Abstract 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‐ μ m 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.

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.

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, NO 3 − 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.

Vertical variations in physical and chemical conditions drive changes in marine zooplankton community 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 1000 m 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 temperature, on the phytoplankton size distribution and the surface large particulate organic matter while oxygen was an additional important factor for structuring zooplankton in the mesopelagic. Our results further suggest that future changes in surface phytoplankton size and taxa composition and mesopelagic oxygen loss might lead to profound shift 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.

Marine plankton mitigate anthropogenic greenhouse gases, modulate biogeochemical cycles, and provide fishery resources. Plankton is distributed across a stratified ecosystem of sunlit surface waters and a vast, though understudied, mesopelagic ‘dark ocean’. In this study, we mapped viruses, prokaryotes, and pico-eukaryotes across 32 globally-distributed cross-depth samples collected during the Tara Oceans Expedition, and assessed their ecologies. Based on depth and O 2 measurements, we divided the marine habitat into epipelagic, oxic mesopelagic, and oxygen minimum zone (OMZ) eco-regions. We identified specific communities associated with each marine habitat, and pinpoint environmental drivers of dark ocean communities. Our results indicate that water masses primarily control mesopelagic community composition. Through co-occurrence network inference and analysis, we identified signature communities strongly associated with OMZ eco-regions. Mesopelagic communities appear to be constrained by a combination of factors compared to epipelagic communities. Thus, variations in a given abiotic factor may cause different responses in sunlit and dark ocean communities. This study expands our knowledge about the ecology of planktonic organisms inhabiting the mesopelagic zone.

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.

Abstract Ocean plankton comprise organisms from viruses to fish larvae that are fundamental to ecosystem functioning and the provision of marine services such as fisheries and CO 2 sequestration. The latter services are partly governed by variations in plankton community composition and the expression of traits such as body size at community-level. While community assembly has been thoroughly studied for the smaller end of the plankton size spectrum, the larger end comprises ectotherms that are often studied at the species, or group-level, rather than as communities. The body size of marine ectotherms decreases with temperature, but controls on community-level traits remain elusive, hindering the predictability of marine services provision. Here, we leverage Tara Oceans datasets to determine how zooplankton community composition and size structure varies with latitude, temperature and productivity-related covariates in the global surface ocean. Zooplankton abundance and median size decreased towards warmer and less productive environments, as a result of changes in copepod composition. However, some clades displayed the opposite relationships, which may be ascribed to alternative feeding strategies. Given that climate models predict increasingly warmed and stratified oceans, our findings suggest that zooplankton communities will shift towards smaller organisms which might weaken their contribution to the biological carbon pump.

rapsus grapsus and Grapsus adscensionis are supralittoral crabs that are known to inhabit oceanic islands and depend on surface currents to recruit in the rocky shores. The ornamentation of the cephalothorax is very distinct among species, but morphological differences are controversial, and integrative studies with different approaches are needed. This study investigated the genetic variation among the populations of G. grapsus from Saint Peter and Saint Paul Archipelago (SP), Fernando de Noronha (FN), Rocas Atoll (RA) and Trindade Island (TR) in the western Atlantic, and G. adscensionis from the islands of Ascension (AI) and Saint Helena (SH) in the mid-Atlantic. Morphology was assessed by geometric morphometric analyses of the carapace and chelae, and numerical analyses of tubercles in the frontal plate of SP, FN, RA, TR and AI populations. In addition, dispersal of the larvae in the Atlantic Ocean was simulated performing a Lagrangian analysis using HYCOM reanalysis dataset as the ocean surface velocity field. The data obtained for the mitochondrial D-loop gene confirmed the distinctness of the two putative species and demonstrated the connectivity between the populations of G. grapsus from the three equatorial islands. The TR population presented unique haplotypes, as well as AI and SH. The geometric morphometric analyses showed differentiation between the carapace shapes for G. grapsus and G. adscensionis, however, the chelae shape does not allow to distinguish between the species or the population. The morphometric and molecular results were consistent with the pattern of particles dispersion in the Atlantic ocean. The larvae of SP, FN and RA mix after two months of drift, while the larvae of TR, AI, SH circulate only around the respective islands. The results reinforce the validation of the two species and the isolation of populations of G. grapsus in TR and of G. adscensionis in AI and SH. The populations of these islands might be maintained by self-recruitment, through larval behavior associated with the local current system, and therefore should be the target of conservation measures.

The influence of oceanographic fronts on the abundance and community composition of invertebrate larvae, mostly of benthic species, along nearly 2000 km of the southwestern Atlantic shelf (21–34°S) was investigated. Meroplankton was sampled through vertical hauls at 89 stations, distributed along 14 cross-shelf transects, during late spring 2010 and early summer 2011. Salinity and temperature were registered with a CTD/rosette system, which provided seawater for chlorophyll-a and nutrient concentrations estimations. Vertical profiles of temperature, salinity, chlorophyll-a and nutrients were used as proxies of the fronts. In addition, high-resolution thermosalinograph data were used to detect surface frontal features. Meroplankton abundance peaks were found at several fronts intersected by the ship, including upwelling zones, estuarine and plume fronts, a shelf-break front, and two cyclonic eddies. Furthermore, meroplankton abundance was also relatively higher at small-scale thermal and/or saline surface fronts observed along the shelf. Such increases in meroplankton abundance are likely to be ascribed to high nutrient input and primary production. Distinct taxa of invertebrate larvae occurred at different types of fronts, besides the coastal realm, which was virtually dominated by decapod, cirripede and bivalve larvae. Small-scale shelf fronts presented high abundances of decapod and gastropod larvae, for instance, while larvae of polychaetes were the most frequent in the estuarine front of Patos Lagoon section.

The ocean is home to myriad small planktonic organisms that underpin the functioning of marine ecosystems. However, their spatial patterns of diversity and the underlying drivers remain poorly known, precluding projections of their responses to global changes. Here we investigate the latitudinal gradients and global predictors of plankton diversity across archaea, bacteria, eukaryotes, and major virus clades using both molecular and imaging data from Tara Oceans. We show a decline of diversity for most planktonic groups toward the poles, mainly driven by decreasing ocean temperatures. Projections into the future suggest that severe warming of the surface ocean by the end of the 21st century could lead to tropicalization of the diversity of most planktonic groups in temperate and polar regions. These changes may have multiple consequences for marine ecosystem functioning and services and are expected to be particularly significant in key areas for carbon sequestration, fisheries, and marine conservation. VIDEO ABSTRACT.

This is the first study in Brazil to propose a trophic relationship between Chaetognatha and Cnidaria based on molecular data (COI gene - DNA barcode). Cnidaria COI sequences were detected in tissue samples from two specimens of Flaccisagitta enflata collected in the surroundings of Fernando de Noronha archipelago, Tropical Western Atlantic. The methodology used followed all the requirements for a safe molecular manipulation, making the possibility of contamination unlikely. A search carried in BLAST showed that both sequences provided here presented a combination of highest query coverage and match identity with Nemopilema nomurai (Cnidaria: Scyphozoa), supporting the hypothesis that they belong to a species of jellyfish. Since gelatinous taxa are known to integrate the broad diet of Chaetognatha, it is possible that F. enflata has ingested this type of item, which was not fully digested until the moment of sampling and was not visualized by the transparency method. Thus, the record of Scyphozoa by the COI marker suggests a trophic relationship between the respective taxon and F. enflata. Future studies using the association of markers as well as specific methodologies may constitute a new perspective on how to evaluate the trophic ecology of key zooplankton species, including the abundant F. enflata in the Atlantic ocean.

Lysmata arvoredensis nov. sp. a new species of shrimp from the south coast of Brazil with a key to species of Lysmata (Caridea: Lysmatidae) recorded in the southwestern Atlantic ABSTRACT Lysmata arvoredensis sp. nov. inhabits temperate waters in the south coast of Brazil and is named in tribute to the Marine Protected Area REBIO Arvoredo. This is the fourth species belonging to the genus Lysmata recorded for the region and the ninth for Brazil. L. arvoredensis sp. nov. can be distinguished from other species of Lysmata by the presence of a nearly completely fused accessory branch with a single free unguiform segment on the outer antennular flagellum; a rostrum with seven dorsal (2+5) and three ventral teeth; a stylocerite with a pointed tip bearing mesial setae; a second pereiopod with 22-24 carpal subsegments and 14-16 subsegments in the merus; a merus of the third pereiopod with five ventrolateral and 12 ventral spines on the propodus; and its color pattern, with red bands and patches in pleonites 2-3 that resemble a mask in dorsal view. Molecular characters demonstrate that L. arvoredensis sp. nov. is most closely related to other species of Lysmata belonging to the Neotropical and Cleaner clades. To support future ecological studies in the region, identification keys to the species of Lysmata recorded in the southwestern Atlantic Ocean are provided.

Plankton was sampled with various nets, from bottom or 500m depth to the surface, in many oceans of the world. Samples were imaged with a ZooScan. The full images were processed with ZooProcess which generated regions of interest (ROIs) around each individual object and a set of associated features measured on the object (see Gorsky et al 2010 for more information). The same objects were re-processed to compute features with the scikit-image toolbox (http://scikit-image.org). The 1,433,278 resulting objects were sorted by a limited number of operators, following a common taxonomic guide, into 93 taxa, using the web application EcoTaxa (http://ecotaxa.obs-vlfr.fr). The archive contains: taxa.csv.gz Table of the classification of each object in the dataset, with columns - objid: unique object identifier in EcoTaxa (integer number). - taxon: taxonomic name. Ambiguous names are made unique by including the name of the parent taxon in parentheses, after the name of the taxon. - lineage: full taxonomic lineage corresponding to this taxon. features_native.csv.gz Table of morphological features computed by ZooProcess. All features are computed on the object only, not the background. All area/length measures are in pixels. All grey levels are in encoded in 8 bits (0=black, 255=white). With columns - objid: same as above - area: area - mean: mean grey - stddev: standard deviation of greys - mode: modal grey - min: minimum grey - max: maximum grey - perim.: perimeter - width,height dimensions - major,minor: length of major,minor axis of the best fitting ellipse - circ.: circularity: 4pi(area/perim.^2) - feret: maximal feret diameter - intden: integrated density: mean*area - median: median grey - skew,kurt: skewness,kurtosis of the histogram of greys - %area: proportion of the image corresponding to the object - area_exc: area excluding holes - fractal: fractal dimension of the perimeter - skelarea: area of the one-pixel wide skeleton of the image - slope: slope of the cumulated histogram of greys - histcum1,2,3: grey level at quantiles 0.25, 0.5, 0.75 of the histogram of greys - nb1,2,3: number of objects after thresholding at the grey levels above - symetrieh,symetriev: index of horizontal,vertical symmetry - symetriehc,symetrievc: same but after thresholding at level histcum1 - convperim,convarea: perimeter,area of the convex hull of the object - fcons: contrast - thickr: thickness ratio: maximum thickness/mean thickness - elongation: elongation index: major/minor - range: range of greys: max-min - meanpos: relative position of the mean grey: (max-mean)/range - cv: coefficient of variation of greys: 100*(stddev/mean) - sr: index of variation of greys: 100*(stddev/range) - perimferet: index of the relative complexity of the perimeter: perim/feret - perimmajor: index of the relative complexity of the perimeter: perim/major features_skimage.csv.gz Table of morphological features recomputed with skimage.measure.regionprops on the ROIs produced by ZooProcess. See http://scikit-image.org/docs/dev/api/skimage.measure.html#skimage.measure.regionprops for documentation. inventory.txt Tree view of the taxonomy and number of images in each taxon, displayed as text. map.png Map of the sampling locations, to give an idea of the diversity sampled in this dataset. imgs Directory containing images of each object, named according to the object id objid and sorted in subdirectories according to their taxon.

Lysmata arvoredensis sp. nov. inhabits temperate waters in the south coast of Brazil and is named in tribute to the Marine Protected Area REBIO Arvoredo. This is the fourth species belonging to the genus Lysmata recorded for the region and the ninth for Brazil. L. arvoredensis sp. nov. can be distinguished from other species of Lysmata by the presence of a nearly completely fused accessory branch with a single free unguiform segment on the outer antennular flagellum; a rostrum with seven dorsal (2+5) and three ventral teeth; a stylocerite with a pointed tip bearing mesial setae; a second pereiopod with 22–24 carpal subsegments and 14–16 subsegments in the merus; a merus of the third pereiopod with five ventrolateral and 12 ventral spines on the propodus; and its color pattern, with red bands and patches in pleonites 2–3 that resemble a mask in dorsal view. Molecular characters demonstrate that L. arvoredensis sp. nov. is most closely related to other species of Lysmata belonging to the Neotropical and Cleaner clades. To support future ecological studies in the region, identification keys to the species of Lysmata recorded in the southwestern Atlantic Ocean are provided.

The aim of this study was to investigate whether the bacterioplankton activity in the meso-eutrophic Conceição Lagoon would increase significantly under allochthonous inputs of inorganic nutrients and organic carbon. Abundance and biomass of bacterioplankton were evaluated under three treatments: light (14 h light/10 h dark), complete darkness (dark-control), and nutrient (C + N + P—dark, 100 : 10 : 1) enrichments during 72 h. Nutrient enrichments promoted a significant increase in abundance (maximum of 19.0 ×10 9 cells·L −1 in the first 32 hours) and biomass of the heterotrophic bacterioplankton, which induced the formation of large clusters. Bacterial biomass remained constant in the non-enriched incubations (dark-control and light). Bacterial growth rates were significantly higher after nutrient additions (1.35 d −1 ), followed by control (0.79 d −1 ), and light (0.63 d −1 ) treatments, which were statistically equal ( p > 0.05 ). Bacterial production rates were also significantly higher under nutrient additions (1.28 d −1 ), compared to the control and light (0.50 d −1 and 0.44 d −1 , respectively), demonstrating that bacterial growth and production in this meso-eutrophic lagoon are under an immediate “bottom-up” regulation, followed by a potential top-down effect. These facts reinforce the urgency on improving the local wastewater management plan in order to prevent further expansion of anoxic waters.

The sally lightfoot crab, Grapsus grapsus (Linnaeus, 1758), has been recorded from the oceanic islands off Brazil, as well as from the Caribbean, in coastal rocky shores in the Eastern Pacific, and in Galapagos. We report the first observation of G. grapsus in freshwater habitat, registered in a stream at Trindade Island, Brazil.

DNA barcoding was used to identify crab larvae from the Marine Biological Reserve of Arvoredo, encompassing a coastal archipelago off the SW Atlantic coast (27°S, 48°W). Partial mitochondrial COI or 16S rRNA gene sequences were obtained for 488 larvae, leading to the identification of 20 species. The COI sequences generated 13 barcode index numbers (BINs) within Barcode of Life Data Systems (BOLD), among which 11 were concordant with single species. DNA from ∼ 6% of the larvae did not amplify using the primers tested; based on external morphological characteristics, these larvae represented four possible additional operational taxonomic units (OTUs) at the family level. Intraspecific variation for the COI and 16S rRNA genes was found to be < 2.6% and < 2.1% respectively (Kimura 2-parameter distance), whereas interspecific divergence ranged from 7.9% to 21.5% and 6.4% to 14.5%, respectively. These results imply that both genes are suitable for use in species identification of brachyuran crabs of this area. Molecular identification of this group successfully enabled the diagnosis of larvae of closely related species, including congeners in Mithrax, Achelous and Callinectes. In addition, eight out of 20 species recognized represent new records for the reserve suggesting that the brachyuran fauna in the area has been underestimated based on traditional biodiversity measures. The availability of primers suited to the targeted species, and the development of a taxonomically comprehensive DNA barcoding database are the major recommendations to improve the accuracy and feasibility of using DNA barcoding for species identification of SW Atlantic brachyuran crabs.

The spatial distribution of a spring/summer community of combined decapod and stomatopod larvae along the southwestern Atlantic shelf, and its possible linkages with hydrographical processes and with parental habitats were investigated. Vertical plankton hauls were performed between 21 °S and 34 °S, at 107 stations, distributed along 2000 km of coastline and up to 400 km offshore, during late spring 2010 and early summer 2011. Salinity and temperature were obtained with a CTD/rosette system, which provided seawater for chlorophyll and nutrient concentrations. A coupling between chlorophyll concentration and abundance of early larvae of benthic species was observed, suggesting that the larval release could be synchronized with phytoplankton maxima. Our findings indicated that the composition and abundance of larvae are strongly influenced by distance from the coast, freshwater sources and water mass distribution. Assemblages of larvae were observed in the most relevant environmental scenarios: (1) coastal environments, dominated by intertidal, shallow water or coastal species; (2) offshore environments, with predominance of holopelagic and deepwater species; and (3) southern continental runoff, mainly represented by benthic neritic crabs and shrimps. In addition, the large-scale distribution of larvae revealed a relationship with the adult’s distribution, shown mainly by the occurrence of larvae transported from the estuaries in the inner shelf and by larvae of the deepwater fauna found mainly along the outer shelf.

We investigated the effect of interannual variation on the biodiversity and distribution of decapod larvae at Saint Paul’s Rocks, an isolated small group of equatorial islands linked to the Atlantic Mid-Ocean ridge. Zooplankton samples were collected between 2003 and 2005 from a range of different environments off the archipelago. Horizontal surface hauls were conducted in the inlet and at three increasing distances from the archipelago, both in the morning and at night, using a 200-lm mesh net. Decapod larvae collected included 11 species and 37 individual taxa. Seven of them have not previously been recorded in the area. Samples taken at the shortest distance from the archipelago exhibited a significantly greater abundance than samples taken further away. No significant differences were detected between months in terms of decapod larvae abundance in the inlet. However, in the open ocean at night, the mean abundance was much higher in May and July. Decapod larvae abundance was much higher at night for both the open ocean and the inlet. Brachyuran and caridean larvae exhibited the greatest abundance and frequency in the inlet, mainly represented by the first-stage larvae of: Grapsus grapsus, Plagusia depressa, Platypodiella spectabilis and Alpheidae. In contrast, Sergestidae holopelagic shrimps were the most abundant and the most frequent larvae in the surface waters of the open ocean. The composition, abundance and distribution of decapod larvae were very similar during the 2 years sampled. Species accumulation curves indicate that even greater diversity exists, demonstrating that long-term research is important for acquiring accurate knowledge about the biodiversity of dynamic ecosystems such as those at Saint Paul’s Rocks.