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.     

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.     

Seasonal population dynamics and production of Microsetella norvegica, a widely distributed but little-studied marine planktonic harpacticoid copepod

Microsetella norvegica is a widely distributed marine planktonicharpacticoid copepod, which is poorly known from the biologicalpoint of view. We investigated the seasonal population dynamicsandproduction of M. norvegica in the central part of the InlandSea of Japan. It occurred throughout the year, whilst its reproductionwas confined to the warm season between May and November. Theproportion of ovigerous females, which carry a single egg sac,was low (mean: 23.1%) in August and September, and high (53.6%)in October. Their brood size attained a maximum (mean: 15.8eggs per sac) in July and August and declined gradually to 6.2eggs in November. Duration time from egg laying to moultingto adulthood was temperature-dependent; it was 31.9 and 14.3days at 20 and 27°C, respectively, under excess food conditionsin the laboratory. An enormously high population abundance (7.32x 10⁴ individuals m^-3), which accounted for 86.5% of the totalcopepods, and biomass (69.6 mg C m^-3) gave an annual maximumproduction rate of 4.90 mg C m^-3 day^-1) in October. Naupliiand copepodites disappeared in December, and the overwinteringpopulation was represented by adults, mainly large females.Associations of M. norvegica with marine snow aggregates, whichhave often been found in oligotrophic waters, were not observedin the food-rich environment of the Inland Sea of Japan.     

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.     

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.     

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 (     

Gas Exchange of the Avian Egg Time, Structure, and Function

Data are presented for oxygen consumption water loss duringincubation water vapor conductance of the shell and pore numberof avian eggs and the way in which these values relate not onlyto egg mass but also to incubation time. It is proposed thatall these functions are proportional to the product of egg massand rate of development where the latter is defined as the inverseof incubation time. These interrelationships account at theend of incubation for similar O₂ and CO₂ tensions in the airspace of eggs utilization of calories (0.5 kcal g^–1) andwater loss (15 g g^–1)     

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.     

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.     

Endosulfan-enhanced crest induction in Daphnia longicephala: evidence for cholinergic innervation of kairomone receptors

Daphnia iongicephala were reared from early embryogenesis inthe presence or absence of crest-inducing kairomones releasedby Anisops gratus (Notonectidae) and from birth in one of fiveconcentrations of the organochiorine pesticide, endosulfan (0,0.1, 1.0, 10 or 100 g 1^–1 The morphologyof the daphnidswas measured 3 days after birth and on production of the firstbrood of eggs. The reproductive parameters, first brood size,age at first reproduction, mean egg volume and total egg volume,were also measured. Endosulfan significantly induced crest developmentin 3-day-old daphnids at 0.1, 1.0 and 10 g ^1– in thepresence and absence of Anisops kairomone, although crest sizein the absence of kairomone was only marginally greater thancontrols. A concentration of 10 pg 1^–1 endosulfan significantlyenhanced crest growth of kairomone-exposed daphnids at maturity.The Anisops kairomone alone induced a large crest in D.iongicephola,reduced first brood size, mean egg volume and total egg volume,and increased age at maturity. The coefficient of variationof mean egg volume was significantly reduced by Anisops kairomone.It is hypothesized that endosulfan enhances crest developmentbyinhibition of .     

Effect of temperature and food concentration on post-embryonic development, egg production and adult body size of calanoid copepod Eurytemora affinis

Post-embryonic development time, egg production rate and adultbody size of calanoid copepod Eurytemora affinis from Lake Ohnuma,Japan, were determined under six temperature-food conditions(10³,5 x 10³, 10⁴ and 5 x 10⁴ cells ml^–1 at 15°C,5 x 10⁴ cells ml^–1 at 10and20°C) in the laboratory.The measured parameters varied with both temperature and foodconcentration. Development time from hatching to adult femalewas 9.2, 11.4 and 22.8 days at 20, 15 and 10°C respectively,at the highest food concentration. The males developed to adultat one to two days earlier than the females. An effect of foodshortage on development time occurred at the lowest food concentration.This development time was 24.8 days even at 15°C, beingtwice as long as that at the highest food concentration. Prosomelength of these food-limited females was approximately 75% ofwell-fed ones, which reduced by only 10% with increasing temperaturefrom 10 to 20°C. Clutch size (C, eggs clutch^–1) ofwell-fed individuals depended on prosome length of the adultfemale (L, mm), and was expressed as an equation: C = 65.2 L³⁸³. Clutch size of individuals reared at less than 10⁴ cellsml^–1, however, mostly laid below the estimated curve,especially at the lowest food concentration being only 10% ofthat at the highest food concentration. These results suggestthat food availability is a more important factor affectingpopulation growth of E.affinis in Lake Ohnuma than variationof temperature.     

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.     

Resting egg production and oviducal cycling in two sympatric species of alpine diaptomids (Copepoda: Calanoida) in relation to temperature and food availability

Resting egg production and oviducal cycling were investigatedfor the calanoid copepods Arctodiaptomus alpinus and Acanthodiaptomusdenticornis both in the laboratory and in a small karstic alpinelake by making a census of the number of eggs produced and theproportion of females in each of four morphologically distinguishedreproductive conditions each day in the laboratory or duringa 2–3 week period in lake enclosures. In the laboratory,individuals were maintained on a mixed diet of natural phytoplanktonat constant temperatures of 4, 10, 15 and 20C, respectively.Both species differed considerably in their temperature requirementsfor reproduction. Lifetime fecundity was highest at 10C in A.alpinusand at 20C in A.denticornis, with up to 327 eggs ^–1 spawnedin the former and up to 582 eggs ^–1 in the latter species.Unfavorable temperatures were further reflected in an increasein egg mortality and the allocation of time spent in a post-reproductivephase, as well as in a decrease of longevity. Increasing temperaturesenhanced egg production rates due to decreasing clutch productionperiods, although clutch size was negatively correlated withtemperature. Maximum rates reached 5.88 and 7.98 eggs ^–1day^–1 in the laboratory, and 0.73 and 0.55 eggs ^–1day^–1 in enclosures in A.alpinus and A.denticornis, respectively.Egg production rates and clutch size were clearly governed bynutritional conditions in the lake, but were less affected byfood supply in the laboratory. Here, rates of egg productionwere adapted to improving food supply by increasing the frequencyof spawning events, rather than the number of eggs per clutch.No correlation was found between female body size and reproductiveparameters in the laboratory. A very low proportion of totalclutch production resulted in clutches composed of subitaneouseggs, i.e. 0.14% in A.denticornis and 1.20% in A.alpinus. Oviducalphase duration allocations indicate that there exists a temperatureoptimum for gamete maturation.     

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.     

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.     

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.     

The estimation of a mackerel (Scomber scombrus L.) spawning stock size by plankton survey

The International Council for the Exploration of the Sea (ICES)Mackerel Working Group recognises two mackerel stocks in Europeanwaters, the North Sea stock and the Western stock. From Marchto July 1977 an Anglo-French plankton survey was carried outin the shelf waters of the Bay of Biscay, Celtic Sea and westof Ireland to estimate the size of the Western stock. Sampleswere collected with a 76 cm Lowestoft pattern high-speed planktonsampler fished to a maximum depth of 100 m. The number of eggsm^–2 at each of five development stages were raised todaily production estimates with development rate coefficientsmeasured with live material collected in March. The mean numberof stage I eggs m^–2 d^–1 was raised to a total dailyproduction estimate for each month by the area within the minimumcontour level, and a production curve was drawn for the spawningseason. The total egg production was estimated at 1.98 x 10¹⁵eggs, + 30%, – 20%. With a mean fecundity of 361 000 anda sex ratio of 0.64 males per female this is equivalent to 8995 million fish.     

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.     

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.     

Planktonic ciliates in the northwestern Indian Ocean: their abundance and biomass in waters of contrasting productivity

The abundance and biomass of planktonic ciliates in the northwesternIndian Ocean ranged from 31 l^–1 and 0.1 µg C l^–1in oligotrophic open-ocean waters to 823 l^–1 and 1.2 µgC l^–1 in more productive waters of the equator, northernArabian Sea and Gulf of Oman. ³Present address: British Antarctic Survey, High Cross, MadingleyRoad, Cambridge, CB3 OET, UK