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The highly productive upwelling zone in the Southeast Pacific provides inputs of C and N to the oceanic deep-water system via lateral transport, although the quality of this organic matter, the organisms being transported, and the controlling mechanisms are unclear. Here, we assessed whether the taxonomic and trophic structure of zooplankton over more oceanic offshore waters differ from that of zooplankton located in the upwelling zone, along with the characterization of the oceanographic processes and variables associated with the lateral transport. For this, epipelagic, mesopelagic, and bathypelagic zooplankton along with oceanographic variables and particulate organic matter (POM) were analyzed at the upwelling zone and adjacent oceanic stations off Central Chile (27–33°S), some of which were located over the Atacama trench, during September 2016. The community structure of size-fractioned zooplankton (2000 μm size classes) was assessed by an automated method, along with the analysis of the isotopic niche computed from their δ15N and δ13C. The isotope signatures of POM indicated that diatoms contributed mostly to production of new organic C, being the main food source for small-sized zooplankton in coastal waters. The zooplankton community structure differed between the upwelling and oceanic areas, but their isotopic signature showed a large overlap (78%). Satellite-derived geostrophic flow and the depth of the mixed layer appeared as the main factors explaining the homogeneity in the isotopic and biogeochemical signatures of zooplankton between the upwelling zone and offshore waters. Cross-shelf advection is thus suggested as a key process promoting zooplankton export to the deep-water ecosystem, including the ultra-deep Atacama Trench., MOPEX Cruise was funded by the Chilean National Agency of Research and Development (ANID) Grant AUB 160003. Funding was also provided by the Millennium Institute of Oceanography (Grant ICN12_019) and ANID-FONDECYT Postdoctoral (3220468) and Initiation (11221079) Projects. We are grateful to D. Toledo, M. Cornejo and K. Aniñir for sampling assistance. The work is a contribution to TROPHONET Project REDES 1039 funded by ANID of Chile and SCOR WG155 on Eastern Boundary Upwelling Systems (EBUS).

Research article, Zooplankton grazing onphytoplankton promotes the release of particulate and dissolved organic matter (DOM) into the water column and therefore plays a key role in organic matter cycling in aquatic systems. Prokaryotes are the main DOM consumers in the ocean by actively remineralizing and transforming it, contributing to its molecular diversification. To explore the molecular composition of zooplankton-derived DOM and its bioavailability to natural prokaryotic communities, the DOM generated by a mixed zooplankton community in the coastal Atlantic off Spain was used as substrate for a natural prokaryotic community and monitored over a ~ 5-d incubation experiment. The molecular composition of solid-phase extracted DOM was characterized via Fourier-transform ion cyclotron resonance mass spectrometry. After ~ 4 d in the zooplankton-derived DOM amended incubation, the prokaryotic community demonstrated a 17-fold exponential increase in cell number. The prokaryotic growth resulted in a reduction in bulk dissolved organic carbon concentration and the zooplankton-derived DOM was considerably transformed at molecular and bulk elemental levels over the incubation period. The C : N ratio (calculated from the obtained molecular formulae) increased while the functional diversity decreased over the incubation time. In addition, molecular indices pointed to a reduced bioavailability of DOM at the end of the experiment. These findings show that zooplankton excreta are a source of labile organic matter that is quickly metabolized by the prokaryotic community. Therefore, a fraction of carbon is shunted from transfer to secondary consumers similarly to the viral shunt, suggesting that the zooplankton–prokaryotic interactions play an important role in the ocean's carbon cycle., IEO, XUNTA DE GALICIA (INGO7A 2018/2), DFG (CO 2218/2-1 and TRR51)

9 pages, 3 figures, 1 table, supplementary data https://doi.org/10.1093/icesjms/fsab189.-- Data availability statement: The original data on sample location, individual fish characteristics and stable isotope composition, including amino acids, can be accessed through the PANGAEA repository (Bode et al., 2021b). Similarly, the original data for copepods can be found in Bode et al. (2018b), The trophic position (TP) of fishes determines their importance in terms of energy flows within food webs. However, accurate estimations of TP are often prevented because of the difficulties in tracing all food sources. This is particularly challenging for omnivorous fishes, such as those from the Order Stomiiformes. In this study, we applied recent developments in stable isotope analysis of amino acids to untangle the contributions of microbial vs. metazoan food webs in 13 species of Stomiiformes. The inclusion of the microbial food web reduced the differences between TP estimates using stable isotopes and those derived from stomach content analysis. In addition, the new estimates allowed to quantify the relative contribution of the microbial food web to each species (6–21%), highlighting the importance of detritus consumption even in piscivorous species (e.g. Stomias boa and Chauliodus danae). The comparison of TP estimates obtained with selected amino acids in fish muscle allowed for the detection of the microbial influence integrated at time scales relevant for net fish growth, even when trophic exchanges in the microbial food web occur at much shorter time scales. The assessment of TP considering the differential contribution of microbial and metazoan food webs challenges our current understanding of marine food webs; yet provides a new quantitative tool for the analysis of their structure and function, This research was funded by projects MAFIA (CTM2012-39587-C04), BATHYPELAGIC (CTM2016-78853-R), and QLOCKS (PID2020-115620RB-100) from the Plan Estatal de I+D+I (Spain), projects SUMMER (grant agreement number 817806) and TRIATLAS (grant agreement number 817578), from the European Union (Horizon 2020 Research and Innovation Programme), and the support through the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

The ocean is the most important subsystem of the Earth’s climate system and functions as its heart, regulating the energy distribution of the planet. It has absorbed more than 90% of the energy accumulated since 1971 and about 30% of the emitted anthropogenic carbon dioxide. As a result, water temperature rises and oceans acidify and deoxygenate, which lead to changes in oceanic circulation and biogeochemistry, to rising sea levels, to more extreme weather events, to shifts in the distribution of species and migratory routes, and to loss of species and habitat diversity. Awareness of the importance of oceans for the sustainability of the global human population is increasing, including the conservation of biodiversity and its legacy to future generations [1]. For instance, oceanic organisms are more vulnerable to warming than terrestrial ones, as the former are generally at temperatures near their upper thermal limits and lack of thermal refuges [2]. Half of the atmospheric carbon fixed annually in natural systems is cycled into the ocean mainly by the biological carbon pump in the open ocean, but some of the main areas capturing and storing this carbon (as mangroves, seagrasses, salt marshes, and coastal upwelling ecosystems) cover less than 3% of the world’s ocean surface [3]. Particularly, eastern boundary upwelling systems are highly productive ecosystems, with up to 40% of the reported global fish catch [4].

Ship-based time-series observations provide crucial data for understanding marine ecosystems, improving decision making in ocean and coastal management. However, only a few time series survive for more than a decade. RADIALES is one of the longest multidisciplinary programs in operation in the northern and northwestern coast of Spain. In the last 30Â years, this program collected monthly data on physical, chemical and plankton observations in an array of five sections of stations representative of upwelling and stratified dynamics. Here, the main achievements, including key contributions to ecosystem conservation policies, are summarized. The development of this program, in line with similar initiatives in other countries, included phases focused on the study of seasonality, on comparative analysis, and lately on the analysis of decadal variability and regime shifts. Furthermore, in recent years there was a substantial improvement in the identification of plankton species by genomics. Among the main findings of RADIALES are the quantification of ocean warming at subsurface layers, the determination of climatologies in thermohaline, biogeochemical and biological variables, the inventory of plankton species (from bacteria to zooplankton) and the identification of regionally coherent regime shifts. Baselines defined by RADIALES series were instrumental for the assessment of environmental impacts (e.g. oil spills) and for the support of environmental policies (e.g. Marine Strategy Framework Directive). By contributing to international databases, data from programs as RADIALES combined with new instrumental observations will help to develop a more coherent and comprehensive understanding of the ocean ecosystems, enhancing our ability to detect and forecast risks., IEO, GAIN (Xunta de Galicia), 3,269

This research was supported by project Malaspina-2010 (CSD2008-00077) funded by program CONSOLIDERINGENIO 2010 (Ministerio de Ciencia e Innovación, Spain), by grant IN607A 2018/2 of the Axencia Galega de Innovación (GAIN, Xunta de Galicia, Spain). Thanks are also due to FCT/MCTES for the financial support to CESAM (UIDP/5 0017/2020+UIDB/50017/2020), through national funds. RA was supported by a Ph.D. fellowship funded by FCT (PD/ BD/113483/2015)., The marine neuston, organisms living in the vicinity of the ocean surface, is one of the least studied zooplankton groups. Neuston occupies a restricted ecological niche and is affected by a wide range of endo- and exogenous processes, while also being a food source to zooplankton, fish migrating from the deep layers and seabirds. In this study, the neustonic communities were characterized along the Malaspina global expedition sampling tropical and subtropical oceanic provinces using stable carbon and nitrogen isotopes to explore their trophic structure and relationships with environmental variables. The differences in stable isotopes mirrored the patterns in environmental characteristics of each province. High δ13C values were associated with continental and atmospheric carbon inputs, while the presence of dinoflagellates, coccolithophorids and upwelling influence are related to low δ13C values. Similarly, provinces presenting high δ15N values were associated with denitrification and nitrate diffusive fluxes, whereas the presence of low δ15N is attributable to nitrogen supplied through N2 fixation by diazotrophs. Neuston showed a large overlap among the isotopic niches of four functional groups, with chaetognaths and detritivore generally exhibiting a smaller degree of overlap compared to carnivores and omnivores. These results support the hypothesis of a common trophic structure in the neuston community across the ocean. However, the size of the niche, small in coastal areas and those influenced by upwelling and large in oligotrophic regions, and their overlap, low in more productive provinces and high in oligotrophic provinces, may be associated with food availability. Small trophic niches are associated with a dominance of specialized over-opportunistic feeding in productive environments.

This study was funded in part by a FundManagement Agreement between the IEO and the Ministerio para la Transición Ecológica (Spain) and by grant number IN607A2018/2 of the Axencia Galega de Innovación (GAIN, Xunta de Galicia, Spain)., The use of top-consumers as bioindicators of the health of food webs is hampered by uncertainties in their effective use of resources. In this study, the abundance of stable nitrogen isotopes in amino acids from homogenised eggs of the Yellow-legged Gull (Larus michahellis) allowed to identify variations in trophic resource exploitation between geographically adjacent nesting colonies in the Ria de Vigo (NW Spain) that exhibited marked differences in pollutants. Eggs from nests in the Cíes Islands (located in a National Park) showed a large variability in stable carbon and nitrogen isotopes in bulk egg content encompassing that of eggs from Vigo city (a major fishing harbour). However, both colonies differed in the relative concentration and abundance of nitrogen isotopes of lysine, an essential amino acid present in marine prey, but also extensively used in feed stocks for poultry and swine. Notwithstanding the similarity in trophic position for both colonies, gulls from Cíes Islands may have acquired a substantial fraction of lysine from garbage dump sites, while those of the urban colony relied on fish discards. This unexpected conclusion is partly supported by the large variability reported for gull's diet in this region and calls for detailed estimations of diet when assessing the conservation status and pollution risks of marine ecosystems.

Research article, We combined flow cytometry, CARD-FISH, and 16S rRNA gene tag pyrosequencing to investigate bacterioplankton dynamics along a transect in shelf waters off A Coruña (Galicia, NW Spain). Over five days (16-20th July 2012) we sampled during the relaxation of a summer upwelling pulse, providing an opportunity to examine the impact of pulses of cold nutrient-rich water into coastal microbial communities. The hydrographic conditions, characterized by intense density stratification of surface waters and the presence of a deep chlorophyll maximum (DCM) at 20–30 m, were relatively maintained over the sampling period. Indeed, bacterial abundance and composition displayed low day to day variation. Alpha diversity analysis suggested that species richness and diversity increased from coastal to shelf stations and from the surface down to the coastal DCM, which could be caused by the mixing of upwelled bacteria with the coastal surface waters. SAR11, SAR86, and Roseobacter were the most abundant bacteria detected in the samples by using CARD-FISH. The assemblages observed by pyrosequencing displayed a strong vertical zonation along the transect. Rhodobacteraceae (under class Alphaproteobacteria) and Bacteriodetes dominated the surface waters and decreased during the upwelling pulse, while SAR 86 (under class Gammaproteobacteria), Actinobacteria and SAR11 clade increased their relative abundance at the coastal DCM with upwelling relaxation, particularly at the shelf stations. Bacterial assemblages from surface waters were associated with higher temperature and light conditions, while coastal DCM assemblages were rather associated to salinity, inorganic nutrients and a diatom-bloom leading to high chlorophyll-a. Our findings suggest that the vertical variability in environmental conditions induced by the intense density stratification, the exportation of warmer and less saline surface water from the rias to the adjacent shelf, and the fertilizing effect of recently upwelled water at the deeper layer, determined the composition of distinct bacterial assemblages at the subsurface and DCM layers., XUNTA DE GALICIA (O9MMA 027604PR, 10MMA604024PR), IEO (PROYECTO RADIALES) y PLAN NACIONAL (MODUPLAN CTM2011-24008), 2,247

In this study, the trophic structure of the pelagic copepod community was analyzed in six geographical zones in the eastern tropical Pacific Ocean. Zooplankton samples were collected on two oceanographic cruises from August to December 2003 using a bongo cone net (333 μm mesh light). The isotopic signatures were measured as δ15N and δ13C in twelve selected species based on their dominance and their feeding types. We observed a significant latitudinal gradient in δ15N values, generally increasing northwards. The values of δ13C isotopes did not show a significant longitudinal gradient, but geographical differences occurred in some species. Pleuromamma robusta was recognized as the species with the highest trophic position in the copepod community. In general, there was a positive relationship between average body size and trophic position, except for the herbivorous species Eucalanus inermis, which was detected in the lowest trophic position. The isotopic niche was similar for each of the 12 species of copepods studied, but in most cases, the niche overlap between each pair of species was low and not higher than 50%. While the differences in isotopic signatures can be attributed mainly to the dominant nutrient sources in each zone, the low trophic niche overlap may be explained by the differential spatial distribution of species, reducing competition for food resources.

The structure of the pelagic food web is a key issue to understand the variability in C and N fluxes in the ocean. The trophic structure of the hyperiid amphipods and its spatial variation in the subtropical area of the Southeast Pacific was studied based on C and N isotope composition. Zooplankton were collected off northern and Central-southern Chile during September-November 2016 at the coastal zone and in oceanic areas. Measurements of δ15N and δ13C of individual hyperiids were used to estimate the trophic position for each species, and to assess the community isotopic niche for 3 pre-defined areas: the coastal upwelling (eutrophic) zone, the coastal transition (mesotrophic) zone and the oceanic (oligotrophic) zone, as well as three depth layers. Our results showed different assemblages of species among the 3 zones, but also a large overlap between the isotopic niche for all zones and depth layers. In the mesotrophic zone there was a noticeable increase in the trophic position of individual species. The trophic diversity of hyperiids over the study region was relatively high, including primary consumers, omnivores and carnivorous feeders, and was associated with the changes in productivity across zones. However, carnivory was more prevalent in the mesotrophic zone and in the mesopelagic (500–1000 m) layer. We suggest that such variations reflect both the biogeochemical processes controlling the productivity in each zone, and the life-traits of individual species.

8 pages, 4 figures, 1 table, supplementary information https://doi.org/10.1038/s41467-020-19875-7.-- Data sets generated during and/or analyzed during the current study are available at PANGAEA (http://www.pangaea.de). The zooplankton biomass from the Malaspina cruise is available in https://doi.org/10.1594/PANGAEA.922974, the deep-sea zooplankton biomass reviewed is available in https://doi.org/10.1594/PANGAEA.923149, the reviewed carbon flux and respiration are available in https://doi.org/10.1594/PANGAEA.923832, the acoustic backscatter is available in https://doi.org/10.1594/PANGAEA.922619, and the stable isotope data in https://doi.org/10.1594/PANGAEA.919314, The biological pump transports organic carbon produced by photosynthesis to the meso- and bathypelagic zones, the latter removing carbon from exchanging with the atmosphere over centennial time scales. Organisms living in both zones are supported by a passive flux of particles, and carbon transported to the deep-sea through vertical zooplankton migrations. Here we report globally-coherent positive relationships between zooplankton biomass in the epi-, meso-, and bathypelagic layers and average net primary production (NPP). We do so based on a global assessment of available deep-sea zooplankton biomass data and large-scale estimates of average NPP. The relationships obtained imply that increased NPP leads to enhanced transference of organic carbon to the deep ocean. Estimated remineralization from respiration rates by deep-sea zooplankton requires a minimum supply of 0.44 Pg C y−1 transported into the bathypelagic ocean, comparable to the passive carbon sequestration. We suggest that the global coupling between NPP and bathypelagic zooplankton biomass must be also supported by an active transport mechanism associated to vertical zooplankton migration, This study was funded by projects Malaspina (CSD2008-00077), Bathypelagic (CTM2016-78853-R) from the Spanish Ministry of Economy and Competitiveness, and to projects SUMMER (Grant Agreement 817806) and TRIATLAS (Grant Agreement 817578) from the European Union (Horizon 2020 Research and Innovation Programme), With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

A 29-year-long time series (1990&ndash, 2018) of phyto- and zooplankton abundance and composition is analyzed to uncover regime shifts related to climate and local oceanography variability. At least two major shifts were identified: one between 1997 and 1998, affecting zooplankton group abundance, phytoplankton species assemblages and climatic series, and a second one between 2001 and 2002, affecting microzooplankton group abundance, mesozooplankton species assemblages and local hydrographic series. Upwelling variability was relatively less important than other climatic or local oceanographic variables for the definition of the regimes. Climate-related regimes were influenced by the dominance of cold and dry (1990&ndash, 1997) vs. warm and wet (1998&ndash, 2018) periods, and characterized by shifts from low to high life trait diversity in phytoplankton assemblages, and from low to high meroplankton dominance for mesozooplankton. Regimes related to local oceanography were defined by the shift from relatively low (1990&ndash, 2001) to high (2002&ndash, 2018) concentrations of nutrients provided by remineralization (or continental inputs) and biological production, and shifts from a low to high abundance of microzooplankton, and from a low to high trait diversity of mesozooplankton species assemblages. These results align with similar shifts described around the same time for most regions of the NE Atlantic. This study points out the different effects of large-scale vs. local environmental variations in shaping plankton assemblages at multiannual time scales.

11 pages, 5 figures, 4 tables, In the last few decades, there has been a remarkable development of niche models to help understand the ecological response of species to current rapid environmental changes. In the present study, we applied niche modelling to the megafauna community of shelf waters of the northwestern (NW) and northern Iberian Peninsula in order to analyse the coexistence of different species taking into consideration their niche preferences. The Spanish Institute of Oceanography conducts the PELACUS multidisciplinary survey annually to assess pelagic fish stocks and collect information on the status of other ecosystem components, such as oceanographic conditions, phytoplankton, zooplankton and marine megafauna. Using data collected from these surveys, we developed niche models for 14 marine megafauna taxa (3 cetaceans, 10 seabirds and 1 fish) incorporating multi-trophic ecological descriptors collected simultaneously during the surveys alongside the more commonly used oceanographic variables (e.g. chlorophyll a and sea surface temperature). Megafauna niche models were developed by pooling observations from 2007 to 2013 and were found to be driven by mean fish biomass and its variability, in addition to sea surface temperature. Hierarchical clustering identified four distinct megafauna assemblages, the first comprising wide-ranging species and the other three associated with shelf-slope waters in Galicia, coastal/shelf waters in Galicia, and the eastern Cantabrian Sea, respectively. Community-level hotspot areas were found in shelf and shelf-break sectors of Galicia, along with small diversity spots scattered throughout the Cantabrian coastal area. Our results showed that synoptically collected survey-based ecological descriptors, especially acoustic-based preyscapes, were among the most important variables explaining megafauna niche preference. These findings highlight the advantage of using integrated ecosystem surveys to collect simultaneous information on a suite of ecosystem components for spatial assessments, Surveys were funded by the Instituto Español de Oceanografía's ECOPEL project, partially co-funded by the European Union Data Collection Framework program. Additional funding was contributed by the CONPELHAB (Marie Curie Career Integration Grant PCIG09-GA-2011-293774), REPRODUCE (MariFish 8120-09-9), LOTOFPEL (Plan Nacional de I+D+I, CTM 2010-16053) and CHALLENGES (CTM2013-47032-R) projects. Maite Louzao was funded by Juan de la Cierva (JCI-2010-07639) and Ramón y Cajal (RYC-2012-09897) postdoctoral contracts

20 pages, 11 figures, 3 tables, supplemental material https://doi.org/10.3389/fmars.2019.00535, Quantification of the actual amount of carbon export to the mesopelagic layer by both zooplankton and micronekton is at present a gap in the knowledge of the biological pump. These organisms perform diel vertical migrations exporting carbon through respiration, excretion, mortality, and egestion during their residence at depth. The role of zooplankton in active flux is nowadays partially assessed. However, micronekton active flux is scarcely known and only a few studies addressed this downward transport. Even less is known about the capacity of both communities to export carbon in the ocean. Here, we show the results of zooplankton and micronekton active flux across a productivity gradient in the tropical and subtropical Atlantic Ocean. Biomass vertical distribution from the surface up to 800 m depth by day and night was studied during April 2015 in a transect from 9°S to 25°N, covering from the quite oligotrophic zone off Brazil to the meso- and eutrophic areas of the equator, Guinea Dome, and the oceanic upwelling off Northwest Africa. Zooplankton and micronekton migrant biomass was estimated from day and night catches at different layers of the water column using MOCNESS-1 (1 m2 mouth area) and Mesopelagos (35 m2) nets, respectively. Respiratory flux was assessed by measuring the enzymatic activity of the electron transfer system (ETS) of organisms at depth. Results showed a close relationship between migrant biomass and respiratory flux in zooplankton and micronekton as expected. Using a rather conservative 50% of efficiency for the net used to capture micronekton, respiratory flux resulted in similar values for both communities. Gravitational (passive) flux measured using sediment traps increased from the oligotrophic toward the meso- and eutrophic zones. Total active flux (including respiration and estimated mortality, excretion, and gut flux) by zooplankton and micronekton accounted for about 25% of total flux (passive plus active) in oligotrophic zones. Total active flux also increased toward meso- and eutrophic zones, reaching about 80% of total flux and being at least twofold higher than passive flux. These results alert about an important underestimation of the ocean biological pump using only passive flux measurements, This work was supported by projects “Migrants and Active Flux in the Atlantic Ocean” (MAFIA, CTM2012-39587-C04), and “Biomass and Active Flux in the Bathypelagic Zone” (Bathypelagic, CTM2016-78853-R) from the Spanish Ministry of Economy and Competitiveness

Malaspina-2010 (CSD2008-00077) funded by program CONSOLIDER-INGENIO 2010; Ministerio de Ciencia e Innovacion (Spain), project EURO-BASIN (FP7-ENV-2010 264933) of the EU; Instituto Espanol de Oceanografia (IEO); PFPI (grant of IEO) to C.M.; FPU grant from the Spanish government (AP2010-5594) to B.F.-C.

Knowledge of the ecology of N 2 -fixing (diazotrophic) plankton is mainly limited to oligotrophic (sub)tropical oceans. However, diazotrophs are widely distributed and active throughout the global ocean. Likewise, relatively little is known about the temporal dynamics of diazotrophs in productive areas. Between February 2014 and December 2015, we carried out 9 one-day samplings in the temperate northwestern Iberian upwelling system to investigate the temporal and vertical variability of the diazotrophic community and its relationship with hydrodynamic forcing. In downwelling conditions, characterized by deeper mixed layers and a homogeneous water column, non-cyanobacterial diazotrophs belonging mainly to nifH clusters 1G (Gammaproteobacteria) and 3 (putative anaerobes) dominated the diazotrophic community. In upwelling and relaxation conditions, affected by enhanced vertical stratification and hydrographic variability, the community was more heterogeneous vertically but less diverse, with prevalence of UCYN-A (unicellular cyanobacteria, subcluster 1B) and non-cyanobacterial diazotrophs from clusters 1G and 3. Oligotyping analysis of UCYN-A phylotype showed that UCYN-A2 sublineage was the most abundant (74%), followed by UCYN-A1 (23%) and UCYN-A4 (2%). UCYN-A1 oligotypes exhibited relatively low frequencies during the three hydrographic conditions, whereas UCYN-A2 showed higher abundances during upwelling and relaxation. Our findings show the presence of a diverse and temporally variable diazotrophic community driven by hydrodynamic forcing in an upwelling system. Xunta de Galicia | Ref. EM2013/021 Ministerio de Economía, Industria y Competitividad | Ref. CTM2016-75451-C2-1-R Ministerio de Educación, Cultura y Deporte | Ref. FPU13/01674 Ministerio de Educación, Cultura y Deporte | Ref. EST16/00142 Universidad de Vigo | Ref. Axudas á investigación 2016

13 pages, 7 figures, 3 tables, supplementary data https://dx.doi.org/10.1093/icesjms/fsx243, Lanternfishes (Myctophidae) constitute the most important component of the daily vertically migrating mesopelagic fish community. This research addresses the estimation of the trophic position and diet of myctophids using stable isotope analyses. Fishes were collected across the central Atlantic, from a very productive zone influenced by the Mauritanian upwelling to the western oligotrophic equatorial waters. The survey also encompassed a zone of low oxygen concentration in the mesopelagic layers. Determinations of δ13C and δ15N values were made on the 20 most frequent and abundant myctophids, from small-sized species (e.g. Notolychnus valdivae) to larger ones (e.g. Myctophum punctatum). Isotope analyses on the seston and several plankton groups were also performed to assess the influence of zonal differences in trophic position (TP) calculations, and to use as food sources in diet estimations. Myctophids displayed a narrow range of trophic positions, being greater than 2 and less than 4, except for N. valdiviae (TP = 1.7). Comparisons of diets estimated through an isotopic mixing model differentiated the smallest species, with a strong seston signature (Diogenichthys atlanticus and N. valdiviae), from the Diaphus species of medium sizes, (D. brachycephalus, D. holti, and D. rafinesquii), which feed on prey of higher TP values, This research has been funded by the Spanish Ministerio de Economía y Competitividad through projects CTM2012-39587-C04 and CTM2016-78853-R

The analysis of a 28-year-long (1989–2016) series of monthly measurements of chlorophyll concentrations and primary production rates at a shelf station off A Coruña (NW Spain) provided evidence of changes at several time scales that were only partly related to upwelling intensity. Chlorophyll determinations were made in acetonic extracts and primary production rates by the measurement of 14C-uptake by natural phytoplankton populations in simulated in situ conditions. Wavelet analysis revealed multiple modes of variation in both series, particularly at high frequencies, but some were only significant for part of the series. For instance, the seasonal cycle was not uniform through the series despite the annual repetition of maxima and minima. At multiannual time scales, both series were divided in three quasi-decadal periods characterized by significant increases in mean values. Fluctuations in chlorophyll and primary production covaried with changes in upwelling intensity at annual scales, but annual means showed low correlation. Changes in dissolved nutrient concentrations from continental sources were the likely drivers of the observed changes in productivity at large time scales. Increases in the decadal mean rates of production and concentrations of chlorophyll were driven by increased intensity of spring blooms associated to increased nutrients and low salinity water in the surface. In contrast, blooms caused by upwelling nutrients remained unchanged along the series. This study illustrates the complexity of interactions in coastal upwelling areas at large time scales, where changes in continental nutrient inputs may affect phytoplankton production more than variations in upwelling intensity.

Atmospheric deposition of semivolatile aromatic hydrocarbons accounts for an important input of organic matter to the surface ocean. Nevertheless, the biogeochemical cycling and sinks of semivolatile aromatic hydrocarbons in the ocean remain largely uncharacterized. Here we present measurements of 64 polycyclic aromatic hydrocarbons in plankton and seawater from the Atlantic, Pacific, Indian and Southern Oceans, as well an assessment of their microbial degradation genes. Concentrations of the more hydrophobic compounds decreased when the plankton biomass was higher, consistent with the relevance of the biological pump. The mass balance for the global oceans showed that the settling fluxes of aromatic hydrocarbons in the water column were two orders of magnitude lower than the atmospheric deposition fluxes. This imbalance was high for low molecular weight hydrocarbons, such as phenanthrene and methylphenanthrenes, highly abundant in the dissolved phase. Parent polycyclic aromatic hydrocarbons were depleted to a higher degree than alkylated polycyclic aromatic hydrocarbons, and the degradation genes for polycyclic aromatic hydrocarbons were found to be ubiquitous in oceanic metagenomes. These observations point to a key role of biodegradation in depleting the bioavailable dissolved hydrocarbons and to the microbial degradation of atmospheric inputs of organic matter as a relevant process for the marine carbon cycle., The authors thank the support of the RV Hespérides staff and UTM technicians during the cruise and Antarctic campaign. G. Caballero and M. J. Ojeda are acknowledged for technical support during the extraction and processing of the samples. This work was funded by the Spanish Government through the Malaspina 2010 (CSD2008-00077), REMARCA (CTM2012-34673), SENTINEL (CTM2015-70535-P) and ISOMICS (CTM2015-65691-R) projects. M.V.-C. acknowledges a Leonardo award from BBVA Foundation. Predoctoral fellowships from the Spanish government (A.M.-V. and P.C.), Catalan government (E.C.-G.), Spanish Oceanography Institute (C.M.) and BBVA Foundation (B.G.-G.) are acknowledged. B.G.-G., A.M.-V., M.V.-C., P.C., E.C.-G., N.B. and J.D. belong to the “Global Change and Genomic Biogeochemistry” research group funded by the Catalan Government (2017SGR800).

The 15N2-tracer assay [Montoya et al. (1996) A simple, high-precision, high-sensitivity tracer assay for N2 fixation. Appl. Environ. Microbiol., 62, 986–993.] is the most used method for measuring biological N2 fixation in terrestrial and aquatic environments. The reliability of this technique depends on the purity of the commercial 15N2 gas stocks used. However, Dabundo et al. [(2014) PLoS One, 9, e110335.] reported the contamination of some of these stocks with labile 15N-labeled compounds (ammonium, nitrate and/or nitrite). The contamination of commercial 15N2 gas stocks with 15N-labeled nitrate and 142 ammonium and consequences for nitrogen fixation measurements. Considering that the tracer assay relies on the conversion of isotopically labeled 15N2 into organic nitrogen, this contamination may have led to overestimated N2 fixation rates. We conducted laboratory and field experiments in order to (i) test the susceptibility of 15N contaminants to assimilation by non-diazotroph organisms and (ii) determine the potential overestimation of the N2 fixation rates estimated in the field. Our findings indicate that the contaminant 15N-compounds are assimilated by non-diazotrophs organisms, leading to an overestimation of N2 fixation rates in the field up to 16-fold under hydrographic conditions of winter mixing.

Stable isotope ratios (SIR) are widely used to estimate food-web trophic levels (TLs). We built systems dynamic N-biomassbased models of diferent levels of complexity, containing explicit descriptions of isotope fractionation and of trophic level. The values of δ15N and TLs, as independent and emergent properties, were used to test the potential for the SIR of nutrients, primary producers, consumers, and detritus to align with food-web TLs. Our analysis shows that there is no universal relationship between TL and δ15N that permits a robust prognostic tool for confguration of food webs even if all system components can be reliably analysed. The predictive capability is confounded by prior dietary preference, intra-guild predation and recycling of biomass through detritus. These matters afect the dynamics of both the TLs and SIR. While SIR data alone have poor explanatory power, they would be valuable for validating the construction and functioning of dynamic models. This requires construction of coupled system dynamic models that describe bulk elemental distribution with an explicit description of isotope discriminations within and amongst functional groups and nutrient pools, as used here. Only adequately confgured models would be able to explain both the bulk elemental distributions and the SIR data. Such an approach would provide a powerful test of the whole model, integrating changing abiotic and biotic events across time and space., Sí

Taurine, an amino acid-like compound, acts as an osmostress protectant in many marine metazoans and algae and is released via various processes into the oceanic dissolved organic matter pool. Taurine transporters are widespread among members of the marine prokaryotic community, tentatively indicating that taurine might be an important substrate for prokaryotes in the ocean. In this study, we determined prokaryotic taurine assimilation and respiration throughout the water column along two transects in the North Atlantic off the Iberian Peninsula. Taurine assimilation efficiency decreased from the epipelagic waters from 55 ± 14% to 27 ± 20% in the bathypelagic layers (means of both transects). Members of the ubiquitous alphaproteobacterial SAR11 clade accounted for a large fraction of cells taking up taurine, especially in surface waters. Archaea (Thaumarchaeota + Euryarchaeota) were also able to take up taurine in the upper water column, but to a lower extent than Bacteria. The contribution of taurine assimilation to the heterotrophic prokaryotic carbon biomass production ranged from 21% in the epipelagic layer to 16% in the bathypelagic layer. Hence, we conclude that dissolved free taurine is a significant carbon and energy source for prokaryotes throughout the oceanic water column being utilized with similar efficiencies as dissolved free amino acids. Electronic supplementary material The online version of this article (10.1007/s00248-019-01320-y) contains supplementary material, which is available to authorized users.

Phytoplankton is a sentinel of marine ecosystem change. Composed by many species with different life-history strategies, it rapidly responds to environment changes. An analysis of the abundance of 54 phytoplankton species in Galicia (NW Spain) between 1989 and 2008 to determine the main components of temporal variability in relation to climate and upwelling showed that most of this variability was stochastic, as seasonality and long term trends contributed to relatively small fractions of the series. In general, trends appeared as non linear, and species clustered in 4 groups according to the trend pattern but there was no defined pattern for diatoms, dinoflagellates or other groups. While, in general, total abundance increased, no clear trend was found for 23 species, 14 species decreased, 4 species increased during the early 1990s, and only 13 species showed a general increase through the series. In contrast, series of local environmental conditions (temperature, stratification, nutrients) and climate-related variables (atmospheric pressure indices, upwelling winds) showed a high fraction of their variability in deterministic seasonality and trends. As a result, each species responded independently to environmental and climate variability, measured by generalized additive models. Most species showed a positive relationship with nutrient concentrations but only a few showed a direct relationship with stratification and upwelling. Climate variables had only measurable effects on some species but no common response emerged. Because its adaptation to frequent disturbances, phytoplankton communities in upwelling ecosystems appear less sensitive to changes in regional climate than other communities characterized by short and well defined productive periods., 1,9530

Growth and survival of two populations of Fucus vesiculosus were studied at its southern limit of distribution on the eastern Atlantic coast. Experimentally denuded areas at an estuarine and a semi-exposed site in an upwelling area (NW Spain) were followed for 17 months. Three different cohorts were detected during the sampling period. Differences among the three cohorts in terms of growth, reproduction and survival were detected. These differences may be due to the different time of appearance of the different cohorts or the presence of the previously established individuals when the second and third cohorts were recruited. Although growth of the cohorts recruiting in autumn was faster than for the cohorts recruiting in spring and summer, individual growth was represented in all cases by a logistic function, as the fastest rates of increase in length occurred during the first 6 months of life, and maximum length was attained after the thallus reached 1 year in age. In the same way, production was...

Stable isotope composition of brown macroalgae has been widely used to monitor N loading during the last decades but some of the required assumptions when using them to detect anthropogenic inputs remain untested. In this study several experiments were run with two key species, A. nodosum and F. vesiculosus , to determine internal nitrogen isotope dynamics. First, the equilibration of the isotopic values of the different parts of the thallus of these species was tested by growing them under different water sources. Then, nitrate uptake capacity and N transport along the frond were tested by 15 N enrichment experiments. The results indicate that although the growing tips had the highest uptake rates, older parts of the frond of both species have the capacity to incorporate N at low rates. No evidence of N transport along the thallus, from the tip to the basal segment of the frond or the converse was found. These results show that the growing tips of these macroalgae can be used to monitor N loadings at time scales from weeks ( F. vesiculosus ) to months ( A. nodosum ). The use of non-growing parts of the thallus to do retrospective studies cannot be recommended because of their measurable exchange of N with the surrounding water.

While it is generally agreed that δ15N of brown macroalgae can discriminate between anthropogenic and natural sources of nitrogen, this study provides new insights on net fractionation processes occurring in some of these species. The contribution of continental and marine sources of nitrogen to benthic macroalgae in the estuaryria system of A Coruña (NW Spain) was investigated by analyzing the temporal (at a monthly and annual basis) and spatial (up to 10 km) variability of δ15N in the macroalgae Ascophyllum nodosum and three species of the genus Fucus (F. serratus, F. spiralis and F. vesiculosus). Total nitrate and ammonium concentrations and δ15N-DIN, along with salinity and temperature in seawater were also studied to address the sources of such variability. Macroalgal δ15Nand nutrient concentrations decreased fromestuarine to marinewaters, suggesting larger dominance of anthropogenic nitrogen sources in the estuary. However, δ15N values of macroalgae were generally higher than those of ambient nitrogen at all temporal and spatial scales considered. This suggests that the isotopic composition of thesemacroalgae is strongly affected by fractionation during uptake, assimilation or release of nitrogen. The absence of correlation between macroalgal and water samples suggests that the δ15N of the species considered cannot be used for monitoring short-term changes. But their long lifespan and slow turnover rates make them suitable to determine the impact of the different nitrogen sources integrated over long-time periods., ANILE (CTM2009-08396 and CTM2010-08804-E) of the Plan Nacional de I+D+i (Spain), and RADIALES of the Instituto Español de Oceanografía., 3,2580

15 pages, 7 figures.-- Open Access under Creative Commons by Attribution Licence. Use, distribution and reproduction are un - restricted; authors and original publication must be credited, The dynamics of individual phytoplankton species are often more variable than predicted for their functional groups. This observation is poorly understood in highly perturbed coastal systems. Trends in the abundance of 54 phytoplankton species spanning a volumetric size range from ~102 to ~105 μm3 in shelf waters of the NW Iberian upwelling system since the late 1980s were modelled as a function of environmental variables. Functional traits were assessed for their ability to explain the across-species changes in occurrence and abundance due to the environmental drivers. Species-specific responses to the environment were heterogeneous, precluding any generalisation by taxonomic groups, although these responses were partially related to cell size. Smaller species showed a high probability of occurrence and higher abundance under high nitrate concentration in the euphotic layer, pointing to a major role of the upwelling of Eastern North Atlantic Central Water but also to inputs of nutrients mineralised over the shelf. However, cell size explained only a small amount of the variability in the individual species’ responses to the environment. Growth rate and maximum rate of nitrogen uptake normalised to cell size provided additional explanation for the individual species’ responses to the environmental drivers. Fast-growing species, though less efficient in taking up upwelled nutrients, thrived in favourable upwelling conditions. These results support the dominance of species of intermediate size during blooms in this upwelling region, and suggest that variability in functional traits among species rather than cell size alone affects the response of phytoplankton to environmental changes in upwelling ecosystems., Project RADIALES conducted and funded by the Instituto Español de Oceanografía (www.seriestemporales-ieo.com). J.O. was supported by a ‘Junta para la Ampliación de Estudios’ Fellowship (JAE-Doc programme 2011) from the CSIC and ESF

Article, The classical paradigm about marine N2 fixation establishes that this process is mainly constrained to nitrogen-poor tropical and subtropical regions, and substained by the colonial cyanobacterium Trichodesmium spp and diatom-diazothroph symbiosis. However, the application of molecular techniques allowed determining a high phylogenic environments where biological N2 fixation may be relevant. Between February 2014 and December 2015, we carried out 10 one-day samplings in the upwelling system off NW Iberia in order to: (1) investigate the seasonal variability in the magnitude of N2 fixation, (2) determine its biogeochemical role as a mechanism of new nitrogen supply, and (3) quantify the main diazotrophs in the region under contrasting hydrographic regimes. Our results indicate that the magnitude of N2 fixation in this region was relatively low (0.001+-0.002 - 0.095+-0.024 micromoles n /m3 d ), comparable to the lower-end of rates described for the subtropical NE Atlantic. Maximum rates were observed at the surface during both upwelling and relaxation conditions. The comparison with nitrate diffusive fluxes revealed the minor role of N2 fixation (

The trophic position of sardine (Sardina pilchardus) in spring was determined from the analysis of the natural abundance of stable nitrogen isotopes in the southern Bay of Biscay between 1998 and 2014. Mesozooplankton (200-500µm) was employed as the reference baseline, as its isotopic signature tracked the seasonal development of phytoplankton blooms. Two methods for estimating trophic position were compared. Scaling the isotopic enrichment per trophic level to the isotopic signature of the reference baseline produced lower trophic position estimates than those using the traditional assumption of a constant enrichment per trophic level. Indeed, the studied sardine population is placed as a mid trophic level consuming prey near the base of the food web. Zonal averages of trophic position increased with decreasing size of the sardine stock and increasing values of the upwelling index, suggesting the interaction of trophic relationships and population dynamics. The effects of changes in the timing of phytoplankton blooms and the release of competition for prey at low population densities were discussed as potential factors causing an increase in mean trophic position of sardine in recent years. The application of the scaled trophic position estimates to the monitoring of sardine populations will facilitate the comparison of different studies and provide new insights on the factors regulating these populations., PELACUS, ECOPEL and PELASSES (Instituto Español de Oceanografía, Spain), SARDYN (5FP grant number QLRT-2001-00818) and REPRODUCE (MariFish Joint Call grant number 8120-09-9).

AC is funded by the Ministry of Science and Innovation of Spain through project CTM2009-08783. AM was part funded by NERC UK project NE/K001345/1. KJF was funded by NERC UK through projects NE/H01750X/1 and NE/F003455/1. This review was supported in part by project EURO-BASIN (Ref. 264933, 7FP, European Union), and by a Leverhulme International Network ‘‘Placing marine mixotrophs in context: modelling mixotrophy in a changing world’’., Exploring climate and anthropogenic impacts on marine ecosystems requires an understanding of how trophic components interact. However, integrative end-to-end ecosystem studies (experimental and/or modelling) are rare. Experimental investigations often concentrate on a particular group or individual species within a trophic level, while tropho-dynamic field studies typically employ either a bottom-up approach concentrating on the phytoplankton community or a top-down approach concentrating on the fish community. Likewise the emphasis within modelling studies is usually placed upon phytoplankton- dominated biogeochemistry or on aspects of fisheries regulation. In consequence the roles of zooplankton communities (protists and metazoans) linking phytoplankton and fish communities are typically under-represented if not (especially in fisheries models) ignored. Where represented in ecosystem models, zooplankton are usually incorporated in an extremely simplistic fashion, using empirical descriptions merging various interacting physiological functions governing zooplankton growth and development, and thence ignoring physiological feedback mechanisms. Here we demonstrate, within a modelled plankton food-web system, how trophic dynamics are sensitive to small changes in parameter values describing zooplankton vital rates and thus the importance of using appropriate zooplankton descriptors. Through a comprehensive review, we reveal the mismatch between empirical understanding and modelling activities identifying important issues that warrant further experimental and modelling investigation. These include: food selectivity, kinetics of prey consumption and interactions with assimilation and growth, form of voided material, mortality rates at different age-stages relative to prior nutrient history. In particular there is a need for dynamic data series in which predator and prey of known nutrient history are studied interacting under varied pH and temperature regimes., FP7, MINECO, NERC, Leverhulme International Network, 3,2690

proyectos RADIALES (IEO) y EURO-BASIN (Ref. 264933, 7FP), Variability in the dominance of copepods vs. gelatinous plankton was analysed using monthly time-series covering the last 55 years and related to changes in climatic, oceanographic, and fishery conditions in the upwelling region of Galicia (NW Spain). Seasonality was generally the main component of variability in all groups, both along the coast and in the nearby ocean, but no common long-term trend was found. Coastal copepods increased since the early 1990s, and gelatinous plankton increased in the ocean during the 1980s. Different trends were found for gelatinous plankton in two coastal sites, characterized by increases in either medusae or tunicates. In all series, multiyear periods of relative dominance of gelatinous vs. copepod plankton were evident. In general, copepod periods were observed in positive phases of the main modes of regional climatic variability. Conversely, gelatinous periods occurred during negative climatic phases. However, the low correlations between gelatinous plankton and climatic, oceanographic, or fishery variables suggest that local factors play a major role in their proliferations., 7FP, IEO

EU Framework Programme 7 (EURO-BASIN, contract No. 264933); the University of Hamburg and the Deutsche Forschungsgemeinschaft.