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.

Summary paragraph: Plankton are essential in marine ecosystems. However, our knowledge of overall community structure is sparse due to inconsistent sampling across their very large organismal size range. Here we use diverse imaging methods to establish complete plankton inventories of organisms spanning five orders of magnitude in size. Plankton community size and trophic structure variation validate a long-held theoretical link between organism size-spectra and ecosystem trophic structures. We found that predator/grazer biomass and biovolume unexpectedly exceed that of primary producers at most (55%) locations, likely due to our better quantification of gelatinous organisms. Bottom- heavy ecosystems (the norm on land) appear to be rare in the ocean. Collectively, gelatinous organisms represent 30% of the total biovolume (8-9% of carbon) of marine plankton communities from tropical to polar ecosystems. Communities can be split into three extreme typologies: diatom/copepod-dominated in eutrophic blooms, rhizarian/chaetognath-dominated in oligotrophic tropical oceans, and gelatinous-dominated elsewhere. While plankton taxonomic composition changes with latitude, functional and trophic structures mostly depend on the amount of prey available for each trophic level. Given future projections of oligotrophication of marine ecosystems, our findings suggest that rhizarian and gelatinous organisms will increasingly dominate the apex position of planktonic ecosystems, leading to significant changes in the ocean’s carbon cycle.

The coastal oceans can be highly variable, especially near ocean fronts. The Ushant Front is the dominant oceanographic feature in the Iroise Sea (NE Atlantic) during summer, separating warm stratified offshore waters from cool vertically-mixed nearshore waters. Mesozooplankton community structure was investigated over an annual cycle to examine relationships with oceanographic conditions. DNA metabarcoding of COI and 18S genes was used in communities from six sites along two cross-shelf transects. Taxonomic assignments of 380 and 296 OTUs (COI and 18S, respectively) identified 21 classes across 13 phyla. Meroplankton relative abundances peaked in spring and summer, particularly for polychaete and decapod larvae, respectively, corresponding to the reproductive periods of these taxa. Meroplankton was most affected by season, while holoplankton varied most by shelf position. Copepods with a mixed feeding strategy were associated with the most offshore sites, especially in the presence of the front, while filter-feeding or carnivorous copepods were associated with nearshore sites. In sum, mesozooplankton communities in well-mixed coastal waters were distinct from those found in the Ushant Front (high thermal stratification and chlorophyll-a). Furthermore, the benthic compartment, through its partial life cycle in the water column, contributed to high heterogeneity in planktonic communities over short temporal and spatial scales.

The South Atlantic Ocean (SAO) is an under‐sampled ocean where the influence of environmental drivers on copepod body size is poorly understood. This study investigated the body size distribution of copepods from 13°S to 64°S to test Bergmann's rule, which predicts the occurrence of smaller organisms in warmer areas. We hypothesized that additional influence of oceanographic features strengthens this pattern. Zooplankton were sampled during the austral summer from the chlorophyll maximum depth up to the surface, at approximately 100 m depth, using a 200‐μm net. The samples were analyzed using the ZooScan imaging system and were classified using the Ecotaxa tool. We estimated copepod family abundance, biomass, and size structure, and their relationships with environmental variables were assessed through generalized linear mixed models (GLMM). Although most copepods increased in size at higher latitudes, not all families followed Bergmann's rule. Small adults of Clausocalanidae, Paracalanidae, Corycaeidae, and Oncaeidae contributed mostly in the Brazil Current (BC), while Calanidae copepodites disproportionately contributed to overall biomass, and Oithonidae adults were the most abundant in high latitudes. Copepod abundance or biomass hotspots were present in the southern limit of the warm BC, at the Subtropical Confluence Zone, and in the subantarctic waters at the Drake passage. Our results suggest that oceanographic features strengthen latitudinal body size relationships due to food availability, and the importance of different life history strategies.

Vertical variations in physical and chemical conditions drive changes in marine zooplankton community diversity and composition. In turn, zooplankton communities play a critical role in regulating the transfer of organic matter produced in the surface ocean to deeper layers. Yet, the links between zooplankton community composition and the strength of vertical fluxes of particles remain elusive, especially on a global scale. Here, we provide a comprehensive analysis of variations in zooplankton community composition and vertical particle flux in the upper kilometer of the global ocean. Zooplankton samples were collected across five depth layers and vertical particle fluxes were assessed using continuous profiles of the Underwater Vision Profiler (UVP5) at 57 stations covering seven ocean basins. Zooplankton samples were analysed using a Zooscan and individual organisms were classified into 19 groups for the quantitative analyses. Zooplankton abundance, biomass and vertical particle flux decreased from the surface to 1000m depth at all latitudes. The zooplankton abundance decrease rate was stronger at sites characterised by oxygen minima (< 5µmol O₂.kg⁻¹) where most zooplankton groups showed a marked decline in abundance, except the jellyfishes, molluscs, annelids, large protists and a few copepod families. The attenuation rate of vertical particle fluxes was weaker at such oxygen-depleted sites. Canonical redundancy analyses showed that the epipelagic zooplankton community composition depended on the community structure of surface phytoplankton and the quantity and the quality of the produced particulate organic matter. We provide a consistent baseline of plankton community structure together with estimates of vertical flux and a contribution to our understanding of global zooplankton dynamics in the upper kilometer of the global ocean. Our results further suggest that future changes in surface phytoplankton composition and mesopelagic oxygen loss might lead to profound changes in zooplankton abundance and community structure in both the euphotic and mesopelagic ocean. These changes may affect the vertical export and hereby the strength of the biological carbon pump