Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs

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2019-08-12 Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs

Jordan, Katherine J.

Jordan, K. J. (2019). Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs (Unpublished master's thesis). University of Calgary, Calgary, AB. http://hdl.handle.net/1880/110722 master thesis

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Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in

Cretaceous Molluscs

Katherine J. Jordan

A THESIS

SUBMITTED TO THE FACULTY OF GRADUATE STUDIES

IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE

DEGREE OF MASTER OF SCIENCE

GRADUATE PROGRAM IN BIOLOGICAL SCIENCES

CALGARY, ALBERTA

AUGUST, 2019

Abstract

Species selection and other macroevolutionary forces are challenging processes to study and quantify when using fossil data. Here, I used the Price equation to analyze changes in geographic range sizes prior to and during a mass extinction event to estimate the relative contribution of three macroevolutionary processes (species selection, anagenesis, and immigration). I also tested the hypothesis that larger geographic range size increases a group’s survivability during mass extinctions. I applied a similar method to Rankin et al. (2015) to study marine gastropods and bivalves of the Gulf and Atlantic Coastal Plain (originally studied by

Jablonski (1987)) over the last 16 million years of the Cretaceous Period. I found three major changes in mean geographic range size shared by both gastropods and bivalves during the end-

Cretaceous: an increase in mean range size during the late Campanian, a decrease in the mid-

Maastrichtian, and an increase near the end of the Cretaceous Period (late Maastrichtian). The

Price Equation indicates that the late Campanian increase in geographic range size was attributable primarily to immigration, the mid-Maastrichtian decrease was due to different combinations of the three processes (species selection, anagenetic change, and immigration) in gastropods and bivalves, and the late Maastrichtian increase was attributable to species selection.

These changes in geographic range size coincide with a marine transgression event, a period of global climate change, and a marine regression event, respectively. A statistically significant correlation between larger geographic range size and increased survivability was found for one time increment (approximately four million years before the KPg boundary). This study shows that the relative contribution of interacting macroevolutionary processes fluctuated over the end-

Cretaceous extinction event and suggests that large geographic range size can increase survivability under certain conditions leading up to a mass extinction.

Keywords: species selection, anagenetic change, immigration, the Price equation, macroevolution, mass extinction.

iii Acknowledgements

I would like to start with an acknowledgement to those who have helped me in my professional development. A heartfelt thank you is much overdue to my advisor Dr. Jeremy Fox whose patience and kindness have gotten me to this point. I am appreciative of every bit of advice, cooking, and funding I have received as member of his lab. I hope that my hard work and future success will be the biggest demonstration of my thanks, Jeremy. I would also like to thank

Dr. Jessica Theodor for being the role model I needed. Her strength and passion are inspiring. I am grateful for all the work she has done to help me become the paleontologist I dreamed I could be. A thank you to Dr. Mindi Summers and Dr. Charles Henderson, as well, for enhancing my invertebrate knowledge to make this thesis possible.

To my twin sister, Rebecca: Her wisdom and love have helped me immensely. I am so glad you are my sister. To my undergraduate professor Dr. Brian Penney: It was Dr. Penney who found out about the graduate opportunity here in Calgary. In a way, I hope I have carried on his legacy by attending graduate school in Alberta as he did. Thank you so much, Dr. Penney.

My friends here in Canada Selina, Colby, Rachel, and all the rest (from campus and Telus

Spark): you all got me through and were more kind than I deserved at times. I appreciate you more than you will ever know. I would also like to extend my gratitude to my mother, Vicki, my partner, Aaron, my dearest friends, Caitlyn, Jess, and Matt. I am the best person I can be because of you all. Thank you for all your support in the last two years. I hope to make you all proud.

iv Dedication

This thesis is dedicated to my grandparents, Leroy and Dorothy Weed of Stonington, ME.

Thank you for giving me a love of the ocean that has stayed with me and brought me, ironically, to the prairies of Alberta.

v Table of Contents

Abstract ...... ii Acknowledgements ...... iv Dedication ...... v Table of Contents ...... vi List of Tables and Equations ...... viii List of Figures and Illustrations ...... ix Epigraph ...... x

CHAPTER ONE: INTRODUCTION ...... 1 1.1: Macroevolution and the Price Equation………………………………………1 1.2: The Cretaceous Mass Extinction and Cretaceous Molluscs…………………10

CHAPTER TWO: METHODS ...... 13 2.1: The Price Equation…………………………………………………………. 13 2.2: Dataset Selection and Preparation for Analysis……………………………. 16

CHAPTER THREE: RESULTS ...... 22 3.1: The Price Equation Results………………………………………………….22 3.2: Statistical Analysis of Datasets……………………………………………. 26 3.3: Further Analysis of Datasets………………………………………………...27

CHAPTER FOUR: DISCUSSION ...... 29 4.1: Macroevolutionary Change Within Time Increment T2-T3………………...29 4.2: Macroevolutionary Change Within Time Increment T6-T7………………...30 4.3: Macroevolutionary Change Within Time Increment T7-T8………………...32 4.4: Periods of Macroevolutionary Stasis in the Data……………………………33 4.5: Differential Survivorship Among Cretaceous Molluscs…………………….35 4.6: Limitations of the Study……………………………………………….….... 38

CHAPTER FIVE: CONCLUSION ...... 41

REFERENCES ...... 43

APPENDIX 1: Table of gastropod ancestor-descendant pairs of the Gulf and Atlantic Coastal Plain. Species names and ranges from Hunt et al. (2005) Appendix B………………….51

APPENDIX 2: Table of bivalve ancestor-descendant pairs of the Gulf and Atlantic Coastal Plain. Species names and ranges from Hunt et al. (2005) Appendix B………………….57

APPENDIX 3: Species used in PAST biostratigraphy analysis. Species were used based on their occurrence and presence in the formations used in the study. Data collected from the Paleobiology Database (PBDB)………………………………………………………….62

APPENDIX 4: Results of the PAST unitary association analysis. Maximal cliques presented from the unitary associated output……………………………………………………….66

vi APPENDIX 5: Gastropod dataset arranged in time bin for Price equation analysis…….67

APPENDIX 6: Bivalve dataset arranged in time bin for Price equation analysis……….77

APPENDIX 7: Gastropod Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants) ……………………………….84

APPENDIX 8: Bivalve Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants) ……………………………….109

vii List of Tables and Equations

Table 1: Results of Permutation Test for Each Time Increment. P values are reported. Null hypothesis: geographic range size and number of descendants left per time bin are independent of each other……………………………………………………………………………………25

Equation 1: Modified from Rankin et al. (2015) ………………………………………….... 13

Equation 2: The extended Price equation modified from Rankin et al. (2015), eq. 2.1………14

viii List of Figures and Illustrations

Figure 1: Total mean geographic range size change (blue line) gives the Price Equation answer for the gastropod analysis. In comparison, the three separate Price Equation terms (species selection, anagenetic change, and immigration) plotted ………………...………………………20

Figure 2: Total mean geographic range size change (blue line) gives the Price Equation answer for the bivalve analysis. In comparison, the three separate Price Equation terms (species selection, anagenetic change, and immigration) plotted ...………………………………………21

Figure 3: Proportion of extinctions in both gastropods and bivalves in each time bin from Campanian to KPg boundary…………………………………………………………………….26

ix Epigraph

PIPPIN: I didn't think it would end this way.

GANDALF: End? No, the journey doesn't end here. Death is just another path, one that we all must take. The grey rain-curtain of this world rolls back, and all turns to silver glass, and then you see it.

PIPPIN: What? Gandalf? See what?

GANDALF: White shores, and beyond, a far green country under a swift sunrise.

PIPPIN: Well, that isn't so bad.

GANDALF: No. No, it isn't.

― J.R.R. Tolkien, The Lord of the Rings

x 1

CHAPTER ONE: INTRODUCTION

1.1: Macroevolution and the Price Equation

The history of life on Earth has been characterized by dramatic directional trends brought about by macroevolutionary processes. These trends included changes in body size, morphological shape, geographic range, or lifestyle, as well as faunal radiations and extinctions, which can be observed in the fossil record dating back to the Ediacaran over 541 million years ago (Butterfield, 2007). For instance, tyrannosaurs increased in body size for millions of years, until the appearance of the dinosaur species, Tyrannosaurs rex, in the Cretaceous Period

(Brusatte et al., 2010). Macroevolution can exhibit decreases in body size (e.g., mammals found in the Bighorn and Clarks Fork Basins of Wyoming during the Paleocene/Eocene boundary)

(Rankin et al., 2015), as well as morphological stasis (e.g., early cyanobacteria (Butterfield,

2007).

Macroevolutionary trends can change through time, and dramatic reversals and new trends have been found following mass extinctions (Gould, 1985). The radiation of the acritarchs, for example, ceased very rapidly by the end of the Ediacaran and led to one of the first observable cases of widespread extinction (Butterfield, 2007). This extinction was followed by one of the largest radiations of eumetazoans: the Cambrian Explosion, which saw the origination and radiation of a completely different and diverse fauna (Butterfield, 2007). Likewise, the radiation of many mammalian groups followed the extinction of non-avian dinosaurs after the

Cretaceous-Paleogene (KPg) mass extinction event around 66 million years ago (Grossnickle &

Newham, 2016). Mass extinctions can also drive the increase of previously rare morphological features. For example, the diversity of crinoids with a more motile lifestyle increased after the

Permian mass extinction, possibly as a result of being able to evade predators such as sea urchins better than stationary stalked crinoids (Baumiller et al., 2010).

Identifying and investigating macroevolutionary trends requires macroevolution to be clearly defined and differentiated from microevolution. Macroevolution involves processes that affect species and higher taxonomic clades, and occurs over a longer period of time and space

(Erwin, 2000). Macroevolution has been decoupled from microevolution because it can account for speciation and extinction events, not just population-level evolutionary changes (Stanley,

1982). Time is also important in differentiating the two methods of evolutionary change. For example, microevolution can be seen as occurring in “ecological time” in which populations change over short periods of time by processes such as natural selection (Bell, 2016). In contrast, macroevolution occurs in “evolutionary time”, affecting higher taxonomic groups over longer periods of time, sometimes even changing the way global communities appear (Bell, 2016). Due to the necessity of long periods of time, macroevolution is hard to replicate in a lab (Bell, 2016).

The fossil record, which encompasses large periods of time, is a valuable asset that can help detect large-scale patterns such as macroevolution (Erwin, 2000; Grantham, 1995).

Species selection has been argued to be one of the most important processes that contribute to macroevolutionary trends. Species selection refers to differential speciation and extinction of species by virtue of their traits. Species selection is the macroevolutionary equivalent of natural selection on the microevolutionary scale (Liberman and Vrba 2005; Rankin et al., 2015; Stanley, 1975). Species selection can act on any trait that varies among species, including both traits that directly reflect the phenotypes of species (e.g., mean adult body size) and emergent characters of a species (e.g., geographic range size) (Lloyd and Gould 1993).

There are two aspects to understanding the complexity of when species selection has occurred: Firstly, species selection is a force that acts distinctly on a macroevolutionary level and may not be the result of cumulative rounds of microevolution. This is because species selection can act on emergent species-level attributes such as geographic distribution (i.e. the latitudinal; diversity gradient) (see Grantham, 2007; Jablonski, 2008). It can be argued that these species- level attributes (e.g. geographic range size) encompass smaller, organismal-level attributes (e.g. ecological lifestyle). For the purposes of this study, species-level attributes are considered as separate and distinct. Secondly, the effect of species selection can be difficult to tease away from the effect of microevolutionary processes. To understand this, the results of the study done by

Simpson (2013) demonstrate that species selection among corals favors corals to be non- photosynthetic, but colonial (which promotes high diversification rates). However, most coral species today are photosynthetic and colonial, or else non-photosynthetic and solitary (Simpson,

2013). Microevolutionary variance among corals constrains species selection and prohibits a macroevolutionary response (i.e. limits the effects of selection) (Simpson, 2013). It can be difficult to determine this kind of phenomenon. Studies such as these indicate the need for a way to quantify the effect of species selection. This is especially true when dealing with fossil data.

To observe the macroevolutionary change in species-level attributes, great spans of time are needed (see Bell, 2016). The fossil record is a good source of observable macroevolutionary change due to species selection because it captures changes occurring over great lengths of time

(see Benton & Pearson, 2001; Vrba, 1984). The need for ancestor-descendant relationship knowledge among species can be a problem as a full record of ancestor-descendant relationships may not be preserved in the fossil record or could be uncertain (Rankin et al., 2015; Vrba, 1984).

So, attempting to quantify species selection may not only be difficult because of the need for

4 species-level attributes, but also because these attributes may be difficult to trace through an incomplete fossil record.

Determining when species selection has occurred and how it affects macroevolutionary change has been done by collecting specimens and tying speciation events to morphological changes in populations by paleontologists (see Benton & Pearson, 2001). Proof of speciation via species selection has often been done by verbal argument rather than by mathematical calculation (see Jablonski, 2008). A concept that was originally defined by itself (e.g. Stanley,

1975), species selection has often been associated with the evolutionary hypotheses of gradualism and punctuated equilibrium (see Eldredge & Gould, 1972; Gould & Eldredge, 1977;

Pennell et al., 2014). More recently, models have been proposed to quantify species selection

(see Jablonski, 2008 for examples). Simpson (2010, 2013) performed two studies that used one model to quantify the decrease in the complexity of crinoid calyxes throughout time, and species selection on nonsymbiotic and symbiotic scleractinian corals. In both studies, Simpson (2010,

2013) made assumptions about the data in order to identify if species selection was present

(regression with a slope of non-zero) and did so by using fossil and extant taxon data with geological time intervals. Most importantly, Simpson (2010, 2013) used the Price equation to separate the effect of species selection away from other macroevolutionary forces (i.e. anagenesis). Goldberg et al. (2010) applied a different approach to identify species selection among plants. The plant family chosen in the study had a significant amount of data on it to construct a phylogenetic tree (Goldberg et al., 2010). A statistical analysis was done on this phylogenetic tree, and Goldberg et al. (2010) found species selection was acting on hermaphroditic plants to promote self-incompatibility (no self-fertilization) and those with self- incompatibility had a higher diversification rate. In conclusion, species selection can potentially

5 be quantified using different methods but to do so requires applicable data and, if using the fossil record, a good history (see Jablonski, 2008; Vrba, 1984).

Anagenetic change, or anagenesis, is another process that affects macroevolution.

Anagenesis is within-lineage evolutionary change (Vaux et al., 2016). It is distinct from cladogenesis (branching of lineages) which might lead to species selection (Aze et al., 2013).

The problem with identifying when anagenesis has occurred is threefold: it can often be confused with cladogenesis, this is done because of vague definitions, and the fossil record may not help in clarification. Firstly, the macroevolutionary signals of anagenesis and cladogenesis often are difficult to distinguish (Gould, 1980). Since both anagenesis and cladogenesis (and by proxy species selection) can lead to speciation, confusion may arise (Vaux et al., 2016). Also, anagenesis and cladogenesis are often discussed together with vague or no definitions (Vaux et al., 2016). The history of this confusion may also be in part due to the hypothesis of punctuated equilibrium; cladogenesis was often equated with punctuated change while anagenesis was equated with phyletic gradualism (see Hallam, 1997 for example; Vaux et al., 2016).

Additionally, without detailed stratigraphic sampling in the fossil record, it may be difficult to determine if anagenesis or cladogenesis have occurred (Simpson, 2013).

The necessity of the fossil record to discern anagenetic change, despite its problems, cannot be underestimated. Examples of anagenesis in both vertebrates and invertebrates are interpreted with fossil evidence (see for example Hallam, 1997; Kimbel et al., 2006; Scannella et al., 2014). Paleontologists hypothesize anagenetic change has occurred when morphospecies are temporally isolated from one another and therefore reproductively isolated (i.e. ancestors will not be found in the same stratigraphic layer as their descendants) (Aze et al., 2013). One limitation of identifying anagenetic change in the fossil record is knowing ancestor-descendant

6 relationships. When ancestor-descendant relationships are known, the change in morphology of temporally isolated species has been interpreted as anagenetic change (e.g. Scannella et al.

(2014) the change in ceratopsian cranial morphology over time as an example). Speciation due to anagenesis most likely is a rare phenomenon (Gingerich, 1985), but here I focus on anagenesis in the sense of within-lineage change, whether or not it leads to speciation.

Anagenesis as a macroevolutionary force can impact many traits of a species including morphology and habitat (Gould, 1988). Interpreting when anagenesis has occurred, especially in the fossil record, has its subjective interpretations by paleontologists. Quantifying anagenesis to separate it from cladogenesis (and species selection) is one way to avoid confusion. This has been done successfully using the Price equation (see Price, 1972; Rankin et al., 2015; Simpson

2010, 2013).

One final process that can affect macroevolutionary change is immigration. Immigrant species migrate into a new area from an ancestral area, and may increase regional biodiversity

(Stigall et al., 2017). The effect of immigration can lead to macroevolutionary change by adding new traits (variance) to a regional area (Rankin et al., 2015). Immigrant descendants will eventually become resident ancestors, and will contribute to the overall mean trait of a group over time (see Rankin et al., 2015). For example, Rankin et al. (2015) found that small-bodied mammal species immigrating into the Bighorn and Clarks Fork Basin during the

Paleocene/Eocene boundary reduced the total mean body size of mammals here over time. The effects of immigration likely depend on the species; immigrant species are typically more cosmopolitan and can contribute to niche partitioning (leading to increased speciation over time) or may lead to extinctions of residents by outcompeting them (Stigall et al., 2017). Immigration of new species can also be associated with environmental changes. These environmental changes

7 can occur rapidly or gradually depending on what is happening in the environment (e.g. abrupt climate change or gradual sea level change), and can affect immigrants and residents alike

(Jackson & Sax, 2010). For example, many species immigrate northward during global warming events (e.g. alpine plants immigrating higher on a mountain) and may outcompete the residents

(Jackson & Sax, 2010).

Immigration in the fossil record can be observed, but is problematic to interpret. It is often difficult to determine when an immigration event has occurred. Observing the spatial patterning and the historical distribution of species is challenging, and this is made even more difficult by an imperfect fossil record (Goldberg et al., 2005). An imperfect fossil record could mean the loss of a potential ancestor and, with it, the loss of being identified as a resident (non- immigrant) (see Goldberg et al., 2005; Springer, 1990). Despite this potential problem, immigration has been identified in the fossil record (see Web, 2006). For example, the Great

American Biotic Interchange (GABI) with the exchange of animals from North and South

America during the late Pliocene is one such example of mass immigration in the fossil record

(Webb, 2006). However, observing and quantifying immigration are two separate matters.

Without high temporal or spatial resolution of a prehistoric group, quantifying immigration can be difficult (Goldberg et al., 2005). Rankin et al. (2015) proved that quantifying immigration could be done using the Price equation, a method demonstrated to have broad applications to macroevolutionary studies.

In summary, species selection is a potentially important macroevolutionary force, conceptually distinct from other macroevolutionary forces such as within-lineage (anagenetic) change and the movement of species to new locations (immigration). Empirically distinguishing species selection, anagenesis, and immigration from one another has proven difficult. The

8 reasons for this are: the controversy over the definition of species selection, data quality issues, and the lack of a method to quantify total macroevolutionary changes and attribute it to different causes. First, if species selection cannot be defined, then it would be difficult to quantify (see

Vrba 1984 and Grantham 1995 for controversies). Second, as mentioned before, data quality for macroevolutionary studies often involves fossil data. With incomplete information of fossil taxa

(i.e. incomplete preservation or sampling bias), it is difficult to determine when a species actually went extinct, which taxa are ancestors and descendants, and if speciation is due to anagenetic change or immigration. To address these issues, a clear and empirical approach is necessary that can take data, regardless of assumptions made, and analyze it to understand the macroevolutionary change affecting a species of interest.

The Price equation is one such empirical approach that can tease apart macroevolutionary effects such as species selection, anagenesis, and immigration by looking at the changes in phenotypic traits or emergent characters between ancestors and their descendants over time

(Rankin et al., 2015). The Price equation, as mentioned above, has been an essential method to distinguish macroevolutionary forces from one another (see Rankin et al., 2015; Simpson 2010,

2013). The Price equation is named for its developer, George Price (Price, 1970). Price's original

(1970) equation provided an abstract, broadly-applicable definition of the response to selection

(Price, 1970; Price, 1972). The original equation allowed the microevolutionary change in the mean phenotypic trait of a group of individual organisms to be calculated by separating the effect of selection (differential survival and reproduction of individuals with different traits) and transmission bias (changes in traits between ancestors and their descendants) (Price, 1972). The

Price equation is a purely descriptive equation (Frank, 1997). It also does not rely on biological assumptions (Rankin et al., 2015). Rather, it starts from information about the traits of ancestral

9 individuals and the traits of their descendants, and then derives from that information the implied values of selection and transmission bias. Because the Price equation makes no assumptions about the nature of the processes giving rise to selection and transmission, it can be interpreted to apply to entities other than individual organisms. Rather than considering individual organisms, the Price equation can be applied to species, which can be thought of as having “phenotypic traits” (measurable attributes such as mean body size or geographic range). Like individual organisms, species can be thought of as surviving, “dying” (going extinct), and leaving

“offspring” (descendant species with their own “traits”). Thus reinterpreted, the Price equation provides a way to calculate species selection (analogous to natural selection in microevolution) and anagenetic change (analogous to transmission bias in microevolution).

One limitation of the original Price equation is it assumes zero immigration: every descendant individual was assumed to be the offspring of another individual in the evolving population, rather than a migrant from elsewhere (see Price, 1972; Rankin et al., 2015). Kerr and

Godfrey-Smith (2009) extended the Price equation to include an immigration term, which can be interpreted as capturing the direct effect of immigration in macroevolution (i.e. macroevolutionary change arising because immigrant species differ from residents in the mean value of the trait of interest). The Price equation has been utilized in the past to look at changes in different properties in species in the fossil record (see for some examples Rankin et al, 2015;

Simpson, 2010, 2013).

For the purposes of this study, the Price equation was applied to groups that lived during the end of the Cretaceous Period, just prior to, and during the Cretaceous mass extinction event.

The benefit of using data around a mass extinction event is the potential to observe the disruption of macroevolutionary forces during a major geological and ecological transition. Changes in

10 geographic range size in Gulf and Atlantic Coastal Plain molluscs will be analyzed. The dataset was created by David Jablonski and used in his studies about Cretaceous gastropods and bivalves from the Gulf and Atlantic Coastal Plain of North America (e.g. Jablonski, 1986; Jablonski

1987). Jablonski hypothesized that large geographic range size was a heritable emergent property of a species that could increase a species’ likelihood of surviving a mass extinction event

(Jablonski, 1986; Jablonski, 1987). Using many lines of evidence, Jablonski demonstrated how this was a possibility and suggested using the Price equation to test this hypothesis (Jablonski,

2008). Therefore, the Price equation was used in this analysis to test whether or not large geographic range size was selected for and what macroevolutionary change affected molluscs during the last sixteen million years of the Cretaceous Period.

1.2: The Cretaceous Mass Extinction and Cretaceous Molluscs

The end-Cretaceous mass extinction event is interesting for many reasons. There have been five mass extinctions identified throughout the history of life on Earth: the late Ordovician, the late Devonian, the end-Permian, the late Triassic, and the end-Cretaceous mass extinctions in order from oldest to youngest in geological time (Raup & Sepkoski, 1982). The Cretaceous mass extinction is best known for the loss of the non-avian dinosaurs, marine reptiles, and flying reptiles (Alvarez et al.,1980). The Cretaceous mass extinction is also notable in that an asteroid impact has been suggested as the cause of an ecological cascading effect that triggered the mass extinction (Alvarez et al., 1980, 1984). The discovery of one of the largest craters on Earth in

Mexico dating to the KPg added to the popularity of this hypothesis (Hildebrand et al., 1991).

Over time, more theories have been suggested for the cause (or causes) of the mass extinction,

11 often citing volcanism and global climate change as mechanisms that triggered the mass dying of many organisms (see Reddin et al, 2019; Burgess et al., 2019). In more recent times, it has been noted that there cannot simply be one explanation for the Cretaceous mass extinction event because of the selectivity in those species that went extinct; more research needs to be done to look at the latest Cretaceous paleoenvironment and geological phenomenon happening at the time (Keller, 2001).

Marine invertebrates living at the end-Cretaceous suffered a loss of 47% diversity at the genus level (Erwin, 1998). The recovery of many invertebrate faunas did not occur until well into the Paleogene (Erwin, 1998). Many dominant Cenozoic and Cretaceous groups such as the ammonites and the rudist bivalves declined in numbers before the end of the Cretaceous and were all lost by the Paleocene (Landman et al., 2014; and Jablonski & Raup, 1993). Many paleontologists have provided theories for the loss of major invertebrate groups that had dominated during the Cretaceous and perhaps the Mesozoic in general. Among these most prolific groups of invertebrates hit by the mass extinction were the molluscs. Molluscs are the second largest invertebrate group after arthropods; and are very widespread in their body morphologies, which have been modified in a variety of ways to allow molluscs to inhabit many different habitats and lifestyles (e.g. terrestrial, marine, freshwater, abyssal zone, continental shelf) (Benton & Harper, 2009). The origin of the phylum extends back to the Ediacaran with evidence of a radula found in stem-mollusc groups and the appearance of a molluscan-like species in the Precambrian (i.e. Kimberella) (Caron et al., 2006; Fedonkin & Waggoner, 1997).

A global trend of molluscan fauna decimation during the Cretaceous can be observed in many localities in modern day Patagonia, Spain, France, and North America (see for example Aberhan

& Kiessling, 2014; Ward et al., 1991; Jablonski, 1986; Jablonski, 1987). There is evidence that

12 different molluscs went extinct at relatively different times, with inoceramids declining steadily before the end-Cretaceous and ammonites going extinct by the Paleogene (Ward et al., 1991).

The decimation of such a successful invertebrate group warrants further study.

A regional group of molluscs were examined by quantifying changes in a specific trait (i.e. geographic range size) before and during the Cretaceous mass extinction event. End-Cretaceous gastropods and bivalves of the Gulf and Atlantic Coastal Plain were the primary focus of this study because of the available dataset and abundance of these two groups in the fossil record. By using the Price equation, macroevolutionary forces such as species selection, anagenetic change, and immigration were detected, and evidence of which of these forces acted on the Cretaceous molluscs during the lead up to the mass extinction were observed.

CHAPTER TWO: METHODS

2.1: The Price Equation

The Price equation is versatile because it includes terms for multiple processes in macroevolution: species selection, immigration, and anagenetic change. The Price equation’s versatility is in its validity without needing to account for biological assumptions. This is especially useful for fossil data where many biological assumptions are made, and as long as certain criteria of the analysis are met, the Price equation can be performed. If we can measure some trait or attribute of each of the species present at some point in time, measure the same attribute of each species present at the same location(s) at a later time point, and determine which of the later species are descended from (or are the same species as) the earlier species, then we can apply the Price equation to analyze change in the mean value of the attribute over time. Any measurable attribute can be used (e.g., body size (Rankin et al., 2015) or scleractinian coral coloniality and photosymbiosis (Simpson, 2013)).

Macroevolutionary change in the mean trait value from one time to a subsequent time can be defined as:

퐚 퐧퐝 퐧 퐢 퐚 횫퐳̅ = ∑ (퐳퐣/퐧퐝) − ∑ (퐳 /퐧 ) 퐣=ퟏ 퐢=ퟏ Equation 1: modified from Rankin et al. (2015)

where the total change in the mean trait value from one time to the next (from ancestor to descendant) is represented by 훥푧̅. On the right-hand side of the equation, 푧푖 is the trait of

a ancestral species i (geographic range size in this study), 푧푗 is the trait of descendant species j, n

d 푖 is ancestral species richness, and n is descendant species richness. An indicator value 퐶푗 can be

14 used to express the relationship between ancestors and descendants. The indicator value will equal one if descendant species (j) is a direct descendant or is the same species as the ancestor

푖 species (i). If not, the indicator value will equal zero. Therefore, an ancestor species has 퐶∗ =

푛푑 푖 푖 ∑푗=1 퐶푗 descendants. In this notation, 퐶∗ represents the absolute fitness of the ancestor species

푖 (i), i.e. how many descendants the ancestor (i) left in a later time. If 퐶∗ is equal to one, then

푖 ancestor has left behind a descendant, and if 퐶∗ is equal to zero, the ancestor has gone extinct without leaving a descendant. Descendants are not considered immigrants in the Price equation if

∗ 푛푎 푖 they have an ancestor (퐶푗 ∑푖=1 퐶푗 = 1) (Rankin et al., 2015).

The right-hand side of Equation 1 can be rewritten in meaningful components that make up the extended Price equation (Equation 2):

푎 푎 ∗ 푎 ∗ ∗ 푖 훥푧̅ = 푐표푣(퐶∗ , 푧 )/(퐶∗ /푛 ) − 푐표푣(퐶푑, 푧푑)/(퐶∗ /푛푑) + 푎푣푒((∆푧)푗)

Equation 2: The extended Price equation modified from Rankin et al. (2015), eq. 2.1

in this version of the equation, cov is the covariance of the population and ave is the average

∗ mean. 퐶∗ represents the total number of ancestors and descendants found at a certain point in time, and is used in calculating the relative fitness value, discussed below. The full derivation of the Price equation can be seen in Kerr & Godfrey-Smith (2008).

푎 푎 ∗ 푎 The first term in the Price equation (푐표푣(퐶∗ , 푧 )/(퐶∗ /푛 )) is the species selection term.

This term is equal to the covariance between the ancestral trait (the variation for selection to act on) and ancestral relative fitness (the number of descendants left by each ancestral species, relative to the number left by the average ancestral species). Species selection equals zero when

15 there is no variation among ancestral species for species selection to act upon, and when variation among ancestral species is independent of the number of descendants they leave.

∗ ∗ The second term (푐표푣(퐶푑, 푧푑)/(퐶∗ /푛푑)), the immigration term, is an addition to the

Price equation proposed by Kerr and Godfrey-Smith (2008). The immigration term accounts for those descendants without ancestors (migrants) which may appear in a population (Kerr &

Godfrey-Smith, 2008). The term equals the covariance between the descendant trait of interest and how many ancestors the descendant species has, which is scaled by the average number of descendants with ancestors (the non-immigrants). This is mathematically equivalent to the difference between the average attribute of all descendant species (including immigrants), and the average attribute of the non-immigrant descendant species. The immigration term equals zero if there are no immigrants in a time bin, or if the immigrant and non-immigrant descendants are identical in terms of their mean attribute.

푖 The last term of the Price equation (푎푣푒((∆푧)푗)) represents anagenetic (within-lineage) change and was a part of Price’s original equation (1970). It is often referred to as the “remainder term” after species selection is parsed out (see Frank, 2012). This term equals the average attribute change from all ancestors to their descendants. Ancestors without descendants

(extinctions) and descendants without ancestors (immigrants) do not contribute to anagenetic change (Rankin et al., 2015). If a speciation event resulted in two descendant species, each descendant’s geographic range size was independently subtracted from the ancestor’s geographic range size. Following this, the average of these two numbers (each descendant minus ancestor) was taken before the average for the whole time bin was calculated. The anagenetic change term can equal zero if there are no changes in geographic range size between the descendants and their

16 ancestors, or if the descendants have the same geographic range size on average as their ancestors.

I used Equation 1 to calculate the contributions of species selection, anagenetic change, and immigration to macroevolutionary change in mean geographic range size from each time bin to the following time bin. Calculations were performed in Microsoft Excel 2019.

2.2: Dataset Selection and Preparation for Analysis

A suitable dataset for the Price equation must include geographically isolated species with traceable ancestor-descendant relationships and a measurable trait of interest. The species must be geographically isolated to allow identification of immigration; if the geographic range size was too large, then immigrant species would be difficult to detect. Ancestor-descendant pairs must be present in order to trace macroevolutionary changes. Lastly, there must be a measurable trait with variation (e.g. body size (Rankin et al., 2015)) to calculate species selection. A fossil dataset with all of these criteria needed to be selected (also with a large enough sample size).

Two datasets were selected for this study: One containing Cretaceous gastropods (Hunt et al., 2005) and another containing Cretaceous bivalves (unpublished dataset provided by D.

Jablonski). Both datasets were originally collected by Jablonski, and were based on museum collections and a review of published literature of the time (see Jablonski, 1987). Both gastropods and bivalves had traceable species-level ancestor-descendant relationships presented in a species-pair list created by Jablonski. This allows for branching events and anagenetic change along branches to be observed. Geographic range sizes were also included in the dataset.

Geographic range size was used as the trait of interest since Jablonski (1986, 1987) had previously used it to predict the survivability of these two molluscan groups.

The biostratigraphy needed to be updated before the Price equation could be used. The biostratigraphy Jablonski had used in his initial studies could not be found. Also, Jablonski’s original studies had been done in the 1980’s and an updated biostratigraphy using modern sources needed to be done. Two methods were used to resolve the biostratigraphy of the Gulf and Atlantic Coastal Plain of the late Cretaceous: First, a review of the literature was done and second a unitary association was done using the PAST software (see Appendix 3 and Appendix

4). The species-pair lists provided by Hunt et al. (2005) and Jablonski were used for the literature review. Occurrence data for the gastropods and bivalves from the datasets was downloaded from the Paleobiology Database (PBDB) (https://paleobiodb.org/navigator/). This occurrence data was limited to the Gulf and Atlantic Coastal Plain region of North America, which includes

Maryland, New Jersey, Alabama, North Carolina, South Carolina, Mississippi, Tennessee,

Georgia, and Texas. Species existing outside of this region were excluded (e.g. those from

Mexico). Formations containing the relevant species were aligned based on published age data and index taxa (see Aurisano, 1989; Farke & Phillips, 2017; Gallagher, 2003; Larina et al., 2016;

Pessagno, 1969; Sohl, 1964; U.S. Geological Survey, 1978; Wade, 1926).

I divided my data into nine time bins of approximately equal lengths of time

(approximately two million years in length) in order to allow analysis of macroevolutionary change over time. Each time bin was given a label of a letter/number pair (e.g. the first time bin is labelled T1). Once these time bins were established, the taxa were assigned to the appropriate time bins based on their temporal ranges. Species that persisted through multiple times bins were counted as range-through species. For example, if a species was present in T1 and T4, then it was

18 also assumed to be present in T2 and T3. Ancestor and descendant species found in the same time bin indicated a branching event.

For the Price equation analysis, the change in the attribute of each species needed to be analyzed from one time bin to the next. The time bins were used to construct time increments with each time increment spanning two adjacent time bins (e.g. T1-T2). To document macroevolutionary change over time, each time bin needed to reflect a transition of geographic range size between ancestor and descendant.

Some assumptions had to be made in order to complete the Price equation analysis.

Firstly, Jablonski organized the data into presumed ancestor-descendant pairs based on accepted literature (Jablonski, 1987). Hunt et al. (2005) updated the gastropod list, but no phylogenetic analysis has been performed for the gastropods or the bivalves. Some of the species (both the ancestors and the descendants) in the bivalve dataset were excluded from the present analysis due to insufficient records. These species (eight in total) had an incomplete record in the PBDB

(either missing formation information or no collection information), so could not be used.

Immigrants were considered to be any first appearance of a species in a time bin where no ancestor had previously existed. Again, there is a possibility there was a sampling error or preservation bias exists where the supposed immigrant could have been a local species the entire time. However, for the purposes of this study, these non-ancestor species were considered immigrants. Immigrants were considered to be under “total descendants” although a separate category labelled “total descendants (w/no immigrants)” was identified in order to aid in calculations.

A species was considered to be a range-through species if it appeared in more than one time bin, even if those time bins were not adjacent. A species was considered to be an immigrant

19 if it appeared without having any ancestors present in previous time bins. Once a species disappeared from the regional fossil record (i.e. no longer appearing in time bins), it was considered to have gone locally extinct. It was assumed the extinction event occurred in the time bin immediately after the last appearance of the species.

Hunt et al. (2005) and Jablonski (unpublished data) reported geographic ranges in kilometers (km) rather than square kilometers (km2). This is because the geographic ranges were originally mapped on a 5000-km discontinuous outcrop belt (Jablonski, 1987). Geographic range size of a species during a time bin was the maximum linear distance between localities in which that species’ fossils were found during that time bin. When discussing geographic range size, I considered a large geographic range to be greater than 500 km while a small range was less than or equal to 500 km. These values were chosen based on the spread of ranges in the data; species tended to have ranges that were very small (less than or equal to 30 km) or very large (greater than 1000 km). Therefore, 500 km was chosen as the division between small and large range sizes.

Dividing the end-Cretaceous into approximately equal two-million year time bins is a difficult process. Any refinement of time using the fossil record is challenging, and this is especially true of the end-Cretaceous where sections of the end-Cretaceous are absent in the Gulf and Atlantic Coastal Plain (the last one million years) (see Jablonski, 1986). In order to get the refinement of two-million year time bins, different formations from different states were used

(e.g. Ripley formation of Mississippi for the T4 time bin). Some of the formations and biozones overlap temporally. For example, the N. alternatum and Ripley formation of Tennessee and

Mississippi respectively overlap. Assumptions were made about these time bins in order to perform a Price equation analysis following Jablonski’s original methods. The formations and

20 biozones used had to be cut to fit a two-million year time bin. For example, the Ripley formation, which is roughly three to four million years, was cut to make up a two-million year time bin.

This meant fossils were recorded in the time bin associated with the Ripley formation.

A unitary association was done using the PAST software as a check of the biostratigraphy aligning the Gulf and Atlantic Coastal Plain formations of the late Cretaceous. A unitary association was chosen because of the lack of strictly regional data in this study (see Guex,

1991). Based on the method used to establish the time bins, the unitary association was expected to differ from the previously established biostratigraphy used to perform the Price equation analysis. The species used in this analysis and the results of the PAST software analysis can be seen in Appendix 3 and Appendix 4.

To set up the analysis, I gathered data on species found in each formation. The formations used for the unitary association were those identified in the initial biostratigraphy analysis. Common marine invertebrate species of the Gulf and Atlantic Coastal Plain of the late

Cretaceous were chosen which included ammonites, gastropods, bivalves, and a species of dinoflagellate. A few endemic species (i.e. those found in one formation or in one states) were included because of their prevalence in their respective localities. Some species selected were part of the datasets used in this study which was unavoidable because of their usefulness in a biostratigraphy analysis (e.g. the bivalve Exogyra costata). Vertebrates were excluded from this analysis. Data from the PBDB was downloaded and input into the PAST software.

Approximately sixty-six species were chosen for this analysis (Appendix 3). The results of the unitary association differed slightly from the original biostratigraphy alignment based on the literature. Eleven cliques were found but because of repeating formations, nine cliques can be observed (equivalent to the number of time bins). The cliques in what would be the middle of the

Maastrichtian agree with the literature review. However, the first and last cliques disagree (e.g. the PAST analysis said E. costata zone was the oldest while the literature review determined it should have been the G. elevata zone). The literature review was used as the primary source of biostratigraphy because of the multiplicity of sources that have lined up these Gulf and Atlantic

Coast formations. The refinement in the biostratigraphy may also not be detected by the unitary association, especially due to the amount of endemic species used in the analysis.

CHAPTER THREE: RESULTS

3.1: The Price Equation Results

The mean geographic range size of the Atlantic and Gulf Coast gastropods and bivalves

from the Campanian through the Maastrichtian fluctuated, rising and falling throughout the eight

time increments (refer to Figure 1 for gastropods and Figure 2 for bivalves). For the gastropods,

the total mean change was negative for the T1-T2 time increment. Both species selection and the

anagenetic change were favoring a larger geographic range size, but the values for both were

small. The immigration term was negative and indicated that those immigrants coming into the

Gulf and Atlantic Coastal Plain had small geographic ranges which brought down the total mean

change value. The bivalves had an increase in their geographic range size during the T1-T2 time

increment. Species selection mainly contributed to this increase while anagenetic change and the

Figure 1: The total mean geographic range size change (blue line) gives the Price Equation answer for the gastropod analysis. In comparison, the three separate Price Equation terms (species selection, anagenetic change, and immigration) plotted as well.

23 immigration both had negative values (decreased geographic range size). Species selection was

Figure 2: The total mean geographic range size change (blue line) gives the Price Equation answer for the bivalve analysis. In comparison, the three separate Price Equation terms (species selection, anagenetic change, and immigration) plotted as well. so high, it nullified the effects of the other two terms. There was one extinction event present for the gastropods but fifteen extinction events (fifteen species went extinct in this time increment) for the bivalves. There were no speciation events for either the gastropods or the bivalves during the T1-T2 time increment.

The first major increase in geographic range size for both the gastropods and the bivalves occurred in the T2-T3 time increment. For both groups, species selection, anagenetic change, and immigration collectively increased geographic range size. Large-ranged immigrants were the main contributors to the increase in geographic range size for the gastropods and bivalves

(immigration term value of 287 km for the gastropods and 526 km for the bivalves). The gastropods had two speciation events while the bivalves had one speciation event during the T2-

T3 time increment. Also, nine species of gastropods went extinct while four species of bivalves went extinct during this time.

Time bins T3-T4, T4-T5, and T5-T6 represent times of limited change in the data where there were no substantial increases or decreases in geographic range size for the gastropods and the bivalves. In the T3-T4 time increment, the gastropods had a decrease in geographic range size mainly attributable to a large number of small-ranged immigrants moving into the area.

Anagenetic change also drove smaller geographic range size while species selection, though small in value, selected for larger geographic range sizes among gastropods. The bivalves experienced an increase in their total mean geographic range though it was slight. Species selection and immigration both increased geographic range size while anagenetic change decreased geographic range size. The result of this was a low, but still present increase in geographic range size. The gastropods experienced two extinction events and five speciation events, while the bivalves experienced five extinction events and three speciation events in the

T3-T4 time increment.

The next time increment (T4-T5) had the highest number of ancestor species under study for both groups (n=88 for the gastropods and n=60 for the bivalves). The increase in numbers throughout the previous time bins indicates the amount of immigration and to a lesser effect speciation (and small number of extinctions) occurring in the late Campanian through the early

Maastrichtian. Both the gastropods and the bivalves experienced increases in their geographic range size during the T4-T5 time increment. For the gastropods, species selection and anagenetic change contributed to an increase in geographic range size. The immigrants moving in had small ranges but their effect was not enough to decrease the total mean change. For the bivalves, total mean change was attributable to species selection. There was no anagenetic change and

25 immigration (both terms were equal to zero) during this time increment. Both groups appeared to have increasing geographic range sizes while simultaneously undergoing more extinction events

(19 for the gastropods and 14 for the bivalves).

Like the previous time increment, during the T5-T6 time increment both gastropods and bivalves had an increase their geographic range size, though the change was not high.

Anagenetic change contributed the most to the increase but species selection also added to the increase in geographic range size for the gastropods. The immigrants at this time had small geographic range sizes but did not decrease the total mean increase in geographic range size among gastropods. There was a total mean increase in the geographic range size of the bivalves mainly because of species selection. Anagenetic change decreased the total mean change a bit while the immigrants did not contribute much to the total mean geographic range size of the bivalves. In terms of extinction events, the bivalves had five extinction events while the gastropods had a total of 26 species that went extinct. There were only two speciation events for the bivalves and no speciation events for the gastropods.

There was a decrease in geographic range sizes for both the gastropods and the bivalves during the next time increment (T6-T7). The gastropods also experienced a high number of extinctions among species (23 in total). The bivalves did not have as many extinctions as the gastropods (losing 10 species). The total mean geographic change during this time bin were about equal between the two groups: -470 km for the gastropods and -485 km for the bivalves.

All three macroevolutionary forces contributed to the decrease in geographic range size for both groups. For the gastropods, the anagenetic change term (-286.05 km) was the main contributor while for the bivalves the main contributor was the immigration term (-199.76 km).

A large increase in the total mean geographic range sizes of both groups occurred in the subsequent time increment (T7-T8). The increase was mainly attributable to species selection for both groups. Anagenetic change increased geographic range further for the gastropods while the effect of immigration was small. For the bivalves, both anagenetic change and immigration decreased the total mean change in geographic size but were not as large as the effect of species selection. The gastropods experienced 19 extinctions in this time increment while the bivalves experienced 14 extinctions. There were no speciation events for either group.

The last time increment (T8-T9) corresponded with the end of the Cretaceous period

(approximately 68-66 million years ago). The T8-T9 time increment saw a decrease in total mean geographic range size for the gastropods in comparison to the last time increment. The bivalves, however, experienced an increase in their total mean geographic range size in comparison to the previous time increment. Species selection had the largest effect for both groups and increased geographic range for both groups (+150 km for the gastropods and +441 km for the bivalves). Anagenetic change increased geographic range sizes for both groups. There were no immigrants for either group in this time increment. No speciation effects occurred for either group during this time bin. The gastropods experienced five extinctions and the bivalves experienced 16 extinction events.

3.2: Statistical Analysis of Datasets

A permutation test for each time increment of both datasets was performed. The null hypothesis predicted there would be no relationship between geographic range size and the number of descendants were left by an ancestor. The results of these permutation tests can be

27 observed in Table 1. A significant relationship (null hypothesis rejected) was observed in T7-T8 for the gastropods and bivalves. In this time increment, an increase in survivability (i.e. leaving descendants) for these molluscs was partly dependent on geographic range size. Further investigation shows that the average descendant geographic range size for the T7-T8 time increment was large (1318.18 km for gastropods and 2346.83 km for bivalves (immigrant ranges not included)).

Table 1: Results of permutation tests for each time increment. P values are reported for tests of the null hypothesis that geographic range size and number of descendants left are independent of each other. Separate tests were conducted for each group (gastropods or bivalves) and time increment. Time Increment Gastropod p-value Bivalve p-value

T1-T2 0.283 0.093

T2-T3 0.217 0.311

T3-T4 0.379 0.894

T4-T5 0.222 0.117

T5-T6 0.616 0.334

T6-T7 0.313 0.328

T7-T8 0.001 0.003

T8-T9 0.207 0.115

3.3: Further Analysis of Datasets

The extinction proportion of each time bin was calculated. This was done by taking the proportion of extinction events over the number of ancestors for each time increment. For

28 example, in the first time increment for the bivalves (T1-T2), there were initially forty ancestors in T1 and in T2 there were fifteen extinction events (i.e. ancestors not found in this time bin).

The proportion of extinction events to ancestors was taken. The results can be observed in

Figure 3. The number of extinction events increased steadily over the Campanian and

Maastrichtian for both the gastropods and the bivalves. The bivalves had a higher percentage of extinction events than the gastropods. Both groups saw the highest amount of extinction events occur during the T7-T8 Proportion of Exintction of Molluscs throughout the End-Cretaceous

Proportion of Extinction (in %) (in of Extinction Proportion 10

0 T1-T2 T2-T3 T3-T4 T4-T5 T5-T6 T6-T7 T7-T8 T8-T9 Time Increment

Gastropods Bivalves

time increment, which corresponds with the last four million years of the Cretaceous.

Figure 3: Proportion of extinctions in both gastropods and bivalves in each time bin from Campanian to KPg boundary.

CHAPTER FOUR: DISCUSSION

Both datasets present three time periods of rapid macroevolutionary change in the geographic range size. These periods of rapid change occur in the same time increments for both mollusc groups. The first increase occurs in the time increment T2-T3 while the other changes (a decrease and increase in geographic range respectively) fall in later time increments T6-T7 and

T7-T8. That both groups exhibit simultaneous periods of rapid change in mean geographic range size in the same direction suggests that these changes reflect biological causes common to both groups, rather than sampling error. For both groups, each major instance of rapid change in mean geographic range size is the result of different combinations of macroevolutionary forces, quantified by the terms of the Price equation. The differences in survivability of ancestor and descendant species (and in some cases genera) can be noted from the data. Species selection, when strong, can be attributed primarily to differential extinction of ancestral species based on their geographic range size.

4.1: Macroevolutionary Change Within Time Increment T2-T3

For both gastropods and bivalves, a major increase in geographic range size occurred at

T2-T3, a time increment that falls in the late Campanian/early Maastrichtian border

(approximately 72 million years ago). The immigration term is the main contributor of this increase for both groups. An influx of immigrants into the Gulf and Atlantic Coastal Plain near the end of the Campanian was most likely the result of a global flooding event. Many studies have analyzed the change in Cretaceous seas prior to the major extinction event, finding a dramatic cycle in sea level changes from the Campanian through the KPg boundary and into the

Danian (Gallagher, 1991; Habib & Miller, 1989). These cycles of sea-level change have been observed from the northern-most points (present-day New Jersey, Delaware, and Maryland) down to the southern points (present-day South Carolina and Georgia) (Gallagher, 1991; Habib

& Miller, 1989 respectively) of the Gulf and Atlantic Coastal Plain. The Atlantic coast in particular during the Campanian experienced alternating periods of marine regression and transgression, with a distinguishable late-Campanian/early-Maastrichtian transgression event

(Habib & Miller, 1989). The T2-T3 time increment falls near the end of this time, so the results most likely represent the aftermath of the transgression event. The free-swimming veliger larva shared by gastropods and bivalves could have enabled rapid dispersal and range expansion during this time (Pechenik, 2005; Scheltema & Williams, 1983). The flooding event and the resulting substantial influx of immigrants with large geographic ranges increased the total mean geographic range size of the molluscan species found during this time.

4.2: Macroevolutionary Change Within Time Increment T6-T7

The next major change in geographic range size occurs at T6-T7, approximately 69-68 million years ago. This time increment falls in the middle to later part of the Maastrichtian.

Unlike the previous major change in the data, different macroevolutionary forces affected the total mean decrease in geographic range size for the gastropods and bivalves. For the gastropods, the decrease in range size was primarily due to anagenetic change and species selection.

Immigration did not significantly contribute to the decrease. There was a higher number of local extinctions events compared the previous time increments of both larger ranged and smaller ranged species.

Global climate change may have facilitated this decrease. Changes in global climate took place throughout the Maastrichtian with nannofossil evidence indicating subtropical and tropical waters cooling, and more sharply defined climate belts appearing during the last stages of the

Cretaceous (Keller, 2001; Worsley, 2012). By the end of the Maastrichtian, this cooling may have been coupled with another regression event in the Cretaceous (Keller, 2001). With a declining sea level, there could have been a subsequent decline in available shelf habitats and potentially increased competition amongst the shelf-dwelling inhabitants (Keller, 2001). It has been found that times of marine regressions correspond with decreases in generic diversity because of decreased available shelf space (Hallam, 1989). In later periods such as in the

Paleogene, marine regression events may have led to delta formations and, with this, detritus matter filling once clear water habitats (Dockery, 1986). Environments restrictions such as these could have led to decreases in geographic range size. The immigrants from the T2-T3 time increment showed a high survival rate during this time with a of 64%. Of these survivors, 67% had a large geographic range (greater than 500 km).

For the bivalves, all three macroevolutionary forces decreased mean geographic range size. Unlike in the gastropods, small-ranged immigrants had the greatest effect in decreasing mean geographic range size. Global climate change and marine regression could have also impacted this decrease in local bivalve ranges. With shrinking available shelf space caused by a marine regression, bivalves may have been developing smaller geographic ranges. The extinction of some large-ranged bivalves (nine species here out of thirty-seven, with an average range of

2800 km) added to this effect, bringing down the total mean range size during this time increment. All immigrants during this time had a range size under 300 km (highest was 240 km and the lowest was 10 km). These small-ranged immigrants either went extinct or underwent a

32 dramatic range reduction before going extinct over the next two million years. The immigrant species that expanded their ranges into the Gulf and Atlantic Coastal Plain during the T2-T3 time increment had a 53% survival rate in the T6-T7 time increment; and these former immigrant species had an average range size of 800 km or larger.

4.3: Macroevolutionary Change Within Time Increment T7-T8

The last major change in geographic range size in both the gastropods and the bivalves occurred in the T7-T8 time increment which is approximately 68-66 million years ago. For the gastropods, species selection and anagenetic change favor a larger geographic range size. The effect of the small-ranged immigrants’ did not impact the total increase in geographic range size.

For the bivalves, the effect of species selection was substantial enough to increase the total mean geographic range size despite anagenetic change and immigration decreasing range size. Many extinction events (19 for the gastropods and 14 for the bivalves in total) occurred during this time increment (see extinction proportions, Figure 3). A majority of the species that survived the T7-

T8 time increment also survived until the end-Cretaceous (T8-T9 time increment for this study).

The results of the Price equation and the statistics done for this time increment show a major upheaval in the Gulf and Atlantic Coastal Plain. This was the only time increment in which there was a significant statistical result that showed survivability was dependent on geographic range size. The average descendant geographic range for both the gastropods and the bivalves was large, indicating survivability (or large-ranged ancestors leaving large-ranged descendants) was dependent on having a large geographic range.

The selective extinction of small-ranged species could be based on a variety of factors.

The end-Maastrichtian was a time of global cooling, even in subtropical waters (Worsley, 2012).

There is evidence for a regression event occurring near the end-Maastrichtian-Danian border

(Habib & Miller, 1989). This decrease in sea level could have precipitated selective extinction of small-ranged species. One might expect that a reduction of the available shelf space would cause species selection to favor a decrease in geographic range size--i.e. reduced mean range size due to anagenetic change, rather than increased mean range size due to species selection. However, the opposite was observed in the data. Species selection favored an increase in geographic range size during this time despite shrinking available shelf space. Small-ranged species were more likely to go extinct while large-ranged species were selected for; because of their larger range sizes, it was less likely that the regression would eliminate their entire range. With the increase in total extinctions, species with a more cosmopolitan distribution could have the opportunity to increase their own range into available space. Assuming prehistoric marine gastropods employed similar feeding and lifestyle strategies as modern prosobranchs, species with a more cosmopolitan distribution could have had the potential to expand their geographic range despite decreasing shelf space. This is what Hansen (1988) referred to as an “initial recovery phase” post-extinction event where survivors begin the initial process of recolonization into empty ecological niche spaces. Hansen (1988) also noted that those local survivors (albeit in Texas) were smaller-bodied deposit feeders from species-poor genera, and carnivores/scavengers. These surviving species with a varying diet and lifestyle may have expanded their range sizes into the vacant niche space for some time until the end- Cretaceous where the data showed a reduction of range mainly driven by species selection.

4.4: Periods of Macroevolutionary Stasis in the Data

Between T3 to T6, three time increments of six million years in total, mean geographic range size of both gastropods and bivalves experienced little change. The terms of the Price equation were all small in magnitude, indicating that modest changes in mean range size were not due to strong opposing macroevolutionary forces canceling one another out. The early to mid-Maastrichtian has been described as a time of change most notably with a rapid cooling of the oceans and the formation of ice in polar regions (Hemleben, Friedrich, & Herrle, 2005).

Despite the environmental shift of this time period, diversification and niche exploitation of some vital marine species, such as photosynthetic foraminifera, were still present (Abramovich et al., 2003). Cycles of transgression and regression events define the time from the late Campanian to the mid-Maastrichtian (encompassing the T3-T6 time increments) (Habib & Miller, 1989).

Despite the environment being in a state of flux, the changes were not enough to cause the extinction of many species and genera. The beginning of this period remained a mix of species with both smaller and larger geographic ranges, immigrants and non-immigrants, and more speciation than extinction events. The extinction events for the gastropods began to increase in frequency during the T5-T6 time increment and continue to exceed the speciation rate for the remainder of the Maastrichtian. As for the bivalves, extinction events also began to occur in the T4-T5 time increment while simultaneously speciation events became less frequent.

Steady immigration for both groups may have contributed to the lack of major change in geographic range size. Immigrants may have replenished species’ numbers (and kept geographic range consistent) post-extinction. Also, despite there being environmental changes, these changes were not enough to cause the mass extinction of many groups (Worsley, 2012). The speciation and extinction events occurring during this period may have been species-specific and not due to any correlation with an emergent property such as large geographic range size. This

35 idea supports Jablonski’s hypothesis about background extinctions; this time may represent a time of “normal” background extinction with large geographic range coupling with species richness to ward off extinction for some species (see Jablonski, 1986).

4.5: Differential Survivorship Among Cretaceous Molluscs

Attempting to determine the central cause of a mass extinction event is difficult because of many potential impacting factors (Hallam, 1989). For the end-Cretaceous mass extinction, global climate change, sea level change, and volcanism are among many environmental conditions that could have impacted marine invertebrates at the time (Hallam, 1989). Some of these conditions have been discussed above as potentially impacting the Gulf and Atlantic Coast molluscs and contributing to their local extinctions. Both gastropods and bivalves suffered great losses (Figure 3), with an increase in local extinctions throughout the end-Cretaceous Period.

Molluscs on a global scale lost great diversity with groups such as inoceramids (bivalves), the reef-building rudists (bivalves), and the ammonites (cephalopods), being lost completely before the extinction event or shortly after (see Marshall & Ward,1996; Lehmann, 1981; Stanley, 1984;

Raup & Jablonski, 1993). Excluding ammonites, bivalves appeared to be the molluscan class that suffered the greatest loss with an extinction of 63% at the genus level (Raup & Jablonski, 1993).

However, the loss was concentrated in of two large groups (the inoceramids and rudists)

(Jablosnki & Raup, 1995).

The cause (or causes) of differential survivorship among moullscs during the end-

Cretaceous has been extrapolated in different ways. Much of the discussion around geographic range size relates the inverse relationship between large geographic range size and extinction rate

(Jablonski & Roy, 2003). Jablonski (1986) hypothesized a difference in survivorship based on

36 developmental mode, large geographic range size, and species richness during times with background extinctions. During background extinctions, species richness and mode of larval development may impact differential survivability, but during a “mass extinction regime” geographic range size of a taxon would be the only extinction deterrent (Jablonski, 1986).

Geographic range size is considered to be an emergent property shared by ancestors and their descendants; and there is a positive correlation between geographic range size and geologic duration of a taxa (Jablonski, 1987). In this study, the connection between geographic range size and a mass extinction event was the primary focus. It has been proposed that a large geographic range size may promote speciation by dividing populations via geological barriers (i.e. mountains) or by forming peripheral isolates (Jablonski & Roy, 2003). Additionally, a broader geographic range could expose species to varying environmental conditions to which their populations can diverge and adapt (Jablonski & Roy, 2003). However, Jablonski and Roy (2003) found that this is not always the case as their gastropod dataset presented a negative relationship between geographic range size and species richness. Generally, though, this is interpreted as a safe estimate among fossil taxa (Jablonski & Roy, 2003).

Gallagher discounts large geographic range being the sole determining factor to prevent the extinction of a group (accounting for the large ranges of ammonites and exogyrids both of which went extinct at the KPg boundary) (Gallagher, 1991). Gallagher proposes that there are two causes of molluscan decimation at the end-Cretaceous: the planktonic population crash at the end-Cretaceous and, consequently, the decrease in bioturbation due to the reduced infauna

(Gallagher, 1991). This line of reasoning is similar to what Alvarez et al. (1980) claims about the decrease in sunlight from an asteroid impact, and how this event triggered a decimation of land and marine photosynthesizers, including plankton and algae. Those mass extinction survivors

37 exhibited a variety of reproductive strategies that were independent of the planktonic food supply

(such as corals and sponges with asexual reproduction, or lophophorates with lecithotrophic larvae) (Gallagher, 1993) Suspension feeders, or those whose trophic requirements were based on the planktonic supply in the oceans, were also more likely to go extinct than deposit feeders

(Gallagher, 1991, 1993). The reduction of planktonic infauna also decreased rates of bioturbation leading to poor substrate for infaunal species but did not affect epifaunal species nearly as much

(Gallagher, 1991). However, it may not have been an advantage being a deposit feeder since benthos is dependent on what is living in the water column, meaning the planktonic crash would also dramatically affect benthic communities (Levinton, 1996).

Other hypotheses have been made regarding what caused differential survival of some molluscs over others. A difference in body size and lifestyle (epifaunal vs. infaunal) among molluscs did not appear to be a determining factor in their survivability (Jablonski & Raup,

1995). Despite all the hypotheses, geographic range size remains an potential explanation for extinction resistance. The data in this study cannot completely disprove other hypotheses stated above. Jablonski’s hypothesis proposing a correlation between large geographic range and increased survivability appears to hold up in this study even when considering the nature of the data collection and assumptions made about the data. Those species with a larger range, and therefore exposed to more variable environmental conditions, would be expected to have a higher survival rate during a mass extinction characterized by sea level change and global climate change. Conversely, when dealing with paleontological data, these larger ranged species are also more likely to be found than endemic species, and therefore may influence the data more strongly. Inferences about the survivability of these larger ranged species over their short-ranged counterparts then must be conservative in nature. Despite an 80% loss of organisms at the

38 species-level during the Cretaceous mass extinction, genera biodiversity was maintained

(Hansen, 1988).

Surviving the mass extinction was not a guarantee for later success. The so-called “Dead

Clade Walking” phenomenon has been suggested as a way in which those survivors can still be impacted by the extinction and may go extinct later after failing to diversify (Jablonski, 2002).

As for the Gulf and Atlantic Coastal Plain gastropod and bivalve survivors of the end-Cretaceous extinction, the aftermath of the extinction meant a long lag in taxonomic recovery that may have lasted millions of years (D’Holdt, 2005). In New Jersey, for example, marine macrofossil diversity in general did not return to its pre-extinction levels until the Thanetian at the end of the

Paleocene epoch (Gallagher, 2003). The Danian saw many Cretaceous mollusc assemblages replaced by sponge-braciopod-coral assemblages, meaning faunal turnover at high levels

(Gallagher, 2003). When looking into the Paleogene with this data, a majority of both gastropod and bivalve species present in the last time bin (T8-T9) went extinct by the beginning of the

Danian (only four species collectively did persist in the Danian). The “Dead Clade Walking” phenomenon may hold some significance despite large geographic range size when it comes to the groups found in this study. Despite losing species members, members of higher taxonomic levels such as genera persisted in the Gulf and Atlantic Coastal Plain throughout the early

Paleocene (Glycymeris for example).

4.6: Limitations of the Study and Significance

As with any study, there are limitations to this one that need to be acknowledged. Firstly, there were very few speciation events that occurred in the data. The maximum number of the speciation events observed in the gastropods and bivalves were 5 and 3 respectively with most

39 time increments having no speciation events at all. Taking this into consideration, speciation events were not discussed at length in comparison to the extinction events that did occur.

The ancestor-descendant relationships used in this study were based on hypothetical relationships (see Jablonski, 1987). Unlike previous applications of the Price equation, phylogenetic tree-based relationships were not used (see Simpson, 2013). The Price equation was able to detect the effects of the Cretaceous mass extinction event on the gastropod and bivalve datasets used. So, even if the results could have been strengthened with additional phylogenetic relationships, the Price equation was able to function with hypothetical relationships As more refinement of molluscan phylogeny is done, these hypothetical relationships along with other ancestor-descendant relationships at lower taxonomic levels (genus and species) can be reaffirmed or reassigned, giving the Price equation more power and applicability. Though the fossil record is difficult to work with in terms of precise tracing of ancestor-descendant relationships, it can nevertheless be used in a quantitative sense to gather a broad understanding of macroevolutionary patterns occurring at a given time.

Hypothesizes about survival based on other attributes (rather than geographic range size) can be discounted slightly. For example, the species that do survive into the last time bin are a mix of both planktonic and non-planktonic in their reproductive strategies (Jablonski unpublished data), which deviates from Gallagher’s hypothesis about the success of the non- planktonic larva over the planktonic larva.

Concerns about the appearance of the data (species) in the geological record can be eliminated. There is generally a lack of data from latest Maastrichtian sites globally because of unconformities in the rock record which can show many rapid extinctions as once (Worsley,

2012). The sudden termination of many branches (ancestors and their descendants) at once

40 cannot be overlooked in light of this. The Price equation is unable to differentiate between gaps in the geological record and actual extinction events. The Gulf and Atlantic Coastal Plain sites, however, combat concerns about conformities because these sites are known to have a rich depositional environment and be very well documented and studied (Jablonski & Roy, 2003).

This area from New Jersey down to Mexico is known to have excellent preservation of organisms down the species-level, and because of this have been studied in extensive detail

(Jablonski & Roy, 2003).

The effect of mass extinctions on species has been well-studied and discussed.

Macroevolutionary change brought about by species selection during a mass extinction may be very different from times of normal background extinction. A good phylogeny of a fossil group could potentially give a different result (more ancestor-descendant pairs). The application of the

Price equation with a larger dataset could prove to be interesting, especially in analyzing mass extinction and post-mass extinction radiations and speciation events.

The results of this study, despite the limitations, show important macroevolutionary changes linked to the emergent character of geographic range size. These changes are observable in both mollucscan groups. Since the Price equation analysis was done on each group separately, this indicates there is a biological pattern present. Also, a major extinction event of these molluscs was quantified before the KPg boundary, distancing the extinction of some marine invertebrates from the asteroid impact. The biostratigraphy completed for this study, though containing assumptions and overlapping in some places, is also beneficial for future analyses of the Gulf and Atlantic Coastal Plain during the end-Cretaceous Period.

CHAPTER FIVE: CONCLUSION

The aim of this thesis was to apply the Price equation to a group of invertebrates to understand how geographic range size impacted their survivability during the Cretaceous mass extinction. Geographic range size has previously been hypothesized to be an emergent character that is heritable between ancestors and descendants on a species level (see Jablonski 1986, 1987).

The Price equation results of this study show three major increases and decreases to geographic range size for both gastropods and bivalves. The first increase at the T2-T3 time increment appears to be driven mainly by an influx of immigrants into the Gulf and Atlantic Coastal Plain as a result of a marine transgression event. The second major event (time increment T6-T7) in which geographic range size decreases may have been brought about by global climate change and a marine regression with decreasing available shelf habitats for both the gastropods and the bivalves. Lastly, a sharp increase of geographic range size occurred in the T7-T8 increment. This increase was brought about primarily by species selection. This time increment, near the end-

Cretaceous, demonstrates selective pressures favoring larger geographic range sizes in both gastropods and bivalves. There was also a statistically significant relationship between survival and geographic range size during this time increment.

There are limitations to this study, especially when tracing ancestor-descendant relationships in the paleontological record at the species level. Species-level macroevolutionary studies are challenging because of small sample sizes and poorly understood evolutionary relationships. As phylogenies are refined, more work can be done using the Price equation on paleontological data. For example, it would be interesting to investigate the effect of the

Cretaceous mass extinction on ammonite body size.

The Price equation can provide an understanding of macroevolutionary changes and lend insight into how groups respond to environmental pressures during climate shifts. This is especially important during the purported sixth mass extinction (see Barnosky et al., 2011), with global climate change impacting many groups, terrestrial and marine alike. While the Price equation is not a method to quantify the possibility of extinctions or the magnitude of them, it can be used to predict likely outcomes and group survivorship.

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APPENDIX 1: Table of gastropod ancestor-descendant pairs of the Gulf and Atlantic Coastal Plain. Species names and ranges from Hunt et al. (2005) Appendix B.

Ancestor Ancestor Range Descendant Descendant (km) Range (km)

Weeksia amplificata (Wade) 690 Weeksia lubbocki 60 Stephenson Planolateralus tuberculosus Sohl 10 Planolateralus conanti 30 Sohl

Planolateralus argenteus Wade 45 Planolateralus 490 microcancelli Sohl

Ataphrus kerri Gabb 160 Ataphrus n. sp. 650 (Providence) Laxispira lumbricalis Gabb 3360 Laxispira monilifera Sohl 3,820 Turritella hilgardi Sohl 2250 Turritella kellumi 10 Stephenson Turritella mcnairyensis Wade 3110 Turritella houstoni 30 Stephenson

Turritella hillensis Stephenson 10 Turritella kerrensis 30 Stephenson Turritella tippana Conrad 3,480 Turritella vertebroides 4020 Morton Haustator? howelli (Harbison) 640 Haustator? 3480 640 paravertebroides (Gardner) Haustator? trilira (Conrad) 4,250 Haustator? bilira 3950 (Stephenson) Nudivagus cooperensis 10 Nudivagus simplicius 10 Stephenson Wade Trobus corona (Conrad) 640 Trobus buboanus 310 Stephenson Cerithiella n. sp. (Coffee) 20 Cerithiella nodolirata 50 (Wade) Alvania (s.l.) costata (Wade) 10 Alvania (s.l.) 2170 tallahatchiensis Sohl

Capulus cuthandensis Stephenson 1,250 Capulus spangleri 1,540 Henderson

Capulus monroei Sohl 10 Capulus? microstriatus 10 Stephenson Capulus monroei Sohl 10 Capulus? microstriatus 10 Stephenson Graciliala calcaris (Wade) 1,860 Graciliala decemlirata 720 (Conrad)

Drepanochilus quadriliiratus 10 Drepanochilus triliratus 1320 (Wade) Stephenson Drepanochilus quadriliiratus 10 Drepanochilus triliratus 1,320 (Wade) Stephenson

Anchura substriata Wade 1,990 Anchura abrupta Conrad 1,540 Anchura lamari Stephenson 300 Anchura bexarensis 10 Stephenson

3,520 Latiala elegans 90 Latiala lobata (Wade) (Stephenson) Pugnellus densatus (Conrad) 3,530 Pugnellus goldmani 3,950 Gardner

Granosolarium coffea Sohl 10 Granosolarium 10 voragiformis (Stephenson)

Pseudomalaxis patens Stephenson 10 Pseudomalaxis? 10 monmouthensis (Gardner)

Pseudomalaxis pillsbryi Harbison 2,250 Pseudomalaxis ripleyana 690 Wade

Echinimathilda unionensis Sohl 640 Echinimathilda corona 2,230 Sohl Clathrobaculus cretacea (Wade) 10 Clathrobaculus parvula 640 Sohl Gegania n. sp. (Eutaw) 10 Gegania n. sp. (Coffee) 10

Gegania bella (Conrad) 1,990 Gegania manzaneti 60 (Stephenson)

Gyrodes major Wade 2,610 Gyrodes supraplicatus 3730 (Conrad) Pseudamaura lepta Sohl 2,120 Pseudamaura lirata 2,050 (Wade)

Morea marylandica Gardner 3,720 Morea transenna 1,920 Stephenson

Morea corsicanensis 1,320 Morea cancellaria Conrad 1,920 corsicanensis Stephenson Schizobasis depressa Wade 10 Schizobasis immersa 690 Wade

Lowenstamia liratus (Wade) 690 Lowenstamia cucullata 650 Sohl Stantonella ripleyana (Conrad) 700 Stantonella interrupta 2,640 (Conrad) Stantonella rugosa (Stephenson) 10 Stantonella subnodosa 10 Wade Protobusycon cretaceum? (Wade) 10 Protobusycon binodosum 10 Sohl

Pyrifusus subliratus Wade 10 Pyrifusus crassus Sohl 650

Pyrifusus ejundicus Sohl 10 Pyrifusus subdensatus 10 Conrad

Deussenia ripleyana Harbison 750 Deussenia bellalirata 2670 bellalirata (Conrad) Deussenia corbis Stephenson 20 Deussenia multilirae 20 Stephenson

Bellifusus angulicostatus Sohl 700 Bellifusus spinosus Sohl 1,390

Drilluta communis Wade 1,230 Drilluta distans (Conrad) 1,930

Drilluta major Wade 750 Drilluta lemniscata Sohl 640 Paleopsephaea pergracilis Wade 10 Paleopsephaea tenuilirata 10 Sohl

Graphidula melanopsis (Conrad) 650 Graphidula multicostata 1,770 Stephenson

Ripleyella elevata (Wade) 80 Ripleyella pulchra 30 (Stephenson)

Pornosis digressa Wade 860 Pornosis modica Sohl 300

Haplovoluta triliratus Sohl 650 Haplovoluta quadriliratus 500 Sohl

Euthriofusus? convexus (Wade) 10 Euthriofusus? mesozoicus 10 (Wade) Remera stephensoni Harbison 700 Remera flexicostata Sohl 1,920

Lupira disparila Sohl 10 Lupira turbinae Sohl 80

Lupira variabilis (Wade) 2,300 Lupira pyriformis 1,930 Stephenson

Pyropsis spinosus (Wade) 10 Pyropsis elongatus 10 (Stephenson)

Pyropsis perlata (Conrad) 650 Pyropsis lanhami 10 Stephenson

Pyropsis perornatus (Wade) 10 Pyropsis prolixa Sohl 10

Hydrotribulus nodosus Wade 10 Hydrotribulus elegans 30 Sohl

Longoconcha imbricatus Sohl 10 Longoconcha 10 tennesseensis (Wade) Volutomorpha splendida Sohl 10 Volutomorpha mutabilis 10 Wade Volutomorpha dumasensis Dall 640 Volutomorpha valida Sohl 30

Volutomorpha eufaulensis 10 Volutomorpha producta 30 (Conrad) Sohl

Liopeplum leiodermum (Conrad) 1,900 Liopeplum canalis 2,030 (Conrad)

Liopeplum thoracicum (Conrad) 860 Liopeplum tabulatum 470 (Stephenson)

Liopeplum spiculatum Sohl 1,960 Liopeplum cretaceum 3,950 (Conrad)

Liopeplum coronatum Sohl 10 Liopeplum nodosum Sohl 610

Parafusus callilateris (Wade) 690 Parafusus saffordi Sohl 80

Cancellaria? mcnairyensis Sohl 10 Cancellaria? matsoni 30 Stephenson Amuletum fasciolatum (Wade) 10 Amuletum wadei Harbison 640

Lutema hubbardi (Stephenson) 60 Lutema simpsonensis 10 Stephenson

Remnita anomalocostata (Wade) 1,630 Remnita biacuminata 1,230 (Wade)

Remnita hastata Sohl 10 Remnita n. sp. cf. R. 10 hastata (Severn)

Beretra gracilis (Wade) 690 Beretra firma Stephenson 40

Fusimilis proxima (Wade) 10 Fusimilis kummeli Sohl 640

Acteon pistilliformis Sohl 690 Acteon cicatricosus Sohl 30

Eoacteon ithyocheilus Sohl 10 Eoacteon ellipticus 1240 (Wade) Eoacteon linteus (Conrad) 2,050 Eoacteon percultus Sohl 950

Nonacteonina tensa Stephenson 10 Nonacteonina deflexa 30 Stephenson

Troostella perimpressa Wade 10 Troostella substriatus 10 (Wade)

Ringicula pulchella Shumard 2,170 Ringicula culbersoni 2,180 Stephenson

Ringicula yochelsoni Sohl 30 Ringicula sufflata 10 Stephenson Oligoptycha americana (Wade) 1,280 Oligoptycha n. sp. (Owl 1,350 Creek) Cylichna secalina Shumard 3,280 Cylichna diversilirata Sohl 1,940

Cylichna intermissa intermissa 80 Cylichna pessumata Sohl 10 Sohl Goniocylichna bisculpturata 690 Goniocylichna elongata 610 Wade Sohl

Cylindrotruncatum carinata 10 Cylindrotruncatum 80 (Stephenson) demersum Sohl

Bullopsis demersus Sohl 10 Bullopsis cretacea Conrad 340

Hemiacirsa americana (Wade) 10 Hemiacirsa cretacea 10 (Wade) Plesioacirsa? gravida Sohl 10 Plesioacirsa? implexa 30 Sohl Plesioacirsa microstriata (Wade) 10 Plesioarcisa wadei 80 Cossman Belliscala crideri Stephenson 3,280 Belliscala rockensis 1,230 Stephenson

Striaticostatum pondi 1,230 Striaticostatum bexarense 1,800 (Stephenson) (Stephenson)

Striaticostatum sparsum Sohl 1,820 Striaticostatum harbisoni 10 Sohl

Opalia? n. sp. (Merchantville) 10 Opalia? fistulosa Sohl 10

APPENDIX 2: Table of bivalve ancestor-descendant pairs of the Gulf and Atlantic Coastal

Plain. Species names and ranges from Hunt et al. (2005) Appendix B.

*not used in this study due to lack of information

Ancestor Ancestor Range Descendant Descendant Range (km) (km)

Cuspidaria jerseyensis 2190 Cuspidaria grandis 2250 (Weller) (Stephenson)

Nucula nacatochana (Stephenson) 1930 Nucula microstriata 3230 (Gardner)

4250 Nucula severnensis 1310 Nucula percrassa (Conrad) (Wingard and Sohl)

10 Nucula stantoni 150 Nucula prepercrassa (Stephenson) (Stephenson)

Nuculana corbetensis (Stephenson) 10 Nuculana coloradoensis 10 (Stephenson)

Nuculana kerrensis 160 Nuculana tarensis 890 (Stephenson) (Stephenson)

Scabrotrigonia eufalensis (Gabb) 3830 Scabrotrigonia 2650 angulicostata (Gabb)

Vetericardiella crenalirata (Conrad) 2300 Vetericardiella subcircula 2300 (Wade)

"Tellina" patula (Stephenson 1941) 30 "Tellina" marcosensis 10 (Stephenson 1941)

Agerostrea falcata (Morton) 4020 Agerostrea mesenterica 4100 (Morton)

Anomia argentaria (Morton) 4250 Anomia tellinoides 3580 (Morton)

Anomia preolmstedi (Stephenson) 50 Anomia olmstedi 890 (Stephenson)

Aphrodina eufaulensis (Conrad) 1900 Aphrodina n. sp. (Coon 2050 Creek) [A. eufaulensis of Wade 1926]

Aphrodina tippana (Conrad) 4250 Aphrodina regia (Conrad) 1470

*Arcopsis nolani (Stephenson) 1240 *Arcopsis nolani 750 (Stephenson)

Botula? ripleyana (Gabb) 2300 Botula? carolinensis 3150 (Conrad)

Brevicardium parahillana (Wade) 2050 Brevicardium fragile 1960 (Stephenson)

Brevicardium tenue (Stephenson) 110 Brevicardium 10 guadalupense (Stephenson)

Caestocorbula crassaplica (Gabb) 4250 Caestocorbula? williardi 1920 (Wade)

Camptonectes burlingtonensis 4020 Camptonectes virgatus 3780 (Gabb) (Nilsson)? [C. argillensis (Conrad)

Caryocorbula? carolinensis 160 Caryocorbula? oxynema 860 (Conrad) (Conrad)

Costellacesta riddlei (Kauffman) 30 Costellacesta sayrei 240 (Stephenson)

Crenella? serica (Conrad) 4100 Crenella? elegantula 4250 (Meek & Hayden)

Cucullaea capax (Conrad) 3780 Cucullaea littlei (Gabb) 2330

Cyclorisma alta (Conrad) 850 Cyclorisma? parva 3950 (Gardner)

Cymbophora apressa (Gabb) 4250 Cymbophora wordeni 4250 (Gardner)

Cymbophora trigonalis 3530 Cymbophora cancellosa 2790 (Stephenson?) (Stephenson)

Cyprimeria coonensis (Stephenson) 2620 Cyprimeria alta (Conrad) 3560

Cyprimeria depressa (Conrad) 3950 Cyprimeria gabbi 860 (Stephenson)

Etea corsicana (Stephenson) 10 Etea peasei (Stephenson) 10

Exogyra cancellata (Stephenson) 3700 Exogyra costata costata 4250 (Say)

Flemingostrea battensis 310 Flemingostrea oleana 160 (Stephenson) (Stephenson)

Flemingostrea pratti (Stephenson) 820 Flemingostrea 2650 subspatulata (Forbes)

Glycymeris lacertosa (Wade) 10 Glycymeris rotundata 3500 [Glycymerita?] (Gabb) [Glycymerita?]

Granocardium alabamense (Gabb) 1300 Granocardium lowei 2220 (Stephenson)

Granocardium dumosum (Conrad) 4100 Granocardium tippanum 2410 (Conrad)

Isognomon williardi (Stephenson) 10 Isognomon holmesi 40 (Stephenson)

Legumen concentricum 760 Legumen ellipticum 3860 (Stephenson) (Conrad)

Leptosolen quadrilaterus 10 Leptosolen elongata 300 (Stephenson) (Weller)

Leptosolen? terminalis (Weller) 10 Leptosolen? levis 30 (Stephenson)

Linearia metastriata (Conrad) 3830 Linearia cribelli 3560 (Stephenson)

Linearia ornatissima (Weller) 50 Linearia n. sp. (Ripley) 10

Linter acutata (Stephenson) 2640 Linter burrana 10 (Stephenson)

Liothyris carolinensis (Conrad) 890 Liothyris n.sp. [L. 1950 carolinensis of Wade]

Miocardiopsis? cliffwoodensis 1610 Miocardiopsis? bulbosa 10 (Weller) (Stephenson)

Miocardiopsis? conradi (Gabb) of 10 Miocardiopsis? shumardi 10 Wade 1926 (Stephenson)

Neithea hartmani (Kniker) 2010 Neithea quinquecostata 1670 (Sowerby of Weller)

Nemodon eufalensis (Gabb) 4250 Nemodon grandis (Wade) 4000

Pachycardium stantoni (Wade) 10 Pachycardium wadei 30 (Stephenson)

Panopea decisa (Conrad) 2250 Panopea monmouthensis 2350 (Gardner)

Parmicorbula? percompressa 2650 Parmicorbula terramaria 3480 (Gardner) (Gardner)

Phelopteria petrosa (Conrad) 2300 Phelopteria linguaeformis 3500 (Evans & Shumard)

*Pleuriocardia marsense 10 *Pleuriocardia 50 (Stephenson) penderense (Stephenson)

*Pleuriocardia ochilleanum 80 *Pleuriocardia 1460 (Stephenson) carolinensis (Conrad)

Plicatula clarki (Stephenson) 3310 Plicatula tetrica (Conrad) 2490

Plicatula urticosa (Morton) 1870 Plicatula mullicaenensis 3960 (Weller)

Radiopecten mississippiensis 3250 Radiopecten weeksi 2300 (Conrad) (Stephenson)

Solyma lineolatus (Conrad) 250 Solyma gardnerae 3200 (Stephenson)

Spondylus guadalupae (Roemer) 120 Spondylus siccus (Elder 10 1996)

*Striarca nolani (Stephenson) 1240 *Striarca richardsi 1360 (Harbison)

Striarca umbonata (Conrad) 2100 Striarca poguei 220 (Stephenson)

Syncyclonema conradi (Whitfield) 300 Syncyclonema archeri 350 (Stephenson)

Tellinimera eborea (Conrad) 2300 Tellinimera buboana 4250 (Stephenson)

Tellinimera elliptica (Conrad) 160 Tellinimera stephensoni 260 (Salisbury)

Tellinimera? gabbi (Gardner) 4250 Tellinimera munda 390 (Stephenson)

Trigonarca inflata (Stephenson) 10 Trigonarca maconensis 900 (Conrad)

Veniella mullinensis (Stephenson) 700 Veniella lineata 100 (Shumard)

Anatimya anteradiata (Conrad) 3560 Anatimya lata (Whitfield) 2300

Brachymeris alta (Conrad) 160 Brachymeris carolinensis 860 (Conrad)

Crassatella hodgei (Stephenson) 4100 Crassatella vadosa 4250 (Morton)

Venericardia subteres (Stephenson) 10 Venericardia uvaldana 10 Stephenson

APPENDIX 3: Species used in PAST biostratigraphy analysis. Species were used based on their occurrence and presence in the formations used in the study. Data collected from the

Paleobiology Database (PBDB).

Species name Phylum Class

Glycymeris sp. Mollusca Bivalvia

Cucullaea (Idonearca) littlei Mollusca Bivalvia

Syncyclonema simplicius Mollusca Bivalvia

Lima (Lima) reticulata Mollusca Bivalvia

Anomia (Anomia) argentaria Mollusca Bivalvia

Exogyra costata Mollusca Bivalvia

Paladmete cancellaria Mollusca Gastropoda

Beretra gracilis Mollusca Gastropoda

Discoscaphites sp. Mollusca Cephalopoda

Inoceramus (Inoceramus) sp. Mollusca Bivalvia

Nemodon sp. Mollusca Bivalvia

Nemodon eufaulensis Mollusca Bivalvia

Cricosia filosa Brachiopoda Terebratulida

Turritella kerrensis Mollusca Gastropoda

Ornopsis sp. Mollusca Gastropoda

Discoscaphites conradi Mollusca Cephalopoda

Nucula percrassa Mollusca Bivalvia

Spondylus (Spondylus) sp. Mollusca Bivalvia

Legumen ellipticum Mollusca Bivalvia

Botula carolinensis Mollusca Bivalvia

Crenella elegantula Mollusca Bivalvia

Crassispira appressa Mollusca Gastropoda

Crassatella alta Mollusca Bivalvia

Flemingostrea subspatulata Mollusca Bivalvia

L. elongata

Linearia metastriata Mollusca Bivalvia

Nuculana kerrensis Mollusca Bivalvia

Pterotrigonia Mollusca Bivalvia (Scabrotrigonia) eufaulensis

Vetericardia crenalirata Mollusca Gastropoda

Thalassiphora patula Myzozoa (Dinoflagellate) Dinophyceae

Agerostrea falcata Mollusca Bivalvia

Anomia preolmstedi Mollusca Bivalvia

Aenona eufalensis Mollusca Bivalvia

Aphrodina tippana Mollusca Bivalvia

Beretra ripleyana Mollusca Gastropoda

Protocardium (Brevicardium) Mollusca Bivalvia parahillana

Camptonectes burlingtinensis Mollusca Bivalvia

Corbula carolinensis Mollusca Bivalvia

Lima (Costellaceta) riddlei Mollusca Bivalvia

Crenella serica Mollusca Bivalvia

Cucullaea capax Mollusca Bivalvia

Cyclothyris alta Mollusca Bivalvia

Capulus cuthandensis Mollusca Gastropoda

Capulus monroei Mollusca Gastropoda

Graciliala calcaris Mollusca Gastropoda

Drepanochilus quadriliratus Mollusca Gastropoda

Anchura substriata Mollusca Gastropoda

Anchura lamari Mollusca Gastropoda

Arrhoges (Latiala) lobata Mollusca Gastropoda

Weeksia lubbocki Mollusca Gastropoda

Plannolaterus (Calliophalus) Mollusca Gastropoda conanti

Pecten Mollusca Bivalvia (Camptonectes) microcancelli

Laxispira monilifera Mollusca Gastropoda

Turritella kellumi Mollusca Gastropoda

Turritella houstoni Mollusca Gastropoda

Turritella kerrensis Mollusca Gastropoda

Turritella vetebroides Mollusca Gastropoda

Haustator (Turritella) Mollusca Gastropoda paravertebroides

Haustator bilira Mollusca Gastropoda

Pecten simplicius Mollusca Bivalvia

Baculites sp. Mollusca Cephalopoda

Baculites vaalsensis Mollusca Cephalopoda Placenticeras placenta Mollusca Cephalopoda

Lima (Costellaceta) appressa Mollusca Bivalvia

APPENDIX 4: Results of the PAST unitary association analysis. Maximal cliques presented from the unitary associated output.

APPENDIX 5: Gastropod dataset arranged in time bin for Price equation analysis.

T1 T2 T3 T4 T5 T6 T7 T8 T9 G. N. G. Ripley/ N. D. Nacotoch Owl G. elevata/Dem hyat gansseri Coon alte conr (Navarro Cree elevata/Dem opolis ti foram Creek/ rna adi Group)/ k/Pra opolis Chalk/Merc 74- zone/Mon H. tum 70- N. irie Chalk/Merc hantville/ 72 mouth bilira (72- 68 rugosum/ Bluff/ hantville/ Eutaw Ma Fm(Nave zone72 69 Ma Corsican E. Eutaw &Black (Lat sink)/Sev +/-1 - Ma) (L a Marl costat &Black Creek e ern Fm 69 Ma Maa 72-70ish a 69- Creek (occuring Ca 73-71 Ma (E s) Ma 66 (occuring Middle mp) (Lcamp- Maas) (Texas E. Ma Middle Camp but E Maas) Maas) Camp but placed in placed in earlies time earlies time slice) 83-78 slice) 83-78 Ma (Late Ma (Late Camp) Camp) B. "" "" "" B. "" "" "" B. spinosus angulicostatus ang ulic osta tus D. communis D. "" "" "" "" "" "" D. Lem major D. "" D. exti dista distans nct ns

P. "" P. P. ex pergr pergra tenu ti acilis cilis ilira nc ta t

G. G. extinct mel m ano ul psis ti oc os tat a

P. "" "" "" "" "" "" R. modica (Orn opsis ) digre ssa R. "" "" R. H. triliratus (Ornop (Ornop sis) sis) elevata pulchra H. quadriliratus

E. extinct conv exus E. extinct meso zoicu s

R. stephensoni R. "" "" "" "" "" "" R. flexicostata steph enson i

P. "' "" P. ex creta bin ti ceum odo nc sum t P. "" P. extinct sublira cr tus as su s P. " "" "" P. extinct ejund su icus bd en sa tu s D. D. extinct rip be

ll be ll D. extinct co rb is , D. m ul ti

S. "" "" "" S. interrupta ripl eya na S. rugosa "" "" S. subnodosa H. "" """" H. nodo elegans sus

G. "" "" "" "" G. decemlirata calcari s D. "" D. " "" D. triliratus quad quad, D. trilirat us A. sub "" "" A. "" "" A. abrupta abrupt a A. lamari, A. bexarensis

L. lobata "" "" "" L. elegans P. P. P. goldmani. densatus goldm P.densatus ani, "" N. N. "" "" N. cooperens simpli cooper is??? cius ensis

(Pierre Shale)

T. "" "" "" "" "" mcna iryen sis T. T. kellumi hilgardi T. "' "" T. tippana, ve T. vert rt H. H. H. paravert parave how rtebroi elli des H. trilia, "" "" H. trilia, H. H. bilira bilira L. lum "" "" "" "" L. m on i Trobus "" T. "" "" "" T. buboanus corona cor ona T. kerrensis

P. tuber "" "" "" P. con ati P. P. mirco, P. "" P. argen P. argen micro, m teus P. ic argent ro eus

W. amp "" "" "" W. lubbocki

A. "" A. kerri n. sp .

G. coffea "" "" "" "" "" G. vorg? P. P. patens mon? P. "" P.rip pills

E. corona "" "" E. "" "" "" E. corona unione nsis G. n.sp, G. n. sp G. "" "" G. bella manz

A. (s.l) A. (s.l.) costata tallahatchiensis

M. mary "" "" "" "" "" M. transenna M. c.c "" "" M. cancellaria S. "" S. dep, S. depre imm ssa,

S. imme rsa L. imbri, L. tenn L. tenn V. splendida V. mutabilis V. valida, V. dum V. eu? V. producta

L. dis L. tur L. "" "" L. var pyri

P. "" "" "" "" P. elongatus spino sus P. "" P. lanhami (Kemp Form) perl ata P. "" "" "" "" "" "" P. prolixa peron atus

L. "" "" "" "" "" L. canalis leiodermu m L. thoracicum? "" "" "" "" "" L. tabulatum L. spiculatum "" "" "" "" "" "" "" L. cretaceum L. L. corona nodosu tum m P. "" "" "" P. saffordi calli

A. fascio "" "" "" A. wad ei? L. "" "" "" "" L. simp hubb ardi R. "" "" "" R. "" "" R. anom anom bi , R. a bia R. hastata, R. n sp. B. "" "" "" "" B. gracil firma is F. "" F. F. proxi kumm kummeli ma eli

A. "" A. pisti cictri E. ithy E. ellip E. "" "" "" E. "" E. E. lint per li lint nt N. "" "" "" "" N. defle tensa, xa N. deflexa T. per, T. sub G. "" G. bis G. bis, big G. elong C. carinata "" "" "" C. dem

A. H. "" H. amer, H. ameri cret cana A. P. gravida "" A.P. implexa A.P. A. P. wadei micro B. "" "" B. crider roc i, B. k Rock S. "" "" S. S. pondi bex be x S. S. "" S. sparsum harbisoni sparsu m, S. harbis oni O. n. sp? , O. fist

C. "" "" "" C. divers seca lina C. "" "" C. pessu in mata in

R. pulchella "" "" R. culb R. "" "" R. sufflata yoch O. "" "" "" "" "" "" O. n. sp. ameri cana

B. dem "" "" "" "" B. cret

C. cuth, C. span C. "" "" "" C. monr m oei ic ro

C. n. sp "" "" C. nodolir atum/r ata

A. A. costata tall

G. major "" G. sup G. sup. "" G. "" "" G. supra G. su major p

P. lepta P. "" P. lirata lirata

C. cret C. parvula

L. liratus "" L. liratus, L. cuc "" "" "" "" L. cuc L. cuc

C. mc C. "" "" "" C. matsoni mcna iryen sis

APPENDIX 6: Bivalve dataset arranged in time bin for Price equation analysis.

T1 T2 T3 T4 T5 T6 T7 T8 T9 G. N. G. Riple N. D. Nacotoch Owl G. elevata/Demop hy gansseri y/Coo alt co (Navarro Cree elevata/Demop olis atti foram n er nr Group)/ k/Pr olis Chalk/Merchan 74- zone/Mo Creek na adi N. airie Chalk/Merchan tville/ Eutaw 72 nmouth / H. tu 70- rugosum/ Bluf tville/ Eutaw &Black Creek Ma Fm(Nave bilira m 68 Corsican f/ E. &Black Creek (occuring (La sink)/Sev zone7 (7 M a Marl cost (occuring Middle Camp te ern Fm 2 +/-1 2- a 72-70ish ata Middle Camp but placed in Ca 73-71 Ma - 69 69 (L Ma 69- but placed in earlies time mp (Lcamp- Ma (E M M (Texas E. 66 earlies time slice) 83-78 Ma ) E Maas) Maas) a) aas Maas) Ma slice) 83-78 Ma (Late Camp) ) (Late Camp) C. grandis C. (w/in C. jer contusa sey foram ens zone is, Georgia) C. gra ndi s N. nacatochana N. N. nacatochana microstria ta (Marylan d) N. N. N. percrassa, per percr N. cra assa severnens ssa is N. prepercrassa, N. prepercrassa, N. stantoni N. stantoni N. N. N. cor col corbetens ben ora is ten do sis ens is

N. kerrensis N. N. kerrensis tarensi s S. eufalensis , S. angulicost ata V. V. crenalirat subcir a cula T. T. pat marcosen ula sis A. falcata A. A. A. falcata mesenteri me ca sen teri ca A. A. argentaria tellino ides A. preolmstedi, A. preolmstedi, A. olmstedi A. olmstedi A. A. eufaul n.s ensis, p. A.n.sp A. A. tippan tippa a, A. na, regia A. regia B. carolinensis B. B. carolinensis ripleyana, B. carolinens is B. B. parahi fra llana gil e B. tenue, B.

guadalup ense C. C. cra williardi ssa pli ca C. burlingtone nsis, C. virgatus/arg illensis C. carolinensis, C. carolinensis, C. oxynema C. oxynema C. sayrei C. riddl ei C. serica C. C. C. elegan ele elega tula ga ntula ntu la C. littlei C. C. littlei capa x C. parva, C. alta C. C. worde ap ni pre ssa , C. wo rde ni C. trigonalis C. C. trigonalis can cell osa C. coonensis C. C. C. coonensis coo alt nen a sis C. depressa, C. C. depressa, C. gabbi gabbi

E. peasei E. E. E. peasei cor corsicana sic ana E. costata E. costata cancel lata F. battensis, F. F. battensis, F. oleana oleana F. pratti F. F. pratti sub spa tul ata G. G. rotundata lacerto sa G. alaba mens e, G. lowe i G. dumosum G. G. dumosum tippa num I. williardi I. I. williardi holm esi L. concentricum, L. L. L. ellip ellipticum concentricum, L. ellip L. L. elo quadrilate ng rus ata L. terminalis L. levis L. terminalis L. L. cribelli me tast riat a L. ornatis sima,

L. n.sp. L. L. acutata L. L. acu acutat acutata, tata a L. burrana L. L. carolinens caroli is nensis, L.n.sp. M. M. bul cliff bos woo a densi s M. M. M. shu conrad shumardi ma i rdi N. hartmani, N. N. N. hartmani, N. quin quinq quin uecost ata N. N. eufalensis euf , N. ale grandis nsi s P. wadei P. stant oni P. decisa P. terramaria P. petro sa P.tetrica P. clark i P. P. mu urtic llic osa aen

ens is R. miss R. R. R. miss R. R. miss mis mi week siss ss si ipp ien sis S. S. gar lineolatus dne rae S. guadalupae, S. guadalupae, S. siccus S. siccus S. umbonata S. S. umbonata pogu ei S. conradi S. S. archeri S. conradi arc her i T. T. T. buboana eborea bu , T. bo buboa ana na T. elliptica, T. T. T. elliptica, T. stephensoni ste stephensoni ph T.munda T. gabbi T. T.munda ga bbi T. inflata, T. T. inflata, T. maconensis maconensis V. mullinensis V. V. mullinensis lineata A. A. lata ant era diat a B. alta, B. B. B. alta, B. carolinenesis car carolinenesis oli nen

esis ? C. hodgei C. C. C. hodgei ho vado dg sa ei? V. uvaladana V. V. uvaladana subter es

APPENDIX 7: Gastropod Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants).

Time Increment 1 Ances An #ances #des rel Des. Range Non-imm Anag. Species Range fitness size desc only Chng

B. 700 1 1 1.0434 700 700 0 angulicostat 78 us n/a 1230 n/a 1930 n/a 750 n/a 10 n/a 860 n/a 10 n/a 10 R. 700 1 1 1.0434 700 700 0 stephensoni 78 n/a 10 n/a 10 S. rugosa 10 1 1 1.0434 10 10 0 78 n/a 10 n/a 10 A. sub 1990 1 1 1.0434 1990 1990 0 78 n/a 3110 L. lum 3360 1 1 1.0434 3360 3360 0 78 P. tuber 10 1 1 1.0434 10 10 0 78 n/a 45 G. coffea 10 1 1 1.0434 10 10 0 78 n/a 2250 E. corona 2230 1 1 1.0434 2230 2230 0 78 M. c.c 1320 1 1 1.0434 1320 1320 0 78 n/a 10

85 n/a 690 L. imbri 10 1 0 0 extinct L. tenn 10 1 1 1.0434 10 10 0 78 V. splendida 10 1 1 1.0434 10 10 0 78 L. dis 10 1 1 1.0434 80 80 70 78 n/a 2300 n/a 10 n/a 10 L. 860 1 1 1.0434 860 860 0 thoracicum? 78 L. 1960 1 1 1.0434 1960 1960 0 spiculatum 78 A. fascio 10 1 1 1.0434 10 10 0 78 n/a 60 n/a 1630 n/a 1230 n/a 690 n/a 10 n/a 690 E. ithy 10 1 1 1.0434 1240 1240 1230 78 n/a 950 n/a 30 n/a 10 10 n/a 690 C. carinata 10 1 1 1.0434 10 10 0 78 n/a 10 A. P. 10 1 1 1.0434 10 10 0 gravida 78 n/a 3280 n/a 1230 n/a 1230 n/a 10 R. pulchella 2170 1 1 1.0434 2170 2170 0 78 S. rugosa 1280 n/a 10

C. n. sp 20 1 1 1.0434 20 20 0 78 G. major 2610 1 1 1.0434 2610 2610 0 78 P. lepta 2120 1 1 1.0434 2050 2050 -70 78 L. liratus 690 1 1 1.0434 690 690 0 78 n/a 10

24 23 1 959.1304348 Sum Sum Average rel fit Average desc w/out Ances Desc immi

anc.mean: 868.33 3 desc.mean: 793.19 7 difference: - SS AC Immi 75.136 6 37.318 53.478260 -165.9337135 84 87

- 75.136 6 TS1 Price Equation Answer

Time Increment 2 Ances Species An #anc #de rel Des. Range Non-imm desc Anag. Range es s fitness size only Chng b. ang 700 1 1 1.1296 700 700 0 3 d. comm 1230 1 0 0 extinct d. distants 1930 1 1 1.1296 1930 1930 0 3

87 d. major 750 1 1 1.1296 750 750 0 3 p. per 10 1 1 1.1296 10 10 0 3 P. (orn) dig 860 1 1 1.1296 860 860 0 3 E. con 10 1 0 0 extinct E. mes 10 1 0 0 extinct R. step 700 1 1 1.1296 700 700 0 3 p. cret 10 1 1 1.1296 10 10 0 3 p. ejun 10 1 1 1.1296 10 10 0 3 S. rug 10 1 1 1.1296 10 10 0 3 h. nod 10 1 1 1.1296 10 10 0 3 d. quad 10 1 1 1.1296 10 10 0 3 a. sub 1990 1 1 1.1296 1990 1990 0 3 n/a 3520 n/a 3530 n/a 10 t. mcnair 3110 1 1 1.1296 3110 3110 0 3 n/a 2250 n/a 3480 n/a 4020 n/a 4250 n/a 3950 L.lum 3360 1 1 1.1296 3360 3360 0 3 n/a 640 P. tuber 10 1 1 1.1296 10 10 0 3 p.argen 45 1 1 2.2592 45 45 0 59 1 490 490 445 n/a 690 G. coffea 10 1 1 1.1296 10 10 0 3

P. pills 2250 1 1 1.1296 2250 2250 0 3 E. corona 2230 1 1 1.1296 2230 2230 0 3 n/a 3720 m.c.c 1320 1 1 1.1296 1320 1320 0 3 s. imm 10 1 1 1.1296 10 10 0 3 s. dep 690 1 1 1.1296 690 690 0 3 l. tenn 10 1 0 0 v. mut 10 1 0 0 l. tur 80 1 0 0 l. var 2300 1 1 1.1296 2300 2300 0 3 p. spin 10 1 1 1.1296 10 10 0 3 p. pero 10 1 1 1.1296 10 10 0 3 n/a 1900 L. thoar 860 1 1 1.1296 860 860 0 3 L.spic 1960 1 1 1.1296 1960 1960 0 3 a. fas 10 1 1 1.1296 10 10 0 3 l. hubb 60 1 1 1.1296 60 60 0 3 r. anom 1630 1 1 1.1296 1630 1630 0 3 rbia 1230 1 1 1.1296 1230 1230 0 3 b. grac 690 1 1 1.1296 690 690 0 3 f. prox 10 1 1 1.1296 10 10 0 3 a. pis 690 1 1 1.1296 690 690 0 3 e. ellip 1240 1 0 0 e. per 950 1 1 1.1296 950 950 0 3 n. def 30 1 1 1.1296 30 30 0 3

89 t. per 10 1 0 0 t. sub 10 1 0 0 g. big 690 1 1 1.1296 690 690 0 3 c. car 10 1 1 1.1296 10 10 0 3 a.h.amer 10 1 1 1.1296 10 10 0 3 a. p. grav 10 1 1 1.1296 30 30 20 3 n/a 10 b. cri 3280 1 1 1.1296 3280 3280 0 3 b. rock 1230 1 1 1.1296 1230 1230 0 3 s. pon 1230 1 1 1.1296 1230 1230 0 3 n/a 10 C. pessumata 10 1 1 1.1296 10 10 0 3 R. pulchella 2170 1 1 1.1296 2170 2170 0 3 O. americana 1280 1 1 1.1296 1280 1280 0 3 C. monroei 10 1 1 1.1296 10 10 0 3 C. n. sp 20 1 1 1.1296 20 20 0 3 G. major 2610 1 1 1.1296 3730 3730 1120 3 P. lirata 2050 1 1 1.1296 2050 2050 0 3 L. liratus 690 1 1 2.2592 690 690 0 59 1 650 650 -40 C. 10 1 1 1.1296 10 10 0 mcnairyensis 3

61 66 54 w/out immigrants

Ances mean 793.19 67

Desc mean 1176.9 85 Difference 383.78 86

SS AC Im mi 68.099 28.6 287 57 1 total 383.78 86

Time Increment 3 Ances Species An #ances #des rel Des. Non-imm Anag. Range fitnes Range desc only Chng s size

B. ang 700 1 1 0.957 700 700 0 746 D. major 1930 1 1 0.957 1930 1930 0 746 D. distans 750 1 1 0.957 750 750 0 746 P. pergracilis 10 1 1 0.957 10 10 0 746 P. (Orn) digressa 860 1 1 0.957 860 860 0 746 n/a 80 R. step 700 1 1 0.957 700 700 0 746 P. cret 10 1 1 0.957 10 10 0 746 n/a 10 P. ejun 10 1 1 0.957 10 10 0 746 S. rugosa 10 1 1 0.957 10 10 0 746 H. nodo 10 1 1 0.957 10 10 0 746 n/a 1860 D. quad 10 1 2 1.915 10 10 0 493

1320 1320 1310 A. sub 1990 1 1 0.957 1540 1540 -450 746 L. lobata 3520 1 1 0.957 3520 3520 0 746 P. densatus 3530 1 2 1.915 3530 3530 0 493 3950 3950 420 N. cooperensis??? 10 1 1 0.957 10 10 0 (Pierre Shale) 746 T. mcnair 3110 1 1 0.957 3110 3110 0 746 T. hilgardi 2250 1 1 0.957 10 10 -2240 746 T. tippana 3480 1 1 0.957 3480 3480 0 746 (t. vert) 4020 1 1 0.957 4020 4020 0 746 n/a 3480 H. trilia 4250 1 1 0.957 4250 4250 0 746 (h. bil) 3950 1 1 0.957 3950 3950 0 746 L. lum 3360 1 1 0.957 3360 3360 0 746 T. corona 640 1 1 0.957 640 640 0 746 n/a 10 n/a 30 P. tuber 10 1 1 0.957 10 10 0 746 P. mirco 45 1 1 0.957 45 45 0 746 (P. argen) 490 1 1 0.957 490 490 0 746 W. amp 690 1 1 0.957 690 690 0 746 n/a 160 G. coffea 10 1 1 0.957 10 10 0 746 P. pills 2250 1 1 0.957 690 690 -1560 746 E. corona 2230 1 1 0.957 2230 2230 0 746

92 n/a 640 n/a 1990 n/a 10 M. mary 3720 1 1 0.957 3720 3720 0 746 M. c.c 1320 1 1 0.957 1920 1920 600 746 S. dep 10 1 1 0.957 10 10 0 746 (S. imm) 690 1 1 0.957 690 690 0 746 n/a 10 L .var 2300 1 1 0.957 2300 2300 0 746 P. spin 10 1 1 0.957 10 10 0 746 P. per 10 1 1 0.957 10 10 0 746 L. leiodermum 1900 1 1 0.957 1900 1900 0 746 L. thor 860 1 1 0.957 860 860 0 746 L. spic 1960 1 1 0.957 1960 1960 0 746 n/a 10 A. fascio 10 1 1 0.957 10 10 0 746 L. hubbardi 60 1 1 0.957 60 60 0 746 R. anom 1630 1 1 0.957 1630 1630 0 746 (r. bia) 1230 1 1 0.957 1230 1230 0 746 B. gracilis 690 1 1 0.957 690 690 0 746 F. proxima 10 1 1 0.957 640 640 630 746 A. pisti 690 1 1 0.957 30 30 -660 746 E. per 950 1 1 0.957 950 950 0 746 N. defelxa 30 1 1 0.957 30 30 0 746

G. bis 690 1 1 0.957 690 690 0 746 C. carinata 10 1 1 0.957 10 10 0 746 A. H. americana 10 1 2 1.915 10 10 0 493 10 10 0 A.P. implexa 30 1 0 0 extinct A.P. micro 10 1 1 0.957 80 80 70 746 B. crideri 3280 1 1 0.957 3280 3280 0 746 (b. rock) 1230 1 1 0.957 1230 1230 0 746 S. pondi 1230 1 1 0.957 1230 1230 0 746 S. harbisoni 10 1 2 1.915 10 10 0 493 1820 1820 1810 n/a 10 n/a 10 C. pessumata 10 1 1 0.957 10 10 0 746 R. pulchella 2170 1 1 0.957 2180 2180 10 746 n/a 30 O. americana 1280 1 1 0.957 1280 1280 0 746 n/a 10 C. monroei 10 1 1 0.957 10 10 0 746 C. n. sp 20 1 1 0.957 50 50 30 746 n/a 10 G. sup 3730 1 2 1.915 3730 3730 0 493 2610 2610 -1120 P. lirata 2050 1 1 0.957 2050 2050 0 746 n/a 10 L. liratus 690 1 0 0 extinct (l. cuc) 650 1 1 0.957 650 650 0 746

C. mcnair 10 1 1 0.957 10 10 0 746

68 71 1 Ances Des Rel fitness total total average Ances mean 1176. Non imm desc averg 985 Desc mean 1054. 1203.59154 213 9 Total mean change - S.S. A.C. Immi 122.7 72 42.8 - - 16.19 149.3780 72 661

-123

Time Increment 4 Ances Species An #ances #des rel Des. Range Non-imm Anag. Range fitness size desc only Chng

B.ang 700 1 1 1.2571 700 700 0 429 D. major 750 1 1 1.2571 750 750 0 429 D. distans 1930 1 0 0 extinct P. pergracilis 10 1 1 1.2571 10 10 0 429 n/a 650 R. (orn) 860 1 1 1.2571 860 860 0 digressa 429 R. (Ornopsis) 80 1 1 1.2571 80 80 0 elevata 429 R. step 700 1 1 1.2571 700 700 0 429 P. cret 10 1 1 1.2571 10 10 0 429 P. subliratus 10 1 1 1.2571 10 10 0 429

P. ejun 10 1 1 1.2571 10 10 0 429 n/a 700 S. subnodosa 10 1 0 0 extinct H. nodo 10 1 1 1.2571 30 30 20 429 G. calcaris 1860 1 1 1.2571 1860 1860 0 429 D. quad 10 1 1 1.2571 10 10 0 429 D. trili. 1320 1 1 1.2571 1320 1320 0 429 A. abrupta 1540 1 1 1.2571 1540 1540 0 429 L. lobata 3520 1 1 1.2571 3520 3520 0 429 P. goldmani 3530 1 1 1.2571 3530 3530 0 429 P. den 3950 1 1 1.2571 3950 3950 0 429 N. simplicius 10 1 1 1.2571 10 10 0 429 T. mcnair 3110 1 1 1.2571 3110 3110 0 429 T. kellumi 10 1 0 0 extinct T. tipp 3489 1 1 1.2571 3480 3480 -9 429 T. vert 4020 1 1 1.2571 4020 4020 0 429 H. 3480 1 1 1.2571 640 640 -2840 paravertebroi 429 des H. trilia 4250 1 1 1.2571 4250 4250 0 429 H. bil 3950 1 1 1.2571 3950 3950 0 429 L. lum 3360 1 1 1.2571 3360 3360 0 429 Trobus corona 640 1 1 1.2571 640 640 0 429 T. kerrensis 30 1 0 0 extinct T. hill 10 1 0 0 extinct P. tuber 10 1 1 1.2571 30 30 20 429

P. micro,P. 45 1 1 1.2571 45 45 0 argenteus 429 P. argen 490 1 1 1.2571 490 490 0 429 W. amp 690 1 1 1.2571 690 690 0 429 A. kerri 160 1 1 1.2571 160 160 0 429 G. coffea 10 1 1 1.2571 10 10 0 429 P.rip 690 1 0 0 extinct E. unionensis 640 1 1 1.2571 640 640 0 429 G. bella 1990 1 1 1.2571 1990 1990 0 429 A. (s.l) costata 10 1 1 1.2571 2170 2170 2160 429 M. mary 3720 1 1 1.2571 3720 3720 0 429 M. cancellaria 1920 1 0 0 extinct S. dep 10 1 0 0 extinct S. imm 690 1 0 0 extinct n/a 640 n/a 30 V. eu? 10 1 1 1.2571 30 30 20 429 L .var 2300 1 1 1.2571 1930 1930 -370 429 P. spin 10 1 1 1.2571 10 10 0 429 n/a 650 P. per 10 1 1 1.2571 10 10 0 429 L. leiodermum 1900 1 1 1.2571 1900 1900 0 429 L. thor 860 1 1 1.2571 860 860 0 429 L. spic 1960 1 1 1.2571 1960 1960 0 429 L. coronatum 10 1 1 1.2571 610 610 600 429 n/a 690 A. fascio 10 1 1 1.2571 640 640 630 429

L. hubbardi 60 1 1 1.2571 60 60 0 429 R. anom 1630 1 1 1.2571 1630 1630 0 429 R. bia 1230 1 1 1.2571 1230 1230 0 429 B. gracilis 690 1 1 1.2571 690 690 0 429 F. kummeli 640 1 1 1.2571 640 640 0 429 A. cictri 30 1 0 0 extinct E. per 950 1 1 1.2571 950 950 0 429 N. defelxa 30 1 1 1.2571 30 30 0 429 G. bis 690 1 2 2.5142 690 690 0 857 640 640 -50 C. carinata 10 1 1 1.2571 80 80 70 429 H. amer. 10 1 0 0 extinct H. cret 10 1 0 0 extinct A. P. wadei 80 1 0 0 extinct B. crideri 3280 1 0 0 extinct B. rock 1230 1 1 1.2571 1230 1230 0 429 S. pondi 1230 1 1 1.2571 1800 1800 570 429 S. sparsum 1820 1 1 1.2571 1820 1820 0 429 S. har 10 1 1 1.2571 10 10 0 429 O. n. sp? 10 1 0 0 extinct O. fist 10 1 0 0 extinct n/a 3280 C. pessumata 10 1 1 1.2571 80 80 70 429 R. culb 2180 1 0 0 extinct R. yoch 30 1 1 1.2571 30 30 0 429 O. americana 1280 1 1 1.2571 1280 1280 0 429 B. dem 10 1 1 1.2571 10 10 0 429

C. monroei 10 1 1 1.2571 10 10 0 429 C. 50 1 0 0 extinct nodoliratum A. costata 10 1 1 1.2571 2170 2170 2160 429 G. sup. 3730 1 1 1.2571 3730 3730 0 429 G. major 2610 1 1 1.2571 2610 2610 0 429 P. lirata 2050 1 0 0 extinct C. cret 10 1 1 1.2571 640 640 630 429 L. cuc 650 1 1 1.2571 650 650 0 429 C. mc 10 1 1 1.2571 10 10 0 429

88 70 1 1185.214286 1040.954545 ances des average rel fit Dec w/out immigrants total total 1163.701299 122.7467532 S.S. AC Immi- 91.67 52.585 21.51298701 4 714

122.7 5 TS4 PE answer

Time Increment 5 Ances Species An #ances #des rel Des. Range Non-imm Anag. Range fitness size desc only Chng

B. ang 700 1 1 1.5098 700 700 0 039 D. major 750 1 1 1.5098 750 750 0 039 P. ten 10 1 0 0 extinct G. mel 650 1 1 1.5098 1770 1770 1120 039

R. (orn) 860 1 1 1.5098 860 860 0 digressa 039 R. (Ornopsis) 80 1 1 1.5098 80 80 0 elevata 039 R. step 700 1 1 1.5098 700 700 0 039 P. bind 80 1 0 0 extinct P. sub 10 1 1 1.5098 650 650 640 039 P. ejun 10 1 1 1.5098 10 10 0 039 n/a 2670 n/a 20 n/a 20 S. rip 700 1 1 1.5098 700 700 0 039 H. elegans 30 1 0 0 extinct G. calcaris 1860 1 1 1.5098 1860 1860 0 039 D. quad 10 1 1 1.5098 10 10 0 039 D. tril 1320 1 1 1.5098 1320 1320 0 039 A. abrupta 1540 1 1 1.5098 1540 1540 0 039 L. lobata 3520 1 1 1.5098 3520 3520 0 039 P. goldmani 3530 1 0 0 extinct P. den 3950 1 0 0 extinct N. simplicius 10 1 1 1.5098 10 10 0 039 T. mcnair 3110 1 1 1.5098 3110 3110 0 039 T. tipp 3480 1 0 0 extinct T. vert 4020 1 1 1.5098 4020 4020 0 039 H. howelli 640 1 1 1.5098 3480 3480 2840 039 H. trilia 4250 1 1 1.5098 4250 4250 0 039 H. bil 3950 1 1 1.5098 3950 3950 0 039 L. lum 3360 1 1 1.5098 3820 3820 460 039

100

T. corona 640 1 1 1.5098 640 640 0 039 P. conati 30 1 0 0 extinct P. micro 45 1 0 0 extinct P. argen 490 1 1 1.5098 490 490 0 039 W. amp 690 1 1 1.5098 690 690 0 039 A. kerri 160 1 1 1.5098 650 650 490 039 G. coffea 10 1 1 1.5098 10 10 0 039 E. union 640 1 1 1.5098 640 640 0 039 n/a 10 n/a 10 G. bella 1990 1 1 1.5098 1990 1990 0 039 A. (s.l) tall 2170 1 0 0 extinct M. mary 3720 1 1 1.5098 3720 3720 0 039 V. dum. 640 1 0 0 extinct V. valid 30 1 0 0 extinct V. prod 30 1 0 0 extinct L. pyri 1930 1 0 0 extinct P. spin 10 1 1 1.5098 10 10 0 039 P. perl 650 1 1 1.5098 650 650 0 039 P. per 10 1 1 1.5098 10 10 0 039 L. 1900 1 1 1.5098 1900 1900 0 leiodermum 039 L. thor 860 1 1 1.5098 860 860 0 039 L. spic 1960 1 1 1.5098 1960 1960 0 039 L. nodo 610 1 0 0 extinct P. calli 690 1 1 1.5098 690 690 0 039 A. wadei 640 1 0 0 extinct L. hubbardi 60 1 1 1.5098 60 60 0 039

101

R. anom, R. 1630 1 0 0 extinct bia R. bia 1230 1 1 1.5098 1230 1230 0 039 n/a 10 n/a 10 B. gracilis 690 1 1 1.5098 690 690 0 039 F. kummeli 640 1 0 0 extinct E. per 950 1 1 1.5098 2050 2050 1100 039 N. deflexa 30 1 1 1.5098 30 30 0 039 G. bis. 690 1 0 0 extinct G. elong 640 1 0 0 extinct C. dem 80 1 0 0 extinct B. rock 1230 1 0 0 extinct S. bex 1800 1 1 1.5098 1800 1800 0 039 S. sparsum 1820 1 1 1.5098 1820 1820 0 039 S. har. 10 1 0 0 extinct C. sel 3280 1 1 1.5098 3280 3280 0 039 C. inn inn 80 1 0 0 extinct R. yoch 30 1 1 1.5098 30 30 0 039 O. americana 1280 1 1 1.5098 1280 1280 0 039 B. dem 10 1 1 1.5098 10 10 0 039 n/a 1250 n/a 1540 C. mon 10 1 1 1.5098 10 10 0 039 A. tall 2170 1 0 0 extinct G. sup 3730 1 1 1.5098 3730 3730 0 039 G. major 2610 1 0 0 extinct C. parvula 640 1 0 0 extinct L. cuc 650 1 1 1.5098 650 650 0 039 C. mc 10 1 1 1.5098 10 10 0 039

102

77 51 1 1347.058824 Sum Sum Average rel fit Average Desc w/out Ances Desc imm Ances Mean 1164.6 1 Desc Mean 1237.3 33 Total Mean 72.722 SS AC Immi- Change 94 52.06 130.32 109.7254902 16

72.72 TS5 PE answer

Time Increment 6 Ances An #ance #des rel Des. Non-imm Anag. Chng Species Rang s fitnes Range desc only e s size

B. ang 700 1 1 1.578 700 700 0 9474 D. major 750 1 1 1.578 750 750 0 9474 G. multi 1770 1 0 0 extinct R. 860 1 1 1.578 860 860 0 digressa 9474 R. elevata 80 1 1 1.578 30 30 -50 9474 R. step 700 1 1 1.578 700 700 0 9474 P. crass 650 1 0 0 extinct P. subd 10 1 0 0 extinct D. bell bell 2670 1 0 0 extinct D. corb 20 1 0 0 extinct D. multi 20 1 0 0 extinct S. rip 700 1 1 1.578 700 700 0 9474 G. calcaris 1860 1 1 1.578 1860 1860 0 9474 D. quad 10 1 0 0 extinct

103

D. tril 1320 1 1 1.578 1320 1320 0 9474 A. abrupta 1540 1 1 1.578 1540 1540 0 9474 n/a 300 n/a 10 L. lobata 3520 1 1 1.578 90 90 -3430 9474 N. 10 1 1 1.578 10 10 0 simplicius 9474 T. mcnair 3110 1 1 1.578 30 30 -3080 9474 T. vert 4020 1 0 0 extinct H. para 3480 1 0 0 extinct H. trilia 4250 1 0 0 extinct H. bil 3950 1 0 0 extinct L. mon 3820 1 0 0 extinct T. corona 640 1 1 1.578 640 640 0 9474 P. micro 490 1 0 0 extinct W. amp 690 1 1 1.578 60 60 -630 9474 A. n. sp 650 1 0 0 extinct G. coffea 10 1 1 1.578 10 10 0 9474 n/a 10 E. union 640 1 1 1.578 640 640 0 9474 G. n. sp 10 1 0 0 extinct G. n. sp x2 10 1 0 0 extinct G. bella 1990 1 1 1.578 60 60 -1930 9474 M. mary 3720 1 1 1.578 3720 3720 0 9474 P. spin 10 1 1 1.578 10 10 0 9474 P. perl 650 1 1 1.578 10 10 -640 9474 P. per 10 1 1 1.578 10 10 0 9474 L. 1900 1 1 1.578 1900 1900 0 leiodermu 9474 m

104

L. thor 860 1 1 1.578 470 470 -390 9474 L. spic 1960 1 1 1.578 1960 1960 0 9474 P. calli 690 1 1 1.578 690 690 0 9474 L. hubb 60 1 1 1.578 10 10 -50 9474 R. bia 1230 1 1 1.578 1230 1230 0 9474 R. has 10 1 0 0 extinct R. n. sp 10 1 0 0 extinct B. grac 690 1 1 1.578 40 40 -650 9474 E. lint 2050 1 1 1.578 2050 2050 0 9474 N. def 30 1 2 3.157 30 30 0 8947 10 10 -20 S. bex 1800 1 0 0 extinct S. spar 1820 1 0 0 extinct C. sel 3280 1 1 1.578 3280 3280 0 9474 R. yoch 30 1 1 1.578 10 10 -20 9474 O. amer 1280 1 1 1.578 1280 1280 0 9474 B. dem 10 1 1 1.578 10 10 0 9474 C. spa 1540 1 0 0 extinct C. cuth 1250 1 0 0 extinct C. micro 10 1 0 0 extinct G. sup 3730 1 1 1.578 3730 3730 0 9474 L. cuc 650 1 1 1.578 650 650 0 9474 C. mcnair 10 1 1 1.578 30 30 20 9474

60 38 1 819.210526 3 Sum Sum Average rel fit Average desc w/out ances desc immi

105

1237.3333 33 767.07317 S.S- AC- Immi- 07 132.070175 286.052 52.13735 4 6 6 - 470.26016 26 - 470.260162 6 TS6 PE answer

Time Increment 7 Ances An #ances #des rel Des. Non-imm Anag. Species Range fitness Range desc only Chng size

B. ang 700 1 1 1.8636 700 700 0 364 D. major 750 1 1 1.8636 750 750 0 364 R. digressa 860 1 1 1.8636 860 860 0 364 R. pulchra 30 1 0 0 extinct n/a 650 n/a 500 R. step 700 1 1 1.8636 700 700 0 364 S. rip 700 1 1 1.8636 700 700 0 364 G. cal 1860 1 1 1.8636 1860 1860 0 364 D. tril 1320 1 0 0 extinct A. abrupta 1540 1 0 0 extinct A. lam. 300 1 0 0 extinct A. bex 10 1 0 0 extinct L. elegans 90 1 0 0 extinct N. coop 10 1 0 0 extinct

106

T. hous 30 1 0 0 extinct T. corona 640 1 1 1.8636 640 640 0 364 W. amp 60 1 0 0 extinct G. vorg 10 1 0 0 extinct P. patens 10 1 1 1.8636 10 10 0 364 E. union 640 1 1 1.8636 2230 2230 1590 364 G. manz 60 1 0 0 extinct M mary 3720 1 1 1.8636 3720 3720 0 364 P.elong 10 1 0 0 extinct P. lan 10 1 0 0 extinct P. per 10 1 1 1.8636 10 10 0 364 L. 1900 1 1 1.8636 1900 1900 0 leiodermu 364 m L. tab 470 1 0 0 extinct L. spic 1960 1 1 1.8636 1960 1960 0 364 P. calli 690 1 1 1.8636 690 690 0 364 L. simp 10 1 0 0 extinct R. bia 1230 1 1 1.8636 1230 1230 0 364 B. firma 40 1 1 1.8636 40 40 0 364 E. lint 2050 1 1 1.8636 2050 2050 0 364 N. def. 30 1 0 0 extinct N. ten 10 1 0 0 extinct C. sel 3280 1 1 1.8636 3280 3280 0 364 R. suff 10 1 0 0 extinct O. amer 1280 1 1 1.8636 1280 1280 0 364 B. dem 10 1 1 1.8636 10 10 0 364 G. sup 3730 1 1 1.8636 3730 3730 0 364 L. cuc 650 1 1 1.8636 650 650 0 364

107

C. matsoni 30 1 0 0 extinct

41 22 1 1318.181818 Sum Sum Desc Average rel fit Average desc w/out Ances of ances immi

S.S. A.C Immi Ances mean 767.0 478.83 72.272727 - 732 592 27 61.93181818 Desc mean 1256. 25 Total mean 489.1 489.17 768 683 TS7 PE answer

Time Increment 8 Ances An #ances #des rel Des. Range Non-imm Anag. Species Range fitness size desc only Chng

B. ang 700 1 1 1.2631 1390 1390 690 579 D. major 750 1 1 1.2631 640 640 -110 579 P. digressa 860 1 1 1.2631 300 300 -560 579 H. tril. 650 1 0 0 extinct H. quad 500 1 0 0 extinct R. step 700 1 1 1.2631 1920 1920 1220 579 S. rip 700 1 1 1.2631 2640 2640 1940 579 G. cal 1860 1 1 1.2631 720 720 -1140 579 T corona 640 1 1 1.2631 310 310 -330 579 P. mon 10 1 0 0 extinct E. corona 2230 1 0 0 extinct M. mary 3720 1 1 1.2631 1920 1920 -1800 579

108

P. per 10 1 1 1.2631 10 10 0 579 L. leio 1900 1 1 1.2631 2030 2030 130 579 L. spic 1960 1 1 1.2631 3950 3950 1990 579 P. calli 690 1 1 1.2631 80 80 -610 579 R. bia 1230 1 1 1.2631 1630 1630 400 579 B. firma 40 1 0 0 extinct E. lint 2050 1 1 1.2631 2050 2050 0 579 C. sel 3280 1 1 1.2631 1940 1940 -1340 579 O. amer 1280 1 1 1.2631 1350 1350 70 579 B. dem 10 1 1 1.2631 340 340 330 579 G. sup 3730 1 1 1.2631 3730 3730 0 579 L. cuc 650 1 1 1.2631 650 650 0 579

24 19 1 1452.631579 Sum Sum Average Rel fit Average desc w/out Ances Desc immi Ances mean 1256.2 5 Desc mean 1452.6 No 32 immigrants Total mean 196.38 SS AC 0 16 150.1 46.315 789

196.4 TS8 PE Answer

109

APPENDIX 8: Bivalve Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants).

Time Increment 1

Ances Species An #ances #d rel Des. Non-imm Anag. Range es fitnes Range Desc Chng s size

N. nacatochana 1930 1 1 1.6 1930 1930 0 N. prepercrassa, 4250 1 0 0 extinct (N. stantoni) N. stantoni 150 1 0 0 extinct n/a 10 N. kerrensis 160 1 1 1.6 160 160 0 n/a 30 A. falcata 4020 1 1 1.6 4020 4020 0 A. preolmstedi, (A. 50 1 0 0 extinct olmstedi) A. olmstedi 890 1 0 0 extinct B. carolinensis 3150 1 1 1.6 3150 3150 0 C. carolinensis, (C. 160 1 0 0 extinct oxynema) C. oxynema 860 1 0 0 extinct C. trigonalis 3530 1 1 1.6 2790 2790 -740 C. coonensis 2620 1 1 1.6 2620 2620 0 C. depressa, (C. 3950 1 0 0 extinct gabbi) C. gabbi 860 1 0 0 extinct E. peasei 10 1 1 1.6 10 10 0 F. battensis, (F. 310 1 0 0 extinct oleana) F. oleana 160 1 0 0 extinct F. pratti 820 1 1 1.6 820 820 0 G. dumosum 4100 1 1 1.6 4100 4100 0 I. williardi 10 1 1 1.6 10 10 0 L. concentricum, 760 1 1 1.6 760 760 0 (L. ellip) L. ellip 3860 1 1 1.6 3860 3860 0 L. terminalis 10 1 1 1.6 10 10 0

110 n/a 2640 n/a 10 n/a 10 N. hartmani, (N. 2010 1 1 1.6 2010 2010 0 quin) N. quin 1670 1 1 1.6 1670 1670 0 R. miss 3520 1 1 1.6 3520 3520 0 n/a 3200 S. guadalupae, (S. 120 1 0 0 extinct siccus) S. siccus 10 1 0 0 extinct S. umbonata 2100 1 1 1.6 2100 2100 0 S. conradi 300 1 1 1.6 350 350 50 T. elliptica, (T. 160 1 1 1.6 160 160 0 stephensoni) T. steph 260 1 1 1.6 260 260 0 T.munda 390 1 1 1.6 390 390 0 T. inflata, (T. 10 1 0 0 extinct maconensis) T. maco. 900 1 0 0 extinct V. mullinensis 700 1 1 1.6 700 700 0 n/a 3560 B. alta, (B. 160 1 0 0 extinct carolinenesis) B. carolinensis 860 1 1 1.6 860 860 0 C. hodgei 4100 1 1 1.6 4100 4100 0 V. uvaladana 10 1 1 1.6 10 10 0 Total Total Desc (no Ancestors immigrants) TS1 40 25 ances mean 1347.5 desc mean 1557.1 875 difference 209.68 75

SS A Immi C 294.9 - - 27. 57.61 6 25

111

PE ANSWER TS1 209.688

Time Increment 2

Ances Species An #ances #des rel Des. Non-imm Anag. Range fitness Range Desc Chng size n/a 2250 N. nacatochana 1930 1 1 1.1071 3230 3230 1300 4286 n/a 4250 n/a 1310 N. corbentensis 10 1 1 1.1071 10 10 0 4286 N. kerrensis 160 1 1 1.1071 160 160 0 4286 n/a 3830 n/a 2650 n/a 2300 T. patula 30 1 1 1.1071 30 30 0 4286 A. falcata 4020 1 1 1.1071 4100 4100 80 4286 n/a 4250 B. carolinensis 3150 1 2 2.2142 3150 3150 0 8571 2300 2300 -850 n/a 4100 n/a 2330 n/a 3950 n/a 850 C. cancellosa 2790 1 0 0 extinct C. coonensis 2620 1 1 1.1071 2620 2620 0 4286 E. corsicana 10 1 0 0 extinct n/a 4250 F. pratti 820 1 1 1.1071 820 820 0 4286

112 n/a 3500 G. dumosum 4100 1 1 1.1071 4100 4100 0 4286 I. williardi 10 1 1 1.1071 10 10 0 4286 L. concentricum, 760 1 0 0 extinct (L. ellip) L. ellip 3860 1 1 1.1071 3860 3860 0 4286 L. acutata 2640 1 1 1.1071 2640 2640 0 4286 M. bulbosa 10 1 1 1.1071 10 10 0 4286 M. shumardi 10 1 1 1.1071 10 10 0 4286 N. hartmani, (N. 2010 1 1 1.1071 2010 2010 0 quin) 4286 N. quin 1670 1 1 1.1071 1670 1670 0 4286 n/a 4250 n/a 4000 n/a 3480 n/a 2490 R. miss 3520 1 1 1.1071 3520 3520 0 4286 S. gard 3200 1 1 1.1071 250 250 -2950 4286 S. umbonata 2100 1 1 1.1071 2100 2100 0 4286 S. archeri 350 1 1 1.1071 350 350 0 4286 T. elliptica, (T. 160 1 1 1.1071 160 160 0 stephensoni) 4286 T. steph 260 1 1 1.1071 260 260 0 4286 T.munda 390 1 1 1.1071 4250 4250 3860 4286 V. mullinensis 700 1 1 1.1071 700 700 0 4286 A. anteradiata 3560 1 1 1.1071 3560 3560 0 4286 B. carolinensis 860 1 0 0 extinct C. hodgei 4100 1 1 1.1071 4100 4100 0 4286

113

V. uvaladana 10 1 1 1.1071 10 10 0 4286 Total Total Desc (no immigrants) Ancestors 31 28 ances mean 1607.0 968 desc mean 2311.7 778 difference 704.68 1

SS AC Immi 126.832 51.42 526.42 857 0635 TS2 PE Answer 704.681

Time Increment 3

Ances An #ances #d rel Des. Range Non-imm Anag. Species Range es fitness size Desc Chng

C. grandis 2250 1 1 1.04255 2250 2250 0 319 N. micro 3230 1 0 0 extinct N. per 4250 1 1 1.04255 4250 4250 0 319 N. severn 1310 1 1 1.04255 1310 1310 0 319 N. corb 10 1 1 1.04255 10 10 0 319 N. kerrensis 160 1 1 1.04255 890 890 730 319 S. euf 3830 1 0 0 extinct S. ang 2650 1 1 1.04255 2650 2650 0 319 V. cren 2300 1 1 1.04255 2300 2300 0 319

114

T. patula 30 1 1 1.04255 30 30 0 319 A. mesen 4100 1 1 1.04255 4100 4100 0 319 A. argen 4250 1 2 2.08510 4250 4250 0 638 3580 3580 -670 n/a 1900 n/a 2050 n/a 4250 n/a 1470 B. rip 2300 1 1 1.04255 2300 2300 0 319 B. car 3150 1 1 1.04255 3150 3150 0 319 n/a 2050 n/a 4020 n/a 3780 C. serica 4100 1 1 1.04255 4250 4250 150 319 C. littlei 2330 1 1 1.04255 2330 2330 0 319 C. parva 3950 1 1 1.04255 3950 3950 0 319 C. alta 850 1 1 1.04255 850 850 0 319 n/a 4250 C. coon 2620 1 1 1.04255 2620 2620 0 319 E. corsicana 10 1 1 1.04255 10 10 0 319 E. cost. Cost 4250 1 1 1.04255 3700 3700 -550 319 F. pratti 820 1 1 1.04255 820 820 0 319 G. rot 3500 1 1 1.04255 10 10 -3490 319 G. dum 4100 1 1 1.04255 4100 4100 0 319 I williardi 10 1 1 1.04255 10 10 0 319 L. ellipticum 3860 1 0 0 extinct L. term 10 1 1 1.04255 10 10 0 319

115

L. acutata 2640 1 1 1.04255 2640 2640 0 319 L. caro 890 1 2 2.08510 890 890 0 638 1950 1950 1060 M. bulb 10 1 1 1.04255 10 10 0 319 M. shumardi 10 1 1 1.04255 10 10 0 319 N. hart 2010 1 0 0 extinct N. quin 1670 1 1 1.04255 1670 1670 0 319 N. euf 4250 1 1 1.04255 4250 4250 0 319 N. grandis 4000 1 1 1.04255 4000 4000 0 319 n/a 2250 P. terr 3480 1 1 1.04255 3480 3480 0 319 P. tet 2490 1 1 1.04255 2490 2490 0 319 R. miss 3250 1 1 1.04255 3250 3250 0 319 S. lineolatus 250 1 0 0 extinct S. umbonata 2100 1 1 1.04255 2100 2100 0 319 S. archeri 350 1 1 1.04255 350 350 0 319 T. buboana 4250 1 2 2.08510 4250 4250 0 638 2300 2300 -1950 T. ellip 160 1 1 1.04255 160 160 0 319 T. steph 260 1 1 1.04255 260 260 0 319 T. gabbi 4250 1 1 1.04255 4250 4250 0 319 V. mull 700 1 1 1.04255 100 100 -600 319 A. anter 3560 1 1 1.04255 2300 2300 -1260 319 C. hodgei 4100 1 1 1.04255 4100 4100 0 319 V. uvaladana 10 1 1 1.04255 10 10 0 319

116

Total Total Desc (no immigrants) Ancestors 49 47 ances mean 2222.8 571 desc mean 2224.4 643 difference 1.6071 429

SS A Immi C 13.9514 - 127.655 14 775 0

TS3 PE Answer 1.60714

Time Increment 4

Ances An #ances #des rel Des. Non-imm Anag. Species Range fitness Range Desc Chng size

C. grandis 2250 1 1 1.3043 2250 2250 0 4783 N. per 4250 1 1 1.3043 4250 4250 0 4783 N. severn 1310 1 1 1.3043 1310 1310 0 4783 N. 10 1 1 1.3043 10 10 0 corbentensi 4783 s N. tarensis 890 1 0 0 extinct

S. anguli 2650 1 1 1.3043 2650 2650 0 4783

117

V. 2300 1 0 0 extinct subcircula T. patula 30 1 1 1.3043 30 30 0 4783 A. mesen 4100 1 1 1.3043 4100 4100 0 4783 A. argen 4250 1 1 1.3043 4250 4250 0 4783 A. tell 3580 1 0 0 extinct A. euf 1900 1 1 1.3043 1900 1900 0 4783 A. n.sp 2050 1 1 1.3043 2050 2050 0 4783 A. tipp 4250 1 1 1.3043 4250 4250 0 4783 A.regia 1470 1 1 1.3043 1470 1470 0 4783 B. 3150 1 1 1.3043 3150 3150 0 carolinensis 4783 B. para 2050 1 1 1.3043 2050 2050 0 4783 C. bur 4020 1 0 0 extinct C. virgatus 3780 1 0 0 extinct C. 4250 1 1 1.3043 4250 4250 0 elegantula 4783 C. litt 2330 1 1 1.3043 2330 2330 0 4783 C. parva 3950 1 1 1.3043 3950 3950 0 4783 C. alta 850 1 1 1.3043 850 850 0 4783 C. wordeni 4250 1 1 1.3043 4250 4250 0 4783 C. coon 2620 1 1 1.3043 2620 2620 0 4783 E. 10 1 1 1.3043 10 10 0 corsincana 4783 E. cos cos 3700 1 1 1.3043 3700 3700 0 4783 E. 3700 1 0 0 extinct cancellata F. pratti 820 1 1 1.3043 820 820 0 4783

118

G. rotunda 3500 1 1 1.3043 3500 3500 0 4783 G. lacertosa 10 1 0 0 extinct G. dum 4100 1 1 1.3043 4100 4100 0 4783 I. williardi 10 1 1 1.3043 10 10 0 4783 L. term 10 1 1 1.3043 10 10 0 4783 L. acutata 2640 1 1 1.3043 2640 2640 0 4783 L. carolin 890 1 1 1.3043 890 890 0 4783 L. orn 50 1 0 0 extinct L. n.sp 10 1 0 0 extinct M.bulb 10 1 1 1.3043 10 10 0 4783 M.shum 10 1 1 1.3043 10 10 0 4783 M. conradi 10 1 0 0 extinct N. quin 1670 1 0 0 extinct N. euf 4250 1 1 1.3043 4250 4250 0 4783 N. grandis 4000 1 1 1.3043 4000 4000 0 4783 P. decisa 2250 1 1 1.3043 2250 2250 0 4783 P. terr 3480 1 1 1.3043 3480 3480 0 4783 P.tet 2490 1 1 1.3043 2490 2490 0 4783 R. miss 3250 1 1 1.3043 3250 3250 0 4783 S. umb 2100 1 1 1.3043 2100 2100 0 4783 S. archeri 350 1 1 1.3043 350 350 0 4783 T. eborea 2300 1 1 1.3043 2300 2300 0 4783 T. buboana 4250 1 1 1.3043 4250 4250 0 4783 T. ellip 160 1 1 1.3043 160 160 0 4783

119

T. steph 260 1 1 1.3043 260 260 0 4783 T. gabbi 4250 1 1 1.3043 4250 4250 0 4783 V. lineata 100 1 0 0 extinct A. anter 3560 1 1 1.3043 3560 3560 0 4783 A. lata 2300 1 0 0 extinct C. hodgei 4100 1 1 1.3043 4100 4100 0 4783 V. sibteres 10 1 0 0 extinct Total Total Desc (no immigrants) Ancestors 60 46

ances 2185.833 mean desc 2363.478 mean differe 177.6449 nce SS AC Immi 177.6449 0 0

TS4 PE Answer 177.6449

Time Increment 5

Ances An #ances #de rel fitness Des. Range Non-imm Anag. Species Range s size Desc Chng

C. grandis 2250 1 2 2.1363636 2250 2250 0 36 0 2190 2190 -60 N. per 4250 1 1 1.0681818 4250 4250 0 18 N. severn 1310 1 0 0 extinct

120

N. 10 1 1 1.0681818 10 10 0 corbentensis 18 n/a 10 S. euf 3830 1 1 1.0681818 3830 3830 0 18 S. anguli 2650 1 1 1.0681818 2650 2650 0 18 T. patula 30 1 1 1.0681818 30 30 0 18 A. mesen 4100 1 1 1.0681818 4100 4100 0 18 A. argen 4250 1 1 1.0681818 4250 4250 0 18 A. euf 1900 1 0 0 extinct A. n.sp 2050 1 1 1.0681818 2050 2050 0 18 A. tipp 4250 1 1 1.0681818 4250 4250 0 18 A.regia 1470 1 1 1.0681818 1470 1470 0 18 B. 3150 1 1 1.0681818 3150 3150 0 carolinensis 18 B. para 2050 1 1 1.0681818 1960 1960 -90 18 n/a 4250 C. 4250 1 1 1.0681818 4250 4250 0 elegantula 18 C. litt 2330 1 1 1.0681818 2330 2330 0 18 C. parva 3950 1 1 1.0681818 3950 3950 0 18 C. alta 850 1 1 1.0681818 850 850 0 18 C. wordeni 4250 1 2 2.1363636 4250 4250 0 36 0 45 45 -4205 C. coon 2620 1 1 1.0681818 3560 3560 940 18 E. 10 1 1 1.0681818 10 10 0 corsincana 18 E. cos cos 3700 1 1 1.0681818 4250 4250 550 18 F. pratti 820 1 1 1.0681818 2650 2650 1830 18

121

G. rotunda 3500 1 1 1.0681818 3500 3500 0 18 G. dum 4100 1 1 1.0681818 4100 4100 0 18 I. williardi 10 1 1 1.0681818 10 10 0 18 n/a 300 L. term 10 1 1 1.0681818 10 10 0 18 n/a 3830 L. acutata 2640 1 1 1.0681818 2640 2640 0 18 L. carloi 890 1 1 1.0681818 890 890 0 18 M.bulb 10 1 1 1.0681818 10 10 0 18 M.shum 10 1 1 1.0681818 10 10 0 18 N. euf 4250 1 1 1.0681818 4250 4250 0 18 N. grandis 4000 1 0 0 extinct P. decisa 2250 1 1 1.0681818 2250 2250 0 18 P. terr 3480 1 1 1.0681818 3480 3480 0 18 P.tet 2490 1 1 1.0681818 2490 2490 0 18 n/a 3960 R. miss 3250 1 1 1.0681818 3250 3250 0 18 S. umb 2100 1 1 1.0681818 2100 2100 0 18 S. archeri 350 1 1 1.0681818 350 350 0 18 T. eborea 2300 1 0 0 extinct T. buboana 4250 1 1 1.0681818 4250 4250 0 18 T. ellip 160 1 0 0 extinct T. steph 260 1 1 1.0681818 260 260 0 18 T. gabbi 4250 1 1 1.0681818 4250 4250 0 18 A. anter 3560 1 1 1.0681818 3560 3560 0 18

122

C. hodgei 4100 1 1 1.0681818 4100 4100 0 18 Total Total Desc (no immigrants) Ancestors 47 44

ances 239 mean 5 desc mean 246 3 difference 68. 48 SS AC Immi 91.228239 - 0.7769016 85 23.522727 7 27

TS5 PE Answer 68.482414 24

Time Increment 6

Ances An Range #ances #des rel Des. Non-imm Anag. Species fitness Range size Desc Chng

C. grandis 2250 1 0 0 extinct C. jer 2190 1 0 0 extinct N. per 4250 1 1 1.24324 4250 4250 0 324 N. colorad 10 1 0 0 extinct N. 10 1 1 1.24324 10 10 0 corbentensi 324 s S. anguli 2650 1 1 1.24324 2650 2650 0 324 T. patula 30 1 1 1.24324 10 10 -20 324 A. mesen 4100 1 0 0 extinct A. argen 4250 1 1 1.24324 4250 4250 0 324

123

A. n.sp 2050 1 0 0 extinct A. tipp 4250 1 1 1.24324 4250 4250 0 324 A.regia 1470 1 1 1.24324 1470 1470 0 324 B. 3150 1 1 1.24324 3150 3150 0 carolinensis 324 B. fragile 1960 1 1 1.24324 1960 1960 0 324 n/a 110 n/a 10 C. crass 4250 1 1 1.24324 1920 1920 -2330 324 n/a 240 C. 4250 1 1 1.24324 4250 4250 0 elegantula 324 C. litt 2330 1 1 1.24324 2330 2330 0 324 C. parva 3950 1 1 1.24324 3950 3950 0 324 C. alta 850 1 1 1.24324 850 850 0 324 C. wordeni 4250 1 0 0 extinct C. app 45 1 1 1.24324 45 45 0 324 C. alta 3560 1 0 0 extinct E. 10 1 1 1.24324 10 10 0 corsincana 324 E. cos cos 3700 1 1 1.24324 3700 3700 0 324 F. sub 2650 1 0 0 extinct G. rotunda 3500 1 1 1.24324 3500 3500 0 324 G. dum 4100 1 1 1.24324 4100 4100 0 324 I. williardi 10 1 1 1.24324 10 10 0 324 L.elongata 300 1 1 1.24324 10 10 -290 324 L. term 10 1 1 1.24324 30 30 20 324 L. meta 3830 1 1 1.24324 3560 3560 -270 324

124

L. acutata 2640 1 2 2.48648 2640 2640 0 649 10 10 -2630 L. carloi 890 1 1 1.24324 890 890 0 324 M.bulb 10 1 1 1.24324 10 10 0 324 M.shum 10 1 1 1.24324 10 10 0 324 N. euf 4250 1 0 0 extinct n/a 30 P. decisa 2250 1 1 1.24324 2250 2250 0 324 P. terr 3480 1 1 1.24324 3480 3480 0 324 P.tet 2490 1 1 1.24324 2490 2490 0 324 P. mull 3960 1 1 1.24324 3960 3960 0 324 R. miss 3250 1 1 1.24324 3250 3250 0 324 S. umb 2100 1 1 1.24324 2100 2100 0 324 S. archeri 350 1 1 1.24324 350 350 0 324 T. buboana 4250 1 0 0 extinct A. anter 3560 1 1 1.24324 3560 3560 0 324 C. hodgei 4100 1 1 1.24324 4100 4100 0 324 Total Total Desc (no immigrants) Ancestors 46 37 ances mean 2430.5435 desc mean 1945.2439 difference -485.2996 SS AC Immi -136.3543 -149.189 - 199.7 56

TS6 PE Answer -485.2996

125

Time Increment 7

Ances An #ances #des rel Des. Non-imm Anag. Species Range fitness Range Desc Chng size

N. per 4250 1 0 0 extinct N. 10 1 0 0 extinct corbentensi s S. anguli 2650 1 1 1.5185 2650 2650 0 185 T. macro 10 1 0 0 extinct A. argen 4250 1 1 1.5185 4250 4250 0 185 A. tipp 4250 1 1 1.5185 4250 4250 0 185 A.regia 1470 1 1 1.5185 1470 1470 0 185 B. carolin 3150 1 1 1.5185 3150 3150 0 185 B. fragile 1960 1 1 1.5185 1960 1960 0 185 B. tenue 110 1 0 0 extinct B. guad 10 1 0 0 extinct C. will 1920 1 0 0 extinct C.sayrei 240 1 1 1.5185 30 30 -210 185 C. 4250 1 1 1.5185 4250 4250 0 elegantula 185 C.littlei 2330 1 1 1.5185 3780 3780 1450 185 C.alta 850 1 1 1.5185 850 850 0 185 C.parva 3950 1 1 1.5185 3950 3950 0 185 C.appressa 45 1 1 1.5185 45 45 0 185 E.corsicana 10 1 0 0 extinct E. cos cos 4250 1 1 1.5185 4250 4250 0 185

126

G. 3500 1 1 1.5185 3500 3500 0 rotundata 185 n/a 1300 n/a 2220 G. dum 4100 1 1 1.5185 2410 2410 -1690 185 I.will 10 1 1 1.5185 40 40 30 185 L. quad 10 1 0 0 extinct L.levis 30 1 0 0 extinct L.cribelli 3560 1 0 0 extinct L. actutata 2640 1 0 0 extinct L. burrana 10 1 0 0 extinct L. carolin 890 1 1 1.5185 890 890 0 185 M.Bulb 10 1 1 1.5185 1610 1610 1600 185 M. shum 10 1 0 0 extinct P.wadei 30 1 1 1.5185 10 10 -20 185 P. decisa 2250 1 1 1.5185 2250 2250 0 185 P.terr 3480 1 1 1.5185 3480 3480 0 185 n/a 2300 P.tet 2490 1 1 1.5185 3310 3310 820 185 P.mull 3960 1 1 1.5185 1870 1870 -2090 185 R.miss 3250 1 1 1.5185 2300 2300 -950 185 S.umb 2100 1 1 1.5185 220 220 -1880 185 S.archeri 350 1 0 0 extinct A.anter 3560 1 1 1.5185 3560 3560 0 185 C.hodgei 4100 1 1 1.5185 4250 4250 150 185 Total Total Desc (no immigrants) Ancestors 41 27

ances 1958.66 mean

127

desc 2346.83 mean differe 388.175 nce SS AC Immi 536.712 - - 103.33 45.203703 333 7

TS7 PE Answer 388.175

Time increment 8

Ances An #ances #des rel Des. Non-imm Anag. Species Range fitness Range Desc Chng size

N. per 4250 1 1 2.0666 4250 4250 0 67 S.ang 2650 1 1 2.0666 2650 2650 0 67 A.argen 4250 1 1 2.0666 4250 4250 0 67 A.tipp 4250 1 0 0 extinct A.regia 1470 1 0 0 extinct B.car 3150 1 1 2.0666 3150 3150 0 67 B.fragile 1960 1 1 2.0666 1960 1960 0 67 C.riddlei 30 1 0 0 extinct C.elegantul 4250 1 0 0 extinct a C.capax 3780 1 0 0 extinct C.alta 3560 1 1 2.0666 3560 3560 0 67 C.parva 3950 1 1 2.0666 3950 3950 0 67

128

C.appressa 45 1 1 2.0666 45 45 0 67 E. cos cos 4250 1 1 2.0666 4250 4250 0 67 G.rotunda 3500 1 1 2.0666 3500 3500 0 67 G.alabam 1300 1 0 0 extinct G.lowei 2220 1 0 0 extinct G.tipp 2410 1 0 0 extinct I.holmesi 40 1 0 0 extinct L.carolin 890 1 1 2.0666 890 890 0 67 M.cliff 1610 1 0 0 extinct P.stantoni 10 1 0 0 extinct P.decisa 2250 1 1 2.0666 2350 2350 100 67 P.terr 3480 1 1 2.0666 2650 2650 -830 67 P.petrosa 2300 1 1 2.0666 3500 3500 1200 67 P.clarki 3310 1 0 0 extinct P.urticosa 1870 1 0 0 extinct R.weeksi 2300 1 0 0 extinct S.poguei 220 1 0 0 extinct A.anter 3560 1 1 2.0666 3560 3560 0 67 C.vadosa 4250 1 0 0 extinct Total Total Desc (no immigrants) Ancestors 31 15 ances mean 2495.6 SS AC Immi 45 desc mean 2967.6 440.69 31.333 0 67 33 difference 472.02 15 TS8 PE Answer 472.02