BIOTIC INTERACTIONS OF BIVALVES FROM THE LATE CRETACEOUS COON CREEK TYPE SECTION OF MCNAIRY COUNTY, TENNESSEE

A thesis submitted to Kent State University in partial fulfillment of the requirements for the degree of Master of Science

by

Elizabeth Carol Rhenberg

December, 2007

Thesis written by Elizabeth Carol Rhenberg B.S., The University of Tennessee, Martin, 2005 M.S., Kent State University, 2007

Approved by

______, Advisor Rodney M. Feldmann

______, Chair, Department of Geology Donald F. Palmer

______, Dean, College of Arts and Sciences Jerry D. Feezel

ii TABLE OF CONTENTS

ACKNOWLEDGMENTS ...... vi

SUMMARY ...... 1

INTRODUCTION...... 2

PREVIOUS WORK ...... 4

METHODS AND MATERIALS ...... 9

BIOTIC INTERACTIONS ...... 12

IMPORTANCE OF LOCATION ...... 33

CONCLUSIONS...... 39

REFERENCES...... 43

APPENDIX 1...... 54

APPENDIX 2...... 76

APPENDIX 3...... 81

iii LIST OF FIGURES

FIGURE 1: Area Map...... 3

FIGURE 2: Stratigraphic column...... 5

FIGURE 3: Entobia interactions ...... 15

FIGURE 4: Entobia, Mycelites, Zapfella, and gastropod interactions...... 16

FIGURE 5: Burrowing bivalve interactions ...... 19

FIGURE 6: Gastropod interactions ...... 20

FIGURE 7: Oyster attachments...... 25

FIGURE 8: Anomia and foraminifera interactions...... 27

FIGURE 9: Bryozoan interactions ...... 29

FIGURE 10: Zapfella, Mycelites, and worm interactions ...... 31

iv LIST OF TABLES

TABLE 1: Bivalves studied ...... 13

TABLE 2: Locations and percentages of interactions...... 17

TABLE 3: Crassatella drilled by gastropods ...... 23

v ACKNOWLEDGMENTS

For allowing me to work with their fossils, I would like to thank the Pink Palace

Museum, the Cleveland Museum of Natural History, and the National Museum of

Natural History at the Smithsonian Institution. I would especially like to thank Ron

Brister and the Pink Palace Museum for allowing me to collect in the field at Coon Creek

Science Center. I would also like to thank Dr. Rodney Feldmann for all his help and support during the course of this project.

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CHAPTER 1: Summary

The bivalves and mollusks of the Late Cretaceous Coon Creek Formation in

Tennessee are well preserved and exhibit biotic interactions with a variety of organisms.

Almost 60% of the bivalves collected in the field show some form of interaction; many show multiple forms of biotic activity. These interactions provide information on how the bivalves lived, and if they were alive at the time the other organisms were interacting with them. Bivalves were used as substrates by several organisms, including sponges, bryozoans, other bivalves, worms, microbial fungi, and foraminifera. Predation of the bivalves by gastropods is also noted. Location of burrows, impressions, and drill holes provides information on the orientation of the living bivalves. Interactions that are seen on the interior of the valves indicates that the bivalve was dead at the time the other organism began living on it.

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CHAPTER 2: Introduction

The molluscan assemblage of the Coon Creek Formation in McNairy County,

Tennessee (Fig. 1) is a classic Late Cretaceous fauna. A large number of species is represented, yet not much work has been done on the fauna itself since Wade (1926) initially described it, save for the work of Sohl (1960; 1964) on the gastropods and of

Bishop (1985) on the decapods.

This study will focus on the biotic interactions of the bivalves with other organisms by looking at the physical marks left behind by organisms including sponges, gastropods, and barnacles, for example. The illustrations found in Wade (1926) under- represent biotic interactions due to his attempt to provide “perfect” specimens to illustrate, and previous work on the Coon Creek Formation has not dealt with these interactions. The frequency of the borings as well as their distribution on the shells will be noted within different bivalve taxa.

This study will provide new information on the paleocological framework of the formation by focusing on biotic interactions. The Coon Creek Formation mollusks exhibit exceptional preservation of original shell material; therefore, traces left from the interactions of other organisms are well documented.

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Enville

Coon Creek

Adamsville

Selmer

McNairy County

Figure 1: Map of the study area in McNairy County, Tennessee

CHAPTER 3: Previous Work

Stratigraphy.— The fossiliferous sands and clays of the Coon Creek Formation are shallow marine sediments that were deposited in between the deeper shelf sediments of the Demopolis Formation and the nearshore, shoreline, and fluvial deposits of the

McNairy Sand (Russell, 1975). These sediments were deposited as a part of a regressive sequence during the early Maastrichtian (Fig. 2).

The type locality of the Coon Creek Formation (35° 21.16'N, 88° 24.88'W) is known as the Dave Weeks Place, which is situated along Coon Creek. The site was discovered by Bruce Wade who worked in the area from 1917-1926. The Weeks Place lies in the northeastern part of McNairy County, 5.6 km south of Enville and 12.1 km north of Adamsville, TN. The type locality is approximately 230 m east of Dave Weeks’ house along the headwaters of Coon Creek (Wade, 1926). Today the Coon Creek

Science Center is located above the flood plain and below Dave Weeks’ old house, and consists of several cabins with a mess hall for lodging and a research laboratory.

Historically, the Coon Creek Formation has been considered to be a tongue of the

Ripley Formation (Fig. 2). The Ripley Formation embraces rocks exposed from Georgia to Illinois. However, in Tennessee, Russell (1975) subdivided the Ripley into the Coon

Creek and McNairy formations on the grounds that they are lithologically distinct and

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Figure 2: Historical stratigraphic column of the Coon Creek Formation, modified from

Russell (1975). When Sohl (1960, 1964) described the gastropods, he followed

Stephenson and King (1942).

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mappable, and that they are the lateral equivalents of the Ripley Formation of

Mississippi. The Coon Creek Formation was separated into two horizons by Wade (1926) based on location and lithology; a ferruginous sand horizon with few or no fossils found in the northern part of Tennessee and a fossiliferous and glauconitic horizon in the southern part. At the type locality, the southern horizon is dominant, typified by massive silty sand beds that are dark- to bluish-grey in color and micaceous, glauconitic, calcareous, and fossiliferous (Sohl, 1960).

The Coon Creek Formation is underlain by the Demopolis Formation which consists of marl, chalky marl, and calcareous clay (Russell, 1975). This contact is exposed at the northernmost end of the Coon Creek Science Center property. The contact between the Demopolis and Coon Creek formations is gradational, which makes it difficult to define a precise boundary between them. Therefore, the base of the Coon

Creek is chosen either at the top of the highest marl unit and the base of the transitional clay or at the base of the lowermost persistent glauconitic sands.

The McNairy Sand overlies the Coon Creek Formation. The contact cannot be seen at the type locality. The contact between the Coon Creek and McNairy formations is easier to recognize than the Demopolis-Coon Creek contact in most places (Russell,

1975). The top of the Coon Creek is identified by thin-bedded, light-grey clays that are overlain by the clean, very fine-grained, heavy-mineral-rich sands that compose the

McNairy Sand. In other places, the Coon Creek becomes less glauconitic, silty, and clayey, closely resembling the McNairy Sand and creating a contact that is indistinct.

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The main focus of this paper is not on the stratigraphy of the area, therefore no stratigraphical work was done while in the field.

Paleontology.— Much of the previous work done on the Coon Creek Formation focused on the stratigraphy. Moore (1974) cited the works of Morton, Troost, Safford, and

Hilgard and their descriptions of the lithologic material, and Russell (1975) cited all the previous authors except Hilgard and cited Sohl’s stratigraphic work (Moore only cited his work with the gastropods). When Safford (1869) described the green sands of the Ripley

Formation, he listed the fossils found, but did not mention finding trace fossils.

Stephenson began his studies of the Atlantic-Gulf Coastal Plain in 1914, establishing the Exogyra biostratigraphic zones which are widely recognized as zonal markers throughout the Gulf and Atlantic Coastal Plains (Cobban and Kennedy, 1994).

The Coon Creek Formation lies within the Exogyra costata zone (Wade, 1926), which includes the Exogyra cancellata subzone (Moore, 1974).Wade (1926) started his work in

1917, publishing a series of shorter papers until 1922. He published his extensive study on the fauna of the Ripley Formation exposed at I 1926, Coon Creek, listing 370 species in 215 genera. Sohl (1960, 1964) updated the gastropod systematics. Foraminifera were studied by Berry and Kelley (1929) and by Cushman (1931). Stratigraphic interpretations were made by Pryor and Glass (1961) and Russell (1975). Moore (1974) conducted a systematic and paleoecologic review of the Coon Creek fauna, in which he briefly reviewed the bivalve systematics. Whetstone (1977) noted a plesiosaur; Bishop

(1985) studied the crabs; Vermeij and Dudley (1982) investigated shell repair and drilling in some gastropods; and Cobban and Kennedy (1994) studied ammonites. Most recently,

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Bandel and Dockery conducted a systematic discussion of the Gastropoda (unpublished).

As noted by these works, only Vermeij and Dudley (1982) have studied any form of biotic interactions, and that was done only with gastropods.

Studies of biotic interactions are as varied are the interactions themselves, and they are too numerous to cover properly here. However, a few examples that were particularly helpful to this study are cited here. Several papers discussed attachment strategies of organisms: Bromley and Heinberg (2006) report on a wide range of attachment strategies of organisms to hard substrates, whereas Johnson and Baarli (1999) focused on rocky-shore interactions. Best and Kidwell (2000) discussed the differences between infaunal and epifaunal bivalves found in death assemblages. Various aspects of gastropod predation have been studied, including the efficiency (Kelley et al., 2001), prey selectivity (Kelley and Hansen, 1996; Kitchell et al., 1981), and drilling intensities by gastropods in the Fox Hills Formation (Harries and Schopf, 2007).

CHAPTER 4: Methods and Materials

During March, 2006, preliminary investigations of the Coon Creek type locality and its fauna were conducted by visiting both the Pink Palace Museum of Memphis, TN, and the Coon Creek Science Center near Enville, TN. At the Pink Palace Museum, the collection of bivalves was examined, and specimens that exhibited some form of borings or biotic interaction were photographed to provide basic information on the types of interactions that were present. This provided an initial assessment of the degree to which different bivalve taxa were susceptible to interactions with other animals.

At the Coon Creek Science Center, specimens were collected by two methods: traversing the creek and collecting loose shells, and collecting within the large sediment piles recently excavated by the workers from the Pink Palace Museum. All collected samples were be cleaned and identified, and were then separated according to whether or not they exhibited physical signs of biotic interactions. The type of interaction was also noted.

In late May, a return trip to the Coon Creek Science Center was made to do more collecting, which provided a more diverse array of bivalve taxa to study. The same methods of collecting were used as before. Additionally, wet sieving of the gravel bars that form along the creek was also done. The sieving did not provide any useable material. Two return trips to the Pink Palace Museum also were made in July to examine the collection there in more detail. The first trip was spent doing a detailed examination

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of each species of bivalve in the collection, whereas the second trip was spent selecting fossils to borrow for the duration of the project.

In November 2006, a trip to the Cleveland Museum of Natural History was made to examine the Coon Creek fossils they had acquired. The collection was not large, but it did provide several species of bivalves that preserved biotic interactions. It also permitted examination of a large number of the benthic bivalve Crassatellites vadosus

Morton, 1834. These were used to mark the position of gastropod borings on each valve to determine whether the gastropods showed preference for a particular valve.

In January, 2007, Wade’s original collection of fossils from Coon Creek was inspected at the National Museum of Natural History at the Smithsonian Institution.

Unfortunately, most of Wade’s collection did not preserve many biotic interactions. The fossils depicted in the monograph were either chosen because they were pristine and photogenic, or the interactions were disguised in the way they were preserved by being soaked in heated paraffin wax (Wade, 1917). Several shells were found that might have shown interactions, but nothing could be discerned with certainty. The collection of

Norman Sohl was also examined, and his fossils proved to be more useful for the purposes of this project. However, the useful bivalves were not found in abundance; his studies focused on gastropods.

The fossils, both those borrowed from museums and those collected, have been scrutinized to find all traces of biotic interactions. All interactions have been categorized and their positions on the bivalves have been noted. Using the categories described below, percentages of the interactions have been made. Only the fossils personally

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collected have been used in computing the percentages because of the bias of museum collections toward pristine fossils.

Determination of the interactions was made by consulting the Treatise on

Invertebrate Paleontology (Häntzschel, 1975). Where the burrows were infilled with sediment, it was difficult to determine whether they were constructed by clionids or burrowing bivalves. Therefore, anything that was within the clionid size range of 1.5 to 4 mm (Bromley, 1970) was categorized as a clionid. Anything larger was considered to have been constructed by a bivalve.

At the completion of this study, the fossils will be stored at the Pink Palace

Museum in Memphis, Tennessee.

CHAPTER 5: Biotic Interactions

Overview.— The bivalve fauna within the Coon Creek Formation is diverse and extensive. In 1917, Wade’s preliminary assessment of the Coon Creek fauna included 49 genera and 110 species of bivalves. His reevaluation of the fauna in 1926 increased the number of bivalve genera to 65 with a total of 114 species. The most recent review of the

Coon Creek fauna (Moore, 1974) modified the numbers slightly, with 68 genera and 111 species. The present study includes 34 species in 22 genera (Table 1), using the most recent identifications of the bivalves which were confirmed using the Treatise on

Invertebrate Paleontology (Cox et al., 1969a, b; Stenzel, 1971). Of the 34 species, 18 were collected at Coon Creek and were not borrowed from museum collections.

Of the 238 specimens collected, 139 showed signs of biotic interactions. All specimens borrowed from the Pink Palace Museum exhibited biotic interactions except

Leptosolen biplicata Conrad, 1858, and Legumen planulatum (Conrad, 1853). Out of an additional 45 specimens from the Cleveland Museum of Natural History, 24 exhibited interactions. Of the 200 or so fossils examined in the United States National Museum of

Natural History, 52 showed biotic activity.

A few infaunal bivalves were collected and examined, and none of them had any form of biotic interactions, so all of the bivalves discussed below are epifaunal.

Overall, 63 percent of the fossils collected showed some form of biotic interactions. The most common interactions displayed are produced by clionid sponges

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TABLE 1—List of the bivalves used in this study. The names that Wade used in his original description are on the left and the updated names are on the right.

Family Wade (1926) Species Moore (1974) Species Nuculidae Nucula percrassa Conrad, 1858 Parallelodontidae Nemodon grandis Wade, 1926 Cuculladeidae Cucullaea vulgaris Morton, 1830 Idonearca vulgaris Arcidae Arca securiculata Wade, 1926 Barbatia fractura Wade, 1926 Glycymerididae Glycimeris subcrenata Wade, 1926 Glycimeris microsulci Wade, 1926 Glycimeris lacertosa Wade, 1926 Inoceramidae Inoceramus proximus Tuomey, 1854 Inoceramus sagensis Owen, 1852 Isognomonidae Pedalion periridescens (Wade, Isognomon periridescens 1926) Bakevelliidae Gervilliopsis ensifromis (Conrad, 1858) Ostreidae Ostrea tecticosta Gabb, 1860 Ostrea falcata Morton, 1829 Gryphaeidae Exogyra costata Say, 1820 Exogyra cancellata Stephenson, 1914 Gryphaea vesicularis (Lamarck, Pycnodonte vesicularis 1806) Trigoniidae Trigonia thoracica (Morton, 1834) Pterotrigonia (Scabrotrigonia) thoracica Pectinidae Pecten quinquenarius Conrad, 1853 Pecten burlingtonensis (Gabb, Camptonectes 1860) burlingtonensis Pecten quinquecostatus(Sowerby, Neithea (Neitheopsis) 1814) quinquecostatus Anomiidae Paranomia scabra (Morton, 1834) Anomia argentaria Morton, 1833 Anomia perlineata Wade, 1926 Pholadomyidae Pholadomya occidentalis Morton, 1833 Arcticidae Veniella conradi (Morton, 1833) Crassatellidae Crassatellites vadosus (Morton, Crassatella vadosus 1834)

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Cardiidae Cardium stantoni (Wade, 1926) Protocardia (Pachycardium) stantoni Veneridae Meretrix cretacea (Conrad, 1870) Aphrodina cretacea Legumen planulatum (Conrad, 1853) Cyprimeria alta Conrad, 1875 Cultellidae Leptosolen biplicata Conrad, 1858 Corbulidae Corbula crassiplica Gabb, 1860 Astartidae Astarte? sp. Sowerby, 1816

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(37.4%), burrowing bivalves (15.6%), drilling gastropods (11.3%), and attachments to other organisms (mostly bivalves, 8.0%), but those are not the only interactions found.

Bryozoans, worms, and a variety of other trace fossils are seen on the bivalves. The types of trace fossils found were determined by using the Treatise on Invertebrate

Paleontology (Häntzschel, 1975).

Clionid sponge borings.—The most common form of interactions is with the boring sponges. Sixty-four percent of the collected fossils collected that showed interactions were bored by Entobia Bronn, 1838. These borings consist of globular chambers

(Häntzschel, 1975) and range in size from 1.5 to 4 mm (Bromley, 1970). The borings are found on a variety of species and in a variety of places (Fig. 3, Figs. 4.1-4.2, 4.4-4.6).

The distribution of the sponge borings on the valves is fairly even with respect to which valve the sponges are boring on. Almost 50% of the sponges (Table 2) are found only on left valves, while 41.6% was found on right valves only. By testing the observed numbers (right valve with 37 interactions, left valve with 44) using the chi-squared analysis (Reyment, 1971), the result of 0.605 with 1 degree of freedom at 0.05 shows that there is no preference in which valve the sponges bored on.

Exclusively external clionid borings were much more common (40.4%) than exclusively internal borings (6.7%). This is due to how difficult it would be for a valve to be exposed in such a way that it would allow for internal boring only. However,

52.8% of the shells with borings are found on both the external and internal surfaces of the bivalve. This number is not surprising as all the bivalves that show borings on both

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Figure 3: All scale bars are equal to 1 cm. 1, 2 – Crassatella vadosus, right valve, showing Enotobia interactions 1 – internal view, 2 – external view; 3 – Glycimeris subcrenata, right valve, showing Enotobia interactions – external view; 4 – Crassatella vadosus, left valve, showing Enotobia interactions. Internal view showing central line of Enotobia; 5 – Idonearca vulgaris, right valve, showing Enotobia interactions – external view; 6 – Veniella conradi, left valve, showing Enotobia interactions – external view showing Enotobia only on ventral, posterior margin; 7, 8 - Veniella conradi, both valves 7 – external view of left valve, 8 – external, posterior view of both valves, showing Enotobia interactions only on the right valve.

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Figure 4: All scale bars are equal to 1 cm. 1, 2 – Exogyra costata, left valve – multiple interactions on one bivalve, including clionid sponge borings, attachment scar, and relicts of other oysters attaching. 1 – external view, 2 – internal view; 3 – Protocardia (Pachycardium) stantoni, left valve – multiple interactions including Zapfella (lower arrows), Mycelites (dorsal arrow), and bivalve burrows (large holes); 4 – Protocardia (Pachycardium) stantoni, right valve – multiple interactions include serpulid worm attachments along the dorsal margin, and clionid sponge borings; 5, 6 - Cyprimeria alta, right valve – multiple interactions include clionid sponge borings and gastropod drill hole.

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TABLE 2—Locations and percentages of the most common evidences of interaction that were collected on site.

Interaction Location Totals Percentages Clionid Sponge 89 Right 3741.6 Left 4449.4 Both 44.5 Unknown 44.5

Internal 66.7 External 3640.4 Both 4752.8

Gastropod drill hole 29 Right 1344.8 Left 1551.7 Unknown 13.4

Success 2379.3 Fail 620.7

Attachment scars on oysters 9 Right 222.2 Left 777.8

Evidence of oyster attached on other bivalves 10 Right 330.0 Left 660.0 Both 110.0

Internal 550.0 External 550.0

Boring Bivalves 37 Right 1232.4 Left 2567.6

Internal 25.4 External 2464.9 Both 1129.7

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surfaces are disarticulated, indicating that after the bivalve died it lay on the seafloor for some time before burial, allowing the sponges time to use the valves as a substrate.

Boring bivalves.— Found in several specimens, boring bivalves are common, comprising 26.6% of the interactions. Most of the borers are small and must be seen with a microscope as only a small portion protrudes from the sediment infill. A few specimens are large enough to see with the unaided eye. The internal side of a

Paranomia scabra (Morton, 1834) (Figs. 5.5-5.6) shows the borings of many bivalves, with one disarticulated boring bivalve still inside its boring. The right valve of an

Exogyra cancellata Stephenson, 1914 (Figs. 5.7-5.8) shows similar large borings from other bivalves, with one bivalve still inside one of the holes. Other bivalves (Figs. 5.1-

5.2) have large holes that may be sites in which smaller bivalves once bored. There is some speculation as to which borings are from clionid sponges and which are from boring bivalves when the holes are either empty or infilled with sediment.

Of the valves found that have bivalve borings, two-thirds (67.6%) are found on the left valves (Table 2). The preference for the left valve may be due to the number of

Exogyra valves collected, as the large left valve provides a good substrate for the bivalves could bore. Most of the burrowing bivalves are found on the external surface of larger bivalves (64.9%), with a significant number (29.7%) found on both internal and external surfaces.

Gastropod drilling.— Predation by gastropods (Fig. 6) is relatively common within the specimens collected (20.9% of interactions). The most commonly drilled species was

Crassatella vadosus, but that is because of the abundance of C. vadosus fossils collected.

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Figure 5: All scale bars are equal to 1 cm unless otherwise noted. 1, 2 – Crassatella vadosus, right valve – large holes are believed to have been caused by boring bivalves that have since been separated. 1 – internal view, 2 – external view; 3, 4 – Crassatella vadosus, left valve – external view showing multiple bivalve holes, many still containing bivalves. 3 – full external view, 4 – close up of the dorsal end, showing bivalves still within the borings (arrows), scale bar equal 0.5 cm; 5, 6 – Paranomia scabra, right valve – internal view showing multiple bivalve holes. 5 – full internal view showing several bivalve holes, 6 – close up of ventral margin showing a disarticulated bivalve (arrow), scale bar equal 0.5 cm; 7, 8 – Exogyra cancellata, right valve – external view. 7 – full view showing multiple borings, 8 – close up of the central area showing several borings as well as one bivalve that is still inside (arrow).

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Figure 6: All scale bars are equal to 1 cm unless otherwise noted. 1 – Crassatella vadosus, left valve – failed drill hole; 2 – Astarte? sp. – scale bar equal to 0.2 cm, successful gastropod drill hole; 3 – Anomia perlineata, left valve, successful gastropod drill hole; 4 – Paranomia scabra, right valve, successful gastropod drill hole; 5 – Veniella conradi, right valve – posterior view showing shape of the drilled hole; 6 – Idonearca vulgaris, right valve – drill hole on the ventral margin; 7 – Exogyra cancellata, left valve, successful gastropod drill hole; 8 – Pecten quinquenarius, left valve, successful gastropod drill hole; 9 – Aphrodina cretacea, right valve – two successful drill holes.

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Other species that exhibit drilling include Idonearca vulgaris (Morton, 1830); Anomia perlineata Wade, 1926; Paranomia scabra (Morton, 1834); Camptonectes burlingtonensis Gabb, 1860; Cyprimeria alta Conrad, 1875; Exogyra cancellata

Stephenson, 1914; Ostrea falcata Morton, 1829; Astarte? sp.; Pterotrigonia

(Scabrotrigonia) thoracica (Morton, 1834); Aphrodina cretacea (Conrad, 1870); and

Veniella conradi (Morton, 1833). Not all drill holes are successful; several failed drill holes were noted (Fig. 6.1).

An uncatalogued Aphrodina cretacea (Fig. 6.9) from the Sohl collection in the

Smithsonian, proved to show the most interesting of all gastropod drill holes. One valve had two drill holes, both of which were successful and are only 0.38 cm apart. This is interesting because the amount of time and energy needed for a gastropod to drill through a bivalve shell can be up to 36 hours (Kitchell et al., 1981). Multiple successful drill holes may be caused by interruption of drilling after penetration but before ingestion could take place or by taphonomic alteration of an incomplete hole (Kelley et al., 2001).

The size range of the bivalves drilled is highly variable. The small Astarte? sp. is approximately 0.38 cm from anterior to posterior margins and has a drill hole that is approximately 0.07 cm in diameter. The largest drill holes are found on specimens of

Cyprimeria alta and Exyogra cancellata. Cyprimeria alta (Figs. 4.5-4.6) is approximately 7.6 cm from the posterior to anterior margins with the drill hole that is 0.5 cm in diameter. Although the E. cancellata (Fig. 5.7) valve is small (about 5 cm from ventral margin to umbo) in comparison to most of the Exogyra specimens collected, the

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shell is still thick and the gastropod was successful, with a drill hole that is 0.8 cm in diameter.

Using all the specimens that were collected on site, the numbers in Table 2 show a relatively even distribution in valve preference (51.7% on the left valve, 44.8% on the right valve); chi-square analysis (Reyment, 1971) shows no preference (0.14). Success rate of the gastropod boring was 79.3%.

Crassatella vadosus was by far the most common species of bivalve that was collected at Coon Creek. More than half (127) of the total specimens collected were of

C. vadosus, and there were 131 specimens in the Cleveland Museum of Natural History.

The number of specimens allows for determination of frequency of gastropod drill holes and observation of the valve on which they occur. Using this count (Table 3), it was hoped that valve preference would be discerned. Of the total 258 valves that were studied, 45 had gastropod drill holes. Drilling on the left valve, whether successful or failed, accounted for 60% of the C. vadosus drilled. Running the chi-square analysis

(Reyment, 1971) the results (1.8) show that there was no preference to which valve C. vadosus lay on.

Two distinct shapes of the drill holes are seen on the bivalves: 1) a wider, beveled drill hole, and 2) a narrower, straight drill hole. The beveled drill holes are generally associated with naticid gastropods and straight drill holes associated with muricid gastropods (Kelley and Hansen, 1996). It appears that naticid holes varied in size as the bivalves drilled into varied, with larger drill holes on larger valves. Muricid holes

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TABLE 3—Frequency and location of drilling found on Crassatella vadosus. Fossil Undrilled Drilled Drilled Failed Failed Total Repository Right Left Right Left Valves: Field 96 9 7 1 5 127 Collection CMNH 108 8 14 0 1 131 Percentage 79% 6.6% 8.1% .39% 2.3% 100%

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remained relatively the same size and were only found on Crassatella vadosus specimens.

Location of the drill holes are varied on the bivalves. The most common location is in the relative center of the bivalve (Figs. 6.1-6.4, 6.7-6.9, Appendix 1). Of the bivalves with a central drill hole, if there is an offset, it usually lies closer to the dorsal margin. Muricid holes occur almost exclusively next to the umbo. Kelley (1988) discussed how there appears to be a highly selective process to finding drilling sites, though she pointed out that there appears to be greater variability of position in the

Ripley Formation. In the bivalves Kelley studied, boreholes most commonly occurred in the central or umbonal region of the valves. The least common location of drill holes can be found on the ventral margin (Fig. 6.5-6.6).

The location of the bore site directly affects the drilling time (Kitchell et al.,

1981). The thickness of the shell varies over the bivalve, causing the standardized siting behavior to cut down on the time needed to drill. Standardization of drill sites may have increased over time (Kelley, 1988). Kitchell et al. (1981) noted that varying drill sites may be in response to prey being either too large or too small for the gastropod’s preferred prey size. These ideas may account for drill sites along the ventral margin.

Either the prey was the wrong size for the gastropod, causing it to drill out of its normal range or the valve is not as thick along the ventral margin, cutting back on the time needed to drill.

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Figure 7: All scale bars are equal to 1 cm unless otherwise noted. 1 – Exogyra cancellata, left valve – dorsal attachment scar; 2 – Ostrea tecticosta, left valve – growth lines of another bivalve can be discerned on the external view; 3 – Pycnodonte vesicularis, left valve – attachment scar; 4 – Crassatella vadosus, right valve – oyster attached; 5 – Exogyra costata, left valve – part of shell that it attached to is present, and a drill hole can be seen on anterior side;6 – Paranomia scabra, left valve – oyster attached; 7 – Paranomia scabra, left valve – cluster of three oysters attached; 8 - Idonearca vulgaris, left valve – cluster of nine oysters attached on internal surface.

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Shell attachment.— Shell attachment is separated into two different groups: 1) attachment scar seen on the oysters, and 2) oysters attached to another bivalve. The attached bivalve may either still be attached or its attachment scar is visible and identifiable as such. One unique interaction between two bivalves that does not fit either category is discussed below.

Attachment scars on oysters.— Some of the oysters (6.5% of total interactions) possess attachment scars indicating their initial attachment to another living organism (usually another bivalve). One in particular, Ostrea tecticosta Gabb, 1860 (Fig. 7.2), preserves the growth lines of another bivalve quite clearly. Other examples include Pycnodonte vesicularis (Lamarck, 1806) (Fig. 7.3) which shows a dorsal attachment, and Exogyra cancellata (Fig. 7.1), which also shows a dorsal attachment.

Most of the attachment scars found on the oysters (Table 2) are on the left valve

(77.8%). This number is not surprising the majority of the oysters found were Exogyra.

The attachment scar Exogyra is found on the umbonal region of the left valve, which is also the most readily preserved valve of the oyster. The number of left valve attachment scars is skewed because so many more Exogyra valves were found.

Oyster attached/scar of oyster attached.— In 7.2% of total interactions the oysters are still attached to another bivalve. Based on the fossils in this study, Crassatella vadosus and Paranomia scabra are the most common attachments for oysters (Figs. 7.4, 7.6) with most of them being a single oyster attached to a single bivalve. However, one P. scabra

(Fig. 7.7) has a cluster of three oysters attached. Another example of oyster attachment

28

Figure 8: All scale bars are equal to 1 cm unless otherwise noted. 1, 2 – Idonearca vulgaris, right valve – Anomia argentaria shell attached to the anterior side of I. vulgaris. 1 – shows where A. argentaria laid, perfectly formed to the shape of the large shell, 2 – close up of I. vulgaris with A. argentaria removed, showing the growth lines of the external side of the other valve, other valve was not attached. Arrow shows the beak impression of A. argentaria; 3, 4, 5 – Crassatella vadosus, right valve with seven nodosariacean foraminifera attached to the internal side, two of which are illustrated, 3 – full view of specimen with arrows indicating position of two foraminifera illustrated are, 4 – one example of the foraminiferan found, scale bar equals 0.1 cm, 5 – another example of the foraminiferan found, scale bar equals 0.1 cm.

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can be seen in Idonearca vulgaris, (Fig. 7.8) which has a large cluster of 9 oysters that attached to the internal part of the shell.

More rare, but still found, are scars from an oyster seen on another oyster or bivalve. The best example is on one Exogyra cancellata (Fig. 4.1) where parts of another attached oyster can be seen.

Of the 10 specimens (Table 2) collected that show bivalve attachment, 60% are found on the left valve, 30% on the right valve, and one specimen of Crassatella vadosus shows evidence of shell attachment on both valves. Internal and external attachment of bivalves is evenly distributed at 50% each.

Other bivalves.— A strange occurrence of two bivalves interacting, and perhaps attaching, can be seen between Idonearca vulgaris (Morton, 1830) and Anomia argentaria Morton, 1833 (Figs. 8.1-8.2). The I. vulgaris was excavated from the spoil pile at the Dave Weeks site and was infilled with sediment. When the sediment was removed, the A. argentaria was found pressed inside the anterior side. A. argentaria had conformed to the shape of the I. vulgaris, which could have been caused either by being molded by the sediment or by its original growth position. When the A. argentaria was removed from I. vulgaris it was noted that A. argentaria was a single left valve, which would support the hypothesis that it was washed in after death and conformed by pressure of sediment. However, the inside of the I. vulgaris supported the hypothesis that it originally grew there, as the impression left by the A. argentaria very clearly shows the beak area and growth lines, neither of which are found on the interior of bivalves. The

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Figure 9: All scale bars are equal to 1 cm unless otherwise noted. 1, 2 – Inoceramus sp. – impression of bryozoans near dorsal end. 1 – full view of specimen (arrow indicating location of close-up), 2 – close up on the bryozoan impression, scale bar equals 0.5 cm; 3, 4 – Exogyra costata, left valve – bryozoan still attached near umbo (arrow). 3 – full view of specimen, 4 – close up showing where bryozoan attached; 5 – Veniella conradi, left valve – posterior, external view showing bryozoans still attached (arrow); 6, 7, 8 – Crassatella vadosus, left valve – Iramena trace fossil found near the teeth (arrow). 6 – full view of specimen, 7 – close up showing trace fossil, scale bar equals 0.3 cm, 8 – closer view of trace fossil, scale bar equals 0.1 cm

31

impression found on the I. vulgaris would match the external side of the right valve of A. argentaria, but that valve no longer exists. The lack of the right valve of the Anomia argentaria is curious as the impression left on the internal surface of the Idonearca vulgaris could not have been made by the internal surface of the valve found. It is unclear what could cause this particular association, but the impression indicates that another bivalve (most likely A. argentaria) grew on the internal surface after the I. vulgaris had died.

Other interactions.— Other occurrences observed are less common than those described above; combined they make up 15.1% of the total specimens collected and 25.9% of all interactions. These are interactions with worms, bryozoans, barnacles, fungi, and others.

Bryozoans.— Interactions with bryozoans (4.3% of total interactions) can be seen as impressions, trace fossil borings, and attachments. Impressions are only seen on smoother, mother-of-pearl surfaces, found either on the internal surface of the valve, or on inoceramid shells (Figs. 9.1-9.2). The trace fossil Iramena Boekshoten, 1970, is described as an irregular network of long stolons with round reniform apertures in alternating positions closely adjacent to, the tunnels (Boekshoten, 1970). The tunnels have been observed on the internal side of two Crassatella vadosus fossils (Figs. 9.6-9.8).

Instances of bryozoans still attached are seen on the posterior side of a V. conradi valve

(Fig. 9.5) and in the dorsal area of an Exogyra costata shell (Figs. 9.3-9.4).

Fungal Borings.— Mycelites Roux, 1887 is described as irregularly branching tunnels about 2 to 6 microns wide. There is some debate as to whether the borings are from fungi or algae (Bromley, 1970). It has been most recently studied in fish teeth and has been

32

Figure 10: All scale bars are equal to 1 cm unless otherwise noted. 1 – Idonearca vulgaris, right valve – protective coating along the posterior muscle scar (arrow) indicating presence of an irritant; 2 – Crassatella vadosus, right valve – ventral margin chipped as if by a decapod; 3 – Veniella conradi, right valve – Mycelites traces along the posterior, exterior side; 4 – Inoceramus sagensis, right valve – Zapfella traces; 5 – Exogyra cancellata, left valve – Scaphopod attached at beak, beneath a part of an old attachment of another oyster; 6 – Idonearca vulgaris, right valve - Dakoticancer overana claw cemented to the external ventral margin; 7 – Exogyra costata, left valve – four worm tubes, two larger ones around the dorsal wall (Serpula sp. and Hamulus squamosus) and two smaller ones inside the muscle scar (both serpulids).

33

described as a fungus (Martill, 1989; Underwood et al., 1999; Underwood and Mitchell,

2004) but is also known in shells of invertebrates (Häntzschel, 1975). In the fossils studied at Coon Creek, Mycelites comprises 7.9% of total interactions. Eleven specimens show some trace of Mycelites, with the best example being found on the posterior side of

Veniella conradi (Fig. 10.3).

Barnacles.— The trace fossil Zapfella de Saint-Seine, 1956 is created by acrothoracian barnacles (Johnson and Baarli, 1999) and is described as sac-like bore holes 1 to 4 mm long with slit-like openings (Häntzschel, 1975). Zapfella (2.3% of total interactions) is only seen on the exterior surfaces of the bivalves collected. They are seen most clearly on Cardium stantoni (Wade, 1926) (Fig. 4.3) and on the dorsal side of Inoceramus sagensis (Fig. 10.4).

Worms.— There are only a few examples of traces attributable to worms, one being found on the external dorsal area of Cardium stantoni (Fig. 4.4) and others on the internal dorsal and central areas of Exogyra costata (Fig. 10.7). The worms found on C. stantoni are small tubes that curl around themselves. There are four worms found on E. costata.

Two are relatively large (compared to the curled ones) and produced tubes along the inner wall of the oyster. Of these two worms, one is smooth (a serpulid), while the other is keeled (Hamulus squamoses Gabb, 1859). The two found in the muscle scar of the oyster are small and similar to the ones found on C. stantoni.

Miscellany.— A few of the fossils show unusual interactions with other, once living, things. A most unusual case is found on Idonearca vulgaris. On the I. vulgaris, a crab claw, from Dakoticancer overana Rathbun, 1917 (Fig. 10.6), has been cemented to the

34

ventral margin of the exterior. This association undoubtedly happened after the death of the bivalve (and most certainly the arthropod), due to diagenetic cementation occurring during diagenesis.

An unusual interaction found on one Crassatella vadosus involved seven different nodesariacean foraminifera, two of which have been illustrated (Figs. 8.3-8.5) cemented to the internal side of the bivalve (Loeblich and Tappan, 1964). One Exogyra cancellata has a scaphopod attached to the dorsal side of the shell (Fig. 10.5). The scaphopod is beneath what appears to be an old attachment of another oyster. Two specimens of C. vadosus appear to have breaks along the ventral margin that could be claw marks from an arthropod (Fig. 10.2).

CHAPTER 6: Importance of Location

Location of the interactions is very important when determining how and when the interacting takes place. In 1975, Feldmann and Palubniak developed a system to categorize and describe biotic interactions associated with oysters. This system is used to determine associations in transported death assemblages to attempt to reconstruct the fossil oyster community. Coon Creek is an in situ assemblage; therefore, all the organisms are associated with each other, but the concepts explored by Feldmann and

Palubniak can be helpful in determining when interactions occurred. This paper uses their basic concepts, with some modifications, to determine when interactions took place.

First order.— First order interactions are living organisms interacting with living bivalves. Evidence for the interactions is found, for example, when hollow, calcareous blisters are formed inside shells to protect the organisms from outside irritants. The best example of this in the Coon Creek assemblage can be seen in an Idonearca vulgaris that has this internal protection around the posterior adductor muscle scar (Fig. 10.1).

Second order.— Second order associations are evidence that one or both of the organisms involved were alive during the time of interaction. While second order associations are less definitive, they are no less important.

External surface only interactions.— When interactions occur only on the external surface of a bivalve, it can not be stated with certainty that the bivalve was alive during

35 36

the time of interaction. The exception to this can be seen in the Exogyra oysters; this will be described later.

Interactions that involve the clionid sponges that occur only on the external surface could easily occur on a living organism, because modern Cliona bore living bivalves today (Krinsley and Schneck, 1964). Pycnodonte vesicularis (Fig. 7.3) is a good example of how the borings occur only on the external surface and not on the interior.

The external surface of the left valve shows clionid borings and an attachment scar, whereas the external surface of the right valve shows clionid borings as well as possible burrowing bivalve borings. The internal surfaces of both valves are clean, which is not too surprising since they are conjoined. It is unlikely that both surfaces could have been used as substrates while P. vesicularis was living; thus, it is hypothesized that this specimen was dead when colonized.

When the valves of a bivalve are still articulated, there is a greater chance that the interactions occurred during life since bivalves usually disarticulate after death. An example of an articulate bivalve with borings is Veniella conradi (Figs. 3.7-3.8). The borings probably were initiated when the V. conradi was living, but whether or not they continued after death is not known since the valves are still conjoined.

Exogyra oysters can be an exception to the ambiguity of when the interactions occurred when they are expressed only on the external surface of the valve. Some

Exogyra grow attached to another shell (the attachment scar seen around the beak), but when they grow larger and heavier than the shell to which they are attached they may break from it and become free-living (Stenzel, 1971). When oysters lie on the sediment,

37

the central portion of the left valve would be on the sea floor. If this area has evidence of borings from sponges or smaller bivalves (Fig. 9.3), it is most likely that the oyster was dead and was flipped over by currents. The epibionts would not be able to breathe and feed if they were in the sediment. When the borings are on the right valve or along the margins of the left valve, then it is similar to other bivalves with only external borings in that they could have been either alive or dead at the time of the borings.

Oysters and the Anomiidae both show xenomorphism, where the outside surface of a valve closely imitates the surface of the substrate (Stenzel, 1971). When the substrate is another living organism, the resulting xenomorphic structure of the attaching oyster will mimic the shape of that living organism. The oysters more readily show the biotic substrates than do the Anomiidae, and these can be seen in several specimens of different species (Figs. 7.1-7.3).

Another interaction involving oysters is that bivalves are sometimes seen with oysters still attached to them (Figs. 7.4, 7.6-7.7), and occasionally the oyster is retained as a part of the original substrate (Fig. 7.5). In all these situations, the oyster was alive while it attached and grew, even if the bivalve to which it was attached was not.

Internal shell only interactions.— Bivalves will protect themselves from irritations from outside sources if the irritant manages to find its way inside the shell. If the bivalve was still alive when an outside irritant began to burrow inside the valve, the bivalve would secrete a calcareous protection against it, and an interaction of the first order would occur. Therefore, it is known that any form of interaction that occurs on the internal surface of a bivalve to which the organism does not respond happens after death.

38

Several interactions have been observed on the internal surface of the bivalves.

Most common are clionid sponge borings (Figs. 3.1, 4.2, 4.6). Burrowing bivalves are found less frequently internally, but they can be seen, either with the burrows started from the internal surface (Figs. 5.5-5.6) or possibly burrows that penetrated through the entire shell thickness (Figs. 5.1-5.2). Bryozoans are found either as impressions or in the form of the trace fossil Iramena (Figs. 9.6-9.8). Oysters used the inside of I. vulgaris

(Fig. 7.8) as a substrate and worms used E. costata (Pl. 6, fig. 7). One specimen of

Crassatella vadosus (Figs. 8.3-8.5) shows seven nodosariacean foraminifera that used the internal surface of the bivalve as a substrate.

Third order.— Feldmann and Palubniak (1975) describe third order associations as the animals are found in the same death assemblage. Since Coon Creek is an in situ formation, third order associations will be defined here as being associations that take place after both organisms are deceased. This would be most evident in larger bivalves that are infilled with sediment and other fossils. As third order associations occur after death, they are not included as biotic interactions in this paper.

Fourth order.— Fourth order associations are strong evidence that the organism found was not a part of the death assemblage. This occurs as wood and leaves in the Coon

Creek Formation (Safford, 1896; Wade, 1917).

Location in life.— In addition to indicating when the interactions took place, the location of the interactions can tell upon which valve the bivalve lived. This information is best utilized when the valves are articulated.

39

With reference to the articulated specimen of Veniella conradi (Figs. 3.7-3.8), clionid sponge borings occur only on the left valve; the right valve is completely clean. It can be determined that this particular clam lived with the right valve in the sediment, and the left valve was exposed to the water column. This is known because the sponge is a filter feeder (Prothero, 1998). Clionid sponges need clean water to filter, therefore if they had bored into the valve that was in the sediment, the sediment would essentially choke the sponge.

One specimen of Idonearca vulgaris (Fig. 3.5) shows a similar pattern as the

Veniella specimen. The right valve is riddled with clionid sponge borings, whereas the left valve only has borings along the ventral margin. This indicates that the bivalve lived with the left valve mostly, but not completely, in the sediment and the right valve entirely in the water column. The presence of borings on the margin of the left valve indicates that the valve was far enough out of the sediment for the sponges to exist here without problems.

It may not always be necessary that both valves be present for living position to be determined. One specimen of Veniella shows clionid sponge borings only on the ventral margin (Fig. 3.6). The location of the borings could mean that the right valve rested in the sediment with the left valve in the water column. With the borings on the margin, the right valve would have been tilted in such a way that the margin was exposed enough for the sponges. However, nothing definite can be determined without examining the left valve.

40

Gastropod drill holes could very well indicate which valve rested in the sediment and which was exposed. It is very likely that the valve with the drill hole was exposed to the water column. This hypothesis is not suppportable with the Exogyra oysters (Figs.

6.7, 7.5) that have been drilled, as the left valve is drilled and it sits on the substrate, but the drills are close to the ventral margin.

CHAPTER 7: Conclusions

The interactions found on the bivalves show that there is a wide range of organisms that rely on the shells to survive. Whether it is using the shell as a substrate to grow (clionid sponges and boring bivalves) or drilling through to find a meal

(gastropods), the shell retains a record of what was happening on the sea floor during the

Cretaceous.

It is important to look at where the interactions take place to learn what the bivalve was doing when the interactions occur. Second order interactions are the most commonly found, and sometimes it is not easy to determine whether the bivalve was alive or not during the time of the interaction. Internal interactions are the only sure sign that the interactions took place after the death of the bivalve. External interactions could have happened during life or after, depending on how the organism died and how long it remained on the seafloor prior to burial. Gastropod drill holes almost assuredly occur during the life of the bivalve.

Future work with biotic interactions found in the Coon Creek Formation will need to include a more diverse range of species. The fossils studied were ones most commonly found on site, but those are quite few compared to the 114 species that Wade described (1926). Only a handful of infaunal bivalves were studied, with none showing

41 42

any form of biotic interactions. It would be interesting to see if interactions are seen on infaunal bivalves if a greater number are collected.

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Underwood, C. J., S. F. Mitchell, and C. J. Veltkamp. 1999. Microborings in mid-

Cretaceous fish teeth. Proceedings of the Yorkshire Geological Society, 52:269-274.

Vermeij, G. J. and E.C. Dudley. 1982. Shell Repair and Drilling in Some Gastropods from the Ripley Formation (Upper Cretaceous) of the South-eastern U.S.A. Cretaceous

Research, 3:397-403.

Wade, B. 1917. A Remarkable Upper Cretaceous Fauna from Tennessee. Johns Hopkins

University Circular, 73-101.

Wade, B. 1926. The Fauna of the Ripley Formation on Coon Creek, Tennessee. U.S.

Geological Survey Professional Paper 137: 272 p.

53

Whetstone, K. N. 1977. A plesiosaur from the Coon Creek Formation (Cretaceous) of

Tennessee. Journal of Paleontology, 51:424-425.

APPENDIX 1: Collected Fossils

x* – Indicates more abundant areas of interactions xL, xV, ect. – Indicates that interactions are found only at these areas L – Left valve R – Right valve B – Both valves I – Internal E – External V - Ventral

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Anomidae Anomia CC12 argentaria Entobia x x x

Anomia CC13 argentaria

Anomia CC119 argentaria

Anomia CC228 argentaria

Anomia CC76 argentaria?

Anomia CC77 argentaria?

Anomia CC5 perlineata Naticid drill x x x x

Anomia CC7 perlineata

54 55

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Astartidae CC231 Astarte? sp. Naticid drill x x x

Corbulidae Corbula CC85 crassiplica

Corbula CC86 crassiplica Crassatellidae Crassatella CC16 vadosus

Crassatella CC17 vadosus

Crassatella CC18 vadosus

Crassatella CC19 vadosus Entobia x x x

Crassatella CC20 vadosus Entobia x x x x x x x x Mycelites x x x Bivalve borings (?) x x x x x x

Crassatella CC21 vadosus Naticid drill x x x x x Entobia (mostly around drillhole) x x* x x x x*

Crassatella CC22 vadosus Entobia x x x* x x Bivale boring (?) x x x

56

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC23 vadosus Bryozoan x x x Bivalve attachment scar (?) x x x

Crassatella CC24 vadosus

Crassatella CC25 vadosus Claw mark (?) x x x

Crassatella CC26 vadosus

Crassatella CC27 vadosus

Crassatella CC28 vadosus

Crassatella CC29 vadosus

Crassatella CC30 vadosus Gastropod drill hole (?) x x x x

Crassatella CC31 vadosus

Crassatella CC32 vadosus Entobia x x x x x x

Crassatella CC33 vadosus Entobia x x x Mycelites x x x

57

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC34 vadosus Anomia shell x x

Crassatella CC35 vadosus

Crassatella CC36 vadosus

Crassatella CC37 vadosus Naticid drill x x x x x

Crassatella CC38 vadosus

Crassatella CC39 vadosus

Crassatella CC40 vadosus Hole x x x x

Crassatella CC41 vadosus

Crassatella CC42 vadosus Entobia x xV x x x x x x Naticid drill x x x x x Bivalve boring? x x x

Crassatella CC43 vadosus

Crassatella CC44 vadosus

Crassatella CC45 vadosus

Crassatella CC46 vadosus

58

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC47 vadosus

Crassatella CC48 vadosus Gastrpod drill (?) x x x

Crassatella CC49 vadosus Entobia x x x x x x Bivalve boring (?) x x x

Crassatella CC50 vadosus Entobia x x x* x x x x x Bivalve boring x x x

Crassatella CC51 vadosus Gastropod drill (failed) x x

Crassatella CC52 vadosus

Crassatella CC53 vadosus Claw marks? x x

Crassatella CC54 vadosus Entobia x x x x x x x

Crassatella CC55 vadosus

Crassatella CC56 vadosus

Crassatella CC57 vadosus

Crassatella CC58 vadosus

59

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC59 vadosus Bivalve boring (?) x x x x Gastropod drill (failed) x x x x Entobia (?) x x x

Crassatella CC60 vadosus Entobia x x x

Crassatella CC61 vadosus Entobia x x x Hole x x x

Crassatella CC62 vadosus Entobia x x x x x x x

Crassatella CC63 vadosus Entobia x x x x x x x Mycelites x x x Bivalve boring x x x x

Crassatella CC64 vadosus Entobia x x x* x x x x xE Bivalve boring x x x Nodosariidae Foram (7) x x x x x

Crassatella CC66 vadosus

Crassatella CC92 vadosus Naticid drill x x x x

Crassatella CC143 vadosus Gastropod drill (failed) x x

60

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC144 vadosus Entobia x xV x* x x x x x x Mycelites x xV x Iramena x xV x

Crassatella CC145 vadosus

Crassatella CC146 vadosus Muricid drill x x x x

Crassatella CC147 vadosus Entobia x x x x x x x x

Crassatella CC148 vadosus

Crassatella CC149 vadosus

Crassatella CC150 vadosus Entobia x x x x Bivalve boring (?) x x x x

Crassatella CC151 vadosus

Crassatella CC152 vadosus Muricid drill x x x x Entobia x x x x x x

Crassatella CC153 vadosus Entobia x x x x x x

Crassatella CC154 vadosus Entobia x x x x x x x x

61

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC155 vadosus Bivalve boring (?) x x x

Crassatella CC156 vadosus Entobia x x x x x Bivalve boring (?) x x x x

Crassatella CC157 vadosus

Crassatella CC158 vadosus

Crassatella CC159 vadosus Entobia x x Bivalve boring x x x x

Crassatella CC160 vadosus Entobia x x* x x x x x Bivavle boring (?) x x x x Attachment of oyster scar x x x x

Crassatella CC161 vadosus

Crassatella CC162 vadosus

Crassatella CC163 vadosus Entobia x x x x x*

Crassatella CC164 vadosus Naticid drill x x x x

Crassatella CC165 vadosus

62

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC166 vadosus Entobia x x x x x x

Crassatella CC167 vadosus Bivalve boring (?) x x x

Crassatella CC168 vadosus Entobia x x x x x x

Crassatella CC169 vadosus Bryozoan x x x

Crassatella CC170 vadosus Muricid drill x x x x Entobia x x x* x* x

Crassatella CC171 vadosus Bivalve boring x x x x x

Crassatella CC172 vadosus Gastropod drill (failed) x x x

Crassatella CC173 vadosus Gastropod drill (failed) x x x

Crassatella CC174 vadosus Bivalve boring x x x x x x

Crassatella CC175 vadosus Entobia x x x x Bivalve boring x x x x

63

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC176 vadosus Muricid drill x x x x Entobia x x x* x x x x

Crassatella CC177 vadosus Muricid drill x x x x Entobia x x x* x x x x

Crassatella CC178 vadosus

Crassatella CC179 vadosus Bivalve boring (?) x x x

Crassatella CC180 vadosus

Crassatella CC181 vadosus

Crassatella CC182 vadosus

Crassatella CC183 vadosus Muricid drill x x x x

Crassatella CC184 vadosus Entobia x x x x x x x

Crassatella CC185 vadosus

Crassatella CC186 vadosus Bivalve boring Mycelites Entobia (sparse)

64

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC187 vadosus Muricid drill x x x x Entobia x x x x x Bivalve boring x x x

Crassatella CC188 vadosus Entobia x x x x x x x x Bivalve boring x x x* x Iramena x x x

Crassatella CC189 vadosus Entobia x x x x

Crassatella CC190 vadosus

Crassatella CC191 vadosus Entobia x x* x x* x x x x

Crassatella CC192 vadosus

Crassatella CC193 vadosus Entobia (sparse) x x x x Gastropod drill (failed) x x x

Crassatella CC194 vadosus

Crassatella CC195 vadosus Bivalve boring x x x x Entobia x x x

Crassatella CC196 vadosus

65

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC197 vadosus Entobia x x x x x x x x

Crassatella CC198 vadosus

Crassatella CC199 vadosus Gastropod drill x x x x Bivalve boring (?) x x x

Crassatella CC200 vadosus

Crassatella CC201 vadosus

Crassatella CC205 vadosus Entobia x x x

Crassatella CC206 vadosus Entobia (sparse) x x x x x

Crassatella CC207 vadosus Entobia x x x x

Crassatella CC208 vadosus Muricid drill x x x x

Crassatella CC209 vadosus Entobia x x x x x x x Bivalve boring x x x x x x x x

Crassatella CC210 vadosus Entobia (sparse) x x x x x

Crassatella CC225 vadosus

66

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Crassatella CC226 vadosus Bivalve boring x x x

Crassatella CC227 vadosus

Crassatella vadosus CC232 fragments Veneridae CC94 Cyprimeria alta

CC204 Cyprimeria alta? Entobia x x x x

Gryphaeidae Exogyra CC8 cancellata Entobia x x x

Exogyra CC9 cancellata Entobia x x x xL xL xL xL Attachment scar x x x Bivalve boring (?) x x

Exogyra CC138 cancellata Attachment scar x x x

Exogyra CC139 cancellata Ostrea sp. attached x x x

Exogyra CC140 cancellata Entobia x x x x Gastropod drill x x x x x Scaphopod x x x Attachment of oyster x x x

67

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Exogyra CC141 cancellata

Exogyra CC69 cancellata? Entobia x x x x

CC4 Exogyra costata Attachment scar x x x Entobia x x x x Attachment of oyster x x x x

CC72 Exogyra costata Entobia x x x x x x x x Bivalve boring x x x x x Ostrea sp. attached x x x

CC82 Exogyra costata Entobia x x x x x* x x Attachment scar x x x Large hole x x x

CC130 Exogyra costata Entobia x x x x Bivalve boring x x x x x

CC131 Exogyra costata Bivalve boring (?) x x x x x x x x Entobia x x

CC132 Exogyra costata Bivalve boring (?) x x x

CC133 Exogyra costata Mycelites x x x Attachment scar x x x Entobia x x x

CC134 Exogyra costata Entobia x x x x x x Bivalve boring (?) x x x x Bryozoam x x x

68

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve CC135 Exogyra costata Bivalve boring x x x x x x Entobia x x x x

CC136 Exogyra costata Attachment scar x x x

CC137 Exogyra costata Bivalve boring x x x x x Entobia x x x x x x

CC78 Exogyra?

Bakevelliidae Gervilliopsis CC203 ensiformis Entobia (?) x x

Cucullaeidae Idonearca CC1 vulgaris Shell impression x x x Entobia x x x* x x x* x x

Idonearca CC10 vulgaris

Idonearca CC79 vulgaris Gastropod drill (?) x x x x

Idonearca CC80 vulgaris Bivalve borings (?) x x x

Idonearca CC81 vulgaris Internal protection x x x

Idonearca CC88 vulgaris Entobia x x* x xI x* x x x

69

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Idonearca CC89 vulgaris

Idonearca CC90 vulgaris

Idonearca CC107 vulgaris

Idonearca CC108 vulgaris Borings (?) x x x

Idonearca CC109 vulgaris Entobia x x x

Idonearca CC110 vulgaris Entobia x x x* x x

Idonearca CC111 vulgaris Gastropod drill (?) x x x x

Idonearca CC112 vulgaris Naticid drill x x x x

Idonearca CC213 vulgaris Bivalve boring x x x x x x x Entobia x x x x x x x x

Inoceramidae Inoceramus CC224 proximus

Inoceramus CC229 sagensis

Inoceramus CC230 sagensis Entobia x

70

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Inoceramus CC75 sp.?

Inoceramus CC84 sp.? Bryozoan x x

Inoceramus CC96 sp.? x x Zapfella

Inoceramus CC97 sp.? Entobia x x Zapfella x x

Inoceramus CC99 sp.?

Cultellidae Leptosolen CC202 biplicata

Pectinidae Neithea (Neitheopsis) quinquecostatu CC65 s

Neithea (Neitheopsis) quinquecostatu CC67 s

Neithea (Neitheopsis) quinquecostatu CC142 s

Nuculidae Nucula CC123 percrassa

71

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Nucula CC124 percrassa Entobia (?) x x x

Ostreidae CC91 Ostrea falcata

CC121 Ostrea falcata

CC122 Ostrea falcata

Ostrea falcatafragment CC233 s

CC120 Ostrea sp.

CC212 Ostrea sp.

Ostrea CC71 tecticosta Attachment scar x x

Paranomia CC2 scabra Attachment scar x x x Entobia x x x x x x

Paranomia CC3 scabra Entobia x x x x* x x x

Paranomia CC6 scabra

Paranomia CC73 scabra Entobia x x x x* x x x x

Paranomia CC95 scabra

Paranomia CC113 scabra

72

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Paranomia CC114 scabra Naticid drill x x x

Paranomia CC115 scabra

Paranomia CC116 scabra

Paranomia CC117 scabra Borings (?) x x x

Paranomia CC118 scabra Mycelites x x x x x x

Paranomia CC214 scabra Attachment scar x x x

Paranomia CC215 scabra Bivalve boring x x x x x x x x Entobia x x x x x x x

Paranomia CC74 scabra?

Cardiidae Protocardia (Pachycardium) CC70 stantoni Bivalve borings x x x Zapfella x x x Entobia x x x Mycelites x x x

Protocardia (Pachycardium) CC93 stantoni Entobia x x x

73

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Protocardia (Pachycardium) CC211 stantoni

Trigoniidae Pterotrigonia (Scabratrigonia) CC68 thoracica Entobia x x x x

Pterotrigonia (Scabratrigonia) CC100 thoracica

Pterotrigonia (Scabratrigonia) CC101 thoracica

Pterotrigonia (Scabratrigonia) CC102 thoracica

Pterotrigonia (Scabratrigonia) CC103 thoracica

Pterotrigonia (Scabratrigonia) CC104 thoracica Zapfella x x x Entobia x x x

Pterotrigonia (Scabratrigonia) CC105 thoracica

Pterotrigonia (Scabratrigonia) CC106 thoracica

Pterotrigonia (Scabratrigonia) CC217 thoracica Entobia x x x x*

74

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Pterotrigonia (Scabratrigonia) CC218 thoracica

Pterotrigonia (Scabratrigonia) CC219 thoracica

Pterotrigonia (Scabratrigonia) CC220 thoracica Entobia x x x x

Pterotrigonia (Scabratrigonia) CC221 thoracica

Pterotrigonia (Scabratrigonia) CC222 thoracica Entobia x x x

Pterotrigonia (Scabratrigonia) CC223 thoracica

Arcticidae CC11 Veniella conradi Entobia x x x x

CC87 Veniella conradi Entobia x x x Bivalve boring (?) x x x

CC125 Veniella conradi Mycelites x x x x Entobia x x x

CC126 Veniella conradi

CC127 Veniella conradi

CC128 Veniella conradi

CC129 Veniella conradi

75

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve CC216 Veniella conradi

Unknown (umbo CC98 only) Entobia x x

APPENDIX 2: Pink Palace Fossils

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Anomiidae KSU- Anomia 3079 argentaria Entobia x x x x x x x

Arcidae KSU- ER13 Barbatia fractura Entobia x x x x x x Mycelites x x x Bivalve boring (?) x x x x

Pectinidae KSU- Camptonectes 3080 burlingtonensis? Gastropod drill x x x x x

Crassatellidae KSU- Crassatella 3086 vadosus Attachment of oyster scar x x x Entobia x x x

Veneridae KSU- 3089 Cyprimeria alta Entobia x x x x x x x x Gastropod drill x x x x

76 77

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Gryphaeidae KSU- Exogyra ER23 cancellata Gastropod drill x x x x Entobia x x x x x KSU- 3074 Exogyra costata Bivalve hole x x x x x x x x Attachment of oyster x x x x x x Attachment scar x x x x Entobia x x x x x x x x Bakevelliidae KSU- Gervilliopsis ER20 ensiformis Entobia x x x x x x x

KSU- Gervilliopsis 3073 ensiformis Attachment of oyster scar x x x ?? x x

Cucullaeidae KSU- Idonearca 3078 vulgaris Attachment of oysters (9) x x x

KSU- Idonearca ER11 vulgaris Entobia x x x x x x x Bivalve boring x x x x x Claw x x x

KSU- Idonearca 3068 vulgaris Entobia x* x x xR xR xR x xR Bivalve boring (?) x x x x x x* x

78

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Inoceramidae KSU- Inoceramus ER19 proximus Entobia x x x

KSU- Inoceramus 3071 sagensis Zapfella x x x x

Isognomonidae KSU- Isognomon ER66 periridescens Shells attached (same and Anomia) x x

KSU- Isognomon ER65 periridescens ?? x x x

Veneridae KSU- Legumen ER63 planulatum

Cutellidae KSU- Leptosolen ER67 bipliticata

Parallelodontidae KSU- Nemodon 3070 grandis Entobia x x x large hole x x

Anomiidae KSU- Paranomia 3082 scabra Gastropod drill x x x x Entobia? x x x x x x

79

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve KSU- Paranomia 3081 scabra A Entobia x x x x x B Entobia x x x x Attachment of oyster scar x x

KSU- Paranomia ER35 scabra Attachment of oyster x x x x Entobia x

Pholodomyidae KSU- Pholadomya ER64 occidentalis Large hole x x

Cardiidae Protocardia KSU- (Pachycardium) 3088 stantoni Entobia x x x x x x x worms x x x x Attachment of oyster scar x x x x

Trigoniidea Pterotrigonia KSU- (Scabratriogonia) ER29 thoracica Entobia x x x x x x x

Gryphaeidae KSU- Pycnodonte 3076 vesicularis Entobia x x x x x x x x Attachment scar x x x Bivalve boring (?) x x x x x x

80

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Arcticidae KSU- 3084 Veniella conradi Entobia x x x x x x

KSU- ER45 Veniella conradi Bivalve boring x x xL x x x Entobia x x x x x

APPENDIX 3: Cleveland Museum of Natural History Fossils

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve

Arcidae 13865 Arca securculata Entobia x x x x x x x x

Pecten Camptonectes 13790 burlingtonensis? Attachment of oyster scar x x

Gryphaeidae Exogyra 13844 cancellata A Bivalve boring x x x Entobia x x x x x x x Mycelites x x x x x x B Entobia x x x x x x x x Attachment scar x x x

Exogyra 13845 cancellata A Attachment scar x x x x Entobia x x B Attachment scar x x x Entobia x x x x x x x

13758 Exogyra costata Attachment scar x x x Entobia x x xL x x x x x x* Attachment of oyster scar x x x

81 82

ID ID Dorsal Family Family Ventral Ventral Central Internal Internal Species Anterior External External Posterior Left Valve Valve Left Right Valve Ostreidae 13779 Ostrea falcata Gastropod drill (failed) x x x

Glycymerididae Glycimeris 13813 lacertosa Entobia x x x x x x

Glycimeris 13817 microsulci Gastropod drill x x x

Glycimeris 13804 subcrenata A Entobia x x x x x x B Entobia x x x Bivalve boring C (?) x x x x x

Cucullaeidae Idonearca 13748 vulgaris Entobia x x x xL x

Idonearca 13818 vulgaris Entobia x x x x x x x

Nuculidae Nucula 13775 percrassa A Entobia x x x B Entobia x x x x C Entobia x x x

Anomiidae Paranomia 13752 scabra Gastropod drill x x x