Community structure of the benthos in Scottish sea-lochs. I. Introduction and species diversity

The diversity of the macrobenthos was measured, using the rarefaction method of Sanders (1968), from bottom samples from Loch Etive, Loch Creran and the Firth of Lorne (Scottish west coast). Each sample (representing 1.6 m² of bottom area) was accumulated as a series of separate hauls taken consecutively in a systematic pattern with a van Veen grab. Two habitats were considered: soft mud and muddy sand. Within-habitat comparison of species diversity clearly indicates that diversity is lower in Loch Etive than in Loch Creran or the Firth of Lorne. The reasons for this are probably connected with the relatively high freshwater runoff into Loch Etive, possibly limiting the survival of planktonic larval stages of the benthos. The values of species diversity measured for the soft-mud areas in Loch Creran and the Firth of Lorne are thought to be representative for this habitat along the west coast of the British Isles. They agree well with the diversity predicted by Sanders for such a boreal inshore area, where a maritime climate prevails, on the basis of his time-stability hypothesis and the results of his sampling elsewhere.     

Community structure of the benthos in Scottish Sea-lochs. IV. Multivariate spatial pattern

Most studies of spatial patterns of invertebrates in soft sediments have concentrated on populations of individual species. Those that examined patterns in communities have tended to employ categorical analytical techniques. Using macrofaunal abundance data from van Veen grab samples collected 20 to 100 m apart in known spatial arrangements from Scottish sea-lochs, the relationships between patterns in macrobenthic species composition and distances between samples were explored using matrix correlations in a non-parametric framework. Using a simple definition of spatial structure, i.e. that intersample distances are monotonically correlated with intersample species-similarity, spatial structure at each of seven stations was assessed using non-parametric Mantel tests based on rank-correlations. Changes in community structure were positively correlated with distance at all sites in Loch Etive, on both current-swept muddy sands and soft deep muds. Different components of the macrobenthos contributed to spatial pattern at each site. Simple spatial structure was also detectable at a muddy-sand site in Loch Creran, but neither on soft mud, nor at the soft mud site in the Firth of Lorne. The concept of rank-correlograms was introduced. These were used to examine the extent and form of spatial structure in different components of the macrobenthos at each site. Relationships between similarity and distance were neither simple nor consistent. Results were compared to previous studies which used the same data, and it was concluded that studies carried out at a particular scale, or on a particular component of the benthos, are unlikely to be successful in predicting spatial relationships at other scales or for other components of the benthos. Correlational rather than categorical analyses are recommended for exploratory studies of spatial relationships in the benthos. Analyses of the spatial structure at these seven sea-loch sites suggests that by ensuring that samples are at least 40 m apart an investigator is unlikely to underestimate variability or otherwise invalidate statistical analyses based on the use of the samples as replicates. Spacing samples up to 100 m apart may increase variability estimates, further reducing the chance of concluding that a difference exists when one does not. Received: 4 May 1999 / Accepted: 8 March 2000     

Diet and respiration of the small planktonic marine copepod Oithona nana

The functional responses of Oithona nana (Giesbr.) to various phytoplankton and zooplankton food species are described. The food species were divided into three size categories, the seasonal abundances of which were measured in Loch Turnaig, a Scottish sea loch in 1977. The seasonal variations in feeding rates in the sea for each size class were derived. The seasonal variation in respiration rate of O. nana was measured, and metabolic requirements were claculated as between 6 and 40% of the food material estimated as being eaten. O. nana differs from other common copepods in having a wide food-particle size spectrum and a low metabolic rate. It is suggested that these adaptations constitute the strategy whereby O. nana maintains its population levels throughout the year.     

Community structure of the benthos in Scottish sea-lochs. II. Spatial pattern

Spatial pattern in the distribution of the macrobenthos was studied from series of van Veen grab samples taken in a systematic pattern in Lochs Etive and Creran, W. Scotland. Each series was collected in a circular pattern at 7 stations. Plots of the variances of the numerical occurrences of the species in the samples against their means revealed greater deviation from the random expectation in the case of the 2 muddy sand stations than for the soft-mud stations, with only 1 exception. Two tests of goodness-of-fit to the Poisson distribution were also applied. The degree of concordance between the 3 measures of non-randomness applied was measured. The best correlation was found between values of the ratio of variance: mean and the test statistic D of the Kolmogorov-Smirnov goodness-of-fit test. It was not possible to discuss the biological basis for the pattern measured, except for a few species shown to be markedly aggregated. No clear trend emerged from attempts to detect differences in pattern with various ecological groupings of the macrobenthos. However, from the use of the test values as sample statistics illustrative of overall community pattern, it was possible to conclude that the fauna of the shallow, usually current-swept areas of muddy sand in the sea-lochs investigated was more aggregated than that on the, usually deeper, soft-mud sediments in quieter water.     

Distribution and behaviour of radiocaesium in Scottish freshwater loch sediments

The distribution and behaviour of radiocaesium have been studied in the sediments of two contrasting freshwater lochs: Round Loch of Glenhead, an acidified loch in south-west Scotland, with organic-rich sediments (≈20%C) and Loch Lomond, 35 km north-west of Glasgow, where sediments are low in organic matter (1–6%C, southern basin), but with a relatively high clay content. In the sediments of Scottish freshwater lochs,¹³⁷Cs [half life (t_1/2) = 30.23 yr] originates from fallout from nuclear weapons’ testing (1950s and 1960s) and from the Chernobyl reactor accident in 1986, which is also the source of the shorter-lived¹³⁴Cs [half life (t_1/2) = 2.05 yr]. Use of the characteristic¹³⁴Cs/¹³⁷Cs activity ratio of radiocaesium emitted from Chernobyl enables resolution of sedimentary radiocaesium profiles into the two component sources. In the organic-rich sediment of Round Loch, downward diffusion of radiocaesium in porewaters obscures its pattern of input to the loch. In the more clay-rich sediments of Loch Lomond, separate radiocaesium concentration peaks, related to atmospheric deposition maxima, are clearly discernible, although an influence of partial mixing is apparent. While the derived Chernobyl fallout inventory of radiocaesium in Round Loch sediments is broadly comparable with that for Loch Lomond, the corresponding weapons testing inventory is an order of magnitude lower than in Loch Lomond. Although Round Loch is situated in an area of known elevated Chernobyl deposition, the inventory is much lower than literature values of atmospheric deposition, indicating significant loss of radiocaesium from this loch. The weapons testing inventory in Round Loch is also lower than reported estimates, whereas in Loch Lomond the established inventories from both sources are similar to, or greater than, fallout deposition. The differences between the distribution and inventories in the two lakes confirms that radiocaesium is much less efficiently bound and is correspondingly much more mobile in the organic sediments of Round Loch of Glenhead than in the more clay-rich sediments of Loch Lomond.     

Organic detritus of a tropical estuary

Organic detritus of the Cochin Backwater (India) is largely composed of fine silt and sand particles around which organic matter adheres and forms aggregates. In addition to plankton, large quantities of benthic algae, rooted plants, animal matter, suspended soft mud and the material brought down by the rivers and land runoff constitute the main sources of detritus. Detritus sedimentation in the estuary attains its maximum from April to June. Detritus forms a major portion of seston, while phytoplankton productivity constitutes 0.1 to 1.0% of settled detritus. Detrital pigments include degraded chlorophyll (phaeophytin), which has a marked seasonal variation; detrital carbon, phosphorus and nitrogen show little change during the year. The caloric value of detritus, calculated from protein, carbohydrate and lipid fractions and also from total carbon, indicates that sedimented detritus does not have a high nutritional value. However, being a readily available material, its entry into the food chain seems to increase the efficiency of energy transfer from one trophic level to another.     

Seasonal variation in some population parameters of Euchaeta species (copepoda: Calanoida)

Studies of the biology of oceanic copepods are few relative to those of coastal species. Females of the genus Euchaeta have spermatophores attached to the genital somite by the male and carry their broods of eggs attached to this somite, so defining the breeding season. Populations of E. norvegica in the fjordic environment of Loch Etive (collected between 1971 and 1974 and from 1978 to 1979), Scotland and in the marginal oceanic region of the Rockall Trough, northeastern Atlantic Ocean (collected between 1973 and 1976) and of a further eight species of Euchaeta in the Trough are examined. Seasonal changes in the incidence of egg masses and spermatophores attached to the females, sex ratio and population numbers are determined. Sampling errors in the estimation of these parameters are larger in the oceanic Rockall Trough. The Loch Etive populations of E. norvegica produce two generations per year with a proportion, variable between years, of the population producing a third generation. This species produces a single annual generation in the Rockall Trough as do E. acuta and E. pseudotonsa. Two generations per year are probably passed through by E. gracilis in the Trough while the large bathypelagic E. sarsi may produce a single generation in the autumn of every second year. The other bathypelagic species, E. barbata, E. scotti, E. abbreviata and E. longissima, breed continuously throughout the year and no estimate of their generation times is possible. Consequently, this approach to population analysis is useful where breeding is seasonal, but resolving generation time in bathypelagic crustaceans remains a problem.     

Deposition Inputs,Water Quality and Liming Effectiveness

Rainfall quantity and quality, and stream and loch water quality have been monitored throughout the Loch Fleet Project. This has allowed the effects of liming of parts of the catchment to be monitored, and assessments made of the effectiveness and duration of these treatments. Rainfall over 1989–1993 fluctuated around the long-term average of 2100 mm. Over this time, year by year variations in acid or sulphate loading from deposition was evident and quite substantial. There was no trend, however, reflecting reductions in national sulphur emissions. Liming parts of the catchment in 1986 increased stream and loch water pH and calcium concentrations and reduced inorganic aluminium concentrations. Improved stream and loch water quality has been maintained for more than eight years since liming.     

Deposition Inputs, Water Quality and Liming Effectiveness

Rainfall quantity and quality, and stream and loch water quality have been monitored throughout the Loch Fleet Project. This has allowed the effects of liming of parts of the catchment to be monitored, and assessments made of the effectiveness and duration of these treatments. Rainfall over 1989-1993 fluctuated around the long-term average of 2100 mm. Over this time, year by year variations in acid or sulphate loading from deposition was evident and quite substantial. There was no trend, however, reflecting reductions in national sulphur emissions. Liming parts of the catchment in 1986 increased stream and loch water pH and calcium concentrations and reduced inorganic aluminium concentrations. Improved stream and loch water quality has been maintained for more than eight years since liming.     

Pre-and Post-Liming Aquatic Communities and Trout Diets

In the period following liming and before the introduction of brown trout to Loch Fleet, there was a reduction in the numbers of invertebrates, including the Chironomidae, Oligochaeta and Hydracarina in the loch, but an overall increase in invertebrates in the inlet and outlet streams. This was followed in the loch by a recovery in the numbers of certain groups (particularly the Trichoptera and Ephemeroptera) to pre-liming levels in spite of the introduction of the trout.

The diets of the trout comprised a wide variety of aquatic and terrestrial organisms ranging in size from Fprotozoans to newts and are dominated by chironomids. Opportunistic type of feeding as found in the present study appears to be typical of the brown trout and is reported for many water bodies by other workers.

Significant differences (P < 0.05 and P < 0.001) were found in the numbers and weights of certain food organisms in the stomachs of brown trout caught by different fishing methods in the summers of 1992 and 1993. Fish caught by gill net contained fewer planktonic crustaceans and chironomids, but greater numbers of trichopterans and terrestrial organisms, than in those caught by fly and spinner. the differences are probably related to differences in the depth at which the fish, caught by the various methods, were feeding.     

Growth,mortality and shell morphology of cultivated mussel (Mytilus edulis) stocks cross-planted between two Scottish sea lochs

One-year-old rope-grown blue mussels (Mytilus edulis L.) were held in experimental lantern nets and cross-transplanted between two sea lochs (Lochs Etive and Leven) on the west coast of Scotland. Growth and mortality were monitored from May 1991 to May 1992, as well as shell morphology in native and transplanted mussels. There were highly significant differences (P0.001) in growth rates between lochs; both native and transplanted mussels performed better in all shell and tissue growth-parameters in Loch Etive than in Loch Leven. Stock also had a significant (P0.05) influence on shell length and live weight, but its contribution to total variance was considerable only in the latter case. Wet and ash-free dry meat weights were governed mainly by site and to some extent by site x stock interaction. Mortality rates were quite low (6 to 7% yr^-1) and did not show any significant variation either between sites or stocks. There were significant morphological differences (ratios of shell length, height and width) between the Loch Etive and Loch Leven populations (P0.001) which persisted for one year after transfer. Stock appeared to be the main factor affecting shell morphology, as opposed to site. Linear regressions of shell height and width on length were significantly different between the native stocks, but became indistinguishable from those of the recipient populations one year after reciprocal transfer. This did not, however, conceal the effect of stock origin on dimensional ratios. It is concluded that site differences related to environmental factors, notably chlorophyll a levels and currents, and also possible stress caused by high levels of Zn and Cu, are major determinants of growth, which are of main interest to aquaculture, while morphological differences most probably result from genotypic variation.     

Skeletal growth zones as age-markers in the sea urchinPsammechinus miliaris

Growth of the inshore sea urchinPsammechinus miliaris (Gmelin) was studied in Loch Creran, western Scotland, using the skeletal growth-marker tetracycline in order to test the validity of natural growth-banding in the coronal test plates as annual age-markers. In order to test whether tetracycline affected the growth ofP. miliaris, an injected and a control group of urchins were held in identical conditions in running sea water aquaria for 21 mo from 1989 to 1991 and measured periodically. A small but significant difference in mean size of injected compared to controls was recorded at 12 mo, but none during subsequent measurements. Size measurements during the trial were consistent with an annual growth cycle, with a maximum in spring and slowing or cessation of growth during autumn/winter. Tetracycline-labelled juveniles were recovered up to 18 mo after initial tagging in mark/recapture experiments undertaken from 1987 to 1989 at two intertidal marked quadrats in Loch Creran. Large numbers ofP. miliaris were also marked with tetracycline and held for 1 yr at 10 m depth in seabed cages in Loch Creran during 1988–1990. All of the intertidal recoveries, and about 69% of the caged specimens that had been successfully labelled, showed a consistent relationship between the position of the tetracycline tag and the pattern of natural growth zones. The remainder were mostly large, slow-growing urchins with the tag positioned near the plate margin. In these the outer growth bands were closely spaced and, particularly if major growth bands seemed to be broken up into double or multiple lines, the major bands were impossible to resolve at the margin. The results support the assumption that in wild populations the dark bands visible with reflected light (translucent in transmitted light) after charring the plate are formed when skeletal growth has stopped or slowed in winter. These lie between wider, lighter coloured (opaque in transmitted light) zones of active plate growth in spring/summer. The dark band formed beyond the tag usually was made up of several closely spaced fine lines, or sometimes of two closely spaced dark bands. The wide growth zones beyond the tag, like those formed previously, usually were broken by fine, dark lines that may represent brief discontinuities in growth. From tagging, the double dark bands can be related to growth over one year; but such anomalous bands, along with the general presence of fine, dark lines interrupting the growth zones, make it difficult reliably to estimate age from the closely spaced peripheral banding on older, slow-growing urchins.     

The Brown Trout Population at Loch Fleet Eight Years After Liming

At the start of the Loch Fleet Project in 1984, the Loch and the upper 7 km of its efferent stream were found to be devoid of trout (Salmo trutta) as a result of acidification. Following the liming treatments applied to the catchment, from 1986 the formerly toxic water quality conditions (pH ∼ 4.5, calcium ∼ 1 mgl^-1, elevated aluminium and heavy metal levels) were eliminated, and trout were reintroduced on two occasions, in 1987 and 1988. A total of 520 fish were stocked, at a combined density equivalent to 5.5 kg ha^-1. Surveys of the loch and stream populations were carried out annually until 1993 to monitor their development, using a range of techniques, including electrofishing, gill-netting, seine-netting, spawner trapping and mark-release recapture methods. Length and scale- analysis were used to investigate fish growth.

The trout population in Loch Fleet expanded rapidly as a result of natural spawning in the loch's main feeder stream, augmented by the use of an artificial spawning bed which was constructed at the loch outlet in 1990. in mid-1983 the stock density, estimated by mark-recapture census methods, had increased to 24.9 kg ha^-1. Poor recruitment in the years 1991-93, however, reduced the rate of expansion and resulted in a population comprising mainly older individuals. the poor recruitment in these years was not fully explained but was not caused by water quality and was most likely a result of fry washout by spring spates.

Fish growth rates were high initially and were estimated on the basis of the Elliott trout growth model to be optimal for the prevailing water temperature regime of the loch. By 1991, growth rates had fallen, probably as a result of competition for food, but showed signs of recovery towards the end of the study period in 1993, following the period of lower population densities of young fish.

Trout rapidly repopulated the loch's outlet stream after 1987 but have remained sparse and have shown no signs of spawning within most of the stream. Water analyses have shown that the liming of the Loch Fleet catchment has minimal impact on downstream waters when flows are high, so that potentially toxic acid episodes have not been prevented.     

The Brown Trout Population at Loch Fleet Eight Years After Liming

Abstract

At the start of the Loch Fleet Project in 1984, the Loch and the upper 7 km of its efferent stream were found to be devoid of trout (Salmo trutta) as a result of acidification. Following the liming treatments applied to the catchment, from 1986 the formerly toxic water quality conditions (pH ～ 4.5, calcium ～ 1 mgl^?1, elevated aluminium and heavy metal levels) were eliminated, and trout were reintroduced on two occasions, in 1987 and 1988. A total of 520 fish were stocked, at a combined density equivalent to 5.5 kg ha^?1. Surveys of the loch and stream populations were carried out annually until 1993 to monitor their development, using a range of techniques, including electrofishing, gill-netting, seine-netting, spawner trapping and mark-release recapture methods. Length and scale- analysis were used to investigate fish growth.

The trout population in Loch Fleet expanded rapidly as a result of natural spawning in the loch's main feeder stream, augmented by the use of an artificial spawning bed which was constructed at the loch outlet in 1990. in mid-1983 the stock density, estimated by mark-recapture census methods, had increased to 24.9 kg ha^?1. Poor recruitment in the years 1991–93, however, reduced the rate of expansion and resulted in a population comprising mainly older individuals. the poor recruitment in these years was not fully explained but was not caused by water quality and was most likely a result of fry washout by spring spates.

Fish growth rates were high initially and were estimated on the basis of the Elliott trout growth model to be optimal for the prevailing water temperature regime of the loch. By 1991, growth rates had fallen, probably as a result of competition for food, but showed signs of recovery towards the end of the study period in 1993, following the period of lower population densities of young fish.

Trout rapidly repopulated the loch's outlet stream after 1987 but have remained sparse and have shown no signs of spawning within most of the stream. Water analyses have shown that the liming of the Loch Fleet catchment has minimal impact on downstream waters when flows are high, so that potentially toxic acid episodes have not been prevented.     

Offshore marine cheilostome Bryozoa from Fort Lauderdale,Florida

Cheilostome Bryozoa were collected upon marine fouling test panels exposed offshore from Fort Lauderdale, Florida, USA between 1961 and 1967. Panels were exposed at certain depths, ranging from the surface to the bottom, in several arrays extending to a distance of 4 miles offshore. Twenty-three species were identified from the 114 panels recovered and examined. Most species were found on panels in shallow water, or on panels near the bottom in deeper, offshore water. A notable exception was a large, arborescent colony of Biflustra savartii which engulfed a panel 50 feet (15 m) deep, 4 miles from shore, in the Florida current. The lack of cheilostome Bryozoa on shallow and mid-water panels at the offshore, deepwater stations was apparently due to the low number of larvae reaching them from the distant inshore colonies.     

Comparison of the sedimentation of a diatom spring bloom and of a subsurface chlorophyll maximum

The seasonal pattern of sedimentation was determined over a 8-mo investigation period covering the productive season at a permanent station in the Southern Kattegat (Denmark) in 1990. The phytoplankton succession was characterised by a 4-mo long subsurface maximum associated with the pycnocline which was entirely dominated by the dinoflagellate Gyrodinium aureolum. The bulk sedimentation of organic matter took place during this period and accounted for >60% of the annual particulate organic carbon (POC) and particulate organic nitrogen (PON) sedimentation. The spring bloom period contributed 60% of the sedimentation of intact phytoplankton cells, but only 20% of the POC and PON sedimentation. A minor fraction of the sedimenting matter from the subsurface phytoplankton maximum consisted of intact phytoplankton (<20%), suggesting that the phytoplankton was processed by heterotrophs and that it was mainly products from this activity which contributed to thevertical flux of organic matter. The variation in oxygen concentration below the pycnocline coincided with the pattern of sedimentation with a delay of 3 to 6 wk.     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.     

Community structure of the benthos in Scottish sea-lochs. III. Further studies on patchiness

Large-scale patchiness in the distribution of the benthic fauna was investigated in Loch Etive (Scottish west coast) by two series of van Veen grab hauls. Each series was taken along a traverse across the width of the loch, with sampling points about 100 m apart. One sample series was taken on sandy mud and the other, at a greater depth, on soft mud. Two approaches in data processing were applied: (1) The variance: mean ratio and the Morisita I tests for significant aggregation were applied to the species abundances in the sample series; (2) 3 measures of sample homogeneity, each involving the calculation of an index of faunal similarity, were applied to the separate samples. The results, however, could not show significant differences between the two series of samples. These findings were compared to results obtained from (a) a previous study, where differences in patchiness between areas were shown for a smaller scale of sampling design in Loch Etive and neighbouring areas, and to which the I method is also applied here; and (b) the results of applying the variance: mean ratio and I tests to data published by Holme (1953). The present results indicate greater aggregation at the present scale of sampling than for the previous, smaller scale of sampling, or for the comparable scale of Holme's sampling. The degree of concordance shown by the values of I for the circular-design sampling to the values of the 3 measures of patchiness applied previously, and to the means of the species abundances, was measured by Spearman's rank correlation coefficient. The results clearly demonstrated the I values, unlike the others measured, to be almost completely independent of the mean. It was concluded that, for comparing the pattern from benthic samples using standard-size bottom samplers, where the mean may vary widely between each set of samples, the I method is probably most useful.     

Further studies of short-term variation in the pigment composition of a spring phytoplankton bloom

Phytoplankton sampled at the chlorophyll maximum during the spring bloom at Loch Ewe, Scotland during 1986, were analysed using high-performance liquid chromatography for chloropigments and carotenoids. The day-to-day proportions of the pigments remained relatively stable throughout the bloom, with good preservation of chlorophyll a even at the sediment surface. Chlorophyllide a and phaeophorbide a appeared towards the end of the bloom. The major carotenoids remained stable throughout the bloom, but became rapidly degraded on sedimentation under the oxic conditions. The pigment data is related to the nutrient status of the bloom, when slow steady growth occurred without nutrient limitation. These data contrast sharply with data from the 1983 spring bloom at Loch Ewe when acute nutrient limitation took place.     

Bacterial activity related to sedimenting particulate matter

The rates of sedimentation of particulate matter were measured at a station in the Eckernförde Bight, Baltic Sea, using sediment traps. Different parameters (C, N, chlorophyll a, phaeopigments, adenosine triphosphate (ATP), heterotrophic activity (glucose maximum uptake velocity, v_max), biological oxygen demand (BOD), and electron-transport-system (ETS) activity) were measured during a 1-year period to evalnate the composition, vertical distribution, and activity of microorganisms associated with sedimenting particulate matter. Measurements indicated that the organic fractions of particulate sedimenting matter were transformed considerably during sedimentation. The seasonal fluctuations showed higher heterotrophic activity durang summer and lower activity during winter. Laboratory experiments also indicated that an important part of planktonic particulate material was susceptible to mineralization within a short period of time: phytoplankton was mineralized by about 35% and zooplankton by about 18% per day, at 20°C; at 5°C mineralization was considerably lower (about 3 and 8%, respectively).Please address all correspondence to: Dr. R. Zimmermann, Institut für Meereskunde an der Universität Kiel, Abteilung Marine Mikrobiologie, Düsternbrooker Weg 20, 2300 Kiel 1, Germany (FRG)