Environmental factors influencing diurnal distribution of zooplankton and ichthyoplankton

The diurnal vertical distribution of a large number of speciesof zooplankton, icbthyoplankton and micronekton were determinedin the top 150 m in three locations in the Shelf Water, on theNova Scotia Shelf, and Slope and on Georges Bank during springand fall periods. Species were categorized as to their trophiclevel and their type of diurnal migration behaviour. The influenceof temperature, salinity, and water density on the diurnal verticaldistribution of the species was examined. Temperature was foundto have the greatest influence on the distribution of the largestnumber of species. Diurnal migration behavior of the same speciesin Shelf and Slope water and at different times of the yearwas examined. Results showed that species changed their behaviorin the two water masses, while some species changed their migrationbehavior at different times of the year. During the night inApril the most abundant copepod species, Calanus finmarchicus,making up about 80% of the biomass, was found concentrated abovethe thermocline and the main chlorophyll layer. The majorityof the less abundant species of copepods were found below thethermocline and the chlorophyll layer. At night in August thetwo most abundant copepod species, Centropoger typicus and Paracalanusparvus, making up at least 80% of the zooplankton biomass, werealso concentrated above the thermocline and the main chlorophyllLayer. Three species of copepods were concentrated at the depthof the main chlorophyll layer and two species were concentratedbelow the chlorophyll layer and thermocline. The vertical distributionof other zooplankton and ichthyoplankton species was examinedin relation to the thermocline and chlorophyll layer. Relationshipsbetween concentrations of six species of fish larvae and allspecies of copepods in the same samples showed a general increasein the numbers of larvae m^–3 as the numbers of copepodsm^–3 increased in a range of 500–4000 m^–3.However, the concentration of Merluccius bilinearis decreasedas the concentration of copepods exceeded 4000 m^–3 suggestingthat high concentrations of copepods may not be a favourableenvironment for the larvae.     

Relationship between functional response and gut transit time in the calanoid copepod Acartia clausi: role of food quantity and quality

The relationship between ingestion rate and gut transit timeof the calanoid copepod Acartia clausi was examined in laboratoryexperiments with five different diets: (i) living cells of thediatom Thalassiosira weissflogii, (ii) detrital cells of thesame diatom, (iii) 50:50 mix of the two previous diets on aprotein basis, (iv) dinoflagellate cells of Prorocentrum micansand (v) Prorocentrum minimum. Ingestion followed a Holling type2 response for diets 1 and 4 and a linear one for diets 2, 3and 5. Gut transit time varied with food abundance only whenthe copepods were fed with the living diatom. The gut evacuationrate increased with the concentration of T. weissflogii withvalues of 0.010, 0.020, 0.032, 0.042 min^–1, correspondingto gut transit time of 97, 50, 31 and 24 min, measured at 50,110, 130 and 275 µg protein L^–1, respectively. Copepodsfed with dinoflagellates, mixed and pure detrital diets exhibitedlonger and similar gut transit times ranging from 85 to 166min, depending on diet. The coupling between ingestion rateand gut transit time measurements is discussed in the contextof copepod feeding strategies.     

Diet changes in the carbon and nitrogen content of the copepod Metridia lucens

Diel changes in the near-surface (0–50 m) abundance, prosomelength, and carbon and nitrogen content of the copepod Metridialucens were measured in Deep Cove, Doubtful Sound, New Zealand(45°27'S, 167°9'E) between 3 and 6 September 1996. Metridialucens showed maximal abundance a night, suggesting a patternof normal diel vertical migration (DVM). The change in abundancesuggested that the descent of the population occurred –1h prior to dawn and the ascent –1 h after dusk. However,a proportion of the population remained near the surface duringthe day. Although there was no diel pattern in the prosome lengthof M.lucens collected near the surface, there was a marked dielcycle in the measured carbon and nitrogen contents, with maximalvalues being measured towards the end of the night prior tothe downward migration. We suggest that this diet cycle in themeasured elemental content was caused by DVM occurring morestrongly in those animals which had a better body condition,i.e. a higher elemental content per unit length.     

The importance of macrophyte bed size for cladoceran composition and horizontal migration in a shallow lake

Cladoceran composition and diel horizontal migration were studiedin 2, 10 and 25 m diameter macrophyte exclosures establishedin the littoral zone of shallow Lake Stigsholm, Denmark. Theexclosures were protected from waterfowl grazing, but open tofish. The macrophyte community cornprized Potwnogeton pectinatus,Potamogeton pusillus and Callitriche hemaphroditica. Cladoceranswere sampled randomly every third hour inside and outside themacrophyte exclosures during a 24 h period. In the 2m exclosure,the pelagic species Ceriodaphnia spp. and Bosmina spp. dominatedduring the day, mean density being as high as 3430 indiv. l^–1During the night, density decreased to 10–20% of the daytimedensity thus indicating diel horizontal migration. In the 10and 25 m exciosures, the daytime mean density of Ceriodaphniaspp. was 865 and 202 indiv. l^–1, respectively, and didnot decrease at night. In contrast to the pelagic species, thedensity of macrophyte-associated species tended to be higherin the 10 and 25 m exclosure than in the 2 m exclosure. In thedaytime, Eurycercus lamellatus density in the 2, 10 and 25 rnmacrophyte exclosures was 7, 28 and 16 indiv. l^–1 respectively,while that of Simocephalus vetulus was 11, 171 and 92 mdiv.l^–1, respectively. There was no thy-night difference inthe density of macrophyte-associated species. We conclude thatcladoceran community composition varies with macrophyte bedsize, and that the edge zone between the bed and open wateris an important daytime refuge for potentially migrating pelagiccladocerans.     

Feeding by larval and post-larval ctenophores on microzooplankton

Feeding by the coastal ctenophorc, Mnemiopsis leidyi, on microplanktonwas investigated. Larval ctenophores (tentaculate stage) grewbest and had the highest survival rates when offered a mixtureof ciliates and copepod nauplii. Larvae did not survive whenoffered phytoplankton alone. Clearing of planktonjc ciliatesby post-larval ctenophores was a function of the ciliate speciesand the size of the predator. Removal of small ciliates (<20µm in size) and phytoplankton was negligible. Small post-larvalctenophores (volume <4 cm³) had higher biovolume-specificclearing rates (0.5–1.5 1 cm^–3 day^–1) thandid larger ctenophores fed the same ciliate species. Duringin situ incubations, adult M. leidyi removed ciliates, rotifersand copepod nauplii from natural microplankton assemblages.The data indicate that non-crustacean microzooplanlctoo arean important component of the diet of larval and post-larvallocate cteoophores, particularly when copepod standing stocksare low.     

Spatio-temporal patterns in the copepod community in Malangen, Northern Norway

The objective of this study was to identify the key copepodspecies and their life cycles, and provide evidence for anyseasonal and spatial changes in the copepod community in Malangen,a fjord located 30 km to the south of Tromsø in NorthernNorway (69°30'N, 18°21'E). As a result of high levelsof freshwater run-off in May, the fjord became highly stratifiedwith a sharp pycnocline at 10–30 m depth from May to August.The generation patterns of six copepod species are described.Calanusfinmarchicus produced one generation during the spring thatyear, whereas two generations appeared to be produced by bothPseudocalanus acuspes and P.minutus: one in spring (March-June)and the other in autumn (August-December). However, it is uncertainto what extent P.minutus regularly produces a second generation.Two peaks of CI-CIII Metridia spp. were found; there were differencesalong the length of the fjord in the timing of these, but therelative contributions of M.longa and M.lucens are uncertain.Chiridius armatus CI-CIII peaked in abundance in the spring,which indicates that one main generation was produced at theouter station of the fjord. The copepod community in Malangencould be grouped into three entities according to their numericalabundance during the year one group of highly abundant forms,generally with maxima >50 000 individuals m^–3 (C.finmarchicus,Microcalanus sp., Oithona similis , Oithona spinirostris, Acartiasp. and Pseudocalanus spp.), a second group of less abundantspecies with a clear seasonality in abundance, varying from500 to 50 000 individuals m^–3 (M.longa, M.lucens, Calanushyperboreus, Carmatus, Tenwra longicornis, Oncaea sp., Euchaetanorvegica and Scolecithrwella minor), and a third group of 14holoplanktonic species, sporadically occurring in the fjord.The study demonstrates clear gradients in the abundance of fivespecies along the length of the fjord: the recruiting generationof C.finmarchicus occurred in higher abundances at the outerstation in May and June compared to the other inner sites. Laterin the season, the reverse situation appeared, in which thepopulation was more abundant in the inner part of the fjorcCalanw hyperboreus increased abruptly in abundance from lowwinter levels to a maximum in April-May, and declined steadilyduring the season (except at the innermost station). Metridialucens, M.longa and C.armatus demonstrated different distributionpatterns in Malangen that matched their preferred areas of distribution.Both M.lucens and C.armatus are known as oceanic and deep-waterspecies, respectively, and these were prevalent at the two outersites in Malangen. Metridia longa is a more nentic species andwas found in highest numbers at the two innermost sites. Themechanisms for the differences in abundance among these specieswithin the fjord are discussed.     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

Vertical distribution of pelagic chaetognaths and feeding of Sagitta enflata in the Central Equatorial Pacific

The vertical distribution and feeding of pelagic chaetognathsat 5°S, 160°W in the Central Equatorial Pacific wereinvestigated using a series of 0–500 m vertical haulswith a VMPS net over a 24 h period between 6 and 7 October 1990.The total number of individuals per haul was between 370 and688. Fourteen species in four genera were found at this station.The most abundant species was Sagitta enflata which comprised32.4–61.1% of the individuals collected from the 0–500m layer. Mesopelagic species made up 9.3–15.1% of thetotal number of individuals. Sagitta enflata and Pterosagittadraco were found in the upper part of the thermocline both byday and at night. The fraction of the population containingfood items (FCF) of S.enflata in the 0–50 m layer variedbetween 4.8 and 12.5% (mean 10.8%) and feeding activity washighest between sunrise and noon. The percentages of Copepoda,Foraminifera, crustacean larvae, Chaetognatha, Pteropoda, Ostracoda,fish and unidentified material in the gut of S.enflata were51.9,6.7,3.8,2.9,1.9,1.9 and 30.9%, respectively. Sagitta enflataconsumed food organisms which were mainly between 0.5 and 1.0mm in length. The daily feeding rate of S.enflata was 1.81 preyper individual, which was equivalent to 8.06 mg C m^–2day^–1. This corresponded to     

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     

Community structures of heterotrophic bacteria of copepod fecal pellets

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

Gut evacuation rates of Antarctic copepods during austral spring

The results of gut evacuation experiments performed on Antarctic copepods during the austral spring are presented and discussed. Four species of large copepods commonly occurring in the Indian sector of the Antarctic Ocean were studied: Calanus propinquus, Calanoides acutus, Rhincalanus gigas and Pleuromamma robusta. For each species two experiments were carried out, one in daytime and one in night-time, except for Calanoides acutus, which was only studied at night. P. robusta showed pigment gut retention in all experiments. The results showed that all species studied had a longer gut passage time than that previously recorded and that gut evacuation rate appears to decrease during daytime. Accepted: 1 October 1998     

Midwater macroplankton of British Columbia studied by submersible PISCES IV

Data are reported from 30 dives during winter and spring 1980–83at sites in the Strait of Georgia and inlets running off it,and in inlets on the west coast of Vancouver Island. Observationswere made from the surface to the bottom (maximum 733 m) butmost attention was given to the midwater plankton community.The vertical distribution and abundance of hydromedusae, siphonophores,ctenophores, euphausiids, pelagic worms and molluscs were recordedsystematically, along with data for one copepod species (Neocalanusplumchrus). The midwater environment was found to be stablein terms of species composition and depth ranges, which permittedthe data for several years and many locations to be pooled.Four categories of plankton are recognized: (a) epipelagic (concentratedin the top 50 m); (b) mesopelagic (50–175 m); (c) bathypelagic(below 175 m); and (d) meso-bathypelagic (forms living in bothmeso- and bathypelagic zones). Species in this last categorybehave like mesopelagic forms at the upper end of their ranges,migrating to the surface at night. Deeper-lying members of thesame species do not migrate. For six such species, the cut-offpoint between migratory and non-migratory components was foundto lie at a mean depth of 175 m. This depth is therefore takenas the demarcation point between the meso- and bathypelagiczones. Taking account of published data on light penetration,it is estimated that, for the whole region, daytime light intensityat 175 m, and hence the effective limit for phototaxis of thespecies in question, lies in the range 10^–8–10^–9µW cm^–2.     

Effects of prey motility and concentration on feeding in Acartia tonsa and Temora longicornis: the importance of feeding modes

Feeding experiments were conducted with the ambush-feeding copepodAcartia tonsa and the feeding-current-generating copepod Temoralongicornis. The copepods were offered a mixed diet of the dinoflagellateHeterocapsa triquetra and the ciliate Balanion comatum of similarcell size. The dinoflagellate was offered at a constant concentrationof 10–15 cells mL^–1, whereas the ciliate was offeredat a variety of concentrations, ranging from 7 to 57 cells mL^–1.Copepods with different feeding modes possess different mechanismsfor prey detection, suggesting that the two copepods would responddifferently to the two prey types. Both copepods had significantlyhigher clearance rates on the highly motile ciliate than onthe less motile dinoflagellate. In encounters between A. tonsaand its prey, we argue that this is due to the higher hydromechanicalsignal generated by the ciliate. The advection feeding copepodT. longicornis fed on the two prey according to their relativeconcentrations; in this case, we suggest that although B. comatumis capable of detecting feeding-current-generating predators,the feeding current velocity generated by T. longicornis isgreater than the escape velocity of this ciliate.     

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.     

Atlantis II Cruise: uniformity of deep copepod assemblages in the Mediterranean Sea

Studies of the community structure and biomass of deep copepodsbelow 500 m from R/V Atlantis II cruises AII49 and AII59 inMay–June 1969 and September–October 1970 denoteuniformity in deep assemblages among all major Mediterraneansubregions. Of the 65 copepod species recorded from 600–2500m, 45 were common for both Western and Eastern basins Eightof these species were dominant for all deep Mediterranean waters,comprising 77% of the deep plankton population The uniformityof deep copepod assemblages was confirmed by the Mantel testapplied to a simulated sampling procedure Only in terms of biomassdid Western and Eastern basins show appreciable regional differenceswith highest values for density and volume characterizing Westernstations ¹Died tragically at sea during the course of a cruise on December15, 1988     

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.     

The use of underwater camera in studies of vertical distribution and swimming behaviour of a sergestid shrimp, Sergia lucens

The vertical distribution and swimming behaviour of Sergia lucenswere investigated by means of underwater photography. Duringthe day the species was abundant from 3 to 7 m above the bottomon the upper continental slope at depths from 210 to 360 m.No shrimp were observed closer than 0.5 m above the bottom.The maximum concentration observed was 6.1 individuals m^–3.It is speculated that in many areas on the continental slopesand rises, the density of zooplankton and micronekton is muchhigher near the bottom in the day than in the overlying midwater,and that this may provide a concentrated pelagic food for predatorsclose to the seafloor.     

Gut evacuation rates and ingestion rates of Eudiaptomus graciloides measured by means of the gut fluorescence method

Journal of Plankton Research, 8, 973–983, 1986 FIg. 2. Time-dependent changes in the gut content (percentageof initial ng pigment) of E. gro.ciloides at different temperaturesunder simultaneous feeding. Fig. 4. The relationship between instantaneous evacuation rateand temperature of E. graciloides. The regresston equation forfeeding animals: y = 0.0044 e(0.141 ) (r² = 0.90). For comparisonthe results of non-feeding animals are indicated with open circles.     

Photoresponses of the copepod Mesocyclops edax

The predatory copepod Mesocyclops edax is an important componentof many zooplankton communities where it typically makes extensivedid vertical migrations. To describe the effect of light onadults we measured their photoresponses in the laboratory. Theresponse spectrum is characterized by a wide plateau of greatestsensitivity from about 480 – 580 nm. These animals areadapted to perceive light during the day since their regionof maximum sensitivity overlaps the spectral region of highestquantal intensity underwater (575 – 700 nm). The thresholdintensity for positive phototaxis by dark adapted animals wasabout 5 x 10^–1 Wm^–2 at 540 nm, and they were positivelyphototactic up to an intensity of 5 x 10^–1 Wm^–2.Above this intensity phototaxis is no longer observed. Light-adaptedanimals were less sensitive than dark-adapted, but their generalpattern of response to light intensity did not differ. Thereis no rhythm in phototaxis. Their photoresponses may providea mechanism for controlling vertical migration so as to minimizeexposure to planktivorous fish. ¹Contribution No. 1375-AEL from UM-CEES, Appalachian EnvironmentalLaboratory.     

Zooplankton, especially calanoid copepods, in the upper 1000 m of the south-east Arabian Sea

Abstract. Vertical distribution of zooplankton in the upper1000 m was studied from the south-east Arabian Sea in orderto determine the variations in zooplankton at different depths.The distribution and migration patterns of calanoid copepodspecies were given special attention. The mean zooplankton standingstock in the upper 1000 m was 2.1 g dry Wt m^–2, of which97.7% was concentrated in the upper 400 m. Herbivores were generallymore abundant at all depths, but did not predominate. An increasein zooplankton at night occurred in the upper 200 m, as wellas at the 600–1000 m stratum. The maximum diversity ofcalanoid copepods also coincided with these two depths. Basedon vertical ranges, the calanoid copepod species were assignedto three groups: (i) species occurring predominantly in theepipelagic layer and forming the bulk of the calanoids; (ii)relatively sparser deeper living species confined below 200m; and (iii) species occurring throughout the water column.Some amount of vertical niche separation among congeneric specieswas indicated. While some species showed active migration, agood number of species were non- migratory.