Influence of the biological and physical environment on the vertical distribution of mesozooplankton and micronekton in the eastern tropical Pacific

The day/night vertical distributions of mesozooplankton and micronekton biomass and that of a large number of copepod species to a depth of 1 000 m are described and contrasted in detail from two eastern tropical Pacific stations, the DOME station, in a region of continuous upwelling and the BIOSTAT station, in a non-upwelling area. The effects of various biological parameters such as the zones of primary production and chlorophyll maxima plus physical parameters, such as temperature, salinity and oxygen concentrations, on the distributions of the species and mesozooplankton and micronekton biomass are examined. The thermocline depth appeared to have the greatest influence of all the physical factors on the vertical distribution of the copepod species. The vertical distribution of many species was truncated in the region of the oxygen minimum layer with very few species found below this layer. The vertical distribution of copepod molts in the top 1 000 m showed that the molts probably originated at the depths at which they were found and that the molts from the euphotic zone did not reach the deep water. Copepod species association at different depths showed that few species had common depth distributions during both day and night. The highest degree of association occurred at night in the region of the thermocline. At depths below the thermocline, the degree of similarity in vertical distributions was greatly reduced. The biomass of mesozooplankton at these two stations is compared to other regions of the tropical oceans described in the literature and shown to be among the highest values recorded. The oxygen minimum layer and the thermocline had the greatest impact on species vertical distribution. The biomass of mesoplankton was about twice as high and the number of copepods between three and four times as high at the DOME site as compared to the BIOSTAT site, and evidence suggested that this was influenced by the higher primary production in this region. The difference in micronekton biomass between sites was less, with DOME having about 1.2 to 2.2 the biomass. This suggests that the influence of the higher level of primary productivity at the DOME was evidence of the higher trophic levels.     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). VI. Conclusions sur le rôle des euphausiacés dans les circuits trophiques de l'océan Pacifique intertropical

The role of euphausiids in the food webs of the Intertropical Pacific Ocean is defined through analysis of their nutrition, vertical distributions and migrations, and their utilization by pelagic predators. It is suggested that the abundance of the group, the extensive vertical migrations of many species and the fact that feeding takes place mainly in subsurface layers, result in a leading role of euphausiids in energy transfer between different bathymetric levels. For night-time feeding predators, they represent a noticeable food source only in the 0 to 300 m water layer, as 97% of the euphausiid biomass concentrates in this layer at night. In the daytime, only the smaller specimens (chiefly genus Stylocheiron), accounting for 10 to 15% of the whole biomass of the group, remain available for epipelagic (0 to 400 m) predators, larger individuals dwelling deeper. Euphausiids account for 8 to 10% of the food ingested by micronektonic fishes, but the species are not the same for different categories of fishes. Migrating fishes caught by pelagic trawls, more or less connected with the deep scattering layer, feed on migrating species in subsurface layers at night as well as in deeper layers during the daytime, and on non-migrating species inhabiting shallower and intermediate layers. On the other hand, fishes which comprise the prey of large long-line tunas, which are not caught by trawls because they are fast swimmers, feed almost solely on species which remain above 400 m in the daytime. These results suggest a certain degree of independance between the trophic webs which concern, on the one hand, epipelagic ichthyofauna (including tuna), and, on the other hand, migrating and deep-living faunas. Migrating populations are able to feed at night upon subsurface organisms, a part of this resource being then transmitted during the day to the deep-living fauna; but the epipelagic ichthyofauna, with a feeding activity restricted to light hours, has few possibilities to benefit from the migrating or deepliving biomass. Therefore, energy transfers seem to be intense only from subsurface (0 to 400 m) to deeper layers. From a more general point of view, these investigations suggest that, in the pelagic system, vertical distributions and migrations, and feeding rhythms, are the main factors determining the structure of the food webs.     

Observations on the diurnal vertical migrations of an oceanic animal community

Diurnal changes in abundance caused by vertical migrations have been examined in populations of copepods, ostracods, euphausiids, amphipods, decapods, chaetognaths, siphonophores and fish. The animals were taken in a series of hauls made over a 24 h period with an opening-closing midwater trawl system (RMT 1+8), consisting of a net of 1 m² mouth area combined in the same frame as one of 8 m² mouth area. The samples were taken at 250 m depth in a position 30°N; 23°W on 7/8 April 1972. The specific composition of the community and the numbers of individuals changed continuously with time. The numbers of fish, decapods and chaetognaths increased at night, but those of copepods, ostracods and euphausiids decreased. More species of fish, decapods and copepods were present by night than by day, whereas the numbers of species per haul for other groups remained fairly constant. The relative abundances of groups caught by the RMT 1 have been analysed, but similar treatment of the RMT 8 samples was impossible as only 3 groups were taken from this net. Non-migrants were a minority in every group except chaetognaths. Migrant species have been put into one of 6 transitory categories according to their patterns of abundance and hence migrations. Within each category, migratory behaviour varied both inter- and intraspecifically. The patterns of abundance of many species were smooth and continuous, suggesting slow migratory cycles of small amplitude. Conversely, extensive migrants had discontinuous patterns and presumably more rapid movements. Few migrants had a steady numerical plateau between their upward and downward migrations, and most apparently moved up or down continuously. The presence of migratory species in the sampled layer depended upon the time of day or night. It is concluded that, in a vertical series of hauls, the depths of occurrence of migrants will vary with the sampling time. Further-more, a vertical series will show a species minimum migration range but not necessarily its maximum. Individuals of some species were out of phase with the migrations of their main populations. There is evidence that the distributions and migrations of some species of decapods, euphausiids, copepods and fish could be related to the distribution of underwater light. Three pairs of congeneric copepod species were both spatially and temporally segregated for at least part of their diurnal cycles. Such an orderly arrangement could provide a means of reducing competition between species. Some species, however, overtook others on their migrations and the pattern of underwater light cannot, therefore, regulate the distribution of all species in the same way.     

Distribution of euphausiid assemblages in the Mediterranean Sea

Eleven species of euphausiids from 24 Isaac-Kidd Midwater Trawl (IKMT) night collections taken at stations throughout the Mediterranean Sea were counted. The frequency of occurrence and dominance of individual species and percent similarity faunal analysis of the euphausiid community were used to describe changes in faunal composition between geographical areas and differences in vertical distribution. Although most species were widespread, three distinct patterns of abundance were apparent: Euphausia krohnii, Nematoscelis megalops, Meganyctiphanes norvegica, and Stylocheiron abbreviatum predominated in western basin areas west of the Tyrrhenian Sea; Euphausia hemigibba, Thysanopoda aequalis, and Stylocheiron longicorne predominated in the Tyrrhenian Sea and eastern Mediterranean Sea; Euphausia brevis and Stylocheiron suhmii predominated in the eastern Mediterranean Sea. Percent similarity analysis of data from the IKMT samples and data from Ruud (1936) indicates the Tyrrhenian Sea fauna at the time of the collections was more similar to eastern Mediterranean areas than to most other areas in the western basin, although the degree of similarity was dependent, to some extent, on the depth at which the samples were collected. The composition of euphausiids living above 150 m at night in this area was more similar to eastern basin areas, while the composition of deeper living forms was more similar to those of the rest of the western basin. Comparison of euphausiids collected at three points over a 60 year time-span in the Balearic Sea shows the similarity in composition to be greater within the area over time than between adjacent areas in the western Mediterranean Sea.Contribution No. 2732 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA. This research was supported by the Atomic Energy Commission Contract AT (30-1)-3862, ref. NYO-3862-49, National Science Foundation Grant GA 29303 and Office of Naval Research Contract N00014-71-C-0284.     

Comparison of the food of Triphoturus mexicanus and T. nigrescens,two lanternfishes of the Pacific Ocean

Prey (chiefly euphausiids and copepods) eaten by two myctophids (lanternfishes) are compared from incidence in fish stomachs and from abundance in the environment. One lanternfish species, Triphoturus mexicanus, lives in the California Current, and the other, T. nigrescens, lives in the central Pacific Ocean. Although these two environments are very different physically and biologically, the feeding habits of the two lanternfishes are surprisingly similar. Prey biomass is 94% euphausiids, 3% copepods, and 3% other organisms for T. mexicanus and 88% euphausiids, 4.5% copepods, and 7.5% other organisms for T. nigrescens; the difference between the fish species is not significant when tested statistically. The two fishes resemble one another in frequency distributions of ingested copepod individuals, copepod species, euphausiid individuals, and euphausiid species. During a single diurnal feeding period, both fishes eat a variety of copepod species but tend to eat only a single species of euphausiid. T. mexicanus grows to twice the length of T. nigrescens and eats proportionally larger euphausiids; however, both fishes eat copepods having the same median size. The frequencies of euphausiid species in the diets of both fishes differ from the frequencies in the environment. The chief differences between the feeding habits of the two lanternfishes are that T. nigrescens, in comparison to its congener, eats a greater variety of organisms during one diurnal feeding period and captures smaller euphausiids. The feeding patterns for each lanternfish species are consistent over distances of hundreds of kilometers and over many years of sampling.     

Trophic strategies of euphausiids in a low-latitude ecosystem

Vertical distribution, diet, and morphology of adults were examined in 27 species of euphausiids occurring in the upper 1000 m in the eastern Gulf of Mexico. Vertical distribution patterns were similar to those found in the central ocean gyres and oceanic equatorial waters of the Atlantic, Indian and Pacific Oceans. Most species migrated vertically from their daytime depths of 300 to 600 m to the upper 300 m at night. Exceptions were the non-migrating species of Stylocheiron, which remained in the epipelagic zone day and night, and Nematobrachion boopis, which remained in the mesopelagic zone. Based on gut-contents analysis, the Gulf euphausiids were largely zooplanktivorous, with cyclopoid and calanoid copepods being the most common items in stomachs. ostracods were especially common in the stomachs of Thysanopoda spp. and phytoplankton in the guts of Euphausia spp. Nearly every species' diet contained a considerable amount of olive-colored debris, which may have been marine snow generated in the epipelagic zone. Cluster analysis grouped the euphausiids into nine diet guilds. Euphausiids with a generalized morphology (i.e., spherical eyes, uniform thoracic appendages) tended to group together and demonstrated little variety in stomach contents among species. Euphausiids with a specialized morphology (i.e., bilobed eyes, elongate thoracic appendages) showed considerable variety in stomach contents among species, and several species had diets that were highly specific. Many of the species that had similar gut contents fed on prey of different sizes, as indicated by the width of the calanoid copepod mandibles found in stomachs. Principal-components analysis of seven morphological characters yielded species groups that were similar, but not identical, to those generated by cluster analysis of stomach contents data. We inferred from this that morphological characters partly determine diet, but that behavior is also important. Using the 20 most abundant species and 3 niche parameters, we attempted to identify the degree of separation among euphausiids based on the level of overlap in vertical distribution and diet composition, and on differences in mean prey size. Overlap of <60% in vertical distribution or diet composition was considered to indicate distinction of that parameter. Of 190 total species pairs, only 4 pairs did not demonstrate niche separation in at least one of these categories. We found that differences in these niche parameters were greatest among species with a specialized morphology and least among species that were morphologically generalized.     

Feeding behavior of the ctenophore <Emphasis Type="Italic">Thalassocalyce inconstans</Emphasis>: revision of anatomy of the order Thalassocalycida

Behavioral observations using a remotely operated vehicle (ROV) in the Gulf of California in March, 2003, provided insights into the vertical distribution, feeding and anatomy of the rare and delicate ctenophore Thalassocalyce inconstans. Additional archived ROV video records from the Monterey Bay Aquarium Research Institute of 288 sightings of T. inconstans and 2,437 individual observations of euphausiids in the Gulf of California and Monterey Canyon between 1989 and 2005 were examined to determine ctenophore and euphausiid prey depth distributions with respect to temperature and dissolved oxygen concentration [dO]. In the Gulf of California most ctenophores (96.9%) were above 350 m, the top of the oxygen minimum layer. In Monterey Canyon the ctenophores were more widely distributed throughout the water column, including the hypoxic zone, to depths as great as 3,500 m. Computer-aided behavioral analysis of two video records of the capture of euphausiids by T. inconstans showed that the ctenophore contracted its bell almost instantly (0.5 s), transforming its flattened, hemispherical resting shape into a closed bi-lobed globe in which seawater and prey were engulfed. Euphausiids entrapped within the globe displayed a previously undescribed escape response for krill (‘probing behavior’), in which they hovered and gently probed the inner surfaces of the globe with antennae without stimulating further contraction by the ctenophore. Such rapid bell contraction could be effected only by a peripheral sphincter muscle even though the presence of circumferential ring musculature was unknown for the Phylum Ctenophora. Thereafter, several live T. inconstans were collected by hand off Barbados and microscopic observations confirmed that assumption.     

Vertical distributions of euphausiids and fish in relation to light intensity in the Northeastern Atlantic

The relationship between the vertical distributions of euphausiids and fish and light intensity has been studied directly by using a photometer in conjunction with an acoustically controlled rectangular midwater trawl. Samples were taken at a position centered on 47°N; 17°W on 15 and 16 May 1978. Five species of euphausiid and six species of fish have been analysed, both groups contained migrant and non-migrant species. The population of each of these species occurred throughout a light regime spanning at least three orders of magnitude of intensity; none of them was restricted to, or followed, and isolume. There were no sexual or size differences in the distributions of the euphausiids, but the population of Argyropelecus hemigymnus was probably stratified during the day, with smaller individuals occurring shallower than large ones. The results are discussed in relation to previous observations and to the theories of photic regulation of distributions and migrations.     

The feeding habits of three species of lanternfishes (family myctophidae) off oregon,USA

The common myctophids Stenobrachius leucopsarus, Diaphus theta and Tarletonbeania crenularis were found to feed primarily on the euphausiid Euphausia pacifica, the copepods Metridia lucens and Calanus spp., and the amphipod Parathemisto pacifica. The diets of these species of fishes were diverse and overlapped broadly, suggesting that they are feeding generalists. Most stomachs contained either all copepods or all euphausiids. Euphausiids were the most important prey on the basis of biomass. They comprised over one-half the weight of the stomach contents in over 40% of individuals of each of the three fishes. Stomach fullness and state of digestion of stomach contents differed over the diel period, but not enough to indicate feeding only during the nighttime. Average stomach fullness was greatest during the morning and night and well-digested material, which predominated in most stomachs, was most prevalent in the morning and afternoon. Either some feeding occurs throughout the day or digestion rates are slow.     

Distribution and diel vertical movements of mesopelagic scattering layers in the Red Sea

The mesopelagic zone of the Red Sea represents an extreme environment due to low food concentrations, high temperatures and low oxygen waters. Nevertheless, a 38 kHz echosounder identified at least four distinct scattering layers during the daytime, of which the 2 deepest layers resided entirely within the mesopelagic zone. Two of the acoustic layers were found above a mesopelagic oxygen minimum zone (OMZ), one layer overlapped with the OMZ, and one layer was found below the OMZ. Almost all organisms in the deep layers migrated to the near-surface waters during the night. Backscatter from a 300 kHz lowered Acoustic Doppler Current Profiler indicated a layer of zooplankton within the OMZ. They carried out DVM, yet a portion remained at mesopelagic depths during the night. Our acoustic measurements showed that the bulk of the acoustic backscatter was restricted to waters shallower than 800 m, suggesting that most of the biomass in the Red Sea resides above this depth.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Planktonic bioluminescence in the pack ice and the marginal ice zone of the Beaufort Sea

An icebreaker cruise into the Beaufort Sea in the fall of 1986 provided a unique opportunity for studying planktonic bioluminescence in ice fields and in the marginal ice zone. Bathyphotometer casts (bioluminescence intensity, seawater temperature, beam attenuation coefficient, and salinity) and biological collections were made to a depth of 100 m. A light budget, which describes the planktonic species responsible for the measured bioluminescence, and a dinoflagellate species budget were constructed from the mean light output from luminescent plankton and plankton counts. The vertical distribution of bioluminescence among the ice stations was similar. The maximum intensities were 2 to 8×10⁶ photons s^-1 cm^-3 in the upper 50 m of the sea-ice interface. The marginal ice zone station (MIZ) exhibited a maximum intensity of 2 to 3×10⁸ photons s^-1 cm^-3 between 5 and 30 m depth. At Ice Station 2, Metridia longa and their nauplii contributed approximately 80% of stimulable bioluminescence in the upper 10 m but, overall, Protoperidinium spp. dinoflagellates contributed most of the light to a depth of 100 m. In the MIZ, Protoperidinium spp. dinoflagellates contributed 90% of the light within the upper 10 m, decreasing to 43% of the contributed light at a depth of 40 m. Below 40 m, dinoflagellate bioluminescence decreased to a few percent of the total to a depth of 90 m. Metridia spp. copepods contributed more than 50% of the light at depths from 40 to 90 m. Ostracods, larvaceans, and euphausiid furcilia contributed <1% of all bioluminescence at all depths sampled. Correlation analyses between measured bioluminescence (photons s^-1 cm^-3), the number of bioluminescent dinoflagellates and the light budget for the MIZ indicated highly significant associations: r=0.919, p=0.001, and r=0.912, p<0.001, respectively (Student's two-tailed t-tests). Bioluminescence was negatively correlated with seawater salinity at all stations (p=0.001). Maximum bioluminescence was measured in the less saline surface waters at all stations.     

Vertical distribution and migration of fishes of the lower mesopelagic zone off Oregon

An exceptionally large midwater trawl (50 m² mouth area) with 5 opening and closing codends was towed horizontally in the lower mesopelagic zone at depths of 500, 650, 800 and 1000 m off Oregon (USA) from 1–6 September, 1978. In comparison to more conventional trawls, ours collected more fish, including rare species and large individuals of common species. Comparison of collections made by day and by night revealed that 12 of the 15 most common species probably migrated vertically. Bathylagus milleri evidently migrates from 650 m during the day to 500 m at night. Cyclothone acclinidens and C. atraria were more abundant by night than by day at 800 m, possibly due to an upward migration from deeper depths at night. C. pseudopallida, C. signata, Chauliodus macouni, Tactostoma macropus and Stenobrachius leucopsarus were more abundant by day than by night at 500 m, suggesting that they migrated out of this depth horizon at night. Lampanyctus regalis, and large individuals of B. pacificus were more abundant by night than by day at 500 m, possibly because they migrated upward from near 650 m. Many species exhibited trends of increasing or decreasing size with depth, and several species showed changes in migratory behavior with size. For example, only small (<240 mm) T. macropus migrated vertically, whereas only large (>110 mm) B. pacificus appeared to migrate. Depths of maximum abundance of congeneric species were usually separated. B. milleri and B. pacificus had similar distributions by day, but the former was shallower at night. S. leucopsarus tended to live shallower than S. nannochir both day and night. Congeners always occurring at the same depth were Cyclothone pseudopallida and C. signata (both most abundant at 500 m) and C. acclinidens and C. atraria (both most abundant at 800 m).     

Microbial activities as a function of water depth in the Ligurian Sea: an autoradiographic study

Fourteen species of sergestid shrimps were collected in the Sargasso Sea between the surface and 1500 m near Bermuda on 4 cruises. The vertical distribution and feeding activity of the most abundant species are discussed in relation to interspecific competition and the adaptive significance of vertical migration. Each species lives within a narrow depth range and exhibits a diel vertical migration. Sergestes splendens migrated as much as 825 m, while S. japonicus migrated less than 100 m. Neither the seasonal nor permanent thermocline influenced the migration range. The only species which occurred together both day and night were S. pectinatus with S. vigilax and S. pectinatus with S. sargassi. Morphological differences in the third maxillipeds of these species suggest differences in feeding. Although most species eat a variety of organisms, the foreguts of S. grandis, S. corniculum, and S. splendens contained euphausiids more often than those of other species, and S. grandis and S. robustus fed more frequently on fishes. In contrast, S. japonicus appears to feed on detritus. Food was found in the foreguts of most species less frequently during the day than night, but no species fed only at night. S. sargassi and S. pectinatus fed equally day and night.     

Species composition, biomass and vertical distribution of micronekton over the mid-slope region off southern Tasmania, Australia

Seasonal sampling was carried out based on day/night, vertically stratified tows (100 or 125 m strata) in the upper 900 m of the water column over the mid-slope commercial fishing grounds south of Tasmania. A large midwater trawl (105 m² mouth area) was used with an opening/closing cod-end. Subtropical convergence and subtropical species dominated the fauna, but many less abundant, more widely-distributed species were also present. Fishes, which contributed 89% of micronekton biomass and 135 of 178 species, were dominated by the Myctophidae (48% biomass and 48 species). Twenty micronekton species made up 80% of the total biomass. Overall, the micronekton fish biomass in this region was 2.2 g m⁻² wet weight. A pronounced day/night shift in the distribution of biomass was attributable to diel migratory species. During the day, <0.2% of the total micronekton biomass was found in 0 to 300 m; most biomass was below 400 m, with peaks at 400 to 525 m and 775 to 900 m. At night, 53% of the biomass was found in 0 to 300 m, with progressively less in each deeper stratum. The vertical ranges of individual species typically exceeded 400 to 500 m during the day and night and were non-coincident, although nyctoepipelagic migrators were concentrated in the surface 200 m at night. Distinct epipelagic, lower and upper mesopelagic assemblages were identified, and patterns of epipelagic migration, limited migration and non-migration were categorised for species from each of the lower and upper mesopelagic assemblages. The vertical distribution of these assemblages was coincident with the primary water masses: subantarctic mode water (∼250 to 600 m) and antarctic intermediate water (below ∼700 m). The flux of migrating micronekton, estimated at 0.94 to 3.36 g C m⁻² yr⁻¹ to the lower mesopelagic and 1.14 to 4.06 g C m⁻² yr⁻¹ to the upper mesopelagic, appeared to be considerably outweighed by the consumption needs of aggregated mid-slope benthopelagic predators. We suggest that advection of mesopelagic prey in antarctic intermediate water may sustain aggregated populations of orange roughy (Hoplostethus atlanticus) and other predators on the micronekton in mid-slope depths at this site. Received: 2 April 1997 / Accepted: 21 August 1997     

Drivers of euphausiid species abundance and numerical abundance in the Atlantic Ocean

Mid-ocean ridges are common features of the world’s oceans but there is a lack of understanding as to how their presence affects overlying pelagic biota. The Mid-Atlantic Ridge (MAR) is a dominant feature of the Atlantic Ocean. Here, we examined data on euphausiid distribution and abundance arising from several international research programmes and from the continuous plankton recorder. We used a generalized additive model (GAM) framework to explore spatial patterns of variability in euphausiid distribution on, and at either side of, the MAR from 60°N to 55°S in conjunction with variability in a suite of biological, physical and environmental parameters. Euphausiid species abundance peaked in mid-latitudes and was significantly higher on the ridge than in adjacent waters, but the ridge did not influence numerical abundance significantly. Sea surface temperature (SST) was the most important single factor influencing both euphausiid numerical abundance and species abundance. Increases in sea surface height variance, a proxy for mixing, increased the numerical abundance of euphausiids. GAM predictions of variability in species abundance as a function of SST and depth of the mixed layer were consistent with present theories, which suggest that pelagic niche availability is related to the thermal structure of the near surface water: more deeply-mixed water contained higher euphausiid biodiversity. In addition to exposing present distributional patterns, the GAM framework enables responses to potential future and past environmental variability including temperature change to be explored.     

Deep Arabian Sea mesozooplankton distribution. Intermonsoon, October 1995

Mesozooplankton (<5 mm) collected by stratified oblique tows with a 1-m² MOCNESS was examined at four stations in the Arabian Sea, with special reference to the bathypelagic zone. The profiles commenced about 20 m above bottom, at 4430 m as a maximum depth. The highest mesozooplankton biomass concentrations (wet weight per cubic meter) were obtained from the surface layer during night. A secondary maximum was situated between 150 and 450 m, with maximum concentrations at daytime. This layer coincided with the daytime residence depth of the deep scattering layer. The standing crop of the mesozooplankton in the upper 1000 m was highest at station WAST at 16°N; 60°E (ca. 47 000 mg m⁻²); station CAST at 14°N; 65°E ranked second (ca. 22 500 mg m⁻²), followed by station SAST at 10°N; 65°E (11 420 mg m⁻²). The differences can be related to different productivity regimes at the sea surface generated by the Findlater Jet during the SW monsoon. The differences in surface production were also reflected below 1000 m depth, in the bathypelagic zone, with mesozooplankton wet weights of 5330 mg m⁻² at WAST, 3210 mg m⁻² at CAST, 3390 mg m⁻² at EAST (15°N; 65°E) and 2690 mg m⁻² at SAST. The decrease of mesozooplankton concentration with depth in the oxygen minimum zone (OMZ) was stronger than in comparable depths of open-ocean areas where an OMZ is absent. Among the discriminated four size classes of mesozooplankton, the largest fraction (2 to 5 mm) indicated a biomass peak at 1200 m depth, which coincided with the lower boundary layer of the OMZ. The rate of decrease of mesozooplankton biomass with depth in the bathypelagic zone was statistically similar between the sites, even though the absolute zooplankton biomass at the sites was different. There is no evidence that the presumed lower carbon degradation rates in the OMZ of the Arabian Sea caused a larger standing crop and less of a decrease in biomass with depth in the bathypelagic zone in comparison to other seas. Received: 16 May 1997 / Accepted: 5 June 1997     

Polonium-210 in euphausiids: A detailed study

A detailed study of ²¹⁰Po, the predominant alpha-emitting nuclide found in most marine organisms, has been undertaken in a particular zooplanktonic species, the euphausiid Meganyctiphanes norvegica. The purpose was to obtain information concerning the origin, the localization and the flux of the nuclide in and through this organism. Measurements of ²¹⁰Po were made in euphausiids of different sizes, in dissected organs and tissues, and in excretion products. The results show higher concentrations in the smaller specimens; this fact cannot be explained on the basis of surface adsorption, but is probably related to the ingestion of food. Dissection results show that the distribution of ²¹⁰Po in euphausiids is not homogeneous, but that the majority is concentrated in the internal organs, the alimentary tract and the hepatopancreas in particular. The natural radiation dose received by these organs is in consequence much higher than that received by the whole animal. Use of a dynamic model allowed the flux of ²¹⁰Po through M. norvegica to be calculated. The calculations confirm that food is the principal source of ²¹⁰Po for this species, and clearly show that fecal pellets constitute the major elimination route. Extrapolation of the data to zooplankton in general leads to the conclusion that zooplankton metabolic activity plays an important role in transporting ²¹⁰Po from the surface layers of the ocean to depth.     

Flux of cadmium through euphausiids

Flux of the heavy metal cadmium through the euphausiid Meganyctiphanes norvegica was examined. Radiotracer experiments showed that cadmium can be accumulated either directly from water or through the food chain. When comparing equilibrium cadmium concentration factors based on stable element measurements with those obtained from radiotracer experiments, it is evident that exchange between cadmium in the water and that in euphausiid tissue is a relatively slow process, indicating that, in the long term, ingestion of cadmium will probably be the more important route for the accumulation of this metal. Approximately 10% of cadmium ingested by euphausiids was incorporated into internal tissues when the food source was radioactive Artemia. After 1 month cadmium, accumulated directly from water, was found to be most concentrated in the viscera with lesser amounts in eyes, exoskeleton and muscle, respectively. Use of a simple model, based on the assumption that cadmium taken in by the organism must equal cadmium released plus that accumulated in tissue, allowed assessment of the relative importance of various metabolic parameters in controlling the cadmium flux through euphausiids. Fecal pellets, due to their relatively high rate of production and high cadmium content, accounted for 84% of the total cadmium flux through M. norvegica. Comparisons of stable cadmium concentrations in natural euphausiid food and the organism's resultant fecal pellets indicate that the cadmium concentration in ingested material was increased nearly 5-fold during its passage through the euphausiid. From comparisons of all routes by which cadmium can be released from M. norvegica to the water column, it is concluded that fecal pellet deposition represents the principal mechanism effecting the downward vertical transport of cadmium by this species.     

Small-scale horizontal distributions of zooplankton taxa

Small-scale (100 to 2 400 m) horizontal distributions of major taxonomic categories (class and order) of zooplankton were measured at a depth of 90 m with an opening-closing plankton net over a 3 d period in October 1978 in the California Current. Some zooplankton categories showed evidence of diurnal vertical migration, while others had long-period temporal changes in mean abundance. Variance-to-mean ratio for large copepods and euphausiids was higher at night than during the day, while the opposite was true for chaetognaths and pteropods. Within a given category, the variance-to-mean ratio generally increased with a category's abundance. Spatial abundance variations were characterized by trends (i.e., fluctuations larger than length of the net hauls) in some taxonomic categories. No consistent differences in scales of variability were found as a function of animal size or from day to night. Correlation analysis of taxonomic counts implied that significant biological interactions occurred. The proportions of counts of taxonomic groups showed no large changes over the time-space scales sampled. However, the proportions of biomass in taxonomic groups differed from day to night due to the large variability of euphausiids. Comparisons of wet weight biomass to taxonomic counts indicated that biomass was usually less variable than taxonomic counts.