Spatial structure of the zooplankton community in the Cabo Frio region (Brazil) influenced by coastal upwelling

Cabo Frio area (Central-southern Brazilian littoral) is characterized by a coastal upwelling due to prevailing E-NE winds, mainly during the spring-summer season. The geographic distribution of the zooplankton was described in order to separate ecological areas in terms of specific communities. Cluster analysis by weighted pair-group method (WPGA) and principal component (PC) analysis was performed on the most important hydrological parameters and zooplankton taxa. The first PC, explaining 38% of the total zooplankton variability, represented the influence of the nearshore-offshore gradient. Zooplankton communities and ecological areas were defined as follows:

1. nearshore neritic: the richest region with more than 5 000 org. m-3. In this very narrow nearshore region we observed 3 groups of taxa deliminated by upwelling effects and trophic conditions: a) Ctenocalanus vanus, Penilia avirostris, Calanoides carinatus and Copepod larva, all herbivorous species associated with the phytoplankton blooms in the upwelling zone; b) Creseis acicula and Temora stylifera; c) Siphonophora, Chaetognaths and a high diversity of Copepoda dominated by Coryceus amazonicus, C. giesbrechti, Eucalanus pileatus, Clausocalanus acuicornis, all carnivorous or omnivorous feeders representing a high degree of complexity of the ecosystem.
2. offshore neritic: characterized by appendicularians, this region constitutes an intermediate zone between coastal and oceanic waters.
3. oceanic: occupied by the Brazil Current where zooplankton is poor, but diversified, with typical copepods from tropical warm water: Clausocalanus furcatus, Mecynocera clausi, Corycella gracilis, Oithona setigera, Coryceus typicus, Oncea conifera, Undinula vulgaris, Calocalanus pavo, Haloptilus longicornis, Lucicutia flavicornis.

     

Zooplankton annual cycle in a Mediterranean coastal area

The annual cycle of the zooplankton community in the SaronikosGulf (Aegean Sea, Greece) was studied over a period of 2 years.The sampling scheme included monthly hauls at three stationsdifferentiated according to depth and neritic character. Maximumvalues of total zooplankton abundance were found in summer monthsup to early autumn and partially in spring. Copepods dominatedduring most of the year (Clausocalanus furcatus, Paracalanusparvus, Temora stylifera, Ctenocalanus vanus, Oithona similis,Oithona plumifera), while cladocerans (Penilia avirostris) wereabundant in summer months and in September. No important fluctuationswere detected between the 2 years of study, while monthly oneswere more significant in the more neritic station during thewinter–spring period. Correspondence analysis showed thatthe seasonal evolution of zooplankton is related to environmentalparameters: temperature, hydrography expressed in open sea influenceand topography.     

Zooplankton assemblages and influence of environmental parameters on them in a Mediterranean coastal area

Temporal dynamics of zooplankton communities and assemblages,as well as the influence of environmental factors on them, werestudied in Saronikos Gulf (Aegean Sea, Greece). Different multivariatetechniques (hierarchical clustering, multidimensional scalingand correspondence analysis) were applied on a 2 year data setof zooplankton species composition, based on samples collectedat five stations of the study area. A clear discrimination oftwo communities was revealed, the first one in the semi-enclosedpolluted northern part (Elefsis Bay) andthe second in SaronikosGulf proper. Within the latter community, five assemblages weredistinguished: (i) the coastal winter assemblage characterizedby Ctenocalanus vanus, Oithona similis, Clausocalanus pergensand Fritillaria pellucida; (ii) the psychrophilic assemblageinfluenced by the open sea and characterized by Oithona plumiferaand Clausocalanus jobei; (iii) the spring assemblage characterizedby Evadne nordmanni and Centropages typicus; (iv) the coastalthermophilic assemblage characterized by Penilia avirostis,Evadne ergesrina and Temora stylifera; (v) the thermophilicassemblage influenced by the open sea and characterized by Clausocalanusfurcatus. Both zooplankton and environmental data were treatedby multiple correspondence analysis which revealed the importanceof some environmental factors on zooplankton community composition(eutrophication-pollution, temperature, water mass circulation,hydrology and topography).     

Les diatomees dans l'upwelling de Cabo Frio (Bresil): liste d'especes et etude ecologique

The diatoms in the Cabo Frio (Brazil) upwelling: check listand ecological study. A qualitative and quantitative study onthe diatoms from the Cabo Frio area was made and a check listproduced. Sampling was carried out over one year at an anchoragestation in the upwelling zone. Data were analysed by means ofa multivariate statistical method, in order to show the mainsynecological aspects of the diatom populations. Diatoms dominatein most samples, except in the Brazil current water where dinoflagellatesare dominant, and where there is a poor population of thermophilicdiatoms. Recently upwelled water has a poor phytoplankton composedof epiphytic and benthic species; these are substituted by intenseblooms of neritic and opportunist diatoms when the temperaturerises. The annual distribution of the diatoms accompanies theoccurrence of upwelling. The vertical thermal gradient or waterstability is the most important factor affecting the specificcomposition and distribution of the diatom communities.     

How could thermal stratification affect horizontal distribution of depth-integrated metazooplankton communities in the Gulf of Gabes (Tunisia)?

The spatial and temporal variability of metazooplankton communities in relation to environmental and trophic factors were studied in the Gulf of Gabes during three periods of increasing thermal stratification in June, July and September. Non-metric multi-dimensional scaling (NMDS) revealed the time and space succession of three zooplankton communities in relation to the degree of thermal stratification. A co-inertia analysis showed the association between these communities and the trophic conditions (abundance and composition of microplankton) driven by stratification. Three divergent communities were evident. The first, characterized by annelid larvae and coastal copepods, was associated with shallow mixed and microplankton-rich water (June, coastal). The second, characterized by euphausiids, amphipods, doliolids and several deep-water copepods, corresponded to highly stratified microplankton-poor water (September). The third, characterized by the most ubiquitous and abundant species, was linked to intermediate water (June deep and July). Temperature and salinity conditions also explained the distribution of key species. Clausocalanus furcatus, Oithona plumifera and Triconia conifera were clearly associated with warm and highly stratified water (September). Enhanced thermal stratification led to higher surface salinity, explaining the emergence of euryhaline taxa such as Oithona plumifera.     

Biogeographic study of the planktonic communities of the Prince Edward Islands (Southern Ocean)

Microphytoplankton and zooplankton composition and distributionin the vicinity of the Prince Edward Islands and at the Sub-antarcticFront (SAF) were investigated in late austral summer (April/May)1996. Samples were collected for analysis of chlorophyll a concentration(Chi a), microphytoplankton and zooplankton abundance. Generally,the highest Chl a concentrations (up to 2.0 µg l^–1)and zooplankton densities (up to 192 ind. m^–3) were recordedat stations within the inter-island area while the lowest values(<0.4 µg l^–1) were observed at stations upstreamof the islands. High Chl a and zooplankton biomass values werealso associated with the SAF. Microphytoplankton were dominatedby chain-forming species of the genera Chaetoceros (mainly C.neglectus),Fragilariopsis spp. and the large diatom Dactyliosolen antarcticus.The zooplankton assemblages were always dominated by mesozooplanktonwhich at times contributed up to 98% of total zooplankton abundanceand up to 95% of total biomass. Among mesozooplankton, copepods,mainly Clausocalanus brevipes and Metridia lucens numericallydominated. Among the macrozooplankton euphausiids, mainly Euphausiavallentini, E.longirostis and Stylocheiron maximum, and chaetognaths(Sagitta gazellae) accounted for the bulk of abundance and biomass.Cluster and ordination analysis did not identify any distinctbiogeographic regions among either the microphytoplankton orzooplankton.     

Le zooplancton de la baie de Sh?diac (Nouveau-Brunswick)

Analyses of boreal zooplankton of Shediac Bay demonstrate theabundance of copepods (81%) and meroplankters (18%). Whetherexclusively pelagic or not, 67 species are mentioned for thefirst time in this Northumberland Strait area, out of 76 recordedwithin 23 higher taxa. The fluctuations were observed from Mayto November and pointed out the dominance of such copepods asAcartia tonsa, A. clausi, Oithona similis and Centropages hamatusin relation with temperature, salinity and food distributions. ¹Adresse actuelle: Station marine de Tul?ar, B.P. 141, Universit?de Madagascar, (R?p. Malgache)     

Pelagic fish and zooplankton species assemblages in relation to water mass characteristics in the northern Bering and southeast Chukchi seas

This research explores the distributions and community composition of pelagic species in the sub-Arctic and Arctic waters of the northern Bering and central and southern Chukchi seas during September 2007 by linking pelagic zooplankton and fish assemblages to water masses. Juvenile saffron cod (Eleginus gracilis), polar cod (Boreogadus saida), and shorthorn sculpin (Myoxocephalus scorpius) were most abundant in warm, low salinity Alaska Coastal Water (ACW) of the central Chukchi Sea, characterized by low chlorophyll, low nutrients, and small zooplankton taxa. Adult Pacific herring (Clupea pallasii) were more abundant in the less stratified Bering Strait waters and in the colder, saltier Bering Shelf Water of the northern Bering and southern Chukchi seas, characterized by high chlorophyll, high nutrients, and larger zooplankton taxa. Juvenile pink (Oncorhynchus gorbuscha) and chum (O. keta) salmon were most abundant in the less stratified ACW in the central Chukchi Sea and Bering Strait. Abundances of large zooplankton were dominated by copepods (Eucalanus bungii, Calanus glacialis/marshallae, Metridia pacifica) followed by euphausiids (juvenile Thysanoessa raschii and unidentified taxa), whereas small zooplankton were dominated by bivalve larvae and copepods (Centropages abdominalis, Oithona similis, Pseudocalanus sp.). Pelagic community composition was related to environmental factors, with highest correlations between bottom salinity and large zooplankton taxa, and latitude and fish species. These data were collected in a year with strong northward retreat of summer sea ice and therefore provide a baseline for assessing the effects of future climate warming on pelagic ecosystems in sub-Arctic and Arctic regions.     

Zooplankton composition and distribution off the coast of Galicia, Spain

During June and September 1984, zooplankton samples were collectedwith other hydrographic and biological data along the Galiciancoast (NW of Spain). In June copepods contributed {small tilde}60%to the total zooplankton community, with larvaceans, siphonophoresand cladocerans also abundant. In September >90% of the zooplanktonsampled were copepods. The dominant species of copepods in bothJune and September were Acartia clausi, Paracalanus parvus andTemora longicornis. The meroplankton was dominated by echinoderms,bryozoans, barnacle larvae and bivalve larvae. In June the averagezooplankton biomass was 31.08 mg C m^–3; the Septemberaverage was 41.69 mg C m^–3. The relationship between theslopes of the regression equations (biomass versus abundance)suggests that the zooplankton assemblage in June was composedby larger animals than in September. The major concentrationof zooplankton was between 0 and 50 m, with both June and Septemberdaytime surface samples having 6–7 times the amount oforganisms than the lower water column (50–100 m). Therewere no distinct differences in total zooplankton abundancesat the inshore and offshore stations; however, the inshore stationsoften had a higher percentage of meroplankton than the offshorestations. In June zooplankton abundance at the northern transectsand the western transects was similar. In September there weregreater concentrations of zooplankton in the western Galicianshelf as compared with the northern shelf. These differencesin the horizontal distribution of the zooplankton were relatedto upwelling events.     

Egg production rates and stage-specific development times of Clausocalanus furcatus (Copepoda, Calanoida) in the northern Gulf of Mexico

The calanoid copepod Clausocalanus furcatus is widely distributedin tropical and subtropical oceans, yet knowledge of its reproductionand development remains limited. We estimated egg productionrates and stage-specific development times from laboratory incubationsand field samples collected at a petroleum platform in the northernGulf of Mexico. Egg production experiments were performed in2002 and 2003 at a laboratory on the platform using freshlycollected female copepods. Median stage development durationswere also measured during the incubations. Logistic regressionmodels were used to estimate median stage durations. The mediandevelopment times of Clausocalanus furcatus from hatching toadulthood ranged from 13 to 20 days. Field samples were collectedfrom waters beneath the platform at 12-h intervals using a 30-LNiskin water bottle during March–April and May–June2003, and the Edmonson egg ratio method was used to estimatethe mean in situ egg production rate. This field-derived meanegg production rate was significantly lower than the 12.08 eggsfemale^–1 day^–1 measured from incubation experiments,and this difference may reflect high in situ egg mortality.In situ egg production rates also showed high variability andwere almost five times higher during March–April comparedto May–June.     

The relationship between plankton-feeding Bonaparte's and Mew Gulls and tidal upwelling at Active Pass, British Columbia

The food habits of Bonaparte's (Larus Philadelphia) and MewGulls (L. canus) were studied at Active Pass, British Columbia,in relation to upwelling of zooplankton. Bonaparte's Gulls fedmostly on planktonic crustaceans during September-November andagain during April-May, while Mew Gulls foraged there chieflyin February and March. Both species ate predominantly the euphausiid,Thysanoessa raschii, in spring, while Bonaparte's Gulls fedmainly on the amphipods, Parathemisto pacifica and Calliopiuslaeviusculus, in fall. Year-round sampling of zooplankton andcollection of temperature and salinity data showed gull numbersto correlate with times of maximum upwelling and abundance ofzooplankton prey in surface waters. Outside the upwelling zonein Active Pass, Bonaparte's and Mew Gulls fed mostly on fishesand intertidal organisms, but also on zooplankton along tidelines.     

Spatio-Temporal Variability of Copepod Abundance along the 20°S Monitoring Transect in the Northern Benguela Upwelling System from 2005 to 2011

Long-term data sets are essential to understand climate-induced variability in marine ecosystems. This study provides the first comprehensive analysis of longer-term temporal and spatial variations in zooplankton abundance and copepod community structure in the northern Benguela upwelling system from 2005 to 2011. Samples were collected from the upper 200 m along a transect at 20°S perpendicular to the coast of Namibia to 70 nm offshore. Based on seasonal and interannual trends in surface temperature and salinity, three distinct time periods were discernible with stronger upwelling in spring and extensive warm-water intrusions in late summer, thus, high temperature amplitudes, in the years 2005/06 and 2010/11, and less intensive upwelling followed by weaker warm-water intrusions from 2008/09 to 2009/10. Zooplankton abundance reflected these changes with higher numbers in 2005/06 and 2010/11. In contrast, zooplankton density was lower in 2008/09 and 2009/10, when temperature gradients from spring to late summer were less pronounced. Spatially, copepod abundance tended to be highest between 30 and 60 nautical miles off the coast, coinciding with the shelf break and continental slope. The dominant larger calanoid copepods were Calanoides carinatus, Metridia lucens and Nannocalanus minor. On all three scales studied, i.e. spatially from the coast to offshore waters as well as temporally, both seasonally and interannually, maximum zooplankton abundance was not coupled to the coldest temperature regime, and hence strongest upwelling intensity. Pronounced temperature amplitudes, and therefore strong gradients within a year, were apparently important and resulted in higher zooplankton abundance.     

Summer coastal zooplankton biomass and copepod community structure near the Italian Terra Nova Base (Terra Nova Bay, Ross Sea, Antarctica)

The structure of the zooplankton biotic community and of copepodpopulation in the coastal area of Terra Nova Bay (Ross Sea,Antarctica) was investigated during the 10th Italian AntarcticExpedition (1994/1995). Zooplankton biotic community consistedmainly of pteropods (Limacina helicina and Clione antarctica),Cyclopoid (Oithona similis), Poecilostomatoid (Oncaea curvata)and Calanoid (Ctenocalanus vanus, Paraeuchaeta antarctica, Metridiagerlachei and Stephos longipes) copepods, ostracods, larvalpolychaetes and larval euphausiids. Zooplankton abundance rangedfrom 48.1 ind m^–3 to 5968.9 ind m^–3, and copepodabundance ranged from 45.2 ind m^–3 to 3965.3 ind m^–3.The highest peak of zooplankton abundance was observed between25 m and the surface and was mainly due to the contributionof O. similis, O. curvata and C. vanus. Zooplankton biomassranged from 5.28 mg m^–3 to 13.04 mg m^–3 dry weight;the maximum value was observed between 25 m and the surface.Total lipid content varied from 216.44 to 460.73 mg g^–1dry weight.     

Biogeography of common dolphins (genus Delphinus) in the Southwestern Atlantic Ocean

• 1 The common dolphins (genus Delphinus) have one of most problematic taxonomies and complex distribution patterns of all cetaceans. Although the taxonomy and the distribution seem to have been clarified somewhat in the eastern North Pacific and Indo‐Pacific Oceans, many questions remain in the Southwestern Atlantic Ocean (SWA). We review the biogeography of Delphinus in the SWA.
• 2 We reviewed data from strandings, incidental catches and sightings since 1922. Systematic surveys were conducted in five major areas. Twenty‐one natural history collections were examined, and 135 skulls were measured.
• 3 A total of 184 records of common dolphins were compiled. Delphinus apparently occurs in three stocks in the SWA: one located in northern Brazil and two from southeastern Brazil (～22°S) to central Argentina (～42°S). Two distinct patterns in habitat use were observed by depth: in southeastern Brazil, sightings were restricted to coastal waters with water depths ranging from 18m to 70m. On the other hand, in the area that extends from southern Brazil to Central Argentina (from 28°S to 42°S), sightings were recorded in deeper waters, ranging from 71m to 1435m, with the exception of occasional coastal sightings. The cranial analyses demonstrated that both short‐beaked common dolphins Delphinus delphis and long‐beaked common dolphins Dephinus capensis occur in the SWA.
• 4 In the SWA, Delphinus seems to occur near areas of high productivity. One stock is associated with the productive waters discharged by the Amazon River and possibily with the coastal upwelling system off the coast of Venezuela, while the other stocks are associated with the Cabo Frio upwelling system and the Subtropical Convergence. Our results indicate that the current taxonomy does not adequately reflect the amount of variation within the genus in the world.

     

Zooplankton community structure and copepod egg production in coastal waters of the central Great Barrier Reef lagoon

We describe zooplankton community structure and copepod eggproduction in the vicinity of the coastal boundary zone of theGreat Barrier Reef lagoon, Australia. The abundance and eggproduction rate of constituents of the zooplankton assemblagecharacteristic of the coastal zone rapidly increase subsequentto events such as flooding and upwelling. Our sampling spannedtwo summer monsoonal seasons, the first of which (1990-91) wasvery wet. The second monsoonal season (1991-92) was very dryand was characterized by intrusive upwelling events from theCoral Sea. Chlorophyll a concentrations did not rise in thewet year, probably because of light limitation, but did riseas a result of upwelling. Terrestrial run-off in the wet yearhad a greater apparent effect on zooplankton abundance patternsthan did upwelling in the dry year, except where coastal trappingallowed sufficient time for increases in zooplankton abundanceto occur. Egg production rates by the copepods Acrocalanus gibberand Acrocalanus gracilis showed haphazard spatial differences.Nitrogen-specific egg production ranged between 0.03 and 0.21day^–1 for A.gibber, and between 0.13 and 0.41 day^–1for A.gracilis. The egg production rate by A.gibber was foodlimited for most of the year and showed a poor correlation withtemperature. ³Present address: Department of Biological Sciences, FloridaTech, 150 W University Boulevard, Melbourne, FL 32901, USA     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

First observations on the biology of Clausocalanus furcatus (Copepoda, Calanoida)

This study represents the first attempt to acquire basic informationon the feeding, reproduction and development of the copepodClausocalanus furcatus. This small oceanic calanoid benefitsfrom low phytoplankton concentrations and has a swimming behaviourwhich differs dramatically from that of other small copepods.Our preliminary results may be indicative of the success ofthis species in oligotrophic waters.     

Evidence for Changes in Estuarine Zooplankton Fostered by Increased Climate Variance

Estuaries are among the most valuable aquatic systems in terms of their services to human welfare. They offer an ideal framework to assess multiscale processes linking climate and food web dynamics through the hydrological cycle. Resolving food web responses to climate change is fundamental to resilience management of these threatened ecosystems under global change scenarios. Here, we examined the temporal variability of the plankton food web in the Mondego Estuary, central Iberian Peninsula, over the period 2003 to 2012. The results pointed out a cascading effect from climate to plankton communities that follow a non-stationary behavior shaped by the climate variance envelope. Concurrent changes in hydrographic processes at the regional, that is, upwelling intensity, and local, that is, estuarine hydrology, scales were driven by climatic forcing promoted by the North Atlantic Oscillation; the influence of which permeated the physical environment in the estuary affecting both autotrophic and heterotrophic communities. The most conspicuous change arose around 2008 and consisted of an obvious decrease in freshwater taxa along with a noticeable increase in marine organisms, mainly driven by gelatinous zooplankton. The observed increase in small-sized cosmopolitan copepods, that is, Clausocalanus arcuicornis, Oithona plumifera, thermophilic species, that is, Penilia avirostris, and gelatinous zooplankton suggests a structural change in the Mondego plankton community. These results provide empirical support to the expectation that expanding climate variance changes plankton structure and functioning, likely fostering trophic interactions in pelagic food webs.     

Seasonal abundance and reproduction of clausocalanid copepods in the northern Gulf of Aqaba (Red Sea)

Clausocalanid copepods dominate subtropical mesozooplanktoncommunities. Their life cycles, however, are generally unknown.Here, we follow the seasonal population development and reproductivebiology of Red Sea clausocalanids in the northern Gulf of Aqaba,between February 2002 and December 2003. Monthly vertical haulstaken with a Nansen net (200-µm mesh size) between 100m and the surface revealed four species, including a new record(Clausocalanus minor) for the area. Ctenocalanus vanus abundancespeaked during winter and spring, followed by Clausocalanus farranipeaking in spring and, additionally, in June 2003. In 2002,the abundance of Clausocalanus furcatus showed no distinct seasonaldevelopment of abundance, whereas in 2003, it reached high densitiesin January, July and November. Overall, C. vanus dominated theclausocalanid community during the first half of the year andC. furcatus during the second half of the year, whereas thepercentage of C. farrani remained fairly stable throughout bothyears. The correlation between the environmental parameters(temperature and chlorophyll a concentration) and the proportionof mature gonads was significant only between C. vanus and temperature.Incubation experiments showed that egg production rates variedbetween 0–3.3 and 1.8–9.2 eggs female^–1 day^–1in C. farrani and C. furcatus, respectively. Endoparasitismby dinoflagellates and the occurrence of intersexes were foundin all three species, with C. farrani being the most affected.     

Capelin larvae (Mallotus villosus) and community structure of zooplankton off the coast of northern Norway

Capelin larvae (Mallotus villosus) and mesozooplankton werecollected in May 2001 in the southern part of the Barents Sea.During the period of hatching and first feeding, capelin larvaeexperience high mortality rates. In order to understand theunderlying mechanisms influencing larval survival, we monitoreda selected site for 10 days using a variety of different samplingapproaches. Hydrological parameters conductivity–temperature–depth–fluorescence(CTDF) were recorded and zooplankton samples obtained simultaneouslyusing MOCNESS at three time periods. The mesozooplankton consistedessentially of copepods (85), dominated by Calanus finmarchicus.The zooplankton community consisted of different assemblagesderived from different areas (i.e. different water masses; Atlanticand coastal). Capelin larvae were found in surficial coastalwaters. Copepod nauplii and C. finmarchicus were less abundantin coastal waters where small copepodites (such as Acartia spp.and Temora longicornis) were found together with early stagesof coastal invertebrates that are possible prey for capelinlarvae. We suggest that capelin larvae base their foraging onsmall prey organisms frequently found among the coastal zooplanktonand that these larvae are less dependent on the propagationof the recruiting generation of C. finmarchicus present fromApril to June.