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The Pennsylvania State University The Graduate School College of Earth and Mineral Sciences A MULTIVARIATE ANALYSIS OF THE RECOVERY OF CALCAREOUS NANNOPLANKTON AND PLANKTONIC FORAMINIFERA FROM THE CRETACEOUS/PALEOGENE (K/P) MASS EXTINCTION A Thesis in Geosciences by Jonathan D. Schueth ©2009 Jonathan D. Schueth Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science December 2009 The thesis of Jonathan D. Schueth was reviewed and approved* by the following: Timothy J. Bralower Professor of Geosciences Head of the Department of Geosciences Thesis Adviser Lee R. Kump Professor of Geosciences Mark E. Patzkowsky Associate Professor of Geosciences Katherine H. Freeman Professor of Geosciences Associate Head of Graduate Programs of the Department of Geosciences *Signatures are on file in the Graduate School ii ABSTRACT The Cretaceous-Paleogene (K/P) bolide impact decimated surface ocean ecosystems. While the extinction has been well studied, there have been few investigations of the recovery of ecosystems from the K/P extinction. The relationship between the post-extinction ecologies of nannoplankton and planktonic foraminifera can lend insight into how ecosystems were rebuilt after the K/P impact. This study utilizes multivariate analysis on abundance counts from five sites with wide global distribution to compare the recoveries of these calcareous plankton groups. The analysis shows there is a distinct geographical differentiation in the recovery of both groups, with assemblages in the northern hemisphere surface oceans diversifying more slowly than southern hemisphere surface oceans. This is most likely a result of higher extinction intensity caused by the concentration of dust created by the bolide impact in the northern hemisphere. The multivariate analysis results suggest a probable long-term influence on the post-extinction marine ecosystem was increased nutrient concentration in the surface ocean resulting from the collapse of the biological pump. Nannoplankton recover before planktonic foraminifera in the open ocean, as expected, but after planktonic foraminifera on the shelf. Dinoflagellates, which were common at many shelf locations across the K/P boundary may have provided another food source for planktonic foraminifera and hindered nannoplankton diversification by invading niche space emptied by the K/P extinction. Ecological recovery of calcareous microplankton took place only after the biologic pump was restored ~300 Kyr after the K/P extinction, removing the enhanced nutrient concentrations from the surface ocean. iii TABLE OF CONTENTS List of Tables…………………………………………………………………………………….. v List of Figures…………………………………………………………………………………… vi Preface...................................................................................................................................…... vii Acknowledgements......................................................................................................................viii MICROPLANKTON ECOSYSTEMS AFTER THE K/P BOUNDARY...…………………….. 1 METHODS………………………………………………………………………………………. 5 Data………………………………………………………………………………………… 5 Brazos River Nannoplankton Counts...…………………………………………………….. 7 Multivariate Analysis………………………………………………………………………. 7 Diversity……………………………………………………………………………………..9 Age Model………………………………………………………………………………….. 9 Paleoecology..…………………………………………………………………………….. 11 RESULTS………………………………………………………………………………………. 12 NMS Results……………………………………………………………………………….12 Diversity and Extinction Intensitiy…………………………………………………………13 DISCUSSION…………………………………………………………………………………... 20 Environmental Stress after the K/P Impact……………………………………………….. 20 Trends in the Recovery of Post K/P Microplankton Ecosystems…………………………. 21 Ecological Recovery after the K/P Extinction……………....……………………………. 23 CONCLUSIONS...............……………......……………………………………...………….......26 References………………………………………………………………………………………. 28 Appendix A: Comprehensive Taxonomy……………………………………………………… 40 Appendix B: Paleoecology Table……………………………………………………………… 52 Appendix C: Supplements to Thesis…………………………………………………………… 59 Detailed NMS Methods……………………………...............…....……………………..60 Age Model ...…..…………………………………………………………………………61 Range/Abundance charts of selected Danian taxa….………….……………………….. 67 iv List of Tables Table 1: Page 6 Location, modern day latitude and longitude and source of data for Shasky Rise, Walvis Ridge, Kerguelen Plateau and El Kef. Table 2: Page 11 Composite section standard ages (after Westerhold et al., 2008) and datums (Martini, 1971; Berggren et al., 1995) used to correlate all sites. v List of Figures Fig. 1 pg. 6 Paleogeographic reconstruction of the Cretaceous-Paleogene boundary (65.5 Ma) showing locations used in the study. Map constructed using software available at http://odsn.de.odsn/services/paleomap/paleomap.html based on paleogreographic reconstructions from Hay et al., 1999. Fig. 2 pg. 14 NMS results (samples) plotted. Plots are coded by time (A: nannoplankton, B: planktonic foraminifera) as well as location (C: nannoplankton, D: planktonic foraminifera) Fig 3 pg. 15 Nannoplankton (A) and planktonic foraminifera (B) applied to the NMS results. Paleocene taxa and survivors are coded by paleoecology. Cretaceous taxa are marked with capital letters. Fig. 4 pg. 16 NMS1 values plotted versus age (in Myr) for nannoplankton (blue) and planktonic foraminifera (red) at all five locations Fig. 5 pg. 17 Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera for Shatsky Rise (ODP Site 1210 and DSDP Site 577) Fig. 6 pg. 17 Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera for Walvis Ridge (ODP Site 1262 and DSDP Site 528) Fig. 7 pg. 18 Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera for El Kef Fig. 8 pg. 18 Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera for Brazos River Fig. 9 pg. 19 Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera for Kerguelen Plateau (ODP Site 738) Fig. 10 pg. 19 Extinction intensity of species over the K/P boundary at all locations with nannoplankton in blue and planktonic foraminifera in red. Error bars were calculated from methods outlined in Raup (1991). At Brazos River, no planktonic foraminiferal samples were counted from the Cretaceous, so extinction intensity was not calculated. vi PREFACE Jonathan Schueth is the first author of this work. This work is coauthored by T. J. Bralower, S. Jiang, M. E. Patzkowsky and L. R. Kump. As the first author, I wrote the text and the associated figures and tables. All of the nannofossil counts and other data collection, multivariate analysis, research done to establish the thesis were done by the primary author. The thoughts in the discussion were primarily ideas of the first author, but were developed further with insight from the other contributing authors. The other authors also edited the manuscript and helped me focus it into a more succint argument. vii AKNOWLEDGEMENTS This work was funded by a NASA “ecoresponse” grant. Primary thanks go to my adviser, T.J. Bralower, my committee members L.R. Kump and M. E. Patzkowsky and S. Jiang for input into the thesis. I would also like to thank T.E. Yancey for providing samples and aid at the Brazos River field site and J. Sessa for help collecting the samples used in this research. I also appreciate M.A. Arthur who gave very valuable insight into the thesis. viii A MULTIVARIATE ANALYSIS OF THE RECOVERY OF CALCAREOUS NANNOPLANKTON AND PLANKTONIC FORAMINIFERA FROM THE CRETACEOUS/PALEOGENE (K/P) MASS EXTINCTION Jonathan D. Schueth, Co authors: Timothy J. Bralower, Shijun Jiang, Mark E.Patzkowsky and Lee R.Kump MICROPLANKTON ECOSYSTEMS AFTER THE K/P MASS EXTINCTION The mass extinction event at the Cretaceous-Paleogene (K/P) boundary (65.68 Ma) devastated oceanic ecosystems. Planktonic protists that thrived in the Cretaceous were almost completely eradicated, with >90% of planktonic foraminifera and calcareous nannoplankton species going extinct across the K/P boundary (Kaiho, 1994; Bown et al., 2004). Oceanic food webs and the biologic pump were disrupted and took millions of years to fully recover (e.g., Zachos and Arthur, 1986; D’Hondt et al., 1998; D’Hondt, 2005). The taxonomic basis of the extinction event has been documented in great detail for both planktonic foraminifera (e.g., Luterbacher and Premoli Silva, 1964; Smit and Hertogen, 1980; Smit, 1982; Keller, 1988; Smit, 1990; Molina 1994, 1995; Molina et al., 1996, 1998) and calcareous nannoplankton (e.g., Bramlette and Martini, 1964; Perch-Nielsen, 1969; Percival and Fischer, 1977; Romein, 1977; Perch-Nielsen et al., 1982; Pospichal, 1994). Post extinction foraminifera were dominated by low-diversity assemblages made up of small taxa with simple morphologies (e.g., Luterbacher and Premoli Silva, 1964; Blow, 1979; Tourmarkine and Luterbacher, 1985). Contemporaneous nannoplankton were dominated by resting cysts likely produced during high environmental stress—such as extremes in salinity or nutrients (e.g., Eshet et al., 1992; Tantawy, 2003). Both planktonic foraminifera and nannoplankton took millions of years to recover from the extinction 1 (Coxall et al., 2006; Fuqua et al., 2008), and diversity did not reach pre-extinction levels for as long as 15 million years after the event (Norris, 2001; Bown et al., 2004). While the extinction has been well documented for both plankton groups, very few quantitative, ecological studies have been performed