Mesopelagic Research Articles

The diversity and ecological significance of microbial traits potentially involved in B12 biosynthesis in the global ocean.

Cobalamin (B12), an essential nutrient and growth cofactor for many living organisms on Earth, can be fully synthesized only by selected prokaryotes in nature. Therefore, microbial communities related to B12 biosynthesis could serve as an example subsystem to disentangle the underlying ecological mechanisms balancing the function and taxonomic make-up of complex functional assemblages. By anchoring microbial traits potentially involved in B12 biosynthesis, we depict the biogeographic patterns of B12 biosynthesis genes and the taxa harboring them in the global ocean, despite the limitations of detecting de novo B12 synthesizers via metagenomes alone. Both the taxonomic and functional composition of B12 biosynthesis genes were strongly shaped by depth, differentiating the epipelagic zones from the mesopelagic layers. Functional genes related to B12 biosynthesis were relatively stably distributed across different oceans, but the taxa harboring them varied considerably, showing clear functional redundancy among microbial systems. Microbial taxa carrying B12 biosynthesis genes in the surface water were influenced by environmental factors such as temperature, oxygen, and nitrate. However, the composition of functional genes was only weakly associated with these environmental factors. Null model analyses demonstrated that determinism governed the variations in B12 biosynthesis genes, whereas a higher degree of stochasticity was associated with taxonomic variations. Significant associations were observed between the chlorophyll a concentration and B12 biosynthesis, confirming its importance in primary production in the global ocean. The results of this study reveal an essential ecological mechanism governing the assembly of microbes in nature: the environment selects for function rather than taxonomy; functional redundancy underlies stochastic community assembly.

Metagenome-assembled genomes from particle-associated microbial communities in the mesopelagic zone of the Pacific Ocean.

We report 10 particle-associated metagenome-assembled genomes (MAGs) from the mesopelagic zone of Pacific Ocean seawaters. MAGs comprise members of Flavobacteria Halomonas, Blastomonas, Brevundimonas, Alteromonas, Shingomonas, Sphingopyxis, Tabrizicola, Proteobacteria, and Gammaproteobacteria. Functional annotation suggests that these bacteria are involved in central particulate organic carbon conversion, nitrogen cycling, and phosphorus cycling.

Morphological characteristics, molecular species validation and spatio-temporal distribution of myctophid eggs off the Pacific coast of northeastern Japan

Myctophids in the world's oceans have a huge biomass and act as an important link between secondary producers and upper trophic levels through their extensive diel vertical migrations from the epipelagic to mesopelagic layers. Dominance of myctophid larvae in oceanic ichthyoplankton has been reported from various parts of the major oceans; however, there are few records concerning their eggs. We established species-specific morphological characteristics for identifying myctophid eggs which were then validated by sequence analysis of mitochondrial 12S rRNA and cytochrome b genes using representative subsamples of the morphotypes, and then used the criteria to determine the species composition and spatio-temporal distribution of myctophid eggs. Samples were collected monthly in the epipelagic layer off the Pacific coast of northeastern Japan, resulting in 664 net tows that yielded a total of 10,172 myctophid eggs, defined by a smooth and delicate chorion, segmented yolk, and single oil globule. Subsamples of 160 small, 34 intermediate, and 20 large-sized eggs were sequenced, resulting in their classification as Diaphus theta, Diaphus garmani, and Myctophum asperum, respectively. Mean egg diameter was smallest in D. theta (0.75 mm), intermediate in D. garmani (1.06 mm), and largest in M. asperum (1.31 mm). Eggs of D. theta had a markedly narrower perivitelline space (0.17 mm) than D. garmani and M. asperum (0.49 and 0.71 mm, respectively). Based on morphology, the myctophid eggs were dominated by D. theta (86.0%), followed by D. garmani (12.6%) and M. asperum (1.4%). Abundances of early-stage eggs of D. theta and D. garmani were high during the daytime to dusk but low at night and dawn, indicating that spawning would not be active in the epipelagic layer at night. Spawning can be considered to occur in the mesopelagic layer during the early night, and then the eggs rise to reach the epipelagic layer in the daytime. High abundances of D. theta and D. garmani eggs were observed in the slope region from spring to summer, corresponding with the larval distribution reported for the study area.

Global biogeography of the smallest plankton across ocean depths.

Tiny ocean plankton (picoplankton) are fundamental for the functioning of the biosphere, but the ecological mechanisms shaping their biogeography were partially understood. Comprehending whether these microorganisms are structured by niche versus neutral processes is relevant in the context of global change. We investigate the ecological processes (selection, dispersal, and drift) structuring global-ocean picoplanktonic communities inhabiting the epipelagic (0 to 200 meters), mesopelagic (200 to 1000 meters), and bathypelagic (1000 to 4000 meters) zones. We found that selection decreased, while dispersal limitation increased with depth, possibly due to differences in habitat heterogeneity and dispersal barriers such as water masses and bottom topography. Picoplankton β-diversity positively correlated with environmental heterogeneity and water mass variability, but this relationship tended to be weaker for eukaryotes than for prokaryotes. Community patterns were more pronounced in the Mediterranean Sea, probably because of its cross-basin environmental heterogeneity and deep-water isolation. We conclude that different combinations of ecological mechanisms shape the biogeography of the ocean microbiome across depths.

Depth significantly affects plastisphere microbial evenness, assembly and co-occurrence pattern but not richness and composition

Microplastics have become one of the hot concerns of global marine pollution. In recent years, diversity and abiotic influence factors of plastisphere microbial communities were well documented, but our knowledge of their assembly mechanisms and co-occurrence patterns remains unclear, especially the effects of depth on them. Here, we collected microorganisms on microplastics to investigate how ocean depth affects on microbial diversity, community composition, assembly processes and co-occurrence patterns. Our results indicated that there were similar microbial richness and community compositions but microbial evenness and unique microbes were obviously different in different ocean layers. Our findings also demonstrated that deterministic processes played dominant roles in the assembly of the mesopelagic plastisphere microbial communities, while the bathypelagic microbial community assembly was mainly shaped by stochastic processes. In addition, the co-occurrence networks suggested that the relationships between microorganisms in the mesopelagic layer were more complex and stable than those in the bathypelagic layer. Simultaneously, we also found that Proteobacteria and Actinobacteriota were the most abundant keystones which played important roles in microbial co-occurrence networks at both layers. This study enhanced our understanding of microbial diversity, assembly mechanism, and co-occurrence pattern on plastisphere surfaces, and provided useful insights into microorganisms capable of degrading plastics and microbial remediation.

Divergent patterns of zooplankton connectivity in the epipelagic and mesopelagic zones of the eastern North Pacific.

Due to historical under-sampling of the deep ocean, the distributional ranges of mesopelagic zooplankton are not well documented, leading to uncertainty about the mechanisms that shape midwater zooplankton community composition. Using a combination of DNA metabarcoding (18S-V4 and mtCOI) and trait-based analysis, we characterized zooplankton diversity and community composition in the upper 1000 m of the northeast Pacific Ocean. We tested whether the North Pacific Transition Zone is a biogeographic boundary region for mesopelagic zooplankton. We also tested whether zooplankton taxa occupying different vertical habitats and exhibiting different ecological traits differed in the ranges of temperature, Chl-a, and dissolved oxygen conditions inhabited. The depth of the maximum taxonomic richness deepened with increasing latitude in the North Pacific. Community similarity in the mesopelagic zone also increased in comparison with the epipelagic zone, and no evidence was found for a biogeographic boundary between previously delineated mesopelagic biogeochemical provinces. Epipelagic zooplankton exhibited broader temperature and Chl-a ranges than mesopelagic taxa. Within the epipelagic, taxa with broader temperature and Chl-a ranges also had broader distributional ranges. However, mesopelagic taxa were distributed across wider dissolved oxygen ranges, and within the mesopelagic, only oxygen ranges covaried with distributional ranges. Environmental and distributional ranges also varied among traits, both for epipelagic taxa and mesopelagic taxa. The strongest differences in both environmental and distributional ranges were observed for taxa with or without diel vertical migration behavior. Our results suggest that species traits can influence the differential effects of physical dispersal and environmental selection in shaping biogeographic distributions.

Comparing the isotopic and molecular composition of dissolved organic carbon between the oligotrophic South China Sea and the adjacent North Pacific Ocean: Signals of biodegradation, conservative mixing, and terrestrial input

The cycling of oceanic dissolved organic carbon (DOC) is a crucial component of the global carbon cycle, yet the identification of sources and the mechanisms of its molecular transformation remain poorly understood. This study compared the isotopic and molecular composition of DOC between the oligotrophic South China Sea (SCS) and the adjacent North Pacific Ocean (NPO), and traced both its allochthonous and autochthonous sources as well as its dynamic cycling processes. DOC was collected through solid-phase extraction (SPE) from water samples of both the SCS and NPO. Carbon content, isotopic ratios, and high-resolution Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR-MS) measurements revealed that SPE-DOC contained both labile and refractory fractions. According to our mass balance model, the labile fraction of SPE-DOC exhibited a decline from 11.5 to 12.6 μM in surface waters to a negligible concentration below 1000 m. Conversely, the refractory fraction of SPE-RDOC maintained a relatively consistent value, ranging from 12.7 to 19.0 μM across the entire water column. The vertical distribution patterns of the molecular composition and carbon isotopic ratios jointly indicated that the DOC distributions are shaped by distinct biological and physical processes within different biogeochemical realms of the water column. The production and transformation of the relatively labile DOC fractions were the dominant processes in the epipelagic and mesopelagic zones (upper 1000 m). The extent of diapycnal mixing between the SCS and NPO explained the different vertical distributions of refractory DOC molecules in the bathypelagic oceans. The molecular indices of polyphenol compounds, aromaticity, double bond saturation state, terrestrial mass peaks, and δ13C ratios of SPE-DOC indicated contributions from terrestrial sources, likely riverine input, in the SCS. This study sheds light on the molecular evidence of DOC sources, as well as their transformation and conservative mixing processes along the overturning circulation in marginal seas.

Community assembly and network stability of picoeukaryotic plankton communities in the northeast Indian Ocean

Picoeukaryotic plankton played as key components of marine microbial communities, and participate in the marine food webs, especially the carbon cycling. Despite their importance, little is known about the mechanisms of assembly and network stability of picoeukaryotic plankton in the ocean along vertical gradients. Here, we obtained new dataset on the dynamics of picoeukaryotic plankton communities in the northeast Indian Ocean with a vertical gradient of 0–2000 m on two cruises. The diversity of picoeukaryotic plankton gradually decreased along the vertical gradient, with photosynthetic plankton mainly distributed in the epipelagic zone and heterotrophs dominating in the mesopelagic zone. Temperature and dissolved oxygen were the main environmental factors affecting the structure of the picoeukaryotic plankton community along the vertical gradient. The community assembly mechanisms also differed among different water layers, with the epipelagic zone dominated by stochastic processes and the mesopelagic zone controlled by deterministic processes, and heterogeneous selection processes were the main drivers shaping the picoeukaryotic plankton community structure. In addition, picoeukaryotic plankton community network stability showed nonlinear changes in vertical gradients, the mesopelagic zone is dominated by deterministic processes and therefore most resistant to environmental disturbances. Our results provide new insights into the plankton ecology of the northeast Indian Ocean and the structure of microbial communities and networks along vertical gradients.

BioGeoChemical‐Argo Floats Reveal Stark Latitudinal Gradient in the Southern Ocean Deep Carbon Flux Driven by Phytoplankton Community Composition

AbstractThe gravitational sinking of particles in the mesopelagic layer (∼200–1,000 m) transfers to the deep ocean a part of atmospheric carbon fixed by phytoplankton. This process, called the gravitational pump, exerts an important control on atmospheric CO2 levels but remains poorly characterized given the limited spatio‐temporal coverage of ship‐based flux measurements. Here, we examined the gravitational pump with BioGeoChemical‐Argo floats in the Southern Ocean, a critically under‐sampled area. Using time‐series of bio‐optical measurements, we characterized the concentration of particles in the productive zone, their export and transfer efficiency in the underlying mesopelagic zone, and the magnitude of sinking flux at 1,000 m. We separated float observations into six environments delineated by latitudinal fronts, sea‐ice coverage, and natural iron fertilization. Results show a significant increase in the sinking‐particle flux at 1,000 m with increasing latitude, despite comparable particle concentrations in the productive layer. The variability in deep flux was driven by changes in the transfer efficiency of the flux, related to the composition of the phytoplanktonic community and the size of particles, with intense flux associated with the predominance of micro‐phytoplankton and large particles at the surface. We quantified the relationships between the nature of surface particles and the flux at 1,000 m and used these results to upscale our flux survey across the whole Southern Ocean using surface observations by floats and satellites. We then estimated the basin‐wide Spring‐Summer flux of sinking particles at 1,000 m over the Southern Ocean (0.054 ± 0.021 Pg C).

Could Deep-Sea Fisheries Contribute to the Food Security of Our Planet? Pros and Cons

Hundreds of millions of people on the planet are affected by malnourishment. This contributes to the vulnerability of large swaths of the population worldwide. Children under five years old and adolescent girls are especially disproportionately vulnerable to diseases and even death in less developed countries. Today, by providing a substantial share of global protein intake, as well as fatty acids and micronutrients, fisheries contribute to global food security. As fish stocks in the upper sea levels are increasingly over-exploited, there is a surge in discussion on the potential contributions of deep-sea fisheries for global food and nutrition security. Some mesopelagic fishes show potential in providing important nutrients. Another way of supplying food security might be in using mesopelagic fish as fish feed. However, fishing in the mesopelagic zone could lead to severe ecological repercussions, especially because the impact on the biological carbon pump is uncertain. This paper highlights and juxtaposes different perspectives regarding exploitation pathways of the fish riches of deep seas, and reviews best practice model projects that deal with uncertainties related to fishery management in the mesopelagic zone. The review concludes that due to the essential role of the mesopelagic zone in the global biological carbon pump and complex interaction patterns between pelagic and mesopelagic species, exploitation of the fish riches of the mesopelagic zone must be based on comprehensive data and rigorous analyses. In the face of the current uncertainty on the respective mechanisms, the authors endorse an international moratorium on deep-sea fisheries and/or the rather small-scale exploitation of mesopelagic biomass.

Reconstructing the ocean's mesopelagic zone carbon budget: sensitivity and estimation of parameters associated with prokaryotic remineralization

Abstract. Through the constant rain of sinking marine particles in the ocean, carbon (C) trapped within is exported into the water column and sequestered when reaching depths below the mesopelagic zone. Atmospheric CO2 levels are thereby strongly related to the magnitude of carbon export fluxes in the mesopelagic zone. Sinking particles represent the main source of carbon and energy for mesopelagic organisms, attenuating the C export flux along the water column. Attempts to quantify the amount of C exported versus C consumed by heterotrophic organisms have increased in recent decades. Yet, most of the conducted estimations have led to estimated C demands several times higher than the measured C export fluxes. The choice of parameters such as growth efficiencies or various conversion factors is known to greatly impact the resulting C budget. In parallel, field or experimental data are sorely lacking to obtain accurate values of these crucial overlooked parameters. In this study, we identify the most influential of these parameters and perform inversion of a mechanistic model. Further, we determine the optimal parameter values as the ones that best explain the observed prokaryotic respiration, prokaryotic production and zooplankton respiration. The consistency of the resulting C budget suggests that such budgets can be adequately balanced when using appropriate parameters.

Suspended particles are hotspots of microbial remineralization in the ocean's twilight zone

The sinking of photosynthetically produced organic carbon from the ocean surface to its interior is a significant term in the global carbon cycle. Most sinking organic carbon is, however, remineralized in the mesopelagic zone (∼100 m–1000 m), thereby exerting control over ocean-atmosphere carbon dioxide (CO2) partitioning and hence global climate. Sinking particles are considered hotspots of microbial respiration in the dark ocean. However, our observations in the contrasting Scotia Sea and the Benguela Current show that >90% of microbial remineralisation is associated with suspended, rather than sinking, organic matter, resulting in rapid turnover of the suspended carbon pool and demonstrating its central role in mesopelagic carbon cycling. A non-steady-state model indicates that temporally variable particle fluxes, particle injection pumps and local chemoautotrophy are necessary to help balance the observed mesopelagic respiration. Temperature and oxygen exert control over microbial respiration, particularly for the suspended fraction, further demonstrating the susceptibility of microbial remineralisation to the ongoing decline in oxygen at mid-ocean depths. These observations suggest a partial decoupling of carbon cycling between non-sinking and fast-sinking organic matter, challenging our understanding of how oceanic biological processes regulate climate.

Diel vertical migration of copepods in the tropical and subtropical Atlantic Ocean

The vertical distribution of the zooplankton community was analyzed from the surface to 800 m depth in the tropical-subtropical Atlantic Ocean in a transect of 12 stations during day and nighttime from Brazilian waters (10°S) to the Canary Islands (27°N). A depth stratified opening-closing 1 m2 mouth MOCNESS was used and 8 layers analyzed. Besides this net, the epipelagic layer (0–200 m) was sampled for microzooplankton (53 µm Calvet net) and mesozooplankton (200 µm WP2 net) communities. An increasing gradient of mesozooplankton abundance was observed from the oligotrophic waters off Brazil to the oceanic upwelling off Northwest Africa. Copepods were always the most abundant group (86 %) and 241 species identified, although only 60 were >1 %. Smaller species predominated and Oncaea was the dominant group (18 %) followed by Clausocalanus (14 %) with C. furcatus dominating. Paracalanus and Oithona were also quite abundant (12 and 9 % respectively). Small copepods such as C. furcatus and Euchaeta marina predominated in the oligotrophic waters while Calanus helgolandicus, Calanoides carinatus, Temora stylifera, Paracalanus parvus, and Pleuromamma robusta dominated in the upwelling area. In the epipelagic layer large copepods (e.g., Rhincalanus, Pleuromamma, Euchirella, and Metridia) were numerous during nighttime and in the mesopelagic zone during day were quantitatively less abundant. The vertical distribution of the dominant copepods (>1 %) showed multiple diel patterns but at least five were distinguished and their dominant taxa defined: (1) Species living in the twilight zone with slight up and down motion. (2) Species dominating the epipelagic layers with weak vertical movements. (3) Strong migrants living in the mesopelagic zone during day and moving upward towards the surface or (4) migrating to the subsurface layer at nighttime, and (5) those organisms characterizing the oceanic upwelling where the calanoids predominated (e.g., C. helgolandicus and C. carinatus). Our findings highlight the complexity in the diel vertical migration patterns of copepods in the tropical-subtropical Atlantic Ocean and the relevant role of the deep-sea copepods and their diel vertical migration. These results could be used as baseline information for low latitudes of the Atlantic Ocean in future studies of the biological carbon pump.

Vertical connectivity of microbiome and metabolome reveals depth-dependent variations across a deep cold-seep water column

Deciphering the vertical connectivity of oceanic microbiome and metabolome is crucial for understanding the carbon sequestration and achieving the carbon neutrality. However, we lack a systematic view of the interplay among particle transport, microbial community, and metabolic trait across depths. Through integrating the biogeochemical, microbial, and metabolic characteristics of a deep cold-seep water column (∼1989 m), we find the altered connectivity of microbial community and dissolved organic matter (DOM) across depths. Both the microbial communities (bacteria and protists) and DOM show a clear compositional connectivity from surface to the depth of 1000 m, highlighting the controls of sinking particle over microbial connectivity from the epipelagic to mesopelagic zone. However, due to the biological migration and ocean mixing, the fecal-associated bacteria and protistan consumers unexpectedly emerge and the degradation index of DOM substantially alters around 1000–1200 m. Collectively, we unveil the significance of multi-faceted particle dispersion, which supports the connectivity and variability of deep ocean microbial communities.

Characterization and genomic analysis of Stutzerimonas stutzeri phage vB_PstS_ZQG1, representing a novel viral genus

Stutzerimonas stutzeri is an opportunistic pathogenic bacterium belonging to the Gammaproteobacteria, exhibiting wide distribution in the environment and playing significant ecological roles such as nitrogen fixation or pollutant degradation. Despite its ecological importance, only two S. stutzeri phages have been isolated to date. Here, a novel S. stutzeri phage, vB_PstS_ZQG1, was isolated from the surface seawater of Qingdao, China. Transmission electron microscopy analysis indicates that vB_PstS_ZQG1 has a morphology characterized by a long non-contractile tail. The genomic sequence of vB_PstS_ZQG1 contains a linear, double-strand 61,790-bp with the G+C content of 53.24% and encodes 90 putative open reading frames. Two auxiliary metabolic genes encoding TolA protein and nucleotide pyrophosphohydrolase were identified, which are likely involved in host adaptation and phage reproduction. Phylogenetic and comparative genomic analyses demonstrated that vB_PstS_ZQG1 exhibits low similarity with previously isolated phages or uncultured viruses (average nucleotide identity values range from 21.7 to 29.4), suggesting that it represents a novel viral genus by itself, here named as Fuevirus. Biogeographic analysis showed that vB_PstS_ZQG1 was only detected in epipelagic and mesopelagic zone with low abundance. In summary, our findings of the phage vB_PstS_ZQG1 will provide helpful insights for further research on the interactions between S. stutzeri phages and their hosts, and contribute to discovering unknown viral sequences in the metagenomic database.

Mesopelagic particulate nitrogen dynamics in the subarctic and subtropical regions of the western North Pacific

Recently, new spatiotemporal-scale particle observations by autonomous profiling floats equipped with bio-optical sensors have revealed that, in addition to gravitational particle sinking, the downward transport of surface particles by physical mixing events, which has been overlooked, contributes to particulate organic carbon export. However, the subsequent behavior of these exported particles in the mesopelagic zone (e.g., particle fragmentation and degradation) remains unclear, although it may influence the efficiency of carbon transport to further depths. This study successfully depicted the new annual mean mesopelagic particulate nitrogen (PN) dynamics with multi-layer, steady-state suspended PN pools by reanalyzing seasonal data on the stable nitrogen isotopic compositions of both suspended and sinking particles, each with different profiles, from subarctic station K2 and subtropical station S1 in the North Pacific, which are both CO2 sinks but in different oceanic settings. As analytical conditions, we assumed that the net loss of sinking PN was entirely due to abiotic fragmentation of particle aggregates to non-sinking particles and that the apparent 15N enrichment associated with heterotrophic degradation in the suspended PN pools was vertically constant. The 15N mass balance for the PN supply to the uppermost mesopelagic pool, derived from such constraints, allowed estimating the PN export by the mixed-layer pump, which was 1.6 times greater at K2 than at S1. However, its contribution to the total export (including gravitational PN sinking) from the surface layer was approximately 20% at both stations. Moreover, the ratio of PN supplied to the uppermost pool by the mixed-layer pump and by the fragmentation of particle aggregates was also similar at both stations, approximately 1:1. Using these ratios, together with separate observations of the mixed-layer pump-driven flux, it may be possible to estimate the efficiency of the particulate organic carbon transport due to the biological gravitational pump responsible for carbon sequestration in the deep sea.

Living in darkness: functional diversity of mesopelagic fishes in the western tropical Atlantic

Mesopelagic zones (200–1,000-m depth) are characterized by relatively low light levels, cold waters, and often limited oxygen, forming a stable yet challenging habitat for their inhabitants. To overcome these constraints, mesopelagic fishes have developed several adaptations that enable them to participate in crucial ecosystem processes such as nutrient cycling, carbon transport, and provisioning of harvestable fish stocks. However, our understanding of the functional diversity of mesopelagic fishes remains limited, while it is of particular importance considering the increase in human threats and possible environmental changes in the deep ecosystems. In this context, functional diversity emerges as a powerful tool and can help assess community assembly rules, including species complementary and redundancy. Here, we take advantage of scientific surveys that collected 200 species taxa identified from over 7,000 specimens to determine the functional diversity of mesopelagic fish assemblages across depths and the day–night period. We created a data set of 17 qualitative functional traits related to feeding, survival, and locomotion functions. Based on this information, functional spaces and diversity indices were calculated, and functional groups were established. Furthermore, the influence on the functional diversity of environmental variables and the day–night period was assessed by generalized additive models (GAMs). The hypothesis of functional complementary was tested. Overall, mesopelagic fishes displayed a high functional diversity and could be grouped into 10 major functional groups. Moreover, 107 species exhibited a unique composition of functional trait values, revealing a vast complementarity of functions within the deep-sea ecosystem. We also showed that functional diversity in mesopelagic ecosystems is neither static nor homogeneous, exhibiting higher values in the deepest layers and varying between day and night. We finally discuss processes that may structure mesopelagic fish assemblages and the implications of our findings for the conservation mesopelagic fishes.

Otolith characterization and integrative species identification of adult mesopelagic fishes from the western North Atlantic Ocean

Fish diversity and ecology in the ocean’s mesopelagic zone are understudied compared to other marine regions despite growing interest in harvesting these potential resources. Otoliths can provide a wealth of taxonomic and life history information about fish, which can help fill these knowledge gaps; however, there has been relatively little research to date on the otoliths of mesopelagic species. Here, a species-specific image library was assembled of sagittal otoliths from 70 mesopelagic fishes belonging to 29 families collected in the western North Atlantic Ocean. Images of adult sagittal otoliths from 12 species were documented and photographed for the first time. The fish were identified to species with a combination of morphological characters and DNA barcoding. Regressions between otolith size and fish length are presented for the six species with the largest sample sizes in this study. This otolith image library, coupled with otolith-length and width to fish-length relationships, can be used for prey identification and back-calculation of fish size, making it a valuable tool for studies relating to food webs in the important yet poorly understood mesopelagic zone. In addition, the 44 fish barcodes generated in this study highlight the benefit of using an integrative taxonomic approach to studies of this nature, as well as add to existing public databases that enable cryptic species and metabarcoding analyses of mesopelagic species.

Ocean-wide comparisons of mesopelagic planktonic community structures

For decades, marine plankton have been investigated for their capacity to modulate biogeochemical cycles and provide fishery resources. Between the sunlit (epipelagic) layer and the deep dark waters, lies a vast and heterogeneous part of the ocean: the mesopelagic zone. How plankton composition is shaped by environment has been well-explored in the epipelagic but much less in the mesopelagic ocean. Here, we conducted comparative analyses of trans-kingdom community assemblages thriving in the mesopelagic oxygen minimum zone (OMZ), mesopelagic oxic, and their epipelagic counterparts. We identified nine distinct types of intermediate water masses that correlate with variation in mesopelagic community composition. Furthermore, oxygen, NO3− and particle flux together appeared as the main drivers governing these communities. Novel taxonomic signatures emerged from OMZ while a global co-occurrence network analysis showed that about 70% of the abundance of mesopelagic plankton groups is organized into three community modules. One module gathers prokaryotes, pico-eukaryotes and Nucleo-Cytoplasmic Large DNA Viruses (NCLDV) from oxic regions, and the two other modules are enriched in OMZ prokaryotes and OMZ pico-eukaryotes, respectively. We hypothesize that OMZ conditions led to a diversification of ecological niches, and thus communities, due to selective pressure from limited resources. Our study further clarifies the interplay between environmental factors in the mesopelagic oxic and OMZ, and the compositional features of communities.

Three major mesoplanktonic communities resolved by in situ imaging in the upper 500 m of the global ocean

AbstractAimThe distribution of mesoplankton communities has been poorly studied at global scale, especially from in situ instruments. This study aims to (1) describe the global distribution of mesoplankton communities in relation to their environment and (2) assess the ability of various environmental‐based ocean regionalizations to explain the distribution of these communities.LocationGlobal ocean, 0–500 m depth.Time Period2008–2019.Major Taxa StudiedTwenty‐eight groups of large mesoplanktonic and macroplanktonic organisms, covering Metazoa, Rhizaria and Cyanobacteria.MethodsFrom a global data set of 2500 vertical profiles making use of the Underwater Vision Profiler 5 (UVP5), an in situ imaging instrument, we studied the global distribution of large (>600 μm) mesoplanktonic organisms. Among the 6.8 million imaged objects, 330,000 were large zooplanktonic organisms and phytoplankton colonies, the rest consisting of marine snow particles. Multivariate ordination (PCA) and clustering were used to describe patterns in community composition, while comparison with existing regionalizations was performed with regression methods (RDA).ResultsWithin the observed size range, epipelagic plankton communities were Trichodesmium‐enriched in the intertropical Atlantic, Copepoda‐enriched at high latitudes and in upwelling areas, and Rhizaria‐enriched in oligotrophic areas. In the mesopelagic layer, Copepoda‐enriched communities were also found at high latitudes and in the Atlantic Ocean, while Rhizaria‐enriched communities prevailed in the Peruvian upwelling system and a few mixed communities were found elsewhere. The comparison between the distribution of these communities and a set of existing regionalizations of the ocean suggested that the structure of plankton communities described above is mostly driven by basin‐level environmental conditions.Main ConclusionsIn both layers, three types of plankton communities emerged and seemed to be mostly driven by regional environmental conditions. This work sheds light on the role not only of metazoans, but also of unexpected large protists and cyanobacteria in structuring large mesoplankton communities.