114F. <strong>Tremolada</strong>, T.J. <strong>Bralower</strong> / Marine Micropaleontology 52 (<strong>2004</strong>) 107–116Fig. 7. Palaeoceanographic model for causes <strong>and</strong> effects of the PETM in greenhouse conditions.combination of somewhat higher oxygenation <strong>and</strong>lower productivity (Fig. 6).The PETM triggered the rapid radiation of theplanktonic foraminifera (e.g., Kelly et al., 1996) <strong>and</strong>l<strong>and</strong> mammals (e.g., Clyde <strong>and</strong> Gingerich, 1998).The rate of extinction in the benthic foraminiferalcommunities varies from tropical to high latitude sites(e.g., Kaiho et al., 1996; Speijer et al., 1996). Theextinction of the northern temperate-latitude <strong>and</strong> highlatitudenannofossil species, Octolithus multiplus, waslikely caused by increasing temperature, <strong>and</strong> theelimination of Fasciculithus possibly resulted fromcompetition with Zygrhablithus bijugatus. Furthermore,palynofloras show no extinctions <strong>and</strong> onlymoderate changes in composition <strong>and</strong> diversity duringthe Paleocene–Eocene transition (e.g., Wing <strong>and</strong>Harrington, 2001).6. ConclusionsNannofloral changes associated with the PETMtransition at DSDP Sites 213 <strong>and</strong> 401 correlate withthose documented from several other locations, suchas ODP Site 690 (<strong>Bralower</strong>, 2002) <strong>and</strong> Alamedilla,southern Spain (Monechi et al., 2000). The PETM canbe subdivided into a lower part <strong>and</strong> a upper part on thebasis of nannofossil assemblages. The lower part ischaracterized by high abundances of Discoaster spp.,Ericsonia spp., <strong>and</strong> Fasciculithus spp., interpreted asindicators of warm surface waters <strong>and</strong> probably oligotrophicconditions (Wei <strong>and</strong> Wise, 1990; Aubry,1992; Kelly et al., 1996; <strong>Bralower</strong>, 2002). The upperpart shows an increase in abundance of Toweius spp.<strong>and</strong> Zygrhablithus bijugatus, which likely reflectsmore eutrophic <strong>and</strong> cooler conditions.The major nannofloral changes at DSDP Site 213seem to be strongly affected by diagenesis. Theincrease in abundance of genera most resistant todissolution, such as Discoaster, Ericsonia, <strong>and</strong> Fasciculithus,inversely correlates with the nannofossilspecies richness at Site 213.The appearance of asymmetrical Discoaster <strong>and</strong>Rhomboaster species might indicate the development<strong>and</strong> availability of new ecological niches, resultingfrom major palaeoceanographic changes during thePETM (<strong>Bralower</strong>, 2002). The last occurrence ofOctolithus multiplus might be a reliable bioevent atnorthern intermediate <strong>and</strong> high latitudes.AcknowledgementsWe gratefully acknowledge K. von Salis, J. Young,J. Lees, <strong>and</strong> an anonymous reviewer for criticalcomments that helped us improve the manuscript.Many thanks to I. Premoli Silva, E. Erba, I. Raffi, J.Backman, <strong>and</strong> G. Dickens for thoughtful discussions.Samples were provided by the Ocean DrillingProgram, <strong>and</strong> we thank the curatorial staff for their
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