Articles | Volume 20, issue 5
https://doi.org/10.5194/bg-20-945-2023
© Author(s) 2023. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/bg-20-945-2023
© Author(s) 2023. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Research article
| 
09 Mar 2023
Research article |
| 09 Mar 2023
| 09 Mar 2023
Zooplankton community succession and trophic links during a mesocosm experiment in the coastal upwelling off Callao Bay (Peru)
Patricia Ayón Dejo
Dirección General de Investigaciones en Oceanografía y Cambio Climático, Instituto del Mar del Perú (IMARPE), Callao, Perú
Elda Luz Pinedo Arteaga
Dirección General de Investigaciones en Oceanografía y Cambio Climático, Instituto del Mar del Perú (IMARPE), Callao, Perú
Anna Schukat
BreMarE Bremen Marine Ecology, University of Bremen, Marine Zoology, Bremen, Germany
Jan Taucher
Biological Oceanography, GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Rainer Kiko
Biological Oceanography, GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Laboratoire d'Océanographie de Villefranche-sur-Mer, Sorbonne Université, Villefranche-sur-Mer, France
Helena Hauss
Biological Oceanography, GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Department of Computer Science, Christian Albrechts University Kiel, Kiel, Germany
Sabrina Dorschner
BreMarE Bremen Marine Ecology, University of Bremen, Marine Zoology, Bremen, Germany
Wilhelm Hagen
BreMarE Bremen Marine Ecology, University of Bremen, Marine Zoology, Bremen, Germany
Mariona Segura-Noguera
Departament de Biologia Marina i Oceanografia, Institut de Ciències del Mar (ICM_CSIC), Barcelona, Spain
Biological Oceanography, GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Department of Computer Science, Christian Albrechts University Kiel, Kiel, Germany
Related authors
No articles found.
Ariadna Celina Nocera, Lars Stemmann, Marcel Babin, Tristan Biard, Julie Coustenoble, François Carlotti, Laurent Coppola, Lucas Courchet, Laetitia Drago, Amanda Elineau, Lionel Guidi, Helena Hauss, Laëtitia Jalabert, Lee Karp-Boss, Rainer Kiko, Manon Laget, Fabien Lombard, Andrew McDonnell, Camille Merland, Solène Motreuil, Thelma Panaïotis, Marc Picheral, Andreas Rogge, Anya Waite, and Jean-Olivier Irisson
Earth Syst. Sci. Data Discuss., https://doi.org/10.5194/essd-2025-522, https://doi.org/10.5194/essd-2025-522, 2025
Preprint under review for ESSD
Short summary
Short summary
Plankton and detritus play a key role in ocean health and climate regulation. We present a large global dataset of images and information collected from 2008 to 2018 using specialized underwater camera (UVP). This publicly available dataset will support more accurate ecological models and help train artificial intelligence tools, improving how scientists track ocean biodiversity and monitor environmental changes.
Yawouvi Dodji Soviadan, Miriam Beck, Joelle Habib, Alberto Baudena, Laetitia Drago, Alexandre Accardo, Remi Laxenaire, Sabrina Speich, Peter Brandt, Rainer Kiko, and Stemmann Lars
Biogeosciences, 22, 3485–3501, https://doi.org/10.5194/bg-22-3485-2025, https://doi.org/10.5194/bg-22-3485-2025, 2025
Short summary
Short summary
Key parameters representing the gravity flux in global models are sinking speed and vertical attenuation of exported material. We calculate, for the first time, these parameters in situ in the ocean for six intermittent blooms followed by export events using high-resolution (3 d) time series of 0–1000 m depth profiles from imaging sensors mounted on an Argo float. We show that sinking speed depends not only on size but also on the morphology of the particles, with density being an important property.
Alexandre Accardo, Rémi Laxenaire, Alberto Baudena, Sabrina Speich, Rainer Kiko, and Lars Stemmann
Biogeosciences, 22, 1183–1201, https://doi.org/10.5194/bg-22-1183-2025, https://doi.org/10.5194/bg-22-1183-2025, 2025
Short summary
Short summary
The open ocean helps mitigate climate change by storing CO2 via the biological carbon pump (BCP), which involves processes like organic carbon production at the surface and transferring it to the deep ocean via various pathways. By deploying an autonomous platform, we found significant marine snow accumulation from the surface to the mesopelagic zone in frontal regions between eddies. We suggest that the coupling of hydrodynamics at eddy edges and biological activity may enhance this process.
Nasrollah Moradi, Lili Hufnagel, Simon Ramondenc, Clara Flintrop, Rainer Kiko, Tim Fischer, Helena Hauss, Arne Körtzinger, Gerhard Fischer, and Morten Iversen
EGUsphere, https://doi.org/10.5194/egusphere-2025-347, https://doi.org/10.5194/egusphere-2025-347, 2025
Short summary
Short summary
Mesoscale eddies are suggested to enhance deep-sea carbon export, but quantifying carbon flux in these eddies remains challenging. This study combines in-situ camera particle profiles, carbon flux data, particle settling velocities, and respiration rates, while accounting for water temperature and oxygen concentration. Applied to Cape Verde's cyclonic eddies, it revealed a funnel-shaped flux pattern with doubled flux at the eddy core, highlighting their regional carbon sequestration impacts.
Philipp Suessle, Jan Taucher, Silvan Urs Goldenberg, Moritz Baumann, Kristian Spilling, Andrea Noche-Ferreira, Mari Vanharanta, and Ulf Riebesell
Biogeosciences, 22, 71–86, https://doi.org/10.5194/bg-22-71-2025, https://doi.org/10.5194/bg-22-71-2025, 2025
Short summary
Short summary
Ocean alkalinity enhancement (OAE) is a negative emission technology which may alter marine communities and the particle export they drive. Here, impacts of carbonate-based OAE on the flux and attenuation of sinking particles in an oligotrophic plankton community are presented. Whilst biological parameters remained unaffected, abiotic carbonate precipitation occurred. Among counteracting OAE’s efficiency, it influenced mineral ballasting and particle sinking velocities, requiring monitoring.
Joelle Habib, Lars Stemmann, Alexandre Accardo, Alberto Baudena, Franz Philip Tuchen, Peter Brandt, and Rainer Kiko
EGUsphere, https://doi.org/10.5194/egusphere-2024-3365, https://doi.org/10.5194/egusphere-2024-3365, 2024
Short summary
Short summary
This study investigates how carbon moves from the ocean surface to the depths in the equatorial Atlantic, contributing to long-term carbon storage. Using an Argo float equipped with a camera, we captured two periods with major carbon export events. By identifying particle types and their sinking behaviors, we found that smaller, compact particles are key drivers of carbon transport. Our findings underscore the value of using imaging tools on autonomous platforms in tracking carbon sequestration.
Mathilde Dugenne, Marco Corrales-Ugalde, Jessica Y. Luo, Rainer Kiko, Todd D. O'Brien, Jean-Olivier Irisson, Fabien Lombard, Lars Stemmann, Charles Stock, Clarissa R. Anderson, Marcel Babin, Nagib Bhairy, Sophie Bonnet, Francois Carlotti, Astrid Cornils, E. Taylor Crockford, Patrick Daniel, Corinne Desnos, Laetitia Drago, Amanda Elineau, Alexis Fischer, Nina Grandrémy, Pierre-Luc Grondin, Lionel Guidi, Cecile Guieu, Helena Hauss, Kendra Hayashi, Jenny A. Huggett, Laetitia Jalabert, Lee Karp-Boss, Kasia M. Kenitz, Raphael M. Kudela, Magali Lescot, Claudie Marec, Andrew McDonnell, Zoe Mériguet, Barbara Niehoff, Margaux Noyon, Thelma Panaïotis, Emily Peacock, Marc Picheral, Emilie Riquier, Collin Roesler, Jean-Baptiste Romagnan, Heidi M. Sosik, Gretchen Spencer, Jan Taucher, Chloé Tilliette, and Marion Vilain
Earth Syst. Sci. Data, 16, 2971–2999, https://doi.org/10.5194/essd-16-2971-2024, https://doi.org/10.5194/essd-16-2971-2024, 2024
Short summary
Short summary
Plankton and particles influence carbon cycling and energy flow in marine ecosystems. We used three types of novel plankton imaging systems to obtain size measurements from a range of plankton and particle sizes and across all major oceans. Data were compiled and cross-calibrated from many thousands of images, showing seasonal and spatial changes in particle size structure in different ocean basins. These datasets form the first release of the Pelagic Size Structure database (PSSdb).
Moritz Baumann, Allanah Joy Paul, Jan Taucher, Lennart Thomas Bach, Silvan Goldenberg, Paul Stange, Fabrizio Minutolo, and Ulf Riebesell
Biogeosciences, 20, 2595–2612, https://doi.org/10.5194/bg-20-2595-2023, https://doi.org/10.5194/bg-20-2595-2023, 2023
Short summary
Short summary
The sinking velocity of marine particles affects how much atmospheric CO2 is stored inside our oceans. We measured particle sinking velocities in the Peruvian upwelling system and assessed their physical and biochemical drivers. We found that sinking velocity was mainly influenced by particle size and porosity, while ballasting minerals played only a minor role. Our findings help us to better understand the particle sinking dynamics in this highly productive marine system.
Jens Hartmann, Niels Suitner, Carl Lim, Julieta Schneider, Laura Marín-Samper, Javier Arístegui, Phil Renforth, Jan Taucher, and Ulf Riebesell
Biogeosciences, 20, 781–802, https://doi.org/10.5194/bg-20-781-2023, https://doi.org/10.5194/bg-20-781-2023, 2023
Short summary
Short summary
CO2 can be stored in the ocean via increasing alkalinity of ocean water. Alkalinity can be created via dissolution of alkaline materials, like limestone or soda. Presented research studies boundaries for increasing alkalinity in seawater. The best way to increase alkalinity was found using an equilibrated solution, for example as produced from reactors. Adding particles for dissolution into seawater on the other hand produces the risk of losing alkalinity and degassing of CO2 to the atmosphere.
Rainer Kiko, Marc Picheral, David Antoine, Marcel Babin, Léo Berline, Tristan Biard, Emmanuel Boss, Peter Brandt, Francois Carlotti, Svenja Christiansen, Laurent Coppola, Leandro de la Cruz, Emilie Diamond-Riquier, Xavier Durrieu de Madron, Amanda Elineau, Gabriel Gorsky, Lionel Guidi, Helena Hauss, Jean-Olivier Irisson, Lee Karp-Boss, Johannes Karstensen, Dong-gyun Kim, Rachel M. Lekanoff, Fabien Lombard, Rubens M. Lopes, Claudie Marec, Andrew M. P. McDonnell, Daniela Niemeyer, Margaux Noyon, Stephanie H. O'Daly, Mark D. Ohman, Jessica L. Pretty, Andreas Rogge, Sarah Searson, Masashi Shibata, Yuji Tanaka, Toste Tanhua, Jan Taucher, Emilia Trudnowska, Jessica S. Turner, Anya Waite, and Lars Stemmann
Earth Syst. Sci. Data, 14, 4315–4337, https://doi.org/10.5194/essd-14-4315-2022, https://doi.org/10.5194/essd-14-4315-2022, 2022
Short summary
Short summary
The term
marine particlescomprises detrital aggregates; fecal pellets; bacterioplankton, phytoplankton and zooplankton; and even fish. Here, we present a global dataset that contains 8805 vertical particle size distribution profiles obtained with Underwater Vision Profiler 5 (UVP5) camera systems. These data are valuable to the scientific community, as they can be used to constrain important biogeochemical processes in the ocean, such as the flux of carbon to the deep sea.
Natalia Belkin, Tamar Guy-Haim, Maxim Rubin-Blum, Ayah Lazar, Guy Sisma-Ventura, Rainer Kiko, Arseniy R. Morov, Tal Ozer, Isaac Gertman, Barak Herut, and Eyal Rahav
Ocean Sci., 18, 693–715, https://doi.org/10.5194/os-18-693-2022, https://doi.org/10.5194/os-18-693-2022, 2022
Short summary
Short summary
We studied how distinct water circulations that elevate (cyclone) or descend (anticyclone) water from the upper ocean affect the biomass, activity and diversity of planktonic microorganisms in the impoverished eastern Mediterranean. We show that cyclonic and anticyclonic eddies differ in their community composition and production. Moreover, the anticyclone may be a potential bio-invasion and dispersal vector, while the cyclone may serve as a thermal refugee for native species.
Mariana Hill Cruz, Iris Kriest, Yonss Saranga José, Rainer Kiko, Helena Hauss, and Andreas Oschlies
Biogeosciences, 18, 2891–2916, https://doi.org/10.5194/bg-18-2891-2021, https://doi.org/10.5194/bg-18-2891-2021, 2021
Short summary
Short summary
In this study we use a regional biogeochemical model of the eastern tropical South Pacific Ocean to implicitly simulate the effect that fluctuations in populations of small pelagic fish, such as anchovy and sardine, may have on the biogeochemistry of the northern Humboldt Current System. To do so, we vary the zooplankton mortality in the model, under the assumption that these fishes eat zooplankton. We also evaluate the model for the first time against mesozooplankton observations.
Gerd Krahmann, Damian L. Arévalo-Martínez, Andrew W. Dale, Marcus Dengler, Anja Engel, Nicolaas Glock, Patricia Grasse, Johannes Hahn, Helena Hauss, Mark Hopwood, Rainer Kiko, Alexandra Loginova, Carolin R. Löscher, Marie Maßmig, Alexandra-Sophie Roy, Renato Salvatteci, Stefan Sommer, Toste Tanhua, and Hela Mehrtens
Earth Syst. Sci. Data Discuss., https://doi.org/10.5194/essd-2020-308, https://doi.org/10.5194/essd-2020-308, 2021
Preprint withdrawn
Short summary
Short summary
The project "Climate-Biogeochemistry Interactions in the Tropical Ocean" (SFB 754) was a multidisciplinary research project active from 2008 to 2019 aimed at a better understanding of the coupling between the tropical climate and ocean circulation and the ocean's oxygen and nutrient balance. On 34 research cruises, mainly in the Southeast Tropical Pacific and the Northeast Tropical Atlantic, 1071 physical, chemical and biological data sets were collected.
Cited articles
Algueró-Muñiz, M., Alvarez-Fernandez, S., Thor, P., Bach, L. T., Esposito,
M., Horn, H. G., Ecker, U., Langer, J. A. F., Taucher, J., Malzahn, A. M.,
Riebesell, U., and Boersma, M.: Ocean acidification effects on
mesozooplankton community development: Results from a long-term mesocosm
experiment, PLOS ONE, 12, 1–21, https://doi.org/10.1371/journal.pone.0175851, 2017. a
Argüelles, J., Lorrain, A., Cherel, Y., Graco, M., Tafur, R., Alegre, A.,
Espinoza, P., Taipe, A., Ayón, P., and Bertrand, A.: Tracking habitat
and resource use for the jumbo squid Dosidicus gigas: a stable isotope
analysis in the Northern Humboldt Current System, Mar. Biol., 159,
2105–2116, https://doi.org/10.1007/s00227-012-1998-2, 2012. a
Aronés, K., Grados, D., Ayón, P., and Bertrand, A.:
Spatio-temporal trends in zooplankton biomass in the northern Humboldt
current system off Peru from 1961-2012, Deep Sea Res. Pt. II, 169, 104656, https://doi.org/10.1016/j.dsr2.2019.104656,
2019. a, b, c, d
Aronés, K., Ayón, P., Hirche, H.-J., and Schwamborn, R.: Hydrographic
structure and zooplankton abundance and diversity off Paita, northern Peru
(1994 to 2004) – ENSO effects, trends and changes, J. Mar.
Syst., 78, 582–598, https://doi.org/10.1016/j.jmarsys.2009.01.002, 2009. a
Auel, H. and Verheye, H. M.: Hypoxia tolerance in the copepod Calanoides carinatus and the effect of an intermediate oxygen minimum layer on copepod
vertical distribution in the northern Benguela Current upwelling system and
the Angola-Benguela Front, J. Exp. Mar. Biol.
Ecol., 352, 234–243, https://doi.org/10.1016/j.jembe.2007.07.020, 2007. a
Ayón, P., Swartzman, G., Bertrand, A., Gutiérrez, M., and Bertrand,
S.: Zooplankton and forage fish species off Peru: Large-scale bottom-up
forcing and local-scale depletion, Prog. Oceanogr., 79, 208–214,
https://doi.org/10.1016/j.pocean.2008.10.023, 2008b. a, b, c, d
Bach, L. T., Boxhammer, T., Larsen, A., Hildebrandt, N., Schulz, K. G., and
Riebesell, U.: Influence of plankton community structure on the sinking
velocity of marine aggregates, Global Biogeochem. Cy., 30, 1145–1165,
https://doi.org/10.1002/2016GB005372, 2016. a
Bach, L. T., Paul, A. J., Boxhammer, T., von der Esch, E., Graco, M., Schulz, K. G., Achterberg, E., Aguayo, P., Arístegui, J., Ayón, P., Baños, I., Bernales, A., Boegeholz, A. S., Chavez, F., Chavez, G., Chen, S.-M., Doering, K., Filella, A., Fischer, M., Grasse, P., Haunost, M., Hennke, J., Hernández-Hernández, N., Hopwood, M., Igarza, M., Kalter, V., Kittu, L., Kohnert, P., Ledesma, J., Lieberum, C., Lischka, S., Löscher, C., Ludwig, A., Mendoza, U., Meyer, J., Meyer, J., Minutolo, F., Ortiz Cortes, J., Piiparinen, J., Sforna, C., Spilling, K., Sanchez, S., Spisla, C., Sswat, M., Zavala Moreira, M., and Riebesell, U.: Factors controlling plankton community production, export flux, and particulate matter stoichiometry in the coastal upwelling system off Peru, Biogeosciences, 17, 4831–4852, https://doi.org/10.5194/bg-17-4831-2020, 2020. a, b, c, d, e, f, g, h, i, j, k, l, m, n, o, p, q
Bakun, A. and Weeks, S. J.: The marine ecosystem off Peru: What are the
secrets of its fishery productivity and what might its future hold?,
Prog. Oceanogr., 79, 290–299, https://doi.org/10.1016/j.pocean.2008.10.027,
2008. a, b, c
Båmstedt, U., Gifford, D., Irigoien, X., Atkinson, A., and Roman, M.:
Feeding, in: ICES Zooplankton Methodology Manual, edited by Harris, R.,
Wiebe, P., Lenz, J., Skjoldal, H. R., and Huntley, M., Academic
Press, London, 297–399, https://doi.org/10.1016/B978-012327645-2/50009-8, 2000. a, b, c
Barlow, R. G., Cummings, D. G., and Gibb, S. W.: Improved resolution of mono-
and divinyl chlorophylls a and b and zeaxanthin and lutein in phytoplankton
extracts using reverse phase C-8 HPLC, Mar. Ecol. Prog. Ser., 161,
303–307, 1997. a
Bernales, A., Sanchez, S., Bach, L. T., Graco, M., Ledesma, J., Chang, F., Franco, A., Walz, K., and Riebesell, U.: Succession of phytoplankton in response to a simulated upwelling event in the Northern Humboldt Current System, in preparation, 2023. a
Bertrand, A., Chaigneau, A., Peraltilla, S., Ledesma, J., Graco, M., Monetti,
F., and Chavez, F. P.: Oxygen: A Fundamental Property Regulating Pelagic
Ecosystem Structure in the Coastal Southeastern Tropical Pacific, PLOS ONE,
6, 1–8, https://doi.org/10.1371/journal.pone.0029558, 2011. a
Cabana, G. and Rasmussen, J. B.: Modelling food chain structure and contaminant
bioaccumulation using stable nitrogen isotopes, Nature, 372, 255–257,
https://doi.org/10.1038/372255a0, 1994. a
Carr, M.-E.: Estimation of potential productivity in Eastern Boundary Currents
using remote sensing, Deep Sea Res. Pt. II, 49, 59–80, https://doi.org/10.1016/S0967-0645(01)00094-7, 2001. a
Chavez, F. P. and Messié, M.: A comparison of Eastern Boundary Upwelling
Ecosystems, Prog. Oceanogr., 83, 80–96,
https://doi.org/10.1016/j.pocean.2009.07.032, 2009. a, b
Chavez, F. P., Bertrand, A., Guevara-Carrasco, R., Soler, P., and Csirke, J.:
The northern Humboldt Current System: Brief history, present status and a
view towards the future, Prog. Oceanogr., 79, 95–105,
https://doi.org/10.1016/j.pocean.2008.10.012, 2008. a, b
Checkley, D. M.: Food limitation of egg production by a marine, planktonic
copepod in the sea off southern California, Limnol. Oceanogr., 25,
991–998, https://doi.org/10.4319/lo.1980.25.6.0991, 1980a. a, b, c
Checkley, D. M.: The egg production of a marine planktonic copepod in relation
to its food supply: Laboratory studies, Mar. Ecol. Prog. Ser. 25,
430–446, https://doi.org/10.4319/lo.1980.25.3.0430, 1980b. a, b, c
Criales-Hernández, M. I., Schwamborn, R., Graco, M., Ayón, P., Hirche,
H. J., and Wolff, M.: Zooplankton vertical distribution and migration off
Central Peru in relation to the oxygen minimum layer, Helgoland Mar. Res., 62, 85–100, https://doi.org/10.1007/s10152-007-0094-3, 2008. a
Cullen, J. J., Lewis, M. R., Davis, C. O., and Barber, R. T.: Photosynthetic
characteristics and estimated growth rates indicate grazing is the proximate
control of primary production in the equatorial Pacific, J. Geophys. Res., 97, 639–654, https://doi.org/10.1029/91JC01320, 1992. a
Dalsgaard, J., St John, M., Kattner, G., Müller-Navarra, D., and Hagen, W.:
Fatty acid trophic markers in the pelagic marine environment, Adv. Mar. Biol.,
46, 225–340, https://doi.org/10.1016/s0065-2881(03)46005-7, 2003. a, b, c
Ehrhardt, M. and Koeve, W.: Determination of particulate organic carbon and
nitrogen, Chap. 17, John Wiley and Sons, Ltd, 437–444,
https://doi.org/10.1002/9783527613984.ch17, 1999. a
Escribano, R.: Population dynamics of Calanus chilensis in the Chilean
Eastern Boundary Humboldt Current, Fish. Oceanogr., 7, 245–251,
https://doi.org/10.1046/j.1365-2419.1998.00078.x, 1998. a
Escribano, R., Hidalgo, P., Fuentes, M., and Donoso, K.: Zooplankton time
series in the coastal zone off Chile: Variation in upwelling and responses of
the copepod community, Prog. Oceanogr., 97, 174–186,
https://doi.org/10.1016/j.pocean.2011.11.006, 2012. a
Espinoza, P. and Bertrand, A.: Revisiting Peruvian anchovy (Engraulis ringens) trophodynamics provides a new vision of the Humboldt Current
system, Prog. Oceanogr., 79, 215–227,
https://doi.org/10.1016/j.pocean.2008.10.022, 2008. a, b, c
Espinoza, P. and Bertrand, A.: Ontogenetic and spatiotemporal variability in
anchoveta Engraulis ringens diet off Peru, J. Fish Biol.,
84, 422–435, https://doi.org/10.1111/jfb.12293, 2014. a, b
Espinoza, P., Bertrand, A., van der Lingen, C. D., Garrido, S., and Rojas de
Mendiola, B.: Diet of sardine (Sardinops sagax) in the northern
Humboldt Current system and comparison with the diets of clupeoids in this
and other eastern boundary upwelling systems, Prog. Oceanogr., 83,
242–250, https://doi.org/10.1016/j.pocean.2009.07.045, 2009. a, b
Espinoza, P., Lorrain, A., Ménard, F., Cherel, Y., Tremblay-Boyer, L.,
Argüelles, J., Tafur, R., Bertrand, S., Tremblay, Y., Ayón, P.,
Munaron, J. M., Richard, P., and Bertrand, A.: Trophic structure in the
northern Humboldt Current system: new perspectives from stable isotope
analysis, Mar. Biol., 164, 86, https://doi.org/10.1007/s00227-017-3119-8, 2017. a
Field, A., Miles, J., and Fields, Z. (Eds.): Discovering Statistics using R,
Sage Publications, Ltd, ISBN 978-1-4462-0046-9, 2012. a
Folch, J., Lees, M. B., and Stanley, G.: A simple method for the isolation and
purification of total lipids from animal tissues, J. Biol. Chem., 2261, 497–509, 1957. a
Franz, J., Krahmann, G., Lavik, G., Grasse, P., Dittmar, T., and Riebesell, U.:
Dynamics and stoichiometry of nutrients and phytoplankton in waters
influenced by the oxygen minimum zone in the eastern tropical Pacific, Deep
Sea Res. Pt I, 62, 20–31,
https://doi.org/10.1016/j.dsr.2011.12.004, 2012a. a, b
Franz, J. M. S., Hauss, H., Sommer, U., Dittmar, T., and Riebesell, U.: Production, partitioning and stoichiometry of organic matter under variable nutrient supply during mesocosm experiments in the tropical Pacific and Atlantic Ocean, Biogeosciences, 9, 4629–4643, https://doi.org/10.5194/bg-9-4629-2012, 2012b. a
Gannes, L. Z., O'Brien, D. M., and del Rio Martínez, C.: Stable isotopes in
animal ecology: assumptions, caveats, and a call for more laboratory
experiments, Ecology, 78, 1271–1276,
https://doi.org/10.1890/0012-9658(1997)078[1271:SIIAEA]2.0.CO;2, 1997. a
García-Reyes, M., Sydeman, W. J., Schoeman, D. S., Rykaczewski, R. R.,
Black, B. A., Smit, A. J., and Bograd, S. J.: Under Pressure: Climate
Change, Upwelling, and Eastern Boundary Upwelling Ecosystems, Front. Mar. Sci., 2, 109, https://doi.org/10.3389/fmars.2015.00109, 2015. a
Gentsch, E., Kreibich, T., Hagen, W., and Niehoff, B.: Dietary shifts in the
copepod Temora longicornis during spring: evidence from stable isotope
signatures, fatty acid biomarkers and feeding experiments, J. Plankton Res., 31, 45–60, https://doi.org/10.1093/plankt/fbn097, 2008. a
Gorsky, G., Ohman, M. D., Picheral, M., Gasparini, S., Stemmann, L., Romagnan,
J.-B., Cawood, A., Pesant, S., García-Comas, C., and Prejger, F.:
Digital zooplankton image analysis using the ZooScan integrated system,
J. Plankton Res., 32, 285–303, https://doi.org/10.1093/plankt/fbp124,
2010. a, b
Graco, M. I., Purca, S., Dewitte, B., Castro, C. G., Morón, O., Ledesma, J., Flores, G., and Gutiérrez, D.: The OMZ and nutrient features as a signature of interannual and low-frequency variability in the Peruvian upwelling system, Biogeosciences, 14, 4601–4617, https://doi.org/10.5194/bg-14-4601-2017, 2017. a, b, c
Graham, B., Koch, P., Newsome, S., McMahon, K., and Aurioles, D.: Using
isoscapes to trace the movements and foraging behavior of top predators in
oceanic ecosystems, in: Isoscapes: Understanding movement, pattern, and
process on Earth through isotope mapping, edited by: West, J., Bowen, G.,
Dawson, T., and Tu, K., Springer Netherlands, Dordrecht, 299–318, 2010. a
Granger, J., Sigman, D. M., Lehmann, M. F., and Tortell, P. D.: Nitrogen and
oxygen isotope fractionation during dissimilatory nitrate reduction by
denitrifying bacteria, Limnol. Oceanogr., 53, 2533–2545,
https://doi.org/10.4319/lo.2008.53.6.2533, 2008. a
Gruber, N., Hauri, C., Lachkar, Z., Loher, D., Frölicher, T. L., and
Plattner, G. K.: Rapid progression of ocean acidification in the California
Current System, Science, 337, 220–223, https://doi.org/10.1126/science.1216773, 2012. a
Hagen, W.: Lipids, in: ICES Zooplankton Methodology
Manual, edited by: Harris, R., Wiebe, P., Lenz, J., Skjoldal,
H. R., and Huntley, M., Academic Press, London, 113–119,
45, https://doi.org/10.1016/B978-012327645-2/50009-8, 2000. a
Hauss, H., Franz, J. M. S., and Sommer, U.: Changes in N:P stoichiometry
influence taxonomic composition and nutritional quality of phytoplankton in
the Peruvian upwelling, J. Sea Res., 73, 74–85,
https://doi.org/10.1016/j.seares.2012.06.010, 2012. a
Hauss, H., Franz, J. M., Hansen, T., Struck, U., and Sommer, U.: Relative
inputs of upwelled and atmospheric nitrogen to the eastern tropical North
Atlantic food web: Spatial distribution of δ15N in
mesozooplankton and relation to dissolved nutrient dynamics, Deep Sea
Res. Pt. I, 75, 135–145,
https://doi.org/10.1016/j.dsr.2013.01.010, 2013. a
Hobson, K. A., Alisauskas, R. T., and Clark, R. G.: Stable-Nitrogen Isotope
Enrichment in Avian Tissues Due to Fasting and Nutritional Stress:
Implications for Isotopic Analyses of Diet, The Condor, 95, 388–394,
https://doi.org/10.2307/1369361, 1993. a
Ingall, E. and Jahnke, R.: Evidence for enhanced phosphorus regeneration from
marine sediments overlain by oxygen depleted waters, Geochim.
Cosmochim. Ac., 58, 2571–2575, https://doi.org/10.1016/0016-7037(94)90033-7, 1994. a
Itoh, H. and Nishida, S.: Life history of the copepod Hemicyclops gomsoensis (Poecilostomatoida, Clausidiidae) associated with decapod burrows
in the Tama-River estuary, central Japan, Plankton Benthos Res., 2,
134–146, https://doi.org/10.3800/pbr.2.134, 2007. a
Itoh, H. and Nishida, S.: Life history of the copepod Hemicyclops spinulosus (Poecilostomatoida, Clausidiidae) associated with crab burrows
with notes on male polymorphism and precopulatory mate guarding, Plankton
Benthos Res., 3, 189–201, https://doi.org/10.3800/pbr.3.189, 2008. a
Judkins, D. C.: Vertical distribution of zooplankton in relation to the oxygen
minimum off Peru, Deep Sea Res., 27, 475–487, https://doi.org/10.1016/0198-0149(80)90057-6, 1980. a
Kalvelage, T., Jensen, M. M., Contreras, S., Revsbech, N. P., Lam, P.,
Günter, M., LaRoche, J., Lavik, G., and Kuypers, M. M. M.: Oxygen
Sensitivity of Anammox and Coupled N-Cycle Processes in Oxygen Minimum
Zones, PLoS ONE, 6, e29299, https://doi.org/10.1371/journal.pone.0029299, 2011. a
Karl, D. M., Letelier, R., Hebel, D., Tupas, L., Dore, J., Christian, J., and
Winn, C.: Ecosystem changes in the North Pacific subtropical gyre attributed
to the 1991–92 El Niño, Nature, 373, 230–234,
https://doi.org/10.1038/373230a0, 1995. a
Kattner, G. and Fricke, H. S.: Simple gas-liquid chromatographic method for the
simultaneous determination of fatty acids and alcohols in wax esters of
marine organisms, J. Chromatogr. A, 361, 263–268,
https://doi.org/10.1016/S0021-9673(01)86914-4, 1986. a
Kiørboe, T., Møhlenberg, F., and Riisgård, H. U.: In situ feeding
rates of planktonic copepods: A comparison of four methods, J. Exp. Mar. Biol. Ecol., 88, 67–81,
https://doi.org/10.1016/0022-0981(85)90202-3, 1985.
a
Kleppel, G. S. and Pieper, R. E.: Phytoplankton pigments in the gut contents
of planktonic copepods from coastal waters off southern California, Mar. Biol., 78, 193–198, https://doi.org/10.1007/BF00394700, 1984. a
Kleppel, G. S., Pieper, R. E., and Trager, G.: Variability in the gut contents
of individual Acartia tonsa from waters off Southern California,
Mar. Biol., 97, 185–190, https://doi.org/10.1007/BF00391301, 1988. a
Konchina, Y.: Trophic status of the Peruvian Anchovy and Sardine, J.
Ichtyol., 31, 240–252, 1991. a
Lee, R., Hagen, W., and Kattner, G.: Lipid storage in marine zooplankton,
Mar. Ecol. Prog. Ser., 307, 273–306, https://doi.org/10.3354/meps307273,
2006. a
Lehette, P. and Hernández-León, S.: Zooplankton biomass estimation from
digitized images: a comparison between subtropical and Antarctic organisms,
Limnol. Oceanogr.-Meth., 7, 304–308,
https://doi.org/10.4319/lom.2009.7.304, 2009. a
Lischka, S. and Hagen, W.: Seasonal lipid dynamics of the copepods
Pseudocalanus minutus (Calanoida) and Oithona similis
(Cyclopoida) in the Arctic Kongsfjorden (Svalbard), Mar. Biol., 150,
443–454, https://doi.org/10.1007/s00227-006-0359-4, 2007. a
Lischka, S., Bach, L. T., Schulz, K.-G., and Riebesell, U.: Ciliate and mesozooplankton community response to increasing CO2 levels in the Baltic Sea: insights from a large-scale mesocosm experiment, Biogeosciences, 14, 447–466, https://doi.org/10.5194/bg-14-447-2017, 2017. a
Lischka, S., Ayón, P., Pinedo Arteaga, E. L., Schukat, A.,
Taucher, J., Kiko, R., Hauss, H., Dorschner, S., Hagen, W., and
Segura-Noguera, M.: Response of zooplankton community succession and
trophic links to simulated upwelling conditions during a mesocosm experiment
in the coastal upwelling off Callao Bay (Peru) in austral summer 2017 (KOSMOS
2017 Peru mesocosm study), https://doi.org/10.1594/PANGAEA.947833, 2022. a, b, c
Löscher, C. R., Bange, H. W., Schmitz, R. A., Callbeck, C. M., Engel, A., Hauss, H., Kanzow, T., Kiko, R., Lavik, G., Loginova, A., Melzner, F., Meyer, J., Neulinger, S. C., Pahlow, M., Riebesell, U., Schunck, H., Thomsen, S., and Wagner, H.: Water column biogeochemistry of oxygen minimum zones in the eastern tropical North Atlantic and eastern tropical South Pacific oceans, Biogeosciences, 13, 3585–3606, https://doi.org/10.5194/bg-13-3585-2016, 2016. a
Mackas, D. and Bohrer, R.: Fluorescence analysis of zooplankton gut contents
and an investigation of diel feeding patterns, J. Exp. Mar. Biol. Ecol., 25, 77–85, https://doi.org/10.1016/0022-0981(76)90077-0,
1976. a, b, c
Marcus, N. H., Richmond, C., Sedlacek, C., Miller, G. A., and Oppert, C.:
Impact of hypoxia on the survival, egg production and population dynamics of
Acartia tonsa Dana, J. Exp. Mar. Biol.
Ecol., 301, 111–128, https://doi.org/10.1016/j.jembe.2003.09.016, 2004. a
Massing, J. C., Schukat, A., Auel, H., Auch, D., Kittu, L., Pinedo Arteaga,
E. L., Correa Acosta, J., and Hagen, W.: Toward a Solution of the “Peruvian
Puzzle”: Pelagic Food-Web Structure and Trophic Interactions in the
Northern Humboldt Current Upwelling System Off Peru, Front. Mar. Sci., 8, 2296–7745, https://doi.org/10.3389/fmars.2021.759603, 2022. a
Minagawa, M. and Wada, E.: Stepwise enrichment of δ15N
along food chains: Further evidence and the relation between
δ15N and animal age, Geochim. Cosmochim. Ac., 48, 1135–1140, https://doi.org/10.1016/0016-7037(84)90204-7, 1984. a
Minas, H. J., Minas, M., and Packard, T. T.: Productivity in upwelling areas
deduced from hydrographic and chemical fields, Limnol. Oceanogr.,
31, 1182–1206, https://doi.org/10.4319/lo.1986.31.6.1182, 1986. a
Mollier-Vogel, E., Ryabenko, E., Martinez, P., Wallace, D., Altabet, M. A., and
Schneider, R.: Nitrogen isotope gradients off Peru and Ecuador related to
upwelling, productivity, nutrient uptake and oxygen deficiency, Deep Sea
Res. Pt. I, 70, 14–25,
https://doi.org/10.1016/j.dsr.2012.06.003, 2012. a
Peters, J., Dutz, J., and Hagen, W.: Role of essential fatty acids on the
reproductive success of the copepod Temora longicornis in the North
Sea, Mar. Ecol. Prog. Ser., 341, 153–163, 2007. a
Peterson, B. J. and Fry, B.: Stable isotopes in ecosystem studies, Annu.
Rev. Ecol. Syst., 18, 293–320,
https://doi.org/10.1146/annurev.es.18.110187.001453, 1987. a
Picheral, M., Colin, S., and Irisson, J.-O.: EcoTaxa, a tool for the taxonomic
classification of images, http://ecotaxa.obs-vlfr.fr (last access: 25 May 2021), 2017. a
R Core Team: R: A Language and Environment for Statistical Computing, R
Foundation for Statistical Computing, Vienna, Austria,
ISBN 3-900051-07-0, 2014. a
Rau, G. H., Ohman, M. D., and Pierrot-Bults, A.: Linking nitrogen dynamics to
climate variability off central California: a 51 year record based on
δ15N/δ14N in CalCOFI
zooplankton, Deep Sea Res. Pt. II, 50,
2431–2447, https://doi.org/10.1016/S0967-0645(03)00128-0, 2003. a
Richmond, C., Marcus, N. H., Sedlacek, C., Miller, G. A., and Oppert, C.:
Hypoxia and seasonal temperature: Short-term effects and long-term
implications for Acartia tonsa dana, J. Exp. Mar. Biol. Ecol., 328, 177–196, https://doi.org/10.1016/j.jembe.2005.07.004, 2006. a, b
Ryther, J. H.: Photosynthesis and Fish Production in the Sea, Science, 166,
72–76, https://doi.org/10.1126/science.166.3901.72, 1969. a
Sandel, V., Kiko, R., Brandt, P., Dengler, M., Stemmann, L., Vandromme, P.,
Sommer, U., and Hauss, H.: Nitrogen Fuelling of the Pelagic Food Web of the
Tropical Atlantic, PLOS ONE, 10, 1–19, https://doi.org/10.1371/journal.pone.0131258,
2015. a
Schmidtko, S., Stramma, L., and Visbeck, M.: Decline in global oceanic oxygen
content during the past five decades, Nature, 542, 335–339,
https://doi.org/10.1038/nature21399, 2017. a, b
Schröder, S.-M., Kiko, R., and Koch, R.: MorphoCluster: Efficient
Annotation of Plankton Images by Clustering, Sensors, 20, 3060,
https://doi.org/10.3390/s20113060, 2020. a
Schukat, A., Auel, H., Teuber, L., Lahajnar, N., and Hagen, W.: Complex trophic
interactions of calanoid copepods in the Benguela upwelling system, J.
Sea Res., 85, 186–196, https://doi.org/10.1016/j.seares.2013.04.018,
2014. a
Schwartzlose, R. A., Alheit, J., Bakun, A., Baumgartner, T. R., Cloete, R.,
Crawford, R. J., Fletcher, W. J., Green-Ruiz, Y., Hagen, E., Kawasaki, T.,
Lluch-Belda, D., Lluch-Cota, S. E., MacCall, A. D., Matsuura, Y.,
Nevárez-Martínez, M. O., Parrish, R. H., Roy, C., Serra, R.,
Shust, K. V., Ward, M. N., and Zuzunaga, J. Z.: Worldwide large-scale
fluctuations of sardine and anchovy populations, S. Af. J. Mar. Sci., 21, 289–347, https://doi.org/10.2989/025776199784125962, 1999. a
Segura-Noguera, M., Blasco, D., and Fortuño, J.-M.: An improved
energy-dispersive X-ray microanalysis method for analyzing simultaneously
carbon, nitrogen, oxygen, phosphorus, sulfur, and other cation and anion
concentrations in single natural marine microplankton cells, Limnol. Oceanogr.-Meth., 10, 666–680, https://doi.org/10.4319/lom.2012.10.666, 2012. a
Segura-Noguera, M., Blasco, D., and Fortuño, J.-M.: Taxonomic and
Environmental Variability in the Elemental Composition and Stoichiometry of
Individual Dinoflagellate and Diatom Cells from the NW Mediterranean Sea,
PLOS ONE, 11, e0154050,
https://doi.org/10.1371/journal.pone.0154050, 2016. a, b
Sharp, J. H.: Improved analysis for “particulate” organic carbon and
nitrogen from seawater, Limnol. Oceanogr., 19, 984–989, https://doi.org/10.4319/lo.1974.19.6.0984, 1974. a
Sigman, D. M., Altabet, M. A., Francois, R., McCorkle, D. C., and Gaillard,
J.-F.: The isotopic composition of diatom-bound nitrogen in Southern Ocean
sediments, Paleoceanography, 14, 118–134, https://doi.org/10.1029/1998PA900018, 1999. a
Stramma, L., Johnson, G. C., Sprintall, J., and Mohrholz, V.: Expanding
Oxygen-Minimum Zones in the Tropical Oceans, Science, 320, 655–658,
https://doi.org/10.1126/science.1153847, 2008.
a, b
Taucher, J., Bach, L. T., Boxhammer, T., Nauendorf, A., Achterberg, E. P.,
Algueró-Muñiz, M., Arístegui, J., Czerny, J., Esposito, M.,
Guan, W., Haunost, M., Horn, H. G., Ludwig, A., Meyer, J., Spisla, C., Sswat,
M., Stange, P., and Riebesell, U.: Influence of Ocean Acidification and Deep
Water Upwelling on Oligotrophic Plankton Communities in the Subtropical North
Atlantic: Insights from an In situ Mesocosm Study, Front. Mar.
Sci., 4, 2296–7745, https://doi.org/10.3389/fmars.2017.00085, 2017. a
Teuber, L., Schukat, A., Hagen, W., and Auel, H.: Trophic interactions and
life strategies of epi- to bathypelagic calanoid copepods in the tropical
Atlantic Ocean, J. Plankton Res., 36, 1109–1123,
https://doi.org/10.1093/plankt/fbu030, 2014. a
van Guelpen, L., Markle, D. F., and Duggan, D. J.: An evaluation of accuracy,
precision, and speed of several zooplankton subsampling techniques, ICES
J. Mar. Sci., 40, 226–236, https://doi.org/10.1093/icesjms/40.3.226,
1982. a
Walsh, J. J.: A carbon budget for overfishing off Peru, Nature, 290,
300–304, https://doi.org/10.1038/290300a0, 1981. a
Ward, B. B., Kilpatrick, K. A., Renger, E. H., and Eppley, R. W.: Biological
nitrogen cycling in the nitracline, Limnol. Oceanogr., 34, 493–513,
https://doi.org/10.4319/lo.1989.34.3.0493, 1989. a
Welschmeyer, N. A.: Fluorometric analysis of chlorophyll a in the presence of
chlorophyll b and pheopigments, Limnol. Oceanogr., 39, 1985–1992,
https://doi.org/10.4319/lo.1994.39.8.1985, 1994. a
Wiebe, P. and Holland, W.: Plankton patchiness: Effects on repeated net tows,
Limnol. Oceanogr., 13, 315–321, https://doi.org/10.4319/lo.1968.13.2.0315,
1968. a
Short summary
Ocean upwelling regions are highly productive. With ocean warming, severe changes in upwelling frequency and/or intensity and expansion of accompanying oxygen minimum zones are projected. In a field experiment off Peru, we investigated how different upwelling intensities affect the pelagic food web and found failed reproduction of dominant zooplankton. The changes projected could severely impact the reproductive success of zooplankton communities and the pelagic food web in upwelling regions.
Ocean upwelling regions are highly productive. With ocean warming, severe changes in upwelling...
Altmetrics
Final-revised paper
Preprint