Plankton of the fladen ground during FLEX 76 I. Spring development of the plankton community

Results from plankton sampling in the northern North Sea with the Continuous Plankton Recorder (CPR) and the Undulating Oceanographic Recorder (UOR) during the Fladen Ground Experiment in 1976 (FLEX 76) are summarised. The first evidence of the spring outbreak of phytoplankton was on 19 April, the day after the first signs of vertical stability of the water column were observed. This was followed by spawning of the euphausiid Thysanoessa inermis and rapid increase in the numbers of Calanus finmarchicus. C. finmarchicus was the most abundant species over the FLEX period (19 March to 3 June) and, together with T. inermis, accounted for over 80% of the dry weight of the zooplankton standing stock. By early June the standing crop of phytoplankton had been depleted and nutrients levels were reduced to very low concentrations in the upper 50 m.JONSDAP Contribution No. 49     

Plankton of the fladen ground during FLEX 76 II. Population dynamics and production of Thysanoessa inermis (Crustacea: Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m^-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m^-2 d^-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m^-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.JOSDAP Contribution No. 50     

The undulating oceanographic recorder — A new instrument system for sampling plankton and recording physical variables in the euphotic zone from a ship underway

The prototype Undulating Oceanographic Recorder Mark I is an instrumented towed vehicle which can be programmed to undulate between a minimum depth of 8 m and a selected maximum depth between 15 and 70 m, with an undulation length between 3 and 30 km, at any speed between 7 and 15 knots (3.6 to 7.7 m/sec). It takes a continuous series of plankton samples, each integrated over the undulation depth range, and records data on magnetic tape, at a preselected rate between 30/min and 30/h, from which salinity, temperature and depth can be derived with accuracies of ±0.1, ±0.1C° and ±0.5 m, respectively. The instrument is automatic, self-contained and generates its own power supplies. It is towed on about 200 m of unfaired steel cable and can be handled with the winching and lifting equipment found on most research vessels. Examples of processed data from three cruises are presented: temperature and salinity contours on a vertical section through a complex stratified area in the N.W. Irish Sea; results from a survey of the waters over the continental shelf and slope to the north of Scotland, showing the distribution of water masses and associated plankton; and hydrographic data taken in Loch Etive on the west coast of Scotland.     

Population dynamics and production of euphausiids II. Thysanoessa inermis and T. raschi in the North Sea and American Coastal Waters

Results from the Continuous Plankton Recorder (CPR) survey for 1966 and 1967 are used to describe seasonal changes in abundance, size and aspects of the population structure of Thysanoessa inermis (Krøyer) and T. raschi (M. Sars) at a depth of 10 m in the North Sea and in American coastal waters from the Grand Banks to the Gulf of Maine. Production and dry weight were estimated from these data. Two year-groups were usually present in the breeding population, the proportion surviving into a second year being higher in American waters than in the North Sea. Annual production for each species was within the range 0.69 to 4.66 mg m^-3 and the ratio between production and biomass (P:B) was between 1.3 and 4.2; values outside these ranges were obtained only for American coastal waters in 1967, when the frequency of sampling was low.     

A marine environmental recorder

An inexpensive Marine Environmental Recorder is described. The instrument system is small, lightweight and of low-power consumption. It is flexible, simple to operate and economical. It can be used remotely in a moored, buoyed or towed instrument system, recording measurements continuously for up to 24 h, or intermittently for 1 min every hour, for a period of up to 60 d. It has been used extensively in the Continuous Plankton Recorder and the Undulating Oceanographic Recorder to measure temperature, depth and occasionally chlorophyll and radiant energy; as a temperature recorder, it has a resolution of 0.1 C^o, an uncertainty of measurement of ±0.1 C^o and a stability of calibration within ±0.1 C^o over a period of several months. With optional additional sensors for pitch, roll, vibration, acceleration and water-flow, the instrument system has been used to measure the performance of underwater towed vehicles and plankton samplers. The Marine Environmental Recorder is being incorporated into an instrument system in a data buoy, for automatically monitoring the marine environment in estuaries around the British Isles.     

Plankton of the fladen ground during FLEX 76 III. Vertical distribution,population dynamics and production of Calanus finmarchicus (Crustacea: Copepoda)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR), the Longhurst Hardy Plankton Recorder (LHPR) and by our colleagues from other participating Institutes during the Fladen Ground Experiment (FLEX 76) were used to describe the vertical distribution and population dynamics of Calanus finmarchicus (Gunnerus) and to provide estimates of the production and carbon budget of the population from 19 March to 3 June, 1976. Total production of the 19 March to 3 June, 1976. Total production of the nauplii and copepodite stages (including adults), during the exponential growth phase in May, was estimated to be in the range of 0.49 to 0.91 g C m^-2 d^-1 or 29.0 to 55 g dry wt m^-2 (14.5 to 27.8 g C m^-2) for the three successive 10 d periods in May. Two gross growth efficiencies (K ₁) (20 and 34%), together with the lower value of C. finmarchicus production, were used to calculate the gross ingestion levels of algae as 2.45 and 1.44 g C m^-2 d^-1 (73.5 and 43.2 g C m^-2 over the May period). These ingestion levels, together with the algae ingested by other zooplankton species, are greater than the estimated total phytoplankton production of 45.9 g C m^-2 over the FLEX period. A number of factors are discussed which could explain the discrepancies between the production estimates. One suggestion is that the vertical distribution of the development stages of this herbivorous copepod and their diel and ontogenetic migration patterns enable it to efficiently exploit its food source. Data from the FLEX experiment indicated that the depletion of nutrients limited the size of the spring bloom, but that it was the grazing pressure exerted by C. finmarchicus which was responsible for the control and depletion of the phytoplankton in the spring of 1976 in the northern North Sea.JONSDAP Contribution No. 51     

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.     

Ecological investigations of the late-stage phyllosoma and puerulus larvae of the western rock lobster Panulirus longipes cygnus

The distribution and abundance of the late-stage phyllosoma larvae of Panulirus longipes cygnus George and the distribution and densities of the final larval stage, the puerulus, both in the plankton and at settlement along the coast, were investigated. A total of 3,617 late-stage phyllosoma (Stages VI to IX) and 301 puerulus larvae were caught at 187 plankton stations during the July to November periods 1974, 1975 and 1976 off the west coast of Australia between 29°00 to 32°30S and 113°30 to 115°00E. The depth range sampled was 0 to 35 m on the continental shelf and 0 to 90 m off the shelf. During onshore/offshore cruises with similar sampling effort on and off the shelf, 1,169 late-stage phyllosoma larvae were taken, of which only 9 were caught on the shelf, and these near the outer edge. A series of cruises sampling two areas beyond the shelf near 29°30 and 32°00S yielded 2448 late-stage phyllosoma, with greater densities of larvae in the northern location. The settlement of puerulus-stage larvae along the coast in the same geographical range was also greater in the north than in the south. The data from the onshore/offshore cruises showed a definite effect of moon phase on numbers of puerulus larvae caught on the shelf, with higher catches near new moon. The low numbers of puerulus larvae (usually 0, 1 or 2 individuals) caught at all stations showed that the puerulus stage is sparsely distributed in the plankton. Fewer puerulus larvae were present at the surface than at lower depths, but it was not possible to determine a depth preference for the puerulus between 10 m and the lowest depths sampled because of the low catch numbers. No relationships were found between puerulus larvae density and surface-water temperature, salinity, or plankton biomass at each station. Data on the larval distributions indicate that, near the end of their planktonic existence, the majority of the late-stage phyllosoma larvae of P. longipes cygnus are not carried onto the shelf, where mixing of oceanic and continental shelf waters occurs only on the outer third, but are transported southward by oceanic circulation beyond the shelf. The puerulus moults from the last phyllosoma stage beyond the shelf and completes the larval cycle by swimming across the shelf and settling in the shallow reef areas.     

Cod end feeding by the euphausiid Meganyctiphanes norvegica

Daytime surface swarms of Meganyctiphanes norvegica in the Bay of Fundy were sampled by hand-held dip nets and by plankton nets. A comparison between the food baskets of the animals from samples obtained by the two collection methods showed that those from plankton nets contained many items whereas those from dip nets rarely contained anything. Samples from towed-nets also had significantly more animals with fuller stomachs than those obtained by dip nets at approximately the same time. It is concluded that the observed differences are due to feeding by the euphausiids in the cod end of plankton nets.     

Dependence of dissolved carbohydrate concentrations upon small scale nanoplankton and bacterioplankton distributions in the western Sargasso Sea

Significant correlations between trophic components of the <20 m microbial plankton and the concentrations of total dissolved carbohydrate (TCHO) and polysaccharide (PCHO) have been found at two drogued buoy stations which were sampled at 3 or 4-h intervals over diel cycles. An attempt was made to sample on two pre-selected isotherms (approximating isopycnals) at each station. Significant inverse correlations of TCHO and of PCHO with numbers of phototrophic nanoplankton (PNAN) were found along both isotherms at a station off the Carolina coast. Off central Florida, TCHO and/or PCHO were significant multiple linear functions of PNAN, heterotrophic bacterioplankton (BAC) and sometimes heterotrophic nanoplankton (HNAN) counts. Partial regression coefficients for PNAN were always negative and those for HNAN were positive. Inverse trends dominated the relations of BAC with PCHO and TCHO. The combined data from the two stations (300 nautical miles apart) produced very similar multiple linear relations. This suggests that the observed relationships are real, resulting from general physiological processes and interactions of the microbial plankton groups rather than from chance occurrences at a particular station, and that the combined activities of the <20 m plankton actively regulate dissolved carbohydrate concentrations in the Sargasso Sea.     

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.     

Aspects of the community ecology of deep-sea,benthopelagic plankton,with special attention to gymnopleid copepods

Benthopelagic plankton was collected 10 to 100 m above the bottom of the deep sea (1100 to 3200 m) with an opening-closing net. Five samples were taken from the San Diego Trough, 2 from the eastern tropical Pacific and 2 from the northeast Atlantic. Roughly 80% of the individual organisms collected were copepods; the next most abundant groups were usually isopods, ostracods, and chaetognaths. Within the depth range studied, absolute abundances of most animal groups decreased with depth, but relative abundances (%) did not change. Of over 100 species of adult gymnopleid copepods collected, a few species were abundant, but most were rare. Copepod exoskeletons were more abundant than living copepods. For the gymnopleids, similarities among the 3 geographical regions were strong at the family and genus level but weak at the species level. Species that were widely distributed horizontally tended to occupy different positions (in terms of rank order of abundance) in each community. Gut content analysis indicated that the most abundant gymnopleids were generalized particle feeders. However, size-frequency distributions provided indirect evidence for niche separation. The deep-sea benthopelagic plankton was divisible into a truly planktonic component and a bottom-associated component. Abundances of some copepods changed relative to distance off the bottom. Several copepods and other animals, previously described as epibenthic or bottom-associated, were collected more that 10 m above the bottom and thus appeared to move freely between the bottom and lower water column.     

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.     

Ecology of a Caribbean coral reef. The Porites reef-flat biotope: Part II. Plankton community with evidence for depletion

A quantitative assessment of drifting net plankton crossing a reef-flat biotope was obtained on a Caribbean coral reef. The spatial distribution and abundance of plankton were sampled to provide estimates of the removal of this potential food resource by suspension-feeding populations. Sampling was largely confined to the reef flat and adjacent waters of Laurel Cay, a flourishing coral reef present on the insular shelf off southwestern Puerto Rico. A prior study provides information on the meteorological and hydrographic characteristics of this area. Evidence for plankton accrual was found in the quantitative depletion of qualitatively similar populations sampled downstream of densely populated reef communities. Numerically, the diatom crop was reduced by 91% and zooplankton by 60% in water streaming off the reef. Significant diel and seasonal variations in plankton abundance were obcerved, as well as notable differences in volume flow, the latter closely related to the local wind regime. A time course of net plankton accrual was calculated, taking into account these various factors. During the summer season (July–August), when zooplankton was relatively abundant and water movement over the reef vigorous, the total gain from plankton reached 0.25 gC/m²/day; 75% of this occurred during a 4 h period at sunrise and sunset. Plankton retained on the reef flat in January of February and in September was around 0.1 g C/m²/day. Zooplankton biomass contributed the greatest share, exceeding that of diatoms by a factor of 10 during the day and 42 in the early evening. A mean annual accrual of 0.18 g C/m²/day is equivalent to 4 to 13% of net community metabolism.     

Polychaete assemblages along a rocky shore on the South Adriatic coast (Mediterranean Sea): patterns of spatial distribution

This study reports on the distribution of polychaete assemblages along a depth gradient from 5 to 25 m, along a stretch of rocky coast near Otranto (South Adriatic sea, Italy). Three depths were sampled in May and November 2000. At each depth three sites (about 100 m apart) were sampled by scraping off three replicate quadrats of 20×20 cm. The experimental design enabled identification of across- and along-shore spatial patterns of variation of polychaete assemblages. A total of 4,168 specimens, belonging to 152 taxa were collected. Multivariate analysis showed that the polychaete assemblages differed significantly among depths with the clearest differences between the shallowest sites (5 m) and the deeper ones (15–25 m). A considerable source of variation among sites at each level of the shore was also exhibited by the analyses, with the greatest differences among sites within depths recorded at 5 m. The species most contributing to the differentiation of assemblages among depths and sites within each depth were identified. Some potential causes of the observed differences are hypothesized and discussed. The importance of quantitative observation to provide the context for studying the underlying ecological processes is also stressed.Communicated by R. Cattaneo-Vietti, Genova     

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.     

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.     

Distribution and community structure of the benthic fauna of the north shore of the Gulf of St. Lawrence described by numerical methods of classification and ordination

The invertebrate benthic macrofauna of the northern part of the Gulf of St. Lawrence was sampled with a van Veen grab along a series of transects from 37 to 285 m in May, 1981. The pattern of spatial distribution was determined by classification and ordinastion techniques and by diversity and evenness statistics. Classification and ordination resulted in eleven station groups which could be divided into three broad faunal zones related to temperature, depth and sediment composition. The proportions of suspension feeders increased with sand content and nonselective deposit feeders increased with mud content. Faunal diversity decreased with depth.     

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.