Ecology of mesozooplankton across four North Atlantic basins

A comparative study of the mesozooplankton in four North Atlantic basins is presented. During a trans-Atlantic expedition with R/V G.O. Sars in May and June 2013, the Norwegian Sea, Iceland Sea, Irminger Sea and Labrador Sea was surveyed twice on a round trip from Bergen, Norway to Nuuk, Greenland. Mesozooplankton samples of biomass, species composition and vertical distribution were obtained with WP2 and MOCNESS plankton nets, in addition to in situ data obtained from a Video Plankton Recorder (VPR) and Optical Plankton Counter (OPC) mounted on a submersible towed vehicle. Size-fractionated biomass samples showed that the Norwegian Sea had the highest biomass of small mesozooplankton (180–1000 μm), while Irminger and Iceland Seas had the highest biomass of the medium (1000–2000 μm) and largest (>2000 μm) size fractions, respectively. The Icelandic Sea large fraction biomass was dominated by Amphipods, Chaetognaths, Krill and Calanus hyperboreus. The Labrador Sea had the lowest total mesozooplankton biomass. A total of 9 different species/groups were found to comprise the 5 most numerically dominant species/groups across all basins, with Oithona spp. being the most common genus in all basins. C. finmarchicus was, as expected, found to be the most numerically common species of the Calanus complex in all basins, but the stage composition varied markedly between basins with young copepodite stages dominating only in the Labrador and central Norwegian Seas. In terms of both abundance and biomass, the Iceland Sea had a higher fraction of dominating mesozooplankton distributed below 200 m. The highest average particle density per 25 m interval was registered in the Norwegian Sea during daytime between 25-50 m (OPC data). In the Labrador and Irminger Seas, total estimated particle densities in the upper 50 m were lower and the particle densities peaked at intermediate sizes (1–3 mm). In all basins there were differences in the particle densities estimated between day and night. Based on VPR data, the Irminger and Iceland Seas had the highest density of copepods registered in the upper 200 m, whereas in the Labrador Sea, the highest average copepod densities were registered at depth. Densities of gelatinous organisms were at least an order of magnitude higher in the Labrador and Irminger Seas than in the Iceland Sea.acceptedVersio

Vertical distribution and migration of mesopelagic scatterers in four north Atlantic basins

We studied vertical distribution and diel vertical migration (DVM) behaviour of mesopelagic acoustic scattering layers in relation to environmental conditions in the Norwegian Sea, the Iceland Sea, the Irminger Sea, and the Labrador Sea. Distinct mesopelagic scattering layers were found in all basins, but the daytime depth of the layers varied between basins. The results suggested that daytime vertical distribution across the four basins are strongly influenced by optical conditions. DVM occurred in all basins, and since daytime vertical distribution was influenced by optical conditions, it affected the amplitude of vertical connectivity. We used the proportion of the acoustic backscatter that migrated vertically into the epipelagic zone as a proxy for active vertical flux to the mesopelagic. The proportion of micronekton backscatter participating in the vertical migrations varied between the basins, with the highest and lowest vertical connectivity in the Norwegian Sea and the Iceland Sea respectively. We conclude that a more than 8-fold reduction in backscatter flux in the Iceland Sea was attributable to optical conditions there.publishedVersio

An acoustic method to observe the distribution and behaviour of mesopelagic organisms in front of a trawl

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Application of an unsupervised clustering algorithm on in situ broadband acoustic data to identify different mesopelagic target types

The mesopelagic zone (200–1000 m depth) contains high fish species diversity but biomass and abundances are uncertain yet essential to understand ecosystem functioning. Hull-mounted acoustic systems (usually 38 kHz) often make assumptions on average target strength (TS) of mesopelagic fish assemblages when estimating biomass/abundance. Here, an unsupervised clustering algorithm was applied on broadband acoustic data (54–78 kHz), collected by a towed instrumented platform in the central Northeast Atlantic, to identify different mesopelagic target types based on similarity of individual TS spectra. Numerical density estimates from echo-counting showed spatial differences in vertical distribution patterns of the different target types and TS spectra data suggested that >30% of the gas-bearing targets had high resonance frequencies (>60 kHz) with low scattering strength at 38 kHz. This conceptual study highlights the importance of separating targets into different target groups to obtain correct backscatter information and to account for all relevant scatterers when estimating average TS at 38 kHz, in order to achieve more accurate biomass/abundance estimates. It furthermore demonstrates the use of a towed broadband acoustic platform for fine-scale numerical density estimates as a complementary method to hull-mounted acoustic data to increase knowledge on mesopelagic ecosystem structure.publishedVersio

Mesopelagic flesh shear viscosity estimation from in situ broadband backscattering measurements by a viscous–elastic model inversion

In fisheries acoustics, target strength (TS) is a key parameter in converting acoustic measurements to biological information such as biomass. Modelling is a versatile tool to estimate TS of marine organisms. For swimbladdered fish, flesh shear viscosity is one of the required parameters to correctly calculate TS around the resonance frequency, where the target scatters most strongly. Resonance of mesopelagic swimbladdered fish can occur over a range of frequencies and can be within commonly used frequencies (e.g. 18, 38, or 70 kHz). Since there is little information on flesh shear viscosity of fish, especially for mesopelagic species, their resonance can bias the biological information extracted from acoustic measurements. Here, first, the applicability of using a spherical model to estimate resonant backscattering of a generic swimbladder is investigated. Subsequently, a viscous–elastic spherical gas backscattering model is used to estimate the flesh shear viscosity of swimbladdered mesopelagic fish (most likely Cyclothone spp., Family: Gonostomatidae) from in situ broadband backscattering measurements. Finally, the effects of flesh shear viscosity on the TS of swimbladdered mesopelagic fish at 18, 38 (a widely used channel to study mesopelagic layers), and 70 kHz are examined.publishedVersio

Mesopelagic fish gas bladder elongation, as estimated from wideband acoustic backscattering measurements

Backscattered acoustic energy from a target varies with frequency and carries information about its material properties, size, shape, and orientation. Gas-bearing organisms are strong reflectors of acoustic energy at the commonly used frequencies (∼18–450 kHz) in fishery surveys, but lack of knowledge of their acoustic properties creates large uncertainties in mesopelagic biomass estimates. Improved knowledge about the volume and elongation (i.e., longest to shortest dimension) of swimbladders of mesopelagic fishes has been identified as an important factor to reduce the overall uncertainties in acoustic survey estimates of mesopelagic biomass. In this paper, a finite element approach was used to model gas-filled objects, revealing the structure of the backscattering, also at frequencies well above the main resonance frequency. Similar scattering features were observed in measured broadband backscattering of several individual mesopelagic organisms. A method is suggested for estimating the elongation of a gas-bubble using these features. The method is applied to the in situ measured wideband (33–380 kHz) target strength (TS) of single mesopelagic gas-bearing organisms from two stations in the North Atlantic (NA) and Norwegian Sea (NS). For the selected targets, the method suggested that the average elongation of gas-bladder at the NA and NS stations are 1.49 ± 0.52 and 2.86 ± 0.50, respectively.publishedVersio

Impacts of a Changing Ecosystem on the Feeding and Feeding Conditions for Atlantic Salmon During the First Months at Sea

During the last decades, many wild Atlantic salmon populations have declined dramatically. One hypothesis for an observed reduction in salmon marine growth and survival is reduced abundance of prey. However, the effect of spatial and temporal variation in marine prey abundance on post-smolt feeding conditions is poorly understood. Here we use stomach content data from 2572 salmon postsmolts sampled during 25 years in the Northeast Atlantic Ocean to examine spatial and temporal changes in diet and stomach fullness. Sandeel larvae west of Scotland and Ireland and in the northern North Sea, herring larvae in the eastern part of the Norwegian Sea, and amphipods in the western part of the Norwegian Sea were particularly important prey species. There was a reduction of fish larvae in the postsmolt stomachs over a large geographic area when comparing the period 1995–2004 to 2008–2019. This may be result of a bottom-up driven process, as increasing extent of Arctic Water masses and zooplankton abundance were positively correlated with postsmolt stomach fullness in the Norwegian Sea. Furthermore, the interspecific competition for fish larvae between post-smolts and mackerel may have increased with a larger mackerel stock expanding the feeding migrations northwards since 2007. Norwegian Sea, fish larvae, zooplankton, mackerel, oceanography, dietpublishedVersio

Estimating target strength and physical characteristics of gas-bearing mesopelagic fish from wideband in situ echoes using a viscous-elastic scattering model

Wideband (38 and 50–260 kHz) target strength of organisms were measured in situ using a towed platform in mesopelagic (200–1000 m depth) layers. Organisms with a gas-inclusion are strong scatterers of sound and acoustically distinct from organisms lacking one. In the mesopelagic zone, some of the fish species and physonect siphonophores have a gas-inclusion. Trawl and multinet biological sampling as well as photographic evidence indicate that in the study area (eastern mid-Atlantic Ocean) the majority of the gas-bearing organisms were fish. Subsequently, using a two-layer viscous-elastic spherical gas backscattering model, physical characteristics such as gas-bladder features and body flesh properties were deduced from the measured backscattering signal of individual gas-bearing fish. Acoustic techniques are non-extractive, can be used for the monitoring and quantification of marine organisms in a time- and cost-effective manner, and suit studies of the mesopelagic zone, which is logistically challenging. Vessel-mounted acoustics, widely used for epipelagic studies, has limitations for mesopelagic studies as the deep organisms are inaccessible to high-frequency (≳100 kHz) acoustic pulses transmitted from the surface due to absorption. Therefore, a towed platform equipped with wideband acoustics has several features that can be utilized for monitoring the mesopelagic dense scattering layers containing mixed species.publishedVersio

Mass estimates of individual gas-bearing mesopelagic fish from in situ wideband acoustic measurements ground-truthed by biological net sampling

A new acoustic approach to estimate the mass of individual gas-bearing fish at their resident depth at more than 400 m was tested on Cyclothone spp.. Cyclothone are small and slender, and possibly numerically underestimated globally as individuals can pass through trawl meshes. A towed instrumented platform was used at one sampling station in the Northeast Atlantic, where Cyclothone spp. dominated numerically in net catches, to measure in situ acoustic wideband target strength (TS) spectra, i.e. acoustic scattering response of a given organism (”target”) over a frequency range (here, 38 + 50–260 kHz). Fitting a viscous–elastic scattering model to TS spectra of single targets resulted in swimbladder volume estimates from where individual mass was estimated by assuming neutral buoyancy for a given flesh density, such that fish average density equals that of surrounding water. A density contrast (between fish flesh and seawater) of 1.020 resulted in similar mass–frequency distribution of fish estimated from acoustics/model and Cyclothone spp. caught in nets. The presented proof of concept has the potential to obtain relationships between TS and mass of individual gas-bearing mesopelagic fish in general.publishedVersio

Dynamics of phytoplankton species composition, biomass and nutrients in the North Atlantic during spring and summer - A trans-Atlantic study

During a return trip in May–June 2013 from Bergen, Norway, to Nuuk, Greenland, on the Institute of Marine Research's R/V “G. O. Sars”, four ocean basins, the Norwegian, Iceland, Irminger and Labrador seas, were surveyed twice. Here we analyse the phytoplankton communities and their dynamics in the four basins, winter nutrient content, potential for new production, consumption of nutrients, phenology and algae biomass (Chlorophyll a). The primary production dynamics of the four basins are compared and related to physical forcing and top-down control and discussed in relation to vertical carbon flux. We focus on the primary production dynamics in the central, mostly Atlantic, water masses of the basin-wide gyres. The survey results are put into a larger scale context by comparison with MODIS surface chlorophyll estimates. The timing of the phytoplankton bloom phase differed among the basins. In the Labrador Sea, there was a strong bloom already occurring during the first crossing in mid-May. This was confirmed by nitrate and silicate reduction, which was much larger in the Labrador Sea compared to the other seas in May. In the Irminger Sea the bloom lagged behind, whereas in the Norwegian and Iceland seas both a pre-bloom and a late bloom were observed during May–June. There was a direct relationship between the mixed layer depth and the use of nitrate at the sampling stations, and a direct relationship between chlorophyll concentrations and the use of nitrate. The two relationships suggest that the use of nitrate can serve as a measure of the developmental state of the phytoplankton bloom, phenology, and that it is related to the shallowing of the mixed layer. Flagellates were the most abundant algae in all regions, outnumbered by diatoms only at a couple of stations west of Greenland. Elevated ciliate numbers were observed in the Iceland and Norwegian seas on both sides of the Arctic front. The algae outnumbered the microzooplankton by at least an order of magnitude.publishedVersio