Inter and Intraspecific Morphological Disparity of Crinoid

Inter- and Intraspecific Morphological Variation of Crinoid Columnals in Relation to Water Depth in the Type Cincinnatian (Upper Ordovician)

A thesis submitted to

The Graduate School of the University of Cincinnati

in partial fulfillment of the requirements for the degree of

MASTER OF SCIENCE

in the Department of Geology of the College of Arts and Sciences

2005 by Bradley Deline

B.S. University of Michigan, Ann Arbor, 2003

Committee Chair David L. Meyer

Abstract

Crinoid columnals are a major constituent in the Upper Ordovician fossil assemblage of the Cincinnati Arch Region. Several species of Cincinnatian crinoids are identifiable based on columnal morphology alone. Disarticulated columnals of two crinoids were measured throughout a 68-meter section of the Kope and lower Fairview

Formations to examine the relationship between columnal morphology and sea level fluctuations. The columnal diameter of two disparid crinoids increased in the upward shallowing sequence. Detrended Correspondence Analysis axis 1 scores computed using columnal measurements of two crinoids correlated significantly with a proxy for depth.

Therefore, crinoid columnals may provide a metric for the study of small-scale sea level fluctuation in a depositional sequence. A larger scale study showed similar morphological shifts in five taxa of crinoids, but to differing degrees. The morphologic shifts in the columnals are likely due to differences in nutrient levels and flow regimes between depths.

Table of Contents

List of Figures 2 List of Tables 3 Acknowledgments 4 Introduction 5 Materials and Methods 8 Results 13 Discussion 17 Summary 24 References 26 Appendix 1 (Species Description) 33 Appendix 2 (Crenularium Area Calculation) 40 Appendix 3 (Localities) 43 Appendix 4 (Data) 44 Figure Captions 54 Figures 59 Table Captions 80 Tables 81

LIST OF FIGURES

Figure 1. SEM of Disparid columnals

Figure 2. The effects of applying a moving average and coarsening on the Lithology and the Faunal gradient of the K445 section

Figure 3. SEM of Monobatherid columnals.

Figure 4. SEM of Cladid and Diplobatherid columnals.

Figure 5. Diagram of columnal morphology and explanation of the calculation for lumen and crenularium circularity.

Figure 6. Diagram showing the differences between columns in lateral view.

Figure 7. Diagram showing the difference between column forms.

Figure 8. Relative depth curve.

Figure 9. Comparison of crinoid morphology, lithology and faunal gradient for K445.

Figure 10. PCA axis 1 compared with PCA axis 2

Figure 11. PCA axis 2 compared with relative depth.

Figure 12. PCA axis 2 compared with relative depth for disparids.

Figure 13. Averages of PCA axis 2 compared with relative depth.

Figure 14. Averages of PCA axis 2 compared with stratigraphic position.

Figure 15. Average depth compared with average PCA axis 2 score.

Figure 16. Range in columnal diameter of Ectenocrinus simplex in the K445 section.

Figure 17. The effects of other crinoids on Cincinnaticrinus and Ectenocrinus.

Figure 18. Comparing Relative Depth as well A PCA axis 2 score between pinnulate and nonpinnulate crinoids.

LIST OF TABLES

Table 1. Crinoid Occurrences.

Table 2. Characters used in the Polar coordinate analysis.

Table 3. Result of the correlations in the K445 analysis.

ACKNOWLEDGMENTS

I would like to foremost thank my advisor David Meyer as well as the other members of my committee Carlton Brett and Arnold Miller. My project was much improved due to discussions with Tomasz Baumiller, Forest Gahn, and Patrick

McLaughlin. I would also like to thank Forest Gahn, Brenda Hanke, and Kendall Hauer for access to museum collections. Steve Holland, Jim Brower and Arnold Miller provided access to data, which allowed me to test ideas and broaden the scope of my project. Stacy Deline provided valuable technical assistance in preparation of several of the figures. Funding for this project was provided in part by the University of Cincinnati

Geology Department and the University of Cincinnati Graduate Student Governance

Association. Finally, I would like to thank my wife and family for their support throughout this project.

Introduction

Traditionally, the preferred method to identify small-scale sea level oscillations

within the alternating shales and limestones of the Upper Ordovician Cincinnatian Series

was to track changes in the ratio of shale to limestone (Weiss and Norman, 1960; Ford,

1967). Recently, the use of event beds and cyclicity has helped to create a better-

resolved picture of small-scale eustatic fluctuations (Brett and Algeo 2001). Lithology

alone can be misleading because differing local patterns of sedimentation or variations in

the frequency and intensity of storms (Holland et al., 2001) could give signals through a section that are unrelated to shifts in sea level. Therefore, it is important to include many different sources of information to track these small-scale patterns. In particular, fossil faunal distributions have been shown to track minor sea level shifts that are not detected by examination of the lithology alone (Holland et al., 2001, Miller et al., 2001). Faunal

gradients are based on the ecological preferences of organisms and during short time

intervals these preferences are assumed to remain static.

Subtle differences in environments (flow rate, nutrients, light penetration) that

favor certain taxa over others in a particular area should also cause intraspecific

differences (ecophenotypic variation) along depth gradients. These effects could

manifest themselves in many different ways, but they may be observable in the

morphology of the animal. Therefore, studying morphology as it related to relative water

depth could provide information independent of lithology to help diagnose small-scale

changes and to provide insights into the biology of the organism.

Several different organisms within the type Cincinnatian have been studied to

detect changes in morphology that are related to water depth. Webber and Hanke

(submitted) have shown that there is an anteromedial shift in the eye location of the

trilobite Flexicalymene graulosa through the Kope and Fairview Formations that tracks the faunal gradient presented by Miller et al. (2001). Daley (1993) showed that shell size of the bivalve Ambonychia tracked water depth based on facies models across the

Cincinnatian, but the brachiopod Rafinesquina showed no such variation. However,

Daley showed that the frequency of geniculation (an upward bend in the shell) was much more frequent in populations from shallower facies than in deep-water facies (Daley,

1993). Different taxa, even within an ecological guild, may react very differently to changes in environment. However, it appears that most taxa are sensitive to small-scale water depth oscillations and thus other taxa should be examined for similar patterns.

Stalked crinoids (echinoderms) are filter feeders that are quite common throughout the fossil record in the shallow to marine environments, but they are mostly restricted to depths greater than 100 meters in today’s oceans (Hess, 1999). Crinoids are thought to be an ideal group for the study of small-scale changes in water depth and velocity because of well-documented relationships in the group between hydrodynamic conditions and morphologies associated with feeding. Crinoids need water flow to feed, but the water velocity cannot be too high or they would be forced to abandon feeding posture (Meyer, 1973, Messing, personal communication) or, in extreme cases, become dislodged (Rasmussen, 1977). Thus, crinoids need a distinctive flow regime to survive in a habitat and it has been documented that their morphology often reflects this (Kammer and Ausich, 1987).

Most studies exploring the relationship between crinoids and their environment focus on how distributions of crinoids change with water depth (Meyer et al. 2002), how

filter morphology relates to the flow regime (Ausich, 1980; Kammer and Ausich, 1987),

how morphology relates to substrate (Sprinkle and Guensburg, 1995), and the

evolutionary consequences of these different morphologies (Baumiller, 1993). The

difficulty with studying changes in filter morphology is that articulated specimens are

rare in the fossil record, making studies of fine scale changes in depth difficult. However,

disarticulated crinoid columnals are exceedingly common and underutilized sources of

information. The difficulty with using individual columnals is in species identification.

One attempt to characterize disarticulated columnals was made by Moore and

Jeffords (1968), in which individual columnals were described as “form taxa” without knowledge of calyx morphology. This is a valuable approach to attempt to realize the actual diversity of crinoids in a poorly preserved assemblage and has obvious uses in biostratigraphy (Le Menn, 1985; Le Menn, 1993). This approach also has significant drawbacks, such as the inability to determine whether form taxa represent single biological species. Moreover, there is a danger of taxonomic overrepresentation among crinoids that have xenomorphic stems (several different types of columnals in a single stem). Tracing the morphology of a single species through a depositional sequence using form taxa is therefore problematic.

Working in the type Cincinnatian, Meyer et al. (2002) showed that in a low diversity, well-studied area many crinoids can be identified to the species level using only columnal morphology, thereby eliminating the need for form taxa. Focusing on the study interval of Meyer et al. (2002), the present study examines fine-scale morphologic change in crinoid columnals in relation to changes in relative water depth. This information has two potential benefits. First, it may provide a new source of information about water

depth changes on the scale of depositional sequences. Second, it also complements recent

work on the distributions of crinoid morphologies along depth gradients, and may help to explain differences in species ranges and size distributions with changing water depth.

Materials and Methods

Meyer et al. (2002) recovered 102 fossiliferous limestone slabs from a 68 m

composite section in Campbell County, northern Kentucky, near Cincinnati, OH. These

sections, known as the K445 and Coney Island, encompass the majority of the Kope

Formation, as well as the lower 8 m of the overlying Fairview Formation. These two

units represent the lowermost formations of the type Cincinnatian (Upper Ordovician).

Both are composed of interbedded mudstone, skeletal packstones and grainstones and the

formations show characteristics of storm depositional processes (Jennette and Pryor,

1993, Brett et al. 2003). The Kope and Fairview formations correspond to the first two of

the six third-order depositional sequences (C1 and a portion of C2) reported in the

Cincinnatian (Holland and Patzkowsky, 1996) and have been further subdivided into

submembers representing fourth-order depositional sequences (Brett and Algeo, 2001).

Holland et al. (1997) present a locality map and a detailed description of the stratigraphic

sections. The 102 samples were taken at major fossiliferous horizons at approximately

half-meter intervals and are currently stored at the University of Cincinnati.

The 102 limestone slabs were inspected, and those that did not contain abundant

(n>20) crinoid columnals, or slabs that were abraded to the extent that columnal

measurements would be inaccurate, were excluded from the study. On each of the

remaining 36 slabs the columnal diameter, lumen diameter, and columnal height (A generalized diagram of columnal morphology is given in figure 5) were measured for approximately 30 columnals of the two most abundant crinoids in the section,

Cincinnaticrinus varibrachialus and Ectenocrinus simplex (Figure 1). On slabs with abundant columnals, a 5 by 5 cm2 template was placed on the surface and all columnals in the template were measured to ensure uniform sampling. If the slabs were less densely covered then all of the columnals on the slab were measured. If any of the columnals were still articulated or obviously associated, only one of the groupings was measured to prevent repeated sampling of the same individual.

A Detrended Correspondence Analysis (DCA) was conducted on the six measurements (columnal diameter, height and lumen diameter for both Cincinnaticrinus varibrachialus and Ectenocrinus simplex) for the 36 slabs, which condenses the morphologic information into a limited number of axes that describe the most variation in the data. Detrended Correspondence Analysis was developed for use on ecological data across gradients to eliminate the arch effect that is caused by normally distributed occurrences of organisms. It is expected that morphology would also be normally distributed across a gradient when the morphology in question is directly related to food gathering capability, as is the case in this study. However, Principle Coordinate

Analysis was also used on the K445 data and similar results were obtained (Pearson’s correlation between DCA and PCA results p = 0.006, Spearman’s rank correlation=0.07).

PC-ORD 4.27 (McCune and Mefford, 1997) was used for this analysis and all of the default settings for the axes were chosen, rescaling of axes (on), rescaling threshold (0),

and the number of segments into which the axes are divided for the detrending organism

algorithm (26).

The three measurements and the plot of the DCA axis 1 for the six measurements

were then compared to lithology (Meyer et. al, 2002) and the faunal gradient of the

section compiled by Miller et al. (2001). It should be noted that the faunal gradient DCA

axis presented in this paper differs slightly from the axis published by Miller et al.

(2001), because the original data set was re-analyzed here using only the sections

described in the present study. The lithology quantifies the proportion of shale, siltstone

and limestone as a moving average (Meyer et al. 2002). A 21-point moving average was

applied to the lithology and the faunal gradient to reduce the amount of noise relative to

the larger scale trend. The lithology and faunal gradient datasets were then limited to

stratigraphically corresponding points to the 36 crinoid morphology samples were taken.

This then created curves that had a similar resolution and could be compared (Figure 2).

To examine the relationship between water depth and columnal morphology on a

larger scale, ten crinoid species (Figure 1,3 and 4 show images of the columnals, the stratigraphic ranges can be seen in Table 1) were examined stratigraphically across the entire type Cincinnatian. The sample set was compiled from specimens at the Cincinnati

Museum Center and the collections at Miami University, as well as a few field collections (Appendix 3). Most samples were taken from large slabs and bulk collections of stem material to obtain a less biased sample than would be obtained measuring only well preserved complete specimens, but the use of columnals attached to the calyx was used when the columnal morphology is not distinctive (such is the case with Pycnocrinus dyeri and Glyptocrinus decadactylus). The crinoids were coded for ten different

characters, which accurately describe the morphology of the columnals, including four multi-state characters (Donovan, 1986,Table 2) and six continuous characters. While this sample set represents well the distribution of the common crinoids of the Cincinnatian in museum collections, it may not necessarily reflect actual distributions throughout the

Cincinnatian. This set of crinoids excludes several rare species that were not well represented in the museum collections examined. Moreover, one relatively common crinoid, Anomalocrinus, was excluded because it is an outlier in size and morphology such that it would have dominated the analyses.

These characters all have plausible functional relationships to strength and or flexibility of the stem (Donovan, 1990; Baumiller, 1997). The six continuous characters included the three mentioned above, as well as the circularity of the lumen and crenularium and the percent coverage of the articulation surface by the crenularium. The circularity of the lumen is equal to the ratio of the maximum and minimum radii of the lumen (Figure 5). The circularity of the crenularium is the ratio of the maximum and minimum distance between the centroid of the columnal and the outer edge of the crenularium. In both these measurements a circular case would give a value of one, while anything irregular (e.g. ellipsoid or pentagonal) would be greater than one. In most cases calculating the percent coverage of crenulation is simply the area between two circles defined as the outer and inner rim of the crenularium divided by the area of the articulation surface. However, when the crenularium is not circular, the area is estimated based on the overall shape of the crenulation. For example, Iocrinus subcrassus has a petaloid areola with the crenulation covering the rest of the articulation surface. Treating

the area between petals as five triangles and dividing that area by the area of the articulation surface in this case estimates the percent crenulation.

The multistate and binomial characters are consistent for all individuals within a species. The first character is the geometry of the column outline (circular, pentagonal, square). The second is the type of articulation displayed on the columnal: symplexial

(crenulated articulation surface) or petaloid (petal pattern on the articulation with very little indication of crenulation as seen in Cincinnaticrinus varibrachialus, figure 1). The third categorical variable is the lateral shape of the columnals (Figure 6): planar (the side of the columnal is perpendicular to both the top and bottom articulation surfaces), round convex (the side of the columnal is rounded so that the middle of the columnal protrudes farther than the intersection with the top or bottom articulation surfaces), or angular convex (the side of the columnal is planar, but the angles between the sides of the columnal and the articulation surfaces are different for the top and the bottom of the columnal). The last categorical character is the column form (Figure 7): holomorphic

(uniform columnal size) or heteromorphic (nodal and one internodal or two or more internodals). To reduce the large range in values in the continuous characters relative to the categorical characters the continuous characters were converted to the number of standard deviations away from the mean for each character. A Principle Component

Analysis (PCA) of this data set was then conducted using PCOrd again using the default settings (cross products matrix was created using correlation, and calculate scores for traits by weighted average (on).

A depth curve (Cuffey, 1992; Pope and Reed, 1997) that was created using difference in facies was used to create a rough estimate of relative depth for each member

of each formation. An average depth was obtained graphically for each member and a straight line was drawn to the bottom of the depth curve from each of the Stratigraphic members. An arbitrary scale was then placed along the bottom of the depth curve with the

Upper Whitewater Formation equal to one and the Fulton member or the Kope Formation equal to 13.5 (Figure 8). This scale was then used to compare trends in columnal morphology to relative depth. All statistics (Mann-Whitney U-test for comparing means and Spearman's rank correlation) conducted were done using PAST (Hammer et al. 2001) and a significance value of p=0.05 was used.

Results

Crinoid columnal morphology through the Kope-Fairview Sequence

An average of 22.6 Ectenocrinus simplex columnals and 17.6 Cincinnaticrinus varibrachialux columnals were measured per slab on the 36 samples from the K445 section of the Kope Formation. All of the slabs measure had both E. simplex and C. varibrachialus, with rare occurrences of Iocrinus subcrassus and Glyptocrinus decadactylus. The average diameter of the columnals increased 61 and 162 percent through the section for Ectenocrinus and Cincinnaticrinus, respectively. The lumen diameter and columnal height also increased for both crinoids upward through the section, and the DCA of these six measurements followed this trend toward larger columnals at the top of the section, as would be expected.

The DCA curve and the columnal measurements were compared with the lithology (percent limestone, shale, and silt) and the faunal gradient curve (Figure 9,

Table 3). The crinoid morphology curve showed a significant correlation with the

smoothed faunal gradient curve (Spearman’s rank Correlation, p=0.001). The columnals showed an increase in size (combination of height, columnal, and lumen diameter) during times that the fossil assemblage was dominated by shallower water taxa, thus it can be inferred that the columnals are showing a trend toward larger columnals in shallower water. These correlations remained significant when the four slabs with smaller sample sizes were removed (Spearman’s Rank Correlation, p=0.004) and when the height measurements were not included in the study (Spearman’s Rank Correlation, p=0.004).

The height consistently had a smaller sample size than the other two measurements because most of the crinoid columnals were preserved with the articular surface parallel to bedding with the majority of the columnal embedded within the slab, such that the height could not be measured. Overall, the correlation indicates that there is a trend for smaller columnals in deeper water and larger columnals in shallower water, particularly for Cincinnaticrinus, as indicated by the greater degree of morphologic change. It must be stressed that this correlation is with a proxy for depth and not a direct measurement of water depth itself.

Crinoid columnal morphology over the entire Cincinnatian

The results of the PCA analysis of the ten different species of crinoids showed two distinct axes that represented 29.74 and 20.07 percent of the variation (figure 10).

PCA Axis 1 was correlated with the type of articulation, areola circularity, and is negatively correlated with the lumen diameter and the circularity of the lumen. The average PCA axis 1 score was calculated for each sample (a single species at a single

stratigraphic occurrence) and was compared to relative depth and stratigraphic position.

PCA Axis 1 showed no relationship to relative depth as determined from figure 8

(Spearman’s rank correlation, p=0.115) or stratigraphic position (Spearman’s rank

correlation, p=0.279) and was thus less informative than axis 2, which correlated with

column diameter, lumen diameter and the height of the individual columnals.

When PCA axis 2 was compared to relative water depth the same trend (Larger

columnals in shallower water) appeared for the disparids as was seen in the K445 sample

(Spearman’s rank correlation, p<0.001, n=104) (figure 11). However, this trend is mostly

driven by Cincinnaticrinus (Spearman’s rank correlation, p<0.001)(figure 12), this is mostly due to Cincinnaticrinus occurring over a larger range of relative depths (n=57).

The monobathrids also showed a trend toward larger columnals in shallower water

(Spearman’s rank correlation between relative depth and Monobathrid PCA axis 2 scores, p<0.001, n=48). The cladids (Merocrinus curtus and Plicodendrocrinus casei) showed a strong correlation between PCA axis 2 and relative depth (Spearman’s correlation, p=0.002, n=16), but the trend is in the opposite direction (larger columnals in deeper water), this is caused by the the crinoids occurring at single stratigraphic intervals, such that the comparison is between the different species of crinoids. Merocrinus curtus has

robust columnals and occurs in deep water , such that the trend in cladids is opposite of

the other groups.

The averages per occurrence (stratigraphic stage) for the six crinoids that occurred

at multiple levels, except Plicodendrocrinus, followed the same pattern when compared

with the relative depth of each of the occurrences, namely, more robust columnals in

shallower water (figure 13). The exception to this pattern is Merocrinus, which occurs in

the deepest water facies, but has very robust columnals. Brett et al. (submitted)

hypothesized that Merocrinus may have grew a long robust column to elevate the crown above a low energy dysoxic seafloor, such that factors that are influencing columnal size in Merocrinus may not be important in other species. In addition, there seems to be a non-significant trend toward larger crinoid columnals in some species over time

(stratigraphic position) (Spearman’s rank correlation, p=0.109), but the signal was weaker than when the crinoids were compared to relative depth and there is less agreement in the trend in different species of crinoids (figure 14), indicating that the trend is responsive to depth and not a general increase in size over time.

When the crinoid distributions are compared to relative depth, several of the crinoids showed significant morphological shifts with a change in depth: Iocrinus, between relative depth 6 and 8.5, Mann-Whitney, p=0.02; Ectenocrinus, between relative depth 12.5 and 13.5, Mann-Whitney, p=0.013; between relative depth 7 and 8.5, Mann-

Whitney, p=0.034. Spearman’s correlation was used for Cincinnaticrinus, because it occurs in a greater number of Stratigraphic intervals (Spearman's correlation, p=0.004).

One crinoid species, Glyptocrinus fornshelli, showed a marginally significant shift between relative depth 6 and 8 (Mann-Whitney p=0.052) toward smaller columnals in deeper water.

The averages for PCA axis 2 and the average relative depth for the ten species of crinoids was also compared (Figure 15). There is no statistically significant relationship between depth and PCA axis two score (Spearman's correlation, p=0.318). However, there is a relationship between depth and the filter morphology. Pinnulate crinoids, mostly camerates in this study, have small branches that occur on each successive

skeletal element on the crinoid’s arms and have much denser filtration fans than non- pinnulate crinoids. The crinoids in this study with pinnules (denser fans, Glyptocrinus,

Pycnocrinus, Ptychocrinus, Xenocrinus) are seen in shallower water (Mann-Whitney, p=0.014) than the non-pinnulate crinoids (Cincinnaticrinus, Ectenocrinus, Iocrinus,

Merocrinus Plicodendrocrinus) in this study and there is a difference in columnal morphology between the two groups as seen in the PCA axis 1 scores (Mann-Whitney, p=0.0367), but no difference was found between pinnulate and non-pinnulate crinoids in regard to their PCA axis 2 scores (Mann-Whitney, p=0.531). In comparing the categorical variables to the average depth for the ten different species of crinoids it was found that the geometry (Mann-Whitney, p=0.897) and lateral shape (Mann-Whitney, p=0.302) did not show any relationship with the average depth of the species, but columns with two or more internodals occurr in shallower water than did species with a single nodal and internodal (Mann-Whitney, p=0.039).

Discussion

Possible Explanations for the Increase in Columnal Size

The increase in size of columnals in shallower water within a species, seen repeatedly in this study, could have several different explanations that are either ecophenotypic, evolutionary or taphonomic. A taphonomic explanation would predict that, in a higher energy setting, larger columnals might be preferentially preserved in fossil lags. This would likely increase the average columnal size and decrease the size range. However, the observed size distributions do not follow the prediction of the loss

of smaller columnals in facies that represent shallower depths. Instead, small columnals

are present in all of the samples and the change in the average between samples is driven

by an increase in the size of the largest columnals (i.e. an increase in the range of

columnal size) in shallower water. The range in columnal diameter is determined by the

difference between the maximum and minimum columnal diamters in a sample. This

measure is inherently related to the size of the sample, therefore, it was tested by

examining samples with 21-26 individuals of the most abundant crinoid in the upward

shallowing Kope formation. The range in columnal diameter of Ectenocrinus simplex is

significantly correlated with stratigraphic position (Figure 16, n=22, Spearman rank

correlation, p=0.03). Therefore, the trend is not a taphonomic artifact and biological

mechanisms must be explored.

The driving force behind the morphological variation is important to determine

for reasons other than understanding the controls on crinoid morphology. If the driving

force is ecophenotypic, then crinoid stem morphology would be particularly useful for

correlation and the detection of subtle water depth fluctuation, whereas, if there was an

evolutionary mechanism controlling the change in morphology, than longer-term trends

unrelated to facies might be anticipated.

The distinction between ecophenotypic variation and evolution is also important in understanding the relative importance of anagenesis and cladogenesis in evolution.

However, establishing a case of gradualism is exceedingly difficult (Jablonski, 2000). It should to be noted that the changes observed in this study are occurring in preexisting morphological features and are not evolutionary innovations that are often associated with cladogenesis. This distinction is important because many studies that examine

evolutionary rates (Schopf et al., 1975; Gingerich, 1983) and how these rates behave over

time (i.e. gradualism versus punctuated equilibrium) are based on character change. The

addition of characters and the morphologic alteration of preexisting characters may

represent differing evolutionary mechanisms and therefore must be examined separately.

A possible way to distinguish between ecophenotypic variation and evolution is to

trace a single bed or thin stratigraphic interval onshore to offshore, and determine if the crinoid columnals become smaller as depth increases. The identification of event beds and sedimentary cycles in the Cincinnatian (Brett et al., 2003) eases the difficulty in tracing a single bed.

The crinoid columnals in this study appear to superficially follow a gradualistic pattern of evolution (slowly and continually shifting stratigraphically), but given that the

relative importance of ecophenotypic and evolutionary variation cannot be assumed, it

needs to be tested, which is an avenue for future research. Considering the trend is in the

size of the columnals and not a change in the shape of an organism as well as the close

mirroring of morphology with water depth, there appears to be a strong ecophenotypic

component to the change in the columnals with depth.

There are three different explanations that will be considered, which could

produce a trend toward larger columnals in shallower water. The first is that in shallower

water there may be increased competition on disparid crinoids from camerate crinoids

with denser filtration fans that only rarely occur in deeper water facies. Increased

competition could drive body size in larger or smaller direction relative to the deeper

water morphology in order to avoid competition and create a tiered community (Ausich,

1980), or stunted growth due to a lower concentration of nutrients in the water column.

Meyer et al. (2002) examined the abundance patterns of the crinoids in the K445 section, which allows comparisons between abundance, first appearances and morphology. The appearance of Iocrinus and Glyptocrinus, as well as the peak abundances of Iocrinus are noted in fig 13. The first appearances have no relation to columnal morphology, but the peak abundance of Iocrinus did correspond to downward shifts in columnal diameter in both Cincinnaticrinus and Ectenocrinus, which is consistent with competition. However, there is also a large shift in the lithology that corresponds to the peak abundances of

Iocrinus, which also could be the cause of the morphology changes in Cincinnaticrinus and Ectenocrinus. The resolution of the curves is too low to distinguish whether either are directly related to the observed shift in morphology. Moreover, shifts seen at these points are short small-scale fluctuations that are opposed to the main trend. The opposition between the overall trend and the small-scale shift and the lack of resolution make it is impossible to connect first appearances and peak abundances with the patterns seen in Cincinnaticrinus and Ectenocrinus. Although competition might have some small effects on morphology, there is no evidence that the trends seen in this section are driven by competition.

Alternatively, changes in size may be a result of differences in resources across a gradient, such that with more nutrients available, the crinoids are able to grow to a larger size than they could in a deeper environment. An increase in runoff from the shoreline creates a higher nutrient level closer to shore. The difficulty with this hypothesis is that it is impossible to establish a nutrient gradient in the Ordovician.

The third option is that in a higher energy environment the crinoids need a larger column to withstand the added force from the elevated flow velocity. Drag is

proportional to the square of the flow velocity (equation 1, Vogel, 2003), such that a

shallower water (higher flow velocity) crinoid would be under a much larger drag force

than a crinoid with similar crown size in deeper water.

(1) Drag =coefficient*density of the fluid*Velocity of the fluid squared*filter area

Cincinnaticrinus and Ectenocrinus show different amounts of morphological

change with changes in depth. These two crinoids have very similar morphologies, but

have large differences in morphological plasticity. It is therefore probable that if the

energy were governing morphology that the more morphologically plastic species (in this

case Cincinnaticrinus) would have a longer stratigraphic range as the depth changes

throughout the Cincinnatian, i.e. the more plastic the species is the longer it will be

present in the shallowing sequence. This is not the case because both species are seen in

the same stratigraphic positions through the Cincinnatian, (provided that the species

designations are correct, see Appendix 1). The negative result could have several different explanations. The average water velocity may not be a driving force in morphological change, either because it is never a factor or influential storm events are rare enough that they do not affect individual morphology. The negative result could also occur because the disappearances of the taxa are not depth related, such that the test is being masked by other factors. Finally, average water velocity may be a factor, but it is not the most influential force in governing crinoid distributions. The flow velocity cannot be discounted in having an effect on crinoid morphology and the trends seen in

this study were most likely caused by changes in nutrient level and flow regimes between depths.

Patterns in Crinoid Morphology

Several different crinoid species in the Cincinnatian showed the same increase in columnal size in shallower water that was seen in the more detailed study of the Kope formation, while only one species showed the reverse trend (Plicodendrocrinus casei).

This trend, therefore, is occurring on a larger scale than a single depositional sequence.

Differences in the trends in size, aside from differences in sampling, show that not all species of crinoids are reacting to these changes in an identical manner. The robustness of the crinoid columnals may not be the only factor governing depth preference. The propensity for a homeomorphic stem (see figure 7 for differences in column form) differs with depth, which may be related to the flexibility of the stem as opposed to the strength of the stem (size). Crinoid species with homeomorphic stems can obtain a greater degree of curvature, which is displayed by the tightly coiled holdfast of Glyptocrinus decadactylus, than non-homeomorphic stems, which are often straight and rigid (e.g.

Merocrinus curtus). However, a pattern in stem characters does not prove a functional response. In a low diversity assemblage it is especially difficult to assess the generality of stem characters in relation to facies and the relative importance of stem robustness verses flexibility would be better tested in a higher diversity assemblage such as many

Mississippian occurrences.

The pattern observed in this study, of pinnulate crinoids appearing more

frequently in shallow water and non-pinnulate crinoids in deeper water, as also seen in

most better-preserved Mississippian crinoid assemblages (Kammer, 1985) corroborates

aerosol filtration theory. However, there was not a difference in columnal size between

pinnulate and non-pinnulate crinoids (Figure 17). This result is not surprising given the

differing responses to changes in water depth between species (Figure 13) and the

unknown importance of stem characters, other than size, to the overall strength and

flexibility of the column.

Branch density, defined by Ausich (1980), is the number of arm branches divided

by crown area and it has been shown to be a good metric for comparing fan density

between many crinoid communities including those of the Cincinnatian (Meyer et al.

2002), but it may also be useful in detecting patterns within species. Brower (2005)

showed in a Trenton population of Cincinnaticrinus varibrachialus that stem diameter is

inversely related to branch density, i.e., individuals with larger stems have a lower branch

density. By applying this relationship to the patterns seen in this study, it can be

extrapolated that crinoids with more open fans occurring in shallower water than those

with denser fans. This is the opposite of patterns observed between species which is

governed by aerosol filtration theory, which predicts larger, shallower water individuals

would have a higher branch density. The differences between branch densities in C.

varibrachialus may be small enough that it would not affect species distributions, but this relationship needs to be examined more closely.

SUMMARY

(1) Most crinoid species examined in this study showed a correlation between

columnal size and proxies for depth, such as the faunal gradient DCA axis 1 for

the K445 section or facies patterns for the larger scale study of the Cincinnatian.

Different species of crinoids reacted differently to changes in depth showing that

columnal size may not be the only stem character important in adjusting to small-

scale water depth oscillation.

(2) The columnals of some crinoid species are sensitive to depth-related changes and

when species identification is possible, columnals may be useful as indicators of

changes in depth.

(3) Morphology of crinoid columnals (columnal diameter, lumen diameter, and

columnal height) thus may be useable as an independent metric to add to studies

of small-scale sea level oscillations based on lithology or faunal gradients.

Moreover, it is feasible to create an independent depth curve based on the

morphology of the organisms within the stratigraphic section to compare with

curves created using other evidence.

(4) There is no evidence that competition plays a major role in the large-scale change

in columnal size, but interactions between species may play a role in small

morphological fluctuations.

(5) The repeated trend observed in the size of crinoid columnals is most likely

ecophenotypic variation, even though the driving force of the variation is unclear.

The mostly likely factors driving a change in morphology are changes in nutrient

supply and flow regime.

(6) Trends in filter morphology based on allometric relationships between branch

density and columnal diameter in Cincinnaticrinus variabrachialus show a pattern

(denser fans in deeper water) within species that is opposite to the pattern

observed between species (Denser fans in shallower water). These results are

preliminary and more study is needed.

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Appendix 1

Systematic Paleontology

Disparida, Moore and Laudon, 1943

Cincinnaticrinus varibrachialus, Warn and Strimple, 1977

Discussion: Cincinnaticrinus varibrachialus is a disparid crinoid that ranges from the base of the Kope Formation to the top of the Fairview Formation. The calyx of C. varibrachialus is conical and slender. The arms branch unevenly with the thicker limb often branching several more times during ontogeny (Warn and Strimple, 1977). The stem may reach lengths of a meter and end in a “lichenocrinus” type multiplated holdfast, which according to Warn and Strimple (1977) in mature individuals may become detached from the stalk. The columnals of C. varibrachialus are composed of five fused plates (pentameres) enveloped by a secondary covering of stereom, which causes the pentagonal columnal and lumen to appear round with each pentamere containing a single petal-shaped articular facet (Warn and Strimple, 1977). In profile, the column of C. varibrachialus can be distinguished from that of Ectenocrinus simplex by having a round lateral shape, as opposed to an angular profile.

Cincinnaticrinus pentagonus, Ulrich, 1882

Discussion: Cincinnaticrinus pentagonus is quite similar morphologically to

Cincinnaticrinus varibrachialus and ranges from the Maysville to the Whitewater

member of the Richmond. There is no addition of plates in the calyx during ontogeny,

such that the plate structure of small individuals is the same as mature individuals (Warn

and Strimple, 1977). The columnals of C. pentagonus are larger than those of C.

varibrachialus, but are virtually identical in form. The most visible difference between

C. pentagonus and C. variabrachialus is the ratio between the diameter of the proximal stem and the diameter of the cup. Cincinnaticrinus variabrachialus flares out from the stem to the cup, but in C. pentagonus the cup is only 1.4 times wider than the proximal stem in uncrushed specimens (Warn and Strimple, 1977) and often in non-pristine specimens the cup is no wider than the diameter of the stem. The distinction of the two species may be artificial because there is a continual gradient of calyx diameter to stem diameter ratio and columnal size. The gradient in columnal size is well illustrated in this study. Warn and Strimple (1977) hypothesized that C. varibrachialus is ancestral to C. pentagonus and the size change might be linked to an increase in current activity.

Ectenocrinus simplex, Hall, 1847

Discussion: Ectenocrinus simplex the most common crinoid in the Kope Formation.

Ectenocrinus simplex has a gracile unornamented cup with 10 straight, unbranching arms

that have small ramules (armlets) on every forth brachial. These ramules are rarely

displayed due to infolding of the arms. Several rapidly tapering columnals are

incorporated into the calyx with the growth zone directly below these columnals (Warn

and Strimple, 1977). The columnals of E. simplex are round with a pentagonal lumen and

a raised circular crenularium. The distal stem is trimeric (Brower, personal

communication) leading into a lichenocrinus holdfast (Warn and Strimple, 1977). The

holdfast of E. simplex differs from those of Cincinnaticrinus in two aspects: they are slightly larger (4-5 mm compared to 2-2.5 mm) and the plates of the upper wall of the holdfast are well differentiated (Warn and Strimple, 1977). As in Cincinnaticrinus, it has

been proposed that mature E. simplex shed their holdfast and the distal section of the stem

lies along the seafloor (Warn and Strimple, 1977).

Iocrinus subcrassus, Hall 1866

Discussion: Iocrinus subcrassus occurs abundantly from the Economy member of the

Kope Formation to the Waynesville Formation. Much like Cincinnaticrinus, I.

subcrassus appears to be a generalist that was able to live throughout the depth ranges of

the Cincinnatian. I. subcrassus has a pentagonal column with lobate facets and a round

lumen. The calyx and arms are small given the size of the column (Meek and Worthen,

1865). The arms branch multiple times, although they do not possess ramules as do other

Cincinnatian disparids. I. subcrassus has a coiled holdfast, which would allow I.

subcrassus to inhabit areas without hard substrates, if objects such as bryozoans were

available for attachment.

Cladida, Moore and Laudon, 1943

Merocrinus curtus, Ulrich, 1879

Discussion: Merocrinus curtus is the dominant crinoid in the Fulton submember of the

Kope Formation. This is the deepest section in the Cincinnatian and the morphology of

M. curtus is representative of deep water. The columnals are large with a large lumen

even though they are extremely thin giving the column a stiff appearance. This stiffness

is indicative of deeper water due to a more unidirectional flow that would not require as

much repositioning as in shallower water. The calyx of M. curtus is exceedingly small

given the size and length (up to 70 cm) of the column. The non-pinnulate arms branch

randomly multiple times, but without pinnules or armlets the density of the filter is quite

low.

Plicodendrocrinus casei, Brower, 1995

Discussion: Plicodendrocrinus casei occurs in the youngest section of the Cincinnatian in the Liberty and Whitewater Formations. P. casei has a stellate ornamented calyx and a large plicate anal sac that appears to be longer than the arms. The arms branch numerous times such that the branch density is high though the arms are non-pinulate (Brower,

1995). The columnals are strongly pentalobate with a round lumen and a fringe of articulation along the inner edge of the lobes.

Diplobathrida, Moore and Laudon, 1943

Ptychocrinus parvus, Hall, 1866

Discussion: Ptychocrinus parvus occurs from the Fairmount through the Corryville

Formation. Articulated individuals are rare though the columnals can be rather abundant in localized beds. The pinnulate arms have a high branching frequency creating a dense fan that is expected for a shallow water crinoid. The columnals are round with a round lumen, but are easily recognizable by having large nodals that that occur every seventh columnal.

Monobathrida. Moore and Laudon, 1943

Glyptocrinus decadactylus, Hall, 1847

Discussion: Glyptocrinus decadactylus is one of the most common shallow water crinoids in the Cincinnatian that ranges from the top of the Kope Formation to the top of the Fairview Formation. The cone-shaped calyx of G. decadactylus has a stellate ornamentation and is very robust compared to the disparids described previously. The rays branch twice before becoming detached from the calyx producing twenty unbranching arms. These arms have long pinnules that occur on each ossicle to produce a dense filtration fan even though the branch density is zero. The stem of mature G. decadactylus is approximately 20 cm in length and ends in a curled holdfast. This holdfast either wraps around an attached object or is heavy enough to act as an anchor lying on the seafloor. The columnals are round with a large lumen and a thin circular crenularium.

Glyptocrinus fornshelli, Miller, 1874

Discussion: Glyptocrinus fornshelli is a bizarre crinoid that is only known from a few horizons within the Liberty Formation. However, when G. fornshelli occurs, it is by far the dominant member in the assemblage. There are several morphological differences between G. fornshelli and G. decadactylus, such that there is some uncertainty as to its affinity. The calyx is highly ornamented with a bistellate pattern (i.e. a double ridged star-shaped ornamentation). Small individuals are cup shaped and have the prominent raised rays that are displayed by G. decadactylus. Large individuals loose the pronounced rays while the calyx is more oblong shaped creating a distinctively different morphology

from G. decadactylus. To compound this difference G. fornshelli has a pentagonal column with very thin columnals and the arm rays branch once as part of the calyx and four additional times after becoming free (Miller, 1883).

Pycnocrinus dyeri, Miller, 1883

Discussion: Pycnocrinus dyeri is very similar morphologically to Glyptocrinus decadactylus with two of distinguishing features. G. decadactylus has two secundibrachials between the first and second braches of the rays while P. dyeri has between 7-10 secundibrachials. The second arm branch of P. dyeri occurs shortly after becoming free resulting in twenty pinulate arms. The other difference is in the number and arrangement of interadial plates, which gives P. dyeri a globular shape as opposed to the angular calyx of G. decdactylus. The columnals of P. dyeri are indistinguishable from those of G. decdactylus.

Xenocrinus baeri, Miller, 1881

Discussion: Xenocrinus baeri can be distinguished from other Cincinnatian crinoids based on the unique characteristics of its stem and interadial areas. The stem of X. baeri is quadrangular with the four sides of the columnal bowing inwards. The lumen retains a round shape while the crenularium roughly follows the shape of the columnal. The interradial areas of X. baeri are slightly depressed and are composed of many small plates (Miller, 1881). The plates in the interradial and inter-secondary radial spaces are so small that the calyx can have as many as 550-600 plates (Miller, 1881). X. baeri has large radial plates that branch once in the fixed brachials and several times after the arms

become free. The pinnules are coarse and are composed of more than twelve plates each

(Miller, 1881), which form a dense filter on both sides of the arms.

Appendix 2

Equations for Percentage Area of the Columnal Covered by Crenularium

A.Terms

C =Thickness of the crenularium

CL = the length of the columnal.

CR = the radius of the columnal.

DA = the distance between the centroid of the columnal and the outer edge of the areola.

DAM = the maximum distance on the columnal between the centroid of the columnal and

the outer edge of the areola.

DC = the distance between the centoid of the columnal and the outer edge of the

crenularium.

LR = the radius of the lumen

PL = the length of the petals on the articulated surface.

B. Ectenocrinus simplex, Glyptocrinus decadactylus, Merocrinus curtus, Pycnocrinus dyeri, and Ptychocrinus parvus

If the columnal, lumen, areola and crenularium are all circular or nearly circular the percentage area of the columnal covered by crenularium (C%) is approximated by the equation:

(2) C%= (π DC2- π DA2) /((π CR2)-(π LR2))

C. Xenocrinus baeri

In the case that the columnal and crenularium are quadrangular and the lumen is circular the C% is approximated by the equation:

(3) C%=(CL*C*4)/(CL2- π LR2)

D. Plicodendrocrinus casei

Plicodendrocrinus casei the columnal is distinctly star-shaped with crenulation

the length of the lumen on the inside of the points of the star. The equation that was used

for P. casei is:

(4) C%=(LD*C*5)/((( π (LR+DC)2)+(LD*(DAM –DC)*1/2)*5)- π LR2)

E. Glyptocrinus fornshelli

Glyptocrinus fornshelli has a columnal the shape of a pentagon with a round

lumen and areola. The equation that was used for G. fornshelli is:

(5) C%= (π DC2- π DA2)/ ((5*1/2*(CL/2)* 0.688 (CL/2))- π LR2)

F. Iocrinus subcrassus

Iocrinus subcrassus has a pentagonal columnal like Glyptocrinus fornshelli and a

petaloid articular surface. The area in between the petals is covered by crenulation.

(6) C%= (5*(1/2)*CT* 0.688 (CL/2))/((5*1/2*(CL/2)* 0.688 (CL/2))- π LR2)

G. Cincinnaticrinus varibrachialus, Cincinnaticrinus pentagonus

Cincinnaticrinus has a round lumen and columnal with a petaloid articular surface. The crenulation is located between the petals, but it is subdued compared to

Iocrinus subcrassus.

(7) C%= (5*(1/2)*CT* 0.688 (LD/2 +PL))/ ((π CR2)-(π LR2))

Appendix 3

Localities

Field collections for the second part of the study were taken at the following localities. All other localities mentioned in appendix 4 were taken from museum records.

Indiana Route 1, IN (Southgate Hill in Franklin county, Indiana) Large road cuts on both sides of the road, 1-1.5 miles north of intersection at the town of South Gate and 1.9-2.4 miles south of intersection of Indiana Route 1 and U.S. Route 52 at Cedar Grove (Davis et al. 1998).

Stonelick Creek, OH. Stonelick Creek cuts upstream and downstream from the Ohio Route 131 bridge, near Newtonsville, Clemont County, Ohio (Davis et al. 1998).

Appendix 4

Data

4a. Dataset of average crinoid morphology from K445. (E: Ectenocrinus, C:

Cincinnaticrinus, D: Columnal Diameter, L: Lumen Diameter, H: Columnal Height)

Stratographic position E D E L E H C D C L C H 0.75 1.372381 0.237222 0.5131 1.214286 0.235714 0.522222 2.1 1.482 0.249 0.564 1.442222 0.242857 0.518889 4 1.721905 0.303 0.512 1.5072 0.260889 0.591282 4.92 1.042308 0.182308 0.401923 1.459091 0.243182 0.56625 5.13 1.163571 0.205714 0.507826 1.4725 0.264211 0.545833 6.12 1.346154 0.261304 0.4925 1.551429 0.285833 0.423333 7.04 1.937692 0.293333 0.762727 2.012 0.337143 0.72 8.8 1.630417 0.2935 0.530833 1.535652 0.271905 0.613571 9.5 1.835714 0.27 0.588929 1.822692 0.249231 0.629091 10.325 1.453333 0.283 0.611429 1.45 0.2383 0.595 11.9 1.941071 0.337857 0.66963 1.902308 0.2925 0.66375 13.5 1.682857 0.254286 0.652 1.902353 0.290667 0.4875 15.2 1.389231 0.243077 0.512727 1.359667 0.2525 0.51 16 1.78 0.321538 0.731667 1.767667 0.31 0.55 17.35 1.346923 0.2708 0.518421 1.616667 0.26069 0.610833 19.75 1.441923 0.266538 0.4875 1.39 0.222222 0.44 20.33 1.465833 0.232917 0.514762 1.523846 0.234545 0.482 21.27 1.31125 0.26 0.514167 1.513636 0.314545 22.45 1.878 0.2875 0.61125 1.607647 0.245882 0.51 22.87 1.312632 0.238824 0.55875 1.635294 0.322353 0.47625 23.75 1.507083 0.2675 0.524167 1.656071 0.291071 0.641111 24.2 1.235 0.222308 0.4635 1.523 0.264667 0.584667 24.25 1.355909 0.2385 0.52 1.613333 0.278889 0.52 24.3 1.37875 0.271667 0.515 1.333182 0.255 0.53375 24.9 1.612692 0.284231 0.559474 1.615 0.248148 0.640714 31.58 1.5825 0.277083 0.497059 1.743077 0.29 0.556923 35.2 1.557308 0.298077 0.55 1.796818 0.261429 0.632778 40.88 1.51375 0.308571 0.622 2.043333 0.406 0.58 41.95 1.355455 0.275455 0.335 1.7167 0.2917 0.685 44.45 1.8475 0.2675 0.571667 2.05875 0.284688 0.606667 51.35 2.034 0.304 0.51 2.4275 0.415 0.725 53.74 2.123529 0.358125 0.757059 2.936667 0.361667 0.748 54.16 1.665789 0.308421 0.426154 2.031667 0.315 0.51 54.26 1.932 0.366 0.613333 2.68125 0.425714 0.755 58.07 2.170476 0.328 0.77875 2.47375 0.42375 0.581667 60.4 2.213902 0.32439 0.716 3.187143 0.438571 0.725

4b. The locality and unit of the individuals used in the larger scale study. Also the continous characters are shown before standardization to number of standard deviations away from the average. (C: Cincinnaticrinus, E: Ectenocrinus simplex, G: Glyptocrinus decadactylus, GF: Glyptocrinus fornshelli, I: Iocrinus subcrassus, M: Merocrinus curtus,

P: Pycnocrinus dyeri, PD: Plicodendrocrinus casei, PP: Ptychocrinus parvus, X:

Xenocrinus baeri, D: Columnal Diameter, H: Columnal Height, L: Lumen Diameter, C%:

Percentage area of the columnal covered by crenularium, LC: Lumen Circularity, AC:

Areola Circularity, CC: Crenularium Circularity)

Member Locality D H L C % LC AC CC C1 Economy K445, KY 1.66 0.3 0.4 0.3140 1.0000 6.3000 1.0000 C2 Economy K445, KY 1.03 0.2 0.12 0.3473 1.0000 4.2000 1.0000 C3 Economy K445, KY 1.7 0.33 0.3 0.3002 1.0000 3.9167 1.0000 C4 Economy K445, KY 1.82 0.4 0.32 0.1402 1.0000 3.0000 1.0000 C5 Economy K445, KY 1.6 0.3 0.25 0.3502 1.0000 9.4000 1.0000 C6 Economy K445, KY 1.81 0.37 0.29 0.2820 1.0000 9.8000 1.0000 C7 Economy K445, KY 2.3 0.59 0.26 0.1634 1.0000 5.1000 1.0000 C8 Economy K445, KY 2.43 0.43 0.3 0.1894 1.0000 4.4000 1.0000 C9 Economy K445, KY 1.04 0.2 0.19 0.4563 1.0000 4.6000 1.0000 C10 Economy K445, KY 1.27 0.37 0.18 0.2111 1.0000 11.3333 1.0000 C11 McMicken K445, KY 2.75 0.5 0.26 0.1020 1.0000 5.4000 1.0000 C12 McMicken K445, KY 2.44 0.6 0.32 0.0588 1.0000 4.4286 1.0000 C13 McMicken K445, KY 2.84 0.79 0.37 0.2569 1.0000 3.2667 1.0000 C14 McMicken K445, KY 2.56 0.44 0.37 0.1489 1.0000 2.7692 1.0000 C15 McMicken K445, KY 2.79 0.51 0.3 0.0774 1.0000 6.5714 1.0000 C16 McMicken K445, KY 2.62 0.55 0.31 0.1539 1.0000 3.6000 1.0000 C17 McMicken K445, KY 2.2 0.52 0.33 0.2594 1.0000 1.7391 1.0000 C18 McMicken K445, KY 2.04 0.65 0.27 0.1885 1.0000 6.2000 1.0000 C19 Liberty Indiana Route 1, IN 5.41 0.7 0.51 0.1450 1.0000 3.9630 1.0000 C20 Liberty Indiana Route 1, IN 5.26 1.66 0.5 0.1109 1.0000 8.4667 1.0000 C21 Liberty Indiana Route 1, IN 3.1 0.7 0.33 0.3042 1.0000 5.0588 1.0000 C22 Liberty Indiana Route 1, IN 3.15 1.15 0.5 0.2018 1.0000 3.8824 1.0000 C23 Liberty Indiana Route 1, IN 2.56 0.68 0.44 0.3088 1.0000 3.7000 1.0000 C24 Liberty Indiana Route 1, IN 5.43 1.2 0.5 0.1544 1.0000 3.0000 1.0000 C25 Liberty Indiana Route 1, IN 4.71 1.09 0.42 0.1622 1.0000 2.8286 1.0000 C26 Liberty Indiana Route 1, IN 2.6 0.89 0.15 0.2413 1.0000 4.0000 1.0000 C27 Liberty Indiana Route 1, IN 4.85 0.8 0.39 0.0987 1.0000 1.5417 1.0000 C28 Liberty Indiana Route 1, IN 2.29 0.65 0.4 0.2196 1.0000 3.2667 1.0000 C29 Corryville Stonelick Creek, OH 2.11 0.5 0.36 0.0970 1.0000 5.0000 1.0000 C30 Corryville Stonelick Creek, OH 2.73 0.5 0.29 0.0871 1.0000 3.0000 1.0000

C31 Corryville Stonelick Creek, OH 3.57 0.5 0.47 0.1855 1.0000 6.0667 1.0000 C32 Corryville Stonelick Creek, OH 2.1 0.3 0.2 0.1003 1.0000 6.4000 1.0000 C33 Corryville Stonelick Creek, OH 4.02 0.52 0.58 0.1701 1.0000 7.0000 1.0000 C34 Arnheim Westwood 2.53 0.77 0.5 0.2320 1.0000 4.1111 1.0000 C35 Waynesville Caesar Creek, OH 3.1 0.81 0.59 0.2341 1.0000 3.6667 1.0000 C36 Waynesville Caesar Creek, OH 2.8 0.47 0.35 0.3322 1.0000 6.6667 1.0000 C37 FULTON K445, KY 2.27 0.5 0.21 0.1391 1.0000 3.7000 1.0000 C38 FULTON K445, KY 1.53 0.35 0.15 0.2173 1.0000 2.8667 1.0000 C39 FULTON K445, KY 1.72 0.25 0.26 0.2007 1.0000 4.4000 1.0000 C40 FULTON K445, KY 1.88 0.3 0.21 0.1393 1.0000 4.6000 1.0000 C41 FULTON K445, KY 1.54 0.3 0.15 0.2224 1.0000 5.4286 1.0000 C42 FULTON K445, KY 2.14 0.33 0.33 0.1611 1.0000 6.0000 1.0000 C43 FULTON K445, KY 1.85 0.47 0.3 0.3005 1.0000 3.0000 1.0000 C44 McMicken K445, KY 3.62 0.9 0.44 0.1190 1.0000 5.6667 1.0000 C45 McMicken K445, KY 3.01 0.6 0.49 0.2020 1.0000 4.3500 1.0000 C46 McMicken K445, KY 2.24 0.5 0.3 0.0978 1.0000 4.9000 1.0000 C47 McMicken K445, KY 3.22 0.6 0.6 0.1541 1.0000 9.0000 1.0000 C48 McMicken K445, KY 3.1 0.64 0.54 0.1606 1.0000 6.4000 1.0000 C49 McMicken K445, KY 2.92 0.82 0.43 0.1145 1.0000 6.8000 1.0000 C50 McMicken K445, KY 3.48 0.55 0.53 0.1692 1.0000 8.1000 1.0000 C51 Southgate K445, KY 2.36 0.49 0.51 0.1708 1.0000 6.2000 1.0000 C52 Southgate K445, KY 2.27 0.6 0.31 0.1297 1.0000 4.7000 1.0000 C53 Southgate K445, KY 2.19 0.6 0.34 0.1382 1.0000 5.1000 1.0000 C54 Southgate K445, KY 2.39 0.5 0.33 0.1708 1.0000 3.9000 1.0000 C55 Southgate K445, KY 1.64 0.4 0.2 0.1653 1.0000 3.3000 1.0000 C56 Southgate K445, KY 1.71 0.6 0.29 0.1453 1.0000 3.9000 1.0000 C57 Southgate K445, KY 1.93 0.57 0.23 0.1818 1.0000 4.3000 1.0000 E1 Fulton K445, KY 2.26 0.85 0.31 0.2347 1.6667 1.0000 1.0000 E2 Fulton K445, KY 2.2 0.56 0.64 0.6250 1.6667 1.0000 1.0000 E3 Fulton K445, KY 2.38 0.72 0.35 0.2930 1.6667 1.0000 1.0000 E4 Fulton K445, KY 2.41 0.8 0.31 0.5238 1.6667 1.0000 1.0000 E5 Fulton K445, KY 1.92 0.43 0.37 0.4165 1.6667 1.0000 1.0000 E6 Fulton K445, KY 1.74 0.5 0.34 0.3875 1.6667 1.0000 1.0000 E7 Fulton K445, KY 1.68 0.3 0.42 0.3525 1.6667 1.0000 1.0000 E8 Fulton K445, KY 1.59 0.3 0.31 0.4086 1.6667 1.0000 1.0000 E9 Fulton K445, KY 2.2 0.34 0.53 0.4927 1.6667 1.0000 1.0000 E10 McMicken K445, KY 2.33 0.3 0.4 0.4949 1.6667 1.0000 1.0000 E11 McMicken K445, KY 2.68 0.48 0.41 0.3834 1.6667 1.0000 1.0000 E12 McMicken K445, KY 2.91 0.62 0.53 0.6868 1.6667 1.0000 1.0000 E13 McMicken K445, KY 2.33 0.3 0.33 0.5089 1.6667 1.0000 1.0000 E14 McMicken K445, KY 1.08 0.15 0.19 0.6370 1.6667 1.0000 1.0000 E15 McMicken K445, KY 2.26 0.6 0.32 0.8522 1.6667 1.0000 1.0000 E16 McMicken K445, KY 1.66 0.35 0.31 0.5574 1.6667 1.0000 1.0000 E17 McMicken K445, KY 2.23 0.64 0.3 0.4696 1.6667 1.0000 1.0000 E18 Fulton K445, KY 2.25 0.5 0.38 0.3762 1.6667 1.0000 1.0000 E19 Fulton K445, KY 1.91 0.4 0.44 0.6921 1.6667 1.0000 1.0000 E20 Fulton K445, KY 1.8 0.25 0.22 0.3685 1.6667 1.0000 1.0000

E21 Fulton K445, KY 1.89 0.4 0.27 0.4630 1.6667 1.0000 1.0000 E22 Fulton K445, KY 1.36 0.2 0.22 0.8314 1.6667 1.0000 1.0000 E23 Fulton K445, KY 1.5 0.28 0.13 0.5320 1.6667 1.0000 1.0000 E24 Fulton K445, KY 2.02 0.35 0.33 0.5813 1.6667 1.0000 1.0000 E25 Fulton K445, KY 1.58 0.25 0.36 0.8898 1.6667 1.0000 1.0000 E26 Fulton K445, KY 1.37 0.5 0.26 0.7641 1.6667 1.0000 1.0000 E27 McMicken K445, KY 3.03 0.9 0.4 0.7678 1.6667 1.0000 1.0000 E28 McMicken K445, KY 2.74 0.5 0.49 0.8438 1.6667 1.0000 1.0000 E29 McMicken K445, KY 2.68 0.4 0.49 0.5641 1.6667 1.0000 1.0000 E30 McMicken K445, KY 3.24 0.4 0.6 0.6477 1.6667 1.0000 1.0000 E31 McMicken K445, KY 3.13 0.63 0.5 0.2467 1.6667 1.0000 1.0000 E32 Southgate K445, KY 2.08 0.3 0.46 0.5133 1.6667 1.0000 1.0000 G1 Bellevue Reidlin Road, OH 4.16 0.54 1.72 0.1379 1.0000 1.0000 1.0000 G2 Bellevue Reidlin Road, OH 3.2 0.35 1.44 0.2688 1.0000 1.0000 1.0000 G3 Bellevue Reidlin Road, OH 4.02 0.56 1.47 0.1068 1.0000 1.0000 1.0000 G4 Bellevue Reidlin Road, OH 2.6 0.4 0.85 0.1141 1.0000 1.0000 1.0000 G5 Bellevue Reidlin Road, OH 2.74 0.48 1.01 0.1887 1.0000 1.0000 1.0000 G6 Bellevue Reidlin Road, OH 2.62 0.37 0.83 0.0769 1.0000 1.0000 1.0000 G7 Bellevue Reidlin Road, OH 4.1 0.45 1.16 0.0875 1.0000 1.0000 1.0000 G8 Bellevue Reidlin Road, OH 4.02 0.47 1.49 0.1280 1.0000 1.0000 1.0000 G9 Bellevue Reidlin Road, OH 4.1 0.65 1.94 0.1605 1.0000 1.0000 1.0000 G10 Bellevue Reidlin Road, OH 1.98 0.35 0.58 0.1406 1.0000 1.0000 1.0000 GF1 Clarksville Jacksonburg, OH 2.52 0.36 0.81 0.2745 2.3597 1.0000 1.0000 GF2 Clarksville Jacksonburg, OH 2.39 0.55 1.42 0.3733 2.3597 1.0000 1.0000 GF3 Clarksville Jacksonburg, OH 3.03 0.24 1.18 0.1764 2.3597 1.0000 1.0000 GF4 Liberty Jacksonburg, OH 3.73 0.3 1.94 0.2725 2.3597 1.0000 1.0000 GF5 Liberty Jacksonburg, OH 3.72 0.33 1.38 0.0763 2.3597 1.0000 1.0000 GF6 Liberty Jacksonburg, OH 4.99 0.44 1.05 0.0716 2.3597 1.0000 1.0000 GF7 Liberty Jacksonburg, OH 5.47 0.47 0.81 0.0710 2.3597 1.0000 1.0000 GF8 Liberty Jacksonburg, OH 3.21 0.3 0.86 0.0870 2.3597 1.0000 1.0000 GF9 Liberty Jacksonburg, OH 4.4 0.4 0.93 0.0762 2.3597 1.0000 1.0000 GF10 Liberty Jacksonburg, OH 4.53 0.35 0.93 0.0558 2.3597 1.0000 1.0000 GF11 Liberty Jacksonburg, OH 4.93 0.39 2.17 0.1177 2.3597 1.0000 1.0000 GF12 Liberty Jacksonburg, OH 4.74 0.31 1.48 0.0377 2.3597 1.0000 1.0000 GF13 Liberty Jacksonburg, OH 3.63 0.36 0.92 0.0654 2.3597 1.0000 1.0000 I1 McMicken K445, KY 2.66 0.3 0.49 0.3553 1.0000 4.4667 1.4305 I2 McMicken K445, KY 3.03 0.35 0.67 0.3433 1.0000 8.0000 1.4305 I3 McMicken K445, KY 2.94 0.25 0.6 3.2712 1.0000 3.5217 1.4305 I4 McMicken K445, KY 3.45 0.35 0.64 0.7373 1.0000 3.4286 1.4305 I5 McMicken K445, KY 3.07 0.71 0.46 1.5677 1.0000 3.1000 1.4305 I6 Liberty Indiana Route 1, IN 4.44 0.6 0.88 0.2271 1.0000 5.5172 1.4305 I7 Liberty Indiana Route 1, IN 3.34 0.61 0.57 0.6000 1.0000 2.4412 1.4305 I8 Corryville Stonelick Creek, OH 3.16 0.54 0.43 0.4579 1.0000 4.7778 1.4305 I9 Corryville Stonelick Creek, OH 2.24 0.53 0.43 8.0753 1.0000 4.1429 1.4305 I10 Corryville Stonelick Creek, OH 3.15 0.76 0.55 0.3618 1.0000 5.1500 1.4305 I11 Corryville Stonelick Creek, OH 4.08 0.67 0.94 -0.2708 1.0000 5.9048 1.4305 I12 Corryville Stonelick Creek, OH 3.91 0.8 0.65 0.2041 1.0000 6.7500 1.4305

I13 Corryville Stonelick Creek, OH 3.06 0.54 0.59 -0.1395 1.0000 4.3333 1.4305 I14 Corryville Stonelick Creek, OH 4 0.62 0.64 0.3304 1.0000 9.5833 1.4305 I15 Corryville Stonelick Creek, OH 4.27 0.64 0.66 0.2082 1.0000 8.4000 1.4305 M1 Fulton Five Mile Creek, OH 3.67 0.36 1.13 0.4766 1.0000 1.0000 1.0000 M2 Fulton Five Mile Creek, OH 4.38 0.43 1.26 0.6321 1.0000 1.0000 1.0000 M3 Fulton Five Mile Creek, OH 3.83 0.6 0.93 0.5772 1.0000 1.0000 1.0000 M4 Fulton Five Mile Creek, OH 4.46 0.57 1.09 0.8165 1.0000 1.0000 1.0000 M5 Fulton Five Mile Creek, OH 3.13 0.57 0.93 0.6228 1.0000 1.0000 1.0000 M6 Fulton Five Mile Creek, OH 2.49 0.4 0.75 0.7846 1.0000 1.0000 1.0000 M7 Fulton Five Mile Creek, OH 3.18 0.54 0.8 0.8729 1.0000 1.0000 1.0000 M8 Fulton Five Mile Creek, OH 4.51 0.5 1.26 0.8268 1.0000 1.0000 1.0000 M9 Fulton Five Mile Creek, OH 2.3 0.34 0.79 0.5451 1.0000 1.0000 1.0000 P1 Bellevue Reidlin Road, OH 3.44 0.8 0.96 0.1095 1.5569 1.0000 1.0000 P2 Bellevue Reidlin Road, OH 2.77 0.51 1.15 0.3423 1.5569 1.0000 1.0000 P3 Bellevue Reidlin Road, OH 2.53 0.45 1.54 0.5320 1.5569 1.0000 1.0000 P4 Bellevue Reidlin Road, OH 3.68 1.06 1.12 0.1679 1.5569 1.0000 1.0000 P5 Bellevue Reidlin Road, OH 2.92 0.38 1.26 0.2983 1.5569 1.0000 1.0000 P6 Bellevue Reidlin Road, OH 2.66 0.86 0.99 0.1740 1.5569 1.0000 1.0000 P7 Bellevue Reidlin Road, OH 3.88 0.66 1.46 0.1217 1.5569 1.0000 1.0000 P8 Bellevue Reidlin Road, OH 2.82 0.66 1.01 0.1696 1.5569 1.0000 1.0000 P9 Bellevue Reidlin Road, OH 3.19 0.82 1.32 0.2895 1.5569 1.0000 1.0000 P10 Bellevue Reidlin Road, OH 2.02 0.25 1.09 0.5402 1.5569 1.0000 1.0000 P11 Corryville Stonelick Creek, OH 3.59 0.43 1.73 0.3708 1.5569 1.0000 1.0000 P12 Corryville Stonelick Creek, OH 2.38 0.51 0.8 0.2556 1.5569 1.0000 1.0000 P13 Corryville Stonelick Creek, OH 2.42 0.62 0.82 0.4714 1.5569 1.0000 1.0000 P14 Corryville Stonelick Creek, OH 3.33 0.47 1.28 0.1959 1.5569 1.0000 1.0000 P15 Corryville Stonelick Creek, OH 2.57 0.64 1.13 0.4050 1.5569 1.0000 1.0000 P16 Corryville Stonelick Creek, OH 2.67 0.42 0.74 0.1808 1.5569 1.0000 1.0000 PD1 Whitewater Clarksville, OH 2.49 0.28 0.39 0.3307 1.0000 3.6800 1.9974 PD2 Whitewater Clarksville, OH 3.56 0.57 0.55 0.2060 1.0000 4.4242 1.9974 PD3 Whitewater Clarksville, OH 2.7 0.22 0.53 0.1266 1.0000 6.2000 1.9974 PD4 Whitewater Clarksville, OH 2.97 0.31 0.59 0.2648 1.0000 6.8667 1.9974 PD5 Liberty unknown 4.1 0.55 0.77 0.2131 1.0000 4.1818 1.9974 PD6 Liberty unknown 2.6 0.36 0.57 0.2240 1.0000 3.3438 1.9974 PD7 Liberty unknown 4.16 0.61 0.8 0.2923 1.0000 4.5161 1.9974 PP1 Mount Hope Fairview Heights, OH 3.49 0.31 1.13 0.3038 1.4083 1.0000 1.0000 PP2 Mount Hope Fairview Heights, OH 3.99 0.52 1.53 0.1889 1.4083 1.0000 1.0000 PP3 Mount Hope Fairview Heights, OH 3.84 0.5 1.47 0.3081 1.4083 1.0000 1.0000 PP4 Mount Hope Fairview Heights, OH 3.13 0.5 1.18 0.3648 1.4083 1.0000 1.0000 PP5 Mount Hope Fairview Heights, OH 3.61 0.49 0.88 0.3052 1.4083 1.0000 1.0000 PP6 Mount Hope Fairview Heights, OH 3.1 0.53 1.26 0.3768 1.4083 1.0000 1.0000 PP7 Mount Hope Fairview Heights, OH 2.9 0.48 1.1 0.4856 1.4083 1.0000 1.0000 PP8 Mount Hope Fairview Heights, OH 3.2 0.42 0.99 0.1879 1.4083 1.0000 1.0000 PP9 Mount Hope Fairview Heights, OH 3.61 0.36 1.41 0.1554 1.4083 1.0000 1.0000 PP10 Mount Hope Fairview Heights, OH 3.52 0.68 1.51 0.3027 1.4083 1.0000 1.0000 X1 Liberty Indiana Route 1, IN 1.3 0.1 0.33 0.3241 1.0000 1.4211 1.3676 X2 Liberty Indiana Route 1, IN 2.23 0.37 0.38 0.7343 1.0000 2.8000 1.3676

X3 Liberty Indiana Route 1, IN 1.34 0.15 0.3 0.8390 1.0000 1.0435 1.3676 X4 Liberty Indiana Route 1, IN 1.56 0.36 0.28 0.6577 1.0000 1.1538 1.3676 X5 Liberty Indiana Route 1, IN 2.36 0.2 0.56 0.5498 1.0000 1.6296 1.3676 X6 Liberty Indiana Route 1, IN 2.02 0.33 0.38 0.7333 1.0000 1.6429 1.3676 X7 Liberty Indiana Route 1, IN 3.31 0.64 0.42 0.4161 1.0000 1.5581 1.3676 X8 Liberty Jacksonburg, OH 2.23 0.42 0.36 0.4761 1.0000 1.0000 1.3676 X9 Liberty Jacksonburg, OH 1.56 0.25 0.4 0.6489 1.0000 1.5667 1.3676

4c. Dataset used in the PCA analysis after standardization of the continuous variables.

(Sym: Column Symmetry, Art: Type of Articulation, Homeo: Presence of

Homeomorphic columns, LS: Lateral Shape)

Sym Art Homeo LS D H L C LC AC C1 1 2 1 2 -1.2242 -0.9528 -0.5961 -0.1072 -0.7133 1.4700 C2 1 2 1 2 -1.8763 -1.4245 -1.2496 -0.0571 -0.7133 0.5846 C3 1 2 1 2 -1.1828 -0.8113 -0.8295 -0.1280 -0.7133 0.4651 C4 1 2 1 2 -1.0586 -0.4812 -0.7828 -0.3688 -0.7133 0.0786 C5 1 2 1 2 -1.2863 -0.9528 -0.9462 -0.0528 -0.7133 2.7770 C6 1 2 1 2 -1.0689 -0.6227 -0.8528 -0.1555 -0.7133 2.9456 C7 1 2 1 2 -0.5617 0.4149 -0.9229 -0.3340 -0.7133 0.9640 C8 1 2 1 2 -0.4272 -0.3397 -0.8295 -0.2948 -0.7133 0.6689 C9 1 2 1 2 -1.8659 -1.4245 -1.0862 0.1068 -0.7133 0.7532 C10 1 2 1 2 -1.6279 -0.6227 -1.1096 -0.2621 -0.7133 3.5921 C11 1 2 1 2 -0.0959 -0.0095 -0.9229 -0.4263 -0.7133 1.0905 C12 1 2 1 2 -0.4168 0.4621 -0.7828 -0.4913 -0.7133 0.6809 C13 1 2 1 2 -0.0028 1.3582 -0.6662 -0.1932 -0.7133 0.1911 C14 1 2 1 2 -0.2926 -0.2925 -0.6662 -0.3558 -0.7133 -0.0187 C15 1 2 1 2 -0.0545 0.0376 -0.8295 -0.4634 -0.7133 1.5844 C16 1 2 1 2 -0.2305 0.2263 -0.8062 -0.3483 -0.7133 0.3316 C17 1 2 1 2 -0.6652 0.0848 -0.7595 -0.1895 -0.7133 -0.4530 C18 1 2 1 2 -0.8308 0.6979 -0.8995 -0.2962 -0.7133 1.4278 C19 1 2 1 2 2.6573 0.9337 -0.3394 -0.3617 -0.7133 0.4846 C20 1 2 1 2 2.5021 5.4615 -0.3628 -0.4130 -0.7133 2.3835 C21 1 2 1 2 0.2663 0.9337 -0.7595 -0.1220 -0.7133 0.9467 C22 1 2 1 2 0.3181 3.0561 -0.3628 -0.2762 -0.7133 0.4506 C23 1 2 1 2 -0.2926 0.8394 -0.5028 -0.1152 -0.7133 0.3738 C24 1 2 1 2 2.6780 3.2919 -0.3628 -0.3474 -0.7133 0.0786 C25 1 2 1 2 1.9328 2.7731 -0.5495 -0.3358 -0.7133 0.0064 C26 1 2 1 2 -0.2512 1.8299 -1.1796 -0.2167 -0.7133 0.5003 C27 1 2 1 2 2.0777 1.4054 -0.6195 -0.4314 -0.7133 -0.5362 C28 1 2 1 2 -0.5721 0.6979 -0.5961 -0.2494 -0.7133 0.1911 C29 1 2 1 2 -0.7584 -0.0095 -0.6895 -0.4340 -0.7133 0.9219 C30 1 2 1 2 -0.1166 -0.0095 -0.8528 -0.4487 -0.7133 0.0786 C31 1 2 1 2 0.7528 -0.0095 -0.4328 -0.3006 -0.7133 1.3716 C32 1 2 1 2 -0.7687 -0.9528 -1.0629 -0.4290 -0.7133 1.5121 C33 1 2 1 2 1.2186 0.0848 -0.1761 -0.3239 -0.7133 1.7651 C34 1 2 1 2 -0.3237 1.2639 -0.3628 -0.2307 -0.7133 0.5471 C35 1 2 1 2 0.2663 1.4525 -0.1527 -0.2276 -0.7133 0.3597 C36 1 2 1 2 -0.0442 -0.1510 -0.7128 -0.0798 -0.7133 1.6246 C37 1 2 1 2 -0.5928 -0.0095 -1.0395 -0.3706 -0.7133 0.3738 C38 1 2 1 2 -1.3587 -0.7170 -1.1796 -0.2528 -0.7133 0.0224 C39 1 2 1 2 -1.1621 -1.1886 -0.9229 -0.2778 -0.7133 0.6689 C40 1 2 1 2 -0.9965 -0.9528 -1.0395 -0.3702 -0.7133 0.7532

C41 1 2 1 2 -1.3484 -0.9528 -1.1796 -0.2452 -0.7133 1.1026 C42 1 2 1 2 -0.7273 -0.8113 -0.7595 -0.3374 -0.7133 1.3435 C43 1 2 1 2 -1.0275 -0.1510 -0.8295 -0.1276 -0.7133 0.0786 C44 1 2 1 2 0.8046 1.8770 -0.5028 -0.4008 -0.7133 1.2029 C45 1 2 1 2 0.1732 0.4621 -0.3861 -0.2758 -0.7133 0.6478 C46 1 2 1 2 -0.6238 -0.0095 -0.8295 -0.4327 -0.7133 0.8797 C47 1 2 1 2 0.3905 0.4621 -0.1294 -0.3479 -0.7133 2.6083 C48 1 2 1 2 0.2663 0.6508 -0.2694 -0.3382 -0.7133 1.5121 C49 1 2 1 2 0.0800 1.4997 -0.5261 -0.4075 -0.7133 1.6808 C50 1 2 1 2 0.6597 0.2263 -0.2928 -0.3253 -0.7133 2.2289 C51 1 2 1 2 -0.4996 -0.0567 -0.3394 -0.3228 -0.7133 1.4278 C52 1 2 1 2 -0.5928 0.4621 -0.8062 -0.3847 -0.7133 0.7954 C53 1 2 1 2 -0.6756 0.4621 -0.7362 -0.3720 -0.7133 0.9640 C54 1 2 1 2 -0.4686 -0.0095 -0.7595 -0.3229 -0.7133 0.4581 C55 1 2 1 2 -1.2449 -0.4812 -1.0629 -0.3310 -0.7133 0.2051 C56 1 2 1 2 -1.1724 0.4621 -0.8528 -0.3612 -0.7133 0.4581 C57 1 2 1 2 -0.9447 0.3206 -0.9929 -0.3062 -0.7133 0.6267 E1 1 0 1 3 -0.6031 1.6412 -0.8062 -0.2267 0.9144 -0.7646 E2 1 0 1 3 -0.6652 0.2734 -0.0361 0.3609 0.9144 -0.7646 E3 1 0 1 3 -0.4789 1.0281 -0.7128 -0.1388 0.9144 -0.7646 E4 1 0 1 3 -0.4479 1.4054 -0.8062 0.2085 0.9144 -0.7646 E5 1 0 1 3 -0.9551 -0.3397 -0.6662 0.0470 0.9144 -0.7646 E6 1 0 1 3 -1.1414 -0.0095 -0.7362 0.0033 0.9144 -0.7646 E7 1 0 1 3 -1.2035 -0.9528 -0.5495 -0.0493 0.9144 -0.7646 E8 1 0 1 3 -1.2966 -0.9528 -0.8062 0.0350 0.9144 -0.7646 E9 1 0 1 3 -0.6652 -0.7642 -0.2928 0.1617 0.9144 -0.7646 E10 1 0 1 3 -0.5307 -0.9528 -0.5961 0.1650 0.9144 -0.7646 E11 1 0 1 3 -0.1684 -0.1039 -0.5728 -0.0028 0.9144 -0.7646 E12 1 0 1 3 0.0697 0.5564 -0.2928 0.4539 0.9144 -0.7646 E13 1 0 1 3 -0.5307 -0.9528 -0.7595 0.1860 0.9144 -0.7646 E14 1 0 1 3 -1.8245 -1.6603 -1.0862 0.3789 0.9144 -0.7646 E15 1 0 1 3 -0.6031 0.4621 -0.7828 0.7028 0.9144 -0.7646 E16 1 0 1 3 -1.2242 -0.7170 -0.8062 0.2590 0.9144 -0.7646 E17 1 0 1 3 -0.6342 0.6508 -0.8295 0.1270 0.9144 -0.7646 E18 1 0 1 3 -0.6135 -0.0095 -0.6428 -0.0137 0.9144 -0.7646 E19 1 0 1 3 -0.9654 -0.4812 -0.5028 0.4618 0.9144 -0.7646 E20 1 0 1 3 -1.0793 -1.1886 -1.0162 -0.0253 0.9144 -0.7646 E21 1 0 1 3 -0.9861 -0.4812 -0.8995 0.1169 0.9144 -0.7646 E22 1 0 1 3 -1.5347 -1.4245 -1.0162 0.6715 0.9144 -0.7646 E23 1 0 1 3 -1.3898 -1.0471 -1.2262 0.2208 0.9144 -0.7646 E24 1 0 1 3 -0.8515 -0.7170 -0.7595 0.2951 0.9144 -0.7646 E25 1 0 1 3 -1.3070 -1.1886 -0.6895 0.7594 0.9144 -0.7646 E26 1 0 1 3 -1.5243 -0.0095 -0.9229 0.5701 0.9144 -0.7646 E27 1 0 1 3 0.1939 1.8770 -0.5961 0.5757 0.9144 -0.7646 E28 1 0 1 3 -0.1063 -0.0095 -0.3861 0.6900 0.9144 -0.7646 E29 1 0 1 3 -0.1684 -0.4812 -0.3861 0.2691 0.9144 -0.7646 E30 1 0 1 3 0.4112 -0.4812 -0.1294 0.3949 0.9144 -0.7646

E31 1 0 1 3 0.2974 0.6036 -0.3628 -0.2085 0.9144 -0.7646 E32 1 0 1 3 -0.7894 -0.9528 -0.4561 0.1926 0.9144 -0.7646 G1 1 0 2 2 1.3635 0.1791 2.4843 -0.3724 -0.7133 -0.7646 G2 1 0 2 2 0.3698 -0.7170 1.8309 -0.1753 -0.7133 -0.7646 G3 1 0 2 2 1.2186 0.2734 1.9009 -0.4191 -0.7133 -0.7646 G4 1 0 2 2 -0.2512 -0.4812 0.4540 -0.4082 -0.7133 -0.7646 G5 1 0 2 2 -0.1063 -0.1039 0.8274 -0.2959 -0.7133 -0.7646 G6 1 0 2 2 -0.2305 -0.6227 0.4073 -0.4641 -0.7133 -0.7646 G7 1 0 2 2 1.3014 -0.2454 1.1775 -0.4482 -0.7133 -0.7646 G8 1 0 2 2 1.2186 -0.1510 1.9476 -0.3872 -0.7133 -0.7646 G9 1 0 2 2 1.3014 0.6979 2.9978 -0.3383 -0.7133 -0.7646 G10 1 0 2 2 -0.8930 -0.7170 -0.1761 -0.3682 -0.7133 -0.7646 GF1 3 0 1 3 -0.3340 -0.6698 0.3607 -0.1668 2.6064 -0.7646 GF2 3 0 1 3 -0.4686 0.2263 1.7842 -0.0180 2.6064 -0.7646 GF3 3 0 1 3 0.1939 -1.2358 1.2241 -0.3144 2.6064 -0.7646 GF4 3 0 1 3 0.9184 -0.9528 2.9978 -0.1698 2.6064 -0.7646 GF5 3 0 1 3 0.9081 -0.8113 1.6909 -0.4651 2.6064 -0.7646 GF6 3 0 1 3 2.2226 -0.2925 0.9208 -0.4722 2.6064 -0.7646 GF7 3 0 1 3 2.7194 -0.1510 0.3607 -0.4730 2.6064 -0.7646 GF8 3 0 1 3 0.3802 -0.9528 0.4774 -0.4490 2.6064 -0.7646 GF9 3 0 1 3 1.6119 -0.4812 0.6407 -0.4652 2.6064 -0.7646 GF10 3 0 1 3 1.7465 -0.7170 0.6407 -0.4959 2.6064 -0.7646 GF11 3 0 1 3 2.1605 -0.5283 3.5345 -0.4027 2.6064 -0.7646 GF12 3 0 1 3 1.9638 -0.9057 1.9243 -0.5232 2.6064 -0.7646 GF13 3 0 1 3 0.8149 -0.6698 0.6174 -0.4814 2.6064 -0.7646 I1 3 0 1 2 -0.1891 -0.9528 -0.3861 -0.0451 -0.7133 0.6970 I2 3 0 1 2 0.1939 -0.7170 0.0340 -0.0632 -0.7133 2.1867 I3 3 0 1 2 0.1007 -1.1886 -0.1294 4.3435 -0.7133 0.2986 I4 3 0 1 2 0.6286 -0.7170 -0.0361 0.5298 -0.7133 0.2593 I5 3 0 1 2 0.2353 0.9809 -0.4561 1.7796 -0.7133 0.1208 I6 3 0 1 2 1.6533 0.4621 0.5240 -0.2381 -0.7133 1.1399 I7 3 0 1 2 0.5147 0.5093 -0.1994 0.3232 -0.7133 -0.1570 I8 3 0 1 2 0.3284 0.1791 -0.5261 0.1092 -0.7133 0.8282 I9 3 0 1 2 -0.6238 0.1320 -0.5261 11.5742 -0.7133 0.5605 I10 3 0 1 2 0.3181 1.2167 -0.2461 -0.0353 -0.7133 0.9851 I11 3 0 1 2 1.2807 0.7922 0.6641 -0.9875 -0.7133 1.3033 I12 3 0 1 2 1.1047 1.4054 -0.0127 -0.2726 -0.7133 1.6597 I13 3 0 1 2 0.2249 0.1791 -0.1527 -0.7898 -0.7133 0.6408 I14 3 0 1 2 1.1979 0.5564 -0.0361 -0.0826 -0.7133 2.8543 I15 3 0 1 2 1.4774 0.6508 0.0106 -0.2665 -0.7133 2.3554 M1 1 0 0 1 0.8563 -0.6698 1.1075 0.1374 -0.7133 -0.7646 M2 1 0 0 1 1.5912 -0.3397 1.4108 0.3715 -0.7133 -0.7646 M3 1 0 0 1 1.0219 0.4621 0.6407 0.2889 -0.7133 -0.7646 M4 1 0 0 1 1.6740 0.3206 1.0141 0.6491 -0.7133 -0.7646 M5 1 0 0 1 0.2974 0.3206 0.6407 0.3575 -0.7133 -0.7646 M6 1 0 0 1 -0.3651 -0.4812 0.2207 0.6010 -0.7133 -0.7646 M7 1 0 0 1 0.3491 0.1791 0.3373 0.7340 -0.7133 -0.7646

M8 1 0 0 1 1.7258 -0.0095 1.4108 0.6645 -0.7133 -0.7646 M9 1 0 0 1 -0.5617 -0.7642 0.3140 0.2405 -0.7133 -0.7646 P1 1 0 2 2 0.6183 1.4054 0.7107 -0.4150 0.6463 -0.7646 P2 1 0 2 2 -0.0752 0.0376 1.1541 -0.0647 0.6463 -0.7646 P3 1 0 2 2 -0.3237 -0.2454 2.0643 0.2208 0.6463 -0.7646 P4 1 0 2 2 0.8667 2.6316 1.0841 -0.3272 0.6463 -0.7646 P5 1 0 2 2 0.0800 -0.5755 1.4108 -0.1309 0.6463 -0.7646 P6 1 0 2 2 -0.1891 1.6884 0.7807 -0.3180 0.6463 -0.7646 P7 1 0 2 2 1.0737 0.7451 1.8776 -0.3967 0.6463 -0.7646 P8 1 0 2 2 -0.0235 0.7451 0.8274 -0.3246 0.6463 -0.7646 P9 1 0 2 2 0.3595 1.4997 1.5509 -0.1442 0.6463 -0.7646 P10 1 0 2 2 -0.8515 -1.1886 1.0141 0.2332 0.6463 -0.7646 P11 1 0 2 2 0.7735 -0.3397 2.5077 -0.0217 0.6463 -0.7646 P12 1 0 2 2 -0.4789 0.0376 0.3373 -0.1951 0.6463 -0.7646 P13 1 0 2 2 -0.4375 0.5564 0.3840 0.1296 0.6463 -0.7646 P14 1 0 2 2 0.5044 -0.1510 1.4575 -0.2851 0.6463 -0.7646 P15 1 0 2 2 -0.2823 0.6508 1.1075 0.0296 0.6463 -0.7646 P16 1 0 2 2 -0.1788 -0.3869 0.1973 -0.3077 0.6463 -0.7646 PD1 3 0 0 2 -0.3651 -1.0471 -0.6195 -0.0821 -0.7133 0.3653 PD2 3 0 0 2 0.7425 0.3206 -0.2461 -0.2699 -0.7133 0.6791 PD3 3 0 0 2 -0.1477 -1.3301 -0.2928 -0.3894 -0.7133 1.4278 PD4 3 0 0 2 0.1318 -0.9057 -0.1527 -0.1814 -0.7133 1.7089 PD5 3 0 0 2 1.3014 0.2263 0.2673 -0.2592 -0.7133 0.5769 PD6 3 0 0 2 -0.2512 -0.6698 -0.1994 -0.2428 -0.7133 0.2236 PD7 3 0 0 2 1.3635 0.5093 0.3373 -0.1399 -0.7133 0.7179 PP1 1 0 2 2 0.6700 -0.9057 1.1075 -0.1226 0.2837 -0.7646 PP2 1 0 2 2 1.1875 0.0848 2.0409 -0.2956 0.2837 -0.7646 PP3 1 0 2 2 1.0323 -0.0095 1.9009 -0.1161 0.2837 -0.7646 PP4 1 0 2 2 0.2974 -0.0095 1.2241 -0.0308 0.2837 -0.7646 PP5 1 0 2 2 0.7942 -0.0567 0.5240 -0.1205 0.2837 -0.7646 PP6 1 0 2 2 0.2663 0.1320 1.4108 -0.0128 0.2837 -0.7646 PP7 1 0 2 2 0.0593 -0.1039 1.0374 0.1509 0.2837 -0.7646 PP8 1 0 2 2 0.3698 -0.3869 0.7807 -0.2971 0.2837 -0.7646 PP9 1 0 2 2 0.7942 -0.6698 1.7609 -0.3460 0.2837 -0.7646 PP10 1 0 2 2 0.7011 0.8394 1.9943 -0.1244 0.2837 -0.7646 X1 2 0 1 3 -1.5968 -1.8961 -0.7595 -0.0921 -0.7133 -0.5871 X2 2 0 1 3 -0.6342 -0.6227 -0.6428 0.5252 -0.7133 -0.0057 X3 2 0 1 3 -1.5554 -1.6603 -0.8295 0.6830 -0.7133 -0.7463 X4 2 0 1 3 -1.3277 -0.6698 -0.8762 0.4100 -0.7133 -0.6997 X5 2 0 1 3 -0.4996 -1.4245 -0.2228 0.2476 -0.7133 -0.4991 X6 2 0 1 3 -0.8515 -0.8113 -0.6428 0.5238 -0.7133 -0.4936 X7 2 0 1 3 0.4837 0.6508 -0.5495 0.0464 -0.7133 -0.5293 X8 2 0 1 3 -0.6342 -0.3869 -0.6895 0.1367 -0.7133 -0.7646 X9 2 0 1 3 -1.3277 -1.1886 -0.5961 0.3968 -0.7133 -0.5257

FIGURE CAPTIONS

Figure 1. Disparid crinoids from the type Cincinnatian. A, B Views of the lateral profile

and articular surface of Cincinnaticrinus varibrachialus. Kope Formation, K445. C, D

Views of the lateral profile and articular surface of Cincinnaticrinus pentagonalus.

Liberty Formation, Indiana route 1. E,F Views of the lateral profile and articular surface of Ectenocrinus simplex. Kope formation, K445. G, H Views of the lateral profile and

articular surface of Iocrinus subcrassus. Kope Formation, K445. Scale bars are equal to 1

mm. A,B,E,F,G,H are modified from Meyer et al. (2002).

Figure 2. Variations in the lithology (Meyer et al. 2002) of the Kope and Lower Fairview

Formations as well as the faunal gradient DCA axis 1 scores (Miller et al. 2001). In the

lithology columns gray is percent shale, black is percent siltstone, and white is percent

limestone. The first column is the raw data collected on a centimeter scale. The second

column has a 21-point moving average applied to it to reduce small-scale variation and

emphasize the overall trend. The third column values were computed using only the

horizons that were sampled in this study.

Figure 3. Monobathrid crinoids from the type Cincinnatian. A, B. Views of the lateral

profile and articular surface of Glyptocrinus fornshelli, Liberty Formation. Jacksonburg,

Ohio, Miami University Museum of Paleontology 28121. B,C Views of the lateral profile

and articular surface of Glyptocrinus decadactylus, Cincinnati Museum of Natural

History 11101. Columnals of Pycnocrinus dyeri show no visible difference from those of

G. decadactylus. D, E Views of the lateral profile and articular surface of Xenocrinus baeri, Liberty formation, Indiana route 1. All scale bars are 1 mm.

Figure 4. Diplobathrid and Cladid crinoids from the type Cincinnatian. A, B Views of the lateral profile and articular surface of Merocrinus curtus, Fulton member of the Kope

Formation, K445. C, D Views of the lateral profile and articular surface of

Plicodendrocrinus casei. E,F Views of the lateral profile and articular surface of

Ptychocrinus parvus. Mount Auburn member, Corryville formation, Fairview Heights,

Cincinnati, Ohio, Cincinnati Museum of Natural History 11090. All scale bars are 1 mm.

A is modified from Meyer et al. (2002).

Figure 5. A. A generalized columnal with a round lumen, crenularium, and areola, with morphology labeled. B. A columnal with a round crenularium and areola and a pentagonal lumen. C. Measurements used to calculate lumen circularity for the columnal in B. Lumen circularity is the ratio of the maximum to minimum lumen radii. D. A generalized columnal with a round lumen and areola and a pentagonal crenularium. E.

Measurements used to calculate crenularium circulatity for the columnal in D.

Crenularium circularity is the ratio of the maximum and minimum distance between the centroid of the columnal and the outer edge of the crenularium.

Figure 6. Differences in lateral shape of the columns examined in this study. A. Planar, all of the columnals are the same size and the side of the column forms a straight line. B.

Round convex, the side of the nodal is rounded so that the middle of the columnal

extends farther than the intersection with the top or bottom articulation surfaces. C.

Angular convex, the side of the columnal is planar, but the angles between the sides of the columnal and the articulation surfaces are different for the top and the bottom of the columnal.

Figure 7. Differences in column form of the columns examined in this study. A.

Homeomorphic, all columnals are the same size. B. Heteromorphic, the column is formed by columnals of two different sizes, called nodals and internodals. C. Heteromorphic, the column is formed by the nodal and two or more internodals.

Figure 8. A depth curve (Cuffey, 1998) was used to graphically create a relative depth scale. A straight line was drawn downward for the average depth of each member. An arbitrary scale was placed at the bottom to create a relative depth for each member.

Figure 9. The lithology (Meyer et al. 2002), faunal gradient DCA axis 1 scores (Miller et al. 2001), and crinoid morphology for the K445 section of the Kope and Lower

Fairview Formations. For the lithology column the pattern is as in Figure 2. The columnal and lumen diameters are measured in mm, and the red line is for

Cincinnaticrinus while the black line is for Ectenocrinus. Error bars are standard error.

Figure 10. Results of the PCA analysis. PCA axis 1 represents 29.738 percent of the variation and is correlated with areola circularity (r=0.768), lumen circularity (r=-0.810), lumen diameter (r=-0.633), and the type of articulation (r=0.837). PCA axis 2 represents

20.068 percent of the variation and correlates with lumen diameter (r=0.642), columnal height (r=0.625), and columnal diameter (r=0.817).

Figure 11. PCA axis 2 compared with relative depth (figure 5) for disparids, monobathrid and diplobathrid camerates and cladid crinoids.

Figure 12. PCA axis 2 compared with relative depth for the disparids: Cincinnaticrinus varibrachialus, Cincinnaticrinus pentagonus, Ectenocrinus simplex, and Iocrinus subcrassus. The two species of Cincinnaticrinus (C. variabrachialus and C. pentagonus) have been grouped together to show the continuous change in columnal morphology between the two species.

Figure 13. Average PCA axis 2 scores for each stratigraphic sample for each crinoid compared with relative depth. Error bars are standard error.

Figure 14. Average PCA axis 2 scores for each sample graphed against stratigraphic postion in the type Cincinnatian. Lower numbers are older members and the complete list of members is given in Table 1. Error bars are standard error.

Figure 15. Average PCA axis 2 scores compared with the average relative depth for each crinoid examined. Error bars are standard error around both the average relative depth and average PCA axis 2 score.

Figure 16. The range in columnal diameter (the difference in columnal diameter between the maximum and minimum columnal diameter in a sample) for samples that had between 21 and 26 individuals of Ectenocrinus simplex in the K445 section of the Kope and Lower Fairview formations.

Figure 17. Lithology (as in Figure 2, Meyer et al. 2002) and crinoid columnal data compared with events in crinoid distributions. The asterisks represent the peak abundance in the deposition sequence of Iocrinus subcrassus. The I is the first appearance of I. subcrassus and the G represents the first appearance of Glyptocrinus decadactylus (from

Meyer et al. 2002).

Figure 18. A comparison of the relative depth and PCA axis 2 score of pinnulate and nonpinnulate crinoids. Error bars are standard error.

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Figure 6.

Figure 7.

Figure 8.

Figure 9.

3.5 3 2.5 2 1.5

1 y = 0.0173x + 1.5314 0.5 R2 = 0.3344 0 Columnal Diameter Range (mm 0 10203040506070 Height above the base of the K445 section (m)

Figure 16.

Figure 17.

TABLE CAPTIONS.

Table 1. The stratigraphic position of the crinoids used in this study. The black boxes

represent where samples have been taken for this study, while the gray boxes represent

reported though unsampled occurrences. FU: Fulton member, EC: Economy member,

SO: Southgate member, MM: McMicken member, MH: Mount Hope member, FA:

Fairmount member, BE: Bellevue member, CO: Corryville member, MA: Mount Auburn

member, SU: Sunset member, OR: Oregonia member, FA: Fort Ancient member, CL:

Clacksville member, BL: Blanchester member, LI: Liberty Formation,WH: Whitewater

Formation.

Table 2. Morphological characters of crinoids in the Cincinnatian. Explainations of the

terms can be found in the text and in Donovan (1986).

Table 3 Probability values for Spearman’s rank correlation between the crinoid data,

lithology: percent limestone, percent shale, and a 21-point moving average applied to

percent limestone, shale, and siltstone (Meyer et al., 2002) and the faunal gradiant DCA

axis one (with and without application of a 21-point moving average, Miller et al., 2001).

Significant values at the p=0.05 level are highlighted in grey.

E D E N M A Y S V I L L E R I C H M O N D K O P E FAIRVIEW CORRYVILLE ARNHEIM WAYNESVILLE FU EC SO MM MH FA BE CO MA SU OR FA CL BL LI WH 13.5 13 10 8 6 4 2 6 3 4 2 8 8 8 6 3 Relative Depth

DISPARIDS Cincinnaticrinus varibrachialus Cincinnaticrinus pentagonus Ectenocrinus simplex Iocrinus subcrassus

MONOBATHRIDS Glyptocrinus decadactylus Glyptocrinus fornshelli Pycnocrinus dyeri Xenocrinus baeri

DIPLOBATHRIDS Ptychocrinus parvus

CLADIDS Merocrinus curtus Plicodendrocrinus casei

Table 1.

Morphology Symmetry Articulation Lateral Shape Column Form Genus Circular Quadrangular Pentagonal Petaloid Symplexial Planar Round convex Angular convex Homeomorphic 1 IN 2 or more IN Cincinnaticrinus X X X X Ectinocrinus X X X X Iocrinus X X X X Merocrinus X X X X Plicodendrocrinus X X X X Ptychocrinus X X X X Glyptocrinus decadactylus X X X X Glyptocrinus fornshelli X X X X Pycnocrinus X X X X Xenocrinus X X X X

Table 2.

% ls %shale % ls 21 % shale 21 % silt 21 faunal Faunal 21

DCA 0.783 0.573 0.833 0.183 0.147 0.942 0.001 DCA no height 0.887 0.787 0.968 0.582 0.457 0.968 0.004 DCA reduced 0.742 0.834 0.836 0.554 0.463 0.975 0.004 Cin D ave 0.215 0.053 0.944 0.094 0.018 0.726 0.042 Cin D Max 0.734 0.661 0.506 0.299 0.21 0.935 0.028 Cin D Min 0.024 0.01 0.051 0.023 0.055 0.381 0.106 Cin Lumen 0.933 0.518 0.618 0.135 0.021 0.648 0.002 Ect D Ave 0.202 0.094 0.086 0.108 0.102 0.505 0.505 Ect D Max 0.162 0.311 0.116 0.241 0.791 0.235 0.729 Ect D Min 0.151 0.056 0.183 0.151 0.159 0.06 0.634 Ect Lumen 0.155 0.052 0.017 0.015 0.124 0.678 0.619

Table 3