Colonisation of South African kelp-bed canopies by the alien mussel Mytilus galloprovincialis: extent and implications of a novel bioinvasion

The Mediterranean mussel Mytilus galloprovincialis is the most significant invasive alien marine species in South Africa and, although not normally found subtidally, has recently been observed colonising heads and stipes of the kelp species Ecklonia maxima in False Bay. We quantified this invasion and explored its ecological implications. Transects laid across kelp beds revealed that 10.34% of kelp individuals surveyed bore canopy mussels, with kelp heads (10.07%) being far more commonly infected than stipes (0.27%). Twenty kelp individuals with infected heads and 20 with infected stipes were separately collected for more-detailed examination. Wet mass of mussels on these kelp heads ranged from 2.5 to 2 462 g (median 86.4 g, interquartile range 14.8–353.8 g) and that of mussels on the stipes from 7.6 to 3 492 g (median 595.5 g, interquartile range 194.0–955.0 g). Mussel clumps consisted mostly of individuals <40 mm in length. Mussel clumps supported a rich biota of 80 invertebrate and 13 algal species. Larger clumps supported more epibiotic species, and those on stipes more species than those of comparable mass on kelp heads. The mussels and their associated epibiotic species negatively affected kelp buoyancy, but rarely enough to overcome natural buoyancy. Some kelp individuals that had been toppled by the weight of mussels and their epibiotic species, however, were encountered in situ. Implications of this invasion include large increases in animal biomass and species richness in the kelp canopy, plus reductions in kelp buoyancy and increased hydrodynamic drag on infected kelps, increasing their probability of being uprooted. Uprooted kelp individuals can raft long distances, potentially transporting both native and alien species to distant sites.     

Spatial and temporal distribution of floating kelp in the channels and fjords of southern Chile

Numerous studies suggest that floating macroalgae contribute to population connectivity of the algae themselves as well as of associated organisms. In order to evaluate the importance of floating macroalgae for population connectivity in southern Chile we analysed their spatial and temporal distributions. We estimated the abundance of floating kelp in the Interior Sea of Chiloé and the Los Chonos Archipelago via ship surveys conducted during austral spring (November) and winter (July) of the years 2002–2005. Highest densities of floating kelp (Macrocystis pyrifera and Durvillaea antarctica) were found during spring. Generally, the density of floating kelp was relatively low in areas with high supply of freshwater, and highest abundances were found in large channels opening to the open ocean. We suggest that this pattern is caused by the scarcity of natural kelp beds combined with rapid degradation of floating kelp in the interior fjords and by seaward surface outflow. The maximum densities of floating kelps were found in an area known to be an extensive convergence zone (i.e. estuarine front), which appears to act as a retention zone. In accordance with this distribution pattern, we propose that connectivity between local populations via floating kelp is higher in the outer channels with oceanic influence than in the inner fjords with high freshwater influx.     

Characteristics of Habitats Used by Enteroctopus dofleini in Prince William Sound and Cook Inlet,Alaska

Abstract. Habitat characteristics associated with the giant Pacific octopus, Enteroctopus dofleini (Wülker 1910), were studied in Prince William Sound and Port Graham, Alaska, from beach walk, SCUBA and submersible surveys at depths of 0 to 197 m. Octopus counts on beach walk transects were positively correlated with soft substrates (sand, gravel or broken rubble), the presence of boulders, and dense kelp cover immediately offshore of the transect; and negatively correlated with depth on SCUBA transects. No significant habitat correlations were found with counts on submersible transects. On beach walks, octopus counts were reduced on hard substrates to 38 % of the counts on soft substrates. Counts increased five‐fold in the presence of boulders over counts in their absence, and increased fifteen‐fold adjacent to dense (> 75 %) kelp cover over counts adjacent to sparse (< 25 %) kelp cover. On SCUBA transects, the average density at less than 5 m depth was over five times that below 5 m. No trends in octopus size or sex ratio were detected with depth. Den use was inversely correlated with depth although there was no indication that den availability declined with depth. Octopuses were found at densities from 0 to 2.5 per 1000 m². These densities were only 1 to 50 % of densities of the same species recorded in British Columbia in the late 1970s and early 1980s. Few data are available to test recruitment, mortality, and habitat selection hypotheses that would account for differences between habitats. However, the presence of the highest octopus densities in intertidal and very shallow subtidal areas indicates the likely importance of near‐shore, shallow‐water habitats, and highlights the vulnerability of octopus populations to changes in these habitats.     

Enhalus acoroides responses to experimental shoot density reductions in Haad Chao Mai National Park,Trang Province,Thailand

The effect of self‐shading and competition for light in the seagrass Enhalus acoroides were investigated with a density reduction experiment in Haad Chao Mai National Park, Trang Province, Thailand. The study was carried out in a monospecific meadow with a natural density of 141.0 ± 8.7 shoots·m^?2. The intent was to determine the response of E. acoroides beds to loss of shoots and thinning, which often occur during typhoons and severe storm activity. Permanent quadrats were manipulated by clipping the seagrass shoots to 140, 72, 36 and 16 shoots·m^?2, to yield natural, 50%, 25% and 10% densities, respectively. Reducing shoot density in E. acoroides increased underwater light intensity below the canopy, generating increased leaf surface area and shoot weight. Seagrass leaf width, growth rate, and number of leaves per shoot also increased with greater light. The extent of flowering varied among treatments with no consistent trend. Our results demonstrate that increasing the available light to E. acoroides produces an increasing leaf size response as self‐shading in the bed is reduced.     

Climate driven changes in subtidal kelp forest communities in NW Spain

Reconstructions suggest a massive decline of nearly 1400 ha of kelp forest in North Western Spain in 2007. In line with global rising temperatures, we hypothesized that Sea Surface Temperature (SST) surpassed a lethal threshold for kelp. We examined whether changes in SST correlated to the proposed decline in kelp forest. All investigated SST characteristics suggested to affect kelp abundance increased significantly during the past thirty years, reaching extreme values during the last decade. In addition over the past two decades, the landscape formerly dominated by both cold and warm temperate canopy forming and understory species changed to one dominated by warm temperate understory species, resulting in a loss of vertical community structure. Fisheries landing data of kelp associated species was used to support the suggested change in kelp abundance. Subsequent recovery of the kelp appears to be occurring in deeper waters.     

Effect of the kelp Laminaria hyperborea upon sand dune erosion and water particle velocities

This paper presents a set of results from a laboratory study on water wave propagation above submerged vegetation growing in the surf zone and the effect of submerged vegetation on dune erosion. The study has focused on the kelp Laminaria hyperborea. The reason is that this kelp is commercially harvested along the Norwegian coast and there is a need to obtain better knowledge on the possible consequences of this harvesting. Experiments were run with irregular waves over a sloping bottom, and a kelp field was simulated by 5000 artificial kelp plants in a 1:10 scale. The experiments primarily focused on the effect of kelp upon erosion of a sand dune, wave damping and water velocities. It was found that the water level is a very important factor to the degree of dune erosion, while the kelp has only a minor effect. The kelp does, however, cause significant wave damping and the degree of wave breaking is reduced. It was also found that the kelp modifies the water velocity profile. In a region above the kelp canopy layer, the time-averaged water velocity was shoreward, while the seaward undertow was confined to a region higher up in the water column.     

Short-term temporal dynamics of algal species in a subtidal kelp bed in relation to changes in environmental conditions and canopy biomass

Understanding temporal variation at the scale of weeks to months is critical to understanding broad temporal patterns in diversity in the same way as understanding diversity across landscapes relies on understanding variation at the scale of meters. However, whereas small-scale spatial variation in temperate reef algal assemblages has been extensively studied, fine-scale temporal changes have not been well addressed. By sampling the macroalgae of a subtidal reef near Perth (Australia), dominated by the small kelp Ecklonia radiata, every ∼40 days over a 2-year period, we were able to test whether temporal changes in species richness, assemblage structure and species turn-over were related to seasonal changes in surface temperature, solar radiation and wave height. A total of 93 macroalgal taxa were identified, and species richness per sampling time ranged from 25 to 64 taxa 1.25 m⁻². Biomass of E. radiata was positively correlated with changes in sea surface temperature and light, and negatively correlated with wave height. Species richness, assemblage structure and turn-over of other macroalgae were more associated with seasonal changes in kelp biomass than environmental variables per se. We conclude that seasonal changes in environmental conditions drive changes in the kelp canopy, which in turn drive changes in species richness and assemblage structure. This suggests that habitat-formers such as kelps can exert a strong temporal influence on associated communities, analogous to well-described spatial influences. Thus, as kelp canopy biomass expands and retracts over time-scales of weeks to months, so does available space for colonization and growth, resulting in a high species turn-over. Species richness is therefore increased and maintained through time, in the same way as canopy-gap mosaics increase and maintain species richness across spatial landscapes.     

桑沟湾夏、秋季悬浮颗粒物的沉降通量及再悬浮的影响

应用锚式悬挂沉积物捕捉器法,研究了我国北方重要海水养殖区域桑沟湾悬浮颗粒物沉降通量的分布特征,并通过金属Al标记法,同步测定了沉降颗粒物再悬浮比率。结果表明,桑沟湾的底层悬浮颗粒物(SPM)、颗粒有机碳(POC)、颗粒态总氮(PTN)和颗粒态总磷(PTP)平均表观沉降通量分别为1 511.4g/(m2·d)、20.01g/(m2·d)、1.497g/(m2·d)和0.474g/(m2·d),显著高于我国其他近岸海域,但底层沉降颗粒物再悬浮比率平均值高达92.8%,认为在养殖内湾,受再悬浮程度的影响,测得的底层表观沉降通量是中层的2.7倍,秋季明显大于夏季,海带和扇贝养殖区大于牡蛎养殖区。经再悬浮比率校正后的净沉降通量,仍存在着显著的空间和季节变化,但受控因素发生了根本转变;这主要表现为净沉降颗粒物质主要源于生物代谢活动强烈中上层水体,贝类的排泄作用使牡蛎和扇贝养殖区的净沉降通量显著大于海带养殖区,养殖贝类个体增大、排泄量增加使秋季净沉降通量高于夏季。在我国近岸海域,再悬浮作用的影响,会对该区域悬浮颗粒物沉降通量的估算带来巨大误差,因此该作用不容忽视。     

First survey of fishes in the Betty's Bay Marine Protected Area along South Africa's temperate south-west coast

This first survey of fish in the Betty's Bay Marine Protected Area (MPA), on the south coast of South Africa, was conducted using baited remote underwater video systems (BRUVs). A total of 58 deployments recorded 42 species in 20?km², including reef, kelp and sand habitats in protected and exploited zones, at between 5 and 40?m depth. Chondrichthyans accounted for 28% of diversity. Teleost diversity was dominated by Sparidae, Cheilodactylidae, Sciaenidae and Ariidae. Diversity (H′) was highest in kelp and lowest over sand. Species composition differed among habitat and depths, but protection had no effect. Among four commercial species, only Pachymetopon blochii responded positively to protection. The apparent failure of protection may attest to poor compliance, but an investigation into fish size might show an effect. Many species were detected at the western extreme of their range. Diversity in Betty's Bay was predictably lower than in the more eastward Stilbaai MPA, but also lower than in the westward Table Mountain National Park MPA. Fish diversity did not follow a linear increase eastwards from Cape Point. Betty's Bay includes the most easterly protected kelp forests and contains seven species not recorded in the other two areas, and is therefore an important element in the MPA network.     

Kelp forests in False Bay: urchins vs macroalgae in South Africa’s southwest coast biogeographical transition zone

Kelp forests along South Africa's southwest coast occur in a region of biogeographical overlap. Commonly referred to as transition-zone kelp forests, these dynamic ecosystems are particularly susceptible to grazing from species such as urchins. This study explores relationships between urchins and macroalgae in transition-zone kelp forests along the western side of False Bay. Six kelp forests in this region were sampled. At each site, the urchin Parechinus angulosus and kelp Ecklonia maxima were counted, percentage cover of understorey algae was recorded, and drift algae were collected. Despite variability in urchin and algal cover across sites, kelps generally increased from north to south, while urchins did the opposite. Urchins were negatively associated with macroalgae, although this relationship was weak. A localised threshold of 50 urchins m^–2 was identified, above which kelp density failed to increase above 10 m^–2 and percentage cover of understorey algae usually remained below 20%. Surprisingly, no relationship was discovered between drift algae and attached algae. The findings highlight the complexity of these cool-water environments. To better understand the role of urchins in this system, particularly under changing climate trends, experimental research into the feeding behaviour and possible effects of urchins on attached algae in the presence/absence of drift algae is advised.     

Diatoms associated with two South African kelp species: Ecklonia maxima and Laminaria pallida

Kelp forests are believed to host a large biomass of epiphytic fauna and flora, including diatoms, which constitute the base of aquatic food webs and play an important role in the transfer of energy to higher trophic levels. Epiphytic diatom assemblages associated with two common species of South African kelps, Ecklonia maxima and Laminaria pallida, were investigated in this study. Primary blades of adult and juvenile thalli of both kelp species were sampled at False Bay in July 2017 and analysed using scanning electron microscopy. Our findings showed that both kelp species hosted relatively low densities of diatoms (ranging from 7 [SD 5] cells mm^?2 on adult specimens of L. pallida to 43 [SD 66] cells mm^?2 on blades of juvenile E. maxima), with Amphora and Gomphoseptatum reaching the highest absolute abundances. Although non-metric multidimensional scaling showed overlapping and largely scattered sample sets, a significant relationship between the diatom communities and the species and age of the host macroalga was detected by two-way PERMANOVA. In general, more abundant and diverse diatom communities were observed on juvenile thalli than on adult thalli, with species belonging to Navicula and Rhoicosphenia contributing significantly to the observed dissimilarity. Due to a significant interaction between species and age effects, however, the overall ability of kelp species, their age, and their interaction to explain the variation in diatom community structure was limited. We suggest that the low densities of epiphytic diatoms were directly related to the sloughing of epithelial cells observed in both kelp species. We further speculate that on such unstable substrata some diatom taxa might adapt to an endophytic life to avoid the antifouling mechanisms developed by their hosts.     

Underwater spreading and surface drifting of oil spilled from a submarine pipeline under the combined action of wave and current

Tremendous economic loss and environmental damages are caused by oil-spilling accidents in sea. Accurate prediction of the underwater spreading and surface drifting of oil spills is important for the emergency response. In the present study, numerical investigation on the underwater spread and surface drift of oil spilled from a submarine pipeline under the combined action of wave and current was carried out to examine the effects of physical ocean environment, leaking flux and spilled oil density and viscosity. Reynolds-Averaged-Navier-Stokes (RANS) equations, realizable k-ε turbulence model and volume of fluid (VOF) model are employed to describe the multiphase flow, and velocity-boundary wave-making technique combined with the sponge layer damping absorber technique realizes the numerical wave flume. Oil spill experiments were conducted to validate the numerical model. The calculation results indicate that compared with the environmental conditions of still water, only current and only wave, a larger scope of underwater spreading and relatively slower rising rate and relatively faster drifting rate of oil droplets are observed under the combined action of wave and current. The leaking flux affects the floating time and dispersion concentration, while the ocean environment affects the horizontal migration and surface drifting. Under the specific conditions of present work, oil density has obvious effect on the underwater spread but limited effect on the surface drifting, while oil viscosity has little effect on both the two processes.     

Do predatory fish of benthic crustaceans vary between kelp and barren reef habitats in northeastern New Zealand?

Fish assemblages that included known predators of benthic crustaceans were compared between kelp and barren habitats in northeastern New Zealand using baited underwater video census methods. The benthic-feeding fish were observed in winter, spring/summer and autumn, as well as during night-time in spring/summer. Overall, the fish assemblages varied between barren and kelp habitat, being most marked in winter. Individual benthic-feeding species, such as grey mao mao (Scorpis lineolatus) and leatherjacket (Meuschenia scaber) were associated with barren habitat, while the spotted wrasse (Notolabrus celidotus) was more strongly associated with kelp habitat. The results suggest that changes in habitats on coastal reefs affect populations of species that are benthic predators, which may in turn influence the distribution and abundance of their prey species, such as juvenile spiny lobsters.     

The seasonal cycle of phytoplankton biomass and primary productivity in the Ross Sea, Antarctica

Phytoplankton standing stocks and carbon assimilation were measured during four cruises to the southern Ross Sea, Antarctica during 1996 and 1997 in order to assess the details of the seasonal cycle of biomass and productivity. The seasonal composite showed that phytoplankton biomass increased rapidly during the austral spring, and integrated chlorophyll reached a maximum during the summer (January 15) and decreased thereafter. Particulate matter ratios (carbon:nitrogen, carbon:chlorophyll) also showed distinct seasonal trends with summer minima. Carbon assimilation increased rapidly in the spring, and reached a maximum of 231 mmol C m⁻² d⁻¹, ca. four weeks earlier than the maximum observed biomass (during early December). It decreased rapidly thereafter, and in austral autumn when ice formed, it approached zero. The time of maximum growth rate coincided with the maximum in C-assimilation, and at 0.66 d⁻¹ equaled predictions based on laboratory cultures. Growth rates over the entire growing season, however, were generally much less. Deck-board incubations suggested that photoinhibition occurred at the greatest photon flux densities, but in situ incubations revealed no such surface inhibition. We suggest that due to the nature of the irradiance field in the Antarctic, assemblages maintained in on-deck incubators received more light than those in situ, which resulted in photoinhibition. This in turn resulted in a 17% underestimate in on-deck productivity relative to in situ determinations. The phytoplankton bloom appeared to be initiated when vertical stability was imparted in austral spring, coincident with greater daily photon flux densities. Conversely, decreased productivity likely resulted from trace metal limitation, whereas biomass declines likely resulted from enhanced loss rates, such as aggregate formation and enhanced vertical flux of larger particles. The seasonal progression of productivity and biomass in the southern Ross Sea was similar to other areas in the ocean that experience blooms, and the cycling of carbon in this region is extensive, despite the fact that the growing season extends no more than five months.     

Secondary production in a Laminaria hyperborea kelp forest and variation according to wave exposure

The secondary production of mobile invertebrate fauna in the Laminaria hyperborea (Gunn.) Foslie kelp forest increases with wave exposure level. This faunal group has a key function in transferring kelp carbon to higher levels in the food web. By using a size-frequency method the calculated production was 68 (±18) g D.W. m⁻² yr⁻¹ (±S.E.) at low, 250 (±57) at medium and 308 (±64) at high exposure levels. The calculations included 30 macrofauna species, which accounted for 96% of the specimens registered, with Gastropods, amphipods and bivalves being the most abundant taxa. The calculated secondary production is high, but comparable to that previously reported from other macrophyte systems and was 3%, 8% and 8% of the total primary production at low, medium and high exposure levels, respectively. Our results indicate that large quantities of Laminaria kelp are exported from the system, although the production of sessile animals was not taken into account. The most important factor in determining faunal densities and secondary production was probably habitat size but at low exposure levels the percentage of egg-carrying crustacean females and juveniles were lower than at medium and high exposure levels, thereby indicating lower fitness for animals at low exposure stations.     

Physical disturbance and subtidal habitat structure on open rocky coasts: Effects of wave exposure, extent and intensity

Canopy-forming algae occur across of range of energy environments (i.e., wave sheltered to exposed coasts) where disturbances are frequent (i.e., gap formation) and benthic patterns largely reflect variation in post-disturbance processes. Disturbances vary in extent (area affected) and intensity (degree of damage), and this may affect recolonisation at local scales. On an open oceanic coast, we tested whether habitat structure (patches of canopy algae) differed between heavy and relatively lighter wave exposure (sheltered vs. exposed sides of islands), and whether wave exposure affected the response of prominent habitat-formers to varying disturbance regimes (different sizes of partial and complete canopy removal). Observations of naturally occurring patterns showed sheltered coasts to be characterised by small patches of fucoids, whereas exposed coasts were characterised by large patches of kelp. Canopy-gaps were larger at exposed than sheltered coasts, and mixed canopies constituted > 24% of the subtidal rocky habitat independently of wave exposure. Experimental disturbances showed the local density of kelps to affect recovery through greater recruitment to partial clearings (80% canopy removal). Fucalean algae, on the other hand, mainly recruited into complete clearings (100% removal), but when their recruits were abundant, they also recruited into partial clearings. The covers of filamentous, turf-forming algae increased in all clearings, and more so at exposed than sheltered sites. Extent of disturbance had no detectable effect on recolonisation by canopy-forming algae across the scales examined (i.e., 1.5 m, 3 m diameter loss of canopy). Recolonisation varied among islands kilometres apart, and correlations (r > 0.85) between cover of canopies and cover of their recruits in clearings at the scale of sites, suggested that differences in propagule supply could account for variation in patterns of recolonisation at scales of kilometres. There was no evidence to suggest that the effect of disturbance depended on wave exposure within the range of exposures tested in this study (i.e. open coasts). We recognise that wave exposure can be fundamental to habitat structure of subtidal rocky coasts, but we suggest that its influence may be mediated by the biological setting (e.g., canopy composition).     

应用生态模型研究近海贝类养殖的可持续发展

建立了一个贝藻混养生态模型，并应用于桑沟湾栉孔扇贝、太平洋牡蛎和海带混养生态系统的模拟。这一模型在个体生长模型的基础上模拟种群的生长情况；通过模拟不同播苗养殖和收获方式下的产量，以及不同混养方式对海洋生态系统的影响来确定养殖容量。采用该模型的研究结果表明：当养殖密度分别增加到目前扇贝和牡蛎放苗量的2倍和15倍时总产量最高(达到养殖容量)，但单位面积产量和产量／播苗比减少，因此效益是下降的；扇贝放苗量增加到目前的15倍，牡蛎增加到30倍时会导致养殖生产崩溃，同时生态系统也发生改变：在目前养殖密度下，桑沟湾向黄海输出初级生产产品，但是当放苗密度增加到15～20倍时，桑沟湾需要黄海向其输入初级生产产品。上述结果表明，该模型可以迅速模拟养殖生物量和生态系统的变化，在多元养殖管理中可作为一种有效的管理工具。     

Spatial Variation in the Structure of Kelp Forest Communities Along a Wave Exposure Gradient

Abstract. Four sites were sampled in kelp (Macrocysiis pyrifera) forests occupying rocky bottom habitats along a wave exposure gradient in central California. Consistent betwecn-site differences were found in the three major structural elements - the surface canopy, the undcrstory assemblage, and the ground cover/turf assemblage - of kelp forest communities. Macrocysiis pyrifera was found at all four sites. Nereucyslis tuelkeana only at the most exposed site. The understory kelps Laminaria setchellii and Pterygopltora californica were also characteristic of exposed sites. Articulated coralline algae were more abundant at exposed sites than protected, while fleshy red algae showed the opposite pattern. All four study sites were located along 8.5 km of coastline, and thus were assumed to have available to them the same species pool for colonization. The substrate composition was the same and the amount of unconsolidated substrate was similar at all four sites. We suggest that exposure to wave-generated water motion, through its influence on the surface canopy and therefore on the amount of light reaching the bottom, is responsible for these between-site differences.     

The effects of spatial targeting of fishing effort on the distribution of the Norway lobster,Nephrops norvegicus,on the Farn Deeps grounds,northeast England

Nephrops norvegicus is an essentially sedentary species of lobster that forms the basis of valuable fisheries in the northwest Atlantic and western Mediterranean. Fishers exploiting a sedentary stock are likely to visit the most profitable (highest catch rate) areas first. Such spatial targeting of fishing effort is likely to have important consequences for stock monitoring and assessment. We used underwater television surveys of Nephrops burrow densities on the Farn Deeps grounds, northeast England, to describe changes in abundance and distribution between the beginning and end of a winter fishing season. Above a threshold of c. 0.6 burrows m^–2, overwinter depletion increased with burrow density, consistent with fishing effort being targeted at the highest densities. A simple simulation model showed that this pattern of mortality is an expected consequence of spatially targeted fishing behaviour. The model also predicted that there is decreased spatial variability in density after fishing. An overall decrease in variability was not evident from the survey data, but geostatistical analysis indicated that there was “flattening” of the density profile along a north‐south axis, consistent with the dominant direction of commercial trawling. We concluded that Nephrops fishers are able to find and exploit the highest densities of their target species. A potential consequence is that catch per unit effort (CPUE) data used to monitor trends in this stock potentially could mask declines in stock abundance. CPUE might be more effective if analysed at finer spatial scales, but this is not currently possible. In the absence of these fine scale commercial data, fishery‐independent surveys (e.g., underwater television) are an important source of information on trends in stock abundance.     

Temporal variations in the concentration and settling flux of carbon and phytoplankton pigments in a deep fjordlike estuary

The weekly mass flux of C and phytoplankton pigments at five depths in the main basin of Puget Sound, a deep (200 m) fjordlike estuary, was sampled for a year with moored sequentially-sampling sediment traps. Flux measurements were compared with weekly samples of suspended pigments in the euphotic zone and bi-monthly samples of total suspended matter and particulate C throughout the water column at the mooring site.Seasonal changes in the total mass flux at all depths were small; instead, physical (river runoff, bottom resuspension) and biological (phytoplankton blooms) events caused occasional sharp increases on a weekly scale. The dry weight concentration of pigments in the trap samples mirrored the concentration of pigments in the euphotic zone suspended matter, increasing from 0·01% in winter to a maximum of 0·65% in late summer. Bloom-induced changes in the pigment concentration were observed almost simultaneously in the euphotic zone and in the traps to a depth of 160 m, indicating a rapid vertical transfer of surface-originating particles by organic aggregates. In contrast to the strong seasonal signal in the pigment concentration, C concentration varied by only a factor of three during the year.The seasonal trend of C/pigment ratios in the C flux arises from at least two sources: (1) a balance between terrestrial sources of C during the high-runoff winter season and in-situ primary production in spring and summer, and (2) cycling of C through the zooplankton population. Budget calculations suggest that the loss of primary-produced C and pigment from the euphotic zone by settling is 5% regardless of season. On an annual basis, this C flux (16 g m⁻²) is sufficient to support previously measured values of benthic aerobic respiration at the mooring site. To account for other C sinks such as burial, predation and chemical oxidation, however, terrestrial C sources and alternate transport pathways, such as vertical advection and sediment movement down the steep basin walls, are necessary.