both copepods were low, even though hatching success ofthe few eggs produced was high. Clearance rates for T. styliferawere higher than for most previous measurements of other copepodsfeeding on Phaeocystis solitary cells at lower cell concentrations.We conclude that even though copepods feed well upon Phaeocystis,resulting poor fecundity on this diet may inhibit copepod populationincreases during blooms, thereby contributing to the perpetuationof blooms. However, the high egg hatching success on this dietargues against Phaeocystis containing chemical compounds thatact as mitotic inhibitors reducing copepod egg viability, suchas those found in some other phytoplankters.     

Pyrosoma atlanticum (Tunicata, Thaliacea): grazing impact on phytoplankton standing stock and role in organic carbon flux

Pyrosomas are the large group of pelagic tunicates whose trophicrole in pelagic communities has not yet been sufficiently studied.We ran across a local area of high concentration of the mostwidespread and commonest species of pyrosomas, Pyrosoma atlanticum,450 miles off the Congo river mouth. The following was estimated:gut pigment content, defecation rate, organic carbon and pigmentcontent of fecal pellets, and sinking rate. Based on these dataand the measured number of pyrosomas colonies the grazing impacton phytoplankton and the fecal pellet flux were calculated.During the night swarms of 50–65 mm P.atlanticum removed53% of phytoplankton standing stock in the 0–10 m layer;sparsely distributed pyrosomas consumed only 4%. The grazingimpact in the 0–50 m layer was only 12.5 and <1% respectively.The fecal pellet flux resulting from nocturnal feeding of P.atlanticumwhile swarming made up 1.4–1.6 x 10⁶ pellets m^–210 h^–1 or 305–1035 mg C m^–2 10 h^–1 and1.4 x 10⁵ pellets m^–2 10 h^–1 or 87.4 mg C m^–210 h^–1 while non-swarming. Incubation experiments showedthe rapid degradation of fecal pellets at 23°C: the lossof pigment and carbon content was {small tilde}60–70%after 45 h. We believe that given the sinking rate of 70 m day^–1the main part of fecal material does not leave the upper watercolumn and is retained in the trophic web of the epipelagiclayer.     

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.     

Relationships between fecal pellet production and feeding in the calanoid copepod Boeckella

The number, size (mm³ pellet ^–1) and total volume of fecalpellets produced (mm³ h^–1) by Boeckella titicacae increasedwith concentration of Chlorella and natural seston in threedifferent lakewaters from Bahia de Puno and Lago Grande of LakeTiticaca and nearby Laguna Arapa. The three lakewaters differedin the size and number of fecal pellets produced at seston concentrations>0.5 mg dry wt l^–1. However, the total volume of fecalpellets egested varied little. Large-sized food (>10 µg)resulted in larger pellets but smaller total fecal production.Food concentration had parallel effects on the production offeces and on feeding rates. Both fecal production and feedingincreased linearly with food concentration with saturation between5 and 7.5 x 10⁵ Chlorella ml^–1 (3.7–5.6 mg dry wtl^–1). Varied responses in fecal production when fed differentsize fractions of lake seston suggest Boeckella has a complexand flexible feeding behavior. Feeding preferences of B.titicacaeand B.occidentalis (Ivlev index) for two seston size fractionsare generally reflected by fecal production.     

Seasonal and fine-scale spatial variations in egg production and triacylglycerol content of the copepod Acartia tonsa in a river-dominated estuary and its coastal plume

We measured egg production rates of the estuarine calanoid copepodAcartia tonsa in Mobile Bay, an estuary in the northern Gulfof Mexico. Two stations were sampled approximately monthly,one at the mouth of the bay and the other just beyond the mouthin the salinity front between bay and coastal waters. Over thewhole year, temperature was the most important environmentalvariable controlling egg production. Rates increased with temperatureup to 30°C and 140 eggs female^–1 day^–1. We foundno evidence of food limitation. There was no correlation betweenegg production and phytoplankton abundance, nor increased eggproduction in response to supplements of phytoplankton addedto natural food, suggesting that non-phytoplankton food wasimportant in the diet. At the highest egg production rates,the amount of the storage lipid triacylglycerol (TAG) in adultfemales was greatly diminished, to <50 ng female^–1.This suggests that lipids in the diet can be very tightly coupledto egg production. Both egg production and TAG content of femalesshowed significant variability on spatial scales of 3–15km, especially in relation to the salinity front separat ingwater outwelling from the bay from open coastal water. For organismsthat are using copepods and their eggs as food, this variabilitywould result in a heterogeneous food environment, both in termsof the abundance of food and its nutritional content.     

In situ egg production rate of the planktonic copepod Acartia steueri in Ilkwang Bay, southeastern coast of Korea

Egg production rate (EPR) of the copepod Acartia steueri wasinvestigated by in situ incubation in Ilkwang Bay, Korea. EPRranged from 3.8 to 10.1 eggs female^–1 d^–1, and weight-specificgrowth rate decreased with increasing body weight of the adultfemale.     

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.     

Seasonal reproductive biology of Centropages tenuiremis (copepoda) in Xiamen waters, People's Republic of China

The egg production rate (EPR) and the effects of environmentalvariations on diapause induction in the copepod Centropagestenuiremis in Xiamen waters were studied in 2002 and 2003. TheEPR ranged from 7.33 ± 8.14 (on 18 February 2003) to91.08 ± 20.61 (on 6 June 2003) eggs female^–1day^–1(mean ± 95% confidence limits), and displayed a significantseasonal pattern. Before the period of maximum egg production,EPR increased with seawater temperature and chlorophyll a concentration;but after the maximum, no significant relationship was foundbetween EPR and chlorophyll a concentration, and there was anegative linear relationship between EPR and temperature. Seawatertemperature and photoperiod, respectively, played a primaryrole in the switch to diapause egg production by the femalesas shown by single-factor probit analysis. However, the differencein effect between the two factors was demonstrated by binarylogistic multivariate analysis. These results suggest that thiscopepod rapidly increases subitaneous EPR by taking full advantageof conditions in favorable seasons (winter and spring). SubitaneousEPR decreases and there is a switch to the production of diapauseeggs that survive in the bottom sediments during unfavorableseasons (summer and autumn) in Xiamen waters.     

Feeding and reproduction of Calanus finmarchicus during non-bloom conditions in the Irminger Sea

Simultaneous ingestion and egg production experiments were conductedwith female Calanus finmarchicus in April/May and July/August2002 in the Irminger Sea. Experimental animals were providedwith natural microplankton food assemblages and incubated underin situ conditions for 24 h. The quantity of food consumed wassignificantly related to the concentration of prey cells, withtotal daily ingestion rates ranging from 0.6 to 8.1 µgof carbon female^–1 day^–1, corresponding to carbon-specificrates of 0.6–4.7% day^–1. Egg production rates (EPRs)remained relatively low (0.3–11 eggs female^–1 day^–1)during both periods of investigation and were not influencedby food availability. The data were used to construct energeticbudgets in which the microplankton carbon ingested, includingciliates, was compared with the carbon utilized for egg productionand respiration. These budgets showed that ingestion alone couldnot provide the necessary carbon to sustain the observed demandsfor growth and metabolism. Although ciliates constituted >80%of the total material ingested at times, they were not sufficientto provide the metabolic shortfall. Indeed, the females weretypically lacking 5 µg of carbon each day, 5% of theircarbon biomass. Our study results highlight the possible importanceof internal reserves in sustaining reproduction in C. finmarchicusduring periods of food scarcity.     

In situ egg production and hatching success of the marine copepod Pseudocalanus newmani in Funka Bay and adjacent waters off southwestern Hokkaido, Japan: associated to diatom bloom

. Clutch size and egg viability were examined for wild femalesof Pseudocalanus newmani collected from four different stationsinside and outside Funka Bay, southwestern Hokkaido, Japan,during pre-phytoplankton-bloom conditions in 1997, and froma single station outside the bay during the period from pre-to post-blooming seasons in 1998, to examine spatio-temporalvariability of the reproductive parameters and to detect deleteriouseffects of diatom blooms on in situ copepod reproduction. Clutchsize and hatching success in P.newmani were relatively similarat the four stations in 1997. The eggs almost always hatchedcompletely, but deformed nauplii occurred in 20–40% ofthe cases. The proportion of deformed nauplii gradually decreasedwhen the females were fed on non-diatom Pavlova sp. in the laboratory,suggesting that the occurrence of deformed nauplii was relatedto the quality of in situ food particles, including diatoms.Clutch size varied from 12 to 30 eggs female^–1 duringthe study period in 1998; more than 90% of the variation couldbe explained by the body size of the females which dependedon ambient water temperature. Hatching success also varied,from 15 to 80%, but was not related to either clutch size, femalebody size, water temperature or diatom biomass. These resultssuggest that not only egg production but also hatchability shouldbe measured routinely when estimating recruitment of the copepodinto the planktonic population, and show that neither clutchsize nor egg viability are directly affected by diatom biomass.Although we could not obtain clear evidence of in situ deleteriouseffects of diatoms on abnormal embryos and nauplii, this mightbe related to methodological problems and properties or characteristicsof prey–predator interactions. Also, deformities of copepodnauplii have not yet been observed for reasons other than feedingon diatoms.     

Zooplankton fecal pellet fluxes and vertical transport of particulate organic material in the pelagic environment

Fecal pellet fluxes were determined using a series of multireplicatetraps set at 35, 65, 150, 500, 750 and 1500 m in the northeastPacific. Fecal pellets appear to be important contributors to total carbonfluxes. In near-surface waters (35–150 m), pellet fluxesranged from 2–3 x 10⁵ pellets m^–2 day^–1. Minimumpellet fluxes were observed at 500m({small tilde}0.4x 10⁵ pelletsm^–2day^–1). In contrast, the 1500 m pellet flux valueincreased to approximately 0.8 x 10⁵ pellets m^–2 day^–1relative to the 500 m depth, and is probably the result of insitu repackaging. Analyses of pellet content suggest multiplesources of "large" particle input throughout the water column.The implications of this phenomenon are discussed in terms ofthe detrital rain and ladder of migration theories.     

Growth and ingestion rates of larval fish populations in the coastal waters of Israel

Clupeoid larvae were collected on eight cruises between February1984 and February 1985 in the coastal waters of Israel. Fromanalysis of daily growth increments of otoliths, growth ratesof the abundant clupeoids, Engraulis encrasicolus, Sardina pilchardusand Sardinella aurita were found to be 0.55 mm day^–1,0.67 mm day^–1 and 0.60 mm day^–1, respectively, duringthe first month after hatching. Ingestion rates were estimatedusing an equation from the literature relating ingestion andgrowth of larval fish. Ingestion calculated for populationsof fish larvae in pelagic waters ranged from 0 to >23 mgC m^–2 day^–1 with maximum rates observed in April.Annual ingestion by larval fish at a pelagic station near Haifawas calculated to be 2.2 g C m^–2 year^–1, 10–20%of annual primary production estimated from ¹⁴C uptake.     

Biomass, feeding and production of Noctiluca scintillans in the Seto Inland Sea, Japan

The abundance and biomass of the large heterotrophic dinoflagellateNoctiluca scintillans, together with the changes in its potentialprey items, were monitored in the Seto Inland Sea, Japan, duringsummer 1997 (17 July-11 August). Growth and grazing rates ofNscintillans fed natural plankton populations were also measuredeight and seven times, respectively, during the survey period.The abundance and biomass of N scintillans averaged over thewater column (19 m) were in the range 1–345 cells 1^–1(temporalaverage = 93 cell1^–1) and 0.1–49.6 µg C l^–1(temporalaverage = 13.8 µg C l^–1; three times higher thanthat of calanoid copepods during the same period). Noctilucascintillans populations followed the changes in phytoplankton:N.scintillans biomass was increasing during the period of diatomblooms and was at a plateau or decreasing during periods oflow chlorophyll a. The growth rates of N.scintillans (µ)were also consistent with the wax and wane of the N.scintillanspopulation: N.scintillans showed highest growth rates duringdiatom blooms. A simple relationship between µ and chlorophylla concentration was established, and the production of N.scintillanswas estimated using this relationship and the measured biomass.The estimated production averaged over the water column wasin the range >0.1–5.2 µg C l^–1 day^–1(temporalaverage = 1.4 µg C l^–1 day^–1; 64% of the productionof calanoid copepods during the same period). Diatom clearancerates by N.scintillans were in the range 0.10–0.35 mlcell^–1 day^–1, and the phytoplankton population clearanceby N.scintillans was >12% day^–1. Thus, although thefeeding pressure of N.scintillans on phytoplankton standingstock was low, N.scintillans was an important member of themesozooplank-ton in terms of biomass and production in the SetoInland Sea during summer.     

Community structures of heterotrophic bacteria of copepod fecal pellets

The total and heterotrophic bacterial microflora of gut andfecal pellets of the copepod Temora stylifera was studied inthe coastal zone of the northwestern Mediterranean basin. Digestivetracts and feces were always found to contain bacteria. Withmean values close to 10¹¹ cells ml^–1and 10¹⁰ colony-formingunits (CFU) ml^–1, both total and heterotrophic communitiesfound in fecal pellets largely exceeded the corresponding bacterialmicroflora observed in surrounding seawater (10⁵ cells ml¹ and10³ CFU ml^–). In the same way, the frequency of dividingcells and mean cell volumes increased, respectively, from 3%and 0.12µm³ in seawater to >6% and 0.24 µm³ infecal pellets. The bacterial community structure was investigatedby carrying out 29 morphological and biochemical tests on 147isolated strains. Although Pseudomonas was always the most frequentlyencountered genus, the bacterial communities found in copepodfecal pellets clearly differ from those inhabiting seawater.Vibrio species, which were not detected in seawater microflora,were always present in fecal pellet communities. The close correspondenceobserved between the potential metabolic characteristics ofthe fecal bacterial communities isolated from a subantarcticdeposit feeder (Mytilus edulis) and from zooplankton in thisMediterranean study suggests that the fecal community differentiationis an even more general feature.