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.     

Photosynthetic rates of benthic marine algae in relation to light intensity and seasonal variations

The effect of light intensity on rate of photosynthesis was measured at irregular intervals over a 12-month period for 22 benthic marine algae in the western Baltic Sea. In most species photosynthesis (mg O₂·g dry weight^-1·h^-1) was highest in spring and summer, corresponding to the seasonal growth pattern of the algae. In winter all the species showed adaptation of the light compensation point. Highest productivity was shown by algae which are: short-lived annual species rather than perennials, eulittoral rather than sublittoral, and which possess sheet-like or filamentous thalli rather than coarsely branched forms. These factors are clearly inter-related.This research was carried out while one of us (R.J.K.) was an Alexander von Humboldt fellow at the University of Kiel, and is part of the programme Sonderforschungsbereich 95, Wechselwirkung Meer-Meeresboden, Universität Kiel.     

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

The vertical distribution of seven sternoptychid species was examined from RMT 1+8 samples collected aboard R. V. Meteor in March-April 1979 and from Royal Research Ship R.R.S. Discovery in July 1974 in the central equatorial Atlantic. During daytime sternoptychids occupied depths between 200 and 1250 m, with Sternoptyx pseudobscura living deepest, centering between 800 and 900 m, and Argyropelecus sladeni most shallow, aggregating predominantly at 300 and 400 m. They are all considered limited or partial migrants, ascending only some 100 and 200 m towards the surface at night. Only A. sladeni was observed to enter the epipelagic zone (0 and 200 m).-Feeding patterns were investigated from stomach content analyses of Sternoptyx diaphana, S. pseudobscura, Argyropelecus sladeni and A. affinis. Additional stomach contents were analysed from samples of S. diaphana, A. hemigymnus and A. olfersi collected in June 1985 from F.R.V. Walther Herwig in the temperate NE Atlantic at 46°N, 17°W by means of the Engel Trawl. The food spectrum of the six species is generally described, and additional dietary evidence regarding calanoid copepod prey is provided for four of these taxa. All sternoptychid species investigated were planktivorous, feeding predominantly on copepods and ostracods, except for the largest size class, which preyed heavily on euphausiids and amphipods. The relationship of predator size towards prey type and prey size is analysed for both Sternoptyx species. Of these, S. pseudobscura in particular exhibits taxonomic selectivity towards polychaete prey. The diet of both species of Sternoptyx included a number of epipelagic or even neustonic calanoid copepod species which contributed more than 50% of the total copepod population by numbers. So far it is not known how the predators find access to prey organisms of the upper 200 m, as netfeeding is considered unlikely. Cyclopoid copepods of the genus Sapphirina were observed as dietary component particular of S. diaphana.     

Diets and bathymetric distributions of the macrourid fish of the Rockall Trough,northeastern Atlantic Ocean

The Macrouridae are the most common fish caught in demersal trawls on the continental slope and rise of the Rockall Trough. They represented 41% of all fish caught, with Coryphaenoides rupestris amounting to 28% of the catch. No previous study of the trophic interaction of these fish has been made over a wide bathymetric range. Samples were obtained at 250-m intervals of depth between 400 and 2 900 m in the period 1975 to 1981. The stomachs of 5 326 fish belonging to 12 species were examined to define their diets. Eight species are primarily benthopelagic feeders while four are primarily epibenthic feeders. The bathymetric centres of distribution of the populations of the benthopelagic feeding Trachyrhynchus murrayi, Malacocephalus laevis, Coryphaenoides rupestis, C. guentheri, Nematonurus armatus, Chalinura brevibarbis, C. leptolepis and C. mediterranea are different from each other. Similar differences in bathymetric distribution occurred among the epibenthic feeding Nezumia aequalis, Coelorhynchus coelorhynchus, C. occa and Lionurus carapinus. These differences decrease competition among species that exploit similar resources.     

Population dynamics and production of euphausiids. I. Thysanoessa longicaudata in the North Atlantic Ocean

Adults and furciliae of Thysanoessa longicaudata (Krøyer) taken in continuous plankton records (CPR) in 1966 and 1967 were examined and measured. The measurements were converted to biomass using a length/weight equation, and net production values were calculated from these data. Two generations were spawned annually in the eastern North Atlantic Ocean and only one in the colder western waters and in the Norwegian Sea. The timing of reproduction in the spring was correlated with the mean sea-surface temperature. The highest annual production values (up to 12.24 mg dry weight m^-3 year^-1) were recorded in the eastern areas, where the ratio between production and biomass was usually between 4:1 and 8:1 compared with ratios usually between 2:1 and 3:1 in the western waters. The values for daily production per individual ranged from 3.34 g dry weight day^-1 to 17.54 g day^-1. Mortality in the early stages of development was calculated as about 98.2% of the eggs produced.     

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.     

Models of fish school size in relation to environmental productivity

A simple energy-balance model, relating energetic requirements of fish schools to food production, was used to predict shoal sizes. Lower limits to school size are unlikely to be set by food but rather by predation. Upper limits depend on both food and school behavior, being greater for schools that break up to feed than for schools that remain continuously cohesive. Faced with a decreasing food supply, a school could either break into smaller schools or change behavior, increasing the area available for foraging. The models suggest that environmental productivity needs to be considered when examining fishery statistics such as (catch per unit effort), where maximum catch may be limited by maximum school size.     

Vertical movement of the littoral fringe periwinkle Nodilittorina exigua in relation to wave height

The relationship between daily vertical distribution of the littoral fringe periwinkle Nodilittorina exigua and daily wave height was investigated at Shirahama, Japan (33°42N; 135°21E) daily, from May 1981 to August 1982. The positions of the snails on a cliff shore were high when the wave-reach was high, and ascended with increasing height of the wave-reach. The snails moved a long distance upwards in late summer, when typhoon swells occurred, and moved gradually downwards again in autumn, parallel to decreasing wave-reach. They displayed more active vertical movement in warmer seasons of the year and this activity was not positively correlated with the frequency of wetting. Downward migration in autumn ensured frequent and constant immersion. Taken in conjunction with results in the literature, the present study emphasizes the adaptive significance of vertical movement by N. exigua in the littoral-fringe environment.     

Genetic and taxonomic relationships among Northeastern Atlantic and Mediterranean populations of the soft coral Alcyonium coralloides

Species boundaries among taxa of colonial marine organisms are often obscured by intraspecific morphological and ecological variation; genetic comparisons of recognized “ecotypes” frequently reveal them to be reproductively isolated species. Based on morphological similarities, it has been proposed that the Mediterranean soft coral Alcyonium (=Parerythropodium) coralloides Pallas, 1766 and its Atlantic congener A. hibernicum belong to one highly variable and geographically widespread species, A. coralloides. I collected A. coralloides from ten Atlantic and three Mediterranean locations in 1990 and 1994, and used differences in colony form, substrate use and color to separate them into five distinct morphotypes. Two occur sympatrically in the Mediterranean (M1, M2) and three have overlapping distributions in the Atlantic (A1, A2, A3). I used allozyme electrophoresis to compare morphotypes genetically at 14 enzyme loci. Where two morphotypes occurred sympatrically, fixed allelic differences at 4 to 6 loci indicated reproductive isolation. In all but one pairwise comparison (M1 and A2), morphotypes whose ranges did not overlap were also separated by large genetic distances. From these results I suggest that the five morphotypes represent four distinct species. A. coralloides comprises two morphotypes (M1, A2) with relatively high genetic identity. A. hibernicum (=A1) is reproductively isolated from A. coralloides and should be retained as a valid species; levels of genetic diversity and heterozygosity within populations support the absence of outcrossing in this reportedly asexual species. Morphotypes M2 and A3 are also reproductively isolated from A. coralloides; they are taxonomically distinct from but belong to the same phylogenetic clade as A. hibernicum. Although preliminary observations suggest that differences in reproductive timing maintain species boundaries in sympatry, wider geographic sampling will be required to elucidate the events leading to speciation within this species complex. Received: 8 May 1998 / Accepted: 8 October 1998     

Lipid content of some Caribbean corals in relation to depth and light

Three procedures for the extraction of coral lipids were compared and a rapid and effective method for future use is suggested. This method was used to measure the lipid content of the branching coral Porites porites (Pallas) and the massive corals Montastrea annularis (Ellis and Solander) and Siderastrea siderea (Ellis and Solander) during July and August 1991. P. porites and M. annularis were sampled on two fringing reefs, each characterised by different water quality affecting light transmission, and at depths of down to 30 m on a barrier reef on the west coast of the island. m. annularis contained, on average, 29% of dry weight as lipid, and there were no significant differences in lipid levels between corals sampled on either fringing reef at 3 and 6 m, or between fringing reefs and the barrier reef at 13, 20 or 30 m depth. Five samples were also taken from a single massive colony of S. siderea at 3 m on a fringing reef and at 13, 20 and 30 m depth on the barrier reef. Values for lipid ranged from 26 to 35% of dry tissue weight. P. porites from 3 and 6 m depth on both fringing reefs contained the same amount of lipid (11% of dry tissue weight). However, at 13 m depth on the barrier reef this coral contained significantly less lipid (8.5% of dry tissue weight). This difference may be attributable to a higher nutritional intake by P. porites on the fringing reefs.     

Morphology of the gastric mill in ten species of euphausiids

SEM observation revealed the detailed morphology of the gastric mill in ten species of euphausiids, Bentheuphausia amblyops, Thysanopoda acutifrons, Meganyctiphanes norvegica, Pseudeuphausia latifrons, Euphausia superba, Tessarabrachion oculatum, Thysanoessa longipes, Nematoscelis microps, Nematobrachion boopis, and Stylocheiron maximum. The well-developed gastric mill of euphausiids consists of a pair of cluster spines and a pair of lateral teeth displaying great diversity in morphology. It is suggested that there are three categories of gastric mill and filterpress (gland filter) within euphausiids. One is the well-developed gastric mill with filter-press of B. amblyops; a second is the well-developed gastric mill of M. norvegica, T. oculatum, E. superba, P. latifrons and T. longipes; and the third is the gastric mill without lateral teeth of T. acutifrons, N. microps, N. boopis and S. maximum.     

Biology of euphausiids in the subarctic waters north of Iceland

The seasonal abundance, maturity, spawning, and population dynamics of Thysanoessa inermis (Krøyer, 1846), T. longicaudata (Krøyer, 1846), and Meganyctiphanes norvegica (M. Sars, 1857) were studied in the subarctic waters north of Iceland from February 1993 to February 1994. The material was sampled at approximately monthly intervals along a transect of eight stations extending from 66°16′ to 68°00′N at 18°50′W. Information on temperature and chlorophyll a concentrations is also presented. Spring warming of the water began in March to April and maximum temperatures were recorded in August (3.8?°C). The spring bloom of the phytoplankton started in late March and highest chlorophyll a concentrations were measured during middle to late April (7.0?mg chlorophyll a m^?3). T. inermis was the dominant species in the samples, constituting 77% of juvenile, male and female euphausiids present. The greatest abundance of juvenile, male and female T. inermis and M. norvegica was observed during autumn and winter, with lower abundance in spring and summer. T. longicaudata showed only limited changes in seasonal abundance. Male T. inermis had spermatophores in their ejaculatory ducts from February to May, while mature females had spermatophores attached during April and May. T. longicaudata males bore spermatophores from February to July, whereas females only bore spermatophores in April and May. M. norvegica males had spermatophores from February to April, while the single female with spermatophores was caught in February. Euphausiid eggs were first recorded during the latter part of April; the highest numbers of eggs were observed in the samples taken in late May. Maximum numbers of nauplii of both Thysanoessa spp. and M. norvegica were recorded in late May. The main spawning of the euphausiids coincided with the phytoplankton spring bloom. Most male T. inermis took part in breeding at 1 yr of age while most females appeared not to mature until 2 yr of age. T. inermis has a life span of just over 2 yr, T. longicaudata appears to live just over 1 yr. Limited data did not allow the life span of M. norvegica to be determined.     

Vertical distribution of zooplankton glutamate dehydrogenase in relation to chlorophyll in the vicinity of the Nantucket Shoals

The vertical distributions of chlorophyll, and zooplankton glutamate dehydrogenase (GDH) and respiratory electron transport system (ETS) activities were measured at five stations in the vicinity of the Nantucket Shoals using a submersible pump-profiling system. Distinctly lower zooplankton GDH activities and chlorophyll concentrations were found at a station showing little vertical stratification as compared to the other stations where vertical stratification was more pronounced. Maximum GDH activity was observed at or immediately above the chlorophyll maximum at the stratified stations. Respiration to excretion (O:N) ratios determined from the enzyme data were lowest in or just above the chlorphyll maximum, suggesting proteolytic catabolism at those depths. Both features had previously been observed in the oligotrophic Gulf of Mexico. Ammonium excretion determined from GDH activity was estimated to provide 20 to 40% of the total algal nitrogen requirement.Bigelow Laboratory Contribution No. 82016     

Deep distributions of oceanic cirripede larvae in the Sargasso Sea and surrounding North Atlantic Ocean

From deep ( 1000 m), multi-depth zooplankton samples taken in the Azores frontal region from November 1980 to June 1981, high numbers of the two larval forms ofLepas pectinata, the commonest lepadid cirripede in this area, were sampled. The nauplii and cyprid larvae are large and long-lived. Nauplii were sampled where they feed, in the upper 150 m, closely associated with the fluorescence maximum. The cyprids were mainly sampled between 300 and 400 m, with a sharp cut-off in numbers just below the distribution peak. This deep distribution is intriguing, as the cyprids, which were not migrating diurnally, are non-feeding. To complete their life cycle they must settle on floating debris at the surface, where very few were sampled. They are negatively buoyant and there is no apparent physical reason for this deep distribution. There was no obvious pattern of changes in individual length or length/dry weight distribution with depth, the population appeared homogeneous. There was also no accumulation of other zooplankton at the same depth, or evidence from gut contents of large carnivores that the distribution pattern was caused by selective predation. A close association exists betweenL. pectinata andSargassum spp. weed, the cyprids preferentially settling on small fragments of weathered weed, the supply of which may be enhanced after winter. During winter, the harsh environment at the surface of the northern Sargasso Sea may discourage cyprid settlement. The deep distribution may be an ontogenetic migration conferring some survival advantage and saving energy due to the lower temperature and oxygen levels at depth. If it is a behavioural mechanism, duration of the cyprid stage will be extended until a more clement time of year, allowing them to settle over a longer period. The cyprids of other species were also sampled at depth, so the phenomenon appears to be widespread. If subsequent research confirms these observations as an ontogenetic migration, our current understanding of the fouling behaviour of oceanic lepadid cirripedes would be considerably modified.     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Concentrations of iron-55 in commercial fish species from the North Atlantic

Concentrations of the weapon-test, fallout radionuclide iron-55 have been reported for migratory species, such as the Pacific salmon, which suggest a marked decrease in concentration with decrease in latitude of the point of capture. This situation has been examined for North Atlantic cod stocks in order to provide data for a species representative of a large geographical area, but divided into more or less distinct stocks each with a restricted territory, in order to provide further data on the importance of latitude in determining iron-55 concentrations. The data obtained show that there is a marked dependence upon latitude, Arctic cod averaging 90 pCi ⁵⁵Fe/mg Fe compared with 15 pCi ⁵⁵Fe/mg Fe for cod from middle latitudes. The mechanisms that could lead to such distribution are examined, and it is concluded that deposition of this radionuclide was exceptionally high over the sea in northern latitudes compared with its deposition on land, and that this pattern of deposition, taken together with the very high biological availability of the material at the time of deposition, has led to the very high specific activities found in northern latitudes and recorded for Arctic cod stocks in particular.     

The nature of the photosynthate in natural phytoplankton populations in relation to light quality

The relative concentrations of a variety of organic compounds in phytoplankton from different depths of the photic zone of Saanich Inlet and Indian Arm, British Columbia, Canada, were determined. In surface phytoplankton the greatest proportion of ¹⁴C in newly-formed organic compounds was in the ethanol-soluble fraction irrespective of the light intensity. The proportion of the label in the insoluble fraction increased with depth. Within the ethanol-soluble fraction the relative importance of carbohydrates decreased and amino acids increased with depth. Experiments in Indian Arm indicated that these observed changes were in response to light quality rather than intensity. The percentage release of dissolved organic carbon as a percentage of total fixation decreased with depth and was directly related to the size of the ethanol-soluble fraction.