Benthic Coupling Research Articles

Ciliated epibionts on the Arctic sympagic amphipod Gammarus wilkitzkii as indicators for sympago–benthic coupling

In September 2003 specimens of the sympagic amphipod Gammarus wilkitzkii were sampled in drifting pack ice above 50 m water depth and in the pelagic/benthic environment beneath in the coastal waters off Svalbard (Norway). Abundance values ranged between 1.5 and 8 individuals m−2 for the pelagic/benthic and the sympagic environments, respectively, and showed corresponding sex ratios of 1:1.4, favoring males. In the ice a significantly higher number of juveniles prevailed. In contrast, ovigerous females were more abundant among pelagic/benthic living specimens. Approximately 25% of the individuals hosted ciliated epibionts of the genera Ephelota, Cryptacineta, Acineta, Podophrya (all suctoria), and Epistylis (peritrichia). Cryptacineta and Ephelota were the most abundant epibionts on this amphipod species. Female specimens of G. wilkitzkii showed the highest degree of infestation (>2,100 individual epibionts per amphipod specimen: indE/A) followed by juveniles (>1,200 indE/A) and males (>220 indE/A). Highest densities of epibionts were found on anterior body parts with the antennae bearing up to 130 individuals. This is the first sighting of epibionts on crustacea from the sympagic environment. We interpret them as biomarkers that indicate the existence of sympago–benthic coupling processes between the ice and the underlying waters and the seafloor. The population structure and the proportion of infested specimens are equal for the amphipods sampled from both the sympagic and the pelagic/benthic environment, indicating the existence of exchange (coupling) processes between the two habitats. Considering the annual ice cycle, local ice drift patterns, and the shallow water depth in the sampling area, we hypothesize that ice amphipods spend ice-free periods near the seafloor where they may serve as basibionts for protozoans and eventually re-colonize the ice with the onset of ice formation. Our observations strongly emphasize that shallow coastal areas serve as both retention (rather than sink) areas for ice fauna during ice-free periods, and as stepping-stones for re-colonizing the ice when the ice is formed in winter.

Mediation of benthic–pelagic coupling by microphytobenthos: an energy- and material-based model for initiation of blooms of Aureococcus anophagefferens

We present a conceptual model for initiation of blooms of the estuarine brown-tide pelagophyte Aureococcus anophagefferens. The model is based on the observation that in addition to its well-documented stimulation by organic nutrients, Aureococcus is pre-adapted to low light levels. Its relatively low maximum (light-saturated) growth rate makes it a poor competitor with other estuarine species at high light under acclimated conditions. Its large photosynthetic antenna and relatively low quota of photoprotective pigments make it more susceptible to photoinhibition than other species to which it is compared. These same characteristics give it a competitive advantage at low light levels. In its shallow habitat, both the light level and the rate of nutrient supply from groundwater and benthic porewater are determined by the degree of benthic coupling. Experimental manipulations in a microcosm and a survey of the literature demonstrate the ability of the sediment-associated microphytobenthos (MPB) to regulate both the light- and nutrient-environment in the overlying water column. The model predicts that the growth dynamics of the MPB are such that the benthic/water column interactions tend towards one of two stable states. In one, a well-developed population of MPB restricts resuspension of particulate material and efflux of dissolved nutrients, resulting in clear and nutrient-poor overlying waters. This condition does not favor growth of Aureococcus. In the alternative state, erosion of the MPB results in turbid, nutrient-rich waters that do favor bloom initiation. Alternation between the states is caused by external physical forcing, through wind-driven mixing of the water column. Field data from Quantuck Bay, New York (USA), failed to document the transition from non-bloom to bloom conditions. Even so, they are consistent with the model’s predictions.

Organic inputs and ecosystem efficiency in the deep Mediterranean sea

Most studies on the benthic–pelagic coupling have focussed on how energy flow and fluxes of organic matter might influence the production and structure of the benthic communities. The classical scenario depicts the input of organic material from the photic zone fertilising the seabed and the consequent benthic nutrient regeneration supplying again the water column to sustain primary production processes. More recent studies highlighted the role of merobenthos as cysts of planktonic organisms that spend part of their life quiescent in the sediments. Such benthic propagules, when environmental conditions are not anymore adverse, can supply back plankton communities. Marine canyons might accumulate cysts derived from shelf export and reverse the direction of the coupling in a bottom up perspective. In the Mediterranean Sea, Eastern and Western basins display different productivity, flux rates and benthic standing stocks, with mass fluxes and organic carbon inputs up to 80 times higher in the Western basin. The low inputs reaching the deep-sea sediments are apparently responsible for the lack of coupling in the oligotrophic regions of the Eastern Mediterranean. The poverty of the organic input below 500 m, thought insufficient to fully support growth at the metazoan level, is partially balanced by a higher organic matter quality (i.e. bioavailability) in the more oligotrophic environments. In the Eastern Mediterranean, benthic organisms (e.g., bacteria and small metazoans) are 2–3 times more efficient than in the Western Mediterranean in exploiting the available food sources. These conditions are reflected also by bioenergetic strategies. In the more oligotrophic eastern basin, meiofauna body composition (as lipid, protein and carbohydrate content) displays an apparent biochemical coupling with the composition of the organic food sources and is apparently able to cope with longer periods of scarce and highly unpredictable food inputs. Finally also climate changes might influence the pelagic-benthic coupling modifying primary productivity and sedimentation rates. Recent changes in deep water mass characteristics of the south Aegean, have determined the formation, apparently related to climate anomalies, of new colder and higher salinity water masses. These waters replaced and lifted up the older nutrient-reach deep waters, thus enhancing primary production and phytodetritus input to the deep sea. Such phenomenon has determined a clear accumulation (up to 10-times) of labile organic carbon in the deep sea. There is increasing evidence that greenhouse effects and transient climate changes might have rapid and profound effects also on pelagic–benthic coupling in deep-sea ecosystems.

Quantifying plant pigments and live diatoms in aphotic sediments of Scandinavian coastal waters confirms a major route in the pelagic–benthic coupling

The fate of pelagic diatoms in marine coastal aphotic sediments was investigated from sediment profiles in western Scandinavian waters. We used three independent methods to estimate pigment pools in the sediment: (1) fluorometry, (2) high-performance liquid chromatography and (3) pigments estimated from germinable diatom cells, using the dilution extinction method. A strong positive relationship with an intercept close to zero was observed between fucoxanthin, a marker of diatoms, and chlorophyll a. The fucoxanthin/chl a ratio was on average 1.05, which was similar to monocultures of dominating diatom taxa, indicating that sedimentary chl a was to a large extent of diatom origin. Chl a and fucoxanthin correlated significantly and positively with, and where within the same order of magnitude as, corresponding substances predicted from live diatom cell numbers obtained with the dilution extinction method. This indicates that a major part of surficial sediment chl a was bound in live cells of pelagic diatoms. There was a consistent change in viable cells with sediment depth and with timing of dominating taxa, with the non-spore-forming Skeletonema costatum dominating in the surface sediment in March and May, while the spore forming Chaetoceros spp. dominated deep in the sediment and during periods outside of the spring bloom (February and August). This indicates that chl a is bound in several different cell pools with different degradation rates, depending on diatom taxonomy. Thus, diatoms originating predominantly from the spring bloom may provide an important direct link in the pelagic–benthic coupling in this area.

Seasonal variation in production, retention and export of zooplankton faecal pellets in the marginal ice zone and central Barents Sea

Vertical distribution and sedimentation of faecal pellets (FPs) as well as the production rates of FPs by larger copepods were studied during three cruises to the Barents Sea in March and May 1998, and July 1999. Three to five 24-h stations were selected during each cruise, where at least one main station was located in Arctic water (ArW), one in the polar front region (PF) and one in Atlantic water (AW). A winter scenario was encountered in March with very low concentrations of FPs in the water column, most of the time well below 0.1 mg faecal pellet carbon (FPC) per cubic meter, and with sedimentation rates below 3 mg FPC m −2 day −1 at all depths and stations. Increased concentrations of FPs were observed in May and the maximum biomass of FPs was found in ArW (4.8 mg FPC m −3). This was reflected in high vertical flux of FPs in the ArW, just below the chlorophyll maximum (∼150 mg FPC m −2 day −1). FPC sedimentation explained ∼40% of the total particulate organic carbon (POC) export at 90 m depth at this station. Copepod FP production was moderate to high in May, reflecting favourable feeding conditions. Large spatial variation in the estimated retention potential of FPs was observed, ranging from 96% in AW to ∼40% in the PF region. The July scenario did not differ very much from that observed in May. The lowest suspended concentrations and vertical flux of FPs were again observed in AW, in spite of the high pellet-production rate. FPC explained 34% of the POC export out of the upper layer in ArW, 40% in the PF region, but only 8% in AW. The calculated retention potential of 70% of the produced copepod FPs in AW decreased to 60% and 47% in the PF region and ArW, respectively. Krill FPs comprised a significant fraction of both suspended and sedimented FPC throughout the central Barents Sea. The data show that spatial and temporal variations in the FP “retention filter” are extensive and evidently of importance for the patterns of vertical flux of organic matter and the regulation of pelagic–benthic coupling in the Barents Sea.

Pelagic—benthic coupling in the abyssal eastern North Pacific: An 8‐year time‐series study of food supply and demand

An 8‐yr time‐series study of the trophic coupling between a pelagic food supply and its utilization by the sediment community was conducted at 4,100‐m depth in the eastern North Pacific between 1989 and 1998. Supply of sinking particulate organic carbon (POC) and particulate total nitrogen (PTN) was estimated from collections made with sediment traps moored at 3,500‐ and 4,050‐m depth (600 and 50 m above bottom). Sediment community oxygen consumption (SCOC), an estimate of aerobic utilization of organic matter, was measured in situ. POC and PTN fluxes at both depths declined significantly from October 1989 through 1996 then increased in 1998. Organic carbon : total nitrogen (C : N) of the sinking particulate matter fluctuated ~10 throughout the study, except for a major peak (84.8) at 50 m above bottom in summer 1995, indicating lateral advection of organic material of terrestrial origin. POC: SCOC declined progressively over the first 7 yr, from a high of 0.99 in 1989–1990 to 0.22 in 1995–1996. In 1998, there was an increase in POC: SCOC to 0.43, suggesting that the food deficit was reduced by an increase in sinking flux. A continuing deficit in food supply cannot be sustained without ultimately affecting the structural and functional characteristics of the sediment community. The importance of undersampling by sediment traps, lateral advection of marine and terrestrial organic matter from the continental slope and shelf, and dissolved organic matter is evaluated. These other sources of nutrients for benthic organisms may be important in explaining some of the observed discrepancy between supply and demand in the abyssal Pacific. Decadal‐scale climate variations that influence marine primary production and terrestrial discharges into the ocean may be extremely important in understanding biogeochemical processes in the deep sea.

Meiofauna response to a dynamic river plume front

The benthic response to a plume front was studied in two areas of the northern Adriatic (Mediterranean Sea) differently influenced by the Po River freshwater input. Sediment samples were collected in June 1996 and February 1997 from 12 stations. The adopted sampling strategy was able to identify the front line in real time by satellite images and to locate sampling stations along an inner–outer plume gradient in order to cover the benthic area beneath the river plume, where enhanced biological production was expected, and open-sea sediments not directly influenced by freshwater inputs. Meiofaunal parameters were compared to the physical conditions and to phytodetritus inputs, organic matter accumulation and bacterial secondary production. The sediments of the Adriatic Sea were characterised by high concentrations of phytopigments (0.6 to 13.9 μg g−1 for chlorophyll a and 1.2 to 17.7 μg g−1 for phaeopigments) and biopolymeric organic carbon (0.15 to 3.02 mg g−1). The plume system extended for a large sector of the northern Adriatic. In the northern area, a large and highly dynamic plume area was coupled with a sediment organic matter concentration significantly higher than in open-sea sediments. In the southern sector, where the plume area and the front line did not change markedly during the year, plume–benthic coupling was evident only in the sediments beneath the front, and corresponded to phaeopigment accumulation. Bacterial parameters and secondary production were high and significantly higher in the frontal area than at open-sea stations. Meiofauna density (1342 to 8541 ind. 10 cm−2) did not change either by season or between areas and was significantly correlated with phaeopigments and bacterial secondary production. Meiofauna displayed different responses to plume inputs in the two sampling areas. In the northern sector, meiofauna density was coupled with organic matter distribution and displayed highest values beneath the plume. In the southern sector, the densities of copepods, turbellarians and kinorhynchs displayed highest values under the front in summer, and the same applied to total meiofauna density in winter. Juvenile decapods and copepod nauplii significantly increased their densities in sediments beneath the front. Data presented in the present study suggest that plume inputs and frontal systems, enhancing phytodetritus accumulation and benthic bacterial response, might influence density, composition and distribution of meiofaunal assemblages. As river plumes are highly variable systems affecting the trophic characteristics of the sediments underneath, their dynamics should be considered when analysing mesoscale spatial changes of meiofaunal assemblages.

Seasonal and spatial variation of suspended and sedimented nutrients (C, N, P) in the pelagic system of the Gulf of Riga

The temporal and spatial variation of suspended and sedimented C, N and P in the pelagic system in the central and southern part of the eutrophicated Gulf of Riga were investigated during late spring, summer and early autumn from 1993–1995. The temporal variability at a central station was as high or greater than the spatial variability. Thus no discernible influence of the highly eutrophicated river Daugava on the plankton dynamics and the pelagic–benthic coupling of the Gulf of Riga was observed. High sedimentation rates of carbon-rich organic material were recorded at the end of the spring bloom, while P and N were retained on average 5 and 1.2 times more effectively, respectively, than C in the pelagic system. Low sedimentation rates of P, indicating effective recycling in the mixed surface layer, were typical for most of the productive season. The low C/N ratios of the settled material during summer were probably caused by accumulation of heterotrophic dinoflagellates into the sediment traps rather than due to increased vertical export of N. It is suggested that the major supply of organic matter to the benthos is determined by the time window characterised by low heterotrophic diversity and biomass in early spring. The trophic complexity of the planktonic community in the southern and central Gulf of Riga with its continuous discharge of nutrients by large rivers such as Daugava, results in a high buffering capacity of the pelagic system towards external perturbations such as increased nutrient supply.

Biochemical composition of sedimentary organic matter and bacterial distribution in the Aegean Sea: trophic state and pelagic–benthic coupling

Biochemical composition of sedimentary organic matter (OM), vertical fluxes and bacterial distribution were studied at 15 stations (95–2270 m depth) in the Aegean Sea during spring and summer. Downward fluxes of labile OM were significantly higher in the northern than in the southern part and were higher in summer than in spring. Primary inputs of OM were not related to sedimentary OM concentrations, which had highest values in summer. Sedimentary chlorophyll-a concentrations were similar in the northern and southern parts. Carbohydrates, the main component of sedimentary OM, were about 1.2 times higher in the southern part than in the northern, without significant temporal changes. Total proteins were higher in summer and about double in the northern part. Sedimentary proteins appeared more dependent upon the downward flux of phytopigment than of proteins. Sedimentary OM was characterised by a relatively large fraction of soluble compounds and showed better quality in the northern part. The lack of a depth-related pattern in sedimentary OM and the similar concentrations in the two areas suggest that differences in sedimentary OM quality in the Aegean basin are dependent on system productivity; the bulk of sedimentary OM is largely conservative. Sedimentary bacterial density was about double in the northern part and higher in spring than in summer, but bacterial size was about three times higher in summer, resulting in a larger bacterial biomass in summer. Bacterial density was coupled with total and protein fluxes, indicating a rapid bacterial response to pelagic production. Bacterial biomass was significantly correlated with sedimentary protein and phytopigment concentrations, indicating a clear response to accumulation of labile OM in the sediments. In all cases bacteria accounted for <5% of the organic C and N pools. The efficiency of benthic bacteria in exploiting protein pools, estimated as amounts of protein available per unit bacterial biomass, indicates a constant ratio of about 70 μg proteins/μg C. This suggests a similar bacterial efficiency all over the area studied, unaffected by different trophic conditions.

Phytodetritus deposited on the Antarctic shelf and upper slope: its relevance for the benthic system

Underwater photography was used to ascertain regional, seasonal, interannual or depth dependent variations in the occurrence of phytodetritus on the Antarctic seafloor in order to explain the patchy distribution patterns of the benthos. The information was obtained from an average of 63 photographs taken at each of 76 stations in the Weddell and Lazarev Sea along a 2300 km coastline during four austral summers between 1986 and 1991. In areas where the shelf was broader than 80 km, the sediment showed a significantly higher phytodetritus cover than on the narrower shelf. This can probably be explained by the lower current velocity on the broader shelf. Significantly higher percentages of phytodetritus cover were also found on the seafloor in areas where the megabenthos cover was relatively low. These results indicate that in areas with a low current velocity, organic particles sink relatively fast onto the seafloor where they are available mainly for deposit feeders. The generally more abundant filter feeders are better adapted to a higher current velocity which transports the particles mainly horizontally over longer periods. No significant relationships were found between other physical parameters and the occurrence of phytodetritus. Therefore, the results are also discussed under the aspect of a weak pelagic–benthic coupling, effected by the long-term development of the benthic system.