Increased prokaryotic diversity in the Red Sea deep scattering layer.

BACKGROUND: The diel vertical migration (DVM) of fish provides an active transport of labile dissolved organic matter (DOM) to the deep ocean, fueling the metabolism of heterotrophic bacteria and archaea. We studied the impact of DVM on the mesopelagic prokaryotic diversity of the Red Sea focusing on the mesopelagic deep scattering layer (DSL) between 450-600 m. RESULTS: Despite the general consensus of homogeneous conditions in the mesopelagic layer, we observed variability in physico-chemical variables (oxygen, inorganic nutrients, DOC) in the depth profiles. We also identified distinct seasonal indicator prokaryotes inhabiting the DSL, representing between 2% (in spring) to over 10% (in winter) of total 16S rRNA gene sequences. The dominant indicator groups were Alteromonadales in winter, Vibrionales in spring and Microtrichales in summer. Using multidimensional scaling analysis, the DSL samples showed divergence from the surrounding mesopelagic layers and were distributed according to depth (47% of variance explained). We identified the sources of diversity that contribute to the DSL by analyzing the detailed profiles of spring, where 3 depths were sampled in the mesopelagic. On average, 7% was related to the epipelagic, 34% was common among the other mesopelagic waters and 38% was attributable to the DSL, with 21% of species being unique to this layer. CONCLUSIONS: We conclude that the mesopelagic physico-chemical properties shape a rather uniform prokaryotic community, but that the 200 m deep DSL contributes uniquely and in a high proportion to the diversity of the Red Sea mesopelagic.

Diel dynamics of dissolved organic matter and heterotrophic prokaryotes reveal enhanced growth at the ocean's mesopelagic fish layer during daytime.

Contrary to epipelagic waters, where biogeochemical processes closely follow the light and dark periods, little is known about diel cycles in the ocean's mesopelagic realm. Here, we monitored the dynamics of dissolved organic matter (DOM) and planktonic heterotrophic prokaryotes every 2 h for one day at 0 and 550 m (a depth occupied by vertically migrating fishes during light hours) in oligotrophic waters of the central Red Sea. We additionally performed predator-free seawater incubations of samples collected from the same site both at midnight and at noon. Comparable in situ variability in microbial biomass and dissolved organic carbon concentration suggests a diel supply of fresh DOM in both layers. The presence of fishes in the mesopelagic zone during daytime likely promoted a sustained, longer growth of larger prokaryotic cells. The specific growth rates were consistently higher in the noon experiments from both depths (surface: 0.34 vs. 0.18 d-1, mesopelagic: 0.16 vs. 0.09 d-1). Heterotrophic prokaryotes in the mesopelagic layer were also more efficient at converting extant DOM into new biomass. These results suggest that the ocean's twilight zone receives a consistent diurnal supply of labile DOM from the diel vertical migration of fishes, enabling an unexpectedly active community of heterotrophic prokaryotes.

The Simrad EK60 echosounder dataset from the Malaspina circumnavigation.

We provide the raw acoustic data collected from the R/V Hesperides during the global Malaspina 2010 Spanish Circumnavigation Expedition (14th December 2010, Cádiz-14th July 2011, Cartagena) using a Simrad EK60 scientific echosounder operating at 38 and 120 kHz. The cruise was divided into seven legs: leg 1 (14th December 2010, Cádiz-13th January 2011, Rio de Janeiro), leg 2 (17th January 2011, Rio de Janeiro-6th February 2011, Cape Town), leg 3 (11th February 2011, Cape Town-13th March 2011, Perth), leg 4 (17th March 2011, Perth-30th March 2011, Sydney), leg 5 (16th April 2011, Auckland-8th May 2011, Honolulu), leg 6 (13th May 2011, Honolulu-10th June 2011, Cartagena de Indias) and leg 7 (19th June 2011, Cartagena de Indias-14th July 2011, Cartagena). The echosounder was calibrated at the start of the expedition and calibration parameters were updated in the data acquisition software (ER60) i.e., the logged raw data are calibrated. We also provide a data summary of the acoustic data in the form of post-processed products.

Sleep walking copepods? Calanus diapausing in hypoxic waters adjust their vertical position during winter.

While hypoxia is generally associated with negative connotations, some animals may also take advantage of reduced oxygen concentrations. However, the dynamics of such processes for zooplankton are poorly understood. We made continuous acoustic studies of Calanus helgolandicus overwintering in hypoxic waters (Oslofjorden, Norway). Their apparent minimum oxygen tolerance was 0.2-0.3 mL O2 L-1 at 8°C. The copepods adjusted their vertical distribution in concert with the upward progression of hypoxia as oxygen contents declined in the course of winter. The hypoxic overwintering habitat largely excluded potential predators and mortality appeared low in early winter. As the copepod distribution shallowed in phase with declining oxygen contents at depth, mortality increased. In contrast to recent predictions, C. helgolandicus had sufficient energy reserves to sustain long-term overwintering. Termination of the overwintering phase in spring was gradual but appeared to accelerate during the development of the spring bloom. Enhanced oceanic deoxygenation with climate change may affect seasonally migrating copepods in unpredictable ways.

Discovery of Afifi, the shallowest and southernmost brine pool reported in the Red Sea.

The previously uncharted Afifi brine pool was discovered in the eastern shelf of the southern Red Sea. It is the shallowest brine basin yet reported in the Red Sea (depth range: 353.0 to 400.5 m). It presents a highly saline (228 g/L), thalassohaline, cold (23.3 °C), anoxic brine, inhabited by the bacterial classes KB1, Bacteroidia and Clostridia and the archaeal classes Methanobacteria and Deep Sea Euryarcheota Group. Functional assignments deduced from the taxonomy indicate methanogenesis and sulfur respiration to be important metabolic processes in this environment. The Afifi brine was remarkably enriched in dissolved inorganic carbon due to microbial respiration and in dissolved nitrogen, derived from anammox processes and denitrification, according to high δ15N values (+6.88‰, AIR). The Afifi brine show a linear increase in δ18O and δD relative to seawater that differs from the others Red Sea brine pools, indicating a non-hydrothermal origin, compatible with enrichment in evaporitic environments. Afifi brine was probably formed by venting of fossil connate waters from the evaporitic sediments beneath the seafloor, with a possible contribution from the dehydration of gypsum to anhydrite. Such origin is unique among the known Red Sea brine pools.

Diel dynamics and coupling of heterotrophic prokaryotes and dissolved organic matter in epipelagic and mesopelagic waters of the central Red Sea.

The ecological status of an ecosystem can be approached by the taxa present but also by the size of individual organisms. In aquatic ecosystems, flow cytometry (FC) allows to study the individual size spectra and broad community composition through the evaluation of cytometric categories. The Red Sea represents a warm oligotrophic environment with a strong diel signal of vertically migrating mesopelagic fish, which feed at night at the surface and release dissolved organic carbon (DOC) at depth during day-time. However, knowledge about how these conditions affect the dynamics of heterotrophic prokaryotes (HP) and their coupling with DOC is lacking. Here, we analyzed a high frequency sampling over 24 h to identify the community structure and compositional changes of HP in the epipelagic and mesopelagic layers of the central Red Sea. Our results show marked vertical and diel changes in HP communities in both layers. Specifically, the relative contribution of high nucleic acid content cells was remarkably linked to changes in DOC concentration and properties. The patterns observed were likely associated to the diel vertical migration of mesopelagic fish. These findings reveal that the structure of microbial communities in warm oligotrophic environments may be more dynamic than previously thought.

Microplastic in the gastrointestinal tract of fishes along the Saudi Arabian Red Sea coast.

This study assesses the presence of microplastic litter in the contents of the gastrointestinal tract of 26 commercial and non-commercial fish species from four difference habitats sampled along the Saudi Arabian coast of the Red Sea. A total of 178 individual were examined for microplastics. In total, 26 microplastic fragments were found. Of these, 16 being films (61.5%) and 10 being fishing thread (38.5%). FTIR analysis revealed that the most abundant polymers were polypropylene and polyethylene. Parascolosps eriomma species sampled at Jazan registered the highest number of ingested microplastic. This fish species is benthic and feeds on benthic invertebrates. Although differences in the abundance of microplastic ingestion among species were not statistically significant, a significant change was observed when the level of ingestion of microplastics particles was compared among the habitats. The higher abundance of microplastics particles may be related to the habitats of fish and the presence of microplastics debris near the seabed. The results of this study represent a first evidence that microplastic pollution represents an emerging threat to Red Sea fishes, their food web and human consumers.

Light penetration structures the deep acoustic scattering layers in the global ocean.

The deep scattering layer (DSL) is a ubiquitous acoustic signature found across all oceans and arguably the dominant feature structuring the pelagic open ocean ecosystem. It is formed by mesopelagic fishes and pelagic invertebrates. The DSL animals are an important food source for marine megafauna and contribute to the biological carbon pump through the active flux of organic carbon transported in their daily vertical migrations. They occupy depths from 200 to 1000 m at daytime and migrate to a varying degree into surface waters at nighttime. Their daytime depth, which determines the migration amplitude, varies across the global ocean in concert with water mass properties, in particular the oxygen regime, but the causal underpinning of these correlations has been unclear. We present evidence that the broad variability in the oceanic DSL daytime depth observed during the Malaspina 2010 Circumnavigation Expedition is governed by variation in light penetration. We find that the DSL depth distribution conforms to a common optical depth layer across the global ocean and that a correlation between dissolved oxygen and light penetration provides a parsimonious explanation for the association of shallow DSL distributions with hypoxic waters. In enhancing understanding of this phenomenon, our results should improve the ability to predict and model the dynamics of one of the largest animal biomass components on earth, with key roles in the oceanic biological carbon pump and food web.

Far-field super-resolution imaging of resonant multiples.

We demonstrate for the first time that seismic resonant multiples, usually considered as noise, can be used for super-resolution imaging in the far-field region of sources and receivers. Tests with both synthetic data and field data show that resonant multiples can image reflector boundaries with resolutions more than twice the classical resolution limit. Resolution increases with the order of the resonant multiples. This procedure has important applications in earthquake and exploration seismology, radar, sonar, LIDAR (light detection and ranging), and ultrasound imaging, where the multiples can be used to make high-resolution images.

Social behaviour in mesopelagic jellyfish.

Gelatinous organisms apparently play a central role in deep pelagic ecosystems, but lack of observational methodologies has restricted information on their behaviour. We made acoustic records of diel migrating jellyfish Periphylla periphylla forming small, ephemeral groups at the upper fringe of an acoustic scattering layer consisting of krill. Groups of P. periphylla were also documented photographically using a remotely operated vehicle (ROV). Although the adaptive value of group formation remains speculative, we clearly demonstrate the ability of these jellyfishes to locate and team up with each other.

The submarine volcano eruption off El Hierro Island: effects on the scattering migrant biota and the evolution of the pelagic communities.

The submarine volcano eruption off El Hierro Island (Canary Islands) on 10 October 2011 promoted dramatic perturbation of the water column leading to changes in the distribution of pelagic fauna. To study the response of the scattering biota, we combined acoustic data with hydrographic profiles and concurrent sea surface turbidity indexes from satellite imagery. We also monitored changes in the plankton and nekton communities through the eruptive and post-eruptive phases. Decrease of oxygen, acidification, rising temperature and deposition of chemicals in shallow waters resulted in a reduction of epipelagic stocks and a disruption of diel vertical migration (nocturnal ascent) of mesopelagic organisms. Furthermore, decreased light levels at depth caused by extinction in the volcanic plume resulted in a significant shallowing of the deep acoustic scattering layer. Once the eruption ceased, the distribution and abundances of the pelagic biota returned to baseline levels. There was no evidence of a volcano-induced bloom in the plankton community.

Vertical distribution and diel vertical migration of krill beneath snow-covered ice and in ice-free waters.

A bottom mounted upward looking Simrad EK60 120-kHz echo sounder was used to study scattering layers (SLs) and individuals of the krill Meganyctiphanes norvegica. The mooring was situated at 150-m depth in the Oslofjord, connected with an onshore cable for power and transmission of digitized data. Records spanned 5 months from late autumn to spring. A current meter and CTD was associated with the acoustic mooring and a shore-based webcam monitored ice conditions in the fjord. The continuous measurements were supplemented with intermittent krill sampling campaigns and their physical and biological environment. The krill carried out diel vertical migration (DVM) throughout the winter, regardless of the distribution of potential prey. The fjord froze over in mid-winter and the daytime distribution of a mid-water SL of krill immediately became shallower associated with snow fall after freezing, likely related to reduction of light intensities. Still, a fraction of the population always descended all the way to the bottom, so that the krill population by day seemed to inhabit waters with light levels spanning up to six orders of magnitude. Deep-living krill ascended in synchrony with the rest of the population in the afternoon, but individuals consistently reappeared in near-bottom waters already <1 h after the ascent. Thereafter, the krill appeared to undertake asynchronous migrations, with some krill always being present in near-bottom waters even though the entire population appeared to undertake DVM.

Seasonal variations in vertical migration of glacier lanternfish, Benthosema glaciale.

The seasonal variations in glacier lanternfish (Benthosema glaciale) vertical distribution and diel vertical migration (DVM) were studied by use of a bottom-mounted upward-facing 38 kHz echo sounder deployed at 392 m depth and cabled to shore in Masfjorden (~60°52'N, ~5°24'E), Norway. Acoustic data from July 2007-October 2008 were analyzed, and scattering layers below ~220 m during daytime were attributed to glacier lanternfish based on net sampling in this, and previous studies, as well as from analysis of the acoustic data. At these depths, three different diel behavioral strategies were apparent: normal diel vertical migration (NDVM), inverse DVM (IDVM), and no DVM (NoDVM). NoDVM was present all year, while IDVM was present in autumn and winter, and NDVM was present during spring and summer. The seasonal differences in DVM behavior seem to correlate with previously established seasonal distribution of prey. We hypothesize that in regions with seasonally migrating zooplankton, such as where calanoid copepods overwinter at depth, similar plasticity in DVM behavior might occur in other populations of lanternfishes.