Prof. Dr. Rainer Kiko

Group leader Plankton Biogeochemistry and Dynamics

Heisenberg Professor and Make Our Planet Great Again Laureat


Research Unit 2: Marine Biogeochemistry


E-Mail: rkiko(at)

Address: Prof. Dr. Rainer Kiko, Wischhofstr. 1-3, 24148 Kiel, Deutschland


Group members

Dr. Joelle Habib (Postdoc at the Laboratoire d'Océanographie de Villefranche)

Dr. Xiangyu Weng (Machine Vision)

Simon-Martin Schröder (Computer scientist)

Claudeilton "Claus" Santana (PhD student)



Research topics

Global change impacts the distribution and activity of marine life at local to basin-wide scales, with major consequences for oceanic oxygen dynamics, nutrient cycles and the transfer of carbon dioxid from the atmosphere to the deep sea. I use augmented image observations that integrate autonomous camera and environmental sensor systems with state-of-the-art artificial intelligence solutions and ecophysiological approaches to yield a novel image of zooplankton and detrital particle (Z&P) dynamics in a changing ocean. Zooplankton organisms play an essential role in these processes as they feed on all kinds of particulate matter, respire oxygen and release faecal matter. Detrital particles (e.g. faeces and decaying phytoplankton aggregates) passively sink through the water column and represent a major export vector of carbon from the sunlit surface ocean to the deep sea. Most of this so-called „marine snowfall” is consumed by microbes and specialised zooplankton, resulting in the attenuation of passive flux with depth. Diel vertical migrations of zooplankton and nekton from the epipelagic at 0 – 200 m depth to the mesopelagic at 200 – 1000 m depth induce an active net downward transport of carbon as zooplankton feed at the surface at nighttime and migrate to depth at daytime, where they continue to respire, excrete and generate faecal pellets. These processes take place in a very dynamic and changing physical environment, requiring the mentioned augmented observation tools to observe them. Building the capacities of the next generation of students to apply these tools for improved monitoring, protection and management of marine resources is another aspect of my work.


Imaging Marine life in an Ocean of Change (IOChange) (gefördert durch das Heisenberg Programm der Deutschen Forschungsgemeinschaft)


Tropical Atlantic Deoxygenation: gateway dynamics, feedback mechanisms and ecosystem impacts (Make Our Planet Great Again project at the Laboratoire d'Oceanographie de Villefranche-sur-mer, France)


I/ITAPINA - Imagine Imaging the Atlantic - A pelagic imaging network approach


The role of zooplankton in N-, P- and C- cycling and for the oxygen budget of tropical OMZs


Combining deep learning, in situ imaging and citizen science to resolve the distribution of giant protists in major upwelling regions


Zooplankton and Nekton: Gatekeepers of the Biological Pump (Frontiers Research Topic)


Zooplankton and Nekton: Gatekeepers of the Biological Pump - Volume 2 (Frontiers Research Topic)



ZoOMin at TENATSO: “Zooplankton of the Oxygen Minimum Zone at the Tropical Eastern Atlantic Time-Series Observatory” 


CUSCO - Coastal Upwelling System in a Changing Ocean: Work Package 5: Particle size distribution, export fluxes and zooplankton-particle interactions


MicroZooImager: An integrated optical system for microzooplankton analysis



Publications in review

8) Richards C, et al (submitted to PNAS) Global environmental challenges require interdisciplinary science programs: Lessons learned from the Franco-German MOPGA initiative.

7) Dugenne M, Corrales-Ugalde M, Luo J, Stemmann L, Irisson J-O, Lombard F, O’Brien T, Stock C, Kiko R (preprint) Key link between iron and the size structure of three main mesoplanktonic groups (Crustaceans, Rhizarians, and colonial N2-fixers) in the Global Ocean. in preparation for submission.

6) Siegel D, Burd A, Estapa M, Fields E, Johnson L, Romanelli E, Brzezinski M, Buesseler K, Clevenger S, Cetinic I, Drago L, Durkin C, Kiko R, Kramer SJ, Maas A, Omand M, Passow U, Steinberg D (preprint) Dynamics of Aggregates and Sinking Carbon Fluxes in a Turbulent Ocean. Submitted to Science advances.

5) Brandt P, Körner M, Moum JN,  Rocha M, Subramaniam A, Czeschel R, Krahmann G, Dengler M, Kiko R (in review, Nature Geosciences) Synchronous drivers of equatorial Atlantic productivity

4) Flintrop CM, Ahmerkamp S, Moradi N, Klawonn I, März J, Hörstmann C, Kiko R, Khalili A, Grossart H-P, Álvarez-Salgado XA, Aristegui J, Iversen MH (preprint) Rethinking sinking: Imaging the flow fields of natural marine aggregates to derive sinking velocity. in preparation for submission.

3) Zhang Z, Xu A, Hathorne E, Gutjahr M, Browning TJ, Gosnell K, Liu T, Steiner Z, Kiko R, Yuan Z, Liu H, Achterberg EP, Frank M (submitted to Nature Communications) Substantial trace metal input from the 2022 Hunga Tonga-Hunga Ha’apai eruption into the South Pacific Gyre

2) Alberto Baudena, Rémi Laxenaire, Camille Catalano, Hervé Claustre, Artemis Ioannou, Edouard Leymarie, Marc Picheral, Antoine Poteau, Sabrina Speich, Lars Stemmann, Rainer Kiko (in preparation for resubmission) A Lagrangian perspective on the carbon and oxygen budget of an oceanic eddy.

1) Yawouvi Dodji Soviadan, Mathilde Dugenne, Laetitia Drago, Tristan Biard, Emilia Trudnowska, Fabien Lombard, Jean-Baptiste Romagnan, Jean-Louis Jamet, Rainer Kiko, Gabriel Gorsky, Lars Stemmann (in preparation for resubmission) Complete zooplankton size spectra re-constructed from in situ imaging and Multinet data in the global ocean.


Peer reviewed articles (accepted or published):

75) Shreya Mehta, Rainer Kiko, Helena Hauss, Narendra Ojha, Arvind Singh (accepted, Global Biogeochemical Cycles) Dust deposition and phosphate influx create zonal and meridional biogeochemical gradient in the Trichodesmium abundance.

74) Diogoul N, Brehmer P, Kiko R, Perrot Y, Lebourges-Dhaussy A, Rodrigues E, Thiam A, Mouget A, El Ayoubi S, Sarré A (accepted, PLOSone) Estimating the Copepod Biomass in the North West African upwelling system using a bi-frequency acoustic approach

73) Dugenne M, Corrales-Ugalde M, Luo J, Kiko R, O’Brien T, Irisson J-O, Lombard F, Stemmann L, Stock C, Anderson CR, Babin M, Bhairy N, Bonnet S, Carlotti F, Cornils A, Crockford ET, Daniel P, Desnos C, Drago L, Elineau A, Fischer A, Grandrémy N, Grondin P-L, Guidi L, Guieu C, Hauss H, Hayashi K, Huggett JA, Jalabert L, Karp-Boss L, Kenitz KM, Kudela RM, Lescot M, Marec C, McDonnell A, Mériguet Z, Niehoff B, Noyon M, Panaïotis T, Peacock E, Picheral M, Riquier E, Roesler C, Romagnan J-B, Sosik HM, Spencer G, Taucher J, Tilliette C, Vilain M (accepted) First release of the Pelagic Size Structure database: Global datasets of marine size spectra obtained from plankton imaging devices. Earth System Science Data Discussions:1–41.

72) Manon Laget, Laetitia Drago, Thelma Panaïotis, Rainer Kiko, Lars Stemmann, Andreas Rogge, Natalia Llopis-Monferrer, Aude Leynaert, Jean-Olivier Irisson, Tristan Biard (2024) A global census of the significance of giant mesopelagic protists to the biogeochemical cycles of carbon and silicon. Nature Communications., 15:3341.

71) Rainer Kiko, Rubens M. Lopes, Dodji Yavouwi Soviadan, Lars Stemmann (2023) Towards a distributed and operational pelagic imaging network. Ocean and Coastal Research, 71,

70) Rainer Kiko, Daniele Bianchi, Helena Hauss, Morten H Iversen M, Amy Maas (2023) Editorial: Zooplankton and nekton: gatekeepers of the biological pump, volume II. Frontiers in Marine Science 10.

69) Anja Engel, Carolina Cisternas-Novoa, Helena Hauss, Rainer Kiko, Frédéric A.C. Le Moigne (2023) Hypoxia-tolerant zooplankton reduce the biological carbon pump efficiency in the Humboldt current system off Peru. Communications Earth & Environment, 4, 458,

68) Bobby Leigh, Víctor Aramayo, Ursula Mendoza, Federico Velazco, Rainer Kiko, Patricia Ayón, Ernesto Fernández and Michelle Graco (2023) Total Mass Flux in the Northern Humboldt Current System: Rates and Contribution Sources from Central Peru (12° S) Hydrobiology, 2, 521-536.

67) Ana Fernández-Carrera, Rainer Kiko, Helena Hauss, Douglas S. Hamilton, Eric P. Achterberg, Joseph P. Montoya, Marcus Dengler, Peter Brandt, Ajit Subramaniam (2023) Nitrogen fixation rates in the Guinea Dome and the Equatorial upwelling regions in the Atlantic Ocean Biogeochemistry,

66) Wokil Bam, Beat Gasser, Kanchan Maiti, Isabelle Levy, Juan Carlos Miquel, Roberta L. Hansman, Jan Scholten, Ruifang C. Xie, Stefan Sommer, Rainer Kiko, Peter Swarzenski (2023) Particulate Organic Carbon Export in the Oxygen Minimum Zone off the Peruvian Coast

65)  Peter Rubbens, Stephanie Brodie, Tristan Cordier, Diogo Destro Barcellos, Paul Devos, Jose A Fernandes-Salvador, Jennifer I Fincham, Alessandra Gomes, Nils Olav Handegard, Kerry Howell, Cédric Jamet, Kyrre Heldal Kartveit, Hassan Moustahfid, Clea Parcerisas, Dimitris Politikos, Raphaëlle Sauzède, Maria Sokolova, Laura Uusitalo, Laure Van den Bulcke, Aloysius T M van Helmond, Jordan T Watson, Heather Welch, Oscar Beltran-Perez, Samuel Chaffron, David S Greenberg, Bernhard Kühn, Rainer Kiko, Madiop Lo, Rubens M Lopes, Klas Ove Möller, William Michaels, Ahmet Pala, Jean-Baptiste Romagnan, Pia Schuchert, Vahid Seydi, Sebastian Villasante, Ketil Malde, Jean-Olivier Irisson (2023) Machine learning in marine ecology: an overview of techniques and applications. ICES Journal of Marine Science.

64) Thelma Panaoitis, Marcel Babin, Tristan Biard, Francois Carlotti, Laurent Coppola, Lionel Guidi, Helena Hauss, Lee Karp-Boss, Rainer Kiko, Fabien Lombard, Andrew MP McDonnell, Marc Picheral, Andreas Rogge, Anya M. Waite, Lars Stemmann, Jean-Olivier Irisson (2023) Three mesoplanktonic worlds resolved by in situ imaging in the upper 500 m of the global ocean. Global Ecology and Biogeography.

63) Daniel J Clements, Simon Yang, Thomas Weber, Andrew McDonnell, Rainer Kiko, Lars Stemmann, Daniele Bianchi (2023) New estimate of organic carbon export from optical measurements reveals the role of particle size distribution and export horizon. Global Biogeochemical Cycles, 37, e2022GB007633.

62) Alberto Baudena, Rainer Kiko, Isabel Jalon-Rojas, Maria Luiza Pedrotti (2023) Low-density plastic debris dispersion beneath the Mediterranean Sea surface. Environmental Science & Technology, doi: 10.1021/acs.est.2c08873.

61) Anna Denvil-Sommer, Erik T Buitenhuis, Rainer Kiko, Fabien Lombard, Lionel Guidi, Corrine Le Quéré (2023) Testing the reconstruction of modelled particulate organic carbon from surface ecosystem components using PlankTOM12 and Machine Learning. Geoscientific Model Development, doi: 10.5194/gmd-16-2995-2023

60) Patricia Ayón Dejo, Elda Luz Pinedo Arteaga, Anna Schukat, Jan Taucher, Rainer Kiko, Helena Hauss, Sabrina Dorschner, Wilhelm Hagen, Mariona Segura-Noguera, Silke Lischka (2023) Zooplankton community succession and trophic links during a mesocosm experiment in the coastal upwelling off Callao Bay (Peru). Biogeosciences 20, 945-969.

59) Kerrie M. Swadling, Andrew J. Constable, Alexander D Fraser, Robert A Massom, Melanie D Borup, Laura Ghigliotti, Antonia Granata, Letterio Guglielmo, So Kawaguchi, Fraser Kennedy, Rainer Kiko, Philippe Koubbi, Ryosuke Makabe, Andrew Martin, Andrew Mcminn, Masato Moteki, Evgeny A. Pakhomov, Ilka Peeken, Jody R Reimer, Phillip Reid, Ken G Ryan, Marino Vacchi, Patti Virtue, Christine K Weldrick, Pat Wongpan and Simon Wotherspoon (2023) Biological responses to change in Antarctic sea ice habitats. Frontiers in Ecology and Evolution. 10, DOI:10.3389/fevo.2022.1073823.

58) Rainer Kiko, Marc Picheral, David Antoine, Marcel Babin, Leo 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, Gaby 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 MP McDonnell, Daniela Niemeyer, Margaux Noyon, Stephanie H O’Daly, Mark Ohman, Jessica L Pretty, Andreas Rogge, Sarah Searson, Masashi Shibata, Yuji Tanaka, Toste Tanhua, Jan Taucher, Emilia Trudnowska, Jessica S Turner, Anya Waite, Lars Stemmann (2022) A global marine particle size distribution dataset obtained with the Underwater Vision Profiler 5. Earth System Science Data. 14: 4315 – 4337.

57) Frank Melzner, Ulrike Findeisen, Christian Bock, Ulrike Panknin, Rainer Kiko, Claas Hiebenthal, Mark Lenz, Marlene Wall (accepted) Oxygen flux limitation drives heat wave vulnerability of a coastal keystone predator. Nature Climate Change.

56) Kelsey M. Bisson, Rainer Kiko, David A. Siegel, Lionel Guidi, Marc Picheral, Emmanuel Boss, B.B. Cael (2022) Sampling uncertainties of particle size distributions and derived fluxes. Limnology & Oceanography:Methods

55) Lars Schmarje, Vasco Grossmann, Claudius Zelenka C, Sabine Dippel, Rainer Kiko, Mariusz Oszust, Matti Pastell, Jenny Stracke, Anna Valros, Nina Volkmann, Reinhard Koch (2022) Is one annotation enough? A data-centric image classification benchmark for noisy and ambiguous label estimation. Accepted at NeurIPS 2022, preprint available at

54) Janina Büscher, Armin Form, Max Wisshack, Rainer Kiko, Ulf Riebesell (2022) Cold-water coral ecosystems under future ocean change – live coral performance versus framework dissolution and bioerosion. Limnology & Oceanography.

53) Lars Schmarje, Monty Santarossa, Simon-Martin Schröder, Claudius Zelenka, Rainer Kiko, Jenny Stracke, Nina Volkmann, Reinhard Koch R (2022) A Data-Centric Approach for Improving Ambiguous Labels with Combined Semi-supervised Classification and Clustering. In: Computer Vision – ECCV 2022. Lecture Notes in Computer Science, Avidan S, Brostow G, Cissé M, Farinella GM, Hassner T (eds) Springer Nature Switzerland, Cham, p 363–380.

52) Laetitia Drago, Thelma Panaoïtis, Jean-Olivier Irisson, Marcel Babin, Tristan Biard, François Carlotti, Laurent Coppola, Lionel Guidi, Helena Hauss, Lee Karp-Boss, Fabien Lombard, Andrew McDonnell, Marc Picheral, Andreas Rogge, Anya M. Waite, Lars Stemmann, Rainer Kiko (2022) Global distribution of zooplankton biomass estimated by in-situ imaging and machine learning. Frontiers in Marine Science 9.

51) Natalia Belkin, Tamar Guy-Haim, Maxim Rubin-Blum, Rainer Kiko, Arseniy R. Morov, Guy Sisma-Ventura, Tal Ozer, Ayah Lazar, Isaac Gertman, Jacob Silverman, Barak Herut, Eyal Rahav (2022) Influence of cyclonic and anti-cyclonic eddies on plankton biomass, activity and diversity in the ultra-oligotrophic southeastern Mediterranean Sea. Ocean Science, 18, 693–715,

50) Timm Schoening, Jennifer M. Durden, Claas Faber, Janine Felden, Karl Heger, Henk-Jan T. Hoving, Rainer Kiko, Kevin Köser, Christopher Krämmer, Tom Kwasnitschka, Klas Ove Möller, David Nakath, Andrea Naß, Tim W. Nattkemper, Autun Purser, Martin Zurowietz (2022) Making marine image data FAIR. Scientific data. 9, 414.

49) Daniel J Clements, Simon Yang, Thomas Weber, Andrew McDonnell, Rainer Kiko, Lars Stemmann, Daniele Bianchi (2022) Constraining the particle size distribution of large marine particles in the global ocean with in situ optical observations and supervised learning. Global Biogeochemical Cycles, doi: 10.1029/2021GB007276.

48) Martin Schröder, Rainer Kiko (2022) Assessing Representation Learning and Clustering Algorithms for computer-assisted Image Annotation—Simulating and Benchmarking MorphoCluster. Sensors. 22:2775.

47) Anja Engel, Rainer Kiko, Marcus Dengler (2022) Organic matter supply and utilization in oxygen minimum zones. Annual Review of Marine Science. 14:355-378.

46) Jonathan Sherman, Ajit Subramaniam, Maxim Y. Gorbunov, Ana Fernandez-Carrera, Rainer Kiko, Peter Brandt, Paul G Falkowski (2022) The photophysiological response of nitrogen-limited phytoplankton to episodic nitrogen supply associated with tropical instability waves in the Equatorial Atlantic Frontiers in Marine Science, 8. Doi:10.3389/fmars.2021.814663.

45) Yawouvi Dodji Soviadan, Fabio Benedetti, Manoela Brandao, Sakina Dorothe Ayata, Jean Olivier Irisson, Jean Louis Jamet, Rainer Kiko, Fabien Lombard, Kissao Gnandi, Lars Stemmann (2022) Patterns of mesozooplankton community composition and vertical fluxes in the global ocean. Progress in Oceanography 200:102717.

44) Clarissa Karthäuser, Soeren Ahmerkamp, Hannah K Marchant, Laura A Bristow, Helena Hauss, Morten H Iversen, Rainer Kiko, Jöran März, Gaute Lavik, Marcel MM Kuypers (2021) Small sinking particles control anammox rates in the Peruvian oxygen minimum zone. Nature communications, 12:3235.  doi:10.1038/s41467-021-23340-4

43) Peter Brandt, Johannes Hahn, Sunke Schmidtko, Franz Philip Tuchen, Robert Kopte, Rainer Kiko, Bernard Bourlès, Rena Czeschel, Marcus Dengler (2021) Atlantic Equatorial Undercurrent intensification counteracts warming-induced deoxygenation. Nature Geoscience, 14:278-282. doi:10.1038/s41561-021-00716-1

42) Lars Schmarje, Johannes Brünger, Monty Santarossa, Simon-Martin Schröder, Rainer Kiko, Reinhard Koch (2021) Fuzzy Overclustering: Semi-Supervised Classification of Fuzzy Labels with Overclustering and Inverse Cross-Entropy. Sensors, 21:6661. doi:10.3390/s21196661

41) 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, Hela Mehrtens (2021) Climate-Biogeochemistry Interactions in the Tropical Ocean: Data Collection and Legacy. Frontiers in Marine Science, 8:1270. doi:10.3389/fmars.2021.723304

40) Mariana Hill Cruz, Iris Kriest, José Jonss, Rainer Kiko, Helena Hauss, Andreas Oschlies (2021) Zooplankton mortality effects on the plankton community of the Northern Humboldt Current System: Sensitivity of a regional biogeochemical model.  Biogeosciences,

39) Jahan Vajedsamiei, Frank Melzner, Michael Raatz, Rainer Kiko, Maral Khosravi, Christian Pansch (2021) Simultaneous recording of filtration and respiration in marine organisms in response to short‐term environmental variability.  Limnology and Oceanography Methods

38) Rainer Kiko, Peter Brandt, Svenja Christiansen, Jannik Fausmann, Iris Kriest, Elizandro Rodrigues, Florian Schütte, Helena Hauss (2020) Zooplankton-Mediated Fluxes in the Eastern Tropical North Atlantic.  Frontiers in Marine Science 7:358

37) Simon-Martin Schröder, Rainer Kiko, Reinhard Koch (2020) MorphoCluster: Efficient Annotation of Plankton images by Clustering.  Sensors 20:3060

36) Henk-Jan Hoving, Phillip Neitzel, Helena Hauss, Svenja Christiansen, Rainer Kiko, Bruce H. Robison, Pericles Silva, Arne Körtzinger (2020) In situ observations show vertical community structure of pelagic fauna in the eastern tropical North Atlantic off Cape Verde.  Scientific reports 10:21798

35) Sarah L.C. Giering, Emma L. Cavan, Sünnje L. Basedow, Nathan Briggs, Adrian B. Burd, Louise J. Darroch, Lionel Guidi, Jena-Olivier Irisson, Morten H. Iversen, Rainer Kiko, Dhugal Lindsay, Catarina R. Marcolin, Andrew M.P. McDonnell, Klas O. Möller, Uta Passow, Sandy Thomalla, Thomas W. Trull, Anya M. Waite (2020) Sinking Organic Particles in the Ocean—Flux Estimates From in situ Optical Devices. Frontiers in Marine Science 6:834

34) Rainer Kiko & Helena Hauss (2019) On the estimation of zooplankton-mediated active fluxes in oxygen minimum zone regions. Frontiers in Marine Science 6:741.

33) Fabien Lombard, Emmanuel Boss, Anya M Waite, Julia Uitz, Lars Stemmann, Heidi M Sosik, Jan Schulz, Jean-Baptiste Romagnan, Marc Picheral, Jay Pearlman, Mark D Ohman, Barbara Niehoff, Klas Ove Möller, Patricia Miloslavich, Ana Lara-Lopez, Raphael M Kudela, Rubens Mendez Lopes, Lee Karp-Boss, Rainer Kiko, Julles S Jaffe, Morten Hvitfeldt Iversen, Jean-Olivier Irisson, Helena Hauss, Lionel Guidi, Gaby Gorsky, Sarah Lou Carolin Giering, Peter Gaube, Scott Gallager, George Dubelaar, Robert K Cowen, François Carlotti, Christian Briseño-Avena, Léo Berline, Kelly J Benoit-Bird, Nicholas J Bax, Sonia Dawn Batten, Sakina-Dorothée Ayata, Ward Appeltans (2019) Globally consistent quantitative observations of planktonic ecosystems. Frontiers in Marine Science 6:196.

32) Soeren Thomsen, Johannes Karstensen, Rainer Kiko, Gerd Krahmann, Marcus Dengler, Anja Engel (2019) Remote and local drivers of oxygen and nitrate variability in the shallow oxygen minimum zone off Mauritania in June 2014. Biogeosciences 16:979-998.

31) Daniele Bianchi, Thomas S Weber, Rainer Kiko, Curtis Deutsch (2018) Global niche of marine anaerobic metabolisms expanded by particle microenvironments. Nature Geoscience, doi:10.1038/s41561-018-0081-0.

30) Svenja Christiansen, Henk-Jan Hoving, Florian Schütte, Helena Hauss, Johannes Karstensen, Arne Körtzinger, Simon-Martin Schröder, Lars Stemmann, Bernd Christiansen, Marc Picheral, Peter Brandt, Bruce Robison, Reinhard Koch, Rainer Kiko (2018) Particulate matter flux interception in oceanic mesoscale eddies by the polychaete Poeobius sp. Limnology and Oceanography, doi: 10.1002/lno.10926.

29) Simon-Martin Schröder, Rainer Kiko, Jean-Olivier Irisson, Reinhard Koch (2018) Low-Shot learning of plankton categories. In: Brox T., Bruhn A., Fritz M. (eds) Pattern Recognition. GCPR 2018. Lecture Notes in Computer Science, vol 11269. Springer, Cham.

28) Henk-Jan Hoving, Svenja Christiansen, Eduard Fabrizius, Helena Hauss, Rainer Kiko, Peter Linke, Philip Neitzel, Uwe Piatkowski, Arne Körtzinger (in review, 2018) The Pelagic In situ Observation System (PELAGIOS) to reveal biodiversity, behavior and ecology of elusive oceanic fauna. Ocean Sci. Discuss.,

27) Rainer Kiko, Arne Biastoch, Peter Brandt, Sophie Cravatte, Helena Hauss, Rebecca Hummels, Iris Kriest, Frederic Marin, Andrew M. P. McDonnell, Andreas Oschlies, Marc Picheral, Franziska U. Schwarzkopf, Andreas M. Thurnherr, Lars Stemmann (2017) Biological and physical influences on marine snowfall at the equator. Nature Geoscience, DOI:10.1038/ngeo3042.

26) Arvind Singh, Lennart T Bach, Tim Fischer, Helena Hauss, Rainer Kiko, Allanah J Paul, Paul Stange, Pieter Vandromme, Ulf Riebesell (2017) Niche construction by non-diazotrophs for N2 fixers in the eastern tropical North Atlantic Ocean. Geophysical Research Letters, DOI:10.1002/2017GL074218.

25) Rainer Kiko, Helena Hauss, Friedrich Buchholz, Frank Melzner (2016) Ammonium excretion and oxygen respiration of tropical copepods and euphausiids exposed to oxygen minimum zone conditions. Biogeosciences, 13:2241-2255.

24) Tristan Biard, Lars Stemmann, Marc Picheral, Nicolas Mayot, Pieter Vandromme, Helena Hauss, Gabriel Gorsky, Lionel Guidi, Rainer Kiko, Fabrice Not (2016) In situ imaging reveals the biomass of giant protists in the global ocean. Nature, 532:504-507.

23) Helena Hauss, Svenja Christiansen, Florian Schütte, Rainer Kiko,  Miryan Edvam Lima, Elizandro Rodrigues, Johannes Karstensen, Carolin Löscher, Arne Körtzinger,  Björn Fiedler (2016) Dead zone or oasis in the open ocean? Zooplankton distribution and migration in low-oxygen modewater eddies. Biogeosciences, 13:1977-1989.

22) Judith Meyer, Carolin. R. Löscher, Sven. C. Neulinger, Anna. F. Reichel, Alexandra N. Loginova, Corinna Borchard, Ruth. A. Schmitz, Helena Hauss, Rainer Kiko, Ulf Riebesell (2016) Changing nutrient stoichiometry affects phytoplankton production, DOP build up and dinitrogen fixation – a mesocosm experiment in the eastern tropical North Atlantic. Biogeosciences, 13, 781-794.

21) Carolin R. Löscher, Hermann W. Bange, Ruth A. Schmitz, Cameron M. Callbeck, Anja Engel, Helena Hauss, Thorsten Kanzow, Rainer Kiko, Gaute Lavik, Alexandra N. Loginova, Frank Melzner, Sven C. Neulinger, Markus Pahlow, Ulf Riebesell, Harald Schunck, Sören Thomsen, Hannes Wagner (2016) Water column biogeochemistry of oxygen minimum zones in the eastern tropical North Atlantic and eastern tropical South Pacific Oceans. Biogeosciences, 13:3585-3606.

20) Björn Fiedler, Damian Grundle, Florian Schütte, Johannes Karstensen, Carolin R. Löscher, Helena Hauss, Hannes Wagner, Alexandra Loginova, Rainer Kiko, Pericles Silva, Arne Körtzinger (2016) Oxygen Utilization and Downward Carbon Flux in an Oxygen-Depleted Eddy in the Eastern Tropical North Atlantic. Biogeosciences, 15:5633-5647.

19) Rainer Kiko, Stefan Kern, Maike Kramer (2016) Colonization of newly formin Arctic sea ice by meiofauna - a case study for the future Artic? Polar Biology, doi:10.1007/s00300-016-2052-5.

18) Rainer Kiko, Helena Hauss, Marcus Dengler, Stefan Sommer, Frank Melzner (2015) The squat lobster Pleuroncodes monodon tolerates anoxic “dead zone” conditions off Peru. Marine Biology, 162:1913-1921.

17) Vera Sandel, Rainer Kiko, Peter Brandt, Marcus Dengler, Lars Stemmann, Pieter Vandromme, Ulrich Sommer, Helena Hauss (2015) Nitrogen Fuelling of the Pelagic Food Web of the Tropical Atlantic. PLoS ONE, 10(6): e0131258.

16) Tim Kalvelage, Gaute Lavik, Marlene M. Jensen, Niels Peter Revsbech, Carolin Löscher, Harald Schunck, Dhwani K. Desai, Helena Hauss, Rainer Kiko, Moritz Holtappels, Julie LaRoche, Ruth A. Schmitz, Michelle I. Graco, Marcel M. M. Kuypers (2015) Aerobic Microbial Respiration In Oceanic Oxygen Minimum Zones. PLoS ONE, 10(7): e0133526.

15) Anja Engel, Corinna Borchard, Alexandra N. Loginova, Judith Meyer, Helena Hauss, Rainer Kiko (2015) Effects of varied nitrate and phosphate supply on polysaccharidic and proteinaceous gel particles production during tropical phytoplankton bloom experiments. Biogeosciences, 12:5647–5665.

14) Alexandra N. Loginova, Corinna Borchard, Judith Meyer, Helena Hauss, Rainer Kiko, Anja Engel (2015) Effects of nitrate and phosphate supply on chromophoric and fluorescent dissolved organic matter in the Eastern Tropical North Atlantic: a mesocosm study. Biogeosciences, 12:6897–6914.

13) Lena Teuber, Rainer Kiko, Francois Seguin, Holger Auel (2013) Respiration rates of tropical Atlantic copepods in relation to the oxygen minimum zone. J Exp Mar Biol Ecol, 448:28-36.

12) Warren R Francis, Lynne M Christianson, Rainer Kiko, Meghan L Powers, Nathan C Shaner, Steven H D Haddock (2013) A comparison across non-model animals suggests an optimal sequencing depth for de novo transcriptome assembly BMC Genomics, 2013, 14:167

11) Kramer M, Swadling KM, Meiners K, Kiko R, Scheltz A, Nicolaus M, Werner I (2011) Antarctic sympagic meiofauna in winter: comparing diversity, abundance and biomass between the western Weddell Sea and the southern Indian Ocean. Deep-Sea Research Part II: Topical Studies in Oceanography, 58:1062-1074.

10) Kramer M, Kiko R (2011) Brackish meltponds on Arctic sea ice – a new habitat for marine metazoans. Polar Biology, 34:603-608.

9) Hu M, Tseng Y-C, Stumpp M, Gutowska MA, Kiko R, Lucassen M, Melzner F (2011) Elevated seawater pCO2 differentially affects branchial acid-base transporters over the course of development in the cephalopod Sepia officinalis. American Journal of Physiology - Regulatory, Integrative and Comparative Physiology, DOI: 10.​1152/​ajpregu.​00653.​2010

8) Fehsenfeld, S., Kiko, R., Appelhans, Y., Towle, D. W., Zimmer, M. and Melzner, F. (2011) Effects of elevated seawater pCO2 on gene expression patterns in the gills of the green crab, Carcinus maenas. BMC Genomics, 12 (1). p. 488. DOI: 10.1186/1471-2164-12-488.

7) Kiko R (2010) Acquisition of freeze protection in a sea-ice crustacean through horizontal gene transfer? Polar Biology, 33:543-556.

6) Siebert S, Anton-Erxleben F, Kiko R, Kramer M (2009) Sympagohydra tuuli (Cnidaria, Hydrozoa) – first report form sea ice of the central Arctic Ocean and insights into histology, reproduction and locomotion. Marine biology, 156:541-554.

5) Kiko R, Werner I, Wittmann A (2009) Osmotic and ionic regulation in response to salinity variations and cold resistance in the Arctic under-ice amphipod Apherusa glacialis. Polar Biology, 32:393-398.

4) Kiko R, Kramer M, Spindler M, Wägele H (2008) Tergipes antarcticus (Gastropoda, Nudibranchia): distribution, life cycle, morphology, anatomy and adaptation of the first mollusc known to live in Antarctic sea ice. Polar Biology, 31:1383–1395.

3) Kiko R, Michels J, Mizdalski E, Schnack-Schiel SB, Werner I (2008) Living conditions, abundance and composition of the metazoan fauna in surface and sub-ice layers in pack ice of the western Weddell Sea during late spring. Deep Sea Research Part II, 55:1000–1014.

2) Augustin R, Franke A, Khalturin K, Kiko R, Hemmrich G, Bosch TCG (2006) Dickkopf related genes are components of the positional value gradient in Hydra. Developmental Biology, 296:62–70.

1) Werner I, Auel H, Kiko R (2004) Occurrence of Anonyx sarsi (Amphipoda: Lysianassoidea) below Arctic pack ice: an example for cryo-benthic coupling? Polar Biology 27:474–481.