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5-2009
Ecological Interactions Across a Plio-Pleistocene Interval of Faunal Turnover: Naticid Cannibalism North and South of Cape Hatteras, North Carolina
Max Christie College of William and Mary
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Recommended Citation Christie, Max, "Ecological Interactions Across a Plio-Pleistocene Interval of Faunal Turnover: Naticid Cannibalism North and South of Cape Hatteras, North Carolina" (2009). Undergraduate Honors Theses. Paper 295. https://scholarworks.wm.edu/honorstheses/295
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Ecological Interactions Across a Plio-Pleistocene Interval of Faunal Turnover: Naticid Cannibalism North and South of Cape Hatteras, North Carolina
A thesis submitted in partial fulfillment of the requirements for Departmental Honors in Interdisciplinary Studies from The College of William and Mary by Max Christie
Accepted for _____Honors______(Honors, High Honors, Highest Honors)
______Dr. Rowan Lockwood, Director
______Dr. Karen M. Layou
______Dr. Patricia H. Kelley
______Dr. Stewart A. Ware
Williamsburg, VA May, 2009
Table of Contents
List of Figures and Tables...... 4
Abstract ...... 6
Introduction ...... 7
Plio-Pleistocene Extinction ...... 8
Taxonomic vs. Ecological Patterns Across Extinctions ...... 9
Naticid Cannibalism ...... 12
Research Questions ...... 15
Geologic Setting...... 15
Methodology ...... 20
Sampling Strategy ...... 20
Field Methods ...... 20
Museum Collections ...... 21
Sample Processing ...... 21
Data Analysis ...... 22
Results ...... 26
Frequency of Cannibalism across the Plio-Pleistocene and Biogeographic Boundaries
...... 26
Frequency of Cannibalism across Plio-Pleistocene Stratigraphic Units ...... 32
2 Frequency of Cannibalism across the Plio-Pleistocene and Biogeographic Boundaries:
Sample Size Standardization Using a Resampling Procedure ...... 35
Changes in Size across the Plio-Pleistocene and Biogeographic Boundaries ...... 37
Effects of Naticid Predator Size on the Frequency of Cannibalism ...... 46
Discussion ...... 48
Conclusions ...... 65
Acknowledgements ...... 67
References ...... 70
Appendix ...... 73
3 List of Figures and Tables
Figure 1: Sample of Naticids ...... 14
Figure 2: Geologic map of the Coastal Plain ...... 14
Figure 3: Correlation Chart ...... 19
TABLE 1: SAMPLES USED IN THIS STUDY ...... 19
Figure 4: Stratigraphic section of Register Quarry, NC ...... 24
Figure 5: Map of field sites and museum collections ...... 24
Figure 6: Diagram of measurements ...... 25
Figure 7: Frequency of cannibalism across the Plio-Pleistocene extinction...... 30
Figure 8: Frequency of cannibalism across Cape Hatteras, North Carolina ...... 30
Figure 9: Frequency of cannibalism among spatiotemporal units ...... 31
Figure 10: Frequency of cannibalism across units north of Cape Hatteras ...... 34
Figure 11: Frequency of cannibalism across units south of Cape Hatteras ...... 34
Figure 12: Frequency of cannibalism among resampled spatiotemporal units ...... 36
Figure 13: Average and maximum size across the Plio-Pleistocene extinction and Cape
Hatteras, North Carolina ...... 40
Figure 14: Average and maximum size among spatiotemporal units ...... 41
Figure 15: Percent drilling among 10mm size bins ...... 42
Figure 16: Outer drillhole diameter vs. prey size ...... 43
Figure 17: Average and maximum size among resampled spatiotemporal units ...... 45
4 Figure 18: Frequency of cannibalism vs. predator size ...... 47
Figure 19: Total and prey item drilling frequency ...... 51
Figure 20: Naticid prey abundance ...... 52
Figure 21: Ratio of predator to prey abundance ...... 52
Figure 22: Frequency of cannibalism across the Plio-Pleistocene extinction in Huntoon
(1999) ...... 54
Figure 23: Naticid to non-naticid predator abundance ...... 54
Figure 24: Abundance and drilling frequency of victims and originators ...... 56
Figure 25: Drilling frequency of victims and originators with cannibalism ...... 56
Figure 26: Drilling frequency of preferred taxa during the Pliocene and Pleistocene ...... 58
Figure 27: Abundance of preferred prey taxa ...... 60
5 Abstract
According to previous work along the western Atlantic Coastal Plain, 70% of molluscan species went extinct during a two pulsed event across the Plio-Pleistocene boundary; yet the nature of this extinction event is different north and south of the biogeographic boundary represented by Cape Hatteras, North Carolina. Although the evolutionary effects of this extinction have been studied, the community-level ecological effects are poorly understood. This research focuses on the confamilial predation of naticid snails and seeks to determine changes in the degree of cannibalism across the biogeographic boundary during the late Pliocene. Float and museum collections of the
Yorktown (Moore House Member) and Duplin Formations (sampling before the extinction) and the lower Waccamaw and Chowan River Formations (sampling after the first pulse of extinction) were used in this study. Frequency of cannibalism per collection, and naticid size were analyzed. Data were collected from 21 localities across the Coastal Plain of Virginia, North Carolina, and South Carolina representing 3 units north of Cape Hatteras, and three units south. 1,951 specimens were analyzed. Results indicate that naticid cannibalism increases across the Plio-Pleistocene extinction and is greater north of the biogeographic boundary. Results also indicate a statistically significant decrease in naticid size across the biogeographic boundary. A statistically significant increase in maximum size occurred south of Cape Hatteras. Additional analysis of community level data from Huntoon (1999) suggests that the observed increase in cannibalism was due to a decrease in preferred prey abundance. These results suggest that naticid competition increased after the extinction event because of reduced preferred prey abundance which lead to an increase in cannibalism.
6 Introduction
Approximately 2 million years ago, the Plio-Pleistocene extinction drastically decreased the diversity of the molluscan fauna in the western Atlantic during a pulsed event. Previous research has suggested that this extinction event may have been due to local cooling as glaciers expanded (Stanley, 1986). Other authors have contended that the Plio-Pleistocene extinction was caused by a decline in primary productivity in the western Atlantic (Allmon, 2001; Todd et al., 2002). Although mollusk diversity has increased in the western Atlantic, many taxa do not inhabit the same geographic range they did before the extinction event. During the Pliocene, sub-tropical species ranged far north of their current habitats, into Virginia and Maryland (Allmon et al., 1993). Yet today, these species are confined south of a biogeographic boundary represented by Cape
Hatteras, North Carolina. This suggests that there may be a fundamental difference in the extinction event north and south of the biogeographic boundary. North of this boundary extinction without origination is thought to have occurred, while south of the boundary, paleocommunities experienced both (Allmon et al., 1996b).
This study reconstructs ecological interactions among members of the Naticidae, a family of marine drilling snails across the Plio-Pleistocene extinction and across the biogeographic boundary at Cape Hatteras, North Carolina. Naticids are a common organism in modern and Plio-Pleistocene marine benthic environment; previous research has shown that common species can be used to identify ecological changes in the makeup of a community because the variability of common species can be detected across space more easily than rare taxa (Pearman and Webber, 2007). This study will use Naticidae to examine changes in cannibalism among Naticidae across the Plio-Pleistocene and
7 biogeographic boundaries. For the purposes of this research cannibalism will be defined as confamilial predation, predation within the family Naticidae.
This research provides an understanding of how ecological interactions change across an extinction event and defines a baseline for confamilial naticid predator-prey interactions across the Plio-Pleistocene and biogeographic boundaries. Environmental changes due to human impact such as global warming have caused diversity to decline in modern marine settings. Understanding the ecological interactions among organisms of these systems prior to human interaction is necessary to manage the response of the ecosystem (Novacek and Cleland, 2001). This information may allow us to predict and react to our current biodiversity crisis in more efficient ways by allowing us to interpret how changes in ecological interactions among organisms will affect an ecosystem.
Plio-Pleistocene Extinction
The Plio-Pleistocene extinction was a regional, 2 pulsed extinction event that affected marine organisms such as mollusks, corals, bryozoans, and planktonic and benthic foraminifera (Budd and Miller, 2001; Cheetham, 1986; Hayward, 2002; Raffi et al., 1985; Svenning, 2003). It resulted in a drastic decrease in species diversity in the
Western Atlantic; 70 percent of mollusk species went extinct during this event (Allmon et al., 1996b). Geographic differences govern the surviving lineages; sub-tropical species are confined to areas south of Cape Hatteras, North Carolina, while temperate species reside in habitats north of this biogeographic boundary. Research performed in Virginia and Florida has shown a discontinuity between surviving taxa; lower species richness
8 exists north of Cape Hatteras (Allmon et al., 1996b). Stanley and Campbell (1981) indicate that almost double the number of species of mollusks appear in the fossil record than exist in the live assemblage in Virginia. Petuch (1995) also found that the species diversity within gastropod families decreased drastically across the Plio-Pleistocene in
Florida. His results indicated a two-phased extinction caused by a combination of cooling and lower productivity.
The molluscan fauna began to recover in species richness in the western Atlantic during the Pleistocene; however, Recent diversity still has not equaled the diversity from the Pliocene. This difference has prompted Allmon et al. (1996b) to suggest that areas south of Cape Hatteras, North Carolina have recovered in diversity, while faunas north of
Cape Hatteras have remained depauperate. This is hypothesized to be due to a diversion of the Gulf Stream close to Cape Hatteras. Warm Gulf Stream water travel up the eastern coast of North America until they reach Cape Hatteras, where the Gulf Steam is rerouted offshore (Berggren and Hollister, 1977). During the Pliocene, warmer waters allowed sub-tropical species to range northward into Virginia, but these species have subsequently have gone extinct in these areas due to cooling. Little work has been done comparing ecological interactions across Cape Hatteras, North Carolina during the Plio-Pleistocene extinction.
Taxonomic vs. Ecological Patterns Across Extinctions
An extinction event affects not only species richness, but also the ecological interactions among organisms. A change in either species richness or ecological
9 interactions can alter community structure; the survivors of an extinction event often change the environment once the victims are gone. To investigate these changes, it is necessary to collect data beyond taxonomic richness (Roy, 1996). This is important because faunal turnover and changes in ecological interactions are non-linear, such that small changes in ecology can occur during large extinction events and large changes in ecology can occur during small extinction events (Plotnick and McKinney, 1993).
Droser et al. (2000) found a decoupling of taxonomic and ecological severity for the end-Ordovician and Late Devonian mass extinctions. The authors developed a ranking system of ecological severity from 1 to 4 to compare different aspects of mass extinctions ranging from minor community level changes (level 4) to the appearance of a new ecosystem (level 1). Analysis showed that while the end-Ordovician mass extinction resulted in the second greatest family level loss of diversity, the ecological changes associated with the extinction event were minor. Victims of the extinction were replaced by successors inhabiting similar ecological roles. In contrast, the Late Devonian mass extinction resulted in a less severe change in species richness but second-level changes.
Investigating the ecological patterns of an ecosystem after an extinction event can provide a more complete understanding of how the extinction event affected the community. For example, while mollusk diversity changed little after the Late Triassic extinction in the United Kingdom, there were large changes in the ecology of the ecosystem (Mander et al., 2008). After the late Triassic extinction event, samples are characterized by assemblages with lower abundances, lower diversity, and lower evenness. In these cases, large ecological change occurred with small faunal turnover.
10 Layou (2009) examined paleocommunity-level data to investigate paleoecological changes during a regional extinction event during the Ordovician. The results showed that, despite a moderate decrease in diversity, evenness did not vary greatly. This suggests that the relative structure of the community remained similar pre- and post- extinction. This shows that spatial scale does not change the decoupling of taxonomic and ecological severity. These differences occur both on a global and a regional scale.
Research in modern benthic systems suggests that small changes in taxonomic diversity or abundance could have large effects on ecology. Solan et al. (2004) used a computer model to determine how rates of bioturbation would vary as benthic species went extinct. They found that the changes in bioturbation were dependent on the order in which the species went extinct and that extinction events representing the same loss of taxonomic diversity would result in different changes in ecology. This study investigates a portion of those changes. While the taxonomic changes of the Plio-Pleistocene extinction have been documented (Allmon et al., 1996b; Petuch, 1995; Stanley, 1986), the ecological interactions associated with the extinction event have not been as extensively studied.
Researchers can also use the fossil record to investigate changes in the behaviors associated with ecological interactions. Recent work examining changes in the behavior of paleo-organisms shows how an extinction event can affect interactions between two organisms without the extinction of either taxon. An example of such work was done by
Dietl et al. (2004) who used fossil data in combination with live experiments to show that feeding behavior of muricid snails changed after the Plio-Pleistocene extinction in
Florida. The researchers placed muricid snails in low-competition and high-competition
11 regimes with bivalve prey. In the low-competition tanks, the muricids exhibited similar behaviors to modern muricid snails in the wild; however, in the high-competition tanks, the snail’s behavior changed from umbo drilling to edge-drilling. Edge-drilling is a faster, but risky procedure because of the possibility that the clam can damage the proboscis by clamping down. Next the researchers investigated the fossil record of muricid predation across the Plio-Pleistocene extinction by calculating the percentage of shells drilled at the umbo or edge in the Pliocene and Pleistocene. While edge-drilling during the Pliocene occurred, no record of edge-drilling existed during the Pleistocene.
Using the information from the live experiments, the researchers inferred that edge- drilling is a sign of high competition and that the lack of edge drills during the
Pleistocene suggests that the competitive regime of the ecosystem was higher during the
Pliocene.
Naticid Cannibalism
According to Polis (1981), cannibalism can be classified as an extreme form of interference competition, lowering the fitness of one individual through the predatory actions of another member of the species. Fox (1975a) indicated that cannibalism is a
“normal phenomenon in many natural populations.” Many taxa including several species of fish, arthropods, and even mammals practice forms of cannibalism (Polis, 1981).
Cannibals gain nourishment, reduce competition for food or shelter resources, and eliminate competitor’s genes from the gene pool by preying on another member of the species (Kelley and Hansen, 2007; Polis, 1981). Theoretical (Kelley, 1991), and experimental work (Claessen et al., 2000; Rudolf, 2008) suggest that cannibalism may be
12 used as a proxy for competition because cannibalism often occurs as essential resources become scarcer. It has been suggested that as prey abundance decreases or predator abundance increases, cannibalism helps reduce the predator population to stable levels
(Claessen et al., 2000). Cannibalism by naticid snails has been shown to take place in nature and may be a result of prey selection to maximize energy gain per unit foraging time as shown by cost-benefit ratios (Kelley, 1991; Kelley and Hansen, 2007). These suggest that it is sometimes more beneficial for naticids to prey upon another member of their family then a species of bivalve.
Naticids are carnivorous, infaunal predators of mollusks, burrowing through marine sediments to drill into the shells of bivalves and gastropods (Kelley et al., 2001)
(Figure 1). Naticids envelop their prey item in their foot and drill a typically beveled hole into the shell using a combination of rasping radula and acids (Kelley et al., 2001).
This process can take anywhere from several hours to several days. The naticid then uses its proboscis to digest the prey item. This drilling behavior is preserved readily in the fossil record and can be used to determine patterns of predation in the past (Kitchell et al.,
1981). Some research has concluded that drilled shells are more likely to be destroyed than whole shells (Roy et al., 1994); however, it is unlikely that the rates of this taphonomic process changed across the Plio-Pleistocene extinction and would affect comparisons of drilling frequency. Dietl and Alexander (2000) investigated changes in drilling location in confamilial naticid prey. The researchers found that drilling position changed over time to both cover the prey’s aperture and reduce the risk of prey escape and subsequent drilling of the predator. This previous work suggests that variation in community-level dynamics can be preserved in the drilling record of naticid snails.
13 Figure 1: Picture of a sample of naticids including Euspira heros and Neverita duplicata.
Figure 2: Geologic map of the Coastal Plain (Grey and Yellow) ranging from New Jersey to Florida. Map from: http://tapestry.usgs.gov/
14 Research Questions
This study addresses several aspects of confamilial ecological interactions among naticid snails. The questions governing this research include:
1. What was the frequency of cannibalism across the Plio-Pleistocene
boundary and across the biogeographic boundary at Cape Hatteras?
2. Does naticid body size change across the Plio-Pleistocene and across the
biogeographic boundary at Cape Hatteras?
3. Does naticid size influence frequencies of cannibalism?
Geologic Setting
Data were collected from several localities across the Atlantic Coastal Plain of
North Carolina, South Caroline, and Virginia (Figure 2). The Atlantic Coastal Plain is a low-lying physiographic province adjacent to the Atlantic Ocean, and stretches from New
Jersey to Florida. During the Plio-Pleistocene, deposition within shallow marine basins along the Coastal Plain was dominated by siliciclastic sedimentation. This project focused on three stratigraphic units north of Cape Hatteras, North Carolina: the
Yorktown, Chowan River, and James City Formations, and three stratigraphic units south of Cape Hatteras: the Duplin, lower Waccamaw, and upper Waccamaw Formations
(Figure 3). The Yorktown Formation correlates with the Duplin Formation, the Chowan
River Formation correlates with the lower Waccamaw Formation, and the James City
Formation correlates with the upper Waccamaw Formation, however, the correlations of
15 these units are not completely understood. For this reason, analyses comparing each unit across the biogeographic boundary were not done. These units represent a time interval from the Late Pliocene to the Early Pleistocene. These stratigraphic units are bounded by unconformities.
The Yorktown Formation extends from Maryland into northern North Carolina, and consists of several varying lithological units (Johnson et al., 2001). Ward and
Blackwelder (1980) divided the Yorktown into four members: the Sunken Meadow,
Rushmere, Mogarts Beach, and Moore House Members. This study focuses on the
Moore House Member of the Yorktown Formation. The measured type section of the
Moore House Member contains a calcareous upper layer of orange shell hash with some fine quartz sand and a very fossiliferous lower layer of tan, clayey calcareous sand
(Johnson et al., 2001; Ward and Blackwelder, 1980). The mollusks of the Moore House
Member include generally sub-tropical taxa; this represents the last appearance of these sub-tropical species in the fossil record north of Cape Hatteras (Krantz, 1990). The
Moore House Member is geographically restricted to southeastern Virginia. Moore
House Member sediments represent shallow water depths with high current and wave energies (Ward, 2008). Molluscan assemblages indicate normal salinities; however, some taxa found in the Moore House Member are known to live in brackish-water environments.
The Chowan River Formation is considered either late Pliocene or early
Pleistocene in age. It is exposed mostly along the Chowan River in North Carolina; however, the Chowan River Formation does occur in southeastern Virginia (Ward, 2008).
The Chowan River Formation is divided into two members, the lower Edenhouse
16 Member and the upper Colerain Beach Member (Blackwelder, 1981). The Edenhouse
Member contains shelly, silty sands that are partially bioturbated. Two layers are present in the Edenhouse, a lower layer of shelly, poorly sorted, blue-green, medium quartz sand, and an upper layer of yellow, slightly shelly, fine to medium quartz sand (Blackwelder,
1981). The Colerain Beach Member contains yellowish gray micaceous clay, silt, and find subangular sand with lenses of fossils. A pebbly to bouldery unconformity rests on
the Yorktown at the base of the Chowan River Formation with interbedded silty fine
sand, clayey silt, and biofragmental sand lying on top of this base (Ward, 2008). Fossils
indicate that the Edenhouse Member was deposited in the shallow subtidal zone during
marine regression. This trend continues into the Colerain Beach Member (Blackwelder,
1981). A warm water fauna is present in the Chowan River Formation, including body
fossils and trace fossils such as burrows (Krantz, 1990).
The James City Formation is considered to be early Pleistocene in age and is
primarily exposed in northern North Carolina. The formation is composed primarily of
unconsolidated calcareous sandy clays and argillaceous sands (DuBar and Solliday,
1963). These fine-to medium-grained quartz sands are blue-grey in color when fresh, and
yellow-brown to red-brown when weathered. All units contain well-preserved fossil
material. The James City Formation was deposited during a transgression related to
global warming and melting ice (Blackwelder, 1981).
The Duplin Formation is present south of Cape Hatteras and corresponds to the
Yorktown Formation, although evidence indicates that it was deposited in warmer waters
(Richards, 1950). It extends from North Carolina into South Carolina. The Duplin is
highly fossiliferous and contains very well-preserved fossils in a matrix of fragmented
17 shells (DuBar et al., 1974). The color of these deposits ranges from golden-yellow to bluish-grey. Noncalcareous sediments occur between lenses of fossils, and are made up of well to poorly sorted clayey sands and clays. It appears that, like the Moore House
Member of the Yorktown Formation, the Duplin Formation was deposited during a regression of sea level.
The Waccamaw Formation is thought to be Pliocene to Pleistocene in age and may correspond to the Chowan River and James City Formations. It occurs in North
Carolina from the Cape Fear River into South Carolina (Johnson et al., 2001). The lower
Waccamaw is moderately well-sorted, clay and very fine to fine sands (DuBar et al.,
1974) and is highly fossiliferous with well-preserved molluscan assemblages. Fresh exposures are bluish-grey while weathered material is orange or brown in color. The lower Waccamaw sediments grade into the upper Waccamaw layer with relatively few fossils. Species richness decreases drastically in the upper Waccamaw. These sediments consist of fine to medium quartz sands interbedded with clayey sands and clays. Faunal evidence indicates that the Waccamaw Formation was deposited in a range of shallow subtidal marine to brackish water environments.
These units represent different depositional environments as transgressions and regressions occurred across these areas. These different environments may affect the frequency of cannibalism across the extinction event and the biogeographic boundary.
However, previous research has suggested that the spatial variability of naticids drilling was not related to substrate type, indicating that drilling frequency is not affected by environment (Hansen and Kelley, 1995). This suggests that naticids cannibalism can be investigated among these units regardless of environmental conditions.
18 Figure 3: Correlation chart showing the stratigraphic units used in this study. Chart from W.B. Harris.
Yorktown Formation Chowan River James City Formation Moore House Member Formation North Chuckatuck: 92, 6 Gomez Pit: 35, 2 Baven Hill: 22, 1 of Cape Rice’s Pit: 73, 6 Neuse River 1432: 33, 2 Hatteras Brant’s Pit: 99, 6 Neuse River 1442: 20, 1 Duplin Formation lower Waccamaw upper Waccamaw Formation Formation South Lumber River: 53, 4 Old Dock: 23, 2 Wilmington: 64, 4 of Cape Rayzor’s Farm: 53, 5 Register Quarry: 64, 4 Walker’s Bluff: 30, 1 Hatteras Tar Heel River: 29, 3 Elizabethtown: 20, 2 Niells Eddy Landing: 78, 3 Kirby Pond: 103, 5 Acme 1446: 99, 5 Acme 112: 117, 5 TU 870: 76, 3 Old Dock (Petit): 115, 4 Crescent Beach: 513, 3 Snake Island: 100, 4
Table 1: Sampling effect for stratigraphic units north and south of Cape Hatteras. Local- ity name: number of specimens, taxonomic richness per locality.
19 Methodology
Sampling Strategy
Samples were collected from stratigraphic units before the Plio-Pleistocene extinction north and south of the biogeographic boundary represented by Cape Hatteras, after the first pulse of extinction north and south of Cape Hatteras, and after the second pulse of extinction north and south of Cape Hatteras. This allowed analysis of ecological interactions across the extinction event.
Field Methods
Naticids were float collected at two sites in North Carolina: Register Quarry
(Figure 4) (lower Waccamaw Formation), and the Tar Heel River (Duplin Formation), and were sieved collected at the Lumber River locality (Duplin Formation) (Table 1).
Register Quarry is located south of Cape Hatteras in North Carolina and is approximately
100m long by 50m wide. Naticids were collected from slump material located around the edges of the quarry. The outcrop at the Tar Heel River is located south of Cape Hatteras in North Carolina and is approximately 40m long and 30m high. The fossiliferous
Duplin Formation material is located at the top of the outcrop. Naticids were collected from the slump material along the length of the outcrop. At Register Quarry and the Tar
Heel River, researchers walked around the outcrop and collected all naticids observed.
The Lumber River outcrop is located south of Cape Hatteras in North Carolina near the town of Lumberton. The locality is a river cut approximately 12m long by 5m high.
20 Naticids were sieved out of the sediment along the river because the outcrop is much smaller. Every specimen was collected regardless of the size or quality of preservation.
Museum Collections
Twenty-one float and bulk-collected samples from museum collections were processed in addition to field-collected samples. Samples from the Paleontological
Research Institute, the Virginia Museum of Natural History, the Florida Museum of
Natural History, and the Smithsonian Museum of Natural History were used. All samples contained at least 20 naticid specimens. A map of all field and museum-based collection localities used in this study is below (Figure 5).
Sample Processing
All naticids were identified to the species level using Coastal Plain monographs
(Campbell, 1996; Ray, 1987). Each specimen was measured using digital calipers to determine length, height, and thickness in millimeters to the hundreds place (Figure 6).
Outer and inner drill hole diameters were measured in specimens exhibiting drill holes
(Figure 6). All specimens judged to be less than 85 percent complete were excluded from this study. Taxa identified in this study included: Neverita duplicata, Euspira heros,
Naticarius canrena, Tectonatica pusilla, Lunatia interna, Lunatia perspectiva, Polinices sp., and Sinum chesapeakensis.
21 Data Analysis
Data were used to determine the frequency of cannibalism, average size, and maximum size for each sample and for size-specific ranges within a sample.
Cannibalism frequency for a sample was calculated by dividing the total number of drilled naticids in a sample by the total number of naticids in the sample. The geometric mean of body size for each individual specimen was calculated as the square root of the shell length multiplied by the shell height. Average and maximum size for the sample were calculated using this geometric mean. All data analyses were done in Microsoft
Excel and SPSS version 16.0.
Cannibalism frequency data were non-normally distributed and a square root transformation was used to normalize the data. The mean cannibalism frequency rates for the Pliocene and Pleistocene and north and south of the biogeographic boundary were calculated.
In order to control for the large variation in sample size (from 75 individuals during the Pleistocene in the north to 1127 individuals during the Pliocene in the south), a resampling procedure was used. Resampling was completed using the sample function in
R (R Core Development Team, 2005). Resampling was done with replacement and was standardized to the smallest sample size of 75 snail specimens. The presented results are based on 1000 replicate samples of 75 snail specimens across all localities within each time bin north and south of the geographic boundary. Samples were compared across geographic province and epoch using t-tests.
22 To test the relationship between naticid size and rate of cannibalism external drill hole diameter was used as a proxy for predator size. Naticid drill hole diameters were grouped into 1 mm size classes (0-1 mm, 1-2 mm, 2-3 mm, 3-4 mm, 4-5 mm, and 5-6 mm) to create different bins of predator size. Because naticids cannot cannibalize individuals that are too large and rarely drill individuals too small to provide sufficient nutrition, a potential prey size range was created for each drill hole class (Dietl, 2008).
The potential prey size range was defined as the specimens with a geometric mean between the smallest and the largest sized drill hole in the predator class. Frequency of cannibalism was then calculated for each size class of drill hole by dividing the number of drilled specimens of a size class by the potential size range. This created a cannibalism frequency for each size class of drill hole. The overall frequency of cannibalism for all drill hole classes was used as the expected value.
23 Figure 4: Photo of a stratigraphic section from Register Quarry, NC (lower Waccamaw Formation)
N
Niells Eddy’s Landing
Figure 5: Sites from field and museum collections. The yellow line represents the biogeographic boundary at Cape Hatteras. Yellow circles represent localities (2 localities are excluded because they lacked specific location information). Map from Google Earth.
Locality A : 1442 Acme, 112 Acme, Old Dock, Old Dock (Petit), TU 870, Snake Island, Register Quarry. Locality B: Neuse River 1432, Neuse River 1442, Baven Hill Locality C: Tar Heel and Elizabethtown Locality D: Lumber River and Razier Farm
24 Figure 6: Euspira heros with a naticid drill hole. Length, height, thickness, inner drill hole diameter, and outer drill hole diameter were measured on each specimen.
25 Results
Frequency of Cannibalism across the Plio-Pleistocene and Biogeographic Boundaries
Results showed a general trend of increasing cannibalism across the Plio-
Pleistocene extinction and a greater frequency of cannibalism in the north across the biogeographic boundary at Cape Hatteras, North Carolina. Frequency of cannibalism data were first divided across the extinction event and across the biogeographic boundary using the entire data set. The frequency of cannibalism analysis across the Plio-
Pleistocene (Figure 7) extinction includes localities both north and south of Cape
Hatteras, North Carolina, and frequency of cannibalism analysis across the biogeographic boundary (Figure 8) includes localities from the Pliocene and the Pleistocene. To test for statistical significance, an ANOVA was used with size defined as a covariate. This attempts to remove any size-specific effects of cannibalism from the analysis. Both of these results were statistically significant, indicating that cannibalism became more common after the extinction event and more common north of Cape Hatteras than south.
These results are suggestive of an increase in competition across these boundaries. An increase in the competitive regime could result from an increase in the number of naticids relative to prey items or a decrease in abundance of alternative prey taxa. Cannibalism can be beneficial or detrimental to a species. During times when food resources are scarce, cannibalism gives an organism another set of resources they can draw upon for nourishment. If one member of a species is cannibalistic and the other is not, then the potential pool of prey becomes larger for the cannibal.
26 When environmental stresses are high, cannibalism can help lower the abundance of predators to an appropriate carrying capacity. This helps reduce the competition for the diminishing food resources. In turn, genes that promote cannibalism would be passed on to the next generation as cannibals survived harsh environmental conditions and non- cannibals died from starvation. This benefit could drive levels of cannibalism to increase. During times when food resources are plenty, however, cannibalism could be harmful to a species. While cannibalism may impart the same advantages during times of plenty as times of scarcity, species that persist at high rates of cannibalism when alternative prey are present could cannibalize themselves to extinction. In addition, cannibalism is inherently dangerous because previous work has demonstrated that prey have the ability to kill predators (Dietl and Alexander, 2000). During times when alternative prey are abundant, attempting to cannibalize another individual could end a naticid’s chance to reproduce. More aggressive naticids which would tend to cannibalize more than others would be at a higher risk of becoming cannibalized because of increased contact with other naticids. This could drive levels of cannibalism to decrease as naticids which practiced high levels of cannibalism were subsequently cannibalized and removed from the gene pool producing fewer offspring than less aggressive naticids. It is interesting to note that this process would not remove cannibalism from naticid populations entirely, only increase or decrease the frequency of cannibalism.
Cannibalism would still be necessary to control more aggressive naticids or populations would be preyed upon highly by a small group of hyper-cannibalistic naticids. These results may suggest that an increase in competitive regime occurred during the
27 Pleistocene and north of Cape Hatteras. This may be due to a decrease in prey abundance across the extinction event.
The increase in cannibalism seen across these boundaries could also be a result of sampling strategy or sample size. While naticids are not a rare species, their abundance in bulk samples is low. A combination of float sampling and bulk sampling was chosen for this study because it was unlikely that a sufficient number of individual naticids would be collected through bulk sampling alone. However, this introduces a problem into the data set; many of the samples were collected non-randomly. While efforts were made to ensure that sampling was completed randomly by collecting all naticids, regardless of preservation or size, bias may have been introduced into the sampling. For example, small naticids are much harder to find than large naticids, small naticids may not have been sampled at localities which were float collected. In addition, most of the samples used in this study were museum samples and the exact procedure for the collection of naticids is unknown. The difference between bulk and float sampling across the Plio-Pleistocene or biogeographic boundaries could be investigated by examining naticid data from bulk sampling.
Sample size could also account for the changes in frequency of cannibalism.
While samples from south of Cape Hatteras during the Pliocene were plentiful, samples north of Cape Hatteras during the Pleistocene were rare, exhibiting only 75 individuals among 3 localities. With such a small sample size, it is possible to inflate the frequency of cannibalism by randomly picking an abnormally high number of drilled specimens.
This may mean that the high rate of cannibalism during the Pleistocene and north of Cape
Hatteras is not a real biological signal and may in fact be due to low sampling size. This
28 problem could be addressed by increasing the sample size or performing a resampling procedure.
Because this analysis spans either the Plio-Pleistocene or biogeographic boundaries, these results are not able to finely distinguish between frequency of cannibalism during the Pliocene and Pleistocene, north and south of Cape Hatteras. The comparison of the Pliocene to the Pleistocene units (Figure 7) includes localities from north and south of Cape Hatteras, and the comparison across Cape Hatteras (Figure 8) includes units from the Pliocene and Pleistocene. Therefore, it is necessary to further subdivide these units to distinguish patterns of cannibalism across these boundaries.
Data were separated into four spatiotemporal groups to test for differences between each epoch and geographic province: Pliocene North, Pliocene South,
Pleistocene North, and Pleistocene South. An ANOVA with size defined as a covariate was also used to test for differences between these groups. Results showed similar trends to using the overall data set; comparisons across the Plio-Pleistocene extinction in the north (Figure 9A) and in the south (Figure 9D) exhibited an increase in cannibalism across Plio-Pleistocene extinction and comparisons across Cape Hatteras, North Carolina during the Pliocene (Figure 9B) and the Pleistocene (Figure 9C) exhibited higher frequency of cannibalism in the north. Each of these results was non-significant; however, these results demonstrate the same patterns as the comparisons across the Plio-
Pleistocene and biogeographic boundaries using the entire data set. The large decrease in sample size that occurs when comparing these groups may be to blame for the lack of statistical significance. The number of localities for the spatiotemporal units ranged from
14-17 for the Pliocene, 3-7 for the Pleistocene, 3-4 for the north, and 3-14 for the south.
29 * * 0.60
0.40 (square root)
0.20 Mean frequency of cannibalism
0.00 Pliocene Pleistocene
Figure 7: Results indicate that cannibalism increased across the Plio-Pleistocene extinction when considering localities north and south of Cape Hatteras, NC. This may suggest that competition is increasing after the extinction event. Error
bars represent +/- 2 SE. F18,6 = 6.045 ; p = 0.008.
* 0.60
0.40 (square root)
0.20 Mean frequency of cannibalism
0.00 North South Figure 8: Results indicate that cannibalism increased across the biogeographic boundary at Cape Hatteras, NC when considering localities from the Pliocene and Pleistocene. This may suggest that levels of competition were higher north of Cape
Hatteras. Error bars represent +/- 2 SE. F 7,17 = 5.291 ; p = 0.014
30 North 1.00 A ns 0.80
0.60
0.40 (square root)
0.20 Mean frequency of cannibalism
0.00 Pliocene Pleistocene
F 17,7 = 5.291 ; p = 0.116. Pliocene Pleistocene 0.60 B 1.00 C ns 0.50 ns 0.80
0.40 0.60
0.30
(square root) 0.40 0.20 (square root)
Mean frequency of cannibalism 0.20
0.10 Mean frequency of cannibalism
0.00 0.00 North South North South F = 1.714 ; p = 0.319. F 4,14 = 1.601 4,14 ; p = 0.234. 3,3 South 0.50 D ns
0.40
0.30
(square root) 0.20
0.10 Mean frequency of cannibalism
0.00 Pliocene Pleistocene
F 14,3 = 1.280 ; p = 0.309 Figure 9: No statistically significant difference was seen between the spatiotemporal units across the Plio-Pleistocene and biogeographic boundaries, however, these patterns mirror the analyses from the entire data set. With a larger sample size these results may suggest that levels of competition were greater north of Cape Hatteras during the Pleistocene. Error bars represent +/- 2 SE.
31 If we consider this trend to be a real biological pattern, then the increased frequency of cannibalism in the north relative to the south indicates that there is some geographic-specific factor that is influencing the cannibalistic tendency of naticids in the north. Even though cannibalism frequency is non-significant for these analyses, cannibalism during the Pleistocene from north to south is roughly 20 percent higher
(Figures 9A and D). When considered with the comparisons across Cape Hatteras during the Pliocene and Pleistocene (Figures 9B and C), the difference between the frequency of cannibalism north and south of the biogeographic boundary is higher during the
Pleistocene than the Pliocene by roughly 10 percent. This may be indicative of a more competitive system north of Cape Hatteras during the Pleistocene. The highest levels of cannibalism occur during this spatiotemporal unit and this may mean that alternative prey taxa were less abundant in the north during the Pleistocene. If the Plio-Pleistocene extinction targeted naticid prey taxa more heavily than naticids, then the relative abundance of prey taxa would decrease across the extinction event, potentially corresponding with an increase in competition. Then, if a geographic factor eliminated taxa north of Cape Hatteras, putting further pressure on naticid populations, we would see the greatest level of competition in the north during the Pleistocene.
Frequency of Cannibalism across Plio-Pleistocene Stratigraphic Units
Because the Plio-Pleistocene extinction occurred as a pulsed event, changes across stratigraphic units north and south of Cape Hatteras were examined. Formations were not compared across Cape Hatteras because correlation of the stratigraphy across
32 the biogeographic boundary is unclear. An ANOVA with a post-hoc Tukey test was used to test for differences among these formations. North of Cape Hatteras no statistically significant difference was seen between the Yorktown Formation and the James City
Formation (Figure 10). The Chowan River Formation is only represented by one sample and was not included in this analysis. South of Cape Hatteras, a statistically significant difference was recorded between the Duplin Formation and the lower and upper
Waccamaw Formations; however, no significant difference was seen between the lower and upper Waccamaw Formations (Figure 11). These results seem to support the general findings of the comparisons across the Plio-Pleistocene and biogeographic boundaries. If the Chowan River sample is not considered, then there is a trend of increasing cannibalism across the pulses of the Plio-Pleistocene extinction. Although these results are non-significant, it may be because the sample size for these analyses is extremely low. When considered with the statistically significant increase in cannibalism across the extinction event (Figure 7), there may be an increase in cannibalism across each formation. We would expect to see a general increase in cannibalism across each formation as the extinction event occurred as more species went extinct or were reduced in abundance. This is supported by Petuch (1995), who found that the Plio-Pleistocene extinction occurred in two distinct phases.
33 0.80
0.60
0.40 (square root)
0.20 Mean frequency of cannibalism
0.00 Yorktown Chowan River James City Formation Figure 10: Results indicate that there is no statistically significant difference in the frequency of cannibalism between the Yorktown and James City Formations. The Chowan River Formation is represented by only one sample and was not tested statistically. F 3,3 = 3.040 ; p = 0.157 Error bars represent +/- 2 SE.
0.40
0.30
0.20 (square root)
0.10 Mean frequency of cannibalism
0.00 Duplin lower Waccamaw upper Waccamaw Formation Figure 11: Results indicate that there is a statistically significant difference in the frequency of cannibalism between the Duplin and lower Waccamaw Formations (F 5,8 = -0.1585 ; p = 0.022) and the Duplin and upper Waccamaw Formation (F 5,3 = -0.2239 ; p = 0.014). No statistically significant difference was seen between the lower Waccamaw and upper Waccamaw
Formations (F 8,3 = -0.0654 ; p = 0.557). Error bars represent +/- 2 SE.
34 Frequency of Cannibalism across the Plio-Pleistocene and Biogeographic Boundaries:
Sample Size Standardization Using a Resampling Procedure
When the data were divided into each province and time bin (North Pliocene,
North Pleistocene, South Pliocene, and South Pleistocene), sample sizes varied greatly
(Table 1) due to difficulty obtaining samples from the Chowan River and James City
Formations. Results based on a resampling to a standard sample size of 75 naticid snails
exhibited the same patterns as the comparisons across epoch and geographic province;
greater cannibalism was seen during the Pleistocene and north of Cape Hatteras, North
Carolina (Figures 12 A-D). The resampling procedure implies that the patterns seen in
this study are not due to the effects of sample size and instead may be caused by a
biological signal. These results are consistent with an increase in competition across the
Plio-Pleistocene extinction and north of Cape Hatteras. These statistically significant differences between levels of cannibalism within spatiotemporal units suggest that competition was rising during the Plio-Pleistocene extinction and from south to north across Cape Hatteras as naticids preyed on confamilials.
35 North 0.50 * * * A 0.40
0.30
0.20
0.10 Mean Frequency of Cannibalism
0.00 Pliocene Pleistocene
t 1000,1000 = -152.151 ; p = 0.000 BCPliocene ns Pleistocene 0.12 0.50 * * *
0.10 0.40
0.08 0.30
0.06 0.20 0.04
0.10 Mean Frequency of Cannibalism
Mean Frequency of Cannibalism 0.02
0.00 0.00 North South North South t = 129.011 ; p = 0.000 t 1000,1000 = -0.016 ; p = 0.987 1000,1000 South * * * 0.15 D
0.10
0.05 Mean Frequency of Cannibalism
0.00 Pliocene Pleistocene
t 1000,1000 = -22.820 ; p = 0.000
Figure 12: Resampling within spatiotemporal units shows a statistically significant difference in cannibalism across the Plio-Pleistocene extinction north and south of Cape Hatteras and across the biogeographic boundary during the Pleistocene. No statistically significant difference was seen north and south of Cape Hatteras during the Pliocene. Error bars represent +/- 2 SE.
36 Changes in Size across the Plio-Pleistocene and Biogeographic Boundaries
The average sizes (calculated as geometric mean) from samples before and after the extinction and north and south of the biogeographic boundary of all drilled and undrilled naticids were compared to determine changes in size. A t-test was used to determine differences in size across the Plio-Pleistocene and biogeographic boundaries.
Across the entire data set, average size and maximum size did not show statistically significant changes across the Plio-Pleistocene boundary (Figure 13 A-B), and no statistically significant difference was seen in average size across the biogeographic boundary (Figure 13 C). However, a statistically significant change was seen across the biogeographic boundary when comparing maximum size; naticid maximum body size increased south of Cape Hatteras (Figure 13D). Overall, results show little to no difference in naticid size across the Plio-Pleistocene and biogeographic boundaries.
An increase in body size may give some naticids cannibalistic advantage over others. Also, in some systems, small individuals are able to outcompete larger individuals for resources, and cannibalism helps improve the fitness of large individuals
(Claessen et al., 2000; Rudolf, 2008). We may expect to see a change in cannibalism not because of a change in competition, but because of a change in body size across the Plio-
Pleistocene extinction or from north to south. However, little or no change in body size was seen in these analyses. In the only analysis returning a statistically significant result, the change in size was opposite of the direction of the change in cannibalism. Naticids are growing larger in the south where less cannibalism is occurring. This is opposite of what we would expect. From this we can infer that body size was not a cause of the changes in frequency of cannibalism seen in the previous results. Instead, this may be
37 due to changes in the levels of competition within the ecosystem. Similarly, when compared across geographic area and epoch, average and maximum naticid body size only showed a significant increase in maximum size south of Cape Hatteras, North
Carolina during the Pleistocene (Figure 14 H). No statistically significant result was found between average or maximum size across the Plio-Pleistocene extinction and during the Pliocene (Figure 14 A-G). A naticid can only prey upon an individual smaller than themselves. This means that an increase in average size may not induce higher cannibalism if all naticids are increasing in size. They still will be drawing from the same cannibalistic prey pool if their size increases proportionately. An increase in maximum size may be able to induce a change in cannibalism because it means that certain individuals were becoming larger. If the entire population is becoming larger proportionately, then the potential prey pool stays the same. However, if maximum size increases while average size remains the same, it means that a small number of individual naticids are becoming much larger than other naticids in the population. This may mean that their potential prey pool increases. They may begin to cannibalize more frequently because they are able to eat a larger proportion of naticids and this could potentially drive up cannibalism. The only statistically significant change in this data is an increase in maximum size in the south during the Pleistocene (Figure 14 H). There is no corresponding change in average size (Figure 14 G). This indicates that individuals remained similar sizes across the biogeographic boundary during the Pleistocene except for naticids that were in the south. This does not support the hypothesis that no change in average size accompanied by an increase in maximum size would cause cannibalism to increase because the frequency of cannibalism increases from south to north across the
38 biogeographic boundary. While this may cause cannibalism frequency to increase in the south of Cape Hatteras during the Pleistocene, it is eclipsed by greater levels of cannibalism in the north.
To determine if size-specific effects of cannibalism influenced cannibalism across the Plio-Pleistocene extinction, naticids were grouped into 10mm size bins and were divided into drilled and non-drilled specimens. They were then graphed as a stacked percent histogram to determine how drilling frequency within size classes changed across the Plio-Pleistocene extinction and Cape Hatteras, North Carolina (Figure 15 A-D). The distribution of drilled specimens remains similar across the Plio-Pleistocene extinction and biogeographic boundary, indicating that size-specific effects were not the cause of the increase in cannibalism across these boundaries. Also, these graphs indicate a higher level of cannibalism during the Pleistocene and north of Cape Hatteras. In addition, outer drill hole diameter was plotted against prey size for all drilled specimens to determine if there was a change in the size of confamilial prey naticids were drilling (Figure 16 A&B).
These results also indicated little change across the Plio-Pleistocene or biogeographic boundaries.
39 Average Size Maximum Size
30.0 B ns A ns 60.0
20.0 40.0
10.0 Mean Average Size (mm) Mean Maximum Size (mm) 20.0
0.00 0.00 Pliocene Pleistocene Pliocene Pleistocene
t 18,6 = 1.013 ; p = 0.322 t 18,6 = 1.259 ; p = 0.221
Average Size Maximum Size 30.0 C ns D * 60.0
20.0 40.0
10.0 Mean Average Size (mm) Mean Maximum Size (mm) 20.0
0.00 0.00 North South North South
t = -2.189 ; p = 0.039 t 7,17 = -1.612 ; p = 0.121 7,17
Figure 13: Results indicate little to no change in naticid body size across the Plio- Pleistocene and biogeographic boundaries. This suggests that changes in cannibalism are not due to changes in naticid size. Error bars represent +/- 2 SE.
40 Average Size Maximum Size 40.0 A 60.0 B ns 30.0 ns
40.0 20.0 North Mean Avgerage Size Mean Maximum Size 20.0 10.0
0.00 0.00 Pliocene Pleistocene Pliocene Pleistocene
t 4,3 = 1.150 ; p = 0.303 t 4,3 = 1.265 ; p = 0.261 40.0 ns C D ns 60.0 30.0
40.0 20.0 South Mean Avgerage Size Mean Maximum Size 20.0 10.0
0.00 0.00 Pliocene Pleistocene Pliocene Pleistocene
t 14,3 = -0.045 ; p = 0.965 t 14,3= -0.045 ; p = 0.967 40.0 ns E 60.0 F 30.0 ns
40.0 20.0 Pliocene Mean Avgerage Size Mean Maximum Size 20.0 10.0
0.00 0.00 North South North South t = -0.964 ; p = 0.350 t 4,14= -0.621 ; p = 0.544 4,14 40.0 * GHns 60.0 30.0
40.0 20.0 Mean Maximum Size
Mean Avgerage Size 20.0 Pleistocene 10.0
0.00 0.00 North South North South
t 3,3= -1.694 ; p = 0.165 t 3,3= -3.243 ; p = 0.032 Figure 14: Results indicate little to no change in naticid body size across spatiotemporal units suggesting that changes in cannibalism are not due to changes in size-specific interactions between naticids. Error bars represent +/- 2 SE.
41 % Stacked Size Frequency (mm) % Stacked Size Frequency (mm) ABPliocene Pleistocene 100% 100%
80% 80%
60% 60% Drilled Drilled 40% 40% Undrilled Undrilled 20% 20%
0% 0% 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71+ 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71+
% Stacked Size Frequency (mm) % Stacked Size Frequency (mm) C North D South 100% 100%
80% 80%
60% 60% Drilled Drilled 40% 40% Undrilled Undrilled 20% 20%
0% 0% 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71+ 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71+
Figure 15: When divided into 10mm size bins, naticid cannibalism exhibits similar distributions before and after the extinction event, and north and south of the biogeographic boundary
42 10.00 Plio-Pleistocene Boundary Pliocene A Pleistocene
8.00
6.00
ODH (mm) 4.00
2.00
Pliocene: R2 Linear = 0.437 Pleistocene: R2 Linear = 0.555
0.00
0.00 10.00 20.00 30.00 40.00 50.00 60.00 Prey Size (mm)
Biogeographic Boundary North 10.00 B South
8.00
6.00 ODH (mm)
4.00
2.00 North: R2 Linear = 0.341 South: R2 Linear = 0.527
.00 0.00 10.00 20.00 30.00 40.00 50.00 60.00 Prey Size (mm)
Figure 16: Outer drillhole diameter (ODH) vs. prey size indicates that naticid predators were preying on similar sized naticids across the Plio-Pleistocene and biogeographic boundaries.
43 When considering resampled data, patterns remained similar for average and maximum size across the Plio-Pleistocene extinction and biogeographic boundary. A t- test was used to test for differences between these groups. Resampled data showed statistically significant differences in size across each spatiotemporal unit (Figures 17 A-
H). Even though these relationships are significant, the amount of change from one unit to the other is small. While this indicates a change in body size across the Plio-
Pleistocene extinction this could be due to a very small increase in general body size of naticids. The largest change occurs in maximum size across the Plio-Pleistocene extinction in the north when naticids decrease in body size by roughly 2 centimeters.
Naticid sizes exhibit the same pattern across the Plio-Pleistocene extinction north and south of the biogeographic boundary. This seems to suggest that naticids do actually decrease in size across the extinction event. It is possible that this change in body size would affect the frequency of cannibalism across these boundaries; however, it seems unlikely because average and maximum size is decreasing proportionately to each other north and south of Cape Hatteras. The average naticid and the largest naticid are becoming smaller together in similar proportions; this suggests that there is not a size- specific factor associated with the increase in cannibalism. It is more likely that factors associated with the extinction event put selective pressures on naticids which caused them to increase their rates of cannibalism.
44 Average Size Maximum Size 30.0 A * * * B * * * 60.0
20.0 40.0 North 10.0 20.0 Mean Avgerage Size (mm) Mean Maximum Size (mm)
0.00 0.00 Pliocene Pleistocene Pliocene Pleistocene t = 129.001 ; p = 0.000 t 1000,1000 = 129.001 ; p = 0.000 1000,1000 * * * C D 30.0 * * * 60.0
20.0 40.0 South 10.0 20.0 Mean Maximum Size (mm) Mean Avgerage Size (mm)
0.00 0.00 Pliocene Pleistocene Pliocene Pleistocene t = 129.001 ; p = 0.000 t 1000,1000 = 129.001 ; p = 0.000 1000,1000 * * * 30.0 E * * * F 60.0
20.0 40.0
Pliocene 10.0 20.0 Mean Average Size (mm) Mean Maximum Size (mm)
0.00 0.00 North South North South t = 129.001 ; p = 0.000 t 1000,1000 = 129.001 ; p = 0.000 1000,1000 20.0 * * * 20.0 * * * GH
15.0 15.0
10.0 10.0
Mean Average Size (mm) 5.00 5.00 Pleistocene Mean Maximum Size (mm)
0.00 0.00 North South North South t = 129.001 ; p = 0.000 1000,1000 t 1000,1000 = 129.001 ; p = 0.000 Figure 17: A statistically significant change in average and maximum size was seen in each spatiotemporal unit. There is a general trend of decreasing size across the Plio-Pleistocene extinction and an increase in size south of Cape Hatteras with the exception of average size during the Pliocene, which was greater north of Cape Hatteras. Error bars represent +/- 2 SE.
45 Effects of Naticid Predator Size on the Frequency of Cannibalism
When naticid predator sizes are compared based on the entire data set using external drill hole diameter as a proxy for size, results indicate that there is no size selectivity for naticid cannibals; smaller individuals cannibalize as readily as larger individuals (Figure 18). External drill hole diameter was compared across 6 size classes of drill holes and a chi-square test was used to determine differences between the drill hole size classes. Results were not statistically significant, indicating that larger individuals do not cannibalize other naticids more than smaller individuals. This result is contrary to cannibalistic tendencies seen in other organisms in which the largest individuals perform the most cannibalism (Claessen et al., 2000; Rudolf, 2008). One of the major differences between naticids and these systems is that naticids are able to cannibalize others as soon as they start feeding as juveniles. This means that cannibalism can occur in any size class as long as the predator is larger than the prey. For young and old naticids, cannibalism is always an option and may be a stable part of their diet which could fluctuate depending on environmental conditions.
46 6.00
5.00
4.00
3.00
2.00 Frequency of cannibalism
1.00
0.00 0 - 1 1 - 2 2 - 3 3 - 4 4 - 5 5 - 6 Drill hole size class (mm)
Figure 18: No statistically significant difference was seen in percent cannibalism between drill hole classes. Chi-square = 2.826 ; p = 0.727
47 Discussion
When including the entire data set, a statistically significant difference was seen between the frequency of cannibalism across the Plio-Pleistocene and biogeographic boundaries suggesting that naticids were performing more acts of cannibalism in the north and after the extinction event. Although examining differences between cannibalism frequency by dividing each area and epoch into groups (North Pliocene,
North Pleistocene, South Pliocene, and South Pleistocene) did not exhibit statistically significant results, this may be due to a large drop in sample size. The resampling procedure indicated significant results across the Plio-Pleistocene extinction and across
Cape Hatteras, North Carolina. This suggests that increasing sample size would produce a statistically significant difference between these groups. Little to no difference was seen in naticid body size across the Plio-Pleistocene and biogeographic boundaries and there seems to be no difference in the frequency of cannibalism between large and small individuals.
These changes in rates of cannibalism may represent a change in the competitive regime of the benthic system after the Plio-Pleistocene extinction and north of Cape
Hatteras. I originally hypothesized that greater cannibalism would be seen during the
Pliocene and south of the biogeographic boundary because the extinction event was more severe north of Cape Hatteras (Allmon et al., 1996b), potentially leaving resource niches open for naticids. These results, however, do not support this hypothesis. Alternatively, changes in the environment after the Pliocene may have caused localized extinction north of Cape Hatteras, removing alternative prey items for naticids. As the abundance of prey
48 items decreased, competition for the remaining prey would increase, potentially driving naticids towards cannibalism.
Cannibalism may increase in a high competition environment because it may result in an advantage for the cannibal. Cannibalism has three major benefits: it provides nourishment for the cannibal, it removes competitors from the environment, and it removes the competitor’s genes from the gene pool (Fox, 1975b; Polis, 1981). However, cannibalism can be dangerous for organisms because the roles of predator and prey can easily switch, and the original cannibal can become cannibalized (Dietl and Alexander,
2000). If the frequency of cannibalism is too high, then species could reduce their own abundance, prey upon potential mates, or prey upon your own progeny, potentially leading to extinction. As discussed earlier, this behavior may be beneficial in environments where food resources are low. When alternative prey are not available naticids may increase their frequency of cannibalism in order to compensate for the loss of prey items. Previous research has shown that there is an inverse relationship between cannibalism and alternative prey abundance; as prey abundance decreases, cannibalism increases (Fox, 1975b; Polis, 1981). Cannibalism could be a mechanism to decrease competition by stabilizing predator populations to the carrying capacity while at the same time gaining needed nourishment. If predation of organisms prevents interspecific competition by reducing the size of the population (Sih et al., 1985), then confamilial predation of naticids would reduce competition among naticids. Therefore, we would expect that if the increase in naticid cannibalism across the Plio-Pleistocene extinction was due to an increase in competitive regime, then the abundances of alternative prey
49 taxa would decrease across the extinction event and the biogeographic boundary, especially during the Pleistocene.
This question was investigated using paleocommunity data from Huntoon (1999).
Huntoon’s data were collected across several Coastal Plain stratigraphic units, including the Waccamaw, James City, Flanner’s Beach, and Neuse Formations. The percent abundance, number of complete drill holes, and drilling frequency were used. For bivalves, the drilling frequency is defined as:
ݏݒ݈ܽݒ݈݈݁݀݁݅ݎ݂͓݀ ݏ݂ݒ݈ܽݒ͓݁ ൬ ʹ ൰
The analysis included here only examined the Waccamaw and James City Formations, which span the Plio-Pleistocene boundary. While the formation names imply that the comparison is across the biogeographic boundary, the James City Formation sample was collected right on the biogeographic boundary. This analysis only examines the community south of Cape Hatteras across the extinction event. Huntoon’s samples were collected as bulk samples and all specimens were identified to species level. This data was imported into Microsoft Excel and SPSS version 16.0 for analysis.
Analysis indicated a decrease in the overall drilling frequency across the Plio-
Pleistocene extinction from 11 percent of the community during the Pliocene to 6 percent during the Pleistocene (Figure 19 A&B); also, when considering only species that naticids drill, the Potential Prey Pool, drilling frequency decreases from 14 percent during the Pliocene to 7 percent during the Pleistocene (Figure 19 C&D). In addition, the relative abundance of prey items increases across the extinction events by 11 percent
(Figure 20 A&B).
50 Drilling Frequency Drilling Frequency Pliocene Pleistocene DRILLED DRILLED 11% 6%
A B
UNDRILLED UNDRILLED SPECIMENS SPECIMENS 89% 94%
Drilling Frequency Drilling Frequency of Prey Items of Prey Items Pliocene Pleistocene
PREY DRILLED 14% PREY DRILLED 7% C D
PREY PREY UNDRILLED UNDRILLED 86% 93%
Figure 19: Total drilling frequency (A&B) and prey item drilling frequency (C&D) decrease across the Plio-Pleistocene extinction
51 Prey Abundance Prey Abundance Pliocene Pleistocene ABUNDANCE ABUNDANCE NON-PREY NON-PREY 10% A 21% B
ABUNDANCE ABUNDANCE PREY PREY 79% 90%
Figure 20: Naticid prey abundance increases across the Plio-Pleistocene extinction.
Predator to Prey Abundance Predator to Prey Abundance Pliocene Pleistocene
PREDATORS PREDATORS 15% 4%
CD
NON- NON- PREDATORS PREDATORS 85% 96%
Figure 21: Predator to prey abundance decreases drastically across the Plio-Pleistocene extinction.
52 This suggests that prey items are becoming more available to predators across the extinction event as they become more abundant in the community. However, the increase in cannibalism documented here could also be due to an increase in the predator to prey ratio. This does not seem to be the case because the ratio of predators to prey decreases across the extinction event by 11 percent (Figure 21 A&B). Predators are becoming less common in the community after the Plio-Pleistocene extinction. However, we also see that cannibalism is increasing in this data set (Figure 22). This may be because naticids are becoming more common relative to other predators. Naticids increase from 1.1 percent relative abundance during the Pliocene to 1.8 relative abundance during the Pleistocene. This represents a 17 percent increase in the ratio of naticids to other predators (Figure 23 A&B) and no species of naticids went extinct across the Plio-Pleistocene boundary. This is odd because naticids are becoming a more prominent member of the predatory as drilling frequency is decreasing overall. These results are not consistent with the hypothesis that increasing competition drove naticids to perform more acts of cannibalism because alternative prey items were not present.
These results suggest it is possible that as naticids were becoming more prominent in the community they began to encounter each other more frequently. As this occurred they began to drill each other more frequently. This would seem to suggest that naticids drill the first individual of a prey species that they come across; however, previous research has claimed that naticids have a preference for certain prey items (Kitchell et al.,
1981). This begs the question, what were naticids eating before and after the extinction event? Victims, survivors, and originators were determined using the Huntoon data.
53 0.25
0.20
0.15
0.10 Frequency of cannibalism
0.05
0.00 Pliocene Pleistocene
Figure 22: Cannibalism increases across the Plio-Pleistocene extinction in the Huntoon (1999) data.
Na cid Predator to Na cid Predator to Non-Na cid Predator Non Predator Abundance Abundance - Pliocene Pleistocene NATICID PREDATORS 14% NATICID PREDATORS 34% AB
NON NON NATICID NATICID PREDATORS PREDATORS 66% 86%
Figure 23: Naticids increase in relative abundance as compared to other predators across the Plio-Pleistocene extinction.
54 All species present in the Pliocene only were classified as victims of the extinction event, all species present in the Pleistocene only were classified as originators, and species in both the Pliocene and Pleistocene were classified as survivors. Seventeen percent of the abundance of the community during the Pliocene was made up of victims of the extinction event while these species represented 20 percent of the naticids drilling during this time interval (Figure 24 A&C). In contrast, originators made up only 5 percent of the community during the Pleistocene and only 4 percent of the drilling (Figure 24 B&D).
This suggests that survivors of the extinction event became a larger component of the naticids diet and originators did not fill the gaps in diet left by the species that went extinct across the extinction event. Also, when considering cannibalism within the diet of naticids, cannibalism represented 1 percent of the predation on survivors during the
Pliocene but 5 percent during the Pleistocene. This corresponds with the increase in cannibalism seen in the research presented in this study (Figure 25 A&B). In addition, while predators represented 10 percent of the abundance of the victims of the extinction event, predators represented 19 percent of the abundance of the originators. This suggests that prey taxa were not replaced after the extinction event. Together, these results may mean that while the group of prey taxa as a whole was increasing across the extinction event, the preferred prey taxa were going extinct or reducing in abundance, creating a smaller prey pool for naticids.
55 Abundance of Survivors vs. Abundance of Survivors vs. Vic ms Originators ORIGINATORS 5% A VICTIMS B 17%
SURVIVORS 83% SURVIVORS 95%
Drilling Frequency Drilling Frequency Survivors vs. Vic ms Survivors vs. Originators
ORIGINATORS 4% C VICTIMS D 20%
SURVIVORS 80% SURVIVORS 96%
Figure 24: Victims of the extinction event made up a larger portion of the community than originators (A&B) and a larger portion of the drilling frequency (C&D).
Drilling Frequency Drilling Frequency with Cannibalism with Cannibalism CANNIBALISM CANNIBALISM 1% ORIGINATORS 5% 4%
A VICTIMS B 20%
SURVIVORS SURVIVORS 79% 91%
Figure 25: When cannibalism is examined within the drilling frequency, cannibalism increases across the Plio-Pleistocene extinction.
56 If naticids do prey on taxa preferentially, then the most frequently drilled taxa would be the preferred taxa during a time period. The 5 taxa with the greatest amount of drill holes are Caryocorbula auroraensis, Chione latilirata, Glycymeris Americana,
Pleuromeris decemcostata, and Oysters during the Pliocene and Crassinella lunulata,
Pleuromeris decemcostata, Caryocorbula auroraensis, Gouldia metastriatum, and
Pleuromeris auroraensis during the Pleistocene. The top 5 species during the Pliocene will be referred to as the Pliocene preferred taxa and the top 5 species during the
Pleistocene will be referred to as the Pleistocene preferred taxa. If these preferred taxa groups are considered, both represent 44 percent of all drill holes during their time period
(Figure 26 A&B). During the Pliocene, 11 percent of the abundance of the Pliocene preferred taxa are victims of the extinction event while 0 percent of originators represent
Pleistocene preferred taxa. This suggests that originators are not being drilled heavily and are not replacing extinct taxa in terms of ecological interactions with naticids.
If the abundance of the preferred taxa decreased across the extinction event, then a large portion of the diet of naticids would not be present during the Pleistocene. Many of the species drilled by naticids only represent a small number of individuals, and may have been drilled by naticids occasionally, but do not represent a group to taxa naticids drill often. If the abundance of these species decreases across the Plio-Pleistocene boundary decreased, then this could have caused an increase in competition for these food resources, and driven the observed increase in cannibalism.
57 % of Drilled Specimens: Top 5 Most Commonly Drilled with ExƟncƟon VICTIMS PLIOCENE 11% PREFERRED A PREY 33%
OTHER 56%
% of Drilled Speciemens: Top 5 Most Commonly Drilled with OriginaƟon ORIGINATION 0%
B
TOP 5 44% OTHER 56%
Figure 26: The 5 most commonly drilled prey from both the Pliocene and Pleistocene represent 44% of drill holes in the sample. However, during the Pliocene, 11% of the drill holes present are in species that go extinct while in the Pleistocene, 0% of the drill holes present are in species that originate.
58 When the abundance of both groups of preferred pray taxa are considered during
the Pliocene and the Pleistocene, the abundance of the Pliocene preferred prey taxa
decreases across the extinction event while the abundance of the Pleistocene preferred
taxa remains the same. The Pliocene preferred taxa decrease from a relative abundance
of 37 percent during the Pliocene to 19 percent during the Pliocene (Figure 27 A&C). In
contrast, the relative abundance of the Pleistocene preferred taxa remains the same at 17
percent during the Pliocene and 17 percent during the Pleistocene (Figure 27 B&D). This
suggests that the Pliocene preferred taxa were affected by the extinction event while
Pleistocene preferred taxa were not.
As the abundance of the Pliocene preferred prey decreased naticids would have
had to switch food sources for survival. They began to prey upon the Pleistocene
preferred taxa, but these species made up a smaller set of prey items because their
abundance was less than the Pliocene preferred prey. This means that the abundance of
preferred taxa, almost half of the drilled specimens during the Pliocene and Pleistocene,
was decreasing across the extinction event. This decrease in the abundance of preferred
prey taxa could have caused an increase in the competitive regime by reducing food
resources. After the extinction event naticids did not switch prey and begin drilling many originating taxa, meaning that the majority of the potential prey pool was made up of the survivors of the extinction event, which represent only 80 percent of the drilled specimens from the Pliocene. At the same time, the abundance of naticids in the community was increasing. More food resources would be needed to sustain this population. These factors would result in increased competition for prey items naticids prefer.
59 Pliocene Preferred Prey Pliocene Preferred Prey Pliocene B Pleistocene A PLIOCENE PREFERRED PREY PLIOCENE 19% COMMON PREY ABUNDANCE 37%
OTHER OTHER 63% 81%
C D Pleistocene Preferred Prey Pleistocene Preferred Prey Pliocene Pleistocene PLEISTOCENE PLEISTOCENE PREFERRED PREFERRED PREY PREY 17% 17%
OTHER OTHER 83% 83%
Figure 27: The Pliocene preferred prey taxa decrease in abundance across the Plio-Pleistocene extinction (A&C) while the Pleistocene preferred prey taxa do not (B&D). This suggests that the Pliocene preferred taxa were affected by the extinction event and reduced the potential prey pool of preferred items for naticids.
60 The decrease in drilling frequency can be explained because when all prey items are included, the majority of which only exhibit a small number of drill holes, the abundance of prey items increases across the extinction event. This inflates the denominator of drilling frequency, lowing drilling frequency in the community after the extinction event.
It is possible that after the extinction event naticids began to prey more heavily on survivors, including confamilials in order to make up for the taxa that went extinct across the Plio-Pleistocene boundary. These results support the results shown in this study, that an increase in cannibalism occurred across the Plio-Pleistocene boundary, and support the hypothesis that this increase was due to a increase in the competitive regime across the extinction event. However, instead of the abundances of all species decreasing, the abundances of species that naticids drill most often decreases. While alternative prey are available, naticids are not switching to these alternative prey items and may be increasing their levels of cannibalism in order to compensate for this loss of prey.
One problem with these analyses concerns the sample size of the two localities used with the Huntoon data. The Pliocene unit from the lower Waccamaw formation contained only 1046 specimens while the Pleistocene unit from the James City Formation contained 4228 specimens. Without controlling for the amount of specimens in a sample, it is possible that metrics like species richness would be inflated in the James City
Formation because the high number of specimens would make it more likely that rare species would be present in the sample. In order to control for this, a rarefaction procedure could be done. Rarefaction helps compare the species richness of two samples of different size by estimating the amount of species that would be found if the two samples had the same number of individuals. This would allow a comparison of how the
61 extinction event affected species across the Plio-Pleistocene boundary and may result in statistically removing rarer species from the potential prey pool, increasing the amount of drilling frequency during the Pleistocene. Also, it is important to increase the number of localities to examine the effect of spatial variability of these results. This data represents one locality from the Pliocene and one locality from the Pleistocene. It would be useful to increase the number of samples to look at how the community responded to the Plio-
Pleistocene extinction and expand the analysis north of the biogeographic boundary to capture information about how systems north of Cape Hatteras responded to the extinction event.
Future work could attempt to determine the cause of the Plio-Pleistocene extinction. The two major proposed causes of the Plio-Pleistocene extinction are ocean cooling due to glaciation and a decrease in primary productivity. The question remains, what factor truly influenced the extinction event across the Plio-Pleistocene boundary?
One way to test these hypotheses would be to use conventional metrics for temperature and productivity. Allmon et al (1996a) investigated vertebrate and invertebrate paleontological data such as size of suspension feeders, along with isotopic fossil data and concluded that productivity decreased in Florida across the Plio-Pleistocene boundary. A similar study could be completed across Cape Hatteras and the Plio-
Pleistocene boundary similar in scope to this study. Carbon-14 and Oxygen-18 isotopes are commonly used as proxies for productivity and temperature respectively, diversity and abundance of microfossil data, especially foraminifera and ostracods are used to reconstruct paleoclimate and nutrient regimes, and the size of a species’ range may positively correlate with their temperature tolerance. However, it may also be possible to
62 determine the cause of the Plio-Pleistocene extinction using community level and geographic range data.
In order to determine if a decrease in productivity or temperature caused this event using naticid cannibalism data then we must ask, why did the abundance of their prey taxa decrease? Naticids have been known to live in cold waters, they exhibit a wide geographic range and are found along the coast of the United States from Massachusetts to Texas (Abbott, 1974). If a cooling event occurred during the Pliocene, then naticids may have been able to cope with the change in temperature themselves. Alternative prey taxa, however, may not have been as able to survive the cooling ocean waters as naticids, decreasing their abundances. As their abundances decreased, competition among naticids for food resources would rise, and could result in an increase in cannibalism. Similarly, if a decrease in productivity was to blame for the Plio-Pleistocene extinction then filter feeding prey taxa would begin to decrease in abundance and naticids would prey upon a greater proportion of prey taxa in order to obtain the nutrients needed for survival. As preferred prey taxa became less abundant, competition would increase, and naticids would resort to cannibalism in order to survive.
By plotting the geographic range of each species during the Pliocene and the
Pleistocene, it is possible to observe how these ranges change over the extinction event.
If ocean cooling caused the extinction event, then we would expect to see naticids prey taxa migrate south into warmer waters, driving up levels of cannibalism in areas the prey taxa migrated from. This could not exclude a combination of temperature and productivity as a cause; however, if the diversity of filter feeding mollusks did not decrease across the extinction event then this would be strong evidence that a decrease in
63 productivity did not cause the Plio-Pleistocene extinction. In contrast, if the geographic ranges of prey species stayed the same or increased northward, then cooling temperature as a cause of the Plio-Pleistocene extinction would not be reasonable because prey species would be moving into colder waters during the Pleistocene. If this coincided with a decrease in the diversity of filter feeding mollusks, then this would be evidence that productivity was the cause of the Plio-Pleistocene extinction.
64 Conclusions
The results from this study indicate that levels of naticid cannibalism differ across the Plio-Pleistocene boundary and the biogeographic boundary at Cape Hatteras, North
Carolina suggesting a change in the ecological interactions between organisms in this system. This study found that frequency of cannibalism of naticid snails was higher during the Pleistocene after the extinction event and north of Cape Hatteras, North
Carolina. In addition, this study found that there was little to no change in naticid body size across these spatiotemporal areas and no difference in the frequency of cannibalism between small and large individuals.
When resampled to standardize for sample size, results remained similar and were statistically significant when comparing frequency of cannibalism across spatiotemporal areas. This suggests that the pattern of naticid cannibalism seen here is not due to the effects of sample size and the original analyses would increase in statistical significance if more samples were collected.
When considered in context with previous research, these results seem to suggest an increase in the competitive regime of the ecosystem after the Plio-Pleistocene extinction and north of Cape Hatteras, North Carolina. This area may have experienced differences in the extinction event that govern the surviving lineages such as the deflection of the Gulf Stream at Cape Hatteras during the Pleistocene. This could have caused preferred naticid prey species to decrease in abundance, putting pressure on surviving naticids which may have lead to a greater frequency of cannibalism.
65 These results suggest that it may be possible to use levels of naticid cannibalism as a proxy for the level of competition experienced in a community. We can use this information to define baselines to compare to modern extinctions. By investigating the amount of cannibalism in modern systems, we can track ecological interactions between organisms as well as abundances of taxa in order to determine the health of an ecosystem.
If naticid cannibalism is particularly high in an area, it may mean that alternative prey taxa are below normal levels of abundance and that the health of that ecosystem is in question. Rates of cannibalism can be used as another metric to help determine changes in competition and track modern levels of extinction.
66 Acknowledgements
I never considered paleontology as a topic I would study. I liked dinosaurs as much as the next third grader, but not enough to pursue fossils into college. Instead, I found myself in the biology department preparing to teach high school science. Then in the spring of 2007 my friend convinced me to take paleontology with her. It would fulfill credit hours for my biology major and she assured me that the professor was fantastic. I agreed and suddenly I was indebted to her for making my take my first paleontology class. When I look back, that was what started me upon the path I am on today.
Over the past two and a half years I’ve been able to delve into the fossil record under the guidance of some of the most intelligent and generous people I’ve ever met.
First, I would like to thank my two mentors Dr. Rowan Lockwood and Dr. Karen Layou.
It is difficult to describe how much my two professors have helped me grow as a scientist and a person. Their doors are always open for a quick question that ends up taking over an hour, or just to listen to me complain about my other classes. They have been incredibly generous, sharing their time and their lives to get to know a student who just likes to play with fossils. They demand excellence, and push me to produce the best work I can accomplish while giving me the guidance I need to do it. The greatest praise I can give them is that they are the best teachers I’ve ever had.
While I started my studies in paleontology at the College of William and Mary, I immersed myself in the fossil record at the University of North Carolina at Wilmington.
During the summer of 2008 I participated in a REU program lead by Dr. Patricia Kelley,
Dr. Greg Dietl, and PhD. Student Christy Visaggi. Over the course of the summer they
67 taught me how to be a paleontologist, and began helping me structure the work that would become this thesis. Their dedication to shaping future paleontology students gave me the knowledge and confidence I needed to pursue my passion. Their guidance allowed me to see what research in the fossil record is really like, and put me on a path to pursue these studies in the future. Dr. Kelley has been extremely supportive of this project, acting as a committee member and giving feedback on this thesis and my project in general. I have worked with Dr. Dietl all along the east coast collecting samples from the field and museums. His knowledge of the Coastal Plain and fossil organisms has been indispensible. Without Christy Visaggi’s help in areas from designing this project to giving directions around Wilmington, North Carolina, this project would not have been completed. Her help and friendship was essential to my success last summer.
I would also like to thank Roger Portell and Dr. Lauck “Buck” Ward for their assistance collecting samples from their fantastic collections at the Florida Museum of
Natural History and the Virginia Museum of Natural History. These collections were critical to the completion of this project. In addition I would like to thank the Staff at the
Paleontological Research Institute for their help collecting samples from the PRI collections.
This research was supported in part by a Howard Hughes Medical Institute grant through the Undergraduate Biological Sciences Education Program to the College of
William & Mary, an Undergraduate Research Grant from the Southeastern Section of
The Geological Society of America, and the SWFFC Brayfield Award.
68 A huge thank you goes to the NSF REU in Biodiversity students from 2009,
Stephen Durham, Pedro Monarrez, Elizabeth Graybill, Jessica Lambert, Jennifer Toledo-
Rivera, Kevin Selders, Brad Parnell, and Josh Poole. From collecting snails in the middle of a thunderstorm, to dollar tacos, they never let me down. My geology and paleo lab mates, Kate McClure, Karin Ohman, Sean Moran, Christina Byrd, Amy Rarig, Alex
Olaya, Claire Still, and Lydia Cuker helped enormously with encouragement, suggestions, and feedback on ideas.
Finally I would like to thank my friends who have been so supportive and accepting of the reasons I never see them, and my family who has helped me pursue this passion to graduate school next year.
Thank you all so much.
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72 Appendix
Rawdatafromeachlocalityispresentedinthisappendix.MeasurementsforLength,
Height,Thickness,OuterDrillholeDiameter,andInnerDrillHoleDiameterarerecordedin
millimeters.HoleSectorcorrespondstoDrillHoleSectorasismeasuredbytheprocedure
describedinKelley,1991.NumbersintheNotessectionindicatetheangleofdrillholes.
Locationinformationisincludednexttothelocalityname.
73 Locality OldDock DateCollected 5/18/2001 SouthwestofColumbus,NC Formation LowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External ODͲ1 Neverita duplicata 63.89 49.61 1.36 ODͲ2 Neverita duplicata 28.59 26.74 0.74 ODͲ3 Euspira heros 19.16 18.65 0.62 74 ODͲ4 Euspira heros 21.88 21.68 0.72 ODͲ5 Euspira heros 56.8 58.52 1.74 ODͲ6 Neverita duplicata 44.24 36.15 0.96 ODͲ7 Euspira heros 24.15 24.3 0.73 3.44 1.73 2 168 ODͲ8 Neverita duplicata 40.45 38.1 0.98 ODͲ9 Neverita duplicata 33.12 29.24 0.68 ODͲ10 Neverita duplicata 34.33 28.55 0.9 ODͲ11 Neverita duplicata 20.84 14.47 0.76 ODͲ12 Neverita duplicata 20.47 18.61 0.74 ODͲ13 Neverita duplicata 80.5 73.92 1.35 ODͲ14 Neverita duplicata 64.5 51.05 1.28 ODͲ15 Euspira heros 21.25 21.77 0.69 ODͲ16 Neverita duplicata 30.87 25.28 0.66 ODͲ17 Euspira heros 8.19 8.72 0.39 ODͲ18 Neverita duplicata 45.05 36.25 0.79 ODͲ19 Euspira heros 33.9 40.9 1.15 ODͲ20 Neverita duplicata 24.08 19.77 0.68 ODͲ21 Neverita duplicata 28.95 22.76 1.04 ODͲ22 Neverita duplicata 47.2 42.67 1.36 ODͲ23 Neverita duplicata 15.05 11.14 0.37 75 Locality LumberRiver DateCollected NearLumberton,NC Formation Duplin mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External LRͲ1 Euspira heros 9.07 9.2 0.34 LRͲ2 Euspira heros 9.12 9.69 0.34 LRͲ3 Euspira heros 9.18 8.21 0.4
76 LRͲ4 Euspira heros 10.62 10.84 0.54 LRͲ5 Euspira heros 11.41 11.12 0.38 LRͲ6 Euspira heros 11.7 11.96 0.42 LRͲ7 Naticarius canrena 13.11 13.08 0.96 4.21 2.96 3 239 LRͲ8 Naticarius canrena 13.45 12.95 0.65 LRͲ9 Euspira heros 13.65 13.77 0.44 2.22 1.27 4 227 LRͲ10 Naticarius canrena 13.73 13.9 0.74 LRͲ11 Naticarius canrena 13.81 12.89 0.56 2.4 1.61 3 206 LRͲ12 Euspira heros 13.98 14.97 0.56 LRͲ13 Euspira heros 15.27 14.35 0.25 LRͲ14 Euspira heros 15.35 15.36 0.35 LRͲ15 Naticarius canrena 15.65 15.19 0.49 LRͲ16 Naticarius canrena 16.52 16.74 0.61 LRͲ17 Euspira heros 18.31 17.9 0.91 LRͲ18 Naticarius canrena 18.38 16.37 0.49 LRͲ19 Sinum 18.71 10.36 0.44 LRͲ20 Naticarius canrena 20.46 19.58 0.71 LRͲ21 Naticarius canrena 20.47 20.63 0.59 LRͲ22 Naticarius canrena 21.29 21.04 0.52 4.11 2.64 2 145 LRͲ23 Neverita duplicata 21.3 16.42 0.68 LRͲ24 Naticarius canrena 21.38 20.19 0.72 LRͲ25 Naticarius canrena 21.78 20.25 0.7 LRͲ26 Euspira heros 22.12 24.97 0.89 LRͲ27 Naticarius canrena 22.29 21.58 0.53 LRͲ28 Euspira heros 22.42 21.78 0.84 LRͲ29 Euspira heros 22.52 24.09 0.29 LRͲ30 Naticarius canrena 22.75 23.4 0.9 LRͲ31 Euspira heros 22.98 23.3 0.52 LRͲ32 Naticarius canrena 23.58 21.74 0.77 3.25 1.96 1 105 LRͲ33 Euspira heros 23.71 20.92 0.52 77 LRͲ34 Naticarius canrena 23.92 22.43 0.38 LRͲ35 Naticarius canrena 24.33 21.9 0.85 LRͲ36 Euspira heros 24.81 25.5 0.35 LRͲ37 Euspira heros 25.24 23.48 0.37 LRͲ38 Naticarius canrena 26.5 27.62 0.78 LRͲ39 Naticarius canrena 26.7 26.53 0.72 LRͲ40 Naticarius canrena 27.35 26.25 0.61 4.61 2.93 2 138 LRͲ41 Euspira heros 28.65 27.07 0.65 LRͲ42 Naticarius canrena 28.71 28.29 0.76 LRͲ43 Euspira heros 29.08 26.75 0.5 LRͲ44 Naticarius canrena 29.67 28.76 0.66 LRͲ45 Naticarius canrena 29.92 28.72 0.92 LRͲ46 Naticarius canrena 31.5 29.82 1.11 LRͲ47 Naticarius canrena 31.59 29.67 0.83 LRͲ48 Naticarius canrena 33.19 34.41 0.75 LRͲ49 Naticarius canrena 33.37 29.42 0.7 LRͲ50 Naticarius canrena 33.96 34.63 1.23 LRͲ51 Naticarius canrena 34.04 34.81 0.97 4.59 3.44 4 214 LRͲ52 Euspira heros 37.08 40.74 0.99 LRͲ53 Naticarius canrena 40.83 38.03 0.96 78 Locality RegisterQuarry DateCollected UNCWREUSummer08 NearColumbus,NC Formation lowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External RQͲ1 Sinum 11.36 8 0.32 RQͲ2 Neverita duplicata 11.98 8.71 0.77 RQͲ3 Neverita duplicata 13.87 10.02 0.53
79 RQͲ4 Naticarius canrena 14.31 14.55 0.7 RQͲ5 Naticarius canrena 14.85 13.86 0.72 RQͲ6 Neverita duplicata 15.38 11.01 0.51 RQͲ7 Neverita duplicata 15.74 11.57 0.86 RQͲ8 Euspira heros 16.32 17.56 0.63 RQͲ9 Euspira heros 16.64 17.55 0.23 RQͲ10 Neverita duplicata 16.82 12.8 0.62 RQͲ11 Naticarius canrena 17.14 15.99 0.85 RQͲ12 Neverita duplicata 17.28 12.13 0.43 RQͲ13 Euspira heros 17.28 16.97 0.44 RQͲ14 Naticarius canrena 18.01 18.12 0.75 RQͲ15 Naticarius canrena 18.07 17.39 0.63 RQͲ16 Neverita duplicata 18.76 14.19 0.92 RQͲ17 Naticarius canrena 18.9 18.71 0.88 RQͲ18 Neverita duplicata 19.15 14.27 0.47 5.6 2.68 2 151 RQͲ19 Neverita duplicata 19.43 14.78 0.53 RQͲ20 Neverita duplicata 19.46 15.81 0.45 4.06 1.61 4 230 RQͲ21 Neverita duplicata 19.52 14.15 0.58 4.92 2.91 2 155 RQͲ22 Euspira heros 19.93 19.64 0.47 RQͲ23 Euspira heros 19.94 22.92 0.54 RQͲ24 Neverita duplicata 20.69 15.86 0.54 RQͲ25 Neverita duplicata 20.8 15.66 0.58 RQͲ26 Neverita duplicata 20.99 16.59 0.65 RQͲ27 Neverita duplicata 21.14 16.04 0.37 4.15 2.07 4 202 RQͲ28 Neverita duplicata 21.18 15.33 0.51 RQͲ29 Neverita duplicata 21.44 15.58 0.67 RQͲ30 Neverita duplicata 22.02 16.55 0.44 4.66 3.47 2 161 RQͲ31 Neverita duplicata 22.22 16.99 0.91 RQͲ32 Neverita duplicata 22.25 16.79 0.67 4.22 2.31 2 144 RQͲ33 Euspira heros 22.79 22.92 0.81 3.93 2.65 2 178 80 RQͲ34 Neverita duplicata 22.92 18.47 0.74 RQͲ35 Neverita duplicata 23.11 17.93 0.81 RQͲ36 Neverita duplicata 23.36 19.23 0.65 RQͲ37 Neverita duplicata 23.61 22.15 0.81 RQͲ38 Neverita duplicata 24.18 18.01 0.71 RQͲ39 Neverita duplicata 24.24 20.94 0.67 RQͲ40 Neverita duplicata 24.83 21.82 0.47 4.42 2.63 4 217 RQͲ41 Naticarius canrena 25.42 26.26 0.84 6.1 3.48 3 192 RQͲ42 Neverita duplicata 26.08 24.02 1.12 RQͲ43 Euspira heros 26.11 28.48 0.51 RQͲ44 Neverita duplicata 26.3 22.9 0.94 RQͲ45 Euspira heros 26.43 26.76 0.84 RQͲ46 Neverita duplicata 27.73 21.85 0.79 RQͲ47 Neverita duplicata 27.85 21.92 0.93 RQͲ48 Neverita duplicata 27.95 21.72 7.2 RQͲ49 Neverita duplicata 28.45 21.68 0.75 RQͲ50 Neverita duplicata 32.97 26.08 0.82 RQͲ51 Neverita duplicata 33.36 31.72 0.87 RQͲ52 Neverita duplicata 33.68 27.3 1.28 RQͲ53 Neverita duplicata 33.81 29.29 1.49 RQͲ54 Neverita duplicata 34.83 28.04 1.28 RQͲ55 Neverita duplicata 35.5 28.48 0.78 RQͲ56 Neverita duplicata 35.67 32.6 1.2 RQͲ57 Naticarius canrena 41.59 40.44 0.67 RQͲ58 Neverita duplicata 41.67 37.86 0.76 RQͲ59 Euspira heros 44.34 45.76 0.56 RQͲ60 Neverita duplicata 47.23 35.72 0.93 RQͲ61 Neverita duplicata 49.3 40.05 1.31 8.17 4.68 2 139 RQͲ62 Neverita duplicata 51.54 45.77 1.03 RQͲ63 Neverita duplicata 51.96 42.74 0.95 81 RQͲ64 Neverita duplicata 82.98 69.33 1.19 Locality Chuckatuck DateCollected Museum(BuckWardVMNH) Near36°52'N76°34'W Formation Yorktown mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External CKͲ1 Neverita duplicata 40.09 35.87 0.87 CKͲ2 Neverita duplicata 25.23 19.73 0.42 CKͲ3 Euspira heros 29.97 31.67 0.63
82 CKͲ4 Neverita duplicata 30.51 25.54 0.57 CKͲ5 Neverita duplicata 31.59 24.78 0.83 CKͲ6 Euspira heros 29.25 30.62 0.8 CKͲ7 Neverita duplicata 27.46 22.36 0.43 CKͲ8 Euspira heros 18.57 19 0.56 4.22 2.18 1 CKͲ9 Euspira heros 60.19 59.86 1.17 CKͲ10 Neverita duplicata 46.96 39.78 1.02 CKͲ11 Neverita duplicata 29.74 24.34 0.65 CKͲ12 Euspira heros 31.45 30.76 1.19 CKͲ13 Euspira heros 30.32 30.75 0.78 CKͲ14 Euspira heros 29.85 31.29 0.95 CKͲ15 Euspira heros 27.16 26.98 0.55 CKͲ16 Sinum maculatum 17.52 16.92 0.63 CKͲ17 Euspira heros 29.45 29.41 0.51 CKͲ18 Neverita duplicata 27.88 22.96 0.44 CKͲ19 Neverita duplicata 39.31 30.71 0.9 CKͲ20 Neverita duplicata 31.17 24.06 0.54 CKͲ21 Neverita duplicata 46.99 38.11 0.95 CKͲ22 Euspira heros 20.33 22.06 0.71 CKͲ23 Neverita duplicata 21.08 16.44 0.55 CKͲ24 Euspira heros 27.5 29.69 0.79 CKͲ25 Neverita duplicata 36.99 29.53 0.79 CKͲ26 Euspira heros 17.53 18.95 0.63 4.43 3.47 2 CKͲ27 Neverita duplicata 16.37 12.03 0.33 3.6 2.22 2 CKͲ28 Euspira heros 20.28 21.39 0.48 CKͲ29 Neverita duplicata 39.26 31.09 0.57 CKͲ30 Neverita duplicata 28.4 22.33 0.84 CKͲ31 Euspira heros 28.03 28.13 0.43 4.6 2.57 1 CKͲ32 Neverita duplicata 24.5 20.69 0.48 CKͲ33 Euspira heros 21.82 22.1 0.42 83 CKͲ34 Neverita duplicata 31.36 25.46 0.74 CKͲ35 Neverita duplicata 48.54 37.63 1.25 CKͲ36 Neverita duplicata 33.58 27.55 0.68 CKͲ37 Neverita duplicata 21.57 15.73 0.35 CKͲ38 Neverita duplicata 32.51 24.54 0.64 CKͲ39 Neverita duplicata 22.61 16.75 0.51 CKͲ40 Neverita duplicata 20.76 16.62 0.48 CKͲ41 Euspira heros 58.16 62.2 1.31 CKͲ42 Neverita duplicata 16.2 12.18 0.26 CKͲ43 Euspira heros 20.85 19.79 0.69 4.53 2.28 2 CKͲ44 Neverita duplicata 26.95 21.32 0.52 CKͲ45 Euspira heros 17.12 16.45 0.52 4.7 2.59 4 CKͲ46 Euspira heros 18.66 20.25 0.55 CKͲ47 Neverita duplicata 21.71 18.51 0.56 CKͲ48 Euspira heros 25.31 25.84 0.49 CKͲ49 Euspira heros 13.18 13.15 0.37 CKͲ50 Euspira heros 28.54 30.94 0.53 CKͲ51 Neverita duplicata 18.59 13.93 0.35 CKͲ52 Euspira heros 18.64 19.39 0.55 4.48 2.84 2 CKͲ53 Euspira heros 18.36 20.19 0.45 CKͲ54 Euspira heros 19.44 18.12 0.77 CKͲ55 Euspira heros 19.65 11.6 0.27 2.55 0.95 3 CKͲ56 Euspira heros 23.19 30.23 0.53 CKͲ57 Lunatia perspectiva 60.27 60.67 0.79 CKͲ58 Lunatia perspectiva 44.33 47.67 0.68 CKͲ59 Lunatia perspectiva 49.33 50.28 0.76 CKͲ60 Lunatia perspectiva 29.6 29.25 0.45 CKͲ61 Lunatia perspectiva 52.93 56.24 0.7 CKͲ62 Neverita duplicata 38.18 26.89 0.6 CKͲ63 Lunatia perspectiva 13.34 15.91 0.48 84 CKͲ64 Lunatia perspectiva 12.36 14.15 0.53 CKͲ65 Lunatia perspectiva 9.06 10.16 0.22 CKͲ66 Lunatia perspectiva 14.08 16.52 0.35 CKͲ67 Lunatia perspectiva 9.36 11.43 0.31 CKͲ68 Lunatia perspectiva 10.69 13.4 0.5 CKͲ69 Lunatia perspectiva 7.85 9.59 0.31 CKͲ70 Lunatia perspectiva 10.11 12.08 0.3 CKͲ71 Lunatia perspectiva 13.17 15.49 0.29 CKͲ72 Lunatia perspectiva 8.41 9.76 0.38 CKͲ73 Lunatia perspectiva 9.26 10.87 0.42 CKͲ74 Lunatia interna 12.3 13.71 0.37 CKͲ75 Lunatia interna 16.99 19.79 0.44 CKͲ76 Lunatia interna 16.85 18.93 0.5 CKͲ77 Lunatia interna 13.11 15.06 0.24 CKͲ78 Lunatia interna 8.95 10.69 0.36 CKͲ79 Lunatia interna 16.69 19.86 0.79 CKͲ80 Lunatia interna 13.57 14.24 0.43 CKͲ81 Lunatia interna 12.41 13.93 0.49 CKͲ82 Lunatia interna 10.85 12.57 0.39 CKͲ83 Lunatia interna 9.39 9.79 0.31 CKͲ84 Lunatia interna 8.05 9.23 0.28 CKͲ85 Lunatia interna 11.46 12.39 0.55 CKͲ86 Lunatia interna 7.51 8.45 0.19 CKͲ87 Lunatia interna 9.41 11.07 0.53 CKͲ88 Lunatia interna 9.07 10.48 0.29 CKͲ89 Lunatia interna 9.38 11.15 0.25 CKͲ90 Tectonatica pusilla 5.67 6.13 0.21 CKͲ91 Tectonatica pusilla 6.4 7.28 0.37 CKͲ92 Tectonatica pusilla 5.16 6 0.19 85 Locality Rice'sPit DateCollected Museum(BuckWardVMNH) NearHampton,VA Formation Yorktown mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External RPͲ1 Euspira heros 30.2 30.89 0.57 4.82 3.02 5 RPͲ2 Euspira heros 32.27 34.14 0.47 RPͲ3 Euspira heros 28.12 29.43 0.85
86 RPͲ4 Lunatia interna 15.55 17.89 0.55 RPͲ5 Lunatia perspectiva 17.39 17.17 0.21 RPͲ6 Euspira heros 29.76 25.64 0.65 4.34 2.63 3 drillonbottom RPͲ7 Lunatia interna 17.91 19.36 0.57 RPͲ8 Lunatia interna 17.07 19.08 0.47 RPͲ9 Euspira heros 17.22 16.83 0.46 3.95 2.53 2 RPͲ10 Euspira heros 28.39 27.18 0.71 RPͲ11 Lunatia perspectiva 43.83 46.55 0.58 RPͲ12 Euspira heros 36.64 38.34 0.58 RPͲ13 Lunatia interna 15.61 15.49 0.61 3.23 1.83 3 RPͲ14 Lunatia perspectiva 17.67 20.77 0.3 RPͲ15 Lunatia interna 15.66 15.5 0.61 4.65 2.61 5 RPͲ16 Lunatia interna 17.69 19.57 0.52 RPͲ17 Lunatia interna 14.91 15.54 0.73 RPͲ18 Euspira heros 17.62 17.81 0.47 2.77 1.84 2 RPͲ19 Lunatia interna 16.36 18.03 0.64 RPͲ20 Lunatia interna 17.26 19.33 0.47 3.38 1.99 2 RPͲ21 Lunatia interna 15.14 16.86 0.5 3.98 1.94 1 RPͲ22 Lunatia interna 16.49 16.97 0.51 4.01 2.93 3 RPͲ23 Lunatia interna 18.69 19.6 0.4 RPͲ24 Lunatia perspectiva 11.86 13.02 0.2 RPͲ25 Lunatia interna 15.28 16.13 0.38 5.16 3.19 2 RPͲ26 Euspira heros 26.6 24.07 0.76 4.36 2.7 2 RPͲ27 Lunatia perspectiva 9.84 10.91 0.46 RPͲ28 Lunatia interna 17.85 19.66 0.55 3.26 1.9 2 RPͲ29 Lunatia interna 14.29 14.81 0.44 3.22 1.63 3 RPͲ30 Euspira heros 23.31 22.77 0.81 4.81 2.39 4 RPͲ31 Lunatia perspectiva 12.43 12.5 0.24 RPͲ32 Lunatia interna 13.37 13.43 0.34 RPͲ33 Lunatia perspectiva 18.81 21.14 0.91 87 RPͲ34 Lunatia interna 15.25 15.81 0.46 RPͲ35 Lunatia interna 11.1 11.35 0.35 RPͲ36 Lunatia perspectiva 16.87 18.68 0.48 RPͲ37 Lunatia interna 15.62 17.16 0.6 RPͲ38 Lunatia interna 16.12 17.07 0.44 RPͲ39 Lunatia interna 15.6 16.7 0.46 RPͲ40 Lunatia interna 16.71 16.78 0.38 4.15 2.59 2 RPͲ41 Lunatia interna 13.3 14.44 0.35 RPͲ42 Lunatia perspectiva 13.46 14.16 0.48 RPͲ43 Lunatia perspectiva 15.15 16.35 0.55 RPͲ44 Lunatia interna 13.11 13.72 0.29 RPͲ45 Lunatia interna 13.35 13.16 0.45 RPͲ46 Lunatia interna 11.07 11.28 0.28 RPͲ47 Lunatia perspectiva 15.95 15.92 0.49 RPͲ48 Lunatia interna 10.13 10.77 0.35 RPͲ49 Lunatia perspectiva 8.75 9.5 0.35 RPͲ50 Lunatia interna 20.01 21.61 0.63 RPͲ51 Lunatia interna 10.09 10.25 0.3 RPͲ52 Lunatia interna 8.73 9.11 0.44 RPͲ53 Lunatia interna 9 9.27 0.32 2.83 1.65 3 RPͲ54 Neverita duplicata 9.09 7.49 0.35 RPͲ55 Neverita duplicata 8.49 7.15 0.75 1.12 0.88 3 RPͲ56 Neverita duplicata 11.59 8.65 0.33 RPͲ57 Euspira heros 45.24 47.25 0.61 RPͲ58 Euspira heros 32.96 31.76 0.67 4.92 3.14 2 RPͲ59 Euspira heros 31.99 29.46 0.48 RPͲ60 Lunatia perspectiva 14.9 16.56 0.67 RPͲ61 Euspira heros 25.07 25.2 0.46 RPͲ62 Lunatia perspectiva 17.67 15.99 0.38 RPͲ63 Lunatia perspectiva 16.56 18.23 0.51 3.45 2.21 1 88 RPͲ64 Lunatia perspectiva 9.09 10.03 0.18 1.63 0.98 2 RPͲ65 Lunatia perspectiva 13.71 13.96 0.31 RPͲ66 Polinices sp. 11.68 13.51 0.53 RPͲ67 Lunatia interna 12.04 13.54 0.43 RPͲ68 Polinices sp. 9.66 10.87 0.28 RPͲ69 Polinices sp. 10.18 11.17 0.36 RPͲ70 Polinices sp. 7.34 8.44 0.29 RPͲ71 Polinices sp. 10.25 11.98 0.31 RPͲ72 Polinices sp. 7.54 8.38 0.3 RPͲ73 Lunatia interna 12.37 13.51 0.37 Locality Brant'sPit DateCollected Museum(BuckWardVMNH) Exactlocationunknown,northofCapeHatteras,NC Formation Yorktown mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External BPͲ1 Euspira heros 23.33 20.42 0.23 4.56 2.94 5 BPͲ2 Lunatia interna 17.79 16.74 0.37 BPͲ3 Lunatia interna 15.94 17.33 0.36
89 BPͲ4 Lunatia interna 16.9 18.12 0.67 BPͲ5 Lunatia interna 18.16 19.64 0.55 3.99 2.12 1 BPͲ6 Sinum chesapeakensis 17.42 17.99 0.39 BPͲ7 Euspira heros 16.12 16.36 0.42 4.13 2.69 2 BPͲ8 Lunatia interna 15.75 15.52 0.49 4.64 2.43 1 BPͲ9 Lunatia interna 14.8 15.99 0.42 3.33 1.59 1 BPͲ10 Lunatia interna 16.98 17.49 0.42 BPͲ11 Lunatia interna 16.24 16.85 0.62 3.87 1.99 1 BPͲ12 Lunatia interna 15.2 16.24 0.41 3.84 2.23 2 BPͲ13 Lunatia interna 14.02 15.48 0.44 3.25 1.84 2 BPͲ14 Lunatia interna 17.7 18.13 0.65 3.55 2.2 2 BPͲ15 Lunatia interna 15.67 16.09 0.57 3.41 1.92 2 BPͲ16 Lunatia interna 14.96 16.07 0.68 BPͲ17 Lunatia interna 13.56 14.22 0.36 1.83 1.15 4 BPͲ18 Lunatia interna 13.61 13.91 0.26 3.16 1.56 2 BPͲ19 Lunatia interna 18.46 20.03 0.69 4.68 2.58 1 BPͲ20 Sinum chesapeakensis 18.14 18.93 0.5 BPͲ21 Lunatia interna 16.64 17.69 0.85 BPͲ22 Sinum perspectivium 15.77 10.03 0.34 BPͲ23 Sinum chesapeakensis 16.56 15.61 0.56 BPͲ24 Sinum chesapeakensis 10.06 9.64 0.29 BPͲ25 Lunatia interna 8.69 9.11 0.28 BPͲ26 Lunatia interna 10.12 9.9 0.28 BPͲ27 Lunatia perspectiva 9.62 10.62 0.23 2.63 1.43 2 BPͲ28 Lunatia interna 14.32 14.79 0.3 BPͲ29 Lunatia interna 21.74 22.97 0.52 4.72 2.68 1 BPͲ30 Lunatia interna 15.89 16.22 0.47 5.29 2.91 1 BPͲ31 Lunatia interna 18.55 18.79 0.53 4.31 3.11 2 BPͲ32 Lunatia interna 15.19 15.73 0.34 2.36 1.21 2 BPͲ33 Lunatia interna 10.23 10.95 0.32 4.17 2.74 5 90 BPͲ34 Lunatia interna 16.38 15.47 0.38 BPͲ35 Lunatia interna 14.26 15.21 0.43 4.42 2.61 1 BPͲ36 Sinum chesapeakensis 12.93 12.47 0.37 BPͲ37 Lunatia interna 12.36 13.04 0.26 2.48 1.47 2 BPͲ38 Sinum chesapeakensis 10.19 10.73 0.43 BPͲ39 Lunatia perspectiva 14.07 14.94 0.41 BPͲ40 Lunatia perspectiva 12.33 13.05 0.33 BPͲ41 Lunatia interna 16.27 16.68 0.4 4.95 2.57 2 BPͲ42 Lunatia interna 10.2 10.93 0.27 2.6 1.6 1 BPͲ43 Polinices sp. 8.75 10.05 0.41 3.13 1.78 2 BPͲ44 Polinices sp. 9.46 11.1 0.35 BPͲ45 Polinices sp. 10.18 10.78 0.3 2.87 1.7 1 BPͲ46 Lunatia perspectiva 9.5 10.06 0.3 BPͲ47 Sinum chesapeakensis 14.96 14.31 0.48 BPͲ48 Polinices sp. 9.97 11.08 0.27 2.99 1.72 2 BPͲ49 Lunatia interna 14.21 13.84 0.43 BPͲ50 Sinum chesapeakensis 7.45 7.85 0.19 BPͲ51 Sinum chesapeakensis 6.56 6.44 0.15 BPͲ52 Lunatia interna 9.98 10.43 0.26 3 1.58 2 BPͲ53 Lunatia interna 10.47 11.08 0.69 4.35 2.38 6 BPͲ54 Polinices sp. 8.32 9.32 0.34 BPͲ55 Lunatia interna 10.12 10.28 0.3 1.67 0.68 2 BPͲ56 Polinices sp. 9.98 11.38 0.35 BPͲ57 Polinices sp. 8.98 10.66 0.45 BPͲ58 Lunatia interna 9.03 9.25 0.3 BPͲ59 Polinices sp. 8.26 9.94 0.3 BPͲ60 Polinices sp. 8.25 8.97 0.33 BPͲ61 Lunatia interna 8.64 9.81 0.34 BPͲ62 Lunatia interna 12.36 13.08 0.34 BPͲ63 Polinices sp. 8.15 9.11 0.27 91 BPͲ64 Polinices sp. 4.89 6.14 0.2 BPͲ65 Lunatia interna 7.32 7.97 0.22 BPͲ66 Sinum chesapeakensis 7.56 7.99 0.24 BPͲ67 Euspira heros 7.24 7.39 0.31 BPͲ68 Lunatia interna 6.73 7.11 0.22 BPͲ69 Lunatia interna 7.09 7.84 0.29 BPͲ70 Sinum chesapeakensis 6.45 6.77 0.2 BPͲ71 Polinices sp. 7.2 7.92 0.37 BPͲ72 Tectonaticapusilla 3.37 4.28 0.22 BPͲ73 Polinices sp. 6.57 7.52 0.45 BPͲ74 Polinices sp. 6.31 7.23 0.22 BPͲ75 Polinices sp. 5.93 6.2 0.24 2.07 0.9 3 BPͲ76 Tectonaticapusilla 5.27 5.96 0.3 2.81 1.88 4 BPͲ77 Euspira heros 5.63 5.95 0.47 BPͲ78 Lunatia interna 5.92 6.42 0.24 BPͲ79 Tectonaticapusilla 4.25 4.61 0.23 2.22 1.44 3 BPͲ80 Lunatia interna 3.32 3.64 0.13 1.74 1.16 4 BPͲ81 Lunatia perspectiva 4.45 5.04 0.33 BPͲ82 Tectonaticapusilla 4.2 4.94 0.28 BPͲ83 Tectonaticapusilla 4.43 4.96 0.25 2.27 1.59 1 BPͲ84 Sinum chesapeakensis 5.96 5.01 0.14 BPͲ85 Sinum chesapeakensis 4.61 4.78 0.2 BPͲ86 Tectonaticapusilla 3.59 4.27 0.13 2.28 1.76 4 BPͲ87 Tectonaticapusilla 4.31 4.83 0.25 BPͲ88 Lunatia interna 4.49 4.6 0.22 BPͲ89 Tectonaticapusilla 3.37 4 0.11 BPͲ90 Tectonaticapusilla 2.93 3.27 0.1 BPͲ91 Euspira heros 2.16 1.99 0.13 BPͲ92 Euspira heros 3.74 3.54 0.14 BPͲ93 Tectonaticapusilla 2.96 3.25 0.16 92 BPͲ94 Lunatia interna 3.68 3.7 0.42 BPͲ95 Euspira heros 3.12 3.37 0.24 BPͲ96 Tectonaticapusilla 2.73 2.95 0.23 BPͲ97 Lunatia interna 3.01 3.58 0.48 BPͲ98 Euspira heros 3.06 2.98 0.12 BPͲ99 Lunatia interna 3.12 3.54 0.23 Locality Elizabethtown,NC DateCollected Museum(BuckWardVMNH) NearElizabethtown,NC Formation ChowanRiver mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External EBTͲ1 Neverita duplicata 27.57 24.29 0.67 EBTͲ2 Neverita duplicata 26.52 22.91 0.7 3.36 2.44 3 EBTͲ3 Neverita duplicata 19.6 17.81 0.64
93 EBTͲ4 Neverita duplicata 20.54 17.62 0.49 4.2 2.79 4 EBTͲ5 Neverita duplicata 18.35 13.9 0.58 EBTͲ6 Neverita duplicata 18.47 14.11 0.96 EBTͲ7 Neverita duplicata 20.43 17.42 0.77 EBTͲ8 Neverita duplicata 23.49 19.67 0.44 EBTͲ9 Neverita duplicata 20.24 17.19 0.84 EBTͲ10 Neverita duplicata 13.17 19.12 0.69 EBTͲ11 Lunatia perspectiva 28.38 32.25 0.73 EBTͲ12 Neverita duplicata 10.64 9.11 0.45 EBTͲ13 Neverita duplicata 32.8 29.9 0.6 EBTͲ14 Neverita duplicata 17.45 12.55 0.79 EBTͲ15 Neverita duplicata 11.34 9.51 0.63 2.57 1.55 7 EBTͲ16 Neverita duplicata 25 19.74 0.66 EBTͲ17 Neverita duplicata 21.93 19.33 0.4 EBTͲ18 Neverita duplicata 22.46 18.08 0.59 EBTͲ19 Neverita duplicata 23.24 23.7 1.06 EBTͲ20 Neverita duplicata 24.47 23.04 0.75 Locality GomezPit DateCollected Museum(BuckWardVMNH) Near36°01'N76°43'W Formation ChowanRiver mm mm mm mm mm Diameter Diameter Hole Hole Sector Notes Internal Hole Specimen Genus Species Length Height Thickness External GPͲ1 Euspira heros 24.93 30.23 0.42 GPͲ2 Neverita duplicata 31.84 31.49 0.69 GPͲ3 Euspira heros 39.45 33.86 0.79
94 GPͲ4 Euspira heros 47.12 51.8 0.6 GPͲ5 Euspira heros 28.15 28.78 0.52 GPͲ6 Neverita duplicata 19.97 15.81 0.81 GPͲ7 Euspira heros 18.55 18.54 0.23 GPͲ8 Neverita duplicata 19.14 23.57 0.64 GPͲ9 Neverita duplicata 28.3 25.38 0.63 GPͲ10 Neverita duplicata 16.7 16.2 0.67 GPͲ11 Neverita duplicata 19.49 14.6 0.51 GPͲ12 Neverita duplicata 40.73 42.04 0.49 GPͲ13 Neverita duplicata 49.91 45.6 1.15 GPͲ14 Euspira heros 44.8 44.35 0.4 GPͲ15 Euspira heros 25.93 26.84 0.61 GPͲ16 Neverita duplicata 37.82 35.63 0.52 GPͲ17 Euspira heros 22.88 23.13 0.48 GPͲ18 Euspira heros 56.75 60.33 2.2 GPͲ19 Neverita duplicata 44.21 42.93 1.61 GPͲ20 Neverita duplicata 45.12 39.45 1.11 GPͲ21 Euspira heros 22.33 21.74 0.45 GPͲ22 Euspira heros 35.7 38.39 0.62 GPͲ23 Neverita duplicata 42.06 10.13 0.79 GPͲ24 Euspira heros 32.95 26.35 1.19 GPͲ25 Neverita duplicata 42.9 40.31 0.45 GPͲ26 Neverita duplicata 44.55 41.44 0.97 GPͲ27 Euspira heros 32.33 34.69 0.51 GPͲ28 Neverita duplicata 40.43 41.54 0.68 14.16 8.06 4 GPͲ29 Neverita duplicata 33.21 25.03 0.69 GPͲ30 Neverita duplicata 31.7 30.17 0.65 GPͲ31 Neverita duplicata 40.03 32.53 1.07 GPͲ32 Neverita duplicata 43.88 36.62 0.85 GPͲ33 Neverita duplicata 41.62 38.29 1.01 95 GPͲ34 Neverita duplicata 31.54 29.34 0.9 GPͲ35 Euspira heros 34.8 37.78 0.49 Locality Acme,NC1446 DateCollected Museum(PRI) SouthwestofColumbus,NC Formation LowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External AC2Ͳ1 Lunatia interna 47.48 45.47 0.98 AC2Ͳ2 Tectonatica pusilla 6.12 6.18 0.27 AC2Ͳ3 Neverita duplicata 26.77 20.99 0.65 96 AC2Ͳ4 Neverita duplicata 20.53 15.72 0.55 AC2Ͳ5 Neverita duplicata 14.81 11.47 0.47 AC2Ͳ6 Naticarius canrena 12.2 11.84 0.58 AC2Ͳ7 Neverita duplicata 14.86 11.92 0.38 3.58 1.59 2 AC2Ͳ8 Neverita duplicata 9.67 7.66 0.41 AC2Ͳ9 Tectonatica pusilla 4.33 4.94 0.23 AC2Ͳ10 Tectonatica pusilla 6.72 7.47 0.27 AC2Ͳ11 Tectonatica pusilla 5.57 6.05 0.28 AC2Ͳ12 Tectonatica pusilla 3.88 4.65 0.29 AC2Ͳ13 Sinum chesapeakens 10.52 6.92 0.28 AC2Ͳ14 Tectonatica pusilla 5.93 6.5 0.26 AC2Ͳ15 Tectonatica pusilla 5.91 6.49 0.29 AC2Ͳ16 Sinum chesapeakens 7.66 5.72 0.25 AC2Ͳ17 Tectonatica pusilla 5.4 5.86 0.27 AC2Ͳ18 Tectonatica pusilla 5.55 5.98 0.22 AC2Ͳ19 Tectonatica pusilla 4.8 5.6 0.26 AC2Ͳ20 Tectonatica pusilla 4.96 5.2 0.32 AC2Ͳ21 Tectonatica pusilla 5.02 5.52 0.35 AC2Ͳ22 Tectonatica pusilla 4.29 4.55 0.26 AC2Ͳ23 Tectonatica pusilla 4.45 5.28 0.24 AC2Ͳ24 Tectonatica pusilla 5.92 5.74 0.34 AC2Ͳ25 Tectonatica pusilla 5.27 5.25 0.24 2.69 1.99 2 AC2Ͳ26 Tectonatica pusilla 5.24 5.84 0.42 AC2Ͳ27 Tectonatica pusilla 5.29 5.52 0.35 AC2Ͳ28 Tectonatica pusilla 5.15 5.97 0.24 AC2Ͳ29 Tectonatica pusilla 4.49 5.19 0.28 AC2Ͳ30 Tectonatica pusilla 5.4 6.24 0.23 AC2Ͳ31 Tectonatica pusilla 4.72 5.36 0.29 AC2Ͳ32 Tectonatica pusilla 4.31 4.38 0.17 97 AC2Ͳ33 Tectonatica pusilla 5.08 5.81 0.29 AC2Ͳ34 Tectonatica pusilla 4.12 4.4 0.23 AC2Ͳ35 Euspira heros 4.24 4.22 0.22 AC2Ͳ36 Tectonatica pusilla 4.79 5.44 0.24 AC2Ͳ37 Tectonatica pusilla 4.23 4.61 0.17 AC2Ͳ38 Tectonatica pusilla 3.85 4.62 0.22 AC2Ͳ39 Tectonatica pusilla 4.25 5.2 0.26 AC2Ͳ40 Tectonatica pusilla 3.42 3.43 0.14 AC2Ͳ41 Tectonatica pusilla 3.64 4.22 0.25 AC2Ͳ42 Lunatia interna 4.07 4.19 0.18 AC2Ͳ43 Tectonatica pusilla 4.27 5.16 0.51 AC2Ͳ44 Tectonatica pusilla 3.1 2.78 0.25 AC2Ͳ45 Tectonatica pusilla 4.18 4.94 0.2 AC2Ͳ46 Tectonatica pusilla 4.13 4.56 0.34 AC2Ͳ47 Euspira heros 32.58 33.7 0.55 4.4 2.05 8 AC2Ͳ48 Euspira heros 23.05 24.99 0.45 4.36 2.43 4 AC2Ͳ49 Euspira heros 26.45 27.59 0.82 AC2Ͳ50 Euspira heros 24.58 24.31 0.73 AC2Ͳ51 Euspira heros 19.22 22.38 0.68 3.43 1.7 4 AC2Ͳ52 Neverita duplicata 24.57 21.6 0.66 AC2Ͳ53 Euspira heros 16.33 19.2 0.44 4.4 2.11 1 AC2Ͳ54 Euspira heros 15.55 16.88 0.35 AC2Ͳ55 Euspira heros 15.3 15.09 0.29 3.86 2.51 1 AC2Ͳ56 Naticarius canrena 18.27 18.79 0.43 AC2Ͳ57 Naticarius canrena 16.48 16.66 0.58 AC2Ͳ58 Naticarius canrena 16.73 16.77 0.55 AC2Ͳ59 Naticarius canrena 14.44 13.52 0.38 AC2Ͳ60 Euspira heros 14.97 16.41 0.48 3.43 2.24 6 AC2Ͳ61 Euspira heros 17.94 17.96 0.43 3.27 1.55 1 98 AC2Ͳ62 Euspira heros 13.7 13.16 0.36 AC2Ͳ63 Euspira heros 14.13 15.21 0.62 AC2Ͳ64 Euspira heros 10.17 10.63 0.32 AC2Ͳ65 Euspira heros 12.71 12.5 0.27 AC2Ͳ66 Lunatia perspectiva 7.13 7.64 0.3 AC2Ͳ67 Euspira heros 13.73 13.56 0.52 AC2Ͳ68 Naticarius canrena 16.42 16.3 0.45 AC2Ͳ69 Naticarius canrena 17.31 16.65 0.61 AC2Ͳ70 Naticarius canrena 15.16 14.08 0.73 AC2Ͳ71 Euspira heros 12.87 12.41 0.3 AC2Ͳ72 Naticarius canrena 19.08 18.76 0.49 AC2Ͳ73 Naticarius canrena 10.98 10.93 0.6 AC2Ͳ74 Naticarius canrena 10.73 9.64 0.39 AC2Ͳ75 Euspira heros 8.93 9.67 0.43 AC2Ͳ76 Euspira heros 8.23 8.56 0.24 AC2Ͳ77 Euspira heros 7.05 7.37 0.24 1.08 0.6 3 Drilledtwice(.90/.50/3 AC2Ͳ78 Euspira heros 5.59 6.38 0.15 2.54 1.91 5 AC2Ͳ79 Euspira heros 7.1 7.55 0.21 AC2Ͳ80 Tectonatica pusilla 4.8 5.5 0.22 AC2Ͳ81 Tectonatica pusilla 6.67 7.25 0.29 AC2Ͳ82 Euspira heros 5.24 5.93 0.38 AC2Ͳ83 Euspira heros 7.22 6.77 0.29 AC2Ͳ84 Euspira heros 6.55 7.1 0.31 AC2Ͳ85 Euspira heros 4.92 5.33 0.16 AC2Ͳ86 Euspira heros 5.85 6.09 0.12 AC2Ͳ87 Euspira heros 5.21 5.72 0.18 AC2Ͳ88 Tectonatica pusilla 5.18 6.12 0.19 AC2Ͳ89 Tectonatica pusilla 3.7 4.17 0.09 AC2Ͳ90 Tectonatica pusilla 5.51 5.82 0.19 99 AC2Ͳ91 Tectonatica pusilla 4.57 5 0.2 AC2Ͳ92 Tectonatica pusilla 4.24 5.15 0.17 AC2Ͳ93 Tectonatica pusilla 4.82 4.87 0.22 AC2Ͳ94 Tectonatica pusilla 4.14 4.77 0.17 AC2Ͳ95 Tectonatica pusilla 3.56 3.78 0.15 AC2Ͳ96 Tectonatica pusilla 3.71 4.31 0.19 AC2Ͳ97 Tectonatica pusilla 3.55 4.03 0.18 AC2Ͳ98 Tectonatica pusilla 3.28 3.75 0.33 AC2Ͳ99 Euspira heros 4.15 4.14 0.11 Locality Acme,NC112 DateCollected Museum(PRI) SouthwestofColumbus,NC Formation LowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External ACͲ1 Naticarius canrena 17.78 17.51 0.58 3.47 1.78 2 ACͲ2 Naticarius canrena 17.8 19.1 0.4 ACͲ3 Naticarius canrena 20.77 21.69 0.47 100 ACͲ4 Naticarius canrena 18.57 18.37 0.42 ACͲ5 Naticarius canrena 16.78 16.92 0.68 ACͲ6 Naticarius canrena 16.58 16.33 0.61 ACͲ7 Naticarius canrena 15.15 15.17 0.77 ACͲ8 Naticarius canrena 19.71 19.37 0.91 ACͲ9 Naticarius canrena 16.49 15.98 0.51 ACͲ10 Naticarius canrena 18.79 20.18 0.5 ACͲ11 Naticarius canrena 20.86 20.76 0.44 ACͲ12 Naticarius canrena 17.29 17.52 0.57 ACͲ13 Naticarius canrena 18.32 17.39 0.73 ACͲ14 Naticarius canrena 20.31 20.33 0.75 ACͲ15 Euspira heros 48.78 49.91 0.89 ACͲ16 Euspira heros 21.08 19.85 0.4 ACͲ17 Neverita duplicata 16.17 12.72 0.76 ACͲ18 Neverita duplicata 27.82 24.33 0.76 ACͲ19 Euspira heros 13.13 13.08 0.33 ACͲ20 Naticarius canrena 16.27 16.43 0.72 ACͲ21 Euspira heros 33.38 34.82 0.58 ACͲ22 Neverita duplicata 51.89 46.79 1.18 ACͲ23 Naticarius canrena 22.18 22.73 0.64 ACͲ24 Euspira heros 8.48 8.84 0.39 ACͲ25 Naticarius canrena 15.13 15.22 0.41 ACͲ26 Tectonatica pusilla 3.59 3.74 0.19 ACͲ27 Euspira heros 17.34 18.05 0.34 2.62 1.48 2 ACͲ28 Euspira heros 21.13 22.47 0.68 ACͲ29 Euspira heros 17.26 18.46 0.31 ACͲ30 Neverita duplicata 17.9 14.61 0.29 ACͲ31 Euspira heros 20.68 20.8 0.24 ACͲ32 Naticarius canrena 14.38 14.5 0.71 101 ACͲ33 Neverita duplicata 20.48 16.82 0.61 ACͲ34 Euspira heros 14.42 17.7 0.64 3.56 1.75 1 ACͲ35 Tectonatica pusilla 3.78 4.06 0.17 ACͲ36 Euspira heros 9.84 10.88 0.35 ACͲ37 Euspira heros 8.83 8.98 0.32 ACͲ38 Tectonatica pusilla 3 2.83 0.17 ACͲ39 Neverita duplicata 22.48 18.87 0.62 ACͲ40 Euspira heros 17.02 16.79 0.43 ACͲ41 Euspira heros 15.33 15.35 0.46 ACͲ42 Euspira heros 14.22 13.05 0.33 ACͲ43 Euspira heros 10.15 10.1 0.37 ACͲ44 Naticarius canrena 13.62 14.75 0.45 ACͲ45 Neverita duplicata 17.62 15.25 0.55 ACͲ46 Euspira heros 14.81 15.17 0.3 ACͲ47 Euspira heros 15.11 15.33 0.35 ACͲ48 Euspira heros 8.13 7.95 0.43 ACͲ49 Tectonatica pusilla 6.04 7.19 0.36 ACͲ50 Tectonatica pusilla 4.87 5.13 0.27 ACͲ51 Tectonatica pusilla 3.85 3.95 0.14 ACͲ52 Tectonatica pusilla 5.74 6.44 0.25 ACͲ53 Tectonatica pusilla 5.8 6.65 0.21 ACͲ54 Tectonatica pusilla 4.82 5.68 0.26 ACͲ55 Tectonatica pusilla 6.23 6.8 0.32 ACͲ56 Tectonatica pusilla 6.06 6.38 0.3 ACͲ57 Lunatia interna 6.73 7.74 0.22 ACͲ58 Tectonatica pusilla 4.65 5.44 0.2 ACͲ59 Tectonatica pusilla 4.14 4.32 0.21 ACͲ60 Tectonatica pusilla 4.94 5.32 0.26 0.52 0.38 8 ACͲ61 Tectonatica pusilla 3.71 4.44 0.13 102 ACͲ62 Lunatia interna 4.84 5.29 0.17 ACͲ63 Tectonatica pusilla 4.92 5.35 0.18 ACͲ64 Tectonatica pusilla 4.34 4.5 0.26 ACͲ65 Tectonatica pusilla 4.85 4.83 0.2 ACͲ66 Tectonatica pusilla 4.5 4.74 0.21 ACͲ67 Tectonatica pusilla 4.47 4.44 0.17 ACͲ68 Tectonatica pusilla 4.08 4.36 0.19 ACͲ69 Tectonatica pusilla 4.07 4.49 0.21 ACͲ70 Tectonatica pusilla 4.05 4.21 0.14 ACͲ71 Tectonatica pusilla 4.33 4.61 0.24 ACͲ72 Tectonatica pusilla 4.24 4.89 0.13 ACͲ73 Tectonatica pusilla 3.83 4.17 0.12 ACͲ74 Tectonatica pusilla 3.62 3.94 0.41 ACͲ75 Tectonatica pusilla 4.07 4.33 0.21 ACͲ76 Tectonatica pusilla 3.87 4.27 0.18 ACͲ77 Tectonatica pusilla 3.42 3.48 0.23 ACͲ78 Tectonatica pusilla 3.84 4.18 0.23 ACͲ79 Tectonatica pusilla 3.7 3.65 0.27 ACͲ80 Tectonatica pusilla 3.68 3.7 0.1 ACͲ81 Tectonatica pusilla 3.7 4.2 0.27 ACͲ82 Tectonatica pusilla 3.79 4.08 0.18 ACͲ83 Tectonatica pusilla 3.78 4.01 0.24 ACͲ84 Tectonatica pusilla 3.35 3.44 0.13 ACͲ85 Tectonatica pusilla 3.57 3.93 0.2 ACͲ86 Tectonatica pusilla 3.72 3.88 0.15 ACͲ87 Tectonatica pusilla 3.56 3.93 0.15 ACͲ88 Tectonatica pusilla 3.77 4.13 0.19 ACͲ89 Tectonatica pusilla 3.13 3.31 0.14 ACͲ90 Tectonatica pusilla 3.76 3.81 0.1 103 ACͲ91 Tectonatica pusilla 3.22 3.14 0.14 ACͲ92 Tectonatica pusilla 3.58 3.73 0.18 ACͲ93 Tectonatica pusilla 3.87 3.81 0.15 ACͲ94 Tectonatica pusilla 3.77 4 0.21 0.94 0.42 2 ACͲ95 Tectonatica pusilla 3.21 3.42 0.12 ACͲ96 Tectonatica pusilla 3.51 3.66 0.21 ACͲ97 Tectonatica pusilla 3.21 3.45 0.17 ACͲ98 Tectonatica pusilla 3.48 3.66 0.2 ACͲ99 Tectonatica pusilla 3.24 3.41 0.2 ACͲ100 Tectonatica pusilla 3.19 3.32 0.2 ACͲ101 Tectonatica pusilla 3.52 3.59 1.19 ACͲ102 Tectonatica pusilla 2.89 2.99 0.19 ACͲ103 Tectonatica pusilla 3.29 3.09 0.18 ACͲ104 Tectonatica pusilla 3.38 3.5 0.19 ACͲ105 Tectonatica pusilla 3.71 3.84 0.17 ACͲ106 Tectonatica pusilla 2.99 2.89 0.24 ACͲ107 Tectonatica pusilla 3.34 3.64 0.15 ACͲ108 Tectonatica pusilla 3.21 3.29 0.17 ACͲ109 Tectonatica pusilla 3.44 3.79 0.1 ACͲ110 Tectonatica pusilla 2.99 3.1 0.16 ACͲ111 Tectonatica pusilla 2.95 3.24 0.16 ACͲ112 Tectonatica pusilla 2.41 2.54 0.18 ACͲ113 Tectonatica pusilla 2.79 2.95 0.19 0.96 0.78 ACͲ114 Tectonatica pusilla 2.85 3.05 0.18 ACͲ115 Tectonatica pusilla 2.77 3.04 0.16 ACͲ116 Tectonatica pusilla 3.2 3.23 0.16 ACͲ117 Tectonatica pusilla 2.94 2.59 0.11 2.011667 104 Locality OldDock(TU870) DateCollected FMNH SouthwestofColumbus,NC Formation lowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External ODTUͲ1 Neverita duplicata 77.88 59.5 1.19 ODTUͲ2 Neverita duplicata 56.08 49.24 1.08 ODTUͲ3 Neverita duplicata 37.12 31.39 0.91 5.18 2.93 2 105 ODTUͲ4 Neverita duplicata 41.85 37.56 0.78 ODTUͲ5 Neverita duplicata 37.67 32.26 1.46 ODTUͲ6 Neverita duplicata 33.77 27.84 0.57 ODTUͲ7 Neverita duplicata 49.26 43.64 1.07 ODTUͲ8 Neverita duplicata 42.4 35.07 1.05 ODTUͲ9 Neverita duplicata 29.63 25.71 1.11 ODTUͲ10 Neverita duplicata 36.08 27.12 0.59 ODTUͲ11 Neverita duplicata 32.22 25.5 0.8 4.64 2.43 1 ODTUͲ12 Neverita duplicata 29.28 23.81 0.78 ODTUͲ13 Neverita duplicata 42.45 35.79 1.11 ODTUͲ14 Neverita duplicata 30.85 25.23 0.85 ODTUͲ15 Neverita duplicata 60.77 56.36 1.19 ODTUͲ16 Neverita duplicata 29.36 23.99 0.58 6.34 3.23 2 ODTUͲ17 Neverita duplicata 22.08 19.72 0.62 ODTUͲ18 Neverita duplicata 21.65 17.56 0.7 ODTUͲ19 Neverita duplicata 62.95 57.78 1.17 ODTUͲ20 Neverita duplicata 15.29 12.74 0.51 4.56 2.16 3 ODTUͲ21 Neverita duplicata 30.76 24.31 0.79 ODTUͲ22 Neverita duplicata 28.75 22.21 0.72 ODTUͲ23 Neverita duplicata 16.22 10.98 0.43 ODTUͲ24 Neverita duplicata 19.33 15.13 0.54 ODTUͲ25 Neverita duplicata 53.94 49.53 1.03 ODTUͲ26 Neverita duplicata 37.23 32.56 0.87 ODTUͲ27 Neverita duplicata 22.92 19.51 0.41 ODTUͲ28 Neverita duplicata 14.05 10.38 0.66 ODTUͲ29 Neverita duplicata 81.59 73.84 1.62 ODTUͲ30 Euspira heros 46.2 48.63 1.14 ODTUͲ31 Euspira heros 50.53 51.42 0.78 5.11 3.6 3 ODTUͲ32 Euspira heros 41.43 44.69 0.45 106 ODTUͲ33 Euspira heros 35.47 40.45 0.76 ODTUͲ34 Euspira heros 38.5 38.74 0.48 ODTUͲ35 Euspira heros 55.34 53.44 0.93 ODTUͲ36 Euspira heros 46.54 47.24 0.99 ODTUͲ37 Naticarius canrena 29.79 31.39 0.58 ODTUͲ38 Naticarius canrena 30.84 30.17 1.31 5.13 2.89 1 ODTUͲ39 Naticarius canrena 29.86 30.42 1.06 ODTUͲ40 Euspira heros 54.96 59.21 1.18 ODTUͲ41 Euspira heros 50.07 54.83 0.65 ODTUͲ42 Euspira heros 36.88 38.64 0.6 ODTUͲ43 Euspira heros 34.99 37.44 0.78 4.58 2.56 3 ODTUͲ44 Euspira heros 58.03 56.97 1.3 ODTUͲ45 Naticarius canrena 34.81 36.83 0.86 ODTUͲ46 Euspira heros 27.2 27.28 0.42 ODTUͲ47 Naticarius canrena 34.69 34.7 0.77 ODTUͲ48 Naticarius canrena 35.18 34.23 0.98 ODTUͲ49 Naticarius canrena 25.37 25.11 0.64 4.37 2.46 2 ODTUͲ50 Naticarius canrena 31.05 32.24 1.05 4.56 2.51 1 ODTUͲ51 Euspira heros 23.19 22.92 0.42 ODTUͲ52 Neverita duplicata 81.56 66.24 1.74 ODTUͲ53 Neverita duplicata 80.56 68.16 1.04 ODTUͲ54 Neverita duplicata 78.59 65.65 1.28 ODTUͲ55 Neverita duplicata 61.52 53.62 0.98 ODTUͲ56 Neverita duplicata 46.23 40.84 0.8 ODTUͲ57 Neverita duplicata 21.91 16.74 0.47 ODTUͲ58 Neverita duplicata 14.74 10.39 0.49 2.31 1.02 3 ODTUͲ59 Neverita duplicata 23.26 20.9 0.68 ODTUͲ60 Neverita duplicata 46.27 37.61 1.5 ODTUͲ61 Neverita duplicata 42.02 36.66 0.58 107 ODTUͲ62 Neverita duplicata 65.37 57.94 1.25 ODTUͲ63 Neverita duplicata 26.95 22.67 0.62 ODTUͲ64 Neverita duplicata 19.18 13.82 0.53 ODTUͲ65 Neverita duplicata 62.29 51.8 1.1 ODTUͲ66 Neverita duplicata 26.65 20.98 0.71 ODTUͲ67 Neverita duplicata 38.55 30.77 0.54 ODTUͲ68 Neverita duplicata 15.77 11.86 0.43 ODTUͲ69 Neverita duplicata 25.81 21.15 0.72 ODTUͲ70 Neverita duplicata 21.04 18.13 0.57 ODTUͲ71 Neverita duplicata 28.52 21.68 0.46 ODTUͲ72 Neverita duplicata 30 24.57 0.87 ODTUͲ73 Neverita duplicata 64.35 51.56 1.15 ODTUͲ74 Neverita duplicata 13.88 9.8 0.59 ODTUͲ75 Neverita duplicata 25.39 19.84 0.52 ODTUͲ76 Neverita duplicata 15.85 14.37 0.78 4.678 Locality OldDock(Petit) DateCollected FMNH SouthwestofColumbus,NC Formation lowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External ODPͲ1 Neverita duplicata 85.38 69.22 1.52 ODPͲ2 Neverita duplicata 78.99 66.85 1.24 ODPͲ3 Neverita duplicata 90.35 76.45 1.34 108 ODPͲ4 Euspira heros 22.43 21.82 0.38 ODPͲ5 Neverita duplicata 58.11 54.4 0.88 ODPͲ6 Euspira heros 36.55 37.39 0.54 5.07 3.61 5 ODPͲ7 Neverita duplicata 26.98 22.98 0.88 ODPͲ8 Euspira heros 57.41 60.57 1.18 ODPͲ9 Neverita duplicata 49.33 42.78 1.09 ODPͲ10 Neverita duplicata 40.95 33.93 0.79 ODPͲ11 Euspira heros 35.38 37.74 0.59 ODPͲ12 Neverita duplicata 22.62 17.85 0.45 ODPͲ13 Naticarius canrena 17.18 17.14 0.66 ODPͲ14 Naticarius canrena 37.37 37.46 1.12 ODPͲ15 Neverita duplicata 15.82 11.42 0.52 ODPͲ16 Euspira heros 11.94 12.72 0.32 3.39 1.39 2 ODPͲ17 Naticarius canrena 33.33 32.12 0.88 ODPͲ18 Neverita duplicata 68.28 60.39 1.3 ODPͲ19 Naticarius canrena 39.41 42.63 1.64 ODPͲ20 Naticarius canrena 39.38 42.44 1.21 ODPͲ21 Euspira heros 28.25 28.83 0.76 ODPͲ22 Euspira heros 17.44 17.96 0.58 ODPͲ23 Naticarius canrena 34.88 38.85 0.89 ODPͲ24 Naticarius canrena 37.67 39.08 0.57 ODPͲ25 Naticarius canrena 30 29.95 0.69 4.77 2.29 2 ODPͲ26 Neverita duplicata 19.26 15.46 0.55 ODPͲ27 Euspira heros 46.41 46.89 0.91 ODPͲ28 Naticarius canrena 53.8 54.49 0.65 ODPͲ29 Neverita duplicata 36.77 31.67 0.92 ODPͲ30 Euspira heros 8.56 9.13 0.18 ODPͲ31 Euspira heros 20.85 19.33 0.31 ODPͲ32 Naticarius canrena 24.74 25.24 0.53 109 ODPͲ33 Naticarius canrena 20.34 20.74 0.59 ODPͲ34 Neverita duplicata 20.76 14.61 0.38 ODPͲ35 Neverita duplicata 24.59 18.52 0.5 5.74 3.25 2 ODPͲ36 Neverita duplicata 28.52 21.37 0.58 ODPͲ37 Neverita duplicata 20.66 15.6 0.45 ODPͲ38 Neverita duplicata 31.08 24 0.4 5.62 3.17 2 ODPͲ39 Neverita duplicata 47.48 45.85 0.85 ODPͲ40 Neverita duplicata 17.15 11.81 0.43 ODPͲ41 Naticarius canrena 23.81 21.31 0.41 ODPͲ42 Euspira heros 25.33 23.88 0.31 4.18 2.33 3 ODPͲ43 Naticarius canrena 35.45 36.64 0.78 ODPͲ44 Euspira heros 30.49 32.52 0.85 ODPͲ45 Euspira heros 18 18.7 0.45 2.78 1.5 3 ODPͲ46 Euspira heros 20.85 22.27 0.45 ODPͲ47 Euspira heros 20.29 19.79 0.37 ODPͲ48 Neverita duplicata 22.92 19.57 0.74 ODPͲ49 Neverita duplicata 53.35 40.99 1.21 ODPͲ50 Euspira heros 18.64 18.72 0.39 2.85 1.62 3 ODPͲ51 Euspira heros 21.98 21.82 0.83 ODPͲ52 Naticarius canrena 19.18 19.08 0.75 ODPͲ53 Naticarius canrena 48.21 46.22 0.65 ODPͲ54 Neverita duplicata 36.49 28.72 0.79 ODPͲ55 Naticarius canrena 39.47 38.94 0.82 ODPͲ56 Euspira heros 21.56 26.25 0.42 ODPͲ57 Euspira heros 20.47 21.66 0.47 ODPͲ58 Euspira heros 15.71 17.45 0.42 ODPͲ59 Naticarius canrena 14.41 13.71 0.77 ODPͲ60 Euspira heros 11.87 13.45 0.24 ODPͲ61 Neverita duplicata 16 12.55 0.43 110 ODPͲ62 Euspira heros 19.67 19.73 0.39 ODPͲ63 Naticarius canrena 18.75 19.17 0.63 ODPͲ64 Neverita duplicata 20.95 16.1 0.45 4.61 2.61 2 ODPͲ65 Euspira heros 33.75 34.26 0.81 4.67 3.19 7 ODPͲ66 Euspira heros 28.37 29.62 0.55 ODPͲ67 Neverita duplicata 18.02 12.95 0.51 ODPͲ68 Naticarius canrena 19.57 19.13 0.58 4.49 2.33 2 ODPͲ69 Neverita duplicata 36.71 31.54 0.68 ODPͲ70 Euspira heros 69.83 73.46 1.71 ODPͲ71 Euspira heros 55.83 57.22 0.65 ODPͲ72 Euspira heros 66.22 75.87 1.36 ODPͲ73 Euspira heros 40.88 43.99 0.83 ODPͲ74 Euspira heros 33.22 32.74 0.64 ODPͲ75 Euspira heros 58.92 55.75 1.16 ODPͲ76 Euspira heros 41.31 46.31 0.99 ODPͲ77 Euspira heros 57.59 57.28 1.69 ODPͲ78 Euspira heros 23.5 24.88 0.4 ODPͲ79 Euspira heros 17.86 17.98 0.24 ODPͲ80 Euspira heros 26.69 27.52 0.5 3.87 2.23 2 ODPͲ81 Euspira heros 16.49 17.47 0.34 ODPͲ82 Euspira heros 56.13 58.71 0.73 ODPͲ83 Euspira heros 56.88 60.08 1.55 ODPͲ84 Euspira heros 49.26 48.89 0.75 ODPͲ85 Euspira heros 28.39 27.57 0.47 3.89 2.14 2 ODPͲ86 Euspira heros 28.9 29.69 0.56 ODPͲ87 Euspira heros 15.18 16.04 0.44 ODPͲ88 Euspira heros 23.67 26.17 0.43 ODPͲ89 Euspira heros 21.43 21.9 0.52 ODPͲ90 Euspira heros 23.91 24.9 0.43 111 ODPͲ91 Euspira heros 33.56 32.84 0.44 4.68 3.56 6 ODPͲ92 Euspira heros 31.81 30.93 0.84 ODPͲ93 Euspira heros 16.21 17.31 0.35 ODPͲ94 Euspira heros 16.03 16.11 0.42 2.98 0.56 3 ODPͲ95 Euspira heros 11.56 12.24 0.31 ODPͲ96 Euspira heros 28.54 20.28 0.46 ODPͲ97 Euspira heros 17.02 17.23 0.55 ODPͲ98 Euspira heros 26.52 27.6 0.43 ODPͲ99 Neverita duplicata 87.6 74.29 1.49 ODPͲ100 Neverita duplicata 67.97 61.22 1.07 ODPͲ101 Neverita duplicata 62.03 49.88 1.06 ODPͲ102 Neverita duplicata 59.88 55.41 1.01 ODPͲ103 Neverita duplicata 54.22 47.69 1.32 ODPͲ104 Neverita duplicata 61.95 51.41 0.97 ODPͲ105 Neverita duplicata 51.18 42.77 1.28 ODPͲ106 Neverita duplicata 33.69 26.43 0.86 5.4 3.4 1 ODPͲ107 Neverita duplicata 50.54 49.97 0.94 ODPͲ108 Lunatia perspectiva 46.96 44.69 0.66 ODPͲ109 Neverita duplicata 30.06 23.88 0.94 ODPͲ110 Lunatia perspectiva 40.42 37.77 0.75 ODPͲ111 Neverita duplicata 46.25 42.47 0.8 ODPͲ112 Neverita duplicata 32.5 30.89 0.88 ODPͲ113 Neverita duplicata 18.64 15.22 0.56 ODPͲ114 Neverita duplicata 17.63 14.93 0.63 ODPͲ115 Neverita duplicata 20.22 14.1 0.42 4.311875 112 Locality CrescentBeachAirport DateCollected FMNH Near33°47'N78°42'W Formation lowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External CBͲ1 Neverita duplicata 36.66 31.97 0.52 CBͲ2 Neverita duplicata 43.63 37.98 0.74 CBͲ3 Neverita duplicata 40.17 37.98 0.74 113 CBͲ4 Neverita duplicata 43.49 35.79 0.96 CBͲ5 Neverita duplicata 38.12 33.12 0.46 CBͲ6 Neverita duplicata 33.86 29.67 0.66 CBͲ7 Neverita duplicata 20.93 16.27 0.61 CBͲ8 Neverita duplicata 22.58 18.1 0.46 CBͲ9 Neverita duplicata 20.5 16.87 0.5 CBͲ10 Neverita duplicata 21.83 17.87 0.47 3.98 2.11 2 CBͲ11 Neverita duplicata 23.62 19.27 0.52 CBͲ12 Neverita duplicata 37.89 31.99 0.71 CBͲ13 Neverita duplicata 36.93 33.01 0.81 CBͲ14 Neverita duplicata 41.35 36.08 0.67 CBͲ15 Neverita duplicata 25.98 20.98 0.43 CBͲ16 Neverita duplicata 41.53 36.14 0.85 CBͲ17 Neverita duplicata 34.72 29.64 0.99 CBͲ18 Neverita duplicata 34.31 30.42 0.72 CBͲ19 Neverita duplicata 31.76 26.76 0.67 CBͲ20 Neverita duplicata 18.09 15.39 0.49 4.56 2.37 2 CBͲ21 Neverita duplicata 29.17 25.81 0.64 CBͲ22 Neverita duplicata 37.28 34.3 0.75 CBͲ23 Neverita duplicata 27.51 24.68 0.62 CBͲ24 Neverita duplicata 20.47 16.46 0.42 CBͲ25 Neverita duplicata 23.41 19.04 0.57 CBͲ26 Neverita duplicata 18.92 16.05 0.34 4.03 2.17 2 CBͲ27 Neverita duplicata 20.19 16.83 0.39 CBͲ28 Neverita duplicata 28.77 23.05 0.65 CBͲ29 Neverita duplicata 18.31 15.34 0.5 CBͲ30 Neverita duplicata 29.45 25.04 0.6 CBͲ31 Neverita duplicata 26.16 21.96 0.52 4.26 1.42 2 CBͲ32 Neverita duplicata 20.06 15.73 0.45 114 CBͲ33 Neverita duplicata 21.74 16.01 0.65 incdrillhole CBͲ34 Neverita duplicata 22.97 18.79 0.54 4.57 2.62 2 CBͲ35 Neverita duplicata 23.79 19.5 0.35 CBͲ36 Neverita duplicata 14.4 11.71 0.41 3.71 1.54 3 CBͲ37 Neverita duplicata 21.25 16.94 0.46 CBͲ38 Neverita duplicata 20.87 15.63 0.51 CBͲ39 Neverita duplicata 35.47 29.46 0.79 CBͲ40 Neverita duplicata 22.29 16.18 0.59 CBͲ41 Neverita duplicata 27.66 22.77 0.52 CBͲ42 Neverita duplicata 16.34 13.4 0.43 CBͲ43 Neverita duplicata 27.47 23.64 0.71 CBͲ44 Neverita duplicata 27.1 24.01 0.54 CBͲ45 Neverita duplicata 39 40.02 0.54 CBͲ46 Neverita duplicata 21.8 16.66 0.62 CBͲ47 Neverita duplicata 24.31 20.27 0.64 CBͲ48 Neverita duplicata 18.67 15.24 0.46 CBͲ49 Neverita duplicata 40.57 37.37 1.02 CBͲ50 Neverita duplicata 33.52 30.35 0.7 CBͲ51 Neverita duplicata 24.86 21.87 0.7 CBͲ52 Neverita duplicata 23.36 18.92 0.57 CBͲ53 Neverita duplicata 19.88 16.71 0.46 CBͲ54 Neverita duplicata 33.18 29.61 0.8 CBͲ55 Neverita duplicata 16.25 12.61 0.39 CBͲ56 Neverita duplicata 24.66 20.47 0.62 CBͲ57 Neverita duplicata 18.03 14.38 0.41 2.76 2.3 2 CBͲ58 Neverita duplicata 23.27 18 0.44 CBͲ59 Neverita duplicata 19.27 14.37 0.46 CBͲ60 Neverita duplicata 22.89 17.5 0.55 CBͲ61 Neverita duplicata 15.51 11.96 0.37 115 CBͲ62 Neverita duplicata 18.72 15.21 0.59 CBͲ63 Neverita duplicata 20.67 16.77 0.45 CBͲ64 Neverita duplicata 22.73 19.36 0.49 CBͲ65 Neverita duplicata 16.56 13.12 0.31 CBͲ66 Neverita duplicata 18.04 13.37 0.54 CBͲ67 Neverita duplicata 32.64 30.36 1.01 CBͲ68 Neverita duplicata 19.57 16.44 0.6 incdrillhole CBͲ69 Neverita duplicata 25.91 22.46 0.74 CBͲ70 Neverita duplicata 25.1 21.7 0.51 CBͲ71 Neverita duplicata 21.91 18.52 0.49 CBͲ72 Neverita duplicata 26.51 24.84 0.55 CBͲ73 Neverita duplicata 22.17 17.39 0.42 CBͲ74 Neverita duplicata 19.29 14.32 0.43 CBͲ75 Neverita duplicata 21.82 16.3 0.61 CBͲ76 Neverita duplicata 29.47 26.13 0.47 CBͲ77 Neverita duplicata 18.31 14.46 0.35 CBͲ78 Neverita duplicata 18.94 15.75 0.54 CBͲ79 Neverita duplicata 19.5 16.8 0.56 CBͲ80 Neverita duplicata 16.65 13.27 0.42 CBͲ81 Neverita duplicata 18.99 16.16 0.48 CBͲ82 Neverita duplicata 19.31 15.42 0.43 CBͲ83 Neverita duplicata 20.27 16.37 0.46 CBͲ84 Neverita duplicata 22.04 19.15 0.58 CBͲ85 Neverita duplicata 29.31 26.4 0.68 CBͲ86 Neverita duplicata 11.87 9.84 0.24 2.98 1.22 3 CBͲ87 Neverita duplicata 20.4 16.34 0.52 CBͲ88 Neverita duplicata 14.28 12.11 0.48 CBͲ89 Neverita duplicata 20.29 16.08 0.32 CBͲ90 Neverita duplicata 18.16 12.68 0.47 116 CBͲ91 Neverita duplicata 21.64 17.67 0.42 CBͲ92 Neverita duplicata 22.23 18.73 0.52 CBͲ93 Neverita duplicata 23.97 21.19 0.58 CBͲ94 Neverita duplicata 16.98 13.13 0.53 4.16 1.86 2 CBͲ95 Neverita duplicata 14.57 12.21 0.59 CBͲ96 Neverita duplicata 18.52 14.7 0.4 CBͲ97 Neverita duplicata 19.98 15.36 0.49 CBͲ98 Neverita duplicata 14.96 12.37 0.37 4.66 2.39 3 CBͲ99 Neverita duplicata 27.34 25.53 0.57 CBͲ100 Neverita duplicata 23.3 18.15 0.65 CBͲ101 Neverita duplicata 27.07 23.63 0.58 CBͲ102 Neverita duplicata 7.45 6.08 0.25 CBͲ103 Neverita duplicata 22.05 19.23 0.54 CBͲ104 Neverita duplicata 22.28 19.46 0.75 CBͲ105 Neverita duplicata 31.4 27.31 0.78 CBͲ106 Neverita duplicata 18.01 14.48 0.61 CBͲ107 Neverita duplicata 18.36 15.03 0.56 CBͲ108 Neverita duplicata 19.06 15.74 0.47 CBͲ109 Neverita duplicata 22.61 19.25 0.6 CBͲ110 Neverita duplicata 22.42 17.99 0.47 CBͲ111 Neverita duplicata 26.66 21.75 0.57 CBͲ112 Neverita duplicata 19.19 15.12 0.4 CBͲ113 Neverita duplicata 11.39 9.31 0.24 2.71 1.39 3 CBͲ114 Neverita duplicata 18.63 14.21 0.31 CBͲ115 Neverita duplicata 16.93 12.94 0.5 3.33 1.52 2 CBͲ116 Neverita duplicata 26.98 25.33 0.64 CBͲ117 Neverita duplicata 23.21 18.27 0.52 CBͲ118 Neverita duplicata 17.49 13.88 0.49 CBͲ119 Neverita duplicata 11.58 9.57 0.48 117 CBͲ120 Neverita duplicata 22.75 17.28 0.77 CBͲ121 Neverita duplicata 33.83 28.86 1.03 CBͲ122 Neverita duplicata 33.39 30.3 0.69 CBͲ123 Neverita duplicata 26.51 23.25 0.68 4.59 2.49 2 CBͲ124 Neverita duplicata 20.26 15.64 0.48 CBͲ125 Neverita duplicata 14.98 12.09 0.49 CBͲ126 Neverita duplicata 20.16 16.89 0.58 CBͲ127 Neverita duplicata 20.15 16.7 0.57 CBͲ128 Neverita duplicata 16.69 12.4 0.44 CBͲ129 Neverita duplicata 19.12 15.47 0.6 CBͲ130 Neverita duplicata 23.09 16.96 0.49 CBͲ131 Neverita duplicata 18.44 14.03 0.44 CBͲ132 Neverita duplicata 20.28 16.49 0.56 CBͲ133 Neverita duplicata 27.78 22.85 0.7 CBͲ134 Neverita duplicata 10.11 8.13 0.22 2.59 1.46 3 CBͲ135 Neverita duplicata 14.8 11.57 0.38 CBͲ136 Neverita duplicata 15.5 11.98 0.35 CBͲ137 Neverita duplicata 31.42 26.37 0.69 CBͲ138 Neverita duplicata 26.04 22.52 0.66 CBͲ139 Neverita duplicata 18.91 14.26 0.47 CBͲ140 Neverita duplicata 14.08 11.76 0.32 CBͲ141 Neverita duplicata 23.33 19.36 0.68 CBͲ142 Neverita duplicata 16.52 13.7 0.29 3.36 1.68 3 CBͲ143 Neverita duplicata 20.54 16.81 0.5 CBͲ144 Neverita duplicata 15.5 10.61 0.33 3.69 1.49 2 CBͲ145 Neverita duplicata 21.57 16.81 0.44 CBͲ146 Neverita duplicata 19.79 15.75 0.48 CBͲ147 Neverita duplicata 14.77 11.21 0.32 3.7 1.59 3 CBͲ148 Neverita duplicata 26.43 23.77 0.63 118 CBͲ149 Neverita duplicata 16.42 13.45 0.49 CBͲ150 Neverita duplicata 13.3 10.81 0.31 CBͲ151 Neverita duplicata 21.28 18.49 0.47 CBͲ152 Neverita duplicata 17.67 14.32 0.42 CBͲ153 Neverita duplicata 18.47 14.85 0.57 CBͲ154 Neverita duplicata 17.26 14.36 0.44 CBͲ155 Neverita duplicata 24.91 20.92 0.62 CBͲ156 Neverita duplicata 18.89 14.92 0.56 CBͲ157 Neverita duplicata 20.2 16.44 0.45 CBͲ158 Neverita duplicata 27.99 21.98 0.85 CBͲ159 Neverita duplicata 35.75 30.79 0.9 CBͲ160 Neverita duplicata 23.07 18.47 0.78 CBͲ161 Neverita duplicata 16.8 12.26 0.39 CBͲ162 Neverita duplicata 26.04 21.02 0.86 CBͲ163 Neverita duplicata 17.73 14.68 0.45 CBͲ164 Neverita duplicata 17.62 14.15 0.37 CBͲ165 Neverita duplicata 26.48 22.6 0.64 CBͲ166 Neverita duplicata 20.06 16.98 0.52 CBͲ167 Neverita duplicata 20.86 17.67 0.58 4.4 2.92 3 CBͲ168 Neverita duplicata 30.54 29.09 0.95 CBͲ169 Neverita duplicata 19.53 15.35 0.69 CBͲ170 Neverita duplicata 13.67 11.17 0.47 CBͲ171 Neverita duplicata 14.87 10.54 0.47 CBͲ172 Neverita duplicata 12.77 10.44 0.34 CBͲ173 Neverita duplicata 15.54 12.64 0.39 4.04 1.85 4 CBͲ174 Neverita duplicata 11.75 8.97 0.42 CBͲ175 Neverita duplicata 25.12 21.26 0.59 CBͲ176 Neverita duplicata 26.51 22.01 0.75 CBͲ177 Neverita duplicata 15.23 13.09 0.85 119 CBͲ178 Neverita duplicata 11.68 10.03 0.3 CBͲ179 Neverita duplicata 19.65 15.49 0.45 CBͲ180 Neverita duplicata 20.59 19.1 0.55 CBͲ181 Neverita duplicata 19.91 15.89 0.5 CBͲ182 Neverita duplicata 17.31 13.15 0.42 CBͲ183 Neverita duplicata 16.48 13.12 0.49 CBͲ184 Neverita duplicata 21.49 17.93 0.54 CBͲ185 Neverita duplicata 20.75 16.81 0.64 CBͲ186 Neverita duplicata 22.85 18.3 0.41 CBͲ187 Neverita duplicata 13.2 10.04 0.34 CBͲ188 Neverita duplicata 13.07 10.87 0.31 2.76 1.18 3 CBͲ189 Neverita duplicata 14.03 11.05 0.33 CBͲ190 Neverita duplicata 24.11 20.55 0.52 5.2 2.64 2 CBͲ191 Neverita duplicata 15.46 12.16 0.38 CBͲ192 Neverita duplicata 17.63 15.47 0.4 CBͲ193 Neverita duplicata 20.47 16.95 0.53 CBͲ194 Neverita duplicata 16.92 14.19 0.42 4.38 2.02 3 CBͲ195 Neverita duplicata 13.11 10.98 0.39 CBͲ196 Neverita duplicata 20.7 16.56 0.38 CBͲ197 Neverita duplicata 15.19 12.33 0.54 CBͲ198 Neverita duplicata 18.45 14.25 0.62 CBͲ199 Neverita duplicata 17.17 12.54 0.46 CBͲ200 Neverita duplicata 18.66 14.36 0.45 CBͲ201 Neverita duplicata 21.62 17.64 0.54 CBͲ202 Neverita duplicata 17.5 13.65 0.75 CBͲ203 Neverita duplicata 17.78 14.57 0.59 CBͲ204 Neverita duplicata 18.12 14.64 0.43 CBͲ205 Neverita duplicata 14.74 11.26 0.27 CBͲ206 Neverita duplicata 9.36 7.36 0.37 120 CBͲ207 Neverita duplicata 24.65 21.29 0.47 CBͲ208 Neverita duplicata 15.94 12.36 0.45 CBͲ209 Neverita duplicata 13.32 10.62 0.42 2.31 1.27 3 CBͲ210 Neverita duplicata 21.88 21.95 19.05 4.24 2.31 2 CBͲ211 Neverita duplicata 16.27 13.37 0.49 3.69 1.79 3 CBͲ212 Neverita duplicata 23.36 17.82 0.57 4.02 1.81 3 CBͲ213 Neverita duplicata 19.77 16.98 0.48 CBͲ214 Neverita duplicata 19.87 16.27 0.59 CBͲ215 Neverita duplicata 17.68 13.91 0.4 CBͲ216 Neverita duplicata 15.31 11.55 0.59 CBͲ217 Neverita duplicata 15.84 13.98 0.51 CBͲ218 Neverita duplicata 13.72 10.24 0.27 CBͲ219 Neverita duplicata 16.67 12.35 0.33 3.98 1.81 2 CBͲ220 Neverita duplicata 17.24 13.05 0.3 CBͲ221 Neverita duplicata 16.52 14.24 0.39 CBͲ222 Neverita duplicata 13.08 11.01 0.31 CBͲ223 Neverita duplicata 16.97 12.84 0.41 CBͲ224 Neverita duplicata 18.37 13.6 0.47 CBͲ225 Neverita duplicata 17.09 13.27 0.63 CBͲ226 Neverita duplicata 31.09 28.46 0.85 CBͲ227 Neverita duplicata 18.68 14.52 0.45 CBͲ228 Neverita duplicata 19.23 15.79 0.31 CBͲ229 Neverita duplicata 10.21 8.96 0.37 CBͲ230 Neverita duplicata 19 15.99 0.32 CBͲ231 Neverita duplicata 22.78 20.43 0.47 CBͲ232 Neverita duplicata 22.53 18.48 0.53 CBͲ233 Neverita duplicata 15.49 12 0.43 CBͲ234 Neverita duplicata 6.6 5.56 0.32 0.98 2 CBͲ235 Neverita duplicata 14.32 10.27 0.31 121 CBͲ236 Neverita duplicata 14.5 12.07 0.32 CBͲ237 Neverita duplicata 16.11 13.37 0.4 1.83 3 CBͲ238 Neverita duplicata 18.57 14.6 0.5 CBͲ239 Neverita duplicata 22.98 19.73 0.51 CBͲ240 Neverita duplicata 16.83 13.08 0.5 CBͲ241 Neverita duplicata 15.11 12.97 0.66 CBͲ242 Neverita duplicata 23.11 20.51 0.61 CBͲ243 Neverita duplicata 18.42 14.56 0.63 CBͲ244 Neverita duplicata 14.86 11.24 0.79 CBͲ245 Neverita duplicata 17.47 14.43 0.63 CBͲ246 Neverita duplicata 21.77 17.22 0.4 CBͲ247 Neverita duplicata 18.57 14.33 0.47 2.12 3 CBͲ248 Neverita duplicata 16.9 13.69 0.52 1.83 2 CBͲ249 Neverita duplicata 17.43 13.82 0.52 CBͲ250 Neverita duplicata 16.66 13.69 0.5 CBͲ251 Neverita duplicata 8.79 7.48 0.48 CBͲ252 Neverita duplicata 11.32 9.78 0.42 CBͲ253 Neverita duplicata 13.71 11.21 0.46 CBͲ254 Neverita duplicata 10.56 8.43 0.32 CBͲ255 Neverita duplicata 14.02 10.72 0.37 CBͲ256 Neverita duplicata 17.89 13.94 0.48 CBͲ257 Neverita duplicata 11.35 9.12 0.36 1.02 2 CBͲ258 Neverita duplicata 16.55 12.21 0.66 1.89 2 CBͲ259 Neverita duplicata 24.69 21.68 0.58 2.05 3 CBͲ260 Neverita duplicata 22 17.82 0.4 CBͲ261 Neverita duplicata 13.89 11.14 0.4 1.74 2 CBͲ262 Neverita duplicata 20.56 17.34 0.57 CBͲ263 Neverita duplicata 17.3 12.65 0.65 CBͲ264 Neverita duplicata 11.32 9.64 0.28 122 CBͲ265 Neverita duplicata 19.39 15.59 0.52 CBͲ266 Neverita duplicata 10.2 7.73 0.29 1.6 1.02 4 CBͲ267 Neverita duplicata 15.9 11.94 0.55 CBͲ268 Neverita duplicata 13.71 10.56 0.66 CBͲ269 Neverita duplicata 14.65 10.9 0.45 CBͲ270 Neverita duplicata 10.59 8.35 0.34 CBͲ271 Neverita duplicata 10.38 8.29 0.32 2.66 1.31 3 CBͲ272 Neverita duplicata 12.25 9.72 0.37 CBͲ273 Neverita duplicata 16.1 13.52 0.46 CBͲ274 Neverita duplicata 14.31 11.3 0.5 CBͲ275 Neverita duplicata 19.33 15.74 0.58 CBͲ276 Neverita duplicata 27.69 23.9 0.68 CBͲ277 Neverita duplicata 13.96 12.27 0.44 CBͲ278 Neverita duplicata 15.15 12.04 0.35 3.3 1.33 3 CBͲ279 Neverita duplicata 18.44 15.01 0.47 CBͲ280 Neverita duplicata 20.11 16.35 0.66 CBͲ281 Neverita duplicata 26.82 22.22 0.51 CBͲ282 Neverita duplicata 22.69 19.06 0.72 CBͲ283 Neverita duplicata 21.41 17.39 0.56 CBͲ284 Neverita duplicata 10.71 8.48 0.47 CBͲ285 Neverita duplicata 17.24 14.12 0.48 CBͲ286 Neverita duplicata 12.64 9.18 0.33 CBͲ287 Neverita duplicata 18.56 14.15 0.5 4.35 1.79 2 CBͲ288 Neverita duplicata 11.36 9.11 0.41 CBͲ289 Neverita duplicata 13.22 11.3 0.4 CBͲ290 Neverita duplicata 18.03 14.82 0.52 CBͲ291 Neverita duplicata 10.68 8.64 0.33 CBͲ292 Neverita duplicata 12.42 10.02 0.46 CBͲ293 Neverita duplicata 18.17 15.2 0.65 123 CBͲ294 Neverita duplicata 12.24 9.34 0.35 CBͲ295 Neverita duplicata 17.77 14.18 0.47 CBͲ296 Neverita duplicata 19.02 15.08 0.48 CBͲ297 Neverita duplicata 16.43 13.42 0.48 CBͲ298 Neverita duplicata 22.45 18.3 0.57 4.49 2.34 2 CBͲ299 Neverita duplicata 17.15 14.67 0.4 CBͲ300 Neverita duplicata 14.41 11.34 0.53 CBͲ301 Neverita duplicata 9.91 7.96 0.29 CBͲ302 Neverita duplicata 15.32 12.3 0.41 CBͲ303 Neverita duplicata 10.86 8.84 0.33 CBͲ304 Neverita duplicata 10.22 7.72 0.65 CBͲ305 Neverita duplicata 23.56 19.43 0.91 CBͲ306 Neverita duplicata 12.75 10.08 0.36 2.51 1.02 3 CBͲ307 Neverita duplicata 14.2 11.01 0.53 CBͲ308 Neverita duplicata 11.74 8.25 0.31 CBͲ309 Neverita duplicata 8.95 7.41 0.24 CBͲ310 Neverita duplicata 10.52 8.54 0.36 CBͲ311 Neverita duplicata 8.48 6.99 0.24 CBͲ312 Neverita duplicata 7.32 6.17 0.23 CBͲ313 Neverita duplicata 7.14 5.64 0.3 CBͲ314 Neverita duplicata 5.49 5.07 0.23 CBͲ315 Euspira heros 5.41 4.89 0.25 CBͲ316 Neverita duplicata 5.36 4.69 0.26 CBͲ317 Euspira heros 64.1 65.11 1.15 CBͲ318 Euspira heros 50.34 51.67 0.75 CBͲ319 Euspira heros 23.89 24.87 0.7 CBͲ320 Euspira heros 38.33 41.07 0.6 CBͲ321 Euspira heros 50.04 52.26 0.83 CBͲ322 Euspira heros 42.92 46.54 0.69 124 CBͲ323 Euspira heros 44.63 47.66 0.73 CBͲ324 Euspira heros 15.91 17.16 0.44 CBͲ325 Euspira heros 34.94 37.59 1 CBͲ326 Euspira heros 13 13.1 0.34 2.82 1.77 1 CBͲ327 Euspira heros 21.21 20.84 0.57 CBͲ328 Euspira heros 26.26 27.23 0.67 CBͲ329 Euspira heros 13.1 13.04 0.3 CBͲ330 Euspira heros 30.45 29.69 0.56 CBͲ331 Euspira heros 14.39 14.77 0.35 CBͲ332 Euspira heros 19.02 18.81 0.36 CBͲ333 Euspira heros 15.72 16.49 0.45 CBͲ334 Euspira heros 15.78 15.05 0.32 CBͲ335 Euspira heros 6.8 7.18 0.25 CBͲ336 Euspira heros 24.48 24.53 0.51 CBͲ337 Euspira heros 23.05 24.69 0.54 CBͲ338 Euspira heros 23.04 23.91 0.47 3.26 1.51 4 CBͲ339 Euspira heros 17.13 17.7 0.29 CBͲ340 Euspira heros 6.12 5.74 0.27 CBͲ341 Euspira heros 18.59 20.05 0.52 2.72 1.37 3 CBͲ342 Euspira heros 17.77 17.47 0.53 CBͲ343 Euspira heros 6.57 7 0.27 CBͲ344 Euspira heros 7.39 7.29 0.28 2.28 1.2 2 CBͲ345 Euspira heros 8.53 9.07 0.29 CBͲ346 Euspira heros 17.1 18.9 0.31 CBͲ347 Euspira heros 4.82 4.81 0.28 CBͲ348 Euspira heros 32.69 32.94 0.88 CBͲ349 Euspira heros 31.35 32.33 0.56 CBͲ350 Euspira heros 17.87 18.85 0.48 3.9 2.3 2 CBͲ351 Euspira heros 22.03 22.12 0.65 125 CBͲ352 Euspira heros 18.87 19.54 0.61 CBͲ353 Euspira heros 15.12 15.44 0.66 CBͲ354 Euspira heros 26.04 27.65 0.7 CBͲ355 Euspira heros 20.51 19.42 0.75 CBͲ356 Euspira heros 15.8 16.05 0.8 CBͲ357 Euspira heros 19.04 19.11 0.31 CBͲ358 Euspira heros 15.81 17.34 0.54 CBͲ359 Euspira heros 21.46 22.06 0.87 CBͲ360 Euspira heros 15.99 15.87 0.43 CBͲ361 Euspira heros 17.82 17.93 0.44 CBͲ362 Euspira heros 20.73 21.96 0.47 3.74 2.33 3 CBͲ363 Euspira heros 20.78 20.72 0.34 3.66 1.88 2 CBͲ364 Euspira heros 19.25 19.36 0.43 CBͲ365 Euspira heros 17.81 18.59 0.84 CBͲ366 Euspira heros 22.66 24.8 0.55 CBͲ367 Euspira heros 16.39 16.29 0.35 CBͲ368 Euspira heros 14.75 15.44 0.33 CBͲ369 Euspira heros 19.37 20.73 0.62 CBͲ370 Euspira heros 14.36 14.1 0.37 CBͲ371 Euspira heros 15.73 16.29 0.47 CBͲ372 Euspira heros 14.31 14.59 0.45 CBͲ373 Euspira heros 15.35 15.74 0.32 CBͲ374 Euspira heros 18.14 17.68 0.52 CBͲ375 Euspira heros 13.84 14.35 0.28 CBͲ376 Euspira heros 11.85 12.32 0.52 CBͲ377 Euspira heros 14.92 15.66 0.56 CBͲ378 Euspira heros 15.39 15.27 0.29 CBͲ379 Euspira heros 13.93 14.11 0.29 CBͲ380 Euspira heros 13.34 2.85 0.28 126 CBͲ381 Euspira heros 14.6 15.29 0.41 CBͲ382 Euspira heros 16.98 16.9 0.57 CBͲ383 Euspira heros 14.14 15.02 0.53 CBͲ384 Euspira heros 11.06 11.67 0.28 2.69 1.23 1 CBͲ385 Euspira heros 17.42 17.63 0.6 CBͲ386 Euspira heros 17.37 18.42 0.62 CBͲ387 Euspira heros 11.66 11.19 0.4 2.4 0.74 1 CBͲ388 Euspira heros 12.47 12.82 0.31 CBͲ389 Euspira heros 12.8 13.08 0.27 CBͲ390 Euspira heros 18.88 19.19 0.71 CBͲ391 Euspira heros 13.15 13.02 0.41 CBͲ392 Euspira heros 13.03 13.46 0.43 CBͲ393 Euspira heros 9.01 9.5 0.29 CBͲ394 Euspira heros 8.59 8.67 0.33 CBͲ395 Euspira heros 12.49 12.05 0.49 CBͲ396 Euspira heros 12.13 12.17 0.43 CBͲ397 Euspira heros 14.45 15.11 0.33 CBͲ398 Euspira heros 9.32 9.61 0.38 CBͲ399 Euspira heros 10.35 11 0.32 CBͲ400 Euspira heros 8.24 7.92 0.23 CBͲ401 Euspira heros 11.18 11.44 0.36 CBͲ402 Euspira heros 9.11 9.88 0.35 CBͲ403 Euspira heros 9.45 9.5 0.35 CBͲ404 Euspira heros 8.78 9.35 0.24 CBͲ405 Euspira heros 14.95 13.69 0.35 4.15 2.02 2 CBͲ406 Euspira heros 16.66 17.5 0.38 CBͲ407 Euspira heros 15.7 17.24 0.82 CBͲ408 Euspira heros 19.02 19.77 0.44 CBͲ409 Euspira heros 15.28 16.21 0.4 127 CBͲ410 Euspira heros 15.75 15.79 0.39 CBͲ411 Euspira heros 14.45 14.61 0.34 CBͲ412 Euspira heros 5.42 5.85 0.35 CBͲ413 Euspira heros 11.78 11.99 0.28 CBͲ414 Euspira heros 16.16 16.73 0.75 CBͲ415 Euspira heros 13.88 14.39 0.56 CBͲ416 Euspira heros 13.83 14.24 0.39 CBͲ417 Euspira heros 11.88 12.11 0.37 CBͲ418 Euspira heros 8.19 8.25 0.27 CBͲ419 Euspira heros 8.22 8.56 0.24 CBͲ420 Euspira heros 9.56 9.52 0.29 CBͲ421 Euspira heros 10.19 10.15 0.31 CBͲ422 Euspira heros 7.46 8.33 0.19 CBͲ423 Euspira heros 8.5 9.39 0.26 CBͲ424 Euspira heros 11.19 10.84 0.32 CBͲ425 Euspira heros 8.51 9.11 0.25 CBͲ426 Euspira heros 10.3 9.91 0.31 CBͲ427 Euspira heros 8.52 7.96 0.24 CBͲ428 Euspira heros 9.48 9.28 0.33 CBͲ429 Euspira heros 7.38 7.32 0.31 CBͲ430 Euspira heros 11.63 11.69 0.34 CBͲ431 Euspira heros 6.83 6.64 0.32 CBͲ432 Euspira heros 7.11 6.88 0.27 CBͲ433 Euspira heros 5.95 6.39 10.4 CBͲ434 Euspira heros 5.8 5.92 0.23 CBͲ435 Tectonaticapusilla 4.3 4.66 0.27 CBͲ436 Naticarius canrena 16.47 17.19 0.52 CBͲ437 Naticarius canrena 18.13 17.89 0.73 CBͲ438 Naticarius canrena 17.83 16.93 0.57 128 CBͲ439 Naticarius canrena 26.59 29.1 0.82 CBͲ440 Naticarius canrena 17.99 17.59 0.58 CBͲ441 Naticarius canrena 14.08 14.34 0.58 CBͲ442 Neverita duplicata 22.45 19.19 0.62 CBͲ443 Neverita duplicata 34 28 0.64 CBͲ444 Naticarius canrena 17.35 17.53 0.58 CBͲ445 Naticarius canrena 12.7 13 0.35 CBͲ446 Naticarius canrena 16.56 16.8 0.46 CBͲ447 Naticarius canrena 18.58 18.39 0.49 CBͲ448 Naticarius canrena 14.74 14.81 0.42 CBͲ449 Naticarius canrena 18.79 19.11 0.66 CBͲ450 Naticarius canrena 14.51 13.59 0.55 CBͲ451 Naticarius canrena 15.32 14.78 0.95 CBͲ452 Naticarius canrena 19.84 21.04 0.61 CBͲ453 Naticarius canrena 18.27 18.73 0.55 CBͲ454 Naticarius canrena 23.01 22.39 0.47 CBͲ455 Naticarius canrena 28.63 29.84 0.82 CBͲ456 Naticarius canrena 20.47 21.09 0.73 CBͲ457 Naticarius canrena 15.01 14.69 0.44 CBͲ458 Naticarius canrena 15.66 15.03 0.46 CBͲ459 Naticarius canrena 12.44 12.13 0.53 CBͲ460 Naticarius canrena 14.58 14.71 0.38 CBͲ461 Naticarius canrena 14.53 12.84 0.44 CBͲ462 Naticarius canrena 17.21 17.04 0.63 CBͲ463 Naticarius canrena 14.16 13.65 0.69 CBͲ464 Naticarius canrena 16.27 16.28 0.47 CBͲ465 Naticarius canrena 17.66 17.48 0.51 CBͲ466 Naticarius canrena 16.22 16.06 0.38 CBͲ467 Naticarius canrena 19 18.98 0.79 129 CBͲ468 Naticarius canrena 15.42 15.65 0.39 CBͲ469 Naticarius canrena 14.57 14.41 0.64 CBͲ470 Naticarius canrena 17.76 18.15 0.73 CBͲ471 Naticarius canrena 17.94 18.63 0.47 CBͲ472 Naticarius canrena 16.06 15.46 0.52 CBͲ473 Naticarius canrena 14.19 13.65 0.59 CBͲ474 Naticarius canrena 14.96 14.42 0.46 CBͲ475 Euspira heros 22.46 23.1 0.5 CBͲ476 Euspira heros 18.77 19.12 0.53 CBͲ477 Euspira heros 24.66 25.28 0.43 CBͲ478 Euspira heros 16.26 17.39 0.52 CBͲ479 Euspira heros 22.03 22.22 0.61 CBͲ480 Euspira heros 19.45 19.32 0.66 CBͲ481 Neverita duplicata 32.35 25.4 0.55 CBͲ482 Neverita duplicata 18.3 15.35 0.49 4.63 1.95 2 CBͲ483 Neverita duplicata 16.46 14.07 0.56 CBͲ484 Naticarius canrena 23.84 24.76 0.68 CBͲ485 Naticarius canrena 18.38 18.18 0.43 CBͲ486 Naticarius canrena 16.85 16.7 0.68 CBͲ487 Naticarius canrena 17.34 17.37 0.77 CBͲ488 Naticarius canrena 15.54 15.13 0.52 CBͲ489 Naticarius canrena 17.35 16.95 0.52 CBͲ490 Naticarius canrena 17.25 17.84 0.71 CBͲ491 Naticarius canrena 14.62 14.19 0.44 CBͲ492 Naticarius canrena 12.9 13.26 0.32 CBͲ493 Naticarius canrena 16.89 16.91 0.44 CBͲ494 Naticarius canrena 14.81 15.13 0.45 CBͲ495 Naticarius canrena 15.26 14.42 0.6 CBͲ496 Naticarius canrena 13.51 13.65 0.98 130 CBͲ497 Naticarius canrena 18.14 18.58 0.69 CBͲ498 Neverita duplicata 17.47 15.03 0.46 CBͲ499 Neverita duplicata 19.4 15 0.36 CBͲ500 Neverita duplicata 17.44 13.82 0.43 CBͲ501 Neverita duplicata 14.6 11.52 0.51 CBͲ502 Neverita duplicata 18.19 14.88 0.49 CBͲ503 Neverita duplicata 14.47 12.65 0.86 CBͲ504 Neverita duplicata 14.06 10.9 0.35 CBͲ505 Neverita duplicata 15.53 12.2 0.9 CBͲ506 Neverita duplicata 13.2 10.76 0.38 CBͲ507 Neverita duplicata 10.86 9.29 0.31 CBͲ508 Neverita duplicata 12.02 9.53 0.31 CBͲ509 Naticarius canrena 12.77 12.15 0.39 CBͲ510 Euspira heros 18.42 20.78 0.32 CBͲ511 Euspira heros 14.28 15.22 0.35 CBͲ512 Euspira heros 13.79 14.06 0.41 CBͲ513 Euspira heros 12.66 12.88 0.34 3.293846 131 Locality RazierFarmNC DateCollected VMNH NearLumberton,NC Formation Duplin mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External RFͲ1 Naticarius canrena 33.96 33.14 0.7 RFͲ2 Neverita duplicata 49.83 44.34 1.11 RFͲ3 Euspira heros 35.23 36.09 0.57 132 RFͲ4 Euspira heros 36.71 36.12 0.88 RFͲ5 Euspira heros 23.07 24.05 0.85 RFͲ6 Euspira heros 31.64 35.09 0.75 RFͲ7 Naticarius canrena 28.73 28.69 0.77 RFͲ8 Neverita duplicata 31.58 28.33 0.62 RFͲ9 Neverita duplicata 39.31 35.17 0.95 RFͲ10 Euspira heros 32.49 33.63 0.71 RFͲ11 Naticarius canrena 25.76 25.84 0.56 RFͲ12 Naticarius canrena 18.84 18.55 0.67 RFͲ13 Euspira heros 16.74 17.58 0.55 RFͲ14 Euspira heros 33.15 33.52 0.69 RFͲ15 Naticarius canrena 29.7 30.6 0.79 RFͲ16 Euspira heros 36.99 36.76 0.69 RFͲ17 Euspira heros 40.55 39.46 1.06 RFͲ18 Euspira heros 28.89 29.67 0.44 RFͲ19 Euspira heros 24.02 23.86 0.35 RFͲ20 Euspira heros 22.07 21.4 0.46 RFͲ21 Euspira heros 41.58 41.42 0.98 RFͲ22 Euspira heros 23.03 24.41 0.55 RFͲ23 Euspira heros 25.32 25.47 0.82 RFͲ24 Euspira heros 27.96 27.42 0.91 RFͲ25 Naticarius canrena 22.33 21.72 0.61 RFͲ26 Euspira heros 25.94 26.66 0.65 RFͲ27 Euspira heros 24.83 26.13 0.53 RFͲ28 Euspira heros 29.43 25.6 0.48 RFͲ29 Euspira heros 23.69 25.65 0.89 RFͲ30 Euspira heros 25.83 26.93 0.87 RFͲ31 Euspira heros 24.94 26.17 0.3 RFͲ32 Naticarius canrena 23.04 22.88 0.69 133 RFͲ33 Naticarius canrena 31.5 31.86 0.67 RFͲ34 Euspira heros 24.51 25.61 0.54 RFͲ35 Neverita duplicata 29.18 25.66 0.76 RFͲ36 Neverita duplicata 19.45 14.52 0.64 RFͲ37 Neverita duplicata 22.07 19.37 0.46 RFͲ38 Neverita duplicata 50.42 44.41 0.92 RFͲ39 Neverita duplicata 49.59 42.74 0.89 RFͲ40 Neverita duplicata 23.66 19.55 0.61 4.09 2.37 3 4.09 Locality TearcoatBranchSC DateCollected VMNH WestofGabel,SC Formation Rayzor's(Duplin) mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External TBͲ1 Neverita duplicata 31.12 26.68 0.35 TBͲ2 Euspira heros 26.16 27.13 0.45 TBͲ3 Euspira heros 21.98 24.57 0.46 134 TBͲ4 Euspira heros 21.57 25.81 0.62 TBͲ5 Euspira heros 17.61 20.46 0.38 TBͲ6 Euspira heros 29.71 30.96 0.98 TBͲ7 Euspira heros 20.25 20.67 0.48 TBͲ8 Euspira heros 25.83 24.81 0.62 TBͲ9 Euspira heros 28.71 31.01 0.69 TBͲ10 Naticarius canrena 21.69 20.7 0.45 TBͲ11 Lunatia perspectiva 7.43 8.64 0.28 TBͲ12 Lunatia interna 5.8 6.12 0.28 TBͲ13 Lunatia interna 9.94 10.6 0.32 TBͲ14 Euspira heros 6.41 6.77 0.27 TBͲ15 Lunatia interna 5.8 5.94 0.21 TBͲ16 Tectonatica pusilla 2.98 3.6 0.16 TBͲ17 Tectonatica pusilla 4.01 4.57 0.23 TBͲ18 Tectonatica pusilla 4.16 4.59 0.15 TBͲ19 Tectonatica pusilla 5.44 6.36 0.21 TBͲ20 Lunatia perspectiva 8.58 10.45 0.39 TBͲ21 Euspira heros 6.9 7.34 0.1 TBͲ22 Neverita duplicata 34.18 31.8 0.5 TBͲ23 Neverita duplicata 21.59 17.62 0.46 4.71 2.3 3 TBͲ24 Neverita duplicata 14.81 10.73 0.31 TBͲ25 Neverita duplicata 10.94 8.9 0.32 TBͲ26 Neverita duplicata 12.81 9.99 0.19 TBͲ27 Neverita duplicata 8.16 6.58 0.18 TBͲ28 Neverita duplicata 27.99 24.2 0.65 TBͲ29 Euspira heros 26.35 29.87 0.73 TBͲ30 Naticarius canrena 33.73 33.07 0.62 TBͲ31 Naticarius canrena 21.54 20.97 0.38 3.19 2.03 2 TBͲ32 Naticarius canrena 21.49 21.35 0.48 135 TBͲ33 Euspira heros 31.05 32.7 0.5 TBͲ34 Euspira heros 34.07 34.89 0.87 TBͲ35 Neverita duplicata 29.01 26.64 0.72 TBͲ36 Euspira heros 37.81 40.15 0.58 TBͲ37 Euspira heros 35.03 36.3 0.59 TBͲ38 Euspira heros 28.9 28.08 1.21 TBͲ39 Naticarius canrena 34.94 33.38 0.78 TBͲ40 Euspira heros 24.12 25.07 0.69 TBͲ41 Euspira heros 30.47 32.18 0.46 TBͲ42 Euspira heros 22.12 24.93 0.55 TBͲ43 Euspira heros 27.86 30.06 0.57 TBͲ44 Euspira heros 27.94 30.15 0.4 TBͲ45 Euspira heros 27.6 27.52 0.57 TBͲ46 Euspira heros 28.4 31.35 0.67 TBͲ47 Euspira heros 22.99 27.15 0.44 TBͲ48 Neverita duplicata 39.06 37.31 1.4 TBͲ49 Neverita duplicata 24.13 20.74 0.59 TBͲ50 Neverita duplicata 27.44 22.9 0.59 TBͲ51 Neverita duplicata 29.2 25.25 0.62 TBͲ52 Neverita duplicata 30.28 25.07 0.77 TBͲ53 Neverita duplicata 33.78 27.59 0.64 3.95 136 Locality Wilmington DateCollected VMNH SiteatRandallParkway,Wilmington,NC Formation upperWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External WLͲ1 Neverita duplicata 36.61 33.81 0.92 WLͲ2 Neverita duplicata 51.05 50.59 0.9 WLͲ3 Neverita duplicata 51.67 45.75 0.71 137 WLͲ4 Neverita duplicata 39.2 35.94 0.96 WLͲ5 Neverita duplicata 28.61 25.91 0.51 WLͲ6 Neverita duplicata 21.34 18.63 0.35 WLͲ7 Neverita duplicata 20.75 16.7 0.73 WLͲ8 Neverita duplicata 24.33 21.1 0.56 WLͲ9 Neverita duplicata 24.9 20.57 0.6 WLͲ10 Neverita duplicata 39.06 38.42 0.68 WLͲ11 Neverita duplicata 43.47 41.6 1 WLͲ12 Neverita duplicata 31.56 28.07 0.53 WLͲ13 Neverita duplicata 36.04 34.99 0.72 WLͲ14 Neverita duplicata 40.45 35.69 0.61 WLͲ15 Neverita duplicata 17.2 15.05 0.47 WLͲ16 Neverita duplicata 27.5 24.37 0.59 WLͲ17 Neverita duplicata 32.29 30.54 0.62 WLͲ18 Neverita duplicata 23.2 16.67 0.51 WLͲ19 Neverita duplicata 30.6 27.31 0.87 WLͲ20 Neverita duplicata 25.22 20.01 0.34 WLͲ21 Neverita duplicata 19.74 14.37 0.49 WLͲ22 Neverita duplicata 31.65 25.73 0.9 WLͲ23 Neverita duplicata 23.64 21.46 0.51 WLͲ24 Neverita duplicata 23.16 19.04 0.41 WLͲ25 Euspira heros 31.11 33.26 0.67 5.08 3.31 3 WLͲ26 Euspira heros 28.5 30.51 0.6 3.38 1.78 3 WLͲ27 Euspira heros 27.48 30.46 0.45 3.8 2.02 2 WLͲ28 Euspira heros 30.9 34.03 0.49 WLͲ29 Neverita duplicata 35.35 33.84 0.95 WLͲ30 Sinum chesapeakensis 15.11 10.65 0.31 WLͲ31 Tectonatica pusilla 4.7 5.25 0.28 WLͲ32 Tectonatica pusilla 4.42 5.35 0.27 138 WLͲ33 Tectonatica pusilla 4.45 4.86 0.24 WLͲ34 Tectonatica pusilla 6.44 7.03 0.21 WLͲ35 Tectonatica pusilla 4.33 4.41 0.26 WLͲ36 Tectonatica pusilla 4.33 4.5 0.12 0.88 0.59 4 WLͲ37 Tectonatica pusilla 3.39 4.47 0.25 WLͲ38 Tectonatica pusilla 4.43 4.76 0.26 WLͲ39 Tectonatica pusilla 3.15 3.26 0.18 0.99 0.64 4 WLͲ40 Euspira heros 5.02 4.31 0.14 WLͲ41 Euspira heros 3.88 3.49 0.19 1.65 1.33 3 WLͲ42 Tectonatica pusilla 4.29 4.31 0.12 1.11 1.06 4 WLͲ43 Tectonatica pusilla 3.35 3.47 0.15 WLͲ44 Tectonatica pusilla 3.52 3.69 0.16 WLͲ45 Tectonatica pusilla 3.32 3.66 0.19 WLͲ46 Tectonatica pusilla 3.24 3.51 0.17 WLͲ47 Tectonatica pusilla 3.57 3.66 0.18 WLͲ48 Euspira heros 3.95 3.1 0.44 1.55 5 Onlyusefor%Canniba WLͲ49 Tectonatica pusilla 3.05 3.08 0.16 0.93 0.58 3 WLͲ50 Tectonatica pusilla 3.04 3.24 0.18 WLͲ51 Tectonatica pusilla 2.72 2.85 0.19 WLͲ52 Tectonatica pusilla 3.23 3.67 0.3 WLͲ53 Tectonatica pusilla 3.35 3.4 0.16 WLͲ54 Tectonatica pusilla 3.78 3.99 0.23 WLͲ55 Tectonatica pusilla 3.41 3.32 0.33 WLͲ56 Tectonatica pusilla 3.37 3.52 0.08 WLͲ57 Tectonatica pusilla 2.75 2.99 0.25 WLͲ58 Tectonatica pusilla 3 3.01 0.22 WLͲ59 Tectonatica pusilla 2.87 2.67 0.16 WLͲ60 Tectonatica pusilla 3.03 3.3 0.16 WLͲ61 Tectonatica pusilla 2.99 3.14 0.21 139 WLͲ62 Tectonatica pusilla 2.78 2.99 0.09 WLͲ63 Tectonatica pusilla 2.96 3 0.11 WLͲ64 Tectonatica pusilla 3 2.88 0.1 2.152222 Locality TarHeelRiver DateCollected REU Near34°42'N78°44'W Formation Duplin mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External THͲ1 Neverita duplicata 34.89 30.27 0.57 THͲ2 Neverita duplicata 31.54 26.37 0.55 THͲ3 Neverita duplicata 37.1 31.94 0.69 140 THͲ4 Neverita duplicata 26.75 22.13 0.67 THͲ5 Euspira heros 19.14 20.91 0.71 THͲ6 Euspira heros 25.27 26.11 0.62 THͲ7 Euspira heros 19.52 19 0.56 THͲ8 Naticarius canrena 22.5 22.15 0.64 THͲ9 Neverita duplicata 23.63 20.45 0.6 THͲ10 Naticarius canrena 20.33 18.61 0.51 THͲ11 Neverita duplicata 21.54 18.07 0.61 THͲ12 Neverita duplicata 20.79 22.85 0.73 THͲ13 Euspira heros 27.45 28.23 0.71 THͲ14 Naticarius canrena 23.1 23.54 0.76 THͲ15 Euspira heros 27.86 29.86 0.39 THͲ16 Euspira heros 23.52 27.59 0.35 THͲ17 Neverita duplicata 33.1 29.47 1.05 THͲ18 Neverita duplicata 21.51 16 0.7 THͲ19 Euspira heros 16.31 15.74 0.68 THͲ20 Neverita duplicata 15.98 15.39 0.55 THͲ21 Neverita duplicata 16.26 11.46 0.82 THͲ22 Neverita duplicata 14.91 11.12 0.68 THͲ23 Neverita duplicata 13.99 13.26 0.63 THͲ24 Euspira heros 12.6 12.16 0.72 THͲ25 Euspira heros 19.88 19.35 0.42 THͲ26 Euspira heros 24.88 24.89 0.61 THͲ27 Euspira heros 21.65 23.2 0.61 THͲ28 Euspira heros 20.59 20.99 0.33 THͲ29 Euspira heros 17.22 17.39 0.82 141 Locality Walker'sBluff DateCollected REU Near34°42'N78°29'W Formation upperWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External WBͲ1 Neverita duplicata 28.29 22.77 0.49 4.76 2.74 3 WBͲ2 Neverita duplicata 31.34 29.19 0.73 WBͲ3 Neverita duplicata 32.16 31.44 1.03 142 WBͲ4 Neverita duplicata 43.67 47.17 1.24 WBͲ5 Neverita duplicata 27.04 21.99 0.67 WBͲ6 Neverita duplicata 25.88 23.65 1.05 4.49 2.66 2 WBͲ7 Neverita duplicata 30.15 26.82 0.74 WBͲ8 Neverita duplicata 15.9 12.85 0.49 WBͲ9 Neverita duplicata 15.5 12.44 0.51 WBͲ10 Neverita duplicata 23.96 20.69 0.78 WBͲ11 Neverita duplicata 49.9 46.06 1.05 WBͲ12 Neverita duplicata 56.41 52.38 1.15 WBͲ13 Neverita duplicata 64.39 52.71 0.73 WBͲ14 Neverita duplicata 25.35 20.77 0.82 WBͲ15 Neverita duplicata 57.22 54.06 1.25 WBͲ16 Neverita duplicata 32.24 26.03 0.63 WBͲ17 Neverita duplicata 22.18 18.46 0.62 WBͲ18 Neverita duplicata 32.18 27.24 0.94 WBͲ19 Neverita duplicata 54.23 46.56 0.74 WBͲ20 Neverita duplicata 21.24 18.23 0.94 WBͲ21 Neverita duplicata 55.12 56.51 1.75 WBͲ22 Neverita duplicata 39.84 37.53 1.42 WBͲ23 Neverita duplicata 47.23 38.89 1.04 WBͲ24 Neverita duplicata 28.72 23.98 0.52 WBͲ25 Neverita duplicata 60.77 56.29 0.96 WBͲ26 Neverita duplicata 25.02 20.16 0.81 WBͲ27 Neverita duplicata 34.02 32.04 0.53 6.32 3.49 1 WBͲ28 Neverita duplicata 63.92 62.13 1.58 WBͲ29 Neverita duplicata 32.08 30.08 1.09 WBͲ30 Neverita duplicata 35.86 33.47 1.23 5.19 143 Locality SnakeIsland DateCollected REU NearSnakeIsland,NC Formation lowerWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External SIͲ1 Euspira heros 61.91 64.59 1.02 SIͲ2 Euspira heros 47.6 46.13 0.45 4.54 2.48 5 SIͲ3 Euspira heros 25.36 26.49 0.94 144 SIͲ4 Euspira heros 32.43 33.38 0.8 SIͲ5 Euspira heros 42.74 44.39 0.42 SIͲ6 Euspira heros 27.13 27.49 0.53 SIͲ7 Euspira heros 24.03 24.55 0.77 SIͲ8 Euspira heros 62.09 62.58 1.27 SIͲ9 Euspira heros 39.56 40.63 0.69 5.1 3.35 1 SIͲ10 Euspira heros 17.51 16.91 0.49 SIͲ11 Naticarius canrena 25.58 25.29 0.52 4.14 2.15 1 SIͲ12 Euspira heros 41.75 42.83 0.8 5.23 2.4 3 SIͲ13 Neverita duplicata 66.33 52.82 0.76 SIͲ14 Neverita duplicata 65.5 50.58 0.76 SIͲ15 Neverita duplicata 56.39 44.87 0.94 SIͲ16 Neverita duplicata 45.04 44.05 0.99 SIͲ17 Neverita duplicata 64.94 54.41 0.92 SIͲ18 Neverita duplicata 41.44 36.67 0.81 SIͲ19 Neverita duplicata 25.67 23.54 0.91 SIͲ20 Neverita duplicata 30.81 25.91 0.61 SIͲ21 Neverita duplicata 57.77 52.87 0.94 SIͲ22 Neverita duplicata 56.31 48.62 1.12 SIͲ23 Neverita duplicata 55.97 47.26 1.96 SIͲ24 Neverita duplicata 57.25 48.02 0.97 SIͲ25 Neverita duplicata 56.81 48.64 1.82 SIͲ26 Neverita duplicata 66.49 62.77 1.35 SIͲ27 Neverita duplicata 30.92 29.96 0.66 SIͲ28 Neverita duplicata 86.68 64.7 1.54 SIͲ29 Neverita duplicata 57.55 48.91 0.7 SIͲ30 Neverita duplicata 27.06 20.46 1 SIͲ31 Neverita duplicata 23.03 17.68 0.75 SIͲ32 Neverita duplicata 24.45 18.9 0.48 145 SIͲ33 Neverita duplicata 32.91 23.88 0.98 SIͲ34 Neverita duplicata 22.29 19.01 0.65 SIͲ35 Neverita duplicata 14.28 11.78 0.8 SIͲ36 Neverita duplicata 23.52 18.38 0.71 5.61 3.05 2 SIͲ37 Neverita duplicata 20.04 16.39 0.91 SIͲ38 Euspira heros 37.91 40.22 0.62 5.8 3.26 5 SIͲ39 Euspira heros 46.23 47.15 0.54 4.36 2.75 2 SIͲ40 Euspira heros 17.24 17.76 0.51 SIͲ41 Naticarius canrena 41.3 43.09 0.76 SIͲ42 Sinum perspectivum 31.68 17.76 0.71 SIͲ43 Euspira heros 46.53 47.49 1.03 SIͲ44 Euspira heros 14.97 16.05 0.26 SIͲ45 Euspira heros 18.47 17.88 0.27 SIͲ46 Neverita duplicata 55.72 49.38 1.76 SIͲ47 Neverita duplicata 42.57 42.95 1.3 SIͲ48 Neverita duplicata 46.01 35.49 1.29 SIͲ49 Neverita duplicata 30.28 25.58 1.08 SIͲ50 Neverita duplicata 19.53 14.89 0.9 SIͲ51 Neverita duplicata 43.17 38.41 0.53 SIͲ52 Neverita duplicata 31.37 25.33 1.01 SIͲ53 Neverita duplicata 29.82 23.32 0.75 SIͲ54 Neverita duplicata 23.62 19.27 0.91 SIͲ55 Neverita duplicata 20.63 15.88 0.97 SIͲ56 Neverita duplicata 15.4 11.73 0.46 SIͲ57 Neverita duplicata 18.82 15.96 0.42 SIͲ58 Neverita duplicata 31.03 25.41 0.87 SIͲ59 Neverita duplicata 22.7 17.85 0.73 SIͲ60 Neverita duplicata 32.16 24.03 0.92 SIͲ61 Neverita duplicata 29.02 24.65 0.81 146 SIͲ62 Neverita duplicata 18.11 13.79 0.9 SIͲ63 Neverita duplicata 24.91 18.78 0.55 SIͲ64 Neverita duplicata 58.1 51.84 1.33 SIͲ65 Euspira heros 30.45 31.7 0.62 3.8 2.19 2 SIͲ66 Euspira heros 18.08 17.95 0.47 SIͲ67 Euspira heros 17.23 16.69 0.79 SIͲ68 Euspira heros 16.28 16.17 0.55 SIͲ69 Naticarius canrena 38.88 37.13 0.83 SIͲ70 Naticarius canrena 17.49 16.78 0.59 SIͲ71 Naticarius canrena 20.17 19.39 0.58 4.18 2.24 2 SIͲ72 Euspira heros 39.23 39.35 0.87 SIͲ73 Euspira heros 33.45 33.56 0.69 SIͲ74 Euspira heros 24.73 24.4 0.76 4.25 2.26 2 SIͲ75 Neverita duplicata 36.14 31.52 0.6 SIͲ76 Neverita duplicata 52.08 41.25 0.61 SIͲ77 Neverita duplicata 48.5 41.65 0.92 SIͲ78 Neverita duplicata 21.39 18.25 0.52 SIͲ79 Neverita duplicata 27.58 23.01 0.55 SIͲ80 Neverita duplicata 50.92 45.97 1.06 SIͲ81 Neverita duplicata 25.36 20.02 0.93 SIͲ82 Neverita duplicata 29.71 21.3 0.52 SIͲ83 Neverita duplicata 23.83 17.85 0.7 SIͲ84 Neverita duplicata 52.19 47.7 1.11 SIͲ85 Neverita duplicata 69.94 60.48 1.24 SIͲ86 Neverita duplicata 54.11 46.5 0.7 SIͲ87 Neverita duplicata 56.58 53.44 1.19 SIͲ88 Neverita duplicata 63.24 50.98 0.94 SIͲ89 Neverita duplicata 33.48 25.19 1.11 SIͲ90 Neverita duplicata 36.44 29.87 0.86 147 SIͲ91 Neverita duplicata 23.69 18.39 0.6 SIͲ92 Neverita duplicata 26.67 21.96 0.31 4.76 3.37 2 SIͲ93 Neverita duplicata 22.92 16.32 0.71 SIͲ94 Naticarius canrena 26.19 23.56 1.1 SIͲ95 Naticarius canrena 36.42 30.69 0.95 2.9 1.76 1 drilledtwice(3.26/1.64 SIͲ96 Naticarius canrena 30.5 33.75 1.18 SIͲ97 Naticarius canrena 43.68 43.12 0.88 SIͲ98 Euspira heros 17.62 17.24 0.53 SIͲ99 Euspira heros 22.35 21.91 0.69 SIͲ100 Euspira heros 26.11 25.61 0.43 3.09 1.55 3 4.443077 Locality KirbyPond DateCollected PRI Exactlocationunknown,SC Formation Duplin mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External KPͲ1 Neverita duplicata 57.84 51.29 1.07 KPͲ2 Neverita duplicata 33.02 28 0.74 KPͲ3 Neverita duplicata 37.59 31.03 1.43 148 KPͲ4 Neverita duplicata 36.8 33.27 1.44 KPͲ5 Neverita duplicata 32.44 28.42 0.93 KPͲ6 Neverita duplicata 37.96 31.07 1.42 KPͲ7 Neverita duplicata 42.73 35.21 1.44 KPͲ8 Neverita duplicata 35.01 32.63 0.77 KPͲ9 Neverita duplicata 35.74 29.32 1.19 KPͲ10 Neverita duplicata 31.2 28.59 1.21 KPͲ11 Neverita duplicata 34.99 28.85 0.81 KPͲ12 Neverita duplicata 36.17 28.34 1.13 KPͲ13 Neverita duplicata 32.47 28.28 0.97 KPͲ14 Neverita duplicata 33.39 27.14 0.79 KPͲ15 Naticarius canrena 31.9 30.09 0.86 KPͲ16 Neverita duplicata 33.84 27.65 0.79 KPͲ17 Neverita duplicata 30.59 24.18 0.96 KPͲ18 Neverita duplicata 34.77 32.15 1.29 KPͲ19 Neverita duplicata 32.41 25.49 1.23 KPͲ20 Neverita duplicata 29.77 23.84 1.08 KPͲ21 Neverita duplicata 28.25 23.68 0.9 KPͲ22 Neverita duplicata 25.11 20.21 0.83 KPͲ23 Neverita duplicata 22.16 17.72 0.75 KPͲ24 Neverita duplicata 29.02 26.41 1.08 KPͲ25 Neverita duplicata 18.2 13.49 0.74 KPͲ26 Neverita duplicata 23.24 18.99 0.67 KPͲ27 Neverita duplicata 26.95 22.57 0.81 KPͲ28 Neverita duplicata 17.56 13.87 0.91 KPͲ29 Neverita duplicata 31.77 25.36 1.06 KPͲ30 Neverita duplicata 31.7 26.8 0.86 KPͲ31 Neverita duplicata 26.45 22.51 1.02 KPͲ32 Neverita duplicata 18.78 15.13 0.58 149 KPͲ33 Neverita duplicata 15.81 11.96 0.73 KPͲ34 Neverita duplicata 5.33 4.05 0.21 KPͲ35 Tectonatica pusilla 3.79 3.84 0.13 KPͲ36 Tectonatica pusilla 2.95 3.14 0.21 KPͲ37 Tectonatica pusilla 3.8 3.3 0.17 KPͲ38 Tectonatica pusilla 2.84 3.2 0.23 KPͲ39 Tectonatica pusilla 3.4 3.86 0.22 KPͲ40 Tectonatica pusilla 3.35 3.47 0.23 KPͲ41 Tectonatica pusilla 3 3.22 0.25 KPͲ42 Tectonatica pusilla 2.28 2.58 0.17 KPͲ43 Tectonatica pusilla 2.2 2.6 0.15 KPͲ44 Tectonatica pusilla 3.05 3.11 0.23 KPͲ45 Tectonatica pusilla 2.23 2.31 0.28 KPͲ46 Naticarius canrena 28.56 29.18 0.71 KPͲ47 Naticarius canrena 31.63 31.83 1.02 KPͲ48 Naticarius canrena 27.17 25.17 1.15 KPͲ49 Naticarius canrena 29.84 29.29 0.64 KPͲ50 Naticarius canrena 19.37 17.43 0.52 KPͲ51 Naticarius canrena 29.79 29.38 0.61 KPͲ52 Naticarius canrena 21.09 20.38 0.53 KPͲ53 Naticarius canrena 19.07 19.36 0.89 KPͲ54 Naticarius canrena 29.54 27.56 0.7 3.19 1.98 1 KPͲ55 Naticarius canrena 15.79 15.07 0.47 KPͲ56 Naticarius canrena 30.09 30.11 0.97 KPͲ57 Naticarius canrena 17.43 17.5 0.69 KPͲ58 Euspira heros 26.62 26.92 0.49 KPͲ59 Euspira heros 27.34 26.81 0.67 KPͲ60 Euspira heros 21.31 21.37 0.93 KPͲ61 Euspira heros 24.78 22.4 0.58 150 KPͲ62 Euspira heros 32.36 32.92 0.63 KPͲ63 Euspira heros 32.12 30.54 0.8 KPͲ64 Euspira heros 18.15 19.75 0.53 KPͲ65 Euspira heros 11.44 13.58 0.54 KPͲ66 Neverita duplicata 32.99 28 1.4 KPͲ67 Neverita duplicata 14.4 12.02 0.79 KPͲ68 Neverita duplicata 38.82 29.8 1.21 KPͲ69 Neverita duplicata 22.83 17.51 0.62 KPͲ70 Neverita duplicata 22.39 16.41 0.6 KPͲ71 Neverita duplicata 21.44 16.46 1 KPͲ72 Neverita duplicata 33.35 26.82 0.98 KPͲ73 Neverita duplicata 30.48 22.99 0.79 KPͲ74 Naticarius canrena 17.63 16.02 1.08 KPͲ75 Naticarius canrena 24.41 25.1 0.74 KPͲ76 Neverita duplicata 29.91 26.94 0.78 KPͲ77 Neverita duplicata 40.41 37.13 1.13 KPͲ78 Neverita duplicata 33.7 27.87 0.67 KPͲ79 Neverita duplicata 33.26 26.53 0.91 KPͲ80 Neverita duplicata 39.37 35.78 1.19 KPͲ81 Neverita duplicata 23.1 18.21 0.59 KPͲ82 Neverita duplicata 19.02 14.79 1 KPͲ83 Neverita duplicata 29.06 23.2 0.64 KPͲ84 Neverita duplicata 35.31 26.44 0.98 KPͲ85 Neverita duplicata 25.76 21.09 1.25 KPͲ86 Euspira heros 22.8 23.08 0.64 KPͲ87 Lunatia perspectiva 23.57 26.2 0.65 2.88 1.54 2 KPͲ88 Naticarius canrena 19.27 21.31 1.11 KPͲ89 Naticarius canrena 28.82 27.75 1.2 KPͲ90 Neverita duplicata 33.93 35.6 1.47 151 KPͲ91 Neverita duplicata 31.34 26.86 0.79 KPͲ92 Neverita duplicata 24.33 17.09 1.29 KPͲ93 Neverita duplicata 39.43 34.5 1.26 KPͲ94 Neverita duplicata 26.19 21.55 1.18 KPͲ95 Euspira heros 39.15 39.11 0.57 KPͲ96 Neverita duplicata 40.81 34.3 1.01 KPͲ97 Neverita duplicata 19.07 14.26 0.68 KPͲ98 Neverita duplicata 29.49 24.18 0.99 KPͲ99 Neverita duplicata 24.38 19.65 0.86 KPͲ100 Naticarius canrena 32.29 30.08 1.4 KPͲ101 Euspira heros 23.33 25.26 0.62 KPͲ102 Euspira heros 22.24 23.29 0.37 KPͲ103 Euspira heros 27.92 30.18 0.71 3.035 Locality BavenHill DateCollected PRI NeuseRiver,NC Formation JamesCity mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External BHͲ1 Neverita duplicata 48.24 41.78 0.89 BHͲ2 Neverita duplicata 41.29 35.25 0.74 BHͲ3 Neverita duplicata 24.95 21.99 0.46 152 BHͲ4 Neverita duplicata 24.49 20.85 0.28 5.28 3.07 3 BHͲ5 Neverita duplicata 23.38 19.11 0.38 5.12 3.52 1 BHͲ6 Neverita duplicata 17.46 15.55 0.36 4.89 2.65 3 BHͲ7 Neverita duplicata 18.5 14.58 0.26 4.7 2.93 2 BHͲ8 Neverita duplicata 12.62 9.43 0.26 3.29 1.69 4 BHͲ9 Neverita duplicata 15.06 12.92 0.17 3.22 1.16 2 BHͲ10 Neverita duplicata 16.39 13.94 0.2 4.57 2.33 2 BHͲ11 Neverita duplicata 10.97 9.31 0.15 3.52 2.11 3 BHͲ12 Neverita duplicata 13.17 10.87 0.32 3.55 1.77 3 BHͲ13 Neverita duplicata 16.16 13.75 0.35 BHͲ14 Neverita duplicata 25.25 19.58 0.65 BHͲ15 Neverita duplicata 15.35 12.03 0.34 2.63 1.17 2 BHͲ16 Neverita duplicata 9.64 8.01 0.17 BHͲ17 Neverita duplicata 10.44 8.35 0.15 3.5 1.68 3 BHͲ18 Neverita duplicata 12.03 9.79 0.22 3.4 1.72 2 BHͲ19 Neverita duplicata 13.94 11.36 0.25 BHͲ20 Neverita duplicata 14.17 12.39 0.31 4.11 2.29 2 BHͲ21 Neverita duplicata 10.22 8.75 0.24 2.46 1.08 2 BHͲ22 Neverita duplicata 12.22 9.75 0.34 3.874286 153 Locality PRI1432NeuseRiver DateCollected PRI NeuseRiver,NC Formation JamesCity mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External NRͲ1 Neverita duplicata 30.36 25.87 0.51 NRͲ2 Neverita duplicata 25.65 21.8 0.41 5.65 3.46 2 NRͲ3 Neverita duplicata 25.28 21.57 0.44 154 NRͲ4 Neverita duplicata 22.43 18.02 0.43 4.42 2.69 2 NRͲ5 Neverita duplicata 22.98 18.36 0.6 NRͲ6 Neverita duplicata 15.22 13.15 0.4 4.91 3.05 4 NRͲ7 Neverita duplicata 28.36 22.61 0.44 4.16 2.44 2 NRͲ8 Neverita duplicata 25.19 20.53 0.35 5.28 2.58 3 NRͲ9 Neverita duplicata 17.8 13.82 0.43 4.48 2.68 3 NRͲ10 Neverita duplicata 20.32 17.58 0.56 NRͲ11 Neverita duplicata 4.35 3.81 0.13 NRͲ12 Neverita duplicata 16.64 13.5 0.37 4.44 2.27 3 NRͲ13 Neverita duplicata 21.63 18.02 0.4 NRͲ14 Neverita duplicata 17.73 15.93 0.26 NRͲ15 Neverita duplicata 18.79 15.27 0.35 4.33 2.44 2 NRͲ16 Neverita duplicata 19.21 15.71 0.7 NRͲ17 Neverita duplicata 14.83 12.38 0.22 NRͲ18 Neverita duplicata 10.4 7.94 0.2 2.3 1.56 3 NRͲ19 Neverita duplicata 12.5 10.47 0.29 3.52 1.78 3 NRͲ20 Neverita duplicata 14.78 12.53 0.35 4.35 2.58 3 NRͲ21 Neverita duplicata 11.8 9.4 0.21 NRͲ22 Neverita duplicata 11.65 10.18 0.25 2.3 1.25 3 NRͲ23 Neverita duplicata 9.22 7.69 0.16 3.23 1.79 2 NRͲ24 Neverita duplicata 5.67 4.9 0.17 NRͲ25 Neverita duplicata 5.72 4.82 0.2 1.58 0.57 4 NRͲ26 Tectonatica pusilla 4.99 5.61 0.27 NRͲ27 Neverita duplicata 5.63 5.11 0.27 NRͲ28 Tectonatica pusilla 4.17 4.52 0.22 NRͲ29 Tectonatica pusilla 4.52 5.1 0.2 NRͲ30 Neverita duplicata 4.46 3.78 0.24 NRͲ31 Neverita duplicata 3.61 3.18 0.29 NRͲ32 Neverita duplicata 3.04 2.5 0.11 0.96 0.74 3 155 NRͲ33 Neverita duplicata 3.95 3.25 0.11 3.727333 Locality Keith'sMarlPitNielsEddyLanding DateCollected PRI Near34°20'N78°05'W Formation upperWaccamaw mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External NLͲ1 Neverita duplicata 49.15 46.88 1.01 NLͲ2 Neverita duplicata 11.96 9.18 0.31 NLͲ3 Neverita duplicata 49.92 45.48 0.92 156 NLͲ4 Neverita duplicata 45.82 44.87 0.66 NLͲ5 Neverita duplicata 13.08 9.59 0.4 3.55 2.06 4 NLͲ6 Neverita duplicata 27.96 25.14 0.42 NLͲ7 Neverita duplicata 34.73 36.27 0.92 NLͲ8 Neverita duplicata 51.02 48.54 0.75 NLͲ9 Neverita duplicata 35.39 31.06 0.82 NLͲ10 Neverita duplicata 31.33 26.99 0.61 4.94 2.94 2 NLͲ11 Neverita duplicata 27.91 22.65 0.82 NLͲ12 Neverita duplicata 30.88 27.22 0.71 NLͲ13 Neverita duplicata 19.74 15.58 0.59 4.89 2.87 4 NLͲ14 Neverita duplicata 54.3 55.95 0.92 NLͲ15 Neverita duplicata 19.2 15.25 3 NLͲ16 Neverita duplicata 25.39 20.88 0.55 NLͲ17 Neverita duplicata 20.66 16.47 0.73 NLͲ18 Neverita duplicata 18.35 15.23 0.39 NLͲ19 Neverita duplicata 18.89 14.96 0.47 NLͲ20 Neverita duplicata 24.44 20.62 0.36 NLͲ21 Neverita duplicata 21.59 17.66 0.42 NLͲ22 Neverita duplicata 19.71 15.12 0.43 NLͲ23 Neverita duplicata 21.11 17.64 0.44 4.62 3.08 3 NLͲ24 Neverita duplicata 13.08 11.02 0.58 NLͲ25 Euspira heros 17.33 17.6 0.44 NLͲ26 Neverita duplicata 17.2 13.58 0.56 NLͲ27 Neverita duplicata 17.03 13.97 0.41 NLͲ28 Neverita duplicata 14.43 10.99 0.43 3.12 1.66 2 NLͲ29 Neverita duplicata 11.85 9.24 0.21 NLͲ30 Neverita duplicata 11.78 9.69 0.57 3.23 1.66 2 NLͲ31 Neverita duplicata 10.68 8.37 0.3 3.03 1.27 3 NLͲ32 Neverita duplicata 13.82 11.27 0.28 157 NLͲ33 Neverita duplicata 11.26 9.14 0.51 NLͲ34 Neverita duplicata 9.77 8.74 0.35 NLͲ35 Neverita duplicata 43.95 41.28 0.65 NLͲ36 Euspira heros 32.22 30.89 0.4 NLͲ37 Naticarius canrena 38.29 38.12 0.81 NLͲ38 Neverita duplicata 16.1 12.19 0.38 3.5 2.05 3 NLͲ39 Neverita duplicata 45.42 44.28 0.72 NLͲ40 Euspira heros 10.49 11.27 0.3 2.67 1.5 3 NLͲ41 Euspira heros 14.07 15 0.28 2.58 1.25 1 NLͲ42 Euspira heros 24.01 23.64 0.51 3.04 1.45 2 NLͲ43 Euspira heros 23.9 24.08 0.53 NLͲ44 Neverita duplicata 50.55 47.02 0.64 NLͲ45 Neverita duplicata 39 36.25 0.73 NLͲ46 Neverita duplicata 28.52 23.96 0.58 NLͲ47 Euspira heros 21.52 21.02 0.46 NLͲ48 Euspira heros 14.61 14.43 0.53 NLͲ49 Neverita duplicata 26.03 20.36 0.5 NLͲ50 Euspira heros 23.16 24.05 0.45 NLͲ51 Euspira heros 17.4 17.7 0.53 2.95 1.4 4 NLͲ52 Neverita duplicata 16.55 13.06 0.43 4.03 2.2 4 NLͲ53 Neverita duplicata 33.21 30.19 0.63 NLͲ54 Euspira heros 20.39 18.05 0.31 NLͲ55 Neverita duplicata 49.4 50.74 1.35 NLͲ56 Neverita duplicata 54.02 48.8 0.89 NLͲ57 Neverita duplicata 18.17 14.05 0.46 NLͲ58 Euspira heros 24.7 25.72 0.46 NLͲ59 Neverita duplicata 26.32 21.73 0.44 NLͲ60 Euspira heros 24.14 23.85 0.59 NLͲ61 Euspira heros 14.85 16.46 0.24 158 NLͲ62 Euspira heros 17.2 17.33 0.51 NLͲ63 Euspira heros 17.17 16.25 0.36 NLͲ64 Neverita duplicata 26.16 19.78 0.81 NLͲ65 Euspira heros 20.39 20.07 0.42 3.86 2.19 1 NLͲ66 Euspira heros 13.75 13.98 0.33 NLͲ67 Neverita duplicata 14.55 12.15 0.25 NLͲ68 Neverita duplicata 18.51 13.95 0.53 NLͲ69 Euspira heros 12.59 12.45 0.33 NLͲ70 Euspira heros 10.92 11.25 0.26 NLͲ71 Neverita duplicata 8.63 7.17 0.24 2.45 1.03 3 NLͲ72 Neverita duplicata 11.78 8.89 0.29 2.23 0.96 3 NLͲ73 Neverita duplicata 14.37 10.96 0.29 3.5 1.73 3 NLͲ74 Neverita duplicata 12.95 10.37 0.25 2.91 1.4 2 NLͲ75 Neverita duplicata 8.78 6.68 0.12 2.13 1.01 3 NLͲ76 Neverita duplicata 8.96 6.94 0.18 NLͲ77 Euspira heros 12.58 12.89 0.22 NLͲ78 Euspira heros 9.67 10.25 0.26 3.327895 159 Locality PRI1442NeuseRiver DateCollected PRI NeuseRiver,NC Formation JamesCity mm mm mm mm mm Diameter Diameter Hole Hole Sector Hole Notes Internal Specimen Genus Species Length Height Thickness External NR2Ͳ1 Neverita duplicata 19.94 17.11 0.63 NR2Ͳ2 Neverita duplicata 21.32 15.85 0.52 NR2Ͳ3 Neverita duplicata 19.24 14.17 0.39 160 NR2Ͳ4 Neverita duplicata 19.37 16.24 0.56 NR2Ͳ5 Neverita duplicata 15.03 13.77 0.35 3.25 2.1 4 NR2Ͳ6 Neverita duplicata 8.77 7.26 0.39 NR2Ͳ7 Neverita duplicata 16.38 12.52 0.43 incDH NR2Ͳ8 Neverita duplicata 18.14 14.51 0.32 4.27 3.06 3 NR2Ͳ9 Neverita duplicata 14.88 11.75 0.61 NR2Ͳ10 Neverita duplicata 8.14 5.77 0.27 3.5 2.96 5 NR2Ͳ11 Neverita duplicata 5.85 5.75 0.33 NR2Ͳ12 Neverita duplicata 20.14 17.75 0.57 NR2Ͳ13 Neverita duplicata 8.82 7.12 0.19 NR2Ͳ14 Neverita duplicata 5.58 5.1 0.17 NR2Ͳ15 Neverita duplicata 5.71 4.72 0.24 1.4 0.92 3 NR2Ͳ16 Neverita duplicata 6.3 5.53 0.13 NR2Ͳ17 Neverita duplicata 10.65 8.23 0.21 NR2Ͳ18 Neverita duplicata 25.07 17.94 0.44 NR2Ͳ19 Neverita duplicata 25.33 21.19 0.52 NR2Ͳ20 Neverita duplicata 15.56 14.11 0.42 3.105 161