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.     

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.     

A 5-year study of the influence of the northeast and southwest monsoons on copepod assemblages in the boundary coastal waters between the East China Sea and the Taiwan Strait

Danshuei River in the northwestern tip of Taiwan dischargesinto the boundary coastal waters between the East China Seaand the Taiwan Strait. We conducted a 5-year study between October1998 and September 2003 to assess the influence of the northeast(NE) and the southwest (SW) monsoons on copepod assemblagesin the area. A total of 110 copepod species were identified.Temora turbinata, Paracalanus aculeatus, Acrocalanus gibber,Parvocalanus crassirostris and Oithona rigida comprised 80%of the copepod assemblages that consisted of coastal speciesfrom the East China Sea and species associated with the KuroshioBranch Current. The effect of the NE monsoon was observed duringa short period of each winter when species such as Calanus sinicuswere transported into the study area by the China Coastal Current.In summer, species such as A. gibber, Acrocalanus gracilis andCanthocalanus pauper may be transported into the study areafrom Southern Taiwan by the combined effect of the SW monsoonand the Kuroshio Current. Influence of the NE monsoon on thecopepod assemblages in terms of introduction of species fromthe Yellow Sea and the East China Sea during winter was observedevery year but was unlikely to represent an important carbonsource into the region due to its short-term nature. Local speciessuch as T. turbinata, P. crassirostris and O. rigida form amajor component of the copepod assemblage. The high copepoddiversity in the area was caused by the year-round presenceof many local species and the aggregation of different speciesfrom subtropical, tropical, and temperate water masses. Monsoon-drivenwater currents and the Kuroshio Branch Current appeared to maskthe effect of river discharge in the region.     

The contribution of ammonia excreted by zooplankton to phytoplankton production in Narragansett Bay

Ammonia excreted by mixed zooplankton populations over an annual(1972–1973) cycle in Narragansett Bay varied from 0.04to 3.21 µg at NH₃-N dry wt^–1 day^–1, exclusiveof two exceptional rates measured one year apart: 11.74 and18.39 µg at NH³-N mg dry wt^–1 day^–1. Grossphytoplankton production integrated over the year (1972–1973)averaged 151 mg C m^–3 day^–1 for an 8 m water column;peaks of 332 and 905 mg C m^–3 day^–1 occurred duringthe winter-spring and summer blooms, respectively. Excretedammonia, integrated seasonally and annually, contributed only0.2% and 4.9% of the nitrogen required for observed gross productionduring the winter-spring and summer blooms, respectively, and4.4% annually. However, excreted ammonia may be an importantsource of the nitrogen required by Skeletonema costatum, thedominant diatom in Narragansett Bay, during the post-bloom periodwhen 186% of the nitrogen required for its net production wasmet by ammonia excretion. A combination of zooplankton ammoniaexcretion and benthic ammonia flux contributed 22% of the nitrogenrequired for the annual gross production (440 g C m^–2)while 51% of the nitrogen required for the net production ofSkeletonema was accounted for by regenerated nitrogen. ¹This research was supported by NSF grant GA 31319X awardedto Dr.T.J.Smayda.     

Primary and bacterial production compared to growth and food requirements of Daphnia longispina in Lake Kvernavatnet, west Norway

Primary production, and bacterial production as measured byincorporation of [³H-methyl]thymidineinto ice cold TCA insolublematerial were investigated during 1984 in Lake Kvernavatnet,west Norway. Primary production averaged 222 mg C m^–2day^–1 and bacterial production averaged 163 mg C m^–2day^–1. The bacterial production in the euphotic pelagiczonecontributed -60% of the total pelagic bacterial production.The zooplankton was dominated byDaphnia longispina. From growthexperiments with animals fed only natural food in coarse filteredlake water, the population daily growth increments were calculated.The average production of D.longispina was 151 mg C m^–2day^–1 during the period investigated. The estimated primaryproduction was too low to sustain both the bacterial productionand the zooplankton food requirements. These results imply thatthe carbon cycle of the lake is dependent on the supply of allochtonousmaterial, or that the current methods for measuring productionrates in aquatic environments are systematical erratic.     

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.     

Direct measurement of the production:biomass ratio of the subtropical calanoid copepod Acrocalanus inermis

I have measured the specific population growth rate or production:biomass(P:B) ratio of the copepod Acrocalanus inermis by an incubationtechnique. The principal advantage of this method is that itis site- and time-specific and can therefore be used to determinespatial and temporal patterns in P:B. The values for P:B determinedby this method ranged from 0.07 day^–1 to 0.36^–1,with a mean of 0.28^–1. P:B appeared constant for incubationsof up to 40 h, and did not change significantly with up to afour-fold increase in concentration of animals during incubation.Variability in P:B between two stations and among differentdates was small. ¹Present address: University of Melbourne, Department of Zoology,Parkvill, Victoria 3052, Australia     

The use of sea temperature in characterizing the mesoscale heterogeneity of phytoplankton in an embayment of the southern Benguela upwelling system

The role of meteorological events and hydrography in determiningchanges to the phytoplankton community was investigated in anembayment exposed to a narrow band of coastal upwelling. Dailysampling demonstrated the importance of advective processesdriven by meteorological forcing in controlling rapid shiftsin the biomass and species composition of the phytoplanktoncommunity. Samples of similar phytoplankton composition wereassociated with different stages of the upwelling cycle, asdefined by an index of biological ageing of upwelled waters.Relationships between the physical, chemical and biologicalfields were defined from time-senes measurements. The time elapsedfollowing upwelling, required for the determination of biologicalrates, was estimated from the rate of heating. A primary productionestimate of 2.14 g C m^–2 day^–1 was derived fromdetermination of the rate of nutrient depletion, whereas a phytoplanktonbiomass-nutrient consumption equation provided an estimate of3 92 g C m ^–2 day^–1. Both rates were within therange of estimates obtained from in vitro tracer methods.     

Feeding and metabolism of the siphonophore Sphaeronectes gracilis

The in situ predation rate of the siphonophore Sphaeronectesgracilis was estimated from gut content analysis of hand-collectedsiphonophores and from laboratory data on digestion rates ofprey organisms. At daytime prey densities of 0.25 copepods 1^–1,S. gracilis was estimated to consume 8.1 – 15.4 prey day^–1siphonophore^–1. From data on abundances of siphonophoresand copepods, S. gracilis was estimated to consume 2–4%of the copepods daily. In laboratory experiments, ingestionrates averaged 13.8 prey day^–1 siphonophore^–1 atprey densities of 5 copepods 1^–1 and 36.9 at 20 copeods1^–1. This was equivalent to a specific ingestion rate(for both carbon and nitrogen) of –17% day^–1 and45% day^–1, respectively, while specific ingestion in situwas only 2% day^–1. Ammonium excretion averaged 0.095 µg-atsiphonophore^–1 day^–1 at 5 prey 1^–1, and 0.162at 20 prey 1^–1. The specific respiration (carbon) andspecific excretion (nitrogen as ammonium) were calculated tobe 3% day^–1 at the lower experimental food level, and5% day^–1 at the higher food level. ¹Contribution from the Catalina Marine Science Center No. 66. ²Present address: Dept. of Biology, University of Victoria,Victoria, B.C., Canada V8W 2Y2.     

Microzooplankton grazing in a coastal embayment off Concepcion, Chile, (~36{degrees} S) during non-upwelling conditions

The impact of grazing by natural assemblages of microzooplanktonwas estimated in an upwelling area (Concepción, Chile)during the non-upwelling season in 2003 and 2004. Seawater dilutionexperiments using chlorophyll a (Chl a) as a tracer were usedto estimate daily rates of phytoplankton growth and microzooplanktongrazing. Initial Chl a concentrations ranged from 0.4 to 1.4mg Chl a m^–3 and phytoplankton prey biomass and abundancewere numerically dominated by components <20 µm. Phytoplanktongrowth and microzooplankton grazing rates were 0.19–0.25day^–1 and 0.26–0.52 day ^–1, respectively.These results suggest that microzooplankton exert a significantremoval of primary production (>100%) during the non-upwellingperiod.     

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 hatching success in the peri-Antarctic copepod Calanus simillimus

In situ rates of egg production and hatching success are reportedfor Calanus simillimus, one of the most abundant calanoid speciesin peri-Antarctic regions, during the Italian ‘Italics’cruise in the Straits of Magellan in March-April 1995. Low fecundity(8.6 eggs female^–1 day^–1) and fecal pellet production(0.8 fecal pellets female^–1 day^–1) in this periodindicate that the species was feeding very poorly. Sixty-sixof the 126 females sampled did not produce eggs and 80 of thesedid not produce any fecal pellets during the 24 h period ofincubation. Striking abnormal naupliar and embryonic developmentwas recorded in 81.8% of the eggs spawned. Aberrant eggs didnot undergo normal cleavage, and failed to develop to hatching.Deformed nauplii were asymmetrical and presented strong anatomicalanomalies in the total body length and number of swimming appendages.These results are discussed in the light of recent findingson the causes of low hatching success in copepods.     

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.     

Zooplankton grazing impact in the plume of dilution of the Gironde estuary (France) prior to the spring bloom

The Bay of Biscay is a coastal area intensively exploited forfishing and is submitted to important human actions (proximityof important industrial and agricultural areas). Thus, the understandingof the materials and energy transfers in such ecosystems isof great interest. However, investigations on zooplankton (animportant component of the marine food web) are very scarcein this area. Our study concerns the grazing impact of the zooplanktonin shelf waters in the plume of dilution of the Gironde estuaryprior to the spring bloom. Samples were taken through the photiclayer in April 1993. Our results have shown that Temora longicornisand Paracalanus parvus dominated the ‘herbivorous’planktonic community. The grazing impact of the zooplanktoncommunity on the total phytoplankton stock and on the totalprimary production was low (9–14 and 17–21% day^–1,respectively) during this period, which is in accordance withmost results in coastal areas. However, due to the small sizeof the algae (     

Reproductive response of the copepods Calanoides carinatus and Calanus agulhensis to varying periods of starvation in the southern Benguela upwelling region

Egg production by the calanoid copepods Calanoides carinatusand Calanus agulhensis fed excess Thalassiosira weissflogiiwas monitored in the laboratory following starvation periodsof 1, 3, 5, 7 and 9 days. Following short (1–3 day) periodsof starvation, egg production by C.agulhensis returned to thesatiated rate (51.1 eggs {female} day^–1) more rapidly(after 0.9–2.4 days of excess food) than that of Ca. carinatus(after 2.8–3.1 days). However, following longer (5–9day) periods of starvation, Ca. carinatus regained satiatedlevels of egg production (55.8 eggs {female}^–1 day^–1)more rapidly (after 3.1–4.0 days of excess food) thanC. agulhensis (after 3.8–5.2 days of feeding following5–7 days of starvation). Moreover, many C. agulhensisfemales did not regain normal rates of egg production after9 days of starvation. For both species, the time required foregg production to recover was proportional to the starvationperiod, although only up to 7 days for C. agulhensis, and wasthe same following 4.25 days of starvation. Previously fed Ca.carinatus terminated egg production more rapidly than C. agulhensiswhen starved. The ability of Ca. carinatus to tolerate, andrecover rapidly from, prolonged periods of starvation, combinedwith a comparatively fast development time and high rate ofegg production, provides this species with a strong competitiveadvantage over C. agulhensis in the highly pulsed food environmentof the southern Benguela upwelling region.     

Quantification of the phosphorus released by zooplankton in an oligotrophic lake (La Caldera, Spain): regulating factors and adjustment to theoretical models

Using an in situ approach, we have evaluated the phosphorusinputs from zooplankton in a high-mountain oligotrophic lake.Values of the specific gross release rate (SGRR) fluctuatedbetween 0.2 and 2.9 µg P mg^–1 dry weight h^–1,and were higher when the nauplii of Mixodiaptomus laciniatusdominated the zooplankton community. The rate of P recyclingby the zooplankton was high, reaching 1.6 µg P l^–1day^–1, and showed a highly consistent seasonal patternfrom one year to the next, with maxima in midsummer. Zooplanktonsize accounted for as much as 85% of the variance obtained inthe measurements of the specific rate of P release, while otherfactors, such as the quality or quantity of food, did not significantlyinfluence the SGRR changes. Among the models tested, only theone proposed by Peters (Int, Ver. Theor. Angew. Limnol. Verh.,19, 273–279, 1975) was useful for predictions in thissystem. The stoichiometric model of Hessen and Andersen (Arch.Hydrobiol., 35, 111–120, 1992), applied in this oligotrophicsystem, adequately predicted the phyto- and zooplankton dynamics,whereas the values of P release estimated using this model werefar higher than the excretion rates obtained experimentally.These differences were related to the type of egestion (formationof faecal pellets) of metazooplankton and to the relative importanceof the food resistance to digestion. We believe that in communitieswhere copepods constitute a substantial percentage of the zooplankton,an evaluation of the P release which is readily available [solublereactive phosphate (SRP), total dissolved phosphate (TDP)] toalgae and bacteria would not fit the predictions of generalmodels of mass balance; under these circumstances, assimilationefficiency proves to be the key parameter for predicting thereadily available P (excreted P).     

Production of Oikopleura longicauda (Tunicata: Appendicularia) in Toyama Bay, southern Japan Sea

Oikopleura longicauda occurred throughout the year in ToyamaBay, southern Japan Sea, and analysis of its size compositionand maturity revealed that reproduction was continuous overtheyear. Somatic growth production (P_g) varied with season from0.03 to 103 mg carbon (C) m^–2day^–1 (annual P_g 4.5g C m^–2), and house production (P_e) from 0.11 to 266 mgC m^–2 day^–1 (annualP_e 11.3 g C m^–2). The annualP_g/B ratio was 176. Compared with production data of some predominantzooplankton species in Toyama Bay, it is suggested that despitetheir smaller biomass, appendicularians are an important secondaryproducer.     

Production of Penilia avirostris in Kingston Harbour, Jamaica

The cladoceran Penilia avirostris is one of the more abundantand widespread members of the crustacean zooplankton in nearshoretropical and subtropical waters. Its abundance, biomass, fecundity,development rate and production were estimated in Kingston Harbour,Jamaica, during an 18 month period. Mean annual abundance ofPenilia was 1821 m^–3, while biomass (excluding eggs/embryos)was 2.87 mg ash-free dry-weight (AFDW) m^–3 (43.1 mg AFDWm^–2), accounting for 13% of the copepod community biomass.Fecundity increased with body size. There was no clear seasonalpattern of abundance, size or fecundity, nor were physical orbiological variables correlated to these variations. Developmenttime averaged 20.5 h for juveniles and 41.4 h for adult femalesduring incubations; there was no clear evidence of a diel patternto molting. Growth rate appeared to be exponential, with correspondingsomatic growth rates, averaging 0.27 day^–1 for juveniles,and 0.34 day^–1 for somatic plus reproductive growth inadult females. Annual production was estimated as 173 kJ m^–2year^–1,     

Egg production, growth and development of the cyclopoid copepod Oithona similis

Egg production, growth and development rates of Oithona similiswere measured in the laboratory as a function of food concentrationand composition. On an optimum diet, development is isochronaland growth is near exponential. The maximum juvenile growthrate at 15C (0.2 day^–1) is similar to juvenile growthin calanoid copepods. The maximum weight-specific egg productionrate (0.1 day^–1), on the other hand, is substantiallyless than in free-spawning calanoids, but similar to that inegg-carrying calanoids. In the Kattegat, Oithona spp. egg productionis strongly limited by food during summer and controlled bytemperature during winter. The seasonal signal in fecundityand population biomass is much weaker than in the co-occurringfree-spawning calanoid genera, where fecundity and populationbiomass undergo dramatic seasonal viaration. ¹Present address: Instituto Nacional de Investigacin y DesarrolloPesquero, INIDEP, CC 175, Playa Grande, Mar del Plata, Argentina