Abstract. The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300–600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg−1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (< 5 µmol O2 kg−1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by ongoing ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv) at 75 kHz (shipboard acoustic Doppler current profiler, ADCP) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers avoided the depth range between approximately 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg−1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time series observations of a moored ADCP (upward looking, 300 kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies followed by zooplankton in response to in response to the eddy OMZ have been identified: (i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids); (ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods); (iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes); and (iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy (i), (ii) and (iv), compression of the habitable volume in the surface may increase prey–predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg−1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Gelatinous zooplankton hold key functions in the ocean and have been shown to significantly influence the transport of organic carbon to the deep sea. We discovered a gelatinous, flux‐feeding polychaete of the genus Poeobius in very high abundances in a mesoscale eddy in the tropical Atlantic Ocean, where it co‐occurred with extremely low particle concentrations. Subsequent analysis of an extensive in situ imaging dataset revealed that Poeobius sp. occurred sporadically between 5°S–20°N and 16°W–46°W in the upper 1000 m. Abundances were significantly elevated and the depth distribution compressed in anticyclonic modewater eddies (ACMEs). In two ACMEs, high Poeobius sp. abundances were associated with strongly reduced particle concentrations and fluxes in the layers directly below the polychaete. We discuss possible reasons for the elevated abundances of Poeobius sp. in ACMEs and provide estimations showing that a single zooplankton species can completely intercept the downward particle flux by feeding with their mucous nets, thereby substantially altering the biogeochemical setting within the eddy.

Zooplankton organisms are a central part of pelagic ecosystems. They feed on all kinds of particulate matter and their egested fecal pellets contribute substantially to the passive sinking flux to depth. Some zooplankton species also conduct diel vertical migrations (DVMs) between the surface layer (where they feed at nighttime) and midwater depth (where they hide at daytime from predation). These DVMs cause the active export of organic and inorganic matter from the surface layer as zooplankton organisms excrete, defecate, respire, die, and are preyed upon at depth. In the Eastern Tropical North Atlantic (ETNA), the daytime distribution depth of many migrators (300-600 m) coincides with an expanding and intensifying oxygen minimum zone (OMZ). We here assess the day and night-time biomass distribution of mesozooplankton with an equivalent spherical diameter of 0.39-20 mm in three regions of the ETNA, calculate the DVM-mediated fluxes and compare these to particulate matter fluxes and other biogeochemical processes. Integrated mesozooplankton biomass in the ETNA region is about twice as high at a central OMZ location (cOMZ; 11 • N, 21 • W) compared to the Cape Verde Ocean Observatory (CVOO; 17.6 • N, 24.3 • W) and an oligotrophic location at 5 • N, 23 • W (5N). An Intermediate Particle Maximum (IPM) is particularly strong at cOMZ compared to the other regions. This IPM seems to be related to DVM activity. Zooplankton DVM was found to be responsible for about 31-41% of nitrogen loss from the upper 200m of the water column. Gut flux and mortality make up about 31% of particulate matter supply to the 300-600 m depth layer at cOMZ, whereas it makes up about 32% and 41% at CVOO and 5N, respectively. Resident and migrant zooplankton are responsible for about 7-27% of the total oxygen demand at 300-600 m depth. Changes in zooplankton abundance and migration behavior due to decreasing oxygen levels at midwater depth could therefore alter the elemental cycling of oxygen and carbon in the ETNA OMZ and impact the removal of nitrogen from the surface layer.

Abstract. The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300–600 m depth. Here, oxygen concentrations rarely fall below 40 μmol O2 kg−1, but are thought to decline in the course of climate change. The recent discovery of mesoscale eddies that harbour a shallow suboxic (< 5 μmol O2 kg−1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. A marked reduction in acoustic target strength (derived from shipboard ADCP, 75kHz) within the shallow OMZ at nighttime was evident. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 μmol O2 kg−1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300 kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified followed by zooplankton in response to the eddy OMZ: (i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), (ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), (iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy (i), (ii) and (iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect zooplankton avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 μmol O2 kg−1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Abstract. There is a need for cost-efficient tools to explore deep-ocean ecosystems to collect baseline biological observations on pelagic fauna (zooplankton and nekton) and establish the vertical ecological zonation in the deep sea. The Pelagic In situ Observation System (PELAGIOS) is a 3000 m rated slowly (0.5 m s−1) towed camera system with LED illumination, an integrated oceanographic sensor set (CTD-O2) and telemetry allowing for online data acquisition and video inspection (low definition). The high-definition video is stored on the camera and later annotated using software and related to concomitantly recorded environmental data. The PELAGIOS is particularly suitable for open-ocean observations of gelatinous fauna, which is notoriously under-sampled by nets and/or destroyed by fixatives. In addition to counts, diversity, and distribution data as a function of depth and environmental conditions (T, S, O2), in situ observations of behavior, orientation, and species interactions are collected. Here, we present an overview of the technical setup of the PELAGIOS as well as example observations and analyses from the eastern tropical North Atlantic. Comparisons to data from the Multiple Opening/Closing Net and Environmental Sensing System (MOCNESS) net sampling and data from the Underwater Vision Profiler (UVP) are provided and discussed.

Abstract. Marine particles of different nature are found throughout the global ocean. The term “marine particles” describes detritus aggregates and fecal pellets as well as bacterioplankton, phytoplankton, zooplankton and nekton. Here, we present a global particle size distribution dataset obtained with several Underwater Vision Profiler 5 (UVP5) camera systems. Overall, within the 64 µm to about 50 mm size range covered by the UVP5, detrital particles are the most abundant component of all marine particles; thus, measurements of the particle size distribution with the UVP5 can yield important information on detrital particle dynamics. During deployment, which is possible down to 6000 m depth, the UVP5 images a volume of about 1 L at a frequency of 6 to 20 Hz. Each image is segmented in real time, and size measurements of particles are automatically stored. All UVP5 units used to generate the dataset presented here were inter-calibrated using a UVP5 high-definition unit as reference. Our consistent particle size distribution dataset contains 8805 vertical profiles collected between 19 June 2008 and 23 November 2020. All major ocean basins, as well as the Mediterranean Sea and the Baltic Sea, were sampled. A total of 19 % of all profiles had a maximum sampling depth shallower than 200 dbar, 38 % sampled at least the upper 1000 dbar depth range and 11 % went down to at least 3000 dbar depth. First analysis of the particle size distribution dataset shows that particle abundance is found to be high at high latitudes and in coastal areas where surface productivity or continental inputs are elevated. The lowest values are found in the deep ocean and in the oceanic gyres. Our dataset should be valuable for more in-depth studies that focus on the analysis of regional, temporal and global patterns of particle size distribution and flux as well as for the development and adjustment of regional and global biogeochemical models. The marine particle size distribution dataset (Kiko et al., 2021) is available at https://doi.org/10.1594/PANGAEA.924375.

Nocturnal migration of mesopelagic fish into surface waters is well-documented. Yet, although there is increasing evidence of individual-based deviations from average population migrations and of the importance of small-scale individual behavior for predator-prey interactions and energetic cycling, little is known about what mesopelagic animals do when in upper waters. Using high-resolution data from an upwardfacing, moored, split-beam echosounder we analyzed the night-time individual vertical swimming behavior of pearlsides (Maurolicus muelleri) over one winter. The population behavior is characterized by migration to the surface after sunset, "midnight-sinking" and another migration to the surface in the morning, followed by return to the daytime habitat. Focusing on individuals unveiled diverse behavioral patterns during different phases of the migration. After ascending to upper layers at dusk, M. muelleri leaves the surface waters, not by sinking, but by actively swimming in a step-wise pattern characterized by relocations alternated by pauses. Following the descent, vertical swimming is sustained at lower levels. Around midnight, the vertical swimming direction changes from predominantly downward to upward. Several hours before dawn, the fish start ascending toward the surface in a step-wise pattern. During population ascent in the afternoon and descent in the morning, some individuals at the fringes of schools migrate without intermittent pauses. This study documents the feasibility of using submerged, stationary echosounders in unveiling the individual behavior of mesopelagic fish.

Variability of mesopelagic scattering layers is often attributed to environmental conditions or multi-species layer composition. Yet, little is known about variation in behaviour among the individuals forming scattering layers. Based on a 10 months high-resolution dataset from stationary echosounders in a Norwegian fjord, we here assess short-term and long-term behaviour of a single mesopelagic fish species, the pearlside Maurolicus muelleri. The daytime vertical extension of the monospecific pearlside scattering layers spanned four orders of magnitude ambient light in the autumn and winter and less than one order of magnitude in summer. While the main layers tracked relatively stable light levels over daytime, some individuals actively crossed light gradients of up to 1.5 orders of magnitude. This included individuals that moved between scattering layers, and apparently bold individuals that made regular upward excursions beyond the main population distribution. During the daytime, M. muelleri mitigated the risk of predation by forming tight groups in the upper scattering layer and, at light levels &gt;10−6 µmol m−2 s−1, by instantly diving into deeper waters upon encounters with predators. Our observations suggest that individual, and probably state-dependent, decisions may extend the pearlsides’ vertical distribution, with implications for predator–prey interactions.