State-dependent vertical distribution of the carnivore copepod Pareuchaeta norvegica

We explored the impact of body condition on vertical distributionof the carnivore copepod Pareuchaeta norvegica. Individualssampled in the upper part of the water column had lower dryweight and slightly shorter prosome than individuals sampleddeeper. Non-ovigerous females occurred higher in the water columnthan ovigerous females. Our results suggest that ovigerous femalesand individuals with large energy reserves to a large extentprioritize predator avoidance in deep water versus feeding inthe upper part of the water column compared to non-ovigerousfemales and individuals with low energy reserves. Adult femalescollected closest to the surface were less pigmented than thosefound deeper. This, however, proved to be a quickly adjustabletrait depending on prevailing light conditions.     

Changes with time after capture in the metabolic activity of the carnivorous copepod Euchaeta norvegica Boeck

The rates of respiration and ammonium excretion of the carnivorous copepod Euchaeta norvegica Boeck decreased by 40 and 70%, respectively, during the first ≈10 h after capture. ETS (electron transport system) activity decreased in parallel with the rate of respiration. Increasing O/N and decreasing N/P atomic ratios during incubation indicated a change from protein to lipid metabolism, suggesting that starvation was the major factor responsible for the depression of the metabolic rates after capture. The adenylate energy charge (EC) ratio was significantly lower during the initial 13 h after capture than subsequently, caused mainly by an elevated AMP level. The total content of adenine nucleotides increased during this period of lowered EC. Starvation could also have been responsible for these changes in the adenine nucleotide levels through degradation of RNA. Our earlier interpretation that the lowered EC values after capture reflected capture stress has not been confirmed. The lactate level of E. norvegica just after capture was low (≈0.1 μg·mg dry wt^?1) and thus not indicative of any oxygen debt.     

Zonal distribution and seasonal vertical migration of copepod assemblages in the Scotia Sea

Large, biomass-dominant Southern Ocean copepod species have been much studied, but small and mesopelagic species also play major rôles in these ecosystems. However, little is known of some basic aspects of their ecology. To address this, the abundances of 23 copepod species and genera were analysed from 72 stations sampled during the Discovery Expeditions in the 1920s to 1950s. Stratified net samples, usually to a depth of 1000?m, provided year-round coverage in the Scotia Sea from the Subantarctic Front to the Weddell-Scotia Confluence. Small copepods (Microcalanus pygmaeus, Ctenocalanus spp., Oncaea spp. and Oithona spp.) formed ～75% of total copepod abundance in the top 1000?m across all major zones. Oithona spp. composed ～40% of copepod numbers in the Polar Front area and to its south: further north their importance declined. All mesopelagic taxa except for the warmer-water species Metridia lucens and Pleuromamma robusta, extended throughout the entire study area, with smaller regional differences than for the shallower-living species. The species showed a continuum of temperature ranges, and there was no evidence that the Polar Front was a major biogeographic boundary to their distribution. Indeed, several important species, including Oithona spp. (mainly Oithona similis), Ctenocalanus spp., Metridia lucens and Rhincalanus gigas reached maximum numbers in this area. Total copepod abundance was thus higher in the vicinity of the Polar Front than in any other region. Only two copepod families made pronounced seasonal vertical migrations: Eucalaniidae (Eucalanus longiceps and R. gigas) and Calaniidae (Neocalanus tonsus, Calanoides acutus, Calanus simillimus and Calanus propinquus). Some evidence for a winter descent was found for Ctenocalanus spp. and some deeper-living groups: Euchaeta spp. and the Metridiidae, although their migrations were not so great as for the eucalanids and calanids.     

Predator-induced diet vertical migration in a planktonic copepod

Predator evasion is the most commonly hypothesized reason fordiel vertical migrations undertaken by a wide variety of planktonicorganisms in lakes and seas, yet direct evidence remains elusive.We tested the predation hypothesis by exposing enclosed populationsof a marine copepod Acartia hudsonica to caged or free-rangingindividuals of their natural predator, the planktivorous fishGasterosteus aculeatus. After little more than a week, adultcopepods changed their vertical distribution and migration behaviordepending on the presence or absence of predation. Only free-rangingfish induced vertical migration in the copepod population. Cagedfish had no effect, indicating that vertical migration was nota simple chemically mediated response of copepods to the predator.Rather, copepods seemed to react to the presence of predatorsby other means, perhaps visual or mechanical stimuli, and toexhibit a downward escape response which, because encounterswith visually orienting fish occur chiefly in the daytime, effectivelylimited the copepods' occurrence in the upper water column tothe night-time hours. Alternatively, because fish imposed heavymortality on copepods, it is possible that selective predationaltered the proportions of individuals with fixed, geneticallydetermined migration behaviors. We suggest experiments to distinguishthese alternatives.     

The fate of discarded appendicularian houses: degradation by the copepod, Microsetella norvegica, and other agents

Despite the potential importance of zooplankton in degradationof marine snow, the association of colonising zooplankton withdiscarded appendicularian houses has not been investigated innorthern areas. We sampled the vertical distributions of appendicularians,houses and potential zooplankton colonisers at two stationsin the central North Sea during late summer. In addition, grazingexperiments were performed with the copepod Microsetella norvegica,which was assumed to be the main contributor to house degradation.The results were used in (i) inverse modelling, to estimatethe factors which were significant in shaping the vertical distributionof houses and (ii) calculations to estimate potential housedegradation rates. M. Norvegica was able to feed on appendicularianhouses, with feeding rates up to 0.42 g C (g C)^–1 day^–1(0.14 µg C ind.^–1 day^–1). The model resultssuggested that the vertical distribution of houses was shapedby sinking of houses, bacterial degradation and feeding of M.norvegica and invertebrate larvae. The estimated community degradationrate by M. norvegica was low, whereas invertebrate larvae haddegradation rates close to bacterial degradation. We concludethat at the typical concentrations of M. norvegica in the NorthSea (10⁴ ind. m^–2), its role in marine snow degradationis likely to be small. Degradation by other zooplankton groups,such as invertebrate larvae, can, however, be substantial.     

Importance of the large copepod Paraeuchaeta antarctica (Giesbrecht, 1902) in coastal waters and the diet of seabirds at Kerguelen, Southern Ocean

The importance of the euchaetid copepod Paraeuchaeta antarcticain the subantarctic pelagic ecosystem was quantified in thecoastal waters of the Golfe du Morbihan at Kerguelen Islandsby comparing food samples from two diving seabirds with concurrentnet samples taken within the predator foraging area. Paraeuchaetaantarctica occurred in very high densities (up to 30 individualsm^-3 and 96 mg dry weight m^-3) in the water column, being moreabundant in the deepest part of the gulf than in shallow watersor at the more offshore shelf stations. The common diving petrelfeeds almost exclusively on crustaceans, its diet being dominatedby the amphipod Themisto gaudichaudii (52% by number and 84%by reconstituted mass) and P. antarctica (33% and 16%, respectively).Rockhopper penguins preyed upon crustaceans and fish, with fourtaxa being important, namely T. gaudichaudii (37% and 23%, respectively),Euphausia vallentini (24% and 41%), postlarval fish (10% and24%), and P. antarctica (13% and 3%). Paraeuchaeta antarcticadominated numerically in 21% of the diving-petrel food samplesand in 12% of penguin samples. The two bird species segregatedby preying upon different developmental stages of P. antarctica,diving petrels fed equally on CV of both sexes and CVI, whilepenguins fed on CVI only. Comparison of P. antarctica foundin net and food samples indicated no prey selection by commondiving petrels that caught the different copepod stages in proportionto their availability in the water column. On the other hand,the diving performance of penguins, which is better than thatof the petrels (mean maximum dive depths 69 m for penguins versus32 m in petrels), allows them to catch CVI in deeper waters,probably near the bottom. Our study shows that P. antarcticais a major component of the coastal macrozooplankton communityand a significant prey for two species of diving seabirds inhabitingKerguelen. This is also the first record of a copepod speciesas a prey for penguin, and the first to highlight P. antarcticain the food of austral seabirds.     

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.     

Hypoxia tolerance in the copepod Calanoides carinatus and the effect of an intermediate oxygen minimum layer on copepod vertical distribution in the northern Benguela Current upwelling system and the Angola-Benguela Front

There is a growing concern that hypoxic and anoxic areas in the sea spread in extent and intensity, posing a severe risk to marine ecosystems and fisheries. Hypoxia may affect fish stocks directly or via detrimental effects on important prey species, such as zooplankton. A unique feature of the northern Benguela Current upwelling region and Angola-Benguela frontal system is a pronounced intermediate oxygen minimum layer (IOML) at 60-500 m depth with oxygen concentrations ≤ 1.4 mg O₂l^− 1 (minimum < 0.7 mg O₂l^− 1). Field studies during February-March 2002 demonstrated that the abundance of calanoid copepods and the biomass of mesozooplankton in general were severely reduced within the IOML. The dominant copepod Calanoides carinatus showed a bimodal vertical distribution with parts of the population either comprising all developmental stages concentrated in the surface layer (0-60 m), or copepodids C5 diapausing below 400 m depth apparently avoiding the IOML. Accordingly, abundances of other calanoid copepods were higher at the surface and below 300 m than in the centre of the IOML. The scarcity of planktonic life within the IOML raises the question whether this layer represents an effective barrier for zooplankton vertical migrations. Especially in C. carinatus, ontogenetic vertical migration plays a key role in the retention of the population within the productive upwelling region and for the rapid re-colonisation of plumes of newly upwelled water. To address this issue, the hypoxia tolerance of C. carinatus was determined in a series of laboratory-based, closed-bottle experiments in January 2005. Copepods were kept in gas-tight bottles and the decreasing oxygen concentrations were monitored to establish their minimum oxygen demands. Although copepodids survived apparently unharmed at surprisingly low oxygen concentrations of ca. 1.5 mg O₂l^− 1, they could not tolerate oxygen levels < 1.1 mg O₂l^− 1, implying that the core of the IOML, where O₂ concentrations are below this threshold, is uninhabitable for C. carinatus. In contrast, the IOML may represent a refuge from competition and predation for other copepod species specifically adapted to hypoxic environments.     

Spatial and temporal distribution of copepod communities in the Adriatic Sea

The distribution of planktonic copepods in the Adriatic Seahas been analyzed on the basis of 132 samples collected at 35fixed stations during 4 seasonal cruises. A total of 127 specieshave been determined and 3 characteristic copepod communitieshave been identified. The distribution of copepod species hasbeen found to reflect the dual physiognomy of the Adriatic.The shallow northern section is characterized by high densityvalues, low species diversity, and the dominance of speciesbelonging to the estuarine and coastal communities. The relativelydeep waters of the southern section are characterized by lowdensity values, high species diversity, and the presence ofmany species belonging to the upper, middle, and lower zonesof the oceanic community. Temporal variations in the horizontalspread of these 3 communities are discussed in relation to seasonalcycles in abundance, vertical migration patterns, and the currentregime in the Adriatic.     

Seasonal size distribution of developmental stages of sub-antarctic copepod

Drepanopus pectinatus, the most numerous species of Copepod in the Kerguelen Archipelago completes 4 generations a year. The succession of generations follows the hydrological seasons. Each generation may be characterized by the cephalothorax length of the developmental stages C1 to C6. The mean lengths of stages C2 to C6 are shown to be significantly different (P > 0.05) between seasons (i.e. between generations). The size-classes have a unimodal distribution from stages C1 to C4. For copepodite C5 and adults, bimodal histograms reflect the differentiation of males and females.A reverse trend of the development pattern of lengths is demonstrated in this subantarctic species: a decrease with decreasing temperatures (6° to 2°) from April to September, and an increase with the rising temperatures (2° to 7°) from October to February.Two populations of females are distinguishable in spring and summer, by their small (0.950–1.299 mm length of cephalothorax, 55%) or large (1.300–1.800 mm, 45%) size, respectively.The differences in mean length cannot be explained by their relation to the mean temperature alone. Pigment content is suspected to play a role.     

Diel vertical migrations ofMeganyctiphanes norvegica in the Kattegat: Comparison of net catches and measurements with Acoustic Doppler Current Profilers

Diel vertical migration of a stable and well-defined population of Nordic krill,Meganyctiphanes norvegica (Crustacea, Euphausiacea) was investigated during eight days in August 1989, in the Läsö-Deep, East of the Danish island Läsö. Net catches with a multi-net (MOCNESS) and measurements with a moored and a shipboard Acoustic Doppler Current Profiler (ADCP) were compared. Backscattered energy as a measure for biomass gave good correlations to the dry weight ofM. norvegica and smaller zooplankton from net catches. Diel migratory patterns matched well, as determined, parallel with both methods. Migratory vertical velocity was determined with ADCP at 2–3 cm sec^?1. The potential for the use of ADCPs for biological investigation is discussed. Vertical migration was dependent on environmental parameters. The krill did not cross a temperature barrier of 14°C, although rich food sources were situated beyond it. Differences in salinity did not play a role. Currents were involved in plankton distribution. Light was an important Zeitgeber (synchronizer) and determined the density of the krill aggregations. Feeding behaviour did not interfere with the light-induced migratory pattern of Nordic krill at the Läsö-Deep.     

Implications of the feeding current structure of Euchaeta rimana, a carnivorous pelagic copepod, on the spatial orientation of their prey

Many marine planktonic organisms create water currents to entrainand capture food items. Rheotactic prey entrained within thesefeeding currents often exibit escape reactions. If the directionof escape is away from the feeding current, the prey may successfullydeter predation. If the escape is towards the center of thefeeding current, the prey will be re-entrained towards its predatorand remain at risk of predation. The direction of escape isdependent on (i) the ability of the prey to escape in a directiondifferent than its pre-escape orientation and (ii) the orientationcaused by the interaction of the prey's body with the movingfluid. In this study, the change in orientation of Acartia hudsonicanauplii as a result of entrainment within the feeding currentof Euchaeta rimana, a planktonic predatory copepod, was examined,When escaping in still water, A.hudsonica nauplii were ableto vary their pre-escape direction by only 10. This allowsonly a limited ability to escape in a direction different thantheir pre-escape orientation. Analyses of the feeding currentof E.rimana show the flow speed to be most rapid in the centralregion with an exponential decrease in speed distally. In contrast,flow vorticity is minimal in the center of the feeding currentand maximal at 1.75 mm along the antennae. As a result, thedegree of rotation of the prey towards the center of the feedingcurrent shows a strong dependency on the prey's location withinthe feeding current. The feeding current of E.rimana rotatedthe prey 14 when near the center of the flow field and up to160 when located more distal in the feeding current Since theprey's escape abilities cannot compensate for the rotation dueto the flow, this mechanism will maintain the escaping preywithin the feeding current of their predator. Therefore, thefeeding current facilitates predatory copepods in capturingprey by (i) increasing the amount of water which passes overtheir sensors and through their feeding appendages and (ii)controlling the spatial orientation of their prey prior to escape.     

Effect of vertical mixing on the vertical distribution of copepods in coastal waters

The vertical distribution of copepod adult stages and naupliihad been studied in a coastal water before and during homogenizationof the water column due to wind. When the water column was stratified,the vertical distribution of the zooplankton was also stratified,as is generally described in the literature. During homogenizationof the water column, different patterns of vertical distributionwere observed. The less active nauplius stage was evenly distributed,even though adult stages always exhibited a stratified distribution.Oithona similis and Microsetclla norvegica exhibited the samedistribution in stratified and unstratified water columns. Temoralongicomis and Pseudocalanus sp. were deeply distributed ina less turbulent area. The causes of these changes are discussed.     

The effect of lunar eclipse on the vertical migration behaviour of Meganyctiphanes norvegica (Crustacea: Euphausiacea) in the Ligurian Sea

The vertical migration of a zooplankton community dominated by the euphausiid Meganyctiphanes norvegica was monitored between 16 and 23 September 1997 with a 153 kHz Acoustic Doppler Current Profiler (ADCP) and a MOCNESS net. The sampling period covered a phase in the lunar cycle when the rise of the moon (full moon) coincided initially with sunset and then became progressively later. On 16 September 1997, a lunar eclipse occurred 45 min after sunset, lasting for 2 h. At dusk, the ADCP observed the upward vertical migration of two principal backscattering bands 10 min apart with vertical velocities of up to 7 cm s^-1. After a period at the surface, a more diffuse band subsequently sank at a slower rate (1-2 cm s^-1) to a depth of 75-100 m. Net samples showed that the earlier band consisted mainly of the pteropod Cavolinia inflexa, whilst the later band was mostly euphausiids, predominantly M.norvegica. This species was also the major constituent of the band that sank. The timing of upward migration was relatively constant over the sampling period, but there was an increasing delay of the secondary sinking until 21 September. This showed as a strong correlation between the onset of sinking and the time of moonrise. The lunar eclipse on 16 September perturbed this pattern, such that animals did not sink soon after their arrival at the surface, as occurred on 17 September but remained at the surface until the end of the umbra. This suggests that M.norvegica can perceive moonlight and that this influences vertical migration. Evidence that the behaviour is not solely mediated by this erogenous factor, however, is seen in the pattern that emerged after 21 September, when midnight sinking occurred at a relatively constant time after sunset and before moonrise. These observations support the hypothesis that moonlight is a Zeitgeber for an endogenous rhythm that synchronizes secondary sinking behaviour with the lunar cycle.      

Bi-modal regulation of a formin by srGAP2

The maintenance of rapid and efficient actin dynamics in vivo requires coordination of filament assembly and disassembly. This regulation requires temporal and spatial integration of signaling pathways by protein complexes. However, it remains unclear how these complexes form and then regulate the actin cytoskeleton. Here, we identify a srGAP2 and formin-like 1 (FMNL1, also known as FRL1 or FRLα) complex whose assembly is regulated by Rac signaling. Our data suggest srGAP2 regulates FMNL1 in two ways; 1) Rac-mediated activation of FMNL1 leads to the recruitment of srGAP2, which contains a Rac-specific GAP domain; 2) the SH3 domain of srGAP2 binds the formin homology 1 domain of FMNL1 to inhibit FMNL1-mediated actin severing. Thus, srGAP2 can efficiently terminate the upstream activating Rac signal while also opposing an important functional output of FMNL1, namely actin severing. We also show that FMNL1 and srGAP2 localize to the actin-rich phagocytic cup of macrophage-derived cells, suggesting the complex may regulate this Rac- and actin-driven process in vivo. We propose that after Rac-dependent activation of FMNL1, srGAP2 mediates a potent mechanism to limit the duration of Rac action and inhibit formin activity during rapid actin dynamics.     

Predation by a carnivorous marine copepod, Euchaeta elongata Esterly, on eggs and larvae of the Pacific hake, Merluccius productus

The predatory behavior of the carnivorous marine copepod, adultfemale Euchaeta elongata Esterly, feeding on eggs and larvaeof the Pacific hake, Merluccius productus, was examined in thelaboratory and in a natural setting. E. elongata did not feedon eggs. Predation on larvae is believed to depend on larvalswimming behavior: (1) predation rates were low on early stageyolk-sac larvae which are inactive swimmers and are relativelyundetected by the predator; (2) rates were high on middle stageyolk-sac larvae which are more active swimmers yet have a poorlydeveloped escape response; and (3) rates were low on largerlarvae which are able to escape the predator effectively. Starvedhake larvae were more vulnerable to predation due to a poorescape ability although they were less active and not easilydetected. The presence of naturally occurring alternative prey,Pseudocalanus sp., depressed the rate of E. elongata predationon hake larvae. In an analysis of field data, hake larvae andE. elongata were found to occupy the same depths in Dabob Bay.A high percentage of E. elongata collected had apparently beenfeeding on hake larvae, as indicated by the presence of pigmentsin their guts. Survival of hake larvae in late spring appearsto be relatively poor compared with early spring; poorer survivalin late spring may be due partly to an increase in the abundanceof invertebrate predators, such as E. elongata. ^*Current address: Northwest and Alaska Fisheries Center, 2725Montlake Blvd, Seattle, WA 98112, USA.     

Spatial and temporal variations in calanoid copepod distribution in the Straits of Malacca

The distribution and abundance of planktonic calanoid copepods were studied from samples collected at 13–20 stations during four oceanographic cruises (pre- and post- monsoons, and during northeast (NE) and southwest (SW) monsoons) performed between 1998 and 2000 in the Straits of Malacca. Space and time variations of calanoid copepods were described using univariate (number of species, diversity indices, abundance) as well as multivariate (MDS, ANOSIM, SIMPER) techniques from the Plymouth Routines in Multivariate Ecological Research (PRIMER) package. There were significant differences in abundance between the cruises. k-Dominance curves also revealed significant differences in the relative species abundance distributions among the monsoon periods, and a decrease in diversity from northern to southern parts of the Straits during each cruise. Multi-dimensional scaling revealed four groups of abundances with differences in species composition. Evidence from analysis of similarity (ANOSIM) suggested that the differences in communities among monsoon periods were significant, although spatial differences among samples in geographic locations in the northern, central and southern parts of the Straits were insignificant. These differences resulted from an overall change in the balance of relative abundance of few dominant species, rather from changes of many species. Similarity percentage analyses (SIMPER) indicated that the major species contributing to the average dissimilarity between monsoons varied temporally.