This (rather long) page gives details of the eight participant research teams: their role and activities within the project, some background on the teams and listing of participating scientists. N.B. This information is from the Research Proposal, and apart from insertion of the names of the appointed young visiting scientists has not been updated. The teams are:

      1. The Natural History Museum, London [NHM]
      2. Eidgenossische Technische Hochschule, Zuerich [ETHZ] - Geologisches Institut
      3. Vrije Universiteit Amsterdam, Faculteit der Aardwetenschapen [FdA-VUA]
      4. Universite de Caen Basse-Normandie (U. Caen)
      5. Alfred-Wegener-Institute for Polar & Marine Research, Bremerhaven [AWI]
      6. Consejo Superior de Investigaciones Cientificias - Instituto de Ciencias del Mar, Barcelona [CSIC]
      7. Museu Nacional de Historia Natural da Universidade de Lisboa [MNHN-UL]
      8. Nederlands Instituut voor Onderzoek der Zee [NIOZ]

      1. The Natural History Museum, London [NHM]

      The NHM team will carry out research on the morphological and geological evidence for the phylogenetic relations of the selected coccolithophorids. In addition it will carry out detailed work on Umbilicosphaera sibogae, Helicosphaera carteri and Syracosphaera pulchra in collaboration with FdA-VUA, The team is also responsivble for network coordination, and for development of light microscopy based image analysis methods and cladistic analysis of phylogenetically relevant data.


      • 1) Ultrastructural analysis of coccoliths of all selected taxa using high resolution SEM and quantitative polarised light microscopy techniques. (RT6)
      • 2) Development of image analysis based biometric techniques using light microscopy including development of specific applications for most of the studied taxa. Provision of training in this technique for MNHN-UL and FdA-VUA teams. (for RT10-14)
      • 3) Geological sample studies: study of microevolution and ecological distribution of Umbilicosphaera and Helicosphaera in the selected DSDP/ODP cores. (RT14)
      • 4) Macroevolutionary studies; special studies of selected cladogenetic events in order to maximise the geological data on phylogenetic relations (particularly divergence times) of the selected taxa. (RT15)
      • 5) Cladistic analysis of phylogenetic relations - using the detailed data on the selected taxa to provide a core analysis and as wide an analysis as practicable of other haptophytes relative to this group of taxa. (RT8)
      • 6) Physiological studies using light-temperature cross gradient cabinets on strains of H. carteri, S. pulchraand U. sibogae, in collaboration with visiting participants from FdA-VUA. Supervision of similar studies by visiting participants from MNHN-UL on C. pelagicus. (RT9)


      The NHM is Europe's largest centre for taxonomy-related research and has outstanding collection, library and research facilities. Of particular relevance are its extensive expertise in, and facilities for, electron microscopy (SEM and TEM), image analysis and biometrics, algal culturing, and cladistic analysis. The team includes members with special expertise in these fields, and additional coccolithophorid experts from University College London (UCL). The team has previously been working on two directly relevant projects - (a) detailed study of Emiliania huxleyi coccolith variation including development of image analysis techniques, within the MAST EHUX project, involving JRY and JAB. (b) study of coccolithophorid biomineralization and phylogeny (NERC funded project involving JRY and PRB). Our work in the CODENET project will be a synthesis and development of these. The team members have widespread, mostly informal, links with members of all the other teams.


      • Dr. Jeremy R. Young, Palaeontology Dept. Team co-ordinator.
      • Dr. Gianfranco Novarino. Zoology Dept.
      • Dr. David M. Williams. Botany Dept.
      • Dr. Paul R. Bown. Dept. of Geological Sciences, UCL.
      • Dr. Jackie A. Burnett (1 month/year). Dept. of Geological Sciences, UCL.
      • Andrew Howard (1 month/year). NHM & UCL, PhD student
      • Markus Geisen CODENET funded young visiting scientist

      Associated researchers - additional collaboration on CODENET objectives has been agreed with

      • Prof. Steven Mann and Dr. Sean Davis of University of Bath, Chemistry Dept., working on cocolith biomineralisation.
      • Prof. Patrick Holligan and Dr. Toby Tyrell, University of Southampton, Dept. of Oceanography.

      2. Eidgenossische Technische Hochschule, Zuerich [ETHZ] - Geologisches Institut

      (Swiss Federal Institute of Technology, Zuerich, Geological Institute)

      The ETHZ team will study the microevolution, ecology and palaeoecology of Calcidiscus leptoporus and Gephyrocapsa. This will be integrated with previous biogeographic studies of these groups, and with the genetic and biochemical results of other teams. They will seek to answer the following specific scientific questions: (1) When did the different morphotypes of the two taxa evolve (Phylogeny)? (2) Can the various morphotypes of coccoliths of Gephyrocapsa and C. leptoporus known from Holocene sediments also be identified on living coccospheres and what are their environmental preferences? (3) How large is the morphological plasticity of monoclonal populations cultured under different environmental conditions? (4) Can populations of the various morphotypes be characterised genetically and does genetic differentiation agree with the known geological ages (calibration of molecular clocks)?

      They will also contribute plankton samples, and water samples for strain isolation work from Bermuda, Puerto Rico, Hawaii, an Atlantic transect and the Canary Islands. They will co-ordinate selection of geological material for the network including sample requests to the ODP and provision of a globally distributed set of Holocene sediment samples.


      • 1) Microevolution and phylogeny of Calcidiscus and Gephyrocapsa. The morphological development of all morphotypes in these two groups will be analysed in selected DSDP/ODP sites since their first appearance. In each of 200-500 samples 50-200 specimens of each taxon will be measured. (RT14)
      • 2) Integration of ecological results from studies of modern coccolithophorid assemblages. Living plankton samples from JGOFS time series stations at Bermuda, Hawaii and Canary Islands will have been morphometrically analysed in our currently funded EC and NSF projects. These results will be incorporated and complemented as necessary by similar samples from Puerto Rico, the Atlantic transect (AMT cruise) and material from other teams. (RT11)
      • 3) Morphological plasticity. Monoclonal populations of the various morphotypes will be cultured at different temperatures, salinities and light levels and their morphologies quantified in order to interpret the potential spatial and temporal continuities of morphologies/species? (RT9, 11)
      • 4) Culture isolation. Strains of Gephyrocapsawill be isolated, particularly from the Canary Islands and Bermuda. Water samples will also be contributed for strain isolation by U. Caen. (RT1)


      The research team has outstanding expertise in geological study of coccoliths and includes two of the leading authorities in the field. It has been at the forefront of application of rigorous scientific approaches to palaeoceanography, and micropalaeontology. The team has previously carried out directly relevant research on Calcidiscus and Gephyrocapsa (PhD theses of Knappertsbusch, 1990; Bollmann, 1995). Currently the coccolithophorid population dynamics at the JGOFS time series stations of Bermuda and Hawaii are being studied by two PhD students. Other related ongoing projects focus on plankton diversity and longevity, high-resolution biochronology, development of computer assisted microscopy and automated microfossil identification.s


      • Hans Thierstein (1 month/year). Professor of Geology. Team co-ordinator.
      • Joerg Bollmann Post doctoral researcher.
      • Christine Klaas Post doctoral researcher.
      • Jorijntje Henderiks PhD Student
      • Katharina von Salis Perch-Nielsen. Professor of Geology (since 1992). Leading authority on nannofossil taxonomy and biostratigraphy.
      • Dr. Sabrina Renaud CODENET funded young visiting scientist.

      3. Vrije Universiteit Amsterdam, Faculteit der Aardwetenschapen [FdA-VUA]

      (Earth Sciences Faculty, Free University Amsterdam)

      The FdA-VUA team will perform analyses of coccolith assemblages from sediment traps with the particular aim of quantifying their role in the global carbon cycle. Also, in collaboration with the NHM team, they will carry out detailed study of the taxa Helicosphaera carteri, Syracosphaera pulchra and Umbilicosphaera sibogae . This study will be based on water, sediment trap and sediment samples. Extensive sample collections are available at FdA-VUA for this work, with global coverage including sediment trap samples from Susumu Honjo and Robert Thunnell. The team will co-ordinate study of sediment trap samples including methodology training and inter-calibration. The team will also organise annual network workshops, to be held either in Amsterdam or France (Ch. de Blagnac).


      • 1. Sediment trap studies of seasonal development of coccolith assemblages; transformation of assemblages during settling through the water column and modern species specific flux rates. Determination of species-specific carbonate flux and accumulation rates in comparison with total carbonate flux and accumulation rates. (RT12)
      • 2. Detailed study of the morphological variability, seasonal distribution, biogeography, and palaeobiogeography of H. carteri, S. pulchra and U. sibogae. This work will be co-ordinated with parallel study by the NHM team of these taxa in culture and in geological samples - including research visits to the NHM to participate in culture experiments. (RT11-13)


      The Geomarine research team under the leadership of Prof. van Hinte has been researching actuomicropalaentology for the last ten years and has played a leading role in developing oceanographic research themes of direct relevance to geological studies with major roles in development of sediment trap techniques, isotope geology and integrated biological-palaeobiological studies of planktonic organisms. Participation in the network will bring essential expertise into the network and allows the team to extend their current research in a much broader context.

      The team played a core role in the Global EmilianiaModelling initiative (including workshop organisation), and consequently worked with the NHM, NIOZ groups. The team has particularly close links with ETHZ including exchange of post-doctoral workers.


      • Prof. Jan van Hinte. Professor of Palaeontology/Stratigraphy and Marine Geology at FdA-VUA. Team co-ordinator.
      • Dr. Patrizia Ziveri. Post-Doctoral researcher (young visiting scientist funded by the project).
      • Prof. Peter Westbroek. Docent Biogeochemistry at Rijks Universiteit Leiden. Co-ordinator of the Global Emiliania Modelling initiative (GEM). Experience in coccolithophorid biochemistry and calcification.
      • Dr Gerald Ganssen. Research Scientist
      • 2 PhD students (12 months/year). The team currently includes 2 PhD students, Anna Lototskaya and Sandra Broerse working on directly relevant research projects. They are both due to finish their theses in Summer 1998. They will be succeeded by 2 further PhD students to be applied for in 1997 as a GEM continuation proposal to the Dutch science foundation (NWO).
      • Dr. Annelies Kleijne, (Amsterdam Microanalysis Laboratory), sub-contractor.

      Associated researchers - additional collaboration on CODENET objectives has been agreed with

      • Dr. Elisabetta Erba, Prof. Maria B. Cita, University of Milan, Earth Sciences Dept.

      4. Universite de Caen Basse-Normandie (U. Caen)

      The U. Caen team's principal research objective is to elucidate and characterise the life-cycles of the studied coccolithophorids and to obtain information on the factors affecting life-cycle changes in these coccolithophorids. In addition the team is responsible for isolation of new coccolithophorid strains and for maintenance of a central culture collection.


      • 1) Culture isolation and maintenance: New cultures of coccolithophorids will be established from field samples (obtained in collaboration with CSIC, MNHN-UL and ETHZ) to extend the range of coccolithophorids already available in culture in Caen for experimental purposes. These cultures will be made available to all network participants. In addition cultures obtained by other participants (particularly ETHZ) will be maintained. (RT1)
      • 2) Life-cycle studies: Experiments will be undertaken to determine the factors controlling reproduction and eventual life-cycle changes in the selected species, and ploidy levels of the isolated phases when applicable. (RT2)
      • 3) TEM fine structural studies: The life-cycle stages will be studied using transmission electron microscopy - with particular emphasis on study of organic scale morphology, scale and/or coccolith production, flagellar apparatus, and investigations of changes in cell organisation during cell division and life-cycle transformations. (RT7)


      The Phycology Group of the Laboratoire de Biologie et Biotechnologies Marines of Caen University has over 25 years of experience in isolating and growing marine microalgae resulting in the establishment of one of the largest collections in the country with over 100 strains (mostly original) including a number of type species. Main areas of research concern biology and systematics of coastal phytoplankton groups (essentially phytoflagellates), including coccolithophorids and non-calcified haptophytes, of which 9 species were described in Caen. Equipment available includes culture rooms, optical and EM facilities (TEM , SEM with freeze fracturing devices) and flow cytometry.


      • Prof. Chantal Billard. Team co-ordinator.
      • Dr. Jacqueline Fresnel
      • Dr. Ian Probert CODENET funded young visiting scientist.

      5. Alfred-Wegener-Institute for Polar & Marine Research, Bremerhaven [AWI]

      The AWI team will investigate the phylogenetic relatedness of keystone coccolithophorid algae with a range of molecular markers useful over different time scales and different taxonomic levels and to estimate divergence times for these taxa by calibrating a molecular clock from first appearance dates in the fossil record.


      • 1) Sequencing of the 18S rRNA gene or other non-coding regions from selected coccolithophorid taxa for comparison with all Haptophytes completed to date to reconstruct the phylogenetic history of the coccolithophorids. (RT3B, AWI)
      • 2) Calculation of rates of evolution and a molecular clock for the coccolithophorids using the 18S ssu rRNA gene and non-coding spacer regions. (RT3B, AWI)
      • 3) Gephyrocapsa microevolution. Identification of regions of the genome, for use at the population level, to evaluate gene flow and dispersal and microevolution of populations within the Gephyrocapsa complex. Microsatellite probes will be developed in conjunction with a similar investigation for Emiliania huxleyi as another population level marker. (RT3C-D, AWI).
      • 4) Determination of the plastid genome size of the selected coccolithophorid algae using PFGE. (RT3A, Uni. Erlangen).
      • 5) Creation of a plastid genomic library to isolate and sequence specific genes and their flanking regions that are well characterised with respect to flanking regions/ gene order and arrangements for comparison. Hybridization experiments with primers specific for certain genes that are usually not found on the plastid genome but have been transferred to the nucleus during evolution will be carried out. (RT3A, Uni. Erlangen)


      The Alfred-Wegener-Institute for Polar and Marine Research is one of 13 national research centres in Germany and plays a major role in sponsoring climate and environmental research. Recently, a molecular biology working group has been established at the AWI, led by Dr. L. Medlin, to study molecular ecology and phylogenetics of marine phytoplankton. Dr. Medlin has used a variety of molecular techniques to investigate problems in molecular systematics and phylogeny and population variation in marine phytoplankton, and pioneered application of PCR techniques to the amplification of ribosomal RNA genes for phylogenetic use. Her working group has concentrated on the use of molecular techniques (DNA sequencing and RAPD fingerprinting) to address broad spectrum phylogenetic questions, species level questions and spatial distribution of genetic diversity in marine phytoplankton. Recent advances in the molecular group include the development of genus-specific probes for Phaeocystis, Emiliania, and Chrysochromulina and calculations of a molecular clock for the diatoms from nuclear and plastid genes.


      • Linda K. Medlin, AWI Senior Research Scientist. Team co-ordinator.
      • Volker A.R. Hus: Privatdozent, Dept. of Botany, Univ. of Erlangen.
      • Alberto Garcia Saez CODENET funded young visiting scientist.

      6. Consejo Superior de Investigaciones Cientificias - Instituto de Ciencias del Mar, Barcelona [CSIC]

      (Spanish Council for Scientific Research - Marine Sciences Institute, Barcelona)

      The CSIC team will study the biochemical composition of the selected taxa and investigate the ecological factors determining the distribution of coccolithophorid assemblages in the marine environment, building on the research tradition of the group in this field. In addition, the CSIC team will provide a Mediterranean base for the network.


      • 1) Ecological characterisation of coccolithophorid assemblages. Study of distribution of phytoplankton assemblages, with particular emphasis on coccolithophorids, in relationship to hydrographic conditions. Samples will be obtained during cruises programmed in the context of other projects, mainly in the Mediterranean. Taxonomic observations will be based on optical and electron microscopy. (RT11)
      • 2) Photosynthetic pigment characterisation Selected strains of each of the species will be cultured under carefully standardised conditions, at varying light levels, in order to determine the composition and total content of chlorophyll. This will be used to estimate the pigment composition of the studied taxa in natural conditions and to calibrate estimates of coccolithophorid contribution to total chlorophyll production (RT5). The results of this culture work will also contribute toward physiological characterisation of the species (RT9).
      • 3) Lipid characterization. Alkenones and other lipids produced by coccolithophorids will be studied based on both laboratory culture experiments and study of field samples. This work will be carried out at the CID laboratory of Dr. J. Grimalt and will form a joint project with NIOZ. The CSIC-CID will focus on ecological determinants of lipid composition including particularly the UK37 index (RT4).


      The CSIC team combines research experience in marine geology, environmental chemistry and the study of plankton dynamics in relation to physical and chemical environmental variables. The members of the team have collaborated on research subjects relevant to CODENET aims. Their research activity has been funded by several Spanish and EC funded projects. The ICM and the CID are provided with all necessary equipment to carry out the proposed work. The ICM has access to use the R/V Hesperides with capacity for 29 scientists, funding applications for cruises directly relevant to CODENET have been made.


      • Marta Estrada "Profesora de investigacion", presently vice-director of the ICM. Team co-ordinator.
      • Mikel Latasa "Profesor de Investigacion" at ICM, since 1996.
      • Joan Grimalt "Profesor de Investigacion" at CID, since 1991.
      • Belen Alonso and Albert Palanques, "Colaboradores Cientificos".
      • Lluisa Cros "Titulada Tecnica".
      • Carles Pelejero PhD student, CID.
      • Ramon Margalef Emeritus Professor of the University of Barcelona
      • Kees van Lenning CODENET funded young visiting scientist.

      7. Museu Nacional de Historia Natural da Universidade de Lisboa [MNHN-UL]

      (National Museum of Natural History, University of Lisbon)

      The Lisbon group will conduct a detailed study of Portuguese shelf coccolithophorids, including water column, surface sediment and Quaternary samples. They will also carry out detailed study of the biogeography, palaeobiogeography and evolution of Coccolithus pelagicus.

      Principal Research Activities

      • 1) Study of Portuguese shelf coccolithophorids. Collection of water samples and monitoring of coccolithophorid assemblages at monthly and/or seasonal periodicity. Particular attention will be given to understand spatially distinct water-mass biocoenoses distributions, to monitoring changes during upwelling conditions and to analysing the influence of river mouths on coccolithophorid development. This study will be carried out directly from water column samples, from onboard filters and from Quaternary piston cores retrieved from the Portuguese continental margin. In association with this water samples will be collected for isolation of new strains of the keystone species from this area.
      • This study will include integration of previous data to produce a 7 year matrix of coccolithophorid distribution relative to other phytoplankton and physico-chemical parameters. This will be used for ecological characterisation of the key coccolithophore species and direct comparison with Quaternary palaeoecological records (RT11,14).
      • 2) Detailed morphometric studies of Coccolithus pelagicus in order to study its biogeography, palaeobiogeography and evolution. This will include study of Holocene sediment samples and Geological sample sets as co-ordinated by ETHZ, and participation in culture work in collaboration with NHM and CSIC. (RT10-14).


      The Portuguese team has 10 years experience in collecting water column samples (on annual, seasonal and monthly bases) to perform biogeographic and ecological studies of phytoplankton communities in general and Coccolithophorids in particular.

      In the early 1990's we started to study surface and piston core sediments from Portuguese continental platform and shelf areas (Tagus estuary, coastal lagoons) in order to characterise coccolith thanato- and taphocoenoses and their changes through time (Late Quaternary) and space. We are also exploring their potential use as marine tracers inside paralic environments (e.g. as tsunami indicators) and as a proxy of particular hydrodynamic conditions (reworking) near the bottom of certain sectors of the Portuguese continental slope (e.g. Mediterranean outflow related contourite on Algarve slope).

      Our location and cruise facilities allow us to contribute to other teams efforts by collecting living Atlantic coccolithophorid specimens and we want to learn from other teams experience in culture experiments to test several hypotheses concerning the behaviour of Coccolithus pelagicus in particular.


      • Mario Cachao Geologist, "Professor auxiliar" at the Dept. of Geology, University of Lisbon, team coordinator
      • Anabela Olivera Geologist, doctoral researcher at IH (Instituto Hidrografico) and at UCTRA (University of Algarve).
      • Maria Teresa Moita Biologist, doctoral researcher at IPIMAR (Instituto Portugues de Investigacao das Pescas e do Mar).
      • Maria da Graca Vilarinho. Biologist, doctoral researcher at IPIMAR.

      8. Nederlands Instituut voor Onderzoek der Zee [NIOZ]

      (Netherlands Institute for Sea Research)

      The NIOZ team will study the biochemistry and biogeochemistry of the selected coccolithophorids, and also of the non-calcifying Haptophyte Isochrysis galbana, including particular lipid and stable isotope studies. The research will constitute a joint project with Dr. Grimalt's group in the CSIC team. The NIOZ work will focus on very detailed analysis of the lipids based on advanced separation techniques (e.g. liquid column chromatography and thin layer chromatography) and including C13 analyses.


      • 1) Free and complex lipids of cultured biomass, including hydrocarbons, alcohols, ketones, fatty acids, sterols, glycerides, ethers, etc. will be analysed qualitatively and quantitatively with GC and GC/HRMS. This will include analyses of each of the taxa of inter-strain differences within the taxa, especially Gephyrocapsa and of cultures grown under varying conditions. (RT4A-C)
      • 2) Selected suites of sediment trap and sediment samples will be analysed for free and complex lipids, structurally and isotopically, with an emphasis on lipids possibly originating from coccolithophorids and compared with microscopic studies performed on the same samples. (RT4D)


      The Netherlands Institute of Sea Research (NIOZ) is the principal oceanographic institute in The Netherlands. It has >30years experience in ecosystem and algal research and routinely participates in major multidisciplinary and multinational studies.

      The team is part of the Marine Biogeochemistry & Toxicology Department of the NIOZ. The team has >25 years of experience mainly built up at Delft University before the whole group moved to the NIOZ, in January 1993. The group has published ca. 400 publications in peer-reviewed scientific journals, including ca.15 in Nature and Science.

      The team presently consists of 25 people; 2 staff members, 4 technicians, trainees, many PhD students and several post-docs as well as some visiting scientists. All the high-quality instrumentation required for the research proposed is present in-house. The major emphasis of the group's research is recognition and isolation of novel low- and high-molecular-weight (macromolecular) lipids, their biosynthesis, fate in the sedimentary record, evolutionary pathways and application to reconstruction of palaeo-environments and the genesis of fossil fuels. Sponsorship for these activities has come from NWO (the Dutch science funding council), Industry, EC and governmental bodies.


      • Dr ir Jaap S. Sinninghe Daamste senior scientist, Team co-ordinator.
      • Prof Dr. Jan W. de Leeuw -.Director of NIOZ and professor in Organic Geochemistry at University of Utrecht.
      • Dr ir Gerard Versteegh post doctoral researcher
      • Wim Pool, Technician, M. Dekker, Technician
      • Dr. Hanno Kinkel CODENET funded young visiting scientist

      Return to: top, Introduction, Phylogeny, Microevolution, Ecology, Research Tasks, Team Details, The Species

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      This page is based on the CODENET proposal, as accepted for funding by TMR, June 1997