ALBERT

Description: Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The production of organic material is fueled by upwelling of nutrient-rich deep waters and high incident light at the sea surface. However, biotic and abiotic factors can modify surface production and related biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions of their future functioning requires understanding of the mechanisms driving the biogeochemical cycles therein. In this field experiment, we used in situ mesocosms as tools to improve our mechanistic understanding of processes controlling organic matter cycling in the coastal Peruvian upwelling system. Eight mesocosms, each with a volume of ∼55 m3, were deployed for 50 d ∼6 km off Callao (12∘ S) during austral summer 2017, coinciding with a coastal El Niño phase. After mesocosm deployment, we collected subsurface waters at two different locations in the regional oxygen minimum zone (OMZ) and injected these into four mesocosms (mixing ratio ≈1.5 : 1 mesocosm: OMZ water). The focus of this paper is on temporal developments of organic matter production, export, and stoichiometry in the individual mesocosms. The mesocosm phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift coincided with a short-term increase in production during the A. sanguinea bloom, which left a pronounced imprint on organic matter C : N : P stoichiometry. However, C, N, and P export fluxes did not increase because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, export fluxes during the study were decoupled from surface production and sustained by the remaining plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant for most of the experiment. We explain this constancy by light limitation through self-shading by phytoplankton and by inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes remained balanced and there were few opportunities for blooms, which represents an event where the system becomes unbalanced. Overall, our mesocosm study revealed some key links between ecological and biogeochemical processes for one of the most economically important regions in the oceans.

Description: Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The production of organic material is fueled by upwelling of nutrient-rich deep waters and high incident light at the sea surface. However, biotic and abiotic factors can modify surface production and related biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions of their future functioning requires understanding of the mechanisms driving the biogeochemical cycles therein. In this field experiment, we used in situ mesocosms as tools to improve our mechanistic understanding of processes controlling organic matter cycling in the coastal Peruvian upwelling system. Eight mesocosms, each with a volume of ∼55 m3, were deployed for 50 d ∼6 km off Callao (12∘ S) during austral summer 2017, coinciding with a coastal El Niño phase. After mesocosm deployment, we collected subsurface waters at two different locations in the regional oxygen minimum zone (OMZ) and injected these into four mesocosms (mixing ratio ≈1.5 : 1 mesocosm: OMZ water). The focus of this paper is on temporal developments of organic matter production, export, and stoichiometry in the individual mesocosms. The mesocosm phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift coincided with a short-term increase in production during the A. sanguinea bloom, which left a pronounced imprint on organic matter C : N : P stoichiometry. However, C, N, and P export fluxes did not increase because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, export fluxes during the study were decoupled from surface production and sustained by the remaining plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant for most of the experiment. We explain this constancy by light limitation through self-shading by phytoplankton and by inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes remained balanced and there were few opportunities for blooms, which represents an event where the system becomes unbalanced. Overall, our mesocosm study revealed some key links between ecological and biogeochemical processes for one of the most economically important regions in the oceans.

Description: Marine nitrogen (N2) fixation supports significant primary productivity in the global ocean. However, in one of the most productive regions of the world ocean, the northern Humboldt Upwelling System (HUS), the magnitude and spatial distribution of this process remains poorly characterized. This study presents a spatially resolved dataset of N2 fixation rates across six coastal transects of the northern HUS off Peru (8°S – 16°S) during austral summer. N2 fixation rates were detected throughout the waters column including within the OMZ between 12°S and 16°S. N2 fixation rates were highest where the subsurface Oxygen Minimum Zone (OMZ, O2 〈20 µmol L-1) was most intense and estimated nitrogen (N) loss was highest. There, rates were measured throughout the water column. Hence the vertical and spatial distribution of rates indicates colocation of N2 fixation with N loss in the coastal productive waters of the northern HUS. Despite high phosphate and total dissolvable iron (TdFe) concentrations throughout the study area, N2 fixation was still generally low (1.19 ± 3.81 nmol L-1 d-1) and its distribution could not be directly explained by these two factors. Our results suggest that the distribution was likely influenced by a complex interplay of environmental factors including phytoplankton biomass and organic matter availability, and potentially iron, or other trace metal (co)-limitation of both N2 fixation and primary production. In general, our results support previous conclusions that N2 fixation in the northern HUS plays a minor role as a source of new N and to replenish the regional N loss. Key Points: A north-to-south pattern in N2 fixation rates was observed implying increased N turnover between 12°S and 16°S where N loss was pronounced Highest N2 fixation rates were measured in coastal productive waters above and within the OMZ, showing no clear relationship with Fe or P The magnitude of N2 fixation was low compared to predictions, estimated to account for ∼0.3% of primary production and 〈2% of local N loss

Description: The northern Humboldt Current upwelling system (HCS) belongs to the most productive marine ecosystems, providing five to eight times higher fisheries landings per unit area than other coastal upwelling systems. To solve this “Peruvian puzzle”, to elucidate the pelagic food-web structure and to better understand trophic interactions in the HCS, a combined stable isotope and fatty acid trophic biomarker approach was adopted for key zooplankton taxa and higher trophic positions with an extensive spatial coverage from 8.5 to 16°S and a vertical range down to 1,000 m depth. A pronounced regional shift by up to ∼5‰ in the δ15N baseline of the food web occurred from North to South. Besides regional shifts, δ15N ratios of particulate organic matter (POM) also tended to increase with depth, with differences of up to 3.8‰ between surface waters and the oxygen minimum zone. In consequence, suspension-feeding zooplankton permanently residing at depth had up to ∼6‰ higher δ15N signals than surface-living species or diel vertical migrants. The comprehensive data set covered over 20 zooplankton taxa and indicated that three crustacean species usually are key in the zooplankton community, i.e., the copepods Calanus chilensis at the surface and Eucalanus inermis in the pronounced OMZ and the krill Euphausia mucronata, resulting in an overall low number of major trophic pathways toward anchovies. In addition, the semi-pelagic squat lobster Pleuroncodes monodon appears to play a key role in the benthic-pelagic coupling, as indicated by highest δ13C’ ratios of −14.7‰. If feeding on benthic resources and by diel vertical migration, they provide a unique pathway for returning carbon and energy from the seafloor to the epipelagic layer, increasing the food supply for pelagic fish. Overall, these mechanisms result in a very efficient food chain, channeling energy toward higher trophic positions and partially explaining the “Peruvian puzzle” of enormous fish production in the HCS.

Description: Marine diazotrophs convert dinitrogen (N-2) gas into bioavailable nitrogen (N), supporting life in the global ocean. In 2012, the first version of the global oceanic diazotroph database (version 1) was published. Here, we present an updated version of the database (version 2), significantly increasing the number of in situ diazotrophic measurements from 13 565 to 55 286. Data points for N-2 fixation rates, diazotrophic cell abundance, and nifH gene copy abundance have increased by 184 %, 86 %, and 809 %, respectively. Version 2 includes two new data sheets for the nifH gene copy abundance of non-cyanobacterial diazotrophs and cell-specific N2 fixation rates. The measurements of N-2 fixation rates approximately follow a log-normal distribution in both version 1 and version 2. However, version 2 considerably extends both the left and right tails of the distribution. Consequently, when estimating global oceanic N-2 fixation rates using the geometric means of different ocean basins, version 1 and version 2 yield similar rates (43-57 versus 45-63 TgNyr (-1); ranges based on one geometric standard error). In contrast, when using arithmetic means, version 2 suggests a significantly higher rate of 223 +/- 30 TgNyr (-1) (mean +/- standard error; same hereafter) compared to version 1 (74 +/- 7 TgNyr (-1)). Specifically, substantial rate increases are estimated for the South Pacific Ocean (88 +/- 23 versus 20 +/- 2 TgNyr 1), primarily driven by measurements in the southwestern subtropics, and for the North Atlantic Ocean (40 +/- 9 versus 10 +/- 2 TgNyr (-1)). Moreover, version 2 estimates the N-2 fixation rate in the Indian Ocean to be 35 +/- 14 TgNyr (-1), which could not be estimated using version 1 due to limited data availability. Furthermore, a comparison of N-2 fixation rates obtained through different measurement methods at the same months, locations, and depths reveals that the conventional N-15(2) bubble method yields lower rates in 69% cases compared to the new N-15(2) dissolution method. This updated version of the database can facilitate future studies in marine ecology and biogeochemistry. The database is stored at the Figshare repository (https://doi.org/10.6084/m9.figshare.21677687; Shao et al., 2022).

Description: The Eastern Tropical South Pacific (ETSP), hosts an extensive Oxygen Minimum Zone (OMZ) in the water column which has a major imprint on local and global marine biogeochemistry. Due to the low oxygen conditions within the OMZ, microbial processes of nitrogen (N) loss, such as anammox and denitrification are sustained in the water column. These processes result in a pronounced N deficit which reduces bioavailable N for primary productivity and thus influences fisheries production in the region. To maintain a balanced marine N inventory regionally in ETSP, the N deficit would have to be compensated by N inputs via upwelling or N2 fixation. A classical assumption is that N2 fixation is favoured by iron (Fe) availability and a surplus of inorganic phosphate relative to inorganic nitrogen (this relativity is defined as P*), both conditions are present in the ETSP. Over the past decades, this assumption has been integrated into most coupled circulation and N-cycle biogeochemical models. These models indicate that there is a close spatial link between areas of high N loss, generally confined to OMZs and N2 fixation. On the contrary, other biogeochemical models have revealed that a close spatial link between N loss and N2 fixation in OMZ areas may give rise to run-away loss of fixed N in the ETSP, ultimately destabilizing the regional marine N inventory. While N loss processes are relatively well understood in the ETSP, the lack of a comprehensive dataset that resolves N2 fixation rates in both space and time constraints an accurate assessment of the regional marine N inventory, potential feedback mechanisms, and their impact on N turnover and productivity. Therefore, the main objective of this doctoral dissertation was to investigate the spatial distribution of N2 fixation relative to N loss in the ETSP, in order to understand potential feedbacks in the regional N cycle.

Description: The Humboldt Upwelling System (HUS) supports high levels of primary production and has the largest single-stock fishery worldwide. The high fish production is suggested to be related to high trophic transfer efficiency in the HUS. Mucous-mesh grazers (pelagic tunicates and gastropods) are mostly of low nutritious value and might reduce trophic transfer efficiency when they are locally abundant. Unfortunately, little is known about the spatial dynamics of mucous-mesh grazers from Peruvian waters, limiting our understanding of their potential ecological role(s). We provide a spatial assessment of mucous-mesh grazer abundance from the Peruvian shelf in austral summer 2018/2019 along six cross-shelf transects spanning from 8.5 to 16° S latitude. The community was dominated by appendicularians and doliolids. Salps occurred in high abundance but infrequently and pelagic gastropods were mostly restricted to the North. At low latitudes, the abundance of mucous-mesh grazers was higher than some key species of crustacean mesozooplankton. Transects in this region had stronger Ekman-transport, higher temperature, lower surface turbidity and a broader oxygenated upper water layer compared to higher-latitude transects. Small-scale lateral intrusions of upwelled water were potentially associated with high abundances of doliolids at specific stations. The high abundance and estimated ingestion rates of mucous-mesh grazers in the northern HUS suggest that a large flux of carbon from lower trophic levels is shunted to tunicates in recently upwelled water masses. The data provide important information on the ecology of mucous mesh grazers and stress the relevance to increase research effort on investigating their functioning in upwelling systems.

Description: This data is part of the BMBF projects CUSCO (Coastal Upwelling Systems in a Changing Ocean) and BioTip subproject Humboldt Tipping. Data was collected during cruise number MSM80 with research vessel Maria S. Merian from 23.12.2018 - 30.01.2019 (from Panama to Valparaiso) in the Humboldt Upwelling system off the Eastern Tropical south Pacific. Samples were taken by CTD- rosette sampler from different depths and analysed onboard for dissolved inorganic nutrients and total dissolved nutrients. Triplicate nutrient samples were analysed for concentrations with an autosampler (XY-2 autosampler, SEAL Analytical) and a continuous flow analyzer (QUAAtro Autoanalyzer, SEAL Analytical) using standard colorimetric and flourometric methods by Kastriot Qelaj. Dissolved organic nutrients were calculated as the difference of the two for respective nitrogen and phosphorous nutrients. Phosphate was measured according to Murphy and Riley (1962). Ammonium was measured fluorometrically based on Holmes et al. (1999). An empty cell means that corresponding nutrient samples were not taken for the respective depth.

Description: This data is part of the BMBF projects CUSCO (Coastal Upwelling Systems in a Changing Ocean) and BioTip subproject Humboldt Tipping. Here we report water column nitrogen fixation, carbon fixation rates and particulate organic matter composition from the upper 300 m. Data was collected during cruise number MSM80 with research vessel Maria S. Merian from 23.12.2018 - 30.01.2019 (from Panama to Valparaiso) in the Humboldt Upwelling system off the Eastern Tropical south Pacific. Samples were taken by CTD- rosette sampler from different depths, injected with 15N labelled N2 gas based on the modified dissolution method (Großkopf et al., 2012 and Mohr et al., 2010) and additionally 13C labeled sodium bicarbonate. Samples were incubated for 24 hours at light intensities that resemble the in situ light conditions. After incubation a volume of the sample (20 - 1500 ml) was filtered onto a pre-combusted Whatman GF/75 filter. Filters were frozen, transported to the institute on dry ice and measured on a mass spectrometer for Delta15N and 13C (Delta V Advantage Isotope Ratio MS, ThermoFisher) with the ConFlo IV interface (ThermoFisher). Nitrogen fixation rates were calculated based on Montoya et al (1996) while carbon fixation rates were calculated based on the equation by Slawyk et al. (1977). Limits of detection (LOD) and minimum quantifiable rates (MQR) for Nitrogen fixation rates were calculated according to the criteria described by White et al. (2020) and standard error propagation methods described in Gradoville et al. (2017) respectively. POP and Biogenic silica concentrations were measured spectrophotometrically following Hansen and Koroleff (1999).

Description: This data is part of the BMBF projects CUSCO (Coastal Upwelling Systems in a Changing Ocean) and BioTip subproject Humboldt Tipping. The file contains total dissolvable trace metal concentrations (Fe, Cd, Ni, Cu, Zn and Co) from various depths of sampled stations. Trace metal concentrations were determined via Inductively coupled plasma mass spectrometry (ICP-MS, Element XR, ThermoFisher Scientific) after pre-concentration as per Rapp et al. (2017).