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Dr. Raphael Morard



Micropaleontology - Paleoceanography


+49 421 218-65979



UFT / 1560

Raphael Morard

Working Group: Micropaleontology – Paleoceanography

Research interests

  • Genetic and morphological diversity in Foraminifera
  • Evolution of microplankton
  • Integration of genetic diversity in paleoceanography

Current research

  • Databases development for permanent genetic resources - PFR²
  • Global pattern of cryptic diversity in planktonic foraminifera
  • Metagenomic analyses


Research positions

Since Jan. 2013Post-Doctoral researcher at MARUM. Molecular genetics of planktonic foraminifera.
2011 -2012Post-Doc at Biological station of Roscoff (CNRS, France). Global genetic diversity in planktonic foraminifera within the TARA Ocean project.
2007 – 2010PhD Student at University Claude Bernard Lyon I. Toward a better understanding of pelagic fossil record: Morphological diversity, biogeography and ecology of cryptic species of planktonic foraminifera.


2007 - 2010PhD degree in Micropaleontology. University Claude Bernard Lyon I, France
2005 - 2007Master degree in Paleontology, Sedimentology and Paleoenvironment. University Claude Bernard Lyon I
2002 - 2005Bachelor degree in Earth sciences. University Claude Bernard Lyon I

Publication List



André, A., Quillévéré, F., Schiebel, R., Morard, R., Howa, H., Meilland, J., Douady, C.J., 2018. Disconnection between genetic and morphological diversity in the planktonic foraminifer Neogloboquadrina pachyderma from the Indian sector of the Southern Ocean. Mar. Micropaleontol. 144, 14–24. doi:10.1016/j.marmicro.2018.10.001

Schmidt, C., Morard, R., Romero, O., Kucera, M., 2018. Diverse Internal Symbiont Community in the Endosymbiotic Foraminifera Pararotalia calcariformata: Implications for Symbiont Shuffling Under Thermal Stress. Front. Microbiol. 9, 1–11. doi:10.3389/fmicb.2018.02018

Schiebel, R., Smart, S.M., Jentzen, A., Jonkers, L., Morard, R., Meilland, J., Michel, E., Coxall, H.K., Hull, P.M., de Garidel-Thoron, T., Aze, T., Quillévéré, F., Ren, H., Sigman, D.M., Vonhof, H.B., Martínez-García, A., Kučera, M., Bijma, J., Spero, H.J., Haug, G.H., 2018. Advances in planktonic foraminifer research: New perspectives for paleoceanography. Rev. Micropaléontologie 1–26. doi:10.1016/J.REVMIC.2018.10.001

Morard, R., Mahé, F., Romac, S., Poulain, J., Kucera, M.,de Vargas, C., 2018. Surface ocean metabarcoding confirms limited diversity in planktonic foraminifera but reveals unknown hyper-abundant lineages. Sci. Rep. 8:2539, 1–10. doi:10.1038/s41598-018-20833-z


Kucera, M., Silye, L., Weiner, A.K.M., Darling, K.F., Lübben, B., Holzmann, M., Pawlowski, J., Schönfeld, J., Morard, R., 2017. Caught in the act: anatomy of an ongoing benthic–planktonic transition in a marine protist. J. Plankton Res. 39, 436–449. doi:10.1093/plankt/fbx018

Morard, R., Lejzerowicz, F., Darling, K.F., Lecroq-bennet, B., Pedersen, M.W., Orlando, L., Pawlowski, J., Mulitza, S., Vargas, C. De, Kucera, M., 2017. Planktonic foraminifera-derived environmental DNA extracted from abyssal sediments preserves patterns of plankton macroecology. Biogeosciences 14, 2741–2754. doi:10.5194/bg-14-2741-2017


Morard, R., Escarguel, G., Weiner, A.K.M., André, A., Douady, C.J., Wade, C.M., Darling, K.F., Ujiié, Y., Seears, H.A., Quillévéré, F., de Garidel-Thoron, T., de Vargas, C., Kucera, M., 2016. Nomenclature for the Nameless: A Proposal for an Integrative Molecular Taxonomy of Cryptic Diversity Exemplified by Planktonic Foraminifera. Syst. Biol. 65, 925–940. doi:10.1093/sysbio/syw031

Morard, R., Reinelt, M., Chiessi, C.M., Groeneveld, J., Kucera, M., 2016. Tracing shifts of oceanic fronts using the cryptic diversity of the planktonic foraminifera Globorotalia inflata. Paleoceanography 31, 1193–1205. doi:10.1002/2016PA002977

Schmidt, C., Morard, R., Prazeres, M., Barak, H., Kucera, M., 2016. Retention of high thermal tolerance in the invasive foraminifera Amphistegina lobifera from the Eastern Mediterranean and the Gulf of Aqaba. Mar. Biol. 163, 228. doi:10.1007/s00227-016-2998-4

Weiner, A.K.M., Morard, R., Weinkauf, M.F., Darling, K.F., André, A., Quillévéré, F., Ujiie, Y., Douady, C.J., de Vargas, C., Kucera, M., 2016. Methodology for single-cell genetic analysis of planktonic foraminifera for studies of protist diversity and evolution. Fontiers Mar. Sci. 3, 1–15. doi:10.3389/fmars.2016.00255


Schmidt, C., Morard, R., Almogi-Labin, A., Weinmann, A.E., Titelboim, D., Abramovich, S., Kucera, M., 2015. Recent Invasion of the Symbiont-Bearing Foraminifera Pararotalia into the Eastern Mediterranean Facilitated by the Ongoing Warming Trend. PLoS One 10, e0132917. doi:10.1371/journal.pone.0132917

Spezzaferri, S., Kucera, M., Pearson, P.N., Wade, B.S., Rappo, S., Poole, C.R., Morard, R., Stalder, C., 2015. Fossil and genetic evidence for the polyphyletic nature of the planktonic foraminifera “Globigerinoides”, and description of the new genus Trilobatus. PLoS One 10, e0128108. doi:10.1371/journal.pone.0128108

De Vargas, C., Audic, S., Henry, N., Decelle, J., Mahe, F., Logares, R., Lara, E., Berney, C., Le Bescot, N., Probert, I., Carmichael, M., Poulain, J., Romac, S., Colin, S., Aury, J.-M., Bittner, L., Chaffron, S., Dunthorn, M., Engelen, S., Flegontova, O., Guidi, L., Horak, A., Jaillon, O., Lima-Mendez, G., Luke, J., Malviya, S., Morard, R., Mulot, M., Scalco, E., Siano, R., Vincent, F., Zingone, A., Dimier, C., Picheral, M., Searson, S., Kandels-Lewis, S., Acinas, S.G., Bork, P., Bowler, C., Gorsky, G., Grimsley, N., Hingamp, P., Iudicone, D., Not, F., Ogata, H., Pesant, S., Raes, J., Sieracki, M.E., Speich, S., Stemmann, L., Sunagawa, S., Weissenbach, J., Wincker, P., Karsenti, E., Boss, E., Follows, M., Karp-Boss, L., Krzic, U., Reynaud, E.G., Sardet, C., Sullivan, M.B., Velayoudon, D., 2015. Eukaryotic plankton diversity in the sunlit ocean. Science. 348, 1–11. doi:10.1126/science.1261605

Morard, R., Darling, K.F., Mahé, F., Audic, S., Ujiié, Y., Weiner, A.K.M., André, A., Seears, H. a., Wade, C.M., Quillévéré, F., Douady, C.J., Escarguel, G., de Garidel-Thoron, T., Siccha, M., Kucera, M., de Vargas, C., 2015. PFR2: a curated database of planktonic Foraminifera18S ribosomal DNA as a resource for studies of plankton ecology, biogeography, and evolution. Mol. Ecol. Resour. 49, 1–14. doi:10.1111/1755-0998.12410


André, A., Quillévéré, F., Morard, R., Ujiié, Y., Escarguel, G., de Vargas, C., de Garidel-Thoron, T., Douady, C.J., 2014. SSU rDNA Divergence in Planktonic Foraminifera: Molecular Taxonomy and Biogeographic Implications. PLoS One 9, e104641. doi:10.1371/journal.pone.0104641


Morard, R., Quillévéré, F., Escarguel, G., Garidel-thoron, T. De, 2013. Ecological modeling of the temperature dependence of cryptic species of planktonic Foraminifera in the Southern Hemisphere. Palaeogeography, Palaeoclimatology, Palaeocology. 391 Part A, 13–33.

Quilllévéré, F., Morard, R., Escarguel, G., Douady, C.J., Ujiié, Y., de Garidel-Thoron, T., de Vargas, C. 2013. Global scale same-specimen morpho-genetic analysis of Truncorotalia truncatulinoides: A perspective on the morphological species concept in planktonic foraminifera. Palaeogeography, Palaeoclimatology, Palaeocology, 391 Part A, 2–12.

André, A., Weiner, A., Quillévéré, F., Aurahs, R., Morard, R., Douady, C.J., Garidel-Thoron, T. de, Escarguel, G., Vargas, C. de, Kucera, M., 2013. The cryptic and the apparent reversed : lack of genetic differentiation within the morphologically diverse plexus of the planktonic foraminifer Globigerinoides sacculifer. Paleobiology 39, 21–39.

Guillou, L., Bachar, D., Audic, S., Bass, D., Berney, C., Bittner, L., Boutte, C., Burgaud, G., de Vargas, C., Decelle, J., Del Campo, J., Dolan, J.R., Dunthorn, M., Edvardsen, B., Holzmann, M., Kooistra, W.H.C.F., Lara, E., Le Bescot, N., Logares, R., Mahé, F., Massana, R., Montresor, M., Morard, R., Not, F., Pawlowski, J., Probert, I., Sauvadet, A.-L., Siano, R., Stoeck, T., Vaulot, D., Zimmermann, P., Christen, R., 2013. The Protist Ribosomal Reference database (PR2): a catalog of unicellular eukaryote small sub-unit rRNA sequences with curated taxonomy. Nucleic Acids Res. 41, D597–604.


Morard, R., Quillévéré, F., Douady, C.J., de Vargas, C., de Garidel-Thoron, T., Escarguel, G., 2011. Worldwide genotyping in the planktonic foraminifer Globoconella inflata: Implications for life history and paleoceanography. PLoS One 6, e26665. doi:10.1371/journal.pone.0026665


Morard, R., Quillévéré, F., Escarguel, G., Ujiie, Y., de Garidel-Thoron, T., Norris, R.D., de Vargas, C., 2009. Morphological recognition of cryptic species in the planktonic foraminifer Orbulina universa.Mar. Micropaleontol. 71, 148–165. doi:10.1016/j.marmicro.2009.03.001

Contribution to Academic conferences and seminars

Morard, R. Constraining the biodiversity of planktonic foraminifera using Next-Generation Sequencing. Eukaryotic-Omics, Geneva , Switzerland, 25-26th April 2014.
Morard, R. Constraining the biodiversity of marine microplankton using Next-Generation Sequencing. Biology Seminar, Kaiserslautern, Germany, 3rd of February 2014.
Weiner, A., Morard, R., Kucera, M., (2014) Extent of cryptic diversity in planktonic foraminifera morphospecies. http://www.sgmeet.com/osm2014/viewabstract.asp?AbstractID=12992. Ocean Sciences Meeting 2014, Honolulu, USA).
Weiner, A., Morard, R., Kucera, M. Genetic diversity and phylogeography of planktonic Foraminifera. In: Qian PY, Kolter R (eds) Gordon Research Conference on Marine Molecular Ecology 2013, Hongkong, 2013.
Morard, R., Mahé, F., Audic, S., Romac, S., Ujiié, Y., Garet-Delmas, M. J., Weiner, A., Kucera, M., de Vargas, C. The potential of Next Generation Sequencing to constrain the global biodiversity of planktonic foraminifera. The Micropalaeontological Society’s Spring Meeting, Prague, Czech Republic, June 19th-22, 2013
Morard R., Le Bescot, N. Mahé, F., Audic S., Romac S., Not F., Ujiié Y., de Vargas C. Reaching the boundaries of global planktonic foraminiferal biodiversity using Next Generation Sequencing. SCOR/IGBP Working group 138, Modern Planktic Foraminifera and Ocean changes, Amsterdam (Netherlands), 1st September 2011.
Morard R., Quillévéré F., Douady C. J., Escarguel G., de Garidel-Thoron T and de Vargas C. The allopatric distribution of Globoconella inflata cryptic species and its potential used for monitoring past movements of the Antarctic subpolar front. FORAMS 2010, Bonn, Germany, September 5th-10th, 2010.
Morard, R. Morphometric analyses of planktonic foraminiferal cryptic species: new perspectives for paleoceanographic reconstructions. Seminar, Tuebingen. 20th of June 2010.
Quillévéré F., Morard R., de Vargas C. Douady C. J., Ujiié Y., de Garidel-Thoron T and Escarguel G. Truncorotalia truncatulinoides and the value of direct morpho-genetic comparisons in planktonic foraminifera. FORAMS 2010, Bonn, Germany, September 5th-10th, 2010.
Morard, R., Quillévéré, F., Escarguel, G., Ujiié, Y., De Garidel Thoron T., Lécuyer C. and de Vargas C. Morphometric and stable isotope (δ13C, δ18O) analyses of planktonic foraminiferal cryptic species: new perspectives for paleoceanographic reconstructions. Bioindicators of past and present environments, The Micropalaeontological Society’s Spring Meeting, Tübingen, Germany, 15th -17th May 2008.
Morard, R. Eco- morpho- genotypes in planktonic foraminifera : new perspectives for paleoceanographic reconstructions. Sarco – Deal, Villefranche sur Mer, 13-15th November 2007.

Cruise reports

Kucera, M., Morard, R., Siccha. M., Weiner, M., Weinkauf, M. Cruise report of RV Sonne Cruise SO226-3 DipFIP ‒ the extent and structure of cryptic diversity in morphospecies of planktonic Foraminifera of the Indopacific Warm Pool: Wellington‒Kaohsiung, 04.03.2013‒28.03.2013. Berichte aus dem MARUM und dem Fachbereich Geowissenschaften der Universität Bremen. 06/2013; 293:1‒39.


MsM39 - F/S MerianNorth West Atlantic (Leg St John's - St John's, Canada, June 2014)
DipFIP - F/S SONNEWestern Equatorial Pacific (Leg Wellington (New Zealand) - Koashiung (Taiwan), March 2013)
Tara Ocean expedition - TARA schoonerNorth East Pacific (Leg Hawaï - San Diego, USA, Sept. - Oct. 2011).
BioMarks projectBallast water sampling (Rotterdam harbour, Netherlands, June 2011).
FORCLIM-7 - R/V Sarmiento de GamboaGulf of Biscay (Leg Vigo – Vigo, Spain, April 2009).
GYRAFOR-A - R/V ALISWestern equatorial Pacific (Leg Nouméa – Nouméa, New Caledonia, Jun.-Jul. 2008).