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Characterization of species diversity of zooplankton is key to understanding, assessing, and predicting the function and future of pelagic ecosystems throughout the global ocean. The marine zooplankton assemblage, including only metazoans, is highly diverse and taxonomically complex, with an estimated ~28,000 species of 41 major taxonomic groups. This review provides a comprehensive summary of DNA sequences for the barcode region of mitochondrial cytochrome oxidase I (COI) for identified specimens. The foundation of this summary is the MetaZooGene Barcode Atlas and Database (MZGdb), a new open-access data and metadata portal that is linked to NCBI GenBank and BOLD data repositories. The MZGdb provides enhanced quality control and tools for assembling COI reference sequence databases that are specific to selected taxonomic groups and/or ocean regions, with associated metadata (e.g., collection georeferencing, verification of species identification, molecular protocols), and tools for statistical analysis, mapping, and visualization. To date, over 150,000 COI sequences for ~ 5600 described species of marine metazoan plankton (including holo- and meroplankton) are available via the MZGdb portal. This review uses the MZGdb as a resource for summaries of COI barcode data and metadata for important taxonomic groups of marine zooplankton and selected regions, including the North Atlantic, Arctic, North Pacific, and Southern Oceans. The MZGdb is designed to provide a foundation for analysis of species diversity of marine zooplankton based on DNA barcoding and metabarcoding for assessment of marine ecosystems and rapid detection of the impacts of climate change.

Mesopelagic fishes are an important component of the world’s oceans in terms of their abundance, biomass, and ecosystem function. These fishes are important contributors to the biological carbon pump via their feeding and behaviors, whereby they facilitate the transfer of carbon from shallow waters to the deep sea. Several species undertake diel vertical migration, feeding in shallower waters at night and moving to deeper waters during the day. This process actively expedites the downward flux of carbon. However, carbon budgets and climate models require accurate information regarding the depth distributions and migration patterns of these fishes, and environmental DNA (eDNA) analyses can provide this information. Here, we utilize eDNA approaches, generating taxonomically-informative COI and 12S reference barcodes for 80 species of mesopelagic fishes, which can be used for species-level identification of eDNA sequences. Using these, along with a publicly available barcodes database, we compare results from eDNA analysis with traditional net sampling, and explore the ability of eDNA techniques to detect diel vertical migration in fishes from samples collected in Northwest Atlantic Slope Water. We found that eDNA and net samples often resulted in different species identifications, demonstrating that eDNA can detect species that would otherwise be missed with traditional methods. In our eDNA samples, we also detected more species (12) in our shallowest depth category (0 - 100 m) from night samples than from day samples (3). This is consistent with increased diversity in shallow waters at night due to diel vertical migration. Based on the variability observed in sample duplicates, we suggest that future mesopelagic eDNA studies incorporate larger sample volumes and scaled-up sampling efforts. We also note the potential applications of eDNA analysis in addressing ecological questions related to predator-prey relationships identification of foraging hotspots, and carbon flow through the ocean’s midwaters.

Scattering structures, including deep (>200 m) scattering layers are common in most oceans, but have not previously been properly documented in the Arctic Ocean. In this work, we combine acoustic data for distribution and abundance estimation of zooplankton and fish with biological sampling from the region west and north of Svalbard, to examine high latitude meso- and epipelagic scattering layers and their biological constituents. Our results show that typically, there was strong patchy scattering in the upper part of the epipelagic zone (440 m) in the Arctic habitat north of Svalbard compared to those south in the Fram Strait west of Svalbard (WMD ~400 m). The surface integrated backscatter (Nautical Area-Scattering Coefficient, NASC, sA [m2∙nmi-2]) was considerably lower in the waters around Svalbard compared to the more southern regions (62-69ºN). Also, the mesopelagic nautical area scattering coefficient was a factor of approximately 6-10 lower around Svalbard compared to the areas in the south-eastern part of the Norwegian Sea ~62º30’N. The documented patterns and structures, particularly the DSL and its constituents, will be key reference points for understanding and quantifying future changes in the pelagic ecosystem at the entrance to the Arctic Ocean.

While sound scattering layers have been described previously from ice-covered waters in the Arctic, the existence of a viable mesopelagic community that also includes mesopelagic fishes in the Arctic has been questioned. In addition, it has been hypothesized that vertical migration would hardly exist in these areas. We wanted to check if deep scattering layers (DSLs) was found to the west and north of Svalbard (79°30’N - 82°10’N) during autumn 2015, and if present; whether organisms in such DSLs undertook vertical migrations. Our null hypothesis was that there would be no evidence of diel vertical migration. Multi-frequency acoustic observations by hull mounted echosounders (18, 38 and 120 kHz) revealed a DSL at depths ~210-510 m in areas with bottom depths exceeding ~600 m. Investigating eight geographical locations that differed with respect to time periods, light cycle and sea ice conditions, we show that the deeper layer of DSL displayed a clear ascending movement during night time and a descending movement during daytime. The high-light weighted mean depth (343-514 m) with respect to backscattered energy was statistically deeper than the low-light weighted mean depth (179-437 m) for the locations studied. This behavior of the DSL was found to be consistent both when the sun was continuously above the horizon and after it started to set on 1 September, and both in open water and sea ice covered waters. The weighted mean depth showed an increasing trend, while the nautical area backscattering strength from the DSL showed a decreasing trend from south to north among the studied locations. Hydrographic observations revealed that the diel migration was found in the lower part of the north-flowing Atlantic Water, and was disconnected from the surface water masses above the Atlantic Water during day and night. The organisms conducting vertical migrations were studied by vertical and oblique hauls with zooplankton nets and pelagic trawls. These data suggest that these organisms were mainly various mesopelagic fishes, some few larger fishes, large zooplankton like krill and amphipods, and various gelatinous forms.

In autumn 2015, several sources reported observations of large amounts of gelatinous material in a large north Norwegian fjord system, either caught when trawling for other organisms or fouling fishing gear. The responsible organism was identified as a physonect siphonophore, Nanomia cara, while a ctenophore, Beroe cucumis, and a hydromedusa, Modeeria rotunda, were also registered in high abundances on a couple of occasions. To document the phenomena, we have compiled a variety of data from concurrent fisheries surveys and local fishermen, including physical samples, trawl catch, and acoustic data, photo and video evidence, and environmental data. Because of the gas-filled pneumatophore, characteristic for these types of siphonophores, acoustics provided detailed and unique insight to the horizontal and vertical distribution and potential abundances (~0.2–20 colonies·m−3) of N. cara with the highest concentrations observed in the near bottom region at ~320 m depth in the study area. This suggests that these animals were retained and accumulated in the deep basins of the fjord system possibly blooming here because of favorable environmental conditions and potentially higher prey availability compared to the shallower shelf areas to the north. Few cues as to the origin and onset of the bloom were found, but it may have originated from locally resident siphonophores. The characteristics of the deep-water masses in the fjord basins were different compared to the deep water outside the fjord system, suggesting no recent deep-water import to the fjords. However, water-masses containing siphonophores (not necessarily very abundant), may have been additionally introduced to the fjords at intermediate depths, with the animals subsequently trapped in the deeper fjord basins. The simultaneous observations of abundant siphonophores, hydromedusae, and ctenophores in the Lyngen-Kvænangen fjord system are intriguing, but difficult to provide a unified explanation for, as the organisms differ in their biology and ecology. Nanomia and Beroe spp. are holopelagic, while M. rotunda has a benthic hydroid stage. The species also have different trophic ecologies and dietary preferences. Only by combining information from acoustics, trawling, genetics, and local fishermen, were the identity, abundance, and the vertical and horizontal distribution of the physonect siphonophore, N. cara, established.

Mesopelagic zones of the NW Atlantic Slope Water support pelagic assemblages with high biomass and high diversity. Species of salps (Phylum Urochordata, Class Thaliacea) are bloom-forming, filter-feeding, gelatinous zooplankton, with significant impacts on pelagic food webs and vertical transport of organic material. Questions remain of their diet diversity, prey selectivity, and timing of feeding with respect to vertical distribution and migration. This study analyzed three salp species, Soestia (Iasis) zonaria, Salpa aspera , and Salpa fusiformis , collected from the NW Atlantic Slope Water during July and August 2018 and 2019. DNA metabarcoding using V4 and V9 regions of 18S rRNA found dinoflagellates to be predominant prey for all three salp species in both years. Analysis of five prey groups detected by metabarcoding revealed differences in proportions of prey. Compound-specific stable isotope analysis of essential amino acids, which integrates over several months, found diatoms to be the dominant end-member source for all three species. Our findings on salp diet diversity broaden our understanding of trophic pathways in the mesopelagic food web, including sources of productivity and possible impacts of climate change.

  Euphausiids are an ecologically significant and abundant group of marine zooplankton that form key links between primary producers and consumers in pelagic food webs throughout the world ocean. The euphausiid species, Thysanoessa inermis and T. raschii , have boreal-Arctic distributions, occurring in the North Atlantic, North Pacific, and Arctic Oceans. The species differ in depth ranges and habitat preferences: T. raschii is found in coastal waters on continental shelf habitats, while T. inermis is abundant in slope and deep water regions. Population genetic analysis based on DNA sequence variation of the mitochondrial cytochrome oxidase I (COI) barcode region was carried out for identified specimens of T. inermis and T. raschii collected in the Arctic (Beaufort/Chukchi and Norwegian Seas, Svalbard Area) and North Atlantic (Gulf of St. Lawrence, Labrador Sea, Iqaluit, Hudson Bay). Populations of T. inermis in the N. Atlantic showed high connectivity, but were genetically isolated from the Beaufort/Chukchi Sea population. Population genetic diversity of T. inermis showed high haplotype and nucleotide diversity and no departures from neutral expectations. In contrast, T. raschii showed lower haplotype and nucleotide diversity, with highly significant departures from neutral expectations. A possible hypothesis is that T. raschii experienced a significant historical demographic event (e.g., population bottleneck), while T. inermis maintained a stable population over recent evolutionary history. The results provide new insights into population dynamics and implications for responses to climate change of these key euphausiid species for the Arctic Ocean.

The Western Antarctic Peninsula (WAP) is a biologically rich area supporting large standing stocks of krill and top predators (including whales, seals and seabirds). Physical forcing greatly affects productivity, recruitment, survival and distribution of krill in this area. In turn, such interactions are likely to affect the distribution of baleen whales. The Southern Ocean GLOBEC research program aims to explore the relationships and interactions between the environment, krill and predators around Marguerite Bay (WAP) in autumn 2001 and 2002. Bathymetric and environmental variables including acoustic backscattering as an indicator of prey abundance were used to model whale distribution patterns. We used an iterative approach employing (1) classification and regression tree (CART) models to identify oceanographic and ecological variables contributing to variability in humpbackMegaptera novaeangliaeand minkeBalaenoptera acutorstratawhale distribution, and (2) generalized additive models (GAMs) to elucidate functional ecological relationships between these variables and whale distribution. The CART models indicated that the cetacean distribution was tightly coupled with zooplankton acoustic volume backscatter in the upper (25 to 100 m), and middle (100 to 300 m) portions of the water column. Whale distribution was also related to distance from the ice edge and bathymetric slope. The GAMs indicated a persistent, strong, positive relationship between increasing zooplankton volume and whale relative abundance. Furthermore, there was a lower limit for averaged acoustic volume backscatter of zooplankton below which the relationship between whales and prey was not significant. The GAMs also supported an annual relationship between whale distribution, distance from the ice edge and bathymetric slope, suggesting that these are important features for aggregating prey. Our results demonstrate that during the 2 yr study, whales were consistently and predictably associated with the distribution of zooplankton. Thus, humpback and minke whales may be able to locate physical features and oceanographic processes that enhance prey aggregation.

DNA metabarcoding and morphological taxonomic (microscopic) analysis of the gut contents was used to examine diet diversity of seven species of fishes collected from mesopelagic depths (200-1000 m) in the NW Atlantic Ocean Slope Water during Summer 2018 and 2019. Metabarcoding used two gene regions: V9 hypervariable region of nuclear 18S rRNA and mitochondrial cytochrome oxidase I (COI). V9 sequences were classified into 14 invertebrate prey groups, excluding fish due to predator swamping. Ecological network analysis was used to evaluate relative strengths of predator-prey linkages. Multivariate statistical analysis revealed consistently distinct diets of four fish species in 2018 and/or 2019: Argyropelecus aculeatus, Chauliodus sloani, Hygophum hygomii , and Sigmops elongatus . Three other species analyzed ( Malacosteus niger, Nemichthys scolopaceus , and Scopelogadus beanii ) showed more variability between sampling years. COI sequences were classified into eight invertebrate prey groups, within which prey species were detected and identified. Considering all predator species together, a total of 77 prey species were detected with a minimum of 1,000 COI sequences, including 22 copepods, 18 euphausiids, and 7 amphipods. Morphological prey counts were classified into seven taxonomic groups, including a gelatinous group comprised of soft-bodied organisms. The ocean twilight zone or is home to exceptional diversity and biomass of marine fish, which are key players in deep sea food webs. This study used integrative morphological-molecular analysis to provide new insights into trophic relationships and sources of productivity for mesopelagic fishes, including identification of key prey species, recognition of the importance of gelatinous prey, and characterization of differences in diet among fish predators in the NW Atlantic Slope Water.