Copepod abundance in the western Irish Sea: relationship to physical regime, phytoplankton production and standing stock

The distinct patterns of stratification in the North Channeland stratified region of the western Irish Sea influence theseasonal abundance of phytoplankton. The 3–4 month productionseason in the stratified region was characterized by productionand biomass peaks in the spring (up to 2378 mg C m² day^–1and 178.4 mg chlorophyll m^–2) and autumn (up to 1280 mgC m^–2 day^–1 and 101.9 mg chlorophyll m^–2).Phytoplankton in the North Channel exhibited a short, late productionseason with a single summer (June/July) peak in production (4483mg Cm^–2 day^–1) and biomass (–160.6 mg chlorophyllm^–2). These differences have little influence on copepoddynamics. Both regions supported recurrent annual cycles ofcopepod abundance with similar seasonal maxima (182.8–241.810³ind. m^–2) and dominant species (Pseudocalanus elongatusand Acartia clausi). Specific rates of population increase inthe spring were 0.071 and 0.048 day¹ for the North Channel andstratified region, respectively. Increased copepod abundancein the stratified region coincided with the spring bloom, andwas significantly correlated with chlorophyll standing stock.Increased copepod abundance preceded the summer production peakin the North Channel. This increase was not correlated withchlorophyll standing crop, suggesting that a food resource otherthan phytoplankton may be responsible for the onset of copepodproduction prior to the spring bloom. Hetero-trophic microplanktonas an alternative food source, and advection of copepods fromthe stratified region, are proposed as possible explanationsfor copepod abundance increasing in advance of the summer peakin primary production.     

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,     

Trophodynamics of the plankton community at Dogger Bank: predatory impact by larval fish

The trophodynamics of a coastal plankton community were studied,focusing on fish larvae and their copepod prey. The major objectiveswere to describe distributional overlap and evaluate the predatoryimpact by larval fish. The study was carried out across DoggerBank in the North Sea, August-September 1991. Sampling transectscrossed tidal fronts off the Bank and plankton at all trophiclevels showed peak abundance within frontal zones. Also Verticallythere was a significant overlap in distributional patterns ofthe plankton. Seven species of fish larvae were abundant, ofthese sprat (Sprattus sprattus) dominated. The abundance ofone group of fish larvae peaked in the shallow water close tothe Bank, whereas other species, including sprat, were foundin deeper water. Prey preference and predation pressure of fishlarvae were assessed using information on prey sizes and growthrates of larvae and the copepod prey. We estimated larval removalof preferred prey sizes to 3–4% day^–1, counterbalancedby a 3–7% day^–1' replenishment from copepod productionand growth. Additional predation pressure on copepods by aninvertebrate predator was estimated to 1–3%day^–1.In conclusion, the dynamics of fish larvae and other zooplankterswere closely linked. At peak abundances of fish larvae (>35mg dry weight m^–2), the accumulated predation on specificsize ranges of copepods, exerted by larvae and other predators,could exceed the ability of copepod replenishment and intra-/interspecificcompetition among predators might take place.     

Response of ciliates and Cryptomonas to the spring cohort of a cyclopoid copepod in a shallow hypereutrophic lake

The impact of a cyclopoid copepod population on the protozoacommunity (two ciliate categories and Cryptomonas) was assessedweekly during the spring cohort of Cyclops vicinus (one monthduration) in hypereutrophic Lake Søbygård by insitu gradient experiments with manipulation of ambient zooplanktonabundance. As C.vicinus always made up >92% of the zooplanktonbiomass, the response of protozoa is assumed to be a resultof predation by the copepod. Significant effects of copepodbiomass on protozoa net population growth rates were obtainedin the four experiments. Copepod clearance rates were significantlyhigher on oligotrichs than on prostomatids and Cryptomonas butdeclined for all three protozoa categories during the firstthree weeks of the copepod cohort, probably because of the changein developmental instar composition of the copepod population.Grazing impact on protozoa at ambient copepod abundance wasconsiderable (range, 0.05–0.87 day^–1) and could,together with the estimated reproductive potential of protozoans(range, –0.20–0.87 day^–1), account for thedecline in abundance and biomass of protozoa during the cohortdevelopment. Carbon flow from the protozoa to C.vicinus (range,2.8–23.5 µg C l^–1 day^–1) documents thepresence of a trophic link between protozoa and the spring cohortof C.vicinus in Lake Søbygård.     

Mesozooplankton Distribution and Copepod Grazing in the Subtropical Atlantic Near the Azores: Influence of Mesoscale Structures

Mesozooplankton distribution and copepod grazing were investigatedin the Subtropical Atlantic Ocean near the Azores during theAZORES I (August) and II (April–May) cruises. Mesozooplanktonbiomass and abundance remained low throughout the region, butsignificant increases were found related to the presence ofthe Azores Front. The Azores Front also exhibited maximum valuesof copepod community ingestion, reaching 250 mg C m^–2ingested daily. This increase in ingestion was related to increasesin copepod abundance, but not in copepod gut contents. No relationshipwas found between gut contents, or ingestion, and phytoplanktonbiomass or production. Daily cycles were found in copepod gutcontents, being higher during the night, but not in copepodabundance. Multi-dimensional scaling analysis revealed differencesin copepod taxonomic composition between both sides of the front.During spring, daily copepod ingestion represents an averageof 6% of the integrated chlorophyll (Chl) a concentration and22% of the primary production. These percentages increase to15% of Chl and 61% of production if we only consider large (>2µm) phytoplankton. No clear influence of the cycloniceddy LETICIA was found in mesozooplankton biomass or grazing.A significant effect of the Great Meteor Tablemount was foundin copepod abundance and grazing, with higher values locatedwest of the mount.     

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.     

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.     

Productivity of Daphnia longispina in a highly humic boreal lake

The development of the Daphnia longispina (O. F. Müller)population in a highly humic boreal lake was followed throughoutone growing season, and the amount of secondary production wasestimated in relation to primary production and available foodresources. The growth rate method was applied in the secondaryproduction measurements. Daphnia longispina did not appear inthe water column until 16 May, after which the animals werepresent throughout the growing season. The population showedthree density peaks; the first appeared in early June, and thesecond and third in mid-July and at the beginning of September,respectively. Somatic production followed a seasonal pattern,with highest production rates in midsummer. The maximum valueof 127 mg C m^–2day^–1 was recorded at the beginningof July. The total annual net production of D. longispina was7.9 g C m^–2. During most of the growing season, the primaryproductivity in the lake was well below 100 mg C m^–2 day^–1and the total annual productivity of photosynthetic algae was5.0 g C m^–2. We conclude that in this lake the zooplanktonpopulation did not rely on phytoplankton primary productionas a sole carbon source, but that most of the carbon must haveoriginated from bacterial production either directly or througha microbial loop.     

Growth and development of Neocalanus flemingeri/plumchrus in the northern Gulf of Alaska: validation of the artificial-cohort method in cold waters

In situ growth and development of Neocalanus flemingeri/plumchrusstage C1–C4 copepodites were estimated by both the artificial-cohortand the single-stage incubation methods in March, April andMay of 2001–2005 at 5–6°C. Results from thesetwo methods were comparable and consistent. In the field, C1–C4stage durations ranged from 7 to >100 days, dependent ontemperature and chlorophyll a (Chl a) concentration. Averagestage durations were 12.4–14.1 days, yielding an averageof 56 days to reach C5, but under optimal conditions stage durationswere closer to 10 days, shortening the time to reach C5 (fromC1) to 46 days. Generally, growth rates decreased with increasingstage, ranging from 0.28 day^–1 to close to zero but weretypically between 0.20 and 0.05 day^–1, averaging 0.110± 0.006 day^–1 (mean ± SE) for single-stageand 0.107 ± 0.005 day^–1 (mean ± SE) forartificial-cohort methods. Growth was well described by equationsof Michaelis–Menten form, with maximum growth rates (G_max)of 0.17–0.18 day^–1 and half saturation Chl a concentrations(K_chl) of 0.45–0.46 mg m^–3 for combined C1–3,while G_max dropped to 0.08–0.09 day^–1 but K_chl remainedat 0.38–0.93 mg m^–3 for C4. In this study, in situgrowth of N. flemingeri/plumchrus was frequently food limitedto some degree, particularly during March. A comparison withglobal models of copepod growth rates suggests that these modelsstill require considerable refinement. We suggest that the artificial-cohortmethod is the most practical approach to generating the multispeciesdata required to address these deficiencies.     

Trophic links in the lowland River Meuse (Belgium): assessing the role of bacteria and protozoans in planktonic food webs

Trophic interactions within the plankton of the lowland RiverMeuse (Belgium) were measured in spring and summer 2001. Consumptionof bacteria by protozoa was measured by monitoring the disappearanceof ³H-thymidine-labelled bacteria. Metazooplankton bacterivorywas assessed using 0.5-µm fluorescent microparticles (FMPs),and predation of metazooplankton on ciliates was measured usingnatural ciliate assemblages labelled with FMPs as tracer food.Grazing of metazooplankton on flagellates was determined throughin situ incubations with manipulated metazooplankton densities.Protozooplankton bacterivory varied between 6.08 and 53.90 mgC m^–3 day^–1 (i.e. from 0.12 to 0.86 g C^–1bacteria g C^–1 protozoa day^–1). Metazooplankton,essentially rotifers, grazing on bacteria was negligible comparedwith grazing by protozoa (1000 times lower). Predation of rotiferson heterotrophic flagellates (HFs) was generally low (on average1.77 mg C m^–3 day^–1, i.e. 0.084 g C^–1 flagellatesg C^–1 rotifers day^–1), the higher contribution ofHF in the diet of rotifers being observed when Keratella cochleariswas the dominant metazooplankter. Predation of rotifers on ciliateswas low in spring samples (0.56 mg C m^–3 day^–1,i.e. 0.014 g C^–1 ciliates g C^–1 rotifers day^–1)in contrast to measurements performed in July (8.72 mg C m^–3day^–1, i.e. 0.242 g C^–1 ciliates g C^–1 rotifersday^–1). The proportion of protozoa in the diet of rotiferswas low compared with that of phytoplankton (<30% of totalcarbon ingestion) except when phytoplankton biomass decreasedbelow the incipient limiting level (ILL) of the main metazooplantonicspecies. In such conditions, protozoa (mainly ciliates) constituted50% of total rotifer diet. These results give evidence thatmicrobial organisms play a significant role within the planktonicfood web of a eutrophic lowland river, ciliates providing analternative food for metazooplankton when phytoplankton becomesscarce.     

The role of faecal material in the particulate organic carbon flux in the northern Humboldt Current, Chile (23{degrees}S), before and during the 1997-1998 El Nino

Sedimentation rates of faecal material, phytoplankton and microzooplanktonand production rates of faecal material from crustaceans andpelagic tunicates were estimated during the austral summer andwinter 1997, and summer 1998, in the northern Humboldt Current(23°S, off Antofagasta, Chile). Sampling periods coveredpre-El Niño (January 1997) and El Niño 1997–98(July 1997 and January 1998). Samples were collected using floatingsediment traps deployed at 65, 100, 200 and 300 m depth in oceanicand coastal areas. Sedimentation rates during January 1997 were,on average, 152 ± 23 and 85 ± 57 mg C m^–2day^–1 at 65 and 300 m depth, respectively. During July,these rates averaged 93 ± 56 mg C m^–2 day^–1at 65 m depth and 35 ± 12 mg C m^–2 day^–1at 300 m depth, while in January 1998 they were 98 and 109 ±37 mg C m^–2 day^–1 at 65 and 200 m depth, respectively.Recognizable faecal material made up the bulk of the sedimentingmatter, accounting for 8 ± 5% (n = 14), 31 ± 26%(n = 16) and 8 ± 5% (n = 5) of the average total organiccarbon recorded from all sediment trap samples collected duringJanuary and July 1997 and January 1998, respectively. However,at300 m depth, the contribution of recognizable faecal materialto total sedimented organic carbon increased to 43 ±33% (n = 4) during July 1997. The remaining sedimenting particlesconsisted mainly of tintinnids, crustacean exuviae, heterotrophicdinoflagellates (both thecated and athecated) and diatom cells.During this study, we estimated that only a minor fraction (average± SD = 5 ± 8%) of the copepod faecal materialproduced within the photic zone sedimented down to 300 m depth,suggesting an efficient recycling within the overlaying watercolumn. On the other hand, an important fraction (47 ±30%) of the euphausiid faecal strings was collected in the 300m depth trap, suggesting that this material would enhance thedownward flux of particulate organic matter (POC). POC fluxesto 65 and 300 m depth traps were in the range of 4–20%and 3–8% of the estimated primary production during thewhole study period. It is postulated that the overall verticalflux of particulates and, in particular, faecal pellets wasdetermined by a combination of three factors. The first wasthe composition of the zooplankton assemblages in the studyarea. When the dominant group was calanoid copepods, their faecesseemed to contribute poorly to the vertical flux of particulates.On the other hand, when the dominant group was euphausiids,a significant proportion of their faecal material was collectedin the sediment trap located at 300 m depth. The second wasthe relatively high abundance of cyclopoid copepods from thegenera Oncaea, Corycaeus and Oithona, which are reported tofeed on aggregates of phytodetritus and faecal pellets producedby calanoid copepods, suggesting that they may act as a naturalfilter to sedimenting particulates. The third was the compositionand size spectrum of the phyto- and microzooplankton assemblageswhich are potential food sources for the meso- and macrozooplankton.These factors were partially modulated by both the 1997–1998El Niño and seasonality.     

Predation and respiration by the small cyclopoid copepod Oithona similisr: How important is feeding on ciliates and heterotrophic flagellates?

Feeding on natural plankton populations and respiration of thesmall cyclopoid copepod Oithona similis were measured duringthe warm season in Buzzards Bay, Massachusetts, USA. AlthoughO.similis did not significantly ingest small autotrophic andheterotrophic flagellates (2–8 µn), this copepodactively fed on >10 µm particles, including autotrophic/heterotrophic(dino)flagel-lates and ciliates, with clearance rates of 0.03–0.38ml animal^–1 h^–1. The clearance rates increased withthe prey size. O.similis also fed on copepod nauplii (mainlycomposed of the N1 stage of Acartia tonsa with a clearance rateof 0.16 ml animal^–1 h^–1. Daily carbon ration fromthe combination of these food items averaged 148 ng C animal^–1day^–1 (41% of body C day^–1), with ciliates and heterotrophicdino-flagellates being the main food source ({small tilde}69%of total carbon ration). Respiration rates were 020–0.23µl O₂ animal^–1 day^–1. Assuming a respiratoryquotient of 0.8 and digestion efficiency of 0.7, the carbonrequirement for respiration was calculated to be 125–143ng C animal^–1 day^–1, close to the daily carbon rationestimated above. We conclude that predation on ciliates andheterotrophic dinoflagellates was important for O.similis tosustain its population in our study area during the warm season.     

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.     

Production of tropical larvaceans in Kingston Harbour, Jamaica: are we ignoring an important secondary producer?

The larvacean community was observed during an 18 month periodat the mouth of eutrophic Kingston Harbour, Jamaica. Duringthis period, larvaceans averaged 3607 m^–3 with a biomassof 2.2mg ash-free dry weight m^–3 (32.6mg AFDW m^–2in a community dominated by Oiko pleura longicauda There wereno relationships between larvacean biornass and any size fractionof chlorophyll, suggesting that other factors must normallyregulate larvacean communities. The evidence indicates thatthis regulation is by predation. Annual production by larvaceanswas 586 kJ m^–3 year ^–1 (29.3 g AFDW m^–2 year^–1);production of houses could represent an added 300–600kJ m^–2 year. While copepod biomass was 10 times higherthan that of the larvaceans during the same period, copepodgrowth rates were only one.third those of larvaceans. Thus,larvacean annual production is at least 30% that of the copepods,due to their rapid growth rates, and at least 50% that of thecopepods when house production is considered. The contributionof larvaceans to plankton production has been underappreciatedhistorically when only their biomass is considered. ¹Present address :Monterey Bay Aquarium Research Institute 7700Sandholdt Road, Moss Landing, CA 95039-0628, USA     

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.     

Seasonal nitrogen assimilation and carbon fixation in a fjordic sea loch

Carbon (C) fixation and nitrogen (N) assimilation rates havebeen estimated from ¹⁴C and ¹⁵N techniques for a 12 month periodin a Scottish sea loch. The maximum rate of nitrogen assimilated(29.92 mmol N m^–2 day^–1) was in April at the mostseaward station; similar high rates were experienced duringMay at the other stations. Carbon fixation rates were maximal(488–4047 mg C m^–2day^–1) at the time of highphytoplankton biomass (maximum 8.3 mg m^–3 chlorophylla) during May, whilst nitrate concentrations remained >0.7µ.mol l^–1. C:N assimilation ratios suggest nitrogenlimitation only during the peak of the spring bloom, althoughat times nitrogen (nitrate and ammonium) concentration fellto 0.2 µmol l^–1 in the following months. The verticalstability of the water column, influenced by tidal and riverineflushing, varied along the axis of the loch, resulting in markeddifferences between sampling stations. Although ammonium waspreferentially assimilated by phytoplankton, >50% of productionwas supported by nitrate uptake and only during the summer monthswas the assimilation of ammonium quantitatively important.     

The seasonal abundance and production of Oithona nana (Copepoda:Cyclopoida) in Southampton Water

Recent studies indicate that Oithona spp. contribute significantlyto total copepod biomass. Little is known, however, about theirecological significance, particularly in the case of the estuarineOithona nana. A study comprising three sites within SouthamptonWater was conducted to evaluate the late-stage copepodite/adult(stages IV–VI) O. nana community, using 120-µm meshnets. Although present throughout the estuary, there was a strikingspatial gradient with O. nana most common in the upper estuary.A clear seasonal pattern was observed with O. nana as the mostabundant copepod species from late summer until early winter.It comprised 61% of all copepods recorded, with a biomass of757.22 mg C m^–3. Production estimates of O. nana werederived from the ‘instantaneous-growth’ approach,using appropriate growth equations. The estimated productionof O. nana ranged from 1.50 mg C m^–3 year^–1 withinthe lower estuary to 146.77 mg C m^–3 year^–1 in theupper estuary. In the upper estuary, this compares with productionrates of 187.47 mg C m^–3 year^–1 for all Acartiacongeners (excluding nauplii), the most common calanoid genus.Throughout the estuary, O. nana annual production represented18% of total copepod production clearly indicating that, atleast in the upper estuary, O. nana production may be directlycomparable with calanoid production.     

Growth and development of Metridia pacifica (Copepoda: Calanoida) in the northern Gulf of Alaska

Juvenile growth and development rates for Metridia pacifica,one of the dominant larger copepods in the subarctic Pacific,were investigated from March through October of 2001–2004in the northern Gulf of Alaska. The relationship between prosomelength (PL, µm) and dry weight (DW, µg) was determined:log₁₀ DW = 3.29 x log₁₀ PL – 8.75. The stage durationsof copepodites ranged from 3 to 52.5 days but were 8–15days under optimal condition. Seasonally, growth rates increasedfrom March to October and typically ranged between 0.004 and0.285 day^–1, averaging 0.114 ± 0.007 day^–1(mean ± SE). After standardization to 5°C (Q₁₀ of2.7), growth rates averaged 0.083 ± 0.005 day^–1and were significantly correlated to chlorophyll a, with saturatedgrowth rates of 0.149 day^–1 for C1–C3, 0.102 day^–1for C4–C5 and 0.136 day^–1 for all stages combined.Measured juvenile growth rates were comparable with specificegg production rates in this species. The comparisons of ourrates in this study with those predicted by the global modelsof copepod growth rates suggested that further refinement ofthese models is required.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

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.