Exuviaella cordata red tide in Bulgarian coastal waters (May to June 1986)

The structure and some functional characteristics of the plankton community at the time of aExuviaella cordata red tide were investigated in Burgas Bay, Bulgaria, in May and June 1986. Characteristics of main plankton components (phyto-, bacterio-, nanoheterotrophic plankton, ciliates and mesoplankton) in the bloom area are presented. Development of theE. cordata bloom was determined by abiotic conditions among which eutrophication and salinity decrease caused the patchy character of its rapid development. Attainment of maximum red tide (ca. 1x10⁹ cells l^-1; 1x10³ g m^-3) from background (500 to 800x10³ cells l^-1; 600 mg m^-3) took 3 to 7d. Growth rate () during that period was 1.2 to 2.2 doublings per day. A sharp bloom decline (3 to 4d) was caused by parasitic flagellates destroying the alga's chloroplast. Diel biomass losses due to grazing remained below 5%. Metabolites and degradation products ofE. cordata revealed no pronounced toxic effects on the other components of the planktonic community. The rapid bloom degradation due to the effects of parasitic flagellates indicates the high potential of ecosystem self-regulation.     

Diurnal vertical migration of Solmissus albescens (Hydromedusae) in the Southern Adriatic

During a study on diurnal vertical migration in the hydromedusa Solmissus albescens (Gegenbaur, 1856), 112 net samples were taken from a depth of 1000 m to the surface. Sampling was performed in July, August, and September, 1969, at a station located 20 miles SSW of Dubrovnik (Yugoslavia). S. albescens migrates vertical distances of over 300 m. No significant influences on migration were noticed due to environmental factors (temperature, salinity, vertical water currents, etc.). The hydromedusa was able to approach surface layers. The recording obtained at various water-depth levels are considered in relation to illumination, mechanisms of evolution, and feeding of S. albescens.     

Comparison of cellular and molecular cytotoxic mechanisms of Cochlodinium polykrikoides in isolated trout and rat hepatocytes

Harmful algal blooms produced by the marine ichthyotoxic dinoflagellate Cochlodinium polykrikoides are responsible for mass mortalities of wild and farmed fish globally. This study compared the cytotoxic mechanisms of C. polykrikoides total extract on both trout and rat liver hepatocytes. Trout hepatocytes were more sensitive than rat hepatocytes against C. polykrikoides extract. The effective concentration 50 after 3 hour incubation (EC50_3hr) concentrations found for C. polykrikoides extract in trout and rat hepatocytes (i.e., 50% membrane lysis in 3 hr) were Eq. 1 cell/ml and Eq. 240 cell/ml, respectively. C. polykrikoides extract exposure in both isolated trout and rat hepatocytes resulted in membrane lysis, reactive oxygen species formation, glutathione depletion, collapse of mitochondrial membrane potential, ATP depletion, increase in adenosine diphosphate (ADP)/adenosine triphosphate (ATP) ratio, cytochrome c release into the hepatocyte cytosol, and activation of caspases cascade. Trout hepatocyte toxicity was also associated with lysosomal membrane injury. Mitochondrial permeability transition in both trout and rat hepatocytes produced cytochrome c release from the mitochondrial intramembrane space into the cytosol. Thus, the cytochrome c release triggered activation of caspase-3 and apoptosis. Finally, data demonstrated that C. polykrikoides extract may induce more apoptotic phenotype in rat than trout hepatocytes, which in the latter favored predominantly necrotic mode of cell death.     

Diurnal vertical migration of an estuarine cladoceran,Podon polyphemoides,in the Chesapeake Bay

Populations of the marine cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) zooplankton typically occur in large patches, extending the length of the estuary. These patches are characterized by well defined maxima, which do not move seaward during a given season. During the day, the bulk of the population is situated in the upper part of the two-layered estuarine circulation, where it is subjected to transport by net non-tidal seaward current. One mechanism for maintaining the population within the estuary is provided by the diurnal behavior of the podonids, which migrate to deeper water during the night. The landward advection of the deeper currents, thus, helps to counterbalance seaward population loss during the day. The extent of the vertical movement of the population can be modified by various environmental factors, but a suggestive correlation with light was found. It is thought that the downward component of the migration is due to passive sinking.     

Diurnal migration and vertical distribution of phyllosoma larvae of the western rock lobster Panulirus cygnus

Studies off the west coast of Australia showed that the phyllosoma larvae of Panulirus cygnus George undergo a diurnal vertical migration, with light as an important factor influencing the depth distribution of all 9 phyllosoma stages. The early stages (I to III) occurred at the surface at night regardless of moonlight intensity, whereas late stages (VI to IX) concentrated at the surface only on nights with less than 5% of full moonlight. Midday peak densities of early-stage larvae occurred in the 30 to 60 m depth range while those of mid (IV to VI) and late stages were in the 50 to 120 m range. Depths of peak densities of larvae increased with distance offshore. The limits of vertical distribution of the phyllosoma remained within ranges of illuminance which were estimated to be in the order of 50 to 250 E m^-2 sec^-1 for early stages, 20 to 200 E m^-2 sec^-1 for mid stages and 5 to 50 E m^-2 sec^-1 for late stages. Minimal rates of net vertical movement were estimated for the larvae. Early stages exhibited mean net rates of ascent and descent of 13.7 and 13.0 m h^-1, respectively, while the rates for mid stages were 16.0 and 16.6 m h^-1 and for late stages 19.4 and 20.1 m h^-1. Diurnal migrations and vertical distribution are shown to have a vital role in the relationships between circulation in the south-eastern Indian Ocean and the transport and dispersal of the phyllosoma larvae. The diurnal migrations of early stages place them at the surface at night, when offshore vectors of wind-driven ocean-surface transport dominate, and below the depth of wind-induced transport during the day, when offshore vectors are small or negative, thus accounting for their offshore displacement. Mid and late stages, because of their deeper daytime distribution and absence from the surface on moonlight nights, are predominantly subject to circulation features underlying the immediate surface layer. This is hypothesized to account for the return of the phyllosoma to areas near the continental shelf edge by subjecting them to a coastward mass transport of water which underlies the immediate surface layer.The western rock lobster is referred to as P. longipes or P. longipes cygnus in some of the literature quoted; these are synonymous with P. cygnus.     

A dispersal model for larvae of the deep sea red crab Geryon quinquedens based upon behavioral regulation of vertical migration in the hatching stage

Behavioral responses to gravity, hydrostatic pressure, and thermoclines are described for Stage I zoeae of the deep sea red crab Geryon quinquedens Smith. Survival and rate of development as a function of temperature is presented for all larval stages. Although temperatures between 10° and 25°C have no direct effect upon survival, development time is five times longer at 10°C than at 25°C. Stage I larvae show strong negative response to gravity. Swimming rate increases with an increase in pressure up to 20 atm above ambient at 11°C, but not at 15°C. Swimming rates at 15°C are higher than those measured at 11°C at each pressure tested. Stage I larvae readily penetrate sharp thermoclines. Potential dispersal ranges of G. quinquedens larvae in the Mid-Atlantic Bight are suggested based on larval behavior, development time, and coastal hydrography. A testable recruitment model is proposed for G. quinquedens.Contribution no. 1365 of the Center for Environmental and Estuarine Studies     

Comparison between remote-sensed data and in situ measurements in coastal waters: The taranto sea case

Monitoring and managing small coastal ecosystems requires a considerable understanding of the temporal dynamics of biophysical factors describing the coastal water systems. For this reason, daily observation from space could be a very efficient tool. The objective of the work described in this paper is to evaluate the contribution of remote sensing to the continuous monitoring of coastal areas. It is well known that in coastal areas, the presence of inorganic suspended sediments and coloured dissolved organic matter can make chlorophyll-concentration measurements from remote sensing difficult. To overcome these difficulties, an alternative approach to the SeaWiFS standard chlorophyll algorithm is presented, based on a semi-analytic model for sea water and on the use of MODIS data as input in a model for atmospheric effects removal. Moreover, land contamination (mixed sea–land pixels) can introduce ambiguities in sea-surface temperature measurements from remote sensing. This paper proposes the use of a hydrodynamic model as a time–space interpolator of in situ campaign data, to extensively validate the temperature values extracted from AVHRR sensor. We validated the proposed approach, using experimental field data collected over a two-year campaign in the Taranto Gulf. The results seem to indicate a good agreement between remote-sensed and in situ data.     

Horizontal and vertical trends in the distributions of larval fishes in coastal waters off central New South Wales,Australia

Larval fishes were sampled across six transects perpendicular to a 50 km section of the coast off Sydney, Australia, in April/May and August/September 1990. Samples were collected at the surface and at depth (20 to 30 m) at three locations across each transect; over the 30, 70 and 100 m depth contours. There was a large level of heterogeneity in the horizontal and vertical distributions of most taxa examined, and no general pattern of distribution was evident for the whole assemblage. Classification analyses revealed that the major differences between assemblages were related to depth. Horizontal trends in the distributions of the abundant taxa were evident in the inshore-offshore direction, but not longshore. Seven taxa belonging to the families Gobiidae, Labridae, Sillaginidae, Sparidae, Ambassidae, Clupeidae and Clinidae/Tripterygiidae were most abundant inshore, whereas 4 taxa of the families Cepolidae, Percichthyidae, Cheilodactylidae and Gonorynchidae were generally more abundant offshore and 24 taxa showed no discrete horizontal trends across transects. More taxa and individuals were generally caught at depth than at the surface and this was evident across all transects. Twenty taxa were more numerous at depth, whereas 4 taxa, the Cheilodactylidae, Gonorynchidae, Mullidae and Scorpididae, were most abundant at the surface and 11 taxa showed no difference in densities between depths. Ontogenetic differences in the distributions of some larvae were evident. The mean size of larval Liza argentea (Mulgilidae) caught was greater offshore than inshore, and greater at the surface than at depth. In contrast, larger Pseudocaranx dentex (Carangidae) occurred in greater numbers at depth than at the surface. The data emphasise that the assemblages of larval fishes in coastal waters off central New South Wales cannot be modelled as a single unit, which concurs with the findings in other temperate and tropical vaters. Furthermore, the data denote the need to spatially stratify sampling in these waters in order to assess seasonal changes in these assemblages.     

Dimethylsulfide and the algaPhaeocystis pouchetii in antarctic coastal waters

The concentrations of dimethylsulfide (DMS), dimethylsulfoniopropionate (DMSP) and dimethylsulfoxide (DMSO) were measured in water collected from the Southern Ocean 10 km offshore from Davis Station, Antarctica, during the period May 1987 to January 1988, inclusive. During winter and spring, when the sea-ice was up to 1.9 m thick, DMS, DMSP and DMSO concentrations were low (0.2 to 1.5 nM), as were phytoplankton numbers. The maximum concentration of the sulfur compounds generally occurred in the top 10 m of the water column. DMS levels rose dramatically from early December onwards, reaching a peak of 290 nM at a depth of 15 m in January. This concentration is higher than reported elsewhere in the ocean. These high concentrations occurred at the same time as a bloom of the algaPhaeocystis pouchetii. A significant correlation occurred between DMS concentration and cell numbers of the alga. The ratio of DMS concentration to the number of cells of the alga was considerably higher than reported for blooms of this species elsewhere in the ocean. Up to 10% of the total global flux of DMS to the atmosphere may emanate from antarctic seas.     

Diurnal variation of nitrogen cycling in coastal,marine sediments

A closed chamber technique was developed to determine the emission of microbially produced N₂O from an estuarine sediment. A diurnal variation was observed; maximum emissions of 0.4 to 4.0 mol N₂O–N m^-2 h^-1 were recorded at night whereas the rates were low or even negative, -0.4 to 0.4 mol N₂O–N m^-2 h^-1, during the day. The bacterial denitrification located in the uppermost centimeter was apparently the major source of the emitted N₂O. The diurnal emission pattern was thus inversely related to the O₂ availability at the sediment surface; in the dark, the lack of O₂ production by benthic photosynthesis allowed the denitrification to occur closer to the sediment-water interface and was likely to enhance the release of N₂O to the water. The daily averages for the emission were about 40 mol N₂O–N m^-2 d^-1 for three investigation periods in autumn (November), winter (February) and spring (April), whereas no significant emission was recorded in the NO ₃ ^- -depleted sediment in early summer (June). In this estuary, the N₂O emissions from the sediment were significant contributions to the overall release of N₂O to the atmosphere.     

Aspects of iron and nitrogen nutrition in the red tide dinoflagellateGymnodinium sanguineum

Iron-stress-mediated effects on biochemical constituents of the red tide dinoflagellateGymnodinium sanguineum Hirasaka were examined in 1988 by comparing Fe-replete and Fe-deplete batch cultures. The influence of nitrogen source (NO₃ or NH₄) on characteristics of Fe-deplete cells was also studied [i.e., Fe-deplete/NO₃-grown (= — Fe/NO₃) vs Fe-deplete/NH₄-grown (= — Fe/NH₄)]. Common to both N sources were reductions of chlorophylla (chla) and Fe quotas (per cell volume) by 75% and ca. 1.5 orders of magnitude, respectively, under Fe depletion. The Fe requirement ofG. sanguineum exceeded those of certain neritic diatoms by one to two orders of magnitude. — Fe/NH₄ cells exhibited 30 to 50% greater N quotas and free amino acid:protein ratios than did Fe-deplete cells grown on NO₃. In vivo fluorescence:chla increased with Fe deficiency particularly in — Fe/NO₃ cultures, surpassing — Fe/NH₄ values by ca. two-fold. Effects of Fe depletion were consistent with this element's essential role in the biosynthesis of chla and components of the photosynthetic electron transport (PET) system, and also in NO₃ utilization. Fe:N ratios were larger (1.5-fold) for iron-deficient NO₃-grown than NH₄-grown cells, likely reflecting the Fe content of NO₃ assimilatory enzymes [nitrate (NR) and nitrite (NiR) reductase] and of electron transport components needed to provide reductant, coupled with a diminished capacity of — Fe/NO₃ cells to acquire and assimilate nitrogen. Indicators of PET efficiency suggested that under iron stress, supply of Fe for NR and NiR is partly at the expense of iron-containing PET components. Utilization of nitrate by NO₃-grown cells was inhibited sufficiently by Fe depletion to yield symptoms bordering on N deficiency. In an ecological context, the most important effect mediated by nitrogen source may be the determination of critical Q_Fe (i.e., Fe required to just sustain maximal growth), thereby regulating the degree of growth limitation for a given subsaturating iron concentration.     

Microzooplankton grazing and selectivity of phytoplankton in coastal waters

Microzooplankton grazing activity in the Celtic Sea and Carmarthen Bay in summer 1983 and autumn 1984 was investigated by applying a dilution technique to high-performance liquid chromatographic (HPLC) analysis of photosynthetic pigments in phytoplankton present within natural microplankton communities. Specific grazing rates on phytoplankton, as measured by the utilisation of chlorophyll a, were high and varied seasonally. In surface waters during the autumn, grazing varied between 0.4 d^-1 in the bay and 1.0 d^-1 in the Celtic Sea, indicating that 30 and 65% of the algal standing stocks, respectively, were grazed daily. Grazing rates by microzooplankton within the thermocline in summer suggest that 13 to 42% of the crop was grazed each day. Microzooplankton showed selection for algae containing chlorophyll b, in spite of a predominance of chlorophyll c within the phytoplankton community. Changes in taxon-specific carotenoids indicated strong selection for peridinin, lutein and alloxanthin and selection against fucoxanthin and diadinoxanthin. This indicates a trophic preference by microzooplankton for dinoflagellates, cryptophytes, chlorophytes and prasinophytes and selection against diatoms, even when the latter group forms the largest crop within the phytoplankton. Interestingly, those algal taxa preferentially grazed also showed the highest specific growth-rates, suggesting a dynamic feed-back between microzooplankton and phytoplankton. Conversion of grazing rates on each pigment into chlorophyll a equivalents suggests firstly, that in only one experiment could all the grazed chlorophyll a be accounted for by the attrition of other chlorophylls and carotenoids, and secondly that in spite of negative selection, a greater mass of diatoms could be grazed by microzooplankton than any other algal taxon. The former may be due either to a fundamental difference in the break-down rates of chlorophyll a compared to other pigments, or to cyanobacteria forming a significant food source for microzooplankton. In either case, chlorophyll a is considered to be a good measure of grazing activity by microzooplankton.     

天津近岸海域微塑料污染现状分析

微塑料(microplastics,MPs)已成为继气候变化、臭氧问题、海洋酸化之后新的全球重大环境问题,为揭示天津近岸海域微塑料分布规律和影响因素,利用表层现场采样、密度悬浮法分离、光学显微镜相结合的方法,研究了天津开阔海域的3个断面微塑料的丰度分布,分析了微塑料的粒径范围、形状类型和化学成分.结果表明,所监测开阔海域的3个断面中,海水样品微塑料丰度在210—1170个·m~(-3)之间,均值为612个·m~(-3);微塑料类型包括细长形的纤维、不规则状的碎片和薄膜,纤维和碎片占到绝大多数,分别占到52.1%和46.8%;而检出微塑料颗粒粒径同大多数研究相同,粒径越小,含量越多,其中55.3%的微塑料颗粒粒径是小于0.5 mm;通过Mphunter软件分析出表层海水中微塑料的颜色以深色系(如:蓝色、黑色、红色)居多,少数有黄色、绿色、白色;利用Bio-Rad Knowltall光谱分析出海水中微塑料主要成分有PP-PE(聚丙烯-聚乙烯)、ABS(苯乙烯-丁二烯-丙烯共聚物)、CE(纤维素塑料)、EP(环氧树脂)、PA(聚酰胺)、PE(聚乙烯)、PET(聚对苯二甲酸乙酸脂)、PP(聚丙烯)、PVC(聚氯乙烯)9种塑料类型,含量由高到低依次为:PETPP-PEEPCEABSPEPAPVCPP.结果分析表明,天津近岸海域微塑料污染受到海洋产业发展的影响,如海洋渔业、港口运输等,同时由于天津属于工业化城市,人口密度也较大,因此海洋微塑料污染也受到陆源污水排放的影响.     

Dissolved nutrient dynamics along the southwest coastal waters of India during northeast monsoon: a case study

Dissolved nutrients, Chl-a and primary productivity were measured from seven transects along the coastal waters of the southeastern Arabian Sea during northeast monsoon. Ten major estuaries were chosen to study the influence of estuarine discharge on the nutrient dynamics in the coastal waters. The mean water discharge of the estuaries in the north (64.8?±?18?×?10⁵?m³?d^?1) was found to be higher than those in the south (30.6?±?21.4?×?10⁵?m³?d^?1), whereas the nutrient concentrations were found to be higher in the estuaries of the south. The results from the offshore waters were discussed in accordance with the depth contour classification, that is, shelf (depth?≤?30?m) and slope waters (depth?≥?30?m). Our results suggest that the estuarine discharge plays a major role in the nutrient distribution in near shore shelf waters, whereas in shelf and slope waters, it was mainly controlled by in situ biological processes. The inorganic form of N to P ratios were found to be higher than Redfield ratio in slope waters when compared with shelf waters, suggesting that PO₄^3? (<0.15?µmol?L^?1) is a limiting nutrient for primary production. The multivariate statistical analysis revealed that the nutrient dynamics in the coastal waters was controlled by both biological and physical processes.     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Use of seaweeds for monitoring trace elements in coastal waters

Concentrations of a wide range of trace elements: arsenic, cadmium, cobalt, chromium, hafnium, nickel, thorium, uranium, zinc and the rare earth elements, cerium, europium, samarium, terbium and ytterbium were determined by instrumental neutron activation analysis in the brown alga,Fucus vesiculosus from Eckwarder Hörne, North Sea and from Rügen, Baltic Sea. Another brown alga,Sargassum filipendula from Sri Lanka, Indian ocean (representing an unpolluted control station) was similarly investigated. Cobalt, chromium and nickel concentrations were highest inF. vesiculosus from the North Sea while zinc was highest in samples from the Baltic Sea, reflecting high levels of these elements in coastal waters of the North and the Baltic sea. Cadmium, cobalt, nickel and zinc levels were lowest inS. filipendula from Sri Lanka, probably demonstrating lower levels of those elements in coastal waters. Concentration levels of hafnium, thorium, uranium, and the rare earth elements were highest inS. filipendula. Two years later in 1994,S. filipendula along withUlva sp. (green alga) was resampled from the same sampling site, and in addition to the above elements, six other trace elements (Ag, Ba, Br, Rb, Se and Sr) were determined.Sargassium filipendula showed a particular affinity for Ag, As, Br and Sr. For the other elements, marginal concentration differences were observed betweenS. filipendula andUlva sp., probably reflecting the regional background levels. Substantially higher concentrations of Hf, Th, U, and the rare earths were found again in the 1994Sargassum andUlva samples, reflecting the effect of a substrate rich in rare earth elements. The brown algae used in this study may be used to monitor trace elements in coastal waters.     

Variability of dissolved organic matter in northern adriatic coastal waters

The results obtained in the four seasonal cruises planned in the PRISMA II project are reported. These concern dissolved and colloidal organic carbon, free amino acids and total dissolved carbohydrates and heterotrophic activity. Main factors controlling organic matter degradation, resulting from laboratory tests not planned in the above project, are also discussed. Dissolved organic matter shows seasonal accumulation, which may be markedly different from year to year, and large contributions by colloidal and saccharide components. Heterotrophic activities play an important role in the carbon cycle, although laboratory runs highlight limitations caused by aging of organic matter and phosphorus deficiency.     

The occurrence of glycolic acid in coastal sea water

A reasonably fast and reliable chemical method for the determination of glycolic acid in sea water is described in detail. It consists of adsorption of glycolic acid by alumina, followed by extraction with sulphuric acid and then colorimetry. The overall efficiency of recovery of this method, which is standardized internally, is approximately 67%, and about 20-fold concentration of glycolic acid is achieved. Using this method, the occurrence of glycolic acid in Ipswich Bay (Gulf of Maine, USA) was studied over a period of 6 months and was found to fluctuate between 0 and 80 g/l. The zero values occurred in January, when the phytoplankton chlorophyll values were also at their winter minimum.     

Oxygen-radical-mediated toxic effects of the red tide flagellate Chattonella marina on Vibrio alginolyticus

Toxic mechanisms of the red tide flagellate, Chattonella marina, collected in 1985 from Kagoshima Bay, Japan, were studied at the subcellular level. C. marina was found to reduce ferricytochrome c at a rate related to the concentration of plankton cells. Ca. 50% of the cytochrome c reduction was inhibited by the addition of 100 U superoxide dismutase ml^-1. These results suggest that a part of the cytochrome c reduction was caused by a superoxide anion which was extracellulary released from C. marina. Moreover, a small amount of hydrogen peroxide was detected in the C. marina suspension using the fluorescence spectrophotometric assay method. The identity of the hydrogen peroxide was confirmed by its reaction with 500 U catalase ml^-1. It is thus proposed that C. marina produces harmful active oxygen radicals and therefore exhibits a toxic effect on surrounding living organisms. In agreement with these results, C. marina strongly inhibited the proliferation of marine bacteria, Vibrio alginolyticus, in a plankton/bacteria co-culture system. The growth inhibition of bacteria caused by C. marina was related to the density and the metabolic potential of C. marina. Ruptured plankton showed no toxic effect on the bacteria. Furthermore, the toxic effect of C. marina on V. alginolyticus was completely suppressed by the addition of catalase and superoxide dismutase. In addition to these radical-scavenging enzymes, a chemical scavenger, sodium benzoate, also had a protective effect. These results suggest that oxygen radicals are important in the toxic action of C. marina.