Fine-scale observations of the predatory behaviour of the carnivorous copepod <Emphasis Type="Italic">Paraeuchaeta norvegica</Emphasis> and the escape responses of their ichthyoplankton prey,Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>)

Paraeuchaeta norvegica (8.5 mm total length) and yolk-sac stage Atlantic cod larvae (4 mm total length) (Gadus morhua) larvae were observed in aquaria (3 l of water) using silhouette video photography. This allowed direct observations (and quantitative measurement) of predator–prey interactions between these two species in 3-dimensions. Tail beats, used by cod larvae to propel themselves through the viscous fluid environment, also generate signals detectable by mechanoreceptive copepod predators. When the prey is close enough for detection and successful capture (approximately half a body-length), the copepod launches an extremely rapid high Reynolds number attack, grabbing the larva around its midsection. While capture itself takes place in milliseconds, minutes are required to subdue and completely ingest a cod larva. The behavioural observations are used to estimate the hydrodynamic signal strength of the cod larva’s tail beats and the copepod’s perceptive field for larval fish prey. Cod larvae are more sensitive to fluid velocity than P. norvegica and also appear capable of distinguishing between the signal generated by a swimming and an attacking copepod. However, the copepod can lunge at much faster velocities than a yolk-sac cod larva can escape, leading to the larva’s capture. These observations can serve as input to the predator–prey component of ecosystem models intended to assess the impact of P. norvegica on cod larvae.     

Correlation between photosensitivity and downwelling irradiance in mesopelagic crustaceans

The current study determined behavioral and electrophysiological photosensitivities for three species of mesopelagic crustaceans: Pasiphaea multidentata Esmark, 1866 (Decapoda: Pasiphaeidae), Sergestes arcticus Kröyer, 1855 (Decapoda: Sergestidae), and Meganyctiphanes norvegica M. Sars, 1857 (Euphausiacea: Euphausiidae). In addition, in situ quantifications of the species vertical distributions in relation to downwelling irradiances were also determined in two locations in the northwest Atlantic Ocean, Wilkinson Basin (WB) and Oceanographer Canyon (OC). Data are from six 2-week cruises between June and September from 1995 to 2001. P. multidentata and M. norvegica were the most abundant large crustaceans in WB, and S. arcticus and M. norvegica were the most abundant large crustaceans in OC. The behavioral light sensitivity thresholds of P. multidentata and M. norvegica from WB were both 10⁷ photons cm^–2 s^–1 and those of S. arcticus and M. norvegica from OC were both 10⁸ photons cm^–2 s^–1. Electrophysiologically, P. multidentata was significantly more sensitive than M. norvegica from either location, S. arcticus was significantly more sensitive than M. norvegica from OC, and M. norvegica from WB was significantly more sensitive than M. norvegica from OC. A correlation was found between electrophysiologically measured photosensitivity and downwelling irradiance, with the most sensitive species, P. multidentata and S. arcticus, associated with the lowest irradiance at daytime depths. The photosensitivities of M. norvegica collected from the clearer waters of OC were significantly lower than those of individuals collected from the more turbid WB waters. These results indicate that downwelling irradiance has a significant impact on interspecies and intraspecies vertical distribution patterns in the mesopelagic realm.Communicated by J.P. Grassle, New Brunswick     

Feeding behavior of the Atlantic mackerel Scomber scombrus on the hydromedusa Aglantha digitale

Under laboratory conditions, the Atlantic mackerel Scomber scombrus feed on Aglantha digitale, a small, transparent medusa common in temperate and boreal waters. The same feeding response to A. digitale was observed whether the medusae were offered alone or together with a mixture of copepod prey. A. digitale mortality in the path of the mackerel school is estimated to be near 100%. Median carbon and nitrogen content of A. digitale was equivalent to about ten small copepods (e.g. Pseudocalanus sp.). A. digitale is abundant in temperate-boreal waters and may be an important source of nutrition to mackerel in the northwest Atlantic.     

Cross-shelf distribution of copepods and fish larvae across the central Great Barrier Reef

The distribution of total dry weight of zooplankton, copepod numbers and ichthyoplankton across the outer continental shelf in the central Great Barrier Reef was examined at bi-weekly intervals for three months over summer of 1983. Copepods were sampled (236 m net) within 10 m of the surface and within 10 m of the bottom. Mean densities in surface waters decreased markedly from the mid-shelf to outer shelf and the Coral Sea, but no cross-shelf gradient occurred in the bottom-water. Densities of copepods on the mid-shelf (surface and bottom waters) and in bottom-waters of the outer shelf were typically ca. 400 m^–3. Significantly lower densities (ca. 100 m^–3) occurred in surface waters of the outer shelf, except during outbursts of Acartia australis, when densities in these waters differed little from those elsewhere on the shelf. In oceanic waters, 10 km from the outer shelf station, copepod densities in surface waters were ca. 40 m^–3. Four of the five most abundant copepod taxa in surface waters, Paracalanus spp., Eucalanus crassus, Acrocalanus gracilis and Canthocalanus pauper, tended to be most abundant at the mid-shelf end of the transect. Acartia australis was sporadically very abundant in surface waters of the outer shelf, as was Paracalanus spp. in bottom-water of the outer shelf. An assemblage of Coral Sea species of copepod occurred in bottom-water of the outer shelf during two major intrusions, but not at other times. Densities of all common species varied considerably between cruises. Maximum densities of all common species except A. australis tended to be associated with diatom blooms linked to intrusions but a bloom did not necessarily mean all common species were abundant. Fish larvae included both reef and non-reef taxa, with reef taxa predominating on the outer shelf (approx 2:1 in density of individuals) and non-reef taxa dominating in nearshore samples (approx 2:1). Nine of the ten most abundant taxa analysed showed highly significant variation in numbers among stations and all but one of these also exhibited significant station x cruise interactions. Interactions generally reflected changes in the rank importance of adjacent stations from one cruise to the next or lack of any significant cross-shelf variation on some cruises where overall abundance of the taxa was low.     

Ecology of the euphausiidsThysanoessa raschi,T. inermis andMeganyctiphanes norvegica in Ísafjord-deep,northwest-Iceland

The seasonal abundance, distribution, maturity, growth and population dynamics of the euphausiidsThysanoessa raschi (M. Sars, 1864),T. inermis (Krøyer, 1846) andMeganyctiphanes norvegica (M. Sars, 1857) were studied in Ísafjord-deep, a fjord in northwest Iceland, from February 1987 to February 1988. Sampling was made at nine stations along the length of the fjord at approximately monthly intervals, along with hydrographic measurements and water sampling for nutrient analysis and measurements of chlorophylla concentrations. Spring warming of the water began in late May and maximum temperatures (8° to 10°C) were observed in late July–September. The phytoplankton spring-bloom started in early April, and the highest chlorophylla levels were measured in early May (7.0 mg m^–3). A small increase was observed in the chlorophylla content in August. The greatest abundance of juveniles and males and females of all three species was observed during January and February 1988, during which period the euphausiids were concentrated in the middle and inner parts of the fjord. Euphausiid eggs were first recorded in the plankton in mid-May, and the greatest abundance ofThysanoessa spp. larvae occurred at the end of May. Larvae ofM. norvegica were not observed in Ísafjord-deep, indicating that recruitment of this species was occurring from outside the fjord.T. raschi andT. inermis had a life span of just over 2 yr; the life span ofM. norvegica was more difficult to determine. Almost all femaleT. raschi were mature at the age of 1 yr, while mostT. inermis females appeared not to mature until 2 yr of age. Most males of both species took part in breeding at 1 yr of age. The maximum carapace length ofT. raschi andT. inermis was 8 to 9 and 9 to 10 mm, respectively. The largestM. norvegica had a carapace length of 9 to 10 mm. The spawning of the euphausiids in Ísafjord-deep appeared to be closely related to the phytoplankton spring bloom; water temperature appeared to have no influence on spawning.     

Vertical distribution and mortality of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during overwintering in oceanic waters southwest of Iceland

The seasonal vertical distribution and the predatory regime encountered by Calanus finmarchicus were studied along a transect across the Reykjanes Ridge in the oceanic area southwest of Iceland from data collected during four cruises between November 1996 and June 1997. The mortality for the overwintering period was estimated using linear regressions of density estimates from November 1996 to April 1997. In addition, we also estimated the mortality of the oldest population stages (C4 and older) in April and June by applying the vertical table method. During winter (November/December–January/February), the animals mainly resided at a depth of ∼300–1,500 m in the water of Atlantic origin. Ascent to upper layers took place mainly during March and April, and continued until May. During all cruises, continuous deep-scattering layers were observed, mainly within the range of 400–500 m to 700–800 m depth. Based on sampling with a Harstad pelagic trawl in April, the scattering was mainly ascribed to jellyfish (mainly Periphylla periphylla), small mesopelagic oceanic fishes (several species but Benthosema glaciale and Maurolicus muelleri were most abundant), euphausiids (mainly Meganyctiphanes norvegica) and shrimps (mainly Sergestes arcticus). These species may represent a predatory threat to overwintering C. finmarchicus. From November to April, daily per capita mortality rates were estimated to be (mean ± 95% CL) 0.004 ± 0.0028 for the total data set, and 0.004 ± 0.0033 and 0.004 ± 0.0023 (day^-1) for the Iceland Basin and Irminger Basin, respectively. Mortality rates were higher later in life (mean ± 95% CL) for C5/females (0.13 ± 0.044) and C5/males (0.19 ± 0.051) than for C4/C5 (0.00 ± 0.035) when averaged over all samples taken in April and June 1997. We discuss how the observed distribution and mortality rates of overwintering C. finmarchicus might be related to predatory regime.     

Nodularin concentrations in Baltic Sea zooplankton and fish during a cyanobacterial bloom

Toxic cyanobacterial blooms, dominated by Nodularia spumigena, are a recurrent phenomenon in the Baltic Sea during late summer. Nodularin, a potent hepatotoxin, has been previously observed to accumulate on different trophic levels, in zooplankton, mysid shrimps, fish as well as benthic organisms, even in waterfowl. While the largest concentrations of nodularin have been measured from the benthic organisms and the food web originating from them, the concentrations in the pelagic organisms are not negligible. The observations on concentrations in zooplankton and planktivorous fish are sporadic, however. A field study in the Gulf of Finland, northern Baltic Sea, was conducted during cyanobacterial bloom season where zooplankton (copepod Eurytemora affinis, cladoceran Pleopsis polyphemoides) and fish (herring, sprat, three-spined stickleback) samples for toxin analyses were collected from the same sampling areas, concurrently with phytoplankton community samples. N. spumigena was most abundant in the eastern Gulf of Finland. In this same sampling area, cladoceran P. polyphemoides contained more nodularin than in the other areas, suggesting that this species has a low capacity to avoid cyanobacterial exposure when the abundance of cyanobacterial filaments is high. In copepod E. affinis nodularin concentrations were high in all of the sampling areas, irrespective of the N. spumigena cell numbers. Furthermore, nodularin concentrations in herring samples were highest in the eastern Gulf of Finland. Three-spined stickleback contained the highest concentrations of nodularin of all the three fish species included in this study, probably because it prefers upper water layers where also the risk of nodularin accumulation in zooplankton is the highest. No linear relationship was found between N. spumigena abundance and nodularin concentration in zooplankton and fish, but in the eastern area where the most dense surface-floating bloom was observed, the nodularin concentrations in zooplankton were high. The maximum concentrations in zooplankton and fish samples in this study were higher than measured before, suggesting that the temporal variation of nodularin concentrations in pelagic communities can be large, and vary from negligible to potentially harmful.     

Zooplankton: pre-southwest and northeast monsoons of 1993 to 1994, from the North Arabian Sea

Zooplankton samples from the North Arabian Sea Environment and Ecosystem Research (NASEER) cruises were analyzed to determine the basic taxonomic composition, biomass (standing stock) and the total and copepod numeric abundance; these characteristics are discussed with reference to the different monsoon periods. Cruises carried out during March 1993 and May 1994, categorized as pre-southwest monsoon periods, and a cruise in December 1994, categorized as a northeast monsoon period, are discussed in detail. The biomass of January 1992 versus August 1992 and August 1992 versus March 1993 differed significantly (F = 6.44, P≤ 0.05). Ranges of highest and lowest biomass from each cruise are also given. Distinct “high” and “low” production areas of statistically significant difference (F = 12.67, P≤ 0.05) were observed. The “high” and “low” production areas were mobile and followed the surface wind circulation patterns (wind reversal pattern) during the northeast and southwest monsoon periods. Overall zooplankton showed a patchy distribution. The overall zooplankton abundance and total copepod counts differed significantly between the Cruises 3 versus 4 and 4 versus 5 (F = 15.67, P≤ 0.05 and F = 34.39, P≤ 0.05, respectively). There was no significant difference (P≥ 0.05) in biomass, between eutrophic and oligotrophic stations, suggesting no difference between near shore and offshore waters. Thirty-eight taxonomic groups were identified from the samples, with copepods being the most dominant group, followed by chaetognaths and siphonophores. Copepods constitute an average of 52.50 to 74.93% of the total zooplankton count and reach maxima of 92.14% of the total zooplankton count at the outset of the southwest monsoon (March 1993) and 91.39% at the outset of the active northeast monsoon (December 1994). Received: 27 February 1998 / Accepted: 8 October 1999     

Feeding behavior of adult <Emphasis Type="Italic">Vinciguerria nimbaria</Emphasis> (Phosichthyidae), in the tropical Atlantic (0°–4°N, 15°W)

Adult Vinciguerria nimbaria are the main prey of tuna during the tuna fishing season (late autumn and winter) in the equatorial Atlantic (0–4°N, and ~15°W). V. nimbaria trophic behavior in the fishing grounds was studied in relation to hydrobiological factors to determine its role in the trophic food web. Sampling stations spaced by 20 nautical miles were set up along a 15°W north–south transect from 4°N to 0°40S. At each station, the temperature and vertical fluorescence profiles were recorded. Nitrate and chlorophyll a analyses were performed on water sampled at different levels in the euphotic zone. Vertical plankton hauls were carried out at depths of 0–100 and 0–200 m using a standard WP2 net fitted with a 200-μm mesh gauze. Vinciguerria nimbaria adults were collected using a young-fish mid-water trawl net (10 × 15 m opening mouth, 10 mm cod end mesh). The weight of the stomach contents, the stomach fullness index, the number of prey, the frequency of occurrence and the prey preponderance were recorded for 20 fish from each haul. An oligotrophic typical tropical structure (TTS) was found between 1° and 4°N where small zooplankton was relatively abundant above or near the thermocline. In the TTS, V. nimbaria behaved as an epipelagic fish, feeding on the dominant small prey during the daytime. In turn, it was a prey for tuna. In the equatorial zone, where zooplankton was more abundant than in the north equatorial zone, V. nimbaria behaved as a mesopelagic fish and as an opportunistic mesozooplankton feeder. It consumed a wide range of sizes of food, feeding on the most abundant species of zooplankton as well as the largest zooplankton species, possibly while migrating towards the surface in the late afternoon or in the deep layer.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Spatial and temporal feeding patterns of copepods from the North Pacific centra gyre

Spatial and temporal feeding patterns (determined from an index of gut fullness) are described for 10 typical species of calanoid copepods collected from the North Pacific central gyre (September 1968 to June 1977), an area where the zooplankton is food limited and there were a-priori reasons to suspect that feeding and competition for food were important in regulating zooplankton community structure. Over 100 samples from 11 cruises to the eastern part of the gyre were examined, and patterns of gut fullness were related to environmental variables and the copepod species structure. The copepods studied all tended to be omnivores and food generalists. Males had lower indices of gut fullness than females but both males and females of a species had similar spatial and temporal feeding patterns. Guts were usually fuller at night than during the day, even in nonmigrating species; however, within nighttime depth distributions, no depths were preferred for feeding. There were also differences between species in mean gut fullness, but different species tended to have similar spatial and temporal feeding patterns. There was considerable spatial variability, and locales could be identified in which most species had higher indices of gut fullness. The copepods were not necessarily more abundant in these locales, nor did these tend to be areas of above average chlorophyll concentration. These patterns were consistent with relatively nonselective feeding, and there was no evidence that these species separate their niches by feeding at differing places or times.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Zooplankton responses to hydrological and trophic variability in a Mediterranean coastal ecosystem (Lesina Lagoon,South Adriatic Sea)

Spatial and temporal variability in zooplankton was studied at eight stations located in the Lesina Lagoon (South Adriatic Sea) Salinity, temperature, dissolved oxygen and chlorophyll a (in the lagoon) at these stations were also assessed. The zooplankton community was characterised by clear seasonal oscillations and mostly represented by copepods and their larvae. The dominant copepod species were Calanipeda aquaedulcis and Acartia tonsa, which exhibited spatial–temporal segregation in the lagoon. C. aquaedulcis copepodites seemed to be better adapted to oligotrophic and oligohaline conditions compared with the A. tonsa population. The invasive species A. tonsa has completely replaced the formerly abundant Acartia margalefi. A positive correlation was found between abundances, total species numbers and trophic state. An increasing abundance trend was shown from the lagoon towards the sea. The highest diversity indices were recorded at the two channel inlets, during high tide due to the absence of a clear dominance of one or more coastal species and the co-occurrence of species of lagoon and marine origin.     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic

Fishes and zooplankton were obtained (March–April 1979 and partly in August 1974) from 45 hauls taken during the day and at night in the central equatorial Atlantic between Latitude 3°N and 2°S from the surface to 1250-m depth, using the RMT 1+8, a combined opening-closing plankton and micronekton trawl. The vertical distribution of 30 myctophid species is described. All species migrate in a diel pattern, Ceratoscopelus warmingii and Lampanyctus photonotus down to at least 1250 m. During daytime most species aggregated at 400-to 700-m depth, therefore only partly occupying the depth of the Deep Scattering Layer (400 to 500 m at 15 kHz). The feeding patterns of seven of the most abundant species were compared, with a total of 1 905 stomach contents being analysed. All seven species are regarded as opportunistic predators, which feed predominantly during the night on calanoid copepods. A total of 66 species of calanoid copepods were identified among the prey items, with smaller species definitely being in the minority. Stomachs of C. warmingii (700 to 1 250 m depth) and Lepidophanes guentheri (500 to 900 m depth) from daytime samples contained copepod species restricted to the upper 150 m of the water column, including Undinula vulgaris, Nannocalanus minor, and Euchaeta marina, thereby confirming an extended vertical migration of predators. Differences in diet and preferences between species in their total food spectrum are described.     

Grazing rates of the Atlantic menhaden Brevoortia tyrannus as a function of particle size and concentration

The feeding behavior of adult Atlantic menhaden (Brevoortia tyrannus) upon 5 species of phytoplankton and 2 species of zooplankton has been studied. Four recognizable feeding stages which were a function of the concentration and size of the food particles were observed. During rapid feeding the fish swam at a constant speed for a prolonged period over a wide range of particle concentrations. Particle and food carbon-concentrations at the threshold for initiation and termination of feeding were inversely related to particle size. Carteria chuii (13.2 μ) was not grazed at a significant rate, while two-cell chains of Skeletonema costatum (16. 5 μ) were filtered from the water, indicating a minimum-size threshold for filtration of between 13 and 16 μ. The most rapid filtering rates were observed for the copepod Acartia tonsa ( \(\bar x\) volume swept clear = 24.8 l/fish/min). The maximum food-particle size acceptable to a menhaden appears to be between Acartia tonsa (1200 μ) and adult Artemia salina (10 mm). These results suggest that the large schools of menhaden found in Atlantic coastal waters could have a significant effect on the plankton, selectively grazing zooplankton, larger phytoplankton, and the longer chains of chain-forming diatoms.     

Food utilisation of pelagic mysids, Mysis mixta and M. relicta, during their growing season in the northern Baltic Sea

The dietary habits of the pelagic mysid Mysismixta were studied during its growing season at an open sea location in the Gulf of Finland, the northern Baltic Sea. Stomach samples were taken twice a month from June to September 1997. The most abundant phytoplankton taxa in the stomachs were diatoms and dinoflagellates, and copepods and cladocerans were the most abundant zooplankton identified. A clear change was found in the diets during the study period. Small mysids (3 to 6 mm) fed on sedimented phytoplankton in the early summer (90% benthic particles in June) but shifted gradually to a more pelagic and carnivorous diet (>40% pelagic particles, consisting of ca. 60% zooplankton in September). Seasonal changes in mysid capture ability as well as food availability were suggested to affect the diet composition of mysids during their growth. The ratio of pelagic and benthic food particles could – irrespective of the season – be explained by mysid size, whereas the zooplankton:phytoplankton ratio was better explained by season. The stomach analysis suggests that the mysids needed to attain a threshold size of 8 to 11 mm to initiate feeding on the more evasive copepods. Mysids also started to grow faster at the same time as the proportion of copepods increased in the diet, which suggests that copepods are an important energy source for M. mixta in late summer. Finally, a comparison was made between the M. mixta diet and that of the less abundant M. relicta. The diets of the two pelagic mysid species overlapped by 75% (Schoener's index). The main difference was due to M. mixta eating more zooplankton and pelagic material than M. relicta. Received: 15 September 1999 / Accepted: 18 January 2000     

The three-dimensional prey field of the northern krill, Meganyctiphanes norvegica, and the escape responses of their copepod prey

In the north Atlantic, Meganyctiphanes norvegica feeds predominantly on copepods, including Calanus spp. To quantify its perceptual field for prey, and the sensory systems underlying prey detection, the responses of tethered krill to free-swimming Calanus spp. were observed in 3D using silhouette video imaging. An attack–which occurred despite the krill’s being tethered—was characterized by a pronounced movement of the krill’s antennae towards the target, followed by a propulsion and opening of the feeding basket. Frequency distributions of prey detection distances were significantly different in the light vs. the dark, with median values of 26.5 mm and 19.5 mm, respectively. There were no significant differences in the angles at which prey were detected by krill (relative to the predator’s longitudinal body axis) in the light vs. the dark. Prey detections were symmetrically distributed on either side of the predator, in both light and dark. However, significant asymmetry was found in the dorsal–ventral direction with 80% of the prey detections located below the midline of the krill’s body axis and, given the placement and orientation of the compound eyes, presumably outside its visual field of view. This indicates that, at least under these conditions, vision was not the main sensory modality involved in the detection of active prey by M. norvegica. However, under some circumstances, vision may provide supplemental information. Avoidance responses of copepod prey were nearly twice the velocity of their nominal background swimming speed (153 ± 48 and 85 ± 75 mm s⁻¹, respectively), on average taking them 43 ± 16 mm away from the predator. This is far beyond the krill’s perceptual range, suggesting that the escape reaction provides an effective deterrent to predation (although perhaps less so for free-swimming krill). This information can be used to parameterize models that assess the role of krill as predators in marine ecosystems.     

Seasonal changes in feeding, gonad development and lipid stores in Calanusfinmarchicus and C. hyperboreus from Malangen, northern Norway

 Seasonal dynamics of feeding activity, oil sac volume, gonad development, sex ratio and spawning periods in the two sibling species Calanusfinmarchicus and C. hyperboreus, the key zooplankton copepod organisms throughout the northern Atlantic waters, were studied simultaneously in Malangen, northern Norway, during 1992. We were also tracing differences in surface- and deep-dwelling components of these populations in terms of oil sac volume and gonad development during the time period when the G₁ is preparing for a subsequent generation (G₂) or hibernation. The main difference in the life cycle strategies of these species is the earlier maturation and spawning of C. hyperboreus. No feeding activity in either of the two species was found in February, but both commenced feeding in March, prior to the spring phytoplankton peak. The larger copepod, C. hyperboreus, had a more intensive energy deposition than C. finmarchicus. The period of active feeding was much shorter for the former species, only from March through July in copepodite stages CIV and CV, and even less in females – March and April. Basically, a similar pattern of seasonal changes in gonad length and lipids was observed in the two species. In June, oil sacs in the surface- and deep-dwelling specimens were about equal, during the rest of the year, lipids in the deep CVs exceeded those in the surface. We propose that as copepods accumulated sufficient lipid reserves, they started to descend, while others, containing less fat, stayed in the upper layers feeding. The mean length of the gonads in the surface-dwelling copepods was consistently less than in their deep counterparts from October to February, so that gonad development at the expense of accumulated reserves during resting stage was confirmed. C. finmarchicus males were found in considerable numbers only in February and March, and were only occasionally found in the upper layers (0–100 m), while adult male C. hyperboreus were present from October to March, but were never found in the surface layers. The differences in life cycle timing among the two species are discussed in relation to tradeoffs with regard to foraging strategies, generation numbers, bioenergetics and predator avoidance. Received: 31 March 1999 / Accepted: 23 November 1999     

Natural diets of vertically migrating zooplankton in the Sargasso Sea

The feeding preferences of three common diel vertically migrating zooplankton were investigated from December 1999 to October 2000 at the U.S. JGOFS Bermuda Atlantic Time-Series Study (BATS) station in the Sargasso Sea. Gut content analysis of the copepods Pleuromamma xiphias (Giesbrecht) and Euchirella messinensis (Claus) and of the euphausiid Thysanopoda aequalis (Hansen) indicated that all three species fed on a wide variety of phytoplankton, zooplankton, and detrital material. Diet changes generally reflected seasonal trends in phytoplankton community structure. However, species-specific feeding preferences and differences in feeding selectivity among the three species were evident, and in general agreement with feeding habits predicted from the analysis of mouthpart morphology. The euphausiid T. aequalis fed equally on more different food types compared to both copepod species. The copepod P. xiphias consumed a diverse assemblage of phytoplankton from late winter through the summer (78-93% of gut items, by number, were phytoplankton) and based its diet more strongly on carnivorous feeding in autumn and early winter (31% and 61% of gut items were phytoplankton, respectively). E. messinensis showed the greatest feeding specialization, with a strong preference for pennate diatoms in winter and spring and for coccolithophorids during late summer and fall (constituting 67-93% of gut items by number). All three species consumed diatoms more than other phytoplankton taxa, even though diatoms form only a small fraction of the phytoplankton biomass in the Sargasso Sea. Although the majority of gut items identified were phytoplankton cells, the relative biomass contribution of these small cells may be lower than that of zooplankton and detritus. Zooplankton on which the three species primarily preyed were protozoans and crustaceans, but also included other metazoans such as chaetognaths and cnidarians. Marine snow was also an important component of the diet in all three species, with typically >50% and rarely <20% of the gut content being olive-green debris. Marine snow from larvacean houses was found in the guts of all three species, while E. messinenis appeared to selectively consume marine snow aggregates enriched with bicapitate Nitzschia spp. Large cyanobacteria (>4 µm in diameter) found in guts were also likely consumed with marine snow. The species-specific differences in the diets of these three migrating species suggest that an individual species approach is important in determining how feeding habits affect the structure of pelagic food webs and carbon cycling in the sea. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0815-8.