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 on the recovery.

The impact of a large bolide on the Yucatan peninsula in Mexico is thought to have

caused the K/P mass extinction (Alvarez et al., 1980; Smit and Hertogen, 1980; Alvarez, 1987;

Erikson and Dickson, 1987; Hildebrand et al., 1991; Smit, 1990; D’Hondt et al., 1994a; Toon et

al., 1997). Environmental effects of the impact led to the mass extinction of planktonic

organisms, and, at the same time, delayed their recovery. The bolide is thought to have struck at

an oblique angle directed towards the north to northwest (Johnson and Hickey, 1990; Alvarez et

al., 1995; Schultz and D’Hondt, 1996), ultimately concentrating ejecta in the northern

hemisphere. Models suggest that the K/P impact did not produce enough dust to cause global

darkness, but ejected sufficient material into the atmosphere to hinder photosynthesis for several

months or years (Toon et al., 1997; Pope, 2002). The impact was also thought to hit a large area

of evaporites which led to surface ocean acidification (Sigurdsson et al., 1991; Brett, 1992;

D’Hondt et al., 1994a; Pope et al., 1997). Global darkness, which lasted at most a few months,

and ocean acidification, which would have been very brief due to the ocean’s buffering capacity

(D’Hondt et al., 1994a), may have led to the mass extinction; however, these factors most likely

would not have had long-term ecological impacts (Pope et al., 1997; Pope, 2002). Another

potentially important result of the impact may have been metal loading of the surface ocean. The

bolide impact may have added toxic levels of metals to the surface oceans (Erickson and

Dickson, 1987). Metal loading could have affected the surface ocean for an extended period of time and delayed ecosystem recovery (Jiang et al., in review).

2 The extinction event had a profound effect on the marine carbon cycle. This is evident

from the collapse of the δ13C gradient between the surface and deep ocean, which suggests a

significantly weakened biologic pump (e.g. Zachos and Arthur, 1986; Stott and Kennett, 1989;

Zachos et al., 1989, 1992; Hilting et al., 2008) and decreased organic matter delivery to the deep ocean (Hsü et al., 1982, Zachos et al., 1989). There were two main recovery intervals in the carbon isotopic gradient after the K/P extinction: a step ~300-500 Kyr after the boundary, when initial recovery of the gradient took place, and a second step ~3 Myr after the extinction, when the gradient reached pre-extinction values (D’Hondt et al., 1998; Adams et al., 2004). The pattern of δ13C collapse and recovery in the surface ocean is seen in foraminifera and fine

fraction globally after the K/P extinction. The final recovery is thought to represent the

evolution of larger plankton or grazers (D’Hondt et al., 1998; D’Hondt, 2005).

The return of diversity and dominance to pre-extinction levels is also a key aspect of the

ecologic recovery of the marine plankton. Jiang et al. (in review) showed there was a distinct geographic differentiation in the recovery of nannoplankton, and proposed that the surface ocean in the northern hemisphere recovered nearly 300 Kyr after recovery in the southern hemisphere.

Nannoplankton did not fully recover in diversity and ecology until the biologic pump was restored 3 Myr after the extinction (Fuqua et al., 2008). In the earliest Paleocene, there were two periods of diversification in the planktonic foraminifera (Coxall et al., 2006). Approximately

60% of total Paleogene diversity was reached 300 Kyr after the extinction, and most Paleocene foraminiferal morphologies evolved by that time. This was followed by another rapid diversification 2-3 million years after the extinction which corresponded to a shift to an assemblage that was oligotrophic and similar in diversity to modern assemblages (D’Hondt et al.,

1994b; Norris, 1996). Both of these events were likely related to the reestablishment of the

3 biological pump (Coxall et al., 2006; D’Hondt, 2005), and the final recovery of planktonic foraminifera occurred at roughly the same time as that of the nannoplankton (Fuqua et al., 2008).

There is some indication that the recovery of planktonic foraminifera was geographically dependant, similar to nannoplankton. Planktonic foraminifera quickly diversified in high latitudes while species richness remained low in low latitudes for several million years (Keller et al., 1993; Barrera and Keller, 1994). However, there has been no comparison on how the recovery of planktonic foraminifera and nannoplankton relate. Theoretically, because nannoplankton are a food source for foraminifera (e.g. Hemleben et al., 1989), foraminiferal diversity should be restored after the return of nannoplankton productivity opens up niche space for the planktonic foraminfera (Solé et al., 2002).

This work expands significantly on the previous work of Jiang et al, who showed there were distinct geographic patterns in the recovery of nannoplankton after the extinction. Whereas

Jiang et al. (in review) focused on the nannofossils, the base of the food web, we compare the recovery of nannoplankton with planktonic foraminifera, thereby expanding the focus to multiple trophic levels. By looking at multiple trophic levels, we can better understand how food webs were rebuilt after the K/P extinction. Therefore, while our work is similar to Jiang et al. (in review), it is a more intricate look into how trophic relationships were rebuilt as the marine ecosystem recovered. Our goal is to utilize multivariate analysis of assemblage data from a transect of latitudes and oceanic environments to better understand how the recovery of nannoplankton and planktonic foraminifera were coupled in the 300 Kyr interval following the

K/P boundary. We constrain the relative timing of the resurgence of nannoplankton and planktonic foraminiferal diversity, and we use the suite of sites to determine how the base of the

marine food web was rebuilt in the wake of the K/P mass extinction.

4 METHODS

Data

We have complied assemblage counts for nannofossils and planktonic foraminifera from

five sites with a wide geographical distribution (Fig. 1; Table 1). These include: Shatsky Rise

(Central Pacific), Walvis Ridge (South Atlantic), Kerguelen Plateau (Indian Ocean), and El Kef,

Tunisia (Tethys Sea). To augment these data, we counted nannofossils from the Cottonmouth

Creek K/P section of the Brazos River, Texas (for a stratigraphic overview of this section, see

Yancey, 1996 and Schulte et al., 2006). Planktonic foraminiferal counts from the Cottonmouth

Creek section were obtained from C. Liu through T. Yancey (unpublished data). Nannofossils and foraminifera in these sections were not sampled at the same levels, and because of this, two separate multivariate analyses were run for each fossil group. These data differ from those used in Jiang et al. (in review) by adding planktonic foraminifera and by adding new counts from the

Brazos River.

A comprehensive taxonomy was established to combine data from multiple authors with differing taxonomic concepts (Appendix A). In most cases, species were combined into genera to facilitate comparison; only those species prevalent at all sites were kept separate.

Early Paleocene nannoplankton samples are dominated by calcispheres (predominately calcareous dinoflagellates) which disaggregate readily and are present almost always as fragments. There is wide discrepancy on how to count calcisphere fragments. Jiang and Gartner

(1986) counted every calcisphere fragment as one specimen, whereas other authors have counted every three fragments greater than eight microns as one individual (e.g. Bernoala and Monechi,

2007). Because in most of our samples, the average size of the calcisphere fragments is around

4-5 microns, one individual was counted out of every three fragments greater than or equal to 5

5 microns. All calcisphere counts from the published data were also corrected to this counting method. Braarudospheres also are found mostly as fragments. For this study, three braarudosphere pentalith segments or one pentalith made from three or more segments were considered one individual braarudosphere.

Fig. 1: Paleogeographic reconstruction of the Cretaceous/Paleogene boundary (65.5 Ma) showing the locations used in the study. Map constructed using software available at http://odsn.de/odsn/services/paleomap/paleomap.html based on paleogeographic reconstructions from Hay et al., 1999.

Site Location ODP/DSDP Latitude Longitude Source of data Site Shatsky Central ODP 1210 32º13’ N 158º16’ E Bown, 2005 Rise Pacific DSDP 577 32º27’ N 157º43’ E D’Hondt and Keller, 1991 Walvis South Atlantic ODP 1262 27º11’ S 1º34’ E Bernoala and Monechi, 2007 Ridge DSDP 528 28º32’ S 2º19’ E D’Hondt and Keller, 1991 Kerguelen Indian Ocean ODP 738 62º43’ S 82º47’ E Huber, 1991; Plateau Pospichal, 1993 El Kef, Tethys Sea N/A 36º11’ N 8º43’ E Pospichal, 1994 Tunisia Arenillas et al., 2000

Table 1: Location, modern day latitude and longitude and source of data for Shatsky Rise, Walvis Ridge, Kerguelen Plateau and El Kef.

6

Brazos River Nannoplankton Counts

The K/P section of Cottonmouth Creek (near Brazos River, Falls County, Texas) was sampled in detail. The Danian section at Cottonmouth Creek was sampled starting at the top of the K/P sedimentary complex thought to be an impact tsunami deposit (Bourgeois et al., 1988;

Yancey, 1996). The lowermost Paleocene of Cottonmouth Creek is typified by calcareous mudstones with intermittent calcareous and iron concretion horizons (a detailed stratigraphy is given in Yancey, 1996 and Schulte et al., 2006). Extensive paleontological, geochemical and sedimentological studies have been done on the Cottonmouth Creek section and the nearby

“Brazos 2” core (Hansen et al., 1987; Hansen et al., 1993a,b; Keller, 1989a,b). Biostratigraphy indicates this section is fairly complete and corresponds to NP1 and lowermost NP2 nannofossil biozones of Martini (1971). Added to the samples from Cottnmouth Creek are nine samples from nearby Frost Bluff (Milam County, Texas) (see Gardner, 1933 and D’Agostino and

Yancey, 1996 for a detailed overview of the stratigraphy of this section). The lower seven meters of the Frost Bluff section contains abundant nannofossils. This section correlates roughly to the top of the Cottonmouth Creek section with some overlap. The sampled section at Frost

Bluff lies entirely in the NP2 biozone. Samples were disaggregated in buffered water and counting slides were produced for each sample using glass beads as a counting standard (after

Okada, 1992).

NMS Analysis

Nonmetric Multidemsional Scaling (NMS) in an ordination method that describes a large, multivariate dataset with a smaller set of variables (or dimensions) using the ranked distance between samples. The use of a ranked distance lessens the biases in the analysis by eliminating

7 many assumptions about the structure of the data (Clarke, 1993; McCune and Grace, 2002).

Many other techniques, such as principal components analysis or detrended correspondence analysis, assume a linear relationship among the datapoints or that distances between samples are related by a set formula. Because NMS avoids these assumptions, it was chosen as the least biased method to extract the structure of the data.

Because most of the data from the early Danian are dominated by a few abundant taxa, the NMS results can be biased. Also, the rarer taxa are useful in making determinations of ecosystem properties. To eliminate the dominance of the abundant taxa in the analysis, the datasets were first standardized by converting number counts into percent abundance for each sample. These percentages were then log transformed. These steps lessen the bias of the abundant post-extinction taxa while still maintaining the underlying structure in the data

(McCune and Grace, 2002). The NMS analysis was run using the Vegan package in R software

(www.r-project.org). Two separate NMS analyses were run on the nannoplankton and planktonic foraminifera datasets, and variance represented in the NMS results was calculated

(Appendix C). Results were then rotated around the origin after the fact to show stronger relationships between the samples and the NMS dimensions. Nannoplankton and plankonic foraminifera taxa were plotted in NMS space in R using the function “wascores,” which is built in the metaMDS algorithm. To understand how ecology factors into the recovery, the paleoecology of each taxa was then color coded. A more detailed outline of the NMS methods is given in Appendix C.

8 Diversity

To visualize the changes in diversity across the recovery interval, several “Whittaker

plots” were constructed by plotting the natural log of each species’ relative percent abundance by

the species rank (Whittaker, 1965). These plots show how each sample’s species richness differs

as well as the evenness of the assemblage. Seven samples were chosen from each site for both nannoplankton and planktonic foraminifera to show how diversity changed throughout the recovery interval. To eliminate possible bias of reworked fossils, all taxa considered “Cretaceous

Taxa” (i.e. Percival and Fischer, 1977; Bown, 2005; Bernoala and Monechi, 2007) were

eliminated from Paleocene\ samples.

To determine how extinction intensity influenced the recovery, the percent of species that

went extinct over the K/P was calculated for all sites. This was done by first selecting the

youngest five samples from the Cretaceous and the oldest five samples of the Paleocene (or 0.5

m above and below the K/P boundary depending on sample spacing). The percent of species that

went extinct was calculated by dividing the number of species that did not cross the K/P boundary in these samples by the number of total Cretaceous taxa. The error of this calculation was determined using the method outlined in Raup (1991).

Age Model

An age model is needed to correlate the different sites and interpret the NMS analysis.

This age model was constructed using established orbital chronologies of Shatsky Rise (ODP

Site 1210) and Walvis Ridge (ODP Site 1262) (Westerhold et al., 2008). These two sites were

correlated to one another using this orbital chronostratigraphy and a composite section was

developed. The holes used for planktonic foraminifera samples at Shatsky Rise (DSDP Site 577)

9 and Walvis Ridge (DSDP Site 528) were then correlated to this composite section based on the biostratigraphy of those sections. Because of their close spatial relation, biostratigraphic datums between ODP Site 1210/DSDP Site 577 and ODP Site 1262/DSDP Site 577 should be practically concurrent, and therefore allow for a precise correlation between the DSDP sections to the composite standard. Well defined orbital chronologies do not exist for Kerguelen Plateau, El

Kef or Brazos River, and so correlating these sites to the composite standard is a problem. The only available correlation method between these sites is biostratigraphy; however, correlating these sites with the composite standard using biostratigraphy relies on the assumption that biostratigraphic events are not time transgressive over great distances. To lessen some of the inaccuracy of this correlation, the precision of the age model at Brazos River, El Kef and

Kerguelen Plateau was kept low. All interpretation comparing the timing of events at these sites must be done with the caveat that the correlations may not be entirely accurate. To correlate these sites with the composite standard, key nannofossil and planktonic foraminiferal datums

(Martini, 1971; Berggren et al., 1995) were tied to the composite standard, and their ages were determined (Table 2) and correlated between Brazos River, El Kef, and Kerguelen Plateau. By assuming that sedimentation rates were constant between these tie points (which also adds to the inaccuracy of the correlation), age models were established for these three sites. An extended and more detailed method of the construction of this age model is in Appendix C

10 CS Age (Ma) Event (after Westerhold et al., 2008) K-P Boundary 65.680 FO Pv. eugubina (P0/Pα) 65.66 LO Pv. eugubina (Pα/P1a) 65.63 FO S. triloculinoides(P1a/P1b) 65.51 FO P. inconstans (P1b/P1c) 65.34 FO C. intermedius (NP1/NP2) 65.32

Table 2: Composite section standard ages (after Westerhold et al., 2008) and datums (Martini, 1971; Berggren et al., 1995) used to correlate all sites.

Paleoecology Another important part of understanding how the NMS results relate to the ecological

recovery is to understand the paleoecology of the organisms used in this study. Some taxa, such

as Braarudosphaera bigelowii, are extant and their ecologies are at least partially known (Kelly

et al., 1993). For the extinct taxa, other methods are needed to estimate paleoecology. One way

of determining paleoecology is based on a taxon’s paleogeographic distribution. This has been

done to estimate the paleoecology of several Cretaceous-aged nannofossils. Taxa that are constrained to areas thought to represent high rates of upwelling, where waters are often eutrophic, are considered as eutrophic taxa (e.g. Roth and Bowdler, 1981; Thierstein, 1981;

Watkins, 1989). Taxa that are commonly found in the open ocean gyres, which today are characterized by low-nutrient levels (Hallock, 1987), are thought to be oligotrophic (i.e.

Thierstein, 1981). In some cases, extant species with similar morphologies were used to determine the paleoecology of extinct taxa, as was the case for Guembelitria cretacea (Keller et al., 2002). Ecology can also be determined through overall skeleton size. Because smaller cells are more efficient at taking up nutrients, they are most likely to be prevalent in eutrophic environments (Hallock, 1987; Reynolds, 2006). All of these factors were combined and used to

11 determine the overall paleoecology of the taxa used in the NMS analysis. A more complete

overview of this process, along with a table showing the paleoecologic affinity of each taxon is

given in Appendix B.

RESULTS

NMS Results

NMS results are shown for samples coded both by time (Fig. 2 a, b) and by geographic

location (Fig. 2 c, d). The variance represented in the nannoplankton analsyis was 54.2% for

NMS axis 1 (NMS1) and 22.1% for NMS axis 2. The variance represented in the foraminifera

was 66.1% for NMS1 and 14.9% for NMS2. The plot of how the taxa plot in NMS space is

shown in Fig. 3. To show how NMS1 changes relate between nannofossils and planktonic

foraminifera, as well as show how they change through time, NMS1 has been plotted against

time for each section (Fig. 4)

Most stress-tolerant disaster taxa (such as Guembelitria and Braarudosphaera) plot with

negative first axis (NMS1) values, and “recovery taxa” (i.e. Cr. intermedius and Pr. inconstans) plot with more positive NMS1 values (Fig. 3). This indicates that NMS1 is a measure of environmental stress with high stress related to negative NMS1 values, and low stress related to high NMS1 values. There is also a distinct relation between the second NMS axis (NMS2) and the age of the sample (Fig. 2 a, b). Older samples correspond to low NMS2 values, while younger samples correspond to high NMS2 values.

12 Diversity and Extinction Intensity

The Whittaker Plots are shown in Figs. 5-9. For each plot the log of the relative

abundance of each species is plotted against species rank (with most abundant species plotted

first, least abundant plotted last). In each plot, the black line corresponds to the base measure of

diversity in the youngest Cretaceous sample. Each subsequent colored line corresponds to a

certain age (Shatsky Rise and Walvis Ridge) or, where the age model is not as accurate, a depth

(Brazos River, El Kef, and Kerguelen Plateau). Also, for an indication of how diversity changes

through time, the range and abundance of all Paleocene taxa was compiled in Appendix C.

Extinction intensities are shown in Figure 10. The nannoplankton show some differences

with northern hemisphere sites (Brazos River, El Kef, and Shatsky Rise) having higher

extinction intensities than the southern hemisphere sites, the differences are often within the error

of the other sites. Likewise, there seems to be no significant difference between locations for the planktonic foraminifera, although the error is large for the planktonic foraminifera because most

of the datasets used do not have very high species richness in the Cretaceous.

13

Fig. 2: NMS results (samples) coded by time (A: nannoplankton, B: planktonic foraminifera) and location (C: nannoplankton, D: planktonic foraminifera)

14

Fig. 3: 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

15

nd planktonic foraminifera (red) at all

five locations five locations Fig. 4: NMS1 values plotted versus age (in Myr) for nannoplankton (blue) a

16

Fig. 5: Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera at Shatsky Rise (ODP Site 1210 and DSDP Site 577)

Fig. 6: Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera at Walvis Ridge (ODP Site 1262 and DSDP Site 528)

17

Fig. 7: Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera at El Kef

Fig. 8: Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera at Brazos River

18

Fig. 9: Log relative abundance vs. species rank for nannoplankton and planktonic foraminifera at Kerguelen Plateau (ODP 738)

Fig. 10: 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.

19 DISCUSSION

Environmental Stress after the K/P Impact

The key to understanding the recovery of oceanic food webs following the K/P extinction

is to constrain the primary stress on the post-extinction ecosystems. Ocean acidification and

heavy metal loading were possible environmental stresses caused by the K/P impact. Sulfide

aerosols ejected into the atmosphere could have acidified surface oceans (Sigurdsson et al., 1991;

Brett, 1992; D’Hondt et al., 1994a; Pope et al., 1997), but ocean acidification may have been

brief due to the surface ocean’s buffering capacity (D’Hondt et al., 1994a). Another possible

source of environmental stress would be loading of metals derived from the bolide (Erickson and

Dickson, 1987). Metals concentrations toxic to plankton could have been maintained for

thousands of years due to the collapse of the biological pump (Jiang et al., in review). Enhanced

concentrations of metals (Cd, Cu, Ni, etc.) are known to diminish modern phytoplankton

productivity by either disrupting reproduction (Brand et al., 1983) or by blocking uptake of

necessary nutrients (Sunda and Huntsman, 1998). Metal loading is difficult to prove as toxic

levels are below what can be detected in algal skeletal materials.

Post-extinction assemblages have lower diversity than Cretaceous assemblages, and most

species that dominate the earliest Paleocene are small or stress-tolerant, shelf-dwelling taxa

(Bown, 2005; D’Hondt, 2005; Fig. 4). Paleocene assemblages are similar to those found in

modern eutrophic environments which are typified by low diversity assemblages dominated by

small taxa (Hallock, 1987). Globally, post K/P extinction assemblages can be considered

eutrophic, and many of the bloom-taxa (i.e. the nannoplankton Braarudosphaera and the

planktonic foraminifer Guembelitria) in the Cretaceous were only abundant on the shelf (Keller et al., 2002; Bown, 2005). Adding to the evidence of increased eutrophy in the early Paleocene

20 surface ocean is the collapse of the carbon isotope gradient following the K/P boundary which

indicates the breakdown of the biological pump (D’Hondt et al., 1998; D’Hondt, 2005). All of

this evidence implies that NMS1 is a measure of nutrient stress in the post-extinction

ecosystems.

While NMS1 may be an indication of nutrient stress on post-impact ecosystems, it is not

necessarily a measure of nutrient concentration. Kerguelen Plateau (ODP Site 738) samples

consistently plot with highly positive, or low stress, NMS1 values (Fig. 3) despite the abundance

of eutrophic taxa (such as the nannoplankton Biscutum, Hornibrookina, and Zeugrhabdotus).

High latitude environments are often eutrophic (Hallock, 1987), so the ecosystem at Kerguelen

Plateau most likely would have been adapted to cope with a global increase in nutrient

concentration. However, open ocean ecosystems, such as at Shatsky Rise, which are typically

oligotrophic, would have been greatly impaired by an increase in nutrients. Nutrient stress

would have decreased only after the biological pump began to recover (~300 Kyr after the K/P)

(D’Hondt, 2005).

Trends in the Recovery of Post K/P Microplankton Ecosystems

There is a strong geographic trend in the recovery of both nannoplankton and planktonic

foraminifera. Nannoplankton in the northern hemisphere sites (Shatsky Rise, El Kef, and Brazos

River) have highly negative NMS1 values (Fig. 3) and are dominated by disaster-bloom taxa

(Fig. 4). However, disaster taxa are rare in the southern latitude sites (Walvis Ridge and

Kerguelen Plateau), and these sites have positive NMS1 values. This is an indication that the

ecosystems in the northern latitudes were more influenced by the post-impact environmental

stresses than those in the southern hemisphere. This trend was noted by Jiang et al. (in review),

21 and thought to be a result of the initial concentration of the K/P bolide ejecta in the northern hemisphere. Building upon the result of Jiang et al. (in review), we show that similar trends exist for the planktonic foraminifera, however the planktonic foraminifera at Walvis Ridge are more like those in northern hemisphere sites in both the relative abundance of taxa and NMS1 values after the extinction (Fig. 3). Comparing trends in diversity between sites may provide insight into the cause of this geographic difference.

Based on the Whittaker plots, nannoplankton diversity (species richness and eveness) drops across the boundary in the northern hemisphere sites and stays at this low point for ~200

Kyr (Figs. 6, 8-9). However, eveness quickly returns to pre-extinction values at both Walvis

Ridge and Kerguelen Plateau (Figs. 7, 10), consistant with the interpretation of a quicker recovery in the southern hemisphere. At Kerguelen Plateau, planktonic foraminifera eveness quickly rebounds (within 30 Kyr, Fig. 10), which is similar to the nannoplankton. Unlike the rapid diversification of nannoplankton at Walvis Ridge, planktonic foraminifera have a subdued species richness and eveness until sometime between 24 to 141 kyr after the extinction (Fig. 6).

At El Kef, planktonic foraminiferal eveness recovers almost instantaneously compared to nannoplankton (Fig. 7). These comparisons indicate a distinct difference in the recovery of nannoplankton and planktonic foraminifera. However, it is impossible to interpret changes in planktonic foraminiferal diversity through time at all sites because no Cretaceous samples were counted at Brazos River and dissolution may have lowered Cretaceous diversity at Shatsky Rise

(D’Hondt and Keller, 1991).

The difference in diversification between nannoplankton and planktonic foraminifera is primarily between the open ocean and shelf sections. NMS1 values return to pre-extinction levels for nannoplankton before planktonic foraminifera at Shatsky Rise and Walvis Ridge,

22 whereas this trend is the opposite for El Kef and Brazos River (Fig. 5). At Shatsky Rise,

mesotrophic nannoplankton, such as Coccolithus, Cruciplacolithus, and Prinsius (Fuqua et al.,

2008) become abundant approximately 100 Kyr before the mesotrophic planktonic foraminifera

Preamurica inconstans (Coxall et al., 2006). However, at El Kef, P. inconstans becomes abundant ~200-300 Kyr before the mesotrophic nannoplankton. This trend suggests that geographical distribution and paleoenvironment affected the recovery of planktonic ecosystems after the K/P boundary.

Ecological Recovery after the K/P Extinction

The Chicxulub crater and the distribution of ejecta shows evidence of an oblique, south to north, impact (Johnson and Hickey, 1990; Alvarez et al., 1995; Schultz and D’Hondt, 1996).

Thus, atmospheric circulation likely would have focused most of the fallout in the northern hemisphere. This is possibly the cause of the apparent higher extinction intensity of nannofossils in the northern hemisphere (Fig. 10; Jiang et al., in review). Earliest Paleocene nannoplankton ecosystems were also different in the northern and the southern hemispheres. Northern hemisphere nannoplankton assemblages were dominated by Neobiscutum, a small, weakly calcified taxon that represents a highly eutrophic environment (see Appendix B) whereas the southern hemisphere sites contained a more diverse assemblage of the larger eutrophic taxa

Biscutum, Cyclagelosphaera, and Markalius. Nannoplankton in modern eutrophic environments are dominated by the small taxa Emiliania huxleyi (e.g. Winter, 1985; Ziveri et al., 1995; Tyrell and Merico, 2004). Similar to the E. huxleyi, the dominance of Neobiscutum in the northern hemisphere may indicate a higher nutrient concentration than in the southern hemisphere where the early Paleocene assemblage was more diverse. It is possible that the biologic pump became

23 less efficient at drawing nutrients out of the surface ocean more so in the northern hemisphere as

a result of the impact. This may have resulted in a higher nutrient concentration in the northern

hemisphere. However, the carbon isotope gradient collapse, at least right at the K/P boundary, is

globally similar (i.e. Zachos et al., 1985; Keller and Lindinger, 1989; Barrera and Keller, 1990;

Stott and Kennett, 1990; Barrera and Keller, 1994; D’Hondt et al., 1998) making it difficult to

prove that the biologic pump was disrupted more in the northern hemisphere.

Because the nannoplankton extinction intensity was higher in the northern hemisphere,

there were little to no eutrophic survivors that could have thrived in a high nutrient, post- extinction environment. Therefore, upon its evolution in the earliest Paleocene, Neobiscutum was able to quickly radiate into the eutrophic environment with little competition from other high-nutrient tolerant nannoplankton. However, because of the survivorship of many eutrophic taxa in the southern hemisphere, niche space was quickly filled, and Neobiscutum was not able to compete with an already established eutrophic ecosystem. This explains the difference in recovery between northern and southern hemispheres without invoking a difference in nutrient concentrations.

Unlike the nannoplankton, the planktonic foraminiferal extinction intensities were globally high (>90%) and nearly uniform (Fig. 10). Post-extinction assemblages are dominated by Guembelitra (a survivor) and Woodrigina, a likely descendent of Guembelitria (Olsson et al.,

1999), both of which are small taxa with simple morphologies indicative of a eutrophic environment. These taxa are abundant almost globally after the K/P boundary (the exception being Kerguelen Plateau). Very low planktonic foraminiferal survival rates opened up vast niche space, but did not provide a diverse eutrophic survivor assemblage like that of the southern hemisphere nannoplankton. Without a diverse assemblage of eutrophic survivors, Guembelitria

24 and its descendents quickly filled global niche space and dominated until the biological pump began to recover (Coxall et al., 2006).

At both Shatsky Rise and Walvis Ridge, nannoplankton diversify before planktonic foraminfera (Fig. 5, 6). Mesotrophic nannoplankton also evolve earlier than mesotrophic planktonic foraminifera. This implies that in the open ocean, primary producers recovered before heterotrophs, which agrees with predictions from ecological modeling (Solé et al., 2002).

Once nutrient concentration began to decrease through the restoration of the biologic pump, the eutrophic nannoplankton were outcompeted by a more diverse, mesotrophic assemblage. This in turn led to the diversification of planktonic foraminifera as more niche space was opened and oceanic food webs began to be rebuilt.

On the shelf (El Kef and Brazos River), however, planktonic foraminifera recovered in diversity before nannoplankton. This is the exact opposite of model predictions (Solé et al.,

2002), and may imply the nannoplankton were not the only food source for planktonic foraminifera on the shelf. Nearly all dinoflagellates survived the K/P mass extinction because they have resting stages that offer a refuge during times of high environmental stress (e.g.,

Wendler and Williams, 2002). Dinoflagellates are common primarily on the shelf over the K/P boundary (Brinkhuis and Leereveld, 1988; Brinkhuis and Zarchariasse, 1988; Eshet et al., 1992;

Moshkovitz and Habib, 1993; Brinkhuis et al., 2000), and are modern food sources for many planktonic foraminifera (Zucker, 1973; Hemleben et al., 1989). Therefore, they could have been a primary food source for shelf planktonic foraminifera in the Cretaceous and early Paleocene.

With low extinction rates, dinoflagellates could have quickly filled the niche space emptied by the extinction of nannoplankton. This would have quickly rebuilt the base of the foodweb on the shelf, and allowed the planktonic foraminifera to diversify. If the dinoflagellates invaded niche

25 space previously occupied by nannoplankton, they could also have hindered the recovery of the nannoplankton. Dinoflagellates may also have thrived in a time of higher nutrients following the

K/P boundary because they are more abundant than nannoplankton in modern high-nutrient environments (Hallock, 1987). However, even with a global increase in nutrient concentration, dinoflagellates failed to radiate into the open ocean, as evidenced by the lack of organic-walled dinocysts in open ocean sites. It is possible that dinoflagellates were not able to readily move into the open ocean. Modern dinoflagellates are known to be tied to very specific environments, and transport in the oceans is thought to be minor (Marret and Zonneveld, 2003). Calcareous dinoflagellages (i.e. Thoracosphaera) are present in the open ocean, but we speculate that they were not a viable food source for the planktonic foraminifera, explaining why there was no rapid diversification of planktonic foraminifera in the open ocean.

CONCLUSIONS

The primary influence on the ecological recovery following the K/P bolide impact was an increased nutrient concentration in the surface ocean. This was caused by the collapse of the biological pump and would have delayed ecosystem recovery. Ecological modeling of ecosystem recovery after mass extinction suggests nannoplankton (autotrophs) should diversify before planktonic foraminifera (heterotrophs) as food webs are rebuilt (Solé et al., 2002). In the open ocean, this is the case, whereas on the shelf planktonic foraminifera recovered before nannoplankton. This is most likely due to increased survivorship of dinoflagellates or other primary producers on the shelf, which opened up more niche space for the planktonic foraminifera and allowed them to rapidly diversify after the extinction.

26 There were many factors involved in reconstructing ecosystems after the K/P mass

extinction. Relative distance from the K/P impact appears to influence the length of recovery with sites in the northern hemisphere taking longer to recover in diversity and ecology after the extinction. This was most likely a result of higher extinction intensities caused by the concentration of impact debris in the northern hemisphere. Ecosystems only began to recover after the biological pump was restored and nutrients were removed from the surface ocean. This allowed nannoplankton, which prefer low nutrient environments, to diversify. Niche space in turn was opened up for the planktonic foraminifera, and the oceanic ecosystems were finally rebuilt. The recovery of marine food webs from the K/P mass extinction was therefore directly influenced by the increase of nutrients in the surface ocean as a result of the disruption of the biological pump.

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39

Appendix A

Comprehensive Taxonomy Used in NMS Analysis

40 COMPREHENSIVE TAXONOMY Organized by generic groups used in the NMS analysis

Nannoplankton

Acturius A. scouts (Risatti, 1973) Wind and Wise in Wise and Wind, 1977

Ahmuellerella A. octoradiata (Górka, 1957) Reinhardt, 1966 A. regularis (Górka, 1957) Reinhardt and Górka, 1967

Arkhangelskiella A. confusa Burnett, 1998 A. cymbiformis Vekshina, 1959

Biantholithus B. hughesii Varol, 1989 B. sparsus Bramlette and Martini, 1964

Biscutum B. castorum Black in Black and Barnes, 1959 B. constans (Górka, 1957) Black in Black and Barnes, 1959 B. coronum Wind and Wise in Wise and Wind, 1977 B. magnum Wind and Wise in Wise and Wind, 1977 B. sp. cf. B. ellipticum (Górka, 1957) Grün in Grün et al., 1974 B. sp. cf. B. hattneri Wise, 1983 B. harrisonii Varol, 1989

Braarudosphaera B. alta Romein, 1979 B. bigelowii (Gran and Braarud, 1935) Deflandre, 1947

Chiastozygus Ch. amphipons (Bramlette and Martini, 1964) Gartner, 1968 Ch. fessus Stover, 1966 Ch. litterarius (Górka, 1957) Manivit, 1971 Ch. ultimus Perch-Nielsen, 1981 Ch. spp.

Coccolithus C. pelagicus (Wallich, 1871) Schiller, 1930 =C. cavus Hay and Mohler, 1967 = Ericsonia cava C. robustus Bramlette and Sullivan, 1961 = Ericsonia robusta

41 C. subpertusus (Hay and Mohler, 1967) van Heck and Prins, 1987 = Ericsonia subpertusa

Cribrosphaerella C. daniae Perch-Nielsen, 1973 C. ehrenbergii (Arkhangelsky, 1912) Deflandre in Piveteau, 1952

Cruciplacolithus Cr. asymmetricus van Heck and Prins, 1987 Cr. edwardsii Romein, 1979 Cr. intermedius van Hck and Prins, 1987 = Cr. tenuis s.l. Cr. primus Perch-Nielsen, 1977 Cr. zachosii Bernoala and Monechi, 2007 Cr. tenuis s.s. (Stradner, 1961) Hay and Mohler in Hay et al., 1967

Cyclagelosphaera C. alta Perch-Nielsen, 1979 C. margerelii Noël, 1965 C. reinhardtii (Perch-Nielsen, 1968) Romein, 1977 C. tabulata (Grün and Zweili, 1980) Cooper, 1987

Eiffelithus E. eximius (Stover, 1966) Perch-Nielsen, 1968 E. gorkae Reinhardt, 1965 E. parallelus Perch-Nielsen, 1973 E. turriseiffelii (Deflandre in Deflandre and Fert, 1954) Reinhardt, 1965

Futyania F. petalosa (Ellis and Lohmann, 1973) Varol, 1989

Hornibrookina H. apellanizii Bernoala and Monechi, 2007 H. spp

Kamptnerius K. magnificus Deflandre, 1959

Lithraphidites L. carniolensis Deflandre, 1963 L. grossopectinatus Bukry, 1969 L. praequadratus Crux, 1981 L. quadratus Bramlette and Martini, 1964

Markalius M. apertus Perch-Nielsen, 1979 M. astroporus (Stradner, 1961) Hay and Mohler in Hay et al., 1967

42 M. inversus (Deflandre in Deflandre and Fert, 1954) Bramlette and Martini, 1964 M. panis Jiang and Gartner, 1986 = Biscutum constans M. walvisensis Bernoala and Monechi, 2007

Micula M. concava (Stradner in Martini and Stradner, 1960) Verbeek, 1976 M. decussata Vekshina, 1959 =M. staurophora M. murus (Martini, 1961) Bukry, 1973 M. prinsii Perch-Nielsen, 1979 M. swastica Stradner and Steinmetz, 1984

Neobiscutum N. parvulum (Romein, 1979) Varol, 1989 N. romeinii (Perch-Nielsen, 1981) Varol, 1989

Neochiastozygus N. modestus Perch-Nielsen, 1971 N. primitivus Perch-Nielsen, 1981 N. “asymmetrical” N. spp.

Neocrepidolithus N. cohenii (Perch-Nielsen, 1968) Perch-Nielsen, 1984 N. cruciatus (Perch-Nielsen, 1979) Perch-Nielsen, 1981 N. dirimosus (Perch-Nielsen, 1979) Perch-Nielsen, 1981 N. fossus (Romein, 1977) Romein, 1979 N. neocrassus (Perch-Nielsen, 1968) Romein, 1979 N. spp.

Nephrolithus N. corystus Wind, 1983 N. frequens Górka, 1957

Octolithus O. multiplus (Perch-Nielsen, 1973) Romein, 1979

Prediscosphaera P. arkhangelskyi (Reinhardt, 1965) Perch-Nielsen, 1984 P. cretacea (Arkhangelsky, 1912) Gartner, 1968 P. grandis Perch-Nielsen, 1979 P. honjoi Bukry, 1969 P. intercisa (Deflandre in Deflandre and Fert, 1954) Shumenko, 1976 P. majungae Perch-Nielsen, 1973 P. stoveri (Perch-Nielsen, 1968) Shafik and Stradner, 1971 P. spinosa Bramlette and Martini, 1964

43

Prinsius P. dimorphosus (Perch-Nielsen, 1969) Perch-Nielsen, 1977 P. cf. P. tenuiculum Okada and Thierstein, 1979 “ Prinsius sp. (small)”

Thoracosphaera (Calcispheres) T. imperforata Kamptner, 1952 = T. hemi T. operculata Bramette and Martini, 1964 T. saxea Stradner, 1961 T. spp.

Watznaueria W. barnesae (Black, 1959) Perch-Nielsen, 1968 W. biporta Bukry, 1969 W. coronata (Gartner, 1968) Bukry, 1969 W. fossacincta (Black, 1971) Bown in Bown and Cooper, 1989 W. manivitae Bukry, 1973

Zeugrhabdotus Z. bricresenticus (Stover, 1966) Burnett in Gale et al., 1996 = Glaukolithus bicrescenticus Z. embergeri (Noël, 1958) Perch-Nielsen, 1985 Z. sigmoides (Bramlette and Sullivan, 1961) Bown and Young, 1977 = Zygodiscus sigmoides; Placozygus sigmoides Z. spiralis (Bramlette and Martini, 1964) Burnett, 1998 = Zygodiscus spiralis Z. spp.

Other Paleocene Taxa Cyclolithella spp. Goniolithus fluckigeri Deflandre, 1957 Lanternithus duocavus Locker, 1967 Holococcolithus spp. Lapideacassis sp. Black, 1971 Micrantholithus aequalis Sullivan, 1964 Micrantholithus fornicatus Martini, 1961 Micrantholithus sp. cf. M. flos Deflandré in Deflandré and Fert, 1954 Micrantholithus spp. Nodosella? spp.

Other Cretaceous Taxa Amphizygus spp. Bukry, 1969 Axopodorhabdus dietzmannii (Reinhardt, 1965) Wind and Wise, 1983 Broinsonia spp. Bukry, 1969

44 Calculites spp. Prins and Sissingh in Sissingh, 1977 = Phanulithus spp. Calculites obscurus (Deflandre, 1959) Prins and Sissingh in Sissingh, 1977 = Phanulithus obscurus Ceratolithoides spp. Bramlette and Martini, 1964 Ceratolithoides aculeus (Stradner, 1961) Prins and Sissingh in Sissingh, 1977 Ceratolithoides kamptneri Bramlette and Martini, 1964 Corollithion completum Perch-Nielsen, 1973 Corollithion completum Perch-Nielsen, 1973 Corollithion exiguum Stradner, 1961 Corollithion kennedyi Crux, 1981 Corollithion madagaskarensis Perch-Nielsen, 1973 Corollithion rhombicum (Strander and Adamiker, 1966) Bukry, 1969 Cretarhabdus conicus Bramlette and martini, 1964 Cribrocorona gallica (Stradner, 1963) Perch-Nielsen, 1973 =Cribrosphaerella gallica Cyclolithus sp. Kamptner, 1948 Cylindralithus echinus Bukry, 1975 = Cribrocorona echinus Lees and Bown, 2005 Cylindralithus nudus Bukry, 1969 Cylindralithus serratus Bramlette and Martini, 1964 Cylindralithus spp. Discorhabdus ignotus (Górka, 1957) Perch-Nielsen, 1968 Discorhabdus spp. = Bidiscus spp. Gartnerago obliquum (Stradner, 1963) Noël, 1970 Gartnerago spp. Gephyrorhabdus coronadventis (Reinhardt, 1966) Hill, 1976 Gorkaea pseudoanthophorus (Bramlette and Martini, 1964) Varol and Girgis, 1994 Gorkaea spp. Varol and Girgis, 1994 Helicolithus anceps (Górka, 1957) Noël, 1970 Helicolithus cf. H. trabeculatus (Górka, 1957) Verbeek, 1977 Loxolithus armilla (Black in Black and Barnes, 1959) Noël, 1965 Lucianorhabdus cayeuxii Deflandre, 1959 Manivitella pemmatoidea (Deflandre and Manivit, 1965) Thierstein, 1971 Microhabdulus attenuatus (Deflandre, 1959) Deflandre, 1963 Microhabdulus belgicus Haye and Towe, 1963 Microhabdulus decoratus Deflandre, 1959 Microhabdulus undosus Perch-Nielsen, 1973 ?Munarinus sp. Risatti, 1973 Parhabdolithus plebeius (Perch-Nielsen, 1973) Crux, 1982 Parhabdolithus spp. Deflandre, 1952 Perchnielseella stradneri (Perch-Nielsen, 1973) Watkins in Watkins and Bowlder, 1984 Placozygus fibuliformis (Reinhardt, 1964) Hoffmann 1970 Podorhabdus elkefensis Perch-Nielsen, 1981

45 Pontosphaera spp. Lohmann, 1902 = Discolithus spp. Huxley, 1868 Prolatipatella multicarinata Gartner, 1968 Pseudomicula quadrata Perch-Nielsen in Perch-Nielsen et al., 1978 Quadrum bengalensis Burnett, 1998 Quadrum gartneri Prins and Perch-Nielsen et al., 1978 Quadrum sissinghii Perch-Nielsen, 1984 Repagalum sp. cf. R. parvidentatum (Deflandre and Fert, 1954) Forchheimer, 1972 Retecapsa angustiforata Black, 1971 Retecapsa crenulata (Bramlette and martini, 1964) Grün in Grün and Allemann, 1975 Retecapsa schizobrachiata(Gartner, 1968) Grün in Grün and Allemann, 1975 = Cretarhabdus schizobraciatus Retecapsa surriella (Deflndre and Fert, 1954) Grün in Grün and Allemann, 1975 Retecapsa ficula (Stover, 1966) Burnett, 1998 Rhagodiscus angustus (Stradner, 193) Reinhardt, 1967 = Parhabdolithus angustus Rhagodiscus indistinctus Burnett, 1998 Rhagodiscus reniformis Perch-Nielsen, 1973 = Parhabdolithus reniformis Rhagodiscus splendens (Deflandre, 1953) Verbeek, 1977 = Parhabdolithus splendens Rhombolithion rhombicum (Stradner and Adamiker, 1966) Black, 1973 Rotelapillus munitus Noël, 1973 Rucinolithus magnus Bukry, 1975 Rucinolithus spp. Stover, 1966 Scapholithus fossilis Deflandre in Deflandre and Fert, 1954 Scapholithus spp. Semiholithus bicomis Perch-Nielsen, 1973 Semiholithus priscus Perch-Nielsen, 1973 Sollasites spp. Black, 1967 Staurolithites spp. Caratini, 1963 = Vekshinella spp. Loeblich and Tappan, 1963 Staurolithites laffittei Caratini, 1963 Staurolithities mielnicensis (Górka, 1957) Perch-Nielsen, 1968 Staurolithites crux (Deflandre and Fert, 1954) Caratini, 1963 = Vekshinella crux Staurolithites aachenus (Buckry, 1969) Burnett, 1998 = Vagalapilla aachena Stoverius cf. S. conangustatus Howe et al. in Howe, 2003 Stradnerlithus regularis Perch-Nielsen, 1985 = Actinozygus regularis (Górka) Gartner, 1986 Stradnerlithus sp. = Actinozygus sp. Teumentum stradneri Thierstein in Roth and Thierstein, 1972 Teichorhabdus ethmos Wind and Wise in Wise and Wind, 1977 Tetrapodorhabdus decorus Deflandre in Deflandre and Fert, 1954

46 Tranolithus macleodiae (Bukry, 1969) Perch-Nielsen, 1984 Tranolithus minimus (Bukry, 1969) Perch-Nielsen, 1985 Uniplanarius trifidius Stradner in Stradner and Papp, 1961

Planktonic Foraminifera

Archaeoglobigerina A. australis Huber, 1990 A. blowi Pessagno, 1967 A. cretacea d’Orbigny, 1980 A. maleola Huber, 1990

Chiloguembelina Ch. midwayensis Cushman, 1940 Ch. morsei Kline, 1943 Ch. taurica Morozova, 1961 Ch. waiparensis Jenkins, 1965 = Zeauvigerina waiparensis Eoglobigerina E. edita Subbotina, 1953 = E. pentagona; E. polycamera E. eobulloides Morozova, 1959 = E. firnga E. praeedita Blow, 1979 E. simplicissima Blow, 1979

Globanomalina G. archeocompressa Blow, 1979 G. imitata Subbotina, 1953 G. planocompressa Shutskaya, 1965 = Planorotalites planocompressa

Globigerinelloides G. alvarezi Eernod Olvera, 1959 G. multispinus Lalicker, 1948 G. prairiehellensis Pessagno, 1967 G. rosebudensis Smith and Pessagno, 1973 G. subcarinatus Broennimann, 1952 G. ultramicrus Subbotina, 1949 G. volutus White, 1928 G. yaucoensis Pessagno, 1960

47 Globoconusa G. alticonsua Li et al., 1995 = Parvularugoglobigerina alabamensis G. daubjergensis Broenimann, 1953 = Globastica daubjergensis G. extensa Blow, 1979 G. fodina Blow, 1979 G. hillebrandti G. minulata Luterbacher and Premoli Silva, 1964 G. tripartita Morozova et al., 1967

Globotruncana G. aegyptiaca Nakkady, 1950 G. arca Cushman, 1926 G. mariei Banner and Blow, 1960 G. orientalis El Nagger, 1966 G. rosetta Carsey, 1926

Globotruncanella G. caravacensis Smit, 1982 G. citae Bolli, 1951 G. havanensis Voorwijk, 1937 G. minuta Caron and Gonalez Donoso, 1984 G. petaloidea Gandolfi, 1955 G. pschadae Keller, 1946

Globotruncanita G. angulata Tiley, 1951 G. conica White, 1928 G. dupeublei Caron et al., 1984 G. fareedi El Naggar, 1996 G. insignis Gandolfi, 1955 G. stuarti de Lapparent, 1918 G. stuartiformis Dalbiez, 1955

Guembelitria G. alabamensis Liu and Olsson, 1992 = Parvularugoglobigerina alabamensis G. cretacea Cushman, 1933 G. danica Hofker, 1978 = Chiloguembelitria danica G. irregularis Morozova, 1961 G. trifolia (Blow, 1979) Keller, 1989

48 Hedbergella H. homdelensis Olsson, 1964 H. monmouthensis Olsson, 1960 H. sliteri Huber, 1990

Heterohelix H. dentata Stenestad, 1968 H. glabrans Cushman, 1938 H. globulosa Ehrenberg, 1840 H. labellosa Nederbragt, 1991 H. navarroensis Loeblich, 1951 H. planata Cushman, 1938 H. pulchra Brotzen, 1936 H. punctulata Cushman, 1938 H. semicostata Cushman, 1938 H. striata Ehrenberg, 1840

Parasubbotina P. moskvini Shutskaya, 1953 = Morozovella moskvini P. pseudobulloides Plummer, 1926 = Morozovella pseudobulloides P. varianta Subbotina, 1953

Parvularugoglobigerina Pv. alabamensis Liu and Olsson, 1992 Pv. eugubina Luterbacher and Premoli Silva, 1964 = Pv. longiapetura Blow, 1979 Pv. extensa Blow, 1979 = Globoconusa extensa Pv. perexigua Li et al., 1995 Pv. umbrica Luterbacher and Premoli Silva, 1964

Planoglobulina Pl. acervulinoides Cushman, 1927 Pl. brazoensis Martin, 1972 Pl. carseyae Plummer, 1931 Pl. manuelensis Martin, 1972 Pl. multicamerata Klasz, 1953 Pl. hankteninoides Broennimann, 1953

Preamurica Pr. inconstans Subbotina, 1953 = Morozovella inconstans Pr. pseudoinconstans Blow, 1979 Pr. taurica Morozova, 1961 = “Eoglobigerina” taurica

49 Pseudoguembelina Ps. costellifera Masters, 1976 Ps. costulata Cushman, 1938 Ps. excolata Cushman, 1926 Ps. harianensis Nederbagt, 1991 Ps. kempensis Esker, 1968 Ps. palpebra Broennimann and Brown, 1953

Pseudotextularia P. elegans Rzehak, 1891 P. intermedia Klasz, 1953 P. nutalli Voorwijk, 1937

Racemiguembelina R. fucticosa Egger, 1902 R. intermedia De Klasz, 1953 R. powelli Smith and Pessagno, 1973

Rugoglobigerina Ru. hexcamerata Broennimann, 1952 Ru. macrocephala Broennimann, 1953 Ru. milamensis Smith and Pessagno, 1973 Ru. pennyi Broennimann, 1952 Ru. reicheli Broennimann, 1952 Ru. rotundata Broennimann, 1953 Ru. rugosa Plummer, 1926 Ru. scotii Broennimann, 1952

Subbotina S. triloculinoides Subbotina, 1953 = Eoglobigerina triloculinoides; Eoglibigerina microcellulosa S. trivialis Subbotina, 1953 = Eoglobigerina trivialis

Woodrigina W. claytonensis Loeblich and Tappan, 1957 W. hornerstownensis Olsson, 1960

Other Paleocene Taxa Bifarina alabamensis Cushman, 1940 Zeauvigerina teuria Finlay, 1947

50 Other Cretaceous Taxa Abathomphalus intermedius Bolli, 1951 Abathomphalus mayorensis Bolli, 1951 Contusotruncana contusa Cushman, 1926 Contustruncana patelliformis Gandolfi, 1955 Contosotruncana pilcata White, 1926 Gublerina acuta Klasz, 1953 Gublerina cuvillieri Kikoine, 1948 50 Plummerita hankteninoides Broennimann, 1952 Rosita contusa Cushman, 1926 Rugotruncana circumnodifer Finlay, 1940 Shackoina multispinata Cushman and Wickenden, 1930

51

Appendix B

Paleoecology Table

52 PALEOECOLOGY TABLE

The paleoecology of taxa used in this study was an important part of interpreting the

NMS results. In some cases, paleoecology can be determined by a fossil’s geographic

distribution. Certain taxa that are commonly found only in high latitudes, for example the

Cretaceous nannoplankton Nephrolithus frequens, most likely preferred a cold-water eutrophic environment (e.g. Thierstein, 1981; Perch-Nielsen, 1985). Prior to the extinction, Guembelitria cretacea, a planktonic foraminifer that bloomed almost globally right after the K-P extinction,

was found only in high-stress shelf sections, and is morphologically similar to modern, high- stress tolerant planktonic foraminifera (Keller et al., 2002) implying that this taxon was able to tolerate high stress. Some taxa are also extant, such as Braarudosphera bigelowii, and their

ecological affinities have been studied and are at least somewhat understood (Kelly et al., 2003).

Finally, in modern ecosystems, cell size (or in this case skeletal size) often directly relates to the

nutrient availability in the environment (Reynolds, 2006). Large cells (>10 μm) are typically

found in oligotrophic environments where large size aids in the competition for scarce nutrients

and fends off predation. Small cells (~0.5-10 μm) are common in eutrophic environments

because small cells have higher surface area to volume ratios which makes them more efficient

at nutrient uptake. Applying this modern trend in cell size to skeletal size can provide some

insight into the paleoecologies of plankton in the earliest Paleocene. The paleoecology of

Danian genera is shown in Table B1 (nannoplankton) and Table B2 (planktonic foraminifera)

with sources and notes.

53 Trophic Level: D = Disaster-bloom; Nannoplankton E = Eutrophic; M = Mesotrophic; O = Oligotrophic (see Hallock, 1987) Trophic Genera Level Notes Sources May be disaster taxa due to Biantholithus ? similarity with Braarudosphaera based on Cretaceous paleoecololgy Roth and Bowdler, 1981; Biscutum E of B. constans Watkins, 1989 Extant, present generally in high e.g. Kelly et al., 2002; Braarudosphaera D stress shelf environments Tantawy, 2003 C. ultimus common only in shelf Chiastozygus e.g., Pospichal, 1993; E? sections Coccolithus O Fuqua et al., 2008 Aubry, 1998; Cruciplacolithus M Fuqua et al., 2008 Cyclagelosphaera E Bown, 2005 Futyania M Gardin, 2002 Hornibrookina E descendant from Biscutum Perch-Nielsen, 1985 Lees, 2002; Markalius E Bernoala and Monechi, 2007 Müler, 1985; Micrantholithus D Eshet et al., 1992 very small cell size, represents Neobiscutum Gardin, 2002 D climatic instability Neochiastozygus E relatively small and simple e.g. Perch-Nielsen, 1985 Neocrepidolithus E? Common in high latitudes Perch-Nielsen, 1985 found mostly in marginal, near-shore Octolithus Gardin, 2002 E environments Due to its small size, P. dimorphosus Aubry, 1998; Prinsius M may be eutrophic Fuqua et al., 2008 Thoracosphaera Eshet et al., 1992; D /Calcispheres Tantawy, 2003

Zeugrhabdotus E Bown, 2005

Table B1: Nannoplankton paleoecology: trophic level as well as notes and references used.

54

Trophic Level: D = Disaster-bloom; E = Eutrophic; Planktonic Foraminifera M = Mesotrophic; O = Oligotrophic (see Hallock, 1987)

Trophic Genera Level Notes Sources Chiloguembelina E prefers deep, unoxygenated water D'Hondt and Keller, 1991 Eoglobigerina E? small and simple morphology Keller, 1993 pl. 2 Globanomalina E? small and simple morphology Keller, 1993 pl. 2 similar morphology and size as Arenillas et al., 2000 Globoconusa E? Parasubbotina pl. 2 Guembelitria D e.g. Keller et al., 2002 Parasubbotina E related to Subbotina Coxall et al., 2006 has disaster-bloom characteristics Parvularugoglobigerina D e.g. Arenillas et al., 2000 after the K-P boundary Similar to Morozovellids Praemurica M (oligotrophs), but lacked algal Coxall et al., 2006 symbionts Subbotina E asymbiotic deep water dweller Coxall et al., 2006 Woodrigina E preferred deep, unoxygenated water D'Hondt and Keller, 1991

Table B2: Paleoecology of planktonic foraminifera with trophic level, notes and references.

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Aubry, M. –P. 1998. Early Paleogene calcareous nannoplankton evolution: A tale of climatic deterioration. In D. R. Prothero and W. A. Berggren, eds. Eocene-Oligocene Climatic and Biotic Evolution. Princeton University Press, Princeton, New Jersey:272-308.

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56 Kelly, D. C., Norris, R. D., and Zachos, J. C. 2003. Deciphering the paleoceanographic significance of Early Oligocene Braarudosphaera chalks in the South Atlantic. Marine Micropaleontology, 49:49-63.

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58

Appendix C

Supplements to the Thesis:

1. Detailed NMS Analysis Methods

2. Age Model

3. Range/Abundance charts of selected Danian taxa

59 DETAILED NMS ANALYSIS METHODS

Nonmetric Multidimensional Scaling (NMS) is a multivariate analysis that seeks to explain a large, multivariate dataset with a smaller set of variables (or dimensions). The NMS analysis dimensions are compared to the original dataset using a metric termed as stress, which is calculated within the NMS algorithm using Eq. C1 using the rank distances of points in the original dataset and the distances of points in the NMS resulting data matrix (McCune and

Grace, 2002). A smaller stress indicates that the NMS analysis represents the structure of the data accurately. More dimensions applied to the NMS lessen the stress, but this can also lessen the interpretability of the results and the variance represented by each dimension.

Eq. C1 (McCune and Grace, 2002):

Where:

S* = Stress dij = distance metric of the original data matrix

= distance metric of resulting NMS data matrix

The NMS was run using R software and the Vegan package (www.r-project.org) with the metaMDS algorithm. For this analysis, the Bray-Curtis distance metric was used to calculate the distances between samples. The metaMDS algorithm runs the NMS analysis by first computing several random starts based on a user input of the number of axes to use as well as the distance metric used. The result with the lowest stress is used as the solution for the analysis. After a

60 solution is determined, the program scales the results by centering the origin of the plot on the

average of each dimension and then comparing the distance of points in NMS space with the

distance of the points in the original data matrix. It then orients the axes by running a principal

components analysis to force the dimension that represents the highest variance of the data to be

plotted as NMS1.

Because metaMDS requires an input number of axes, the first step in running the analysis

was to determine the number of axes use. One way to do this is to run the analysis several times,

each time increasing the number of dimensions used in the final solution. The stress of each

result can then be plotted against the number of dimensions. This plot shows the point where

adding dimensions to the analysis no longer reduces the stress by a considerable amount, and can be used to interpret the dimensionality of the dataset (McCune and Grace, 2002). For both the nannoplankton and planktonic foraminiferal datasets, these plots were made (Fig C1), and from these plots, it appears that adding any more dimensions than four does not decrease the stress by

a considerable amount. Because of this, the NMS analyses were run with a dimensionality of

four.

Figure C1: Dimensionality versus stress of the NMS analysis for both nannoplankton and planktonic Foraminifera. Stress does not decrease by any considerable amount after four dimensions, indicating that the NMS analysis will represent most of the variance of the original data with four dimensions.

61 The metaMDS algorithm was run again with a dimensionality of four using 35 iterations.

The final stress of the analysis was 5.814 (nannoplankton) and 4.506 (planktonic foraminifera).

Determining the variance represented by the four NMS axes was done using an after-the-fact

method. This method involved first calculating the Bray-Curtis distance for the original data

matrix as well as a Euclidean distance for each of the NMS axes (as outlined in McCune and

Grace, 2002). The percent variance is explained by finding the r2 correlation between these two distances. Also after the analysis, the resulting points in NMS space were rotated about the origin for both plots. This was done using a simple geometric rotation found in any basic trigonometry text (Eq. C2) at an angle so that age/depth of each sample and NMS2 were more closely correlated (because NMS2 is most likely related to the age or depth of the sample, see

Discussion).

Eq. C2

cossin

sincos

x' and y' are the rotated datapoints x and y are the original datapoint θ = rotation angle

AGE MODEL An age model was necessary to correlate the different sites and to plot the NMS results

with time. Well defined cyclostratigraphy exists for Shatsky Rise (ODP Site 1210) and Walvis

Ridge (ODP Site 1262) (Westerhold et al., 2008); however, cyclostratigraphy is not well defined

for the other sections used in this study. Because of this, the most accurate correlation is ODP

Site 1210 to ODP Site 1262 (although this correlation is also not entirely accurate due to a lack

of early Paleocene cycles at ODP Site 1210). All other correlations must be done based on

62 biostratigraphy, and while not accurate, it can be used for a relative comparison. For this study,

Westerhold et al.’s (2008) age “option two” was used to establish age models because option two was based on what was thought to be a more accurate age of the Paleocene/Eocene boundary

(55.93 Ma). First, a composite section was constructed using cyclostratigraphy for ODP Site

1210 and ODP Site 1262 (Westerhold et al., 2008) (Fig. C2). Microfossil datums were added to this correlation to produce a composite standard section used to correlate all sections (Table C1).

The composite section was correlated to the sections sampled for foraminifera at Walvis

Ridge (DSDP Site 528) and Shatsky Rise (DSDP Site 577) (Figs. C3 for DSDP Site 577; Fig. C4 for DSDP Site 528). For these sections, both foraminifera and nannoplankton biostratigraphy is known (e.g. D’Hondt and Keller, 1991), and the K/P boundary can also be correlated between these sites. The composite standard was then correlated to biostratigraphic datums for

Kerguelen Plateau (ODP Site 738), El Kef and Brazos River (Figs. C5, C6 and C7 respectively).

For Cretaceous ages at each site, sedimentation rates found in the literature were used (Table

C2). Age models were constructed under the assumption that sedimentation rates were constant between tie points (also an inaccurate assumption).

63

Figure C2: Correlation of events between Shatsky Rise (ODP Site 1210) and Walvis Ridge (ODP Site 1262) where K/P is the Cretaceous/Paleogene boundary and Pcene 5-8 correspond to major Paleocene cycles outlined in Westerhold et al. (2008).

Age Event (Ma) K-P boundary 65.680 P0/Pa 65.656 Pa/P1a 65.628 Pcne2 65.556 P1a/P1b 65.509 Pcne3 65.456 P1b/P1c 65.340 Pcene4 65.348 NP1/NP2 65.323 Pcne5 65.248 Pcne6 65.168

Table C1: Composite standard section showing major Paleocene cycles and biostratigraphic datums with age in millions of years. P0/Pa = FO of P. eugubina, Pa/P1a= LO P. eugubina, P1b/P1c = FO S. triloculinoides, NP1/NP2 = FO C. intermedius; Pcene2-6 correspond to major Paleocene cycles listed in Westerhold et al (2008).

64

Location ODP Site Sed. rate (m/Kyr) Reference Shatsky Rise ODP 1210 .010575 Bralower et al., 2002 ODP 1212 (DSDP 577 0.003625 Bralower et al., 2002 equivalent) Walvis Ridge ODP 1262 0.0263 Westerhold et al., 2008 Kerguelen Plateau ODP 738 .007737 Huber, 1991 El Kef 0.045 Vonhof and Smit, 1997 Brazos River N/A No Cretaceous Samples counted

Table C2: Sedimentation rate used to establish Cretaceous age models for each site and references.

Fig. C3: Correlation of DSDP Site 577 (Shatsky Rise) to the composite standard; Paleocene cyclostratigraphy listed derived from depth of cycles at ODP Site 1212, a re-drill of DSDP Site 577.

65

Fig. C4: Correlation of DSDP Site 528 (Walvis Ridge) with the composite standard ages of selected microfossil datums.

Fig. C5: Correlation of ODP Site 738 (Kerguelen Plateau) with the composite standard ages of selected microfossil datums. FO of E. firnga corresponds to the start of the APα zone of Stott and Kennet (1990) and is approximately equivalent to the Pα zone of Berggren et al. (1995). FO G. daubjergensis is approximately equivalent to the beginning of the P1a zone of Berggren et al. (1995). Therefore, these two datums are estimated by the datums given for the equivalent bioevents in the Berggren et al. (1995) biostratigraphy.

66

58

Fig. C6: Correlation of El Kef with the composite standard section ages of selected datums.

Fig C7: Correlation of the Brazos River depths with the composite standard section ages of selected microfossil datums.

67

RANGE AND RELATIVE ABUNDANCE OF PALEOCENE TAXA

Range and abundances of several Paleocene taxa were plotted to aid in interpretation of

NMS results. Each diagram is plotted against the age model (after Westerhold et al., 2008).

Also plotted are sample locations to show where ranges were estimated through widely spaced samples. This is done for each site (Fig. C8-12).

Fig C8: Range and relative abundance for nannofossils and planktonic foraminifera at Walvis Ridge

68

Figure C9: Range and relative abundance for nannofossils and planktonic foraminifera at Shatsky Rise

69

Figure C10: Range and relative abundance for nannofossils and planktonic foraminifera at El Kef

70

Figure C11: Range and relative abundance for nannofossils and planktonic foraminifera at Cottonmouth Creek

71

Figure C12: Range and relative abundance for nannofossils and planktonic foraminifera at Kerguelen Plateau

72 References:

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Huber, B. T. 1991. Maestrichtian planktonic foraminifer biostratigraphy and the Cretaceous/Tertiary boundary at hole 738C (Kerguelen Plateau, Southern Indian Ocean). In J. Barron, B. Larsen, et al. eds. Proceedings of the Ocean Drilling Program, Scientific Results, 119:451-465.

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