The Significance of Extant Coccolithophores As Indicators of Ocean Water Masses 95

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The Significance of Extant Coccolithophores As Indicators of Ocean Water Masses 95 Paläontologische Zeitschrift 1 2005, Vol. 79/1, p. 93–112, 31–03–2005 2 3 4 5 6 The significance of extant coccolithophores as indicators of ocean water 7 masses, surface water temperature, and paleoproductivity: a review 8 9 KARL-HEINZ BAUMANN, Bremen; HARALD ANDRULEIT, Hannover; BABETTE BÖCKEL, Bremen; 10 MARKUS GEISEN, Bremerhaven & HANNO KINKEL, Kiel 11 12 with 13 figures 13 14 BAUMANN, K.-H.; ANDRULEIT, H.; BÖCKEL, B.; GEISEN, M. & KINKEL, H. 2005. The significance of extant cocco- 15 lithophores as indicators of ocean water masses, surface water temperature, and paleoproductivity: a review. – Palä- 16 ontologische Zeitschrift 79 (1): 93–112, 13 figs., Stuttgart, 31. 3. 2005. 17 Abstract: Coccolithophores are one of the main groups of marine phytoplankton playing key roles in the marine eco- 18 system as primary producers and in marine biogeochemistry. These organisms have gained considerable attention as 19 they play a unique role in the global carbon cycle because of their combined effects on both the organic carbon and the 20 carbonate pump. Although steady advances in research on coccoliths as biogeochemical agents and palaeontological 21 proxies were obtained knowledge of the biology of these organisms has only progressed considerably in recent years. It 22 has been confirmed that holococcolithophores are not autonomous but stages in the life cycle of heterococcolithophores. A general introduction to the taxonomy and biology of extant coccolithophores is followed by a brief overview of the 23 environmental parameters affecting these phytoplanktonic organisms and their biogeography. Another chapter summa- 24 rizes the investigations on coccolithophores in the Arabian Sea which have been conducted during the past years to pro- 25 duce a synthesis of the production of living coccolithophores in the photic zone, their transformation to settling assem- 26 blages, their accumulation on the seafloor, and their final burial in the sediments. In the following, applications of 27 coccolithophores to palaeoenvironmental analyses are provided as case studies. In particular, the usage of both ecolog- 28 ically restricted species, such as Florisphaera profunda, for palaeoproductivity studies and of a new coccolithophore- 29 based palaeothermometry for surface-water reconstructions are presented. In addition, haptophyte-specific biomarkers 30 (long-chain alkenones) are reviewed and their applicability for palaeoceanographical reconstructions is demonstrated. 31 Keywords: coccolithopores • phytoplankton • Recent • Quaternary • biology • ecology • paleoproxy • alkenones • 32 carbonate producer 33 Kurzfassung: Coccolithophoriden, einzelliges marines Phytoplankton, stellen eine der Hauptprimärproduzenten in 34 den Ozeanen dar. Diesen haptophyten Algen kommt als Karbonatproduzenten und photoautotrophe Organismen eine 35 bedeutende Rolle im Kohlenstoffkreislauf zu, den sie sowohl über die Karbonat- als auch über die biologische Pumpe 36 beeinflussen. Trotz der intensiven Fortschritte in der Bearbeitung als Träger biogeochemischer Informationen und 37 ihrer steigenden Bedeutung als Paläoumweltindikatoren ist der Kenntnisstand über die Biologie von Coccolithopho- 38 riden erst in jüngster Zeit wesentlich erweitert worden. Es hat sich gezeigt, dass Holococcolithophoriden keine auto- 39 nomen Organismen, sondern Phasen im Lebenszyklus von Heterococcolithophoriden sind. Nach einer Einführung in 40 die Taxonomie und Biologie von Coccolithophoriden werden die Umweltparameter, die diese Phytoplanktongruppe 41 beeinflussen, vorgestellt und ihr Vorkommen sowohl geographisch als auch ökologisch beschrieben. Das folgende 42 Kapitel stellt beispielhaft die Bildung des geologischen Signals von Coccolithophoriden dar. Hierzu wurden Plank- 43 tongemeinschaften aus dem Oberflächenwasser mit Sinkgemeinschaften aus Partikelfallen sowie den unterlagernden Oberflächensedimentgemeinschaften verglichen und bewertet. In den folgenden Fallstudien wird auf die Bedeutung 44 von Coccolithophoriden als Paläoumweltindikatoren eingegangen. Insbesondere werden die Anwendung einzelner, 45 ökologisch relevanter Arten, wie die in der tieferen photischen Zone lebende Florisphaera profunda, für Paläopro- 46 duktivitätsrekonstruktionen sowie eine neue, auf Coccolithophoridenarten basierende Rekonstruktion von Oberflä- 47 chenwassertemperaturen dargestellt. In vielen paläozeanographischen Arbeiten werden Haptophyten-spezifische 48 Biomarker (langkettige Alkenone) intensiv für Temperaturrekonstruktionen genutzt. Im Vergleich zu anderen plank- 49 tischen Karbonatproduzenten zeigt sich, dass Coccolithophoriden heute und im Pleistozän einen wesentlichen Anteil 50 am Eintrag von (anorganischen) Kohlenstoff in das geologische Archiv ausmachen. 51 Schlüsselwörter: Coccolithoporiden • Phytoplankton • rezent • Quartär • Biologie • Ökologie • Paläoproxy • 52 Alkenone • Karbonatproduzenten 53 54 55 Addresses of the authors: KARL-HEINZ BAUMANN and BABETTE BÖCKEL, FB Geowissenschaften, Univ. Bremen, Postfach 330440, D-28334 Bremen. – HARALD ANDRULEIT, Federal Institute for Geosciences and Natural Resources, Postfach 510153, D-30631 56 Hannover. – MARKUS GEISEN, Alfred Wegener Institute for Polar and Marine Research, Am Handelshafen 12, D-27570 Bremer- 57 haven. – HANNO KINKEL, Institute for Geosciences, Univ. Kiel, Olshausenstr.40, D-24118 Kiel. 58 0031– 0220/05/0079– 0093 $ .00 © 2005 E. Schweizerbart’sche Verlagsbuchhandlung, D–70176 Stuttgart 94 KARL-HEINZ BAUMANN, HARALD ANDRULEIT, BABETTE BÖCKEL, MARKUS GEISEN, HANNO KINKEL 1 Introduction Biology of coccolithophores 2 3 The living coccolithophores are marine, unicellular phy- Their long evolutionary record, high evolutionary turn- 4 toplankton, belonging to the phylum Haptophyta and the over and enormous abundance in sediments have made 5 division Prymnesiophyceae (JORDAN & CHAMBERLAIN them ideal biostratigraphical marker fossils and hence 6 1997). Their morphology is characterised by an exoskel- they have been widely used for dating of sediments and 7 eton composed of numerous minute calcite platelets – classical palaecological work. It was however only in 8 the coccoliths – that are readily preserved in the sedi- the beginning of the 20th century that researchers (e. g. 9 mentary record. KAMPTNER 1927; LOHMANN 1902; MURRAY & BLACK- 10 Coccolithophores are one of the main groups of MAN 1898) started observing and exploring the biology 11 marine phytoplankton playing key roles in the marine of the organisms. Although steady advances in research 12 ecosystem as primary producers and in marine biogeo- on coccoliths as biogeochemical agents and palaeonto- 13 chemistry, owing to their great abundance, fast turnover logical proxies were obtained, these have not been par- 14 rates, and their capability to carry out photosynthesis alleled by information in the biology of the species. 15 and calcification (WINTER & SIESSER 1994; BOWN Knowledge of the biology of coccolithophores has, 16 1998). They are, from a quantitative point of view, however, progressed considerably in recent years due to 17 among the most important pelagic calcifying organisms increased numbers of culture studies and meticulous 18 in the modern ocean (BAUMANN et al. 2004; HAY observations in plankton samples and are most recently 19 2004). The evolution of calcareous nannoplankton re- reviewed and summarized by BILLARD & INOUYE 20 sulted in a major shift in the locus of global calcification (2004). 21 from the continental shelves towards deep oceans, Coccolithophores include all haptophyte algae pos- 22 which fundamentally changed ocean chemistry and glo- sessing calcified scales (coccoliths) at some stage in 23 bal sedimentation patterns. Recently, coccolithophores their life cycle. Following the taxonomic revision of the 24 have gained increased attention as they play a unique division Haptophyta recently proposed by EDVARDSEN 25 role in the global carbon cycle and are peculiar because et al. (2000), coccolithophores belong to the class Prym- 26 of their combined effects on both, the biological carbon nesiophyceae, which also features non-calcifying organ- 27 and the carbonate pump. Because of their biochemical isms. Coccolithophores generally occur as single cells 28 effects (dimethylsulfide – which act as a source mole- and their typical features have been previously compiled 29 cule for cloud nucleation) they are likely to produce ad- in earlier reviews (PIENAAR 1994; INOUYE 1997; BIL- 30 ditional feedback to climate change (WESTBROEK et al. LARD & INOUYE 2004; summarized in Fig. 1). 31 1993). The taxonomy of coccolithophores is based on the 32 In addition, coccoliths are phenomenally abundant morphology of the coccoliths that cover their cell. Two 33 in sea-floor sediments, and preserve the composition of major groups exist, the holo- and heterococcoliths (Fig. 34 the overlying photic-zone communities due to protect- 2), of which the heterococcolithophore stage is more 35 ed and accelerated sedimentation in faecal pellets or in common. Whereas holococcoliths are formed out of nu- 36 marine snow. Therefore, they are valuable indicators of merous, minute euhedral calcite crystallites the hetero- 37 palaeoceanographical change. Coccolithophores have coccoliths are formed of a radial array of variably 38 one of the most abundant fossil records of any phylum shaped crystal units (see e. g. YOUNG et al. 1992, 1997). 39 and this fossil record is continuous from their first oc- Formation of the heterococcoliths takes place intracel- 40 currence in the Late Triassic
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