A Crinoid Lagerstatte from Maysville, Kentucky: Paleoautecology And

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A crinoid Lagerstätte from Maysville, Kentucky: paleoautecology and taphonomy

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

McMicken College of Arts and Sciences

Mason Jane Milam

B.A. Hunter College

City University of New York

1993

Committee Chair: David L. Meyer, Ph.D.

Abstract

A Lagerstätte of Glyptocrinus decadactylus collected from the Upper Fairview Formation at

Maysville, Kentucky, USA, yields new insights into the paleoautecology of camerate crinoids of the Late Ordovician. The Lagerstätte represents an autochthonous community comprised of a single mudstone interval representing an obrution deposit and containing over 400 individual glyptocrinids, including over 250 calyces. The crinoids had colonized a rise of relatively high energy within the deep subtidal zone where they were partially sheltered by plant matter to which they were attached, as suggested by columns terminating distally in coiled holdfasts of relatively consistent diameter. Turbidity flows originating in storm events led to the suspension of fine sediments by a lofting plume that intruded into and buried the crinoids in their habitat. The crinoid and plant stand on the rise may have created additional friction that slowed the sediment plume and induced deposition, signifying burial facilitated by a feedback process. Some specimens had been dead for a period of time prior to burial, suggesting more than one earlier killing event, likely related to the storms that ultimately caused the obrution. Other specimens were apparently killed by the obrution event and subjected to little or no scavenging in the resulting anoxic conditions, leading to excellent preservation. Most specimens are in trauma position, although more than 6% were preserved in spread-fan feeding posture, aboral surface up. The latter position is indicative of very rapid burial, and is often seen in stalked crinoids capable of neither crawling nor autotomizing their stems to escape.

The high density of the crinoid community was accommodated by tiering, in which, in addition to the plant matter, the crinoids employed the columns of earlier-settled specimens as attachment sites; differing column lengths positioned the crinoids at various

i levels within the community. Further strategy for enduring crowded conditions may have been the maintenance of small body size. Taphonomic evidence and the crinoids' own adaptive strategies suggest a community that had quickly and opportunistically colonized a zone with ample attachment possibilities, high enough current velocities for sufficient feeding, yet sheltered enough to accommodate the crinoids' weak attachment capacity in all but the strongest storms.

ii iii Acknowledgements

From the beginning, Dave Meyer has gone above and beyond in championing my efforts to further my education this late in the game. His kindness and support have been invaluable.

Brenda Hunda provided friendship, advice, and access to fossil material. Carlton Brett’s invertebrate paleontology classes were consistently exciting and inspiring. Dan Cooper discovered and gently excavated the slab. Steve Felton knew where he’d found it and took us back there, while Bob Bergstein provided financial assistance; both of these fine Dry

Dredgers were full of good advice and encouragement. Roger Cuffey graciously prepped and identified the bryozoans. Ben Dattilo very generously instructed in thin-sectioning, sanding, and polishing of mudstones, as well as providing the XRF scans, which Barry

Maynard kindly helped to interpret. Patrick was very patient with my frustrations when it came to certain computer applications, and for that, as well as every kind of support imaginable, thank you.

iv Table of Contents

Title Page

Abstract i

Acknowledgements iv

Table of Contents v

List of Figures vii

List of Tables x

Introduction 1 Stratigraphic Setting 4 Maysville West 7 The Lagerstätte 10

Materials and Methods 14

Taphonomy: Introduction 30 Obrution and Autochthony 32 Paleoenvironment 35 Crinoid Remains: Introduction 38 Measurements: Materials and Methods 39 Results and Discussion 39 Preservation: Materials and Methods 47 Results and Discussion 53 Implications of Early Decay 65 Orientation: Materials and Methods 68 Results and Discussion 71 Incidental Elements 82 Geochemistry 88 Mechanism for Obrution 94 Sediment Capture 100

v Paleoautecology 102 Morphology of Glyptocrinus decadactylus 104 Implications for Paleobathymetry 104 Implications for Preservation Position 107 Implications for Attachment 110 The Question of Autotomy 111 The Community 114 The Holdfast Forest 118 Community Strategy 123 Comparison with Other Lagerstätten 129

Conclusions 134

References 136

Appendices 145

vi List of Figures

Figure 1. A) General location of study area. B) Excellent illustration of Glyptocrinus decadactylus by John Agnew. C) Stratigraphic column of Maysville West with arrow indicating approximate site of crinoid slab. Column modified from T. Schramm, 2011 Masters.

Figure 2. The principal sedimentary environments of the Cincinnatian. Illustration after Steve Holland in A Sea Without Fish, Indiana University Press, 2009.

Figure 3. A) The Maysville West roadcut is the NE-SW trending gray streak above the center of this Google Earth photograph. GPS of cut: 38° 40.407’ North, 83° 47.830’ West. B) Northwestern side of the Maysville West roadcut. The Bellevue Limestone is at top and the Fairview Formation is roughly the next three benches down. The light gray limestone bed indicated by arrow is a seismite horizon interpreted as part of the seafloor heavily disturbed by a paleoearthquake. The crinoid bed was 2-3 meters above this horizon. C) Site of Lagerstätte indicated by arrow. Photo by D.L. Meyer. Photographs A and B and information provided by James St. John ("A" via Google Terrametrics).

Figure 4. Two views of the reassembled slab. Most pieces fit together perfectly. Note the frequent overlap of individual slab pieces: measurements for depth were taken for the deposit as a single unit, not as individual pieces.

Figure 5. Each individual section of the slab was assigned a letter from A to T, with the largest, first-prepared piece designated "M" or M-slab. As M-slab was originally five pieces, these are depicted as M-1 through on M-5. Not every piece of the slab was prepared.

Figure 6. Schematic artwork from photograph of slab piece. Non-crinoidal elements appear yellow in artwork. A) Slab piece A. B) Slab piece B. C) Slab piece C. D) Slab piece D, which was originally in two pieces, later rejoined. E) Slab piece E. F) Slab piece F. There are no non-crinoidal elements. G) Slab piece G. There are no non-crinoidal elements. H) Slab piece H. I) Slab piece I, which was still being prepped at time of photo. The brown spot at left is the lower surface of the slab as it appeared prior to preparation. J) Slab piece J. K) Slab piece K. Photograph currently unavailable. L) Slab piece M. M) Slab piece T. Photograph currently unavailable.

Figure 7. Histogram comparing calyx heights of G. decadactylus from both Maysville slab and local Cincinnatian.

Figure 8. Scatterplot comparing the stem lengths and calyx heights of the Maysville slab G. decadactylus.

Figure 9. Idealized examples of taphonomic grades.

vii Figure 10. Examples of calyx orientation. A) Crinoid in oblique orientation directly above another specimen in lateral orientation. B) Examples of crinoids in oral-aboral position, the “starburst.” C) Crinoids in lateral position, the “modified shaving brush."

Figure 11. Schematic diagram showing hypothesized stalked crinoids in: A) parabolic filter fan (feeding) position, B) trauma position of crinoid with non-muscular arm articulations (the “modified” shaving brush), C) trauma position of crinoid with well-muscled arm articulations (the shaving brush). Modified after Baumiller, et al., in Echinoderm Paleontology, 2008.

Figure 12. Examples of preservation effects. A) Well-preserved specimen missing stem at distal end. B, C) Specimens showing marks of scavenging on their calyces. D,E) Specimens in which the arms are lost distal to physical control. F) Heavily decayed specimens.

Figure 13. Grid system from which orientation was plotted for slab pieces M, A, B, C, D, H, I, J, and K as a single unit.

Figure 14. Detail of slab.

Figure 15. Orientation from slab pieces M, A, B, C, D, H, I, J, and K as a single unit. Total count: 397 elements. A) Laterally-oriented calyces in opening direction. B) Columns. Those without a coil are arbitrarily directed. C) Combined A and B.

Figure 16. Orientation of calyces and stems for three defined sections, as highlighted on page right. Number of specimens in each dataset listed on rose diagram, lower right.: A) Section 1. B) Section 2. (C) Section 3.

Figure 17. Orientation of fossil material for four defined areas, as highlighted on page right. Number of specimens in each dataset listed on rose diagram, lower right. Note that areas overlap. A) North. B) South. C) East. D) West.

Figure 18. These crowns are relatively consistent for orientation, though not direction.

Figure 19. Two crinoids, "A" in starburst, "B" in modified shaving brush, preserved at the same level on the slab.

Figure 20. Other elements from the slab. Note that as the slab is prepared from underneath, these are the undersides of the specimens as preserved. A) Moult remains of Isotelus. B) Cyclonema bilix lata. C) Dalmanella (Onniella). D) Hebertella. E) Rafinesquina with encrusted inarticulate Orbiculoidea. F) Cyrtolites minor.

Figure 21. Trepostome bryozoans from the slab. Atactoporella schucherti, Batostoma implicatum (or possibly Batostoma prosseri ), Batostomella gracilis, Parvohallopora ramosa, and Parvohallopora subplana.

viii Figure 22. Other elements from the slab. A) Ambonychia robusta. B) Rhodophyta. C) Thin section revealing Skolithos-type burrow in vicinity of crinoid remains.

Figure 23. XRF scan revealing sulfur, calcium, iron, and phosphorous as major elements of the slab. Bottom center is a crinoid fossil.

Figure 24. Extracted spectra from XRF, with enlargement for manganese (Mn). Note Mn concentration at upper surface.

Figure 25. Crinoid column fragment with dark pigmentation.

Figure 26. Schematic of lofting turbidity current into distal environment encountering the crinoid community on the rise. Not to scale. Modified from Potter, et al. in Mud and Mudstones: Introduction and Overview, 2005, and Wetzel and Meyer, 2006.

Figure 27. Construction of G. decadactylus A) Free arms B) Tegmen (hidden within arms) C) Fixed arms (brachials) D) Secundibrachs E) Radials (in black) F) Basals G) Distal coil H) Stem I) Interrradials (in blue) J) Calyx K) Crown (calyx + arms). Modified from Hess, H., Ausich W.I., Brett, C.E., and Simms, M.J. in Fossil Crinoids, 1999.

Figure 28. Schematic representing response of stalked crinoid with non-muscular arm articulations to increased current. Modified from Baumiller, et al., in Echinoderm Paleontology, 2008.

Figure 29. Crinoid crowns that appear to have separated from their stems during collection, rather than by autotomy.

Figure 30. Examples of crinoid stems anchored to other crinoids.

Figure 31. Artist's rendition of the Ordovician seafloor with an emphasis on seaweeds, from geology.wisc.edu.

Figure 32. Disparity in column length between similarly-sized crinoids. A) Short column. B) Long column. The size of the distal coils is a good indication of scale.

ix List of Tables

Table 1. Calyx heights and column lengths. Isolated column sections not measured if under two centimeters. Final 35 measurements from Echinoderm Paleobiology, Indiana University Press, 2008, provided by C. Brett.

Table 2. Definitions of taphonomic grades including number of specimens within each grade.

x Introduction Concentrations of fossils are always of special interest. They often display unusual or exceptional preservation, and they may provide a rare glimpse into the ecology of an ancient community. In 1970 Seilacher defined these concentrations as Fossil-Lagerstätten, and further refined the term as Konzentrat-Lagerstätten, for high fossil concentrations, and

Konservat-Lagerstätten, for those with unusually well-preserved fossil material. This last term best describes the subject of this study.

Lagerstätten are highly prized for their value to paleontology and paleoecology, but only recently has their genesis been subject to scrutiny. Long thought to be simple deposits resulting from rapid burial or stagnation alone, such concentrations are now recognized as the products of complex processes that can cover a variety of preservation effects (Brett et al., 1997).

Crinoids disarticulate quickly, and so are typically preserved as isolated ossicles

(Meyer, 1971; Meyer et al., 1989; Kidwell and Baumiller, 1990; Brett and Seilacher, 1991;

Ausich and Sevastupolo, 1994; Brett et al., 1997; Ausich, 1998, 2001; Gahn and Baumiller,

2004). As widespread as they were during the Paleozoic, this amounts to crinoids being some of the most important producers of carbonate sediments at the time. Resulting deposits of crinoidal limestone can be tens of meters thick and cover thousands of kilometers, and yet very few of the remains are recognizable to species level (Ausich, 1997,

1999). Therefore it is all the more remarkable to find very well-preserved crinoids and especially so with their community structure intact.

A Lagerstätte of Glyptocrinus decadactylus Hall, 1847 representing an obrution deposit was collected in the 1990's from a roadcut known as Maysville West, situated in

1 Upper Ordovician rocks outside Maysville, Kentucky (Fig. 1, A-C). This deposit presents an excellent opportunity for both taphonomic analysis and for further understanding of the paleoautecology of camerate crinoids of the Late Ordovician. Therefore it has been the purpose of this research project to (1) continue the preparation of the Lagerstätte deposit begun in 2003 so as to make more raw data available, both for this study and future research; (2) reconstruct the paleoenvironment based on multiple factors, including lithologic structures and crinoid morphology; (3) investigate the origin and results of the killing mechanism, both for lithologic and taphonomic effects; (4) gain insight into the paleoautecology of Glyptocrinus decadactylus, particularly regarding community structure and specific life strategies employed. It is hoped that these findings will prove useful to understanding not just the Maysville deposit, but will add to the growing database of

Lagerstätten worldwide which are proving invaluable to paleontology.

To be noted: throughout the paper, the term "Lagerstätte" is often substituted with

"the deposit," "the slab," the Maysville slab," or the "Maysville deposit." Unless stated otherwise, all of these terms should be considered synonymous. Likewise, “column” and

“stem” are used interchangeably, although “crown” is not interchangeable with “calyx”: rather, crown refers to all structures above the stem, i.e., the calyx, tegmen, and arms.

2 B

B C

FIGURE 1 — A) General location of study area. B) Excellent illustration of Glyptocrinus decadactylus by John Agnew. C) Stratigraphic column of Maysville West with arrow indicating approximate site of crinoid slab. Column modified from Schramm, 2011.

3 Stratigraphic Setting

The paleocontinent of Laurentia was rotated roughly 45° clockwise relative to its current position during the Late Ordovician, with the Cincinnati Arch located between 20° and 25° south latitude. The resulting climate was very warm and semi-arid, as regions in similar positions are today (Dattilo et al., 2008; Holland, 2009; Meyer and Davis, 2009). The bulk of the terriginous sediment laid down in the area at the time was derived from the Taconic

Orogeny forming the Taconic Mountains to the east; these sediments were deposited on and created a northeastward-dipping mixed carbonate-siliciclastic ramp of a shallow epicontinental sea between 451 and 443 million years ago (Wahlman, 1992; Jennette and

Pryor, 1993; Holland, 1993, 2008; Cuffey, 1998; Holland et al., 2001, 2006; Meyer et al.,

2002; Holland and Patzkowsky 2007; Dattilo et al., 2008; Dudei, 2009; Meyer and Davis,

2009).

The Cincinnatian Series are the uppermost rocks of the Ordovician System, and are defined by the rocks exposed in the vicinity of southwestern Ohio, northern Kentucky, and southeastern Indiana (Peck, 1966; Coogan, 2002; Holland, 2008). Several facies occur in this series, but four dominate: these are the offshore, deposited below wave base where the seafloor was affected by only the most severe storms; the deep subtidal, deposited in a shallower environment, the transition zone between storm and fair-weather wave base; and the shallow subtidal, deposited above normal wave base (Fig. 2). The fourth environment, the peritidal or tidal flat, is barely fossiliferous and less common, and it does not occur at the Maysville West roadcut (Meyer et al., 2002; Holland, 2008, 2009).

These rocks are divided into six large-scale depositional sequences, each comprising a basal transgressive systems tract with a thicker highstand systems track overlying it;

4 these formed as the ramp was repeatedly building northward over time. Facies within transgressive systems tracts indicate a deepening upward trend, while those of highstand show a shallowing upward trend (and lowstand system tracts, indicating sea level fall to the point of exposure, are not found in the Cincinnatian). As most of the Cincinnatian sequences consist primarily of highstand system tracts, the Cincinnatian is said to display an over-all shallowing-upward trend. The depositional sequences are numbered C1-C6, oldest to youngest, deepest to shallowest. Each consists of a 1-10 million year time unit, and as such are considered third-order sequences (Holland, 1993, 2008; Holland and

Patzkowsky, 1996; Holland et al., 2001, 2006).

The C1 and C2 sequences in the lower part of the Cincinnatian Upper Ordovician comprise, firstly, the Kope Formation, considered an offshore facies. Above it lies the

Fairview, a deep-subtidal facies, and overlying both is the Bellevue Limestone, a shallow- subtidal facies. The Kope, Fairview, and Bellevue succession are the first of three major units that compose the Cincinnatian Series, and they are well exposed at Maysville West.

Sea level

Tidal flat Fairweather wave base

Storm wave base Shallow subtidal

Deep subtidal Offshore

FIGURE 2 — The principal environments of the Type-Cincinnatian. Illustration after Steve Holland in A Sea Without Fish, Indiana University Press, 2009.

6 Maysville West

The Maysville West roadcut is on the KY Rte 62/68 bypass, approximately two miles northwest of Maysville, Mason County, on the Kentucky side of the Harsha Bridge. (Fig. 3A)

It is one of the best-exposed roadcuts in the Cincinnatian, impressively wide, high, and laterally extensive; its towering faces expose the middle and upper Kope Formation

(Edenian Stage, lower Cincinnatian Series), the Fairview Formation, and the Bellevue

Limestone (the latter two lower-to-middle Maysvillian Stage, middle Cincinnatian Series)

(Jennette and Pryor, 1993; Holland, 1993, 2008, 2009; Holland et al., 2001, 2006; Meyer et al., 2002; Dattilo et al., 2008; Fig. 3).

The Kope and Fairview Formations are the lowermost in the upward-shoaling

Cincinnatian Series and consequently in the roadcut as well. The Kope at roadcut base consists of over 80% mudstones or shales. The overlying Fairview is composed of interbedded terrigenous mudstone, calcisiltite, skeletal packstone, and grainstone, averaging out to a ratio of roughly 50/50 shale to limestone (Peck, 1966; Meyer et al., 2002;

Holland, 2008); it is more than 25 meters thick in the area around Maysville (Peck, 1966).

The Bellevue Limestone exists at the top of the roadcut as a single shelf (Fig. 3C). The original site of the Maysville Lagerstätte was approximately 2-4 meters above a conspicuous seismite horizon in the Fairmount, the upper member of the Fairview, and just below the unconformity that distinguishes the Fairview from the Bellevue Member, as depicted in Figure 3C. Glyptocrinus has been known to appear in the Upper Kope, but this is rare. It is generally found in the Upper Fairview, most often in the Fairmount member, still deep subtidal but bordering on the shallow subtidal Bellevue (Meyer et al., 2002). The

Lagerstätte's position has been interpreted as occurring in the Miamitown Shale equivalent

7 (the Miamitown Shale, an arguably deeper water facies, does not extend into Kentucky)

(Brett et al., 2008). Further investigation would be required to verify this.

8 A B

FIGURE 3. A) The Maysville West roadcut is the NE-SW trending gray streak above the center of this Google Earth photograph. GPS of cut: 38° 40.407’ North, 83° 47.830’ West. B) Northwestern side of the Maysville West roadcut. The Bellevue Limestone is at top and the Fairview Formation is roughly the next three benches down. The light gray limestone bed indicated by arrow is a seismite horizon interpreted as part of the seafloor heavily disturbed by a paleoearthquake. The crinoid bed was 2-3 meters above this horizon. C) Site of Lagerstätte indicated by arrow. Photo by D.L. Meyer. Photographs A and B and information provided by James St. John ("A" via Google Terrametrics).

9 The Lagerstätte

The Lagerstätte slab was collected in 24 pieces, which, when fitted together, total more than 1.3 m. The shape is roughly that of an elongated right-angle triangle, averaging approximately 4-10 cm thickness (Fig. 4). The thickness is relatively consistent throughout until what is designated the "south" margin, or the "point" at the end of the lens-triangle.

This designation is for convenience only, as the deposit was collected without orientation data. This unfortunate deficiency necessitates a terminology: therefore, the right angle of the right-angle triangle is designated the "west" side, the hypotenuse of the triangle the

"east" side, and the remaining side the "north" side; the "south" side is better described as a long point, as previously stated. It is this “point” that most convincingly displays signs of being the outer edge of the deposit, as it contains fewer and more widely spaced crinoid remains, as if the Lagerstätte is literally "tapering off."

A subsequent field trip in 2010, more than ten years after the bulk of the slab was initially collected, yielded a small amount of new material. The original site has become greatly overgrown in the intervening years, however, and once more than 100 cm of overburden and a great deal of overgrowth, including several young trees, had been removed, the resulting ditch quickly filled with deep, soupy mud. All new prospecting was therefore done by feel rather than sight. Interestingly, this was easier than imagined, for reasons that underscore the Lagerstätte deposit's dissimilarity in density and texture from the more indurated underlying limestone, from which it was quite distinct and came away easily.

The new material was collected in five small pieces. The largest is roughly 9 X 9 cm and 4 cm thick, 1 cm at the thinnest edge; the smallest is 7 X 6 cm and 2 cm thick. The new

10 samples are lithologically identical to the material already collected, and only differ in the thinness and sparseness of crinoid remains, only 1-3 animals per piece. However, these new pieces are consistent with material from the south point of the slab as per both their shape and thickness, suggesting that this was their original position and that the deposit is indeed thinning at an outer margin. However, the years between collecting times make it impossible to determine this absolutely.

Despite the thinning of the mudstone and the decreased fossil density of the new material, suggestive of a margin, it is difficult to know if the shape and lateral extent of the deposit is complete. It may be noted that some similar fossil deposits from other locations display "deep, angular indentations along the margins" (Meyer and Milsom, 2001). This has been attributed to "tearing" or separation of the crinoid layer where the fossil concentration is high at thinning margins; margins may also show more disarticulated fossil material, such as in the case of a slab from the Senckenburg Museum (Neugebauer,

1978) and a lens of Pentacrinites fossilis from the Jurassic of Britain (Simms, 1986). These features are not evident on the Maysville slab, however.

The north edge of the slab is consistent with the center with regards to thickness and fossil density. There is nothing at the north end to indicate that the deposit is tapering off. The lack of the aforementioned indentations at the north end, however, are not diagnostic, as they are also nonexistent at the tapering south end. And certainly similar crinoid deposits have proven very extensive in similar conditions. The Maynes Creek

Formation contained a lenticular deposit of ~6 meters in diameter (Gahn and Baumiller,

2004), while Meyer and Milsom (2001) found a lens in Colorado of over 88 m. In 1901,

Springer described a lens measuring 6 X 15 meters.

11 The underside of the slab is smooth and slightly undulating and sharply bounded at its contact with the underlying condensed limestone; this limestone is fossiliferous and contains disarticulated crinoid parts, mainly stems. Glyptocrinids consistent with the preservation and particular description of those within the Lagerstätte have not been found in the underlying limestone, however. The upper surface of the deposit is slightly compromised, with a weakened, crumbly structure; this has likely been exacerbated by the deposit's position at the erosive top of the roadcut, resulting in its being overlain by recent soils. The slab's upper surface also shows evidence for having been at the sediment-water interface for a period of time ("Geochemistry," this document).

FIGURE 4 — Two views of the reassembled slab. Most pieces fit together perfectly. Note the frequent overlap of individual slab pieces; measurements for depth were taken for the deposit as a whole, not as individual pieces.

13 Materials and Methods

Preparation with vibratory and air abrasive tools was begun in 2003 by Scott Vergiels, who, as well as revealing a large section, fitted and fixed together five separate pieces into one which was subsequently displayed at Cincinnati Museum Center. Preparation was resumed by the author in 2008 using similar methods, including that of preparing from the slab's lower surface, where more fossil material is concentrated. Each individual section of the slab was assigned a letter from A to T, with the largest, first-prepared piece from 2003 designated "M" or M-slab. As M-slab was originally five pieces, these are depicted as M-1 through on M-5 in Fig. 5. Measurements were taken with calipers for calyx height, column length, and distal coil width; these measurements were compared within the dataset and to other G. decadactylus calyces from the type-Cincinnatian. Orientation of crinoid elements was calculated in relation to one another. Taphonomic stages were assessed and graded on a 32-point scale. Photographs of the deposit were altered in Photoshop to create detailed outlines for clarity and the results divided into Photoshop "layers" representing orientation and measurement data, as well as a separate layer for non-crinoidal elements (Fig. 6A-L); the non-crinoidal elements were thin-sectioned and acetone peels made for identification, when necessary. Preparation for fabric analysis was conducted at the Dept. of Geosciences,

Indiana-Purdue University Ft. Wayne, on material collected in 2010: these were cut into .5-

1 cm thin sections and analyzed for primary sedimentary structures, ichnofabric, and lithologic changes and anomalies; elemental composition of thin sections was determined through X-ray fluorescence (XRF), the process of bombarding a sample with gamma rays in order to elicit the emission of fluorescent x-rays, utilizing a Bruker M4 Tornado scanner

(Van Grieken and Markowitz, 2002; Dattilo, pers. comm.). All other preparation was done

14 at the Geier Center, Cincinnati Museum Center, also the repository of the material studied

(CMC 50668). Not every piece of the slab was prepared, and it is hoped that the largest unprepared high-density piece, designated "N," is to undergo magnetic resonance image

(MRI) scanning, a process with excellent implications for the preservation of taphonomic features. Materials and Methods are outlined in greater detail at the beginning of relevant chapters throughout the document.

FIGURE 5 — Each individual section of the slab was assigned a letter from A to T, with the largest piece designated "M" or M-slab. As M-slab was originally five pieces, these are depicted here as M-1 through on M-5. Not every piece is prepared.

FIGURE 6A — Schematic artwork from photograph of slab piece A. Non-crinoidal elements appear yellow in artwork.

FIGURE 6B — Schematic artwork from photograph of slab piece B. Non-crinoidal elements appear yellow in artwork.

FIGURE 6C — Schematic artwork from photograph of slab piece C. Non-crinoidal elements appear yellow in artwork.

FIGURE 6D — Schematic artwork from photograph of slab piece D, which was in two pieces, later rejoined. Non-crinoidal elements appear yellow in artwork.

FIGURE 6E — Schematic artwork from photograph of slab piece E. Non-crinoidal elements appear yellow in artwork.

FIGURE 6F — Schematic artwork from photograph of slab piece F. There are no non-crinoidal elements.

FIGURE 6G — Schematic artwork from photograph of slab piece G. There are no non-crinoidal elements.

FIGURE 6H — Schematic artwork from photograph of slab piece H. Non-crinoidal elements appear yellow in artwork.

FIGURE 6I — Schematic artwork from photograph of slab piece I, which was still being prepped at time of photograph. The brown spot at left is the lower surface of the slab as it appeared prior to preparation. Non-crinoidal elements appear yellow in artwork.

FIGURE 6J — Schematic artwork from photograph of slab piece J. Non-crinoidal elements appear yellow in artwork.

FIGURE 6K — Schematic artwork from photograph of slab piece K. Non-crinoidal elements appear yellow in artwork. Photograph currently unavailable.

FIGURE 6L — Schematic artwork from photograph of slab piece M. Non-crinoidal elements appear yellow in artwork.

FIGURE 6M — Schematic artwork from photograph of slab piece T. Non-crinoidal elements appear yellow in artwork. Photograph currently unavailable.

29 Taphonomy: Introduction

Obrution deposits present an unusual opportunity for the study of taphonomy. Those with high concentrations of well-preserved fossils are of special interest not in the least because they provide a "snapshot" or record of a brief moment in the life and death of an ancient marine community (Brandt, 1989; Brett and Seilacher, 1991; Taylor and Brett, 1996;

Briggs, 2003). For most of the history of the discipline, taphonomy was, in fact, considered a branch of paleoecology, and its processes were seen as primarily destructive; it was considered necessary to "strip away the taphonomic overprint to make ecological inferences" (Brett, 1986), as if the less "taphonomy" a site had undergone, the better.

Taphonomy has undergone a minor revolution and become a discipline in itself. The preservational processes which were formally seen as taking information away from a fossil are now known to contain valuable information regarding the depositional environment and the environmental processes which originally produced the fossil. Fossils are now considered very sensitive paleoenvironment indicators (Brett, 1986, 1995;

Thomas, 1986; Brett and Seilacher, 1991; Taylor and Brett, 1996; Gahn and Baumiller,

2004; Thomka, 2011). This is not an entirely new idea: William Buckland made note of fossil preservation as an indicator of sediment rate in his Bridgewater Treatise in 1836.

Nevertheless, the concept was still under-utilized and underdeveloped enough that David

L. Meyer and Carlton Brett saw reason to organize a symposium at the 1984 GSA meeting devoted to exploring "The Positive Aspects of Taphonomy." Although the symposium's focus was on the value of taphonomic processes for reconstructing a paleoenvironment, it is recognized that this information, in turn, says much about where and therefore how an animal lived — and so paleoecology is brought back into the equation.

30 The study of taphonomy is generally divided into two categories, summarized as the mechanical and chemical. The mechanical is biostratinomy, which is concerned with the preservational interactions and processes an organism undergoes from the moment of its death to its burial. These would include anything that could have modified the remains, such as the mode of death, scavenging, transport, and disarticulation, among others. The chemical or diagenetic processes are what occur post-burial; the most important of these including lithification and compaction of the sediment, and leaching and mineral replacement of hard parts of the animal. The preservation of a fossil primarily depends on many versions of these two factors in varying amounts; the final result is a fossil that retains the characteristics of the environment where the animal lived and/or died

(Fagerstrom, 1964; Lawrence, 1968; Behrensmeyer and Kidwell, 1985; Brett and Baird,

1986; Gahn and Baumiller, 2004). In this way a paleoenvironment can be reconstructed in the absence of diagnostic sedimentary structures.

The taphonomy of a site such as that of the Maysville Lagerstätte thus yields data which mirror the conditions of the depositional environment, the nature of such depositional events in general, and the paleoecology of the animals preserved (Brett and

Baird, 1986; Brandt, 1989; Taylor, 1996). The following chapter focuses primarily on the depositional profile of the slab and the processes that created it, and the preservation of the fossils and its implications for the mode and rate of sedimentation. This leads to a reconstruction of the environment in which the crinoids lived and the final burial event; although it is not the focus of this chapter, overlap into the paleoecology of the crinoid community is inevitable.

31 Obrution and Autochthony

The Maysville Lagerstätte is comprised of a thin, medium gray-brown mudstone lens. The sediment is compacted and lithified to a homogenous mud fabric, that is, it is ungraded and consistent for grain size throughout. The mudstones are densely packed with extraordinarily well-preserved fossil material of an otherwise delicate animal, prone to rapid disarticulation and dissolution very quickly under normal circumstances. All of these features are consistent with rapid episodic sedimentation, which has a strong association with obrution, although in this case without the strong current disturbance associated with high-energy storm deposits.

The preservation of the fossil material is the strongest evidence for obrution. Brett et al. (2012) distinguished between two types of obrution deposit: Type I, representing rapid burial of fossil assemblages preserved without bottom disruption within their habitat, and Type II, representing deposition by bottom-flowing currents, leaving behind signature sedimentary structures and fossils which had been transported. The Maysville deposit is of the first category.

Fully articulated, fragile, multi-element skeletons in life position are generally diagnostic of rapid, in situ or autochthonous burial (Brett and Baird, 1986; Kidwell et al.,

1986; Speyer and Brett, 1986; Meyer et al., 1989; Allison, 1998; Datillo et al., 2008). The exceptional preservation and lack of size sorting of the Maysville slab crinoids indicates little or no transportation (Brett and Baird, 1986; Speyer and Brett, 1986; Brandt, 1989), and their spatial distribution is similar to what is seen in living Recent assemblages ("The

Community," this document). All this, as well as the fact that monospecific assemblages are strongly associated with intact colonies, indicates that the crinoids were buried within

32 their habitat (Messing et al., 1990).

Episodic sedimentation is generally associated with random and severe events such as large storms or seismic shocks. However, while the obrution that generated the slab was almost certainly storm-generated, it can be difficult to recognize precisely the type of storm-generated event represented, particularly in distal environments (Brandt, 1989;

Wheatcroft, 1990; Brett and Seilacher, 1991; Thomka, 2010). The Maysville slab does not display the poorly-sorted, fining-upward, graded bed characteristic of turbidites, nor does it show the aligned tool marks and flute casts, current ripples, or cross lamination which are typical indicators of turbidity currents (Kreisa, 1981; Brett and Baird, 1986; Holland,

2009).

Rather, the slab's lack of internal lamination, primary sedimentary structures, or fossil evidence of strong current alignment appears to be the product of a lofting turbidity current (Traykovsi, 2000; Wright, 2001; Mulder, 2003; Potter et al., 2005; Gladstone and

Pritchard, 2010): having shed its coarser and heavier sediments more proximally, it is not uncommon for the distal reaches of such a current to loft into a plume from which the finest sediments are shed far from the current's site of origin. The low-energy, rapid deposition that is usually associated with distal areas can be the result of fine suspended sediments which may have traveled great distances before their final deposition (Taylor and Brett, 1996; Brett et al., 2012; "Mechanism of Obrution," this document).

This leads to the question of whether or not the slab represents a single depositional event. Temporal resolution is often difficult to determine in mud deposits (Brett and Baird,

1993; Brett and Allison, 1998; Meyer et al., 1989). Yet despite the density of crinoids, it is suggested that only one event created the Lagerstätte (although pre-obrution effects left an

33 apparent signature: "Preservation: results and discussion," this document). The crinoids are entombed within a single lens, primarily within the lower third of the mudstone slab yet with the occasional fossil element crossing through to the top, and more than 80% are oriented parallel to the bedding plane; this, combined with the thinness of the deposit suggests a single smothering event (Fagerstrom, 1964; Brett et al., 2012). Likewise, the slab's sharply-bounded lower surface is thought not to result from storm scouring, as is often the case, but from the rapid onset of deposition of fresh sediments (Kreisa, 1981).

The fossiliferous underlying limestones are another indicator, as obrution horizons are often found on the upper surface of such shelly limestone beds (Datillo et al., 2013).

34 Paleoenvironment

The sediments of the Maysville Lagerstätte are interpreted as having been deposited on a moderately energetic rise elevated slightly above the deep subtidal regions. This interpretation is based upon an analysis of the depositional setting and the morphology of the crinoids themselves.

To reiterate, the Fairview represents a deep-subtidal environment deposited between fair-weather and storm-wave base, a transition zone from the offshore of the Kope to the shallow-subtidal of the Bellevue Limestone. Only the most powerful storms would have extended deep enough to disturb the crinoids in their habitat. However, if modern climate systems and those of the Ordovician are similar, then the position of the area and the tilt of the continent would have meant that very large storms were a frequent occurrence, making storm deposits the most prominent feature of the deep subtidal in the

Cincinnatian (Dattilo et al., 2008; Holland, 2009).

The very fine sediments shed far from the source of coarser sediments in the proximal areas, the thin sediment blanketing, and the slight suggestion of multi-directional currents are consistent with markers for deposition having occurred in a distal area.

Furthermore, the "smothered bottom" obrution horizon, composed of suspended sediments winnowed from proximal areas, is an almost exclusively distal horizon phenomenon (Kreisa, 1981; Brett and Baird, 1986; Thomka, 2010).

Observations by Kreisa (1981) in the Recent as well as in paleoenvironments indicate that water depth is positively correlated with finer sediments and weaker current activity, and also that storm layers tend to be thicker in shallower water and thinner at

35 depths, rarely more than 10 cm in deep offshore environments. This corresponds with the relative thinness of the slab.

The preservation of the fossils is another strong indicator of depth. Many storm event beds are associated with very good fossil preservation, but distance from the shoreline often leaves a characteristic imprint. The energy of an environment has a direct effect on the condition of skeletal elements. Fossil material from proximal or near-shore beds, because it is deposited under high-energy conditions and subject to wave scour, is usually heavily-tumbled, fragmented, and well-sorted. Fossil condition from beds farther offshore or medial beds is usually better, with less breakage and sorting, and there is usually a degree of preferred orientation. Distal beds such as the Maysville slab show the best preservation of all, with a high proportion of articulated, unbroken, and unsorted fossils showing little evidence of transportation or orientation (Brett and Baird, 1986;

Speyer and Brett, 1986; Brandt, 1989). Therefore, although a proximal storm event likely began the process that resulted in the crinoids' burial, it also seems likely that the energy of the event had greatly dissipated by the time it reached the crinoids' habitat.

Glyptocrinid fossils are typically found in association with deep subtidal facies, and the Maysville Lagerstätte is no exception (Meyer et al., 2002). Yet there is some indication that the Maysville community was elevated as well. G. decadactylus is one of the more robust forms of crinoid, with a feeding structure best adapted to a more energetic setting, and there are indications that it was a generalist and an opportunist that took advantage of more energetic settings within its deep subtidal environment (""Morphology, " this document; "Community strategy," this document). Many bioclastic limestones representing

36 such settings are, in fact, largely composed of the distinctive elements of glyptocrinids

(Brett et al., 2008).

There is another consideration for the crinoids' habitat as well which would have implications for water depth: it is proposed that the crinoids were using plant matter, probably seaweeds, as holdfast anchors. There is evidence of plant matter throughout the

Fairview, indicating that it was within the photic zone (Meyer and Davis, 2009). Yet what appears to be preferential growth at the site implies that elevation above the deep subtidal might have been conducive to the density of plant growth that the associated density of crinoids suggests ("The Holdfast Forest," this document).

37 Crinoid Remains: Introduction

The Maysville Lagerstätte is composed of complete specimens as well as isolated crowns, brachials, and stem elements totaling more than 400 individual fossil crinoids, all belonging to Glyptocrinus decadactylus. There are over 250 crowns exposed to various degrees, and more than 10% of the crinoids are intact from crown to holdfast with at least half their arms. Most fossil elements took on the color of the ambient mudstones, although a small number are preserved black ("Geochemistry, " this document). They are smaller on average than others of their species found in neighboring areas but are physiologically mature

("Community Strategy," this document). All complete and many partial stems terminate in the distal coil that served as the holdfast in Glyptocrinus.

The crinoids do not appear to have been transported, but are still preserved in positions which indicate the energy of the burial event, and their orientations suggest the nature of its currents.

There are abundant crinoid remains at varying stages of disarticulation, ranging from lightly fragmented specimens to a great number of stem pieces. Multiple factors can influence preservation: these include physical reworking through tumbling, etc., scavenging/predation, and decay (Meyer, 1971; Kidwell and Baumiller, 1990; Baumiller and Ausich, 1992; Baumiller et al., 1995). Not all of these are necessarily factors for the

Maysville Lagerstätte. There is little indication of physical reworking. Predation is a possibility, but the fossil remains provide little evidence either for or against it

(Gluchowski, 2005). The evidence suggests that scavenging and decay account for the discrepancies in preservation, and these will be examined in more detail.

38 Measurements

Materials and methods

Measurements were taken using calipers and an ocular micrometer: crown height was measured from the stem attachment site at the basals to the top of the secundibrachs, the point from which the tertibrachs extend and pass on to free arms (Fig. 27); all stems articulated to a calyx were measured, although isolated stem segments were included when over two centimeters. String was used to include the distal coil in the stem measurement, and the diameter of the inside of the coil was taken for 56 specimens. Data for this study was added to measurements taken earlier from M-slab and published in Echinoderm

Paleobiology (Brett et al., 2008), with the exception of distal coil width, which were taken according to a different criteria in the earlier study and so are not used here. Further measurements for crown height were taken of all available specimens of G. decadactylus in the collections at Cincinnati Museum Center and compared to corresponding data from the

Maysville slab.

Results and discussion

The Lagerstätte crinoids are relatively small when compared to other specimens collected from the areas around Maysville ("Community Strategy," this document). Maysville specimens display a mean calyx height of 10.4 mm (SD = 2.7 mm). In contrast, specimens from the CMC collections show a mean calyx height of 16.8 mm (SD = 2.8 mm; Fig. 7). Stem length was found to have a relatively strong correlation to calyx height, in agreement with earlier measurements taken by Brett et al. (2008) on M-slab, yet with "considerable

39 scatter" (r = 0.44, Pearson's correlation: p < 0.001; Fig. 8). More than 10% of the specimens are complete, and complete stems range from 4.3 to over 16 cm long (Table 1).

All complete specimens terminate in Glyptocrinus' characteristic distal coils, with an average loop diameter of 4.32 mm (SD = 1.786 mm). The stems show a slight distal taper, and the columnals are cuneiform at the coils, widest in diameter at the outside of the curve.

FIGURE 7 — Histogram comparing calyx heights of G. decadactylus from both Maysville slab and local Cincinnatian ("Other").

Calyx height in mm in height Calyx

Stem length in mm

FIGURE 8  Scatterplot comparing the stem lengths and calyx heights of the Maysville slab G. decadactylus.

Calyx Height (mm) Stem Length (cm) 9.4 1.4 TABLE 1— Calyx heights and stem lengths. — 8.2 Isolated stem sections not measured if 13.7 — under two cm. Final 35 measurements 9.5 — 9.3 — from Echinoderm Paleobiology, Indiana 11.3 2.4 University Press, 2008, provided by C. — 5.2 Brett. — 2.2 — 6.2 16.5 — 15.3 — 8.4 — 6.7 — — 5 10.7 — 8 14.2 11.3 10.4 complete 9.2 5. 5 6.9 3.5 7.5 4.3 complete — 9.6 12 — 13.7 — 11.6 — 6.9 2.1 11.5 1.5 13.9 3.2 13.6 6.7 12 7.2 — 6 — 6.5 10.5 6.8 — 2.6 — 3.2 4.2 1.9 12.8 7 17.1 3.3 — 3.6 6.6 2 — 4.6 6.5 2.4 11 9.4 complete — 2.2 — 2.6 — 2.2 8.1 2 9.1 .7 10.4 1 16.3 .9 11.4 1.3 12.5 7 — 2.5 10 2 11.8 — — 8.4 — 3.7 — 8.2 — 2.6 9.5 4.2 11.5 13.6 complete 13.2 2.8 — 4.6 12.9 .5 7.3 2.5 10.7 11.6 complete 7.7 .6 8.1 8.5 complete — 12.2

43 Calyx Height (mm) Stem Length (cm — 10 TABLE 1 continued. — 8 — 8.1 9.5 5 8.9 7.4 complete 8.1 5 — 2.6 6.3 5.9 complete — 3.8 — 3.5 — 5.9 12.8 11.6 complete 9.8 6.6 complete — 14.2 10.1 7.6 complete — 6.6 — 12 9.4 6.4 complete — 12.2 4.9 7.1 8.1 1.9 — 13.6 — 9 — 2.8 6.2 2.9 — 2.8 — 4.6 — 6.6 — 5.3 8.9 9 complete — 5.2 — 5.6 12.3 4.4 — 2.9 — 2.6 — 3.4 8 5 13.2 — — 3 — 10.5 — 12 — 4.4 — 4.7 12.6 2.7 10 — 8.4 .9 — 4.3 — 7.3 — 7.8 — 7.6 — 3.5 10.6 2.2 8.8 .3 8.4 2.6 17.4 — 10.4 .3 — 10.7 11.3 1 — 6.3 — 6 — 7.1 — 11 10 3.3 8.3 1.5 — 8 11 8.6 5

44 Calyx Height (mm) Stem Length (cm) 9.5 3.7 TABLE 1 continued. — 6 8.8 1.3 9.8 — — 4.5 12.6 — 12.2 .4 10.4 — 9.4 4.8 10.1 — 4.2 6.1 complete — 5.8 8.7 6.2 — 7 11.5 6.5 — 3 11.2 10.7 9.1 .7 10 1.1 — 2.5 — 5.7 — 5.6 — 5 10.3 6.9 — 9.7 9.1 3.7 — 10 7.4 4.7 9.4 .8 — 5.3 — 4.6 6.1 2.4 6.2 3.5 10.7 — — 4.6 9.1 4 10.6 10.3 complete 11.4 10.2 complete 12.1 7.8 complete 9.5 10.1 complete 11.8 9.8 complete 8.2 14.1 complete 13.3 9.8 complete 13.3 11.1 complete 13 11.3 complete 12 11.2 10.8 8.3 complete 8.1 3.9 complete 15.6 14 complete 8.6 4.3 complete 12.3 10.9 complete 13.4 10.7 complete 7.6 9 complete 12.4 8.5 complete 7 2.9 complete 14.6 13.1 complete 7.2 4.2 complete 11.2 11.4 complete 15.3 7.5 9.7 6.3 complete 15.9 11.7 complete 12.2 6.3 15.5 16.6 complete 7.9 12.5 9.3 7.7 complete 12 7.7 complete 10 6.6 complete 9.5 9 complete

Calyx Height (mm) Stem Length (cm) TABLE 1 continued. 9.6 8.5 complete 9.3 8.2 14.7 10.5 complete

46 Preservation

Materials and methods

The Maysville Lagerstätte consists of many outstandingly well-preserved glyptocrinids, but there are also many decayed and damaged specimens; moreover, the preservation can be very uneven, even within individual specimens. Many studies have categorized degrees of crinoid preservation with regard to the entire animal (Brandt, 1989; Meyer, 1989; Kidwell and Baumiller, 1990; Ausich and Sevastopulo, 1994): for the purposes of this study, and the inconsistencies of the preservation in particular, a taphonomic grading system was devised to more accurately classify crinoid specimens according to the preservation of individual elements, with techniques borrowed from earlier studies (Meyer et al., 1989; Taylor and

Brett, 1996; Gahn and Baumiller, 2004; Thomka, 2011).

Two-hundred and fifty-three specimens were sufficiently exposed to include in the dataset, and each specimen was assigned a grade based on three particular factors, these being the condition and completeness of the stem, the calyx, and the arms (Fig. 9). Stems were numbered according to whether they were (1) complete, (2) partial long, (3) partial short, or (4) absent. After Gahn and Baumiller (2004), partial long was defined as an incomplete stem equal to or longer than the calyx; this is thought to more accurately reflect the stem's degree of completeness than a linear measurement, considering the variability of stem length. Partial short was defined as an incomplete stem shorter than the height of the calyx. A complete stem was that attached to a calyx at one end and concluding in a distal coil.

Calyces were assigned one of two grades. They were described as complete when the plates were all or almost all intact (allowances were made for a missing plate or two).

47 Calyces were partial when the calyx appeared to have collapsed inward upon itself, a condition that compromised the plates into a fragmented and/or jumbled state. A partial calyx may have a "compressed" look, but because such specimens frequently appear closely alongside fully-rounded calyces, their appearance was deemed not to result from actual compression. Calyces with multiple missing plates also received a rating of partial.

Arms received one of four grades: (1) complete/near complete; (2) 50% or less complete; (3) absent distal to physical control; and (4) absent/near absent. "Near complete" and "complete" received the same designation because arm-overlap could make it impossible to state with certainty the condition of every arm even where all appeared intact, although a reasonable inference could be made. "Absent distal to physical control" refers to the particular anatomy of glyptocrinids and its implications for limited control over arm movement and flexibility ("Morphology," this document). Finally, "absent/near absent" refers to both total absence and to the presence of short "nubbins" of arms.

Finally, each individual crinoid was designated a grade based on a combination of the preceding factors of four stem categories, two calyx categories, and four arm categories.

This results in a 32-point grading system, not every grade of which contains a specimen.

Each level of preservation was assigned a letter or double-letter grade, from, very generally, most complete (A) to least (FF), based on the typical patterns by which crinoids disarticulate after death according to previous workers (Meyer et al., 1989; Ausich and

Baumiller, 1993; Ausich and Sevastopulo, 1994; Taylor and Brett, 1996; Gahn and

Baumiller, 2004).

As the dissolution of the distal arms is the first stage in crinoid decay from which disarticulation proceeds proximally toward the calyx (Baumiller, 2003), arms were chosen

48 FIGURE 9 — Idealized examples of taphonomic grades.

49 as the best benchmark for decay. This resulted in four broad categories, each containing eight grades: Category 1, grades A-H, are all specimens with complete/near complete arms;

Category 2, grades I-P, are specimens with arms that are 50% or less complete; Category 3, grades Q-X, have arms lost distal to physical control; and Category 4, grades Y-FF, are missing arms entirely, or nearly so. The resulting grades within categories may be imperfect, but still reveal patterns not otherwise obvious.

Calyx position/angle was considered separately, under "Orientation," despite its importance to similar preservational grading systems in other studies (Thomka et al.,

2011; Gahn and Baumiller, 2004; Wetzel and Meyer, 2006). This is because, overwhelmingly, the lateral position, in which the long axis of the calyx lies parallel to the substrate, predominates (Fig. 10C). Occurrences of oblique position, wherein the axis of the calyx is at an oblique angle to the substrate, were deemed inconsequentially slight as to both the oblique angle of the calyces and the apparent lack of consequences for preservation (Fig. 10A). The strong exceptions to the ubiquitous lateral position are the specimens preserved in what Baumiller et al. (2008) called the "starburst," in which the long axis of the calyx is perpendicular to the substrate with brachials radially extended or splayed (Fig. 10B). This position reveals neither the calyx nor the stem (at least so that the stem could be paired with the specimen) although, the arms can be considered

"complete/near complete" by default, as anything but intact arms arrayed around a calyx would not register as a "starburst." Therefore, crinoids in starburst position were not graded according to their individual elements.

The calyces are well-rounded, that is, they are not collapsed or compacted where decay is not obvious. Incidental elements such as brachiopods and gastropods are likewise

50 largely well-rounded, as is a single mollusk preserved as an internal mold.

Finally, only specimens with calyces were included in the dataset, as the presence of at least a partial calyx was judged necessary to identify an individual crinoid. Isolated ossicles, including many detached arms and pluricolumnals (stem segments), were not included. Stem length was considered too variable to attempt to surmise missing crinoid numbers by counting stem fragments and adding them up to equal a complete crinoid.

Also, calyces too hidden from view were not included, for obvious reasons; "too hidden" meant at least half obscured by matrix or other fossil elements, the removal of which may have proven more destructive than instructive. These same calyces were, however, included in the body count of inferred glyptocrinid numbers, and result in the proposed

400+ count for total crinoids within the Lagerstätte.

B C

FIGURE 10 — Examples of calyx orientation. A) Crinoid in oblique position directly above another specimen in lateral position. B) Crinoids in oral-aboral position, the "starburst." C) Crinoids in lateral position, the “modified shaving brush."

52 Preservation: results and discussion

In considering the degrees of preservation displayed by the fossil crinoids, both intrinsic and extrinsic effects must be taken into account. Intrinsic effects would consist firstly of the disarticulation resistance of the crinoids, and secondly of the potential for preservation afforded by their behaviors. Extrinsic factors would be any outside forces acting upon the crinoids, such as scavenging. Effects for both factors and the abundance in which the effects appear are outlined in Table 2.

It has been demonstrated that the arms, calyx, and stems of crinoids disarticulate at independent rates. Crinoid decay experiments under normal conditions with Recent specimens show disarticulation is seen first in the loss of the distal arms, beginning with the pinnules and the tips of the arms and proceeding proximally to the calyx, followed by the collapse of the calyx and plate loss, leaving the stems the final element. Complete disarticulation into isolated ossicles usually requires close to two weeks time (Meyer,

1971; Meyer and Meyer, 1986; Meyer et al., 1989; Kidwell and Baumiller, 1990; Baumiller and Ausich, 1992; Taylor and Brett, 1996).

Disarticulation rates in Recent stalked crinoids are taken as analogs for the same in

Ordovician glyptocrinids in this study, despite that Ausich and Baumiller (1993) hypothesized that the arms and stem of the latter might actually disarticulate at the same rate; while sampling results seemed to bear this out, they were ultimately inconclusive.

However, when comparing disarticulation rates among different crinoid taxa, Gahn and

Baumiller (2004) found disarticulation in fossil crinoids to be directly proportional to the surface area of the articular surfaces and therefore inversely proportional to the non- articular surface area subject to extrinsic effects. This would support Recent and

53 Ordovician crinoids disarticulating at similar rates. Finally, the combination of both intrinsic and extrinsic effects on preservation may shed some light on the timing of the event(s) that led to the community's demise.

Each taphonomic category and grade is depicted in simplified form in Figure 9. The excellent preservation for which the Lagerstätte is known is reflected in Category 1, grades

A-H, consisting of crinoids with complete or near-complete arms. These are generally the best-preserved crinoids and, at 85 specimens, the single largest category on the slab. The grade with the greatest abundance is C (44 specimens), consisting of animals with complete/near complete arms and calyces and partial-long stems, followed by G (17 specimens ); A (14 specimens); and E (7 specimens). All four of these grades (within the category of eight grades) show the best arm preservation coupled with the best calyx preservation, as might be expected. By contrast, of the four grades with partial calyces, three have each a single specimen, and one (H) is the only grade out of 32 with no specimen at all (although this may not be surprising when considering that its other factors are a partial calyx and an absent stem: disarticulation rates and intuition suggest that for both of these to occur in a specimen with well-preserved arms would be unlikely).

The crinoids of Category 1 are interpreted to have been killed by the obrution event that created the Lagerstätte. Their preservation reveals very little evidence of either decay or scavenging, suggesting that these were hindered by anoxic conditions below the sediment/water interface. Thin sections from other areas of the slab reveal scavenging close to heavily-decayed crinoid elements; however, as no thin sectioning was done on slab pieces containing well-preserved elements, it cannot be stated definitively that scavenging ended with the final obrution event, but the preservation in Category 1 supports the idea.

Grade Calyx Stem Arms Total

Category 1 A complete complete complete/near complete 14 B partial complete complete/near complete 1 C complete partial long complete/near complete 44 D partial partial long complete/near complete 1 E complete partial short complete/near complete 7 F partial partial short complete/near complete 1 G complete absent complete/near complete 17 H partial absent complete/near complete 0 Category 2 I complete complete 50% or less complete 8 J partial complete 50% or less complete 4 K complete partial long 50% or less complete 8 L partial partial long 50% or less complete 9 M complete partial short 50% or less complete 6 N partial partial short 50% or less complete 1 O complete absent 50% or less complete 4 P partial absent 50% or less complete 4 Category 3 Q complete complete absent distal to physical control 10 R partial complete absent distal to physical control 1 S complete partial long absent distal to physical control 22 T partial partial long absent distal to physical control 1 U complete partial short absent distal to physical control 6 V partial partial short absent distal to physical control 1 W complete absent absent distal to physical control 2 X partial absent absent distal to physical control 5 Category 4 Y complete complete absent /near absent 3 Z partial complete absent /near absent 7 AA complete partial long absent/near absent 18 BB partial partial long absent/near absent 19 CC complete partial short absent/near absent 9 DD partial partial short absent/near absent 12 EE complete absent absent/near absent 6 FF partial absent absent/near absent 2

TABLE 2 — Definitions of taphonomic grades including number of specimens within each grade.

55 Considering, then, that nearly all within the category display well-preserved arms coupled with a well-preserved calyx, the preservation of the stem comes into question.

Only 14 specimens are in life condition, complete from pinnules to coil. Of the best preserved, then, 68 out of 85 are missing parts of their stems, 44 of them at the distal end

(Fig 12A). Yet stems usually have the highest preservation potential, primarily for their relative hardiness and to a lesser degree, their palatability for scavengers. Ausich and

Baumiller (1993) found that stems remained complete for up to five or six days in Recent crinoids, followed by separation into pluricolumnals over a 1-2 day period before finally disarticulating into isolated columnals after another week. It is therefore doubtful that the stems were lost to decay, and preferential scavenging for stems, which lack viscera or muscle tissue, is exceedingly unlikely. The stems were, therefore, probably broken in the obrution event, which in some cases, curiously, did no damage to the rest of the animal, a condition that initially suggested preferential shedding of the distal coil, or autotomy.

The stems of Recent crinoids are comprised of multiple pluricolumnal segments bound together by a "through-going" ligament and an intercolumnal ligament. The former is lacking at the articulation point between segments of an approximately constant length.

Baumiller and Ausich (1992) found some consistency in the lengths of isolated pluricolumnals in Recent crinoids, strongly suggesting that when stems disarticulate, they separate first along these points of relative weakness, both post-mortem and as autotomous behavior in life (Ausich and Baumiller, 1993). Similar patterns in the pluricolumnals of Mississippian crinoids led Baumiller and Ausich (1992) and Baumiller et al. (1995) to assert that these were likely, like the Recent specimens, capable of autotomizing their stalks. It is not known if Ordovician crinoids had the same ability, and

56 the non-preservable nature of the ligaments necessary makes it impossible to determine either way. However, a survey of pluricolumnal lengths by Ausich and Baumiller (1998) collected from the Ordovician suggests that they did have the similar points of weakness along the stem. Although the patterns of regularity for pluricolumnal length were not so pronounced as in the Mississippian or Recent crinoids studied, Ordovician specimens did show a pattern that was non-random at the 95-99% level. Pluricolumnal measurements for

Glyptocrinus fornshelli, for example, suggested that the segments of the stem might consist of 12 or 13 columnals each (Ausich and Baumiller, 1998).

The 44 distally-broken stems in grade C seem to indicate that breakage occurred preferentially down the stem, possibly at the attachment coil. The wide range of stem lengths seen on the complete crinoids, however, makes it difficult to determine whether breakage occurred immediately before the coil or if there was originally more length to the stem. It's also questionable whether or not this would indicate autotomy. It's possible that the behavior of autotomy had yet to evolve, and it would seem to have limited benefit for crinoids without reattachment capability (Baumiller et al., 2008; "Morphology," this document). It may be more likely that the stem simply snapped under stress at one of the naturally occurring points of weakness along the stem. Furthermore, the shape of the columnals at the distal coil may have been a contributing factor, assuming that the breakage point was, indeed, at the coil: the wedge-shaped columnals of the coil would likely have had less structural integrity than the rounded columnals of the stem, resulting in the coil representing the weakest point on the stem.

It is worth noting that despite the “starburst” crinoids dis-inclusion from this set of measurements due to the lack of data regarding the calyx and stem, the completeness of the

57 arms alone would place them within Category 1. Were they counted as such, the percentage of Category 1 crinoids, i.e., crinoids killed in the final obrution event, would be roughly 2/3 of the slab specimens.

Results are mixed for Category 2 (I-P, arms 50% or less complete) and Category 3

(Q-X, arms absent distal to physical control). Both categories show similar numbers (44 and 48 specimens, respectively). The abundance distribution, however, is very different.

Both categories are interpreted as consisting of crinoids that died shortly prior to the final obrution event.

The number of occurrences within Category 2 range from 1 to 9 crinoids per grade,

with a mean of 5.5 crinoids. Interestingly, crinoids with complete calyces show a mean

abundance of 6.5 mm, while crinoids with partial calyces show the same at 4.5 mm. In other words, there is not a great disparity in preservation abundances per grade, although there is disparity in the degree of preservation. The relative completeness of the arms suggests that they may have been dead only a few days at most if they were lying exposed on the seafloor, while the calyces are often more damaged than the arm preservation can account for (Fig. 12B, C). It is therefore suggested that scavenging accounts for the partial calyces. This conclusion is based primarily on the fact that calyces would not, under normal circumstances, decay faster than arms, but calyces would be targeted first by scavengers.

Thomka et al. (2011) distinguished between calyx damage from scavenging in the form of plate loss/disruption in the proximal tegmen, and calyx damage due to soft-tissue decay, which often shows the calyx collapsed near the base of the arms due to post-burial decay of the visceral mass (Allison, 1988; Kidwell and Baumiller, 1990). However, it is typical within Category 2 for calyces to be most heavily disrupted well below the base of

58 the arms, and the tegmen is rarely ever visible for study, hidden as it is within the encircling arms (and while the tegmen is visible in many starburst specimens, these fall almost entirely into Category 1 which, as previously stated, show little or no evidence of scavenging, including preferential damage to the tegmen). Both plate disruption at and below the base of the arms are seen on the slab, and distinguishing between the two is impossible in cases of severe dissolution (as often seen in Category 4). However, it is the disparity between good arm preservation and poor calyx preservation that most points to scavenging rather than decay as a primary preservation factor for Category 2.

Category 3, distinguished by arms absent distal to physical control, requires further explanation. Tumbling experiments with Recent crinoids show that unaccustomed high water velocity and/or increased sediment load in the water provokes a defensive response in which the arms compress into a cylinder, what Seilacher called the shaving brush position (Meyer, 1997; Baumiller et al., 2008; Fig. 11). Assuming that glyptocrinid feeding position echoed that of Recent stalked crinoids, the Maysville slab crinoids are in trauma/defensive position, at least to what the limits of their morphology would allow: glyptocrinids lacked control of the arms above a certain point, that point appearing just distal to where the arms become free, roughly a third again the height of the calyx. In laterally-oriented specimens, this is the point at which the arms cease to be compressed and instead splay outward at apparent random, a "modified" shaving brush (Fig. 11). This is interpreted to be the body position in Category 3, as it is in most specimens in all categories on the slab. As the orientational bias is not strong, the modified shaving brush is most likely in response to an influx of sediment rather than greatly increased water

59 velocity. The fact that the great majority of the intact crinoids are in this position is commonly seen where obrution was the killing mechanism (Baumiller et al., 2008).

This position appears to have consequences for preservation. Category 3 ranges from Q to X. Grades Q and S show a sharp spike in occurrence, 10 specimens and 22, respectively. These are the grades with the best preservation overall within this category, and the calyx is complete both times; this is as opposed to grades R, T, V, and X, where it is not, with abundance totals of 1, 1, 1, and 5, respectively. Grade R, between Q and S, and grade T, immediately following them, have partial calyces: the stem is complete (Q) and partial-long (S) in these two grades, whereas it is partial-short in two grades (U, V) and absent in two (W, X). The stem is complete in R and partial-long in T, but, as stated, these are the grades with the partial calyces. Therefore, it seems that the arms were more vulnerable to disarticulation from the point where arm control was lost than proximal to it

(Fig. 12D, E); the trauma position seemed to impart some protection, if not from obrution, then from post-mortem disarticulation (Thomka, 2010). It is interesting to note that the modified shaving brush is just as often the preserved position of the decayed crinoids. This can't, of course, be determined where arms are lacking entirely. But clearly, in specimens where the preservation allows for identification of the shaving brush, even the crinoids that died previous to the obrution event were responding to stress. Yet they were buried and preserved before decay progressed enough for the trauma position to relax again, suggesting deaths a very short time prior to final burial. This could also be an argument for the trauma position in glyptocrinids being ligament-induced or enhanced, rather than only

(limited) muscular, as the slower decay rate of ligaments could have held the trauma positions intact for a day or so longer ("Morphology," this document).

60 The final category (4, Y-FF) consists of specimens lacking arms entirely, or where the arms are greatly reduced (Fig. 12F). These crinoids had apparently been dead the longest, so it not surprising to find little difference in abundance whether the calyx is preserved or not. The highest number of occurrences, for example, is in grade AA, 18 specimens (complete calyx, partial long stem, no arms) and grade BB, 19 specimens (same characteristics, but with a partial calyx). Next-highest number of occurrences is in grade

DD, 12 specimens, again with a partial calyx. Category 4 has the highest abundance (76 specimens) after the final-obrution crinoids of Category 1. Partial calyces within this final category are generally too damaged to determine whether scavenging or decay is the greatest factor in their condition.

Of the three crinoid elements rated for preservation, it seems clear that only one can be considered diagnostic for time spent in the taphonomically active zone. The inconsistencies in preservation of the stem mean that stem completeness cannot be taken as a taphonomic effect, and more likely indicates a morphological characteristic, i.e. points of naturally-occurring weakness along the stem leading to breakage. Calyx condition is indicative of decay in many cases, but in Category 2, discrepancies in preservation between the arm and calyx condition seem to indicate the effects of scavenging; this may also be the case in Category 4, although decay is usually advanced enough in these specimens to make causal identification difficult. Arm preservation is, therefore, the most reliable effect for considering time of death prior to and during obrution.

Preservation suggests that at least two mortality episodes occurred prior to the obrution event. The Category 4 crinoids are the most severely decayed, and must have had more time to reach that condition; there is also the matter of the many pluricolumnals of

61 varying lengths which were not included in this preservation assessment. However, the final stage of preservation is all but lacking, in that there are almost no isolated columnals.

As a stem usually takes close to a week to deteriorate into isolated columnals, the lack of them suggests deaths relatively close to the final obrution time, at which point the decay process was virtually arrested before the final stage of decay could be reached.

Another spike in decay can be seen in Category 3, which is indicative of a mortality episode occurring perhaps only days prior. The actions of scavengers blur results somewhat in Category 2, and finally there is the mass mortality final obrution event, after which scavenging and decay were no longer a strong factor, and the preservation indicates crinoids as they may have appeared in the final moments of life.

FIGURE 11 — Schematic diagram showing hypothesized stalked crinoids in A) parabolic filter fan (feeding) position B) trauma position of crinoid with non-muscular arm articulations (the “modified” shaving brush) C) trauma position of crinoid with well-muscled arm articulation (the shaving brush). Modified after Baumiller, et al., in Echinoderm Paleontology, 2008.

A B

C D

E F

FIGURE 12 — Examples of preservation effects. A) Well-preserved specimen missing stem at distal end. B, C) Specimens showing marks of scavenging on their calyces. D,E) Specimens in which the arms are lost distal to physical control. F) Heavily decayed specimens.

64 Implications of Early Decay

Carcasses are recycled through decay in most environments. The controls on decay are, then, an important factor in fossil and site preservation (Briggs, 2003).

Fossilization is the end-result of a delicate balance between decay and diagenetic mineral formation. In most depositional settings, organic remains will be broken down quickly. Decay inhibition is unusual and usually occurs in unusual conditions, such as peat bogs or amber. The anoxic conditions that can be created by a blanket of sediment may have a dampening effect on decay, but do not stop it. Most decay beneath the sediment/water interface is anaerobic, as a great deal of oxygen is needed for aerobic decay, and the rates of diffusion through tissue and sediment are not high enough to supply the respiring decay organisms (Allison, 1988). Anoxia does, however, limit scavenging and sediment turnover by bioturbators; between this and the lower rate of bacterial activity, decay processes may be slowed enough for diagenetic mineral formation to be enhanced

(Brett and Baird, 1986; Brigg, 2003), leading to excellent preservation.

Trawling and submersible excursions in the study of Recent crinoid assemblages found numerous upright stems, some with, some without crowns, alongside broken, scattered stems in deep-sea habitats (Neumann et al., 1977; Meyer et al., 1978). The crownless stems were attributed largely to predation (Meyer et al., 1978; Messing et al.,

1990), which may have been a factor for the Maysville crinoid community in life, or crown autotomy, which may not have been an option for glyptocrinids. It is possible that upright, crownless stems were recently dead animals, whether victims of predation or not, and were in the decay process, which could leave the stem the last element standing. Llewellyn and Messing's (1993) count of crinoid-ossicle content from the sediment of modern stalked

65 crinoid communities found stem material to dominate the ossicles, comprising 9-52% of the bioclast.

Fossil assemblages are typically made up of two separate fossil groupings: an event assemblage, composed of specimens that were buried in one or more distinct events, and a background assemblage, resulting from assorted hard parts that accumulated over time during normal sedimentation. This second assemblage is characterized primarily by skeletal elements which have evidently spent more time in the taphonomically active zone and are in a greater state of corruption than those of the event assemblage (Brett and

Baird, 1986; Speyer and Brett, 1986; Thomka, 2010). The slab fossils' positions in the matrix and relative to one another indicate they were all buried together, and the decayed crinoids were neither buried alive nor were they dead for very long.

The number of dead crinoids at varying stages of decay is far beyond what has been observed in Recent crinoid colonies (Llewellyn and Messing, 1993; Messing et al., 1990), even allowing for the possible ossicle-trapping effect of community density. Therefore, it is proposed that environmental conditions related to storms, very likely the oncoming final- burial storm event, could have been the killing mechanism for the crinoids that pre- deceased the final obrution. The conditions of Type I obrution put forth by Brett et al.

(2012) include similar patterns of mortality, and suggest the cause to have been changes in water temperature, oxygen levels, or salinity, among other possible storm-related effects

(Thomka, 2010; Taylor and Brett, 1996; Datillo, 2008). The slab cannot, then, be said to possess a true background assemblage. The deaths were close enough together to infer a single killing event, which occurred as more than one punctuated episode. The smothering muds which comprise the fabric of the single lens are apparently from the same source.

66 The entombed crinoids represent a single community of animals which lived contemporaneously and died within as little as a few days of one another. The fact that the crinoids didn't all die at once but were buried together is further evidence of autochthony as well (Lane, 1963).

67 Orientation

Materials and methods

Because the original orientation of the slab is not known, an arbitrary position was chosen, designated as "north," and used consistently for all orientation measurements. These measurements were taken on a 360 axis by superimposing a transparent compass over photographs of slab pieces in Photoshop (Fig. 13). As the largest individual piece, M-slab was designated the base to which all other pieces were fitted: only individual pieces that unquestionably fit very snugly and precisely to M-slab were included in the orientation dataset. Therefore, orientation was plotted on a rose diagram for slab pieces M, A, B, C, D, H,

I, J, and K as a single unit. The unit was then divided into 17.5 cm squares on an archeological-type grid in the manner of Thomka (2010), and orientation was replotted for the most densely-fossiliferous combinations of the 37 resulting squares: squares 9, 10, 17, and 18 were designated Section A; squares 11, 12, 19, and 20 were designated Section B; and squares 13, 14, 21, and 22 were designated Section C. The unit was then redivided into areas labeled North, South, West, and East, according to the artificial orientational data point previously established: orientation was then plotted for squares 1-23 as North; 24-37 as South; 1-3, 8-11, 16-19, 24-27, and 31-37 as West; and 4-7, 12-15, 20-23, and 28-30 as

East. There is overlap between the four areas of this last category.

Crowns were included only when exposed enough to allow an accurate reading, and attached stems were only included in the measurements for the most proximal two centimeters. This was to avoid the obvious data distortions possible when encountering a crown with an encurved stem. However, as the crown was initially considered the element most likely “caught” by currents, due to its adaptive features specific to that purpose,

68 measurement was based on its orientation, with the compass directed toward the arms.

Stems without crowns but with coils were measured with the compass directed away from the coil, toward the missing crown. If curved, the stem was measured for its average trend, excluding the coil itself. Only stems four centimeters or longer were included.

Stems without crowns or coils were likewise measured; however, without a coil or crown as indicators, their directionality is necessarily random. Averages were taken for curved stems. Again, only stems four centimeters or longer were considered.

As previously stated, calyx position within the matrix was considered for lateral and starburst positions only. The few slightly oblique specimens were classified as lateral.

A total of 397 elements comprise the orientation dataset. This is a higher number than the 253 studied for preservation due to the inclusion of stems and calyces which were exposed enough to plot the orientation but not enough to assess the preservation.

FIGURE 13 — Grid system from which orientation was plotted for slab pieces M, A, B, C, D, H, I, J, and K as a single unit.

70 Orientation: results and discussion

Casual observation of the Maysville slab reveals little preferred orientation to the crinoid remains, as Figure 14 illustrates. Yet some degree of alignment exists, particularly in

"patches" of preferred orientation seen where a few elements in close proximity exhibit a similar trend. This indicates some current action, if minimal, and implies small eddying currents which affected the final resting positions of the crinoids.

It is generally acknowledged that paleocurrents are difficult to reconstruct from fossil crinoid orientation alone (Brett and Baird, 1986; Thomka, 2010). Different crinoid elements, such as crowns or stems, frequently behave differently in the same currents, although the specifics of that behavior and its interpretation are in some dispute;

Schwarzacher (1963), for example, asserted that crinoid stems will roll perpendicular to current flow for a period of time before turning into parallel alignment with the flow.

Moreover, crinoid stems may be deeply curved, making orientation difficult to ascertain and inspiring conflicting ideas: Wetzel and Meyer (2006) interpreted curved stems as having been dragged across the seafloor by strong currents, while Baumiller and Ausich

(1996) saw them as normal stem posture preserved by weak currents, arguing that strong unidirectional currents would result in an alignment of straight, parallel stems.

Strong unidirectional currents are not, however, a general attribute of distal environments, and so the absence of any evidence for them is not surprising in the

Maysville slab. Although it is typical for lithologic structures resulting from storm events to register ambiguously in such environments (Kreisa, 1981; Brett an Baird, 1986; Brandt,

1989; Wheatcroft, 1990; Brett and Seilacher, 1991), patterns did emerge when orientation of the fossils was examined more closely.

FIGURE 14 — Detail of M-slab.

72 Long-axis orientation measurements were first taken for all calyces and stems on

all nine of the close-fitting slab pieces, provided the elements fit the criteria presented in the previous chapter. The combined data for all elements shows a decided trend in an approximately north-south direction, with maxima at 350 and 190 degrees (Fig 15C). As earlier studies have found discrepancies between calyx and stem orientation, both elements were also plotted separately (Fig. 15A, B). Yet there appears to be almost no difference between the orientation of the two elements on the Maysville slab, with the very slight exception that more calyces seem to be directed northward. The greater southward trend seemingly exhibited by the stems alone (Fig. 15B) is, however, largely nullified by the fact that stem direction was random, unlike calyces (measured toward the arms, in opening direction) or stems with coils (measurement directed away from the coil). Lacking a calyx or coil meant that a stem was measured with the compass arbitrarily directed toward either end, meaning that a value of, for example, 0 degrees could just as well have been 180 degrees. When this is taken into consideration, it becomes clear that stem orientation is not significantly different from calyx, and so both were combined for all further measurements.

Thirty-seven squares comprise the grid system used for the next measurements, none of which alone contain enough material for a meaningful dataset. Groupings of squares were instead plotted, firstly in sets of four, chosen for their density of crinoid material, designated Sections A, B, and C. Section A (59 specimens; Fig. 16A) shows considerable variation in orientation to the north, east and southerly directions but very little toward the west. Section B (40 specimens; Fig. 16B) by contrast is primarily oriented east, west, and south, but very little is directed to the north. Section C (37 specimens; Fig.

16C) is directed largely north-west and south, with little trending east. The most significant

73 aspect of the section measurements seems to be not where the crinoids are oriented so much as where they are not; in each of the three sections, there is a direction where the crinoids are not, for the most part, trending, with much variation in orientation in the areas where they are.

Similar patterns are seen in the next set of measurements, conducted on groupings of squares corresponding to north, south, east, and west on the slab. These are larger areas than those defined as sections in the last set of measurements, and contain considerably more fossils. "North" is made up of 23 squares containing 242 elements (Fig. 17A). The results are very similar to what is seen for all elements on the entire slab: a trend north and southward with, for "North," maxima at 0 and 190 degrees. "West" shows similar results

(Fig. 17D), with two maxima northward at 340 and 10 degrees, and two southward at 190 and 160 degrees. It also has a large number of specimens, with 275 over 23 squares.

"South," on the other hand, shows a definite trend predominantly southward (Fig. 17B) although it also has the lowest number of specimens represented, with 154 elements spread over 14 squares. Its southerly maximum is 160 degrees, with roughly 12 specimens, while its northerly maximum, at 350 degrees, contains 7 specimens. "East," (Fig. 17C) interestingly, shows a pattern that suggests a mirror image to that of Section A, the most westerly of the three sections. The 122 specimens over 15 squares that comprise "East" are scattered to the north, west, and south with little orientation to the east, while those of

Section A show a similar pattern to every direction but west.

The slab crinoids, then, do show a general trend toward the north but also toward the south, indicating that currents were active though not very strong and that neither crowns nor stems were preferentially caught and oriented by the current (Fig. 18).

74 A. Calyces B. Stems

C. Calyces and stems

Calyces and stems

All

FIGURE 15 — Orientation from slab pieces M, A, B, C, D, H, I, J, and K as a single unit. Total count: 397 elements. A) Laterally-oriented calyces in opening direction. B) Stems. Those without a coil are arbitrarily directed. C) Combined A and B.

75 A .

No. = 59 B

No. = 40 C

No. = 37

FIGURE 16 — Orientation of calyces and stems for three defined sections, as highlighted on page right. Number of specimens in each dataset listed on rose diagram, lower right. A) Section 1 B) Section 2 (C) Section 3.

76 North

South

East

West

FIGURE 17 — Orientation of fossil material for four defined sections on the slab, as highlighted on page right. Number of specimens in each dataset: North) 242. South) 154. East) 122. West) 275. Note that sections overlap.

77 Furthermore, "patches" of preferred orientation as represented by the grid squares imply oscillatory currents, which are a common feature of shelves or rises in distal environments during storms (Fagerstrom, 1964; Ewing, 1973; Butman, et al., 1979; Kreisa, 1981; Wright,

2001; Webber, et al., 2008; Thomka, 2010).

Even as the sediments were shed over the crinoids, there may have been vestiges of a storm overhead. Storm energy is often transferred downward through the water column and induces oscillatory currents to depths as much as 200 meters (Butman et al., 1979;

Taylor and Brett, 1996). These currents can themselves be powerful agents of sediment suspension and resettling, because even very weak oscillatory currents are many times more effective than unidirectional currents of the same magnitude (Ewing, 1973; Kreisa,

1981). Such currents and turbulence naturally decrease with depth, but they needn't be very strong to induce the slight reorientation patterns seen in the slab crinoids.

Evidence of oscillatory currents also comes in the brachiopod valves, which are almost entirely preserved in concave-up position (Fig. 20C-E). Ordinary current action would have positioned the valves into hydrodynamically-stable concave-down position, and the fact that they were buried concave-up highlights the speed of the depositional event. Clearly the valves were buried immediately prior to deposition, or the muds were sufficiently stirred during or immediately after obrution to cause repositioning of the valves. A Rafinesquina valve includes encrusting inarticulate Orbiculoidea on its underside, a position that would have been smothering in life. (Brett and Baird, 1986; Taylor and

Brett, 1996).

Several of the slab crinoids were buried and preserved in full-fan feeding position, the starburst configuration. As the preservation of many of these starburst crinoids is

FIGURE 18 — These crowns are relatively consistent for orientation, though not direction.

79 among some of the best on the slab, they were likely still alive when the final burial event occurred. Preservation of this type is indicative of rapid burial, deposition occurring at a faster rate than the crinoids' reaction time ("Morphology," this document). Yet while starbursts often appear in close association with one another (Fig. 10B), they are also alongside crinoids that did have time to react by folding themselves into defensive position, the modified shaving brush. Furthermore, the two positions are generally on the same level within the bedding plane, implying that they were both buried at the same time (Fig. 19). A possible explanation could be that oscillating currents affected some crinoids and not others, perhaps disturbing one crinoid enough to induce trauma position while sparing its near neighbor, which maintained feeding position. Conversely, some starburst crinoids may have been caught in localized oscillating currents which spared their near neighbors, allowing their neighbors the extra moments to achieve defensive position. What is clear is that the conditions that created the Maysville Lagerstätte were complex and involved more than a simple rain of fine sediment with mild unidirectional flow.

FIGURE 19 — Two crinoids, "A" in starburst, "B" in modified shaving brush, preserved at the same level on the slab.

81 Incidental Elements

The generally excellent preservation of the crinoids is not mirrored in the preservation of many of the other elements found within the deposit. An admixture of mostly poorly preserved and fragmentary fossil material is jumbled at apparent random amongst the crinoids, and it seems unlikely that most of these non-crinoidal elements were resident to the crinoid colony and/or living in association with them.

Degree of corrasion is related largely to time of exposure, and most of the incidental elements display the results of varying lengths of time in the taphonomically active zone, although this did not necessarily occur within the crinoids' habitat. Storm events that winnow sediments from proximal areas and transport them into deeper water horizons can do the same for other, heavier, material such as shells (Datillo et al., 2008). This is not to say that the final obrution event carried all of these other elements in; rather, it seems more likely that they arrived earlier. A single crinoid's distal coil wrapped around a broken chunk of bryozoan supports this: glyptocrinid columns were not prehensile, and instead the distal end of the stem would develop its encircling coil upon contact with an available hard object, growing its anchor much as does a climbing vine ("Morphology," this document). Therefore, the bryozoan with an attached crinoid was almost certainly put to service as a holdfast anchor after its death, fragmentation, and transportation into the crinoid community. The rounded and tumbled condition of the bryozoan is not mirrored in the articulated condition of the crinoid. This would seem to indicate that at least some of the elements had been in the crinoids' habitat for some time.

The poor condition of most of the fossil debris strongly suggests it to have been reworked prior to its deposition amongst the crinoids and likewise prior to the final burial

82 event. Bryozoans, for example, are consistently well-tumbled and fragmentary. With few exceptions, most brachiopods are also in poor condition. The non-crinoidal elements may have been earlier weakened by boring or other scavenging (Gluchowski, 2005), and the currents which deposited them among the crinoids were sufficient to further damage them, as is common (Brett and Baird, 1986; Taylor and Brett, 1996). It is, however, fairly certain that the violence and/or time of exposure they had undergone would have left the crinoids in far worse condition, had the crinoids been subjected to it.

Incidental faunal elements include moult remains of the trilobite Isotelus (Fig. 20A) consisting of two isolated librigenae or free cheeks, which often separate from the cephalon along the facial sutures when trilobites moult or die (Speyer and Brett, 1986; Gon, 2008;

Brenda Hunda, pers. consult.). Five types of trepostome bryozoans (Fig. 21), all fragments and showing evidence of much tumbling (well-rounded corners, etc.), are scattered throughout the deposit: these include Atactoporella schucherti, Batostoma implicatum (or possibly Batostoma prosseri ), Batostomella gracilis, Parvohallopora ramosa, and

Parvohallopora subplana (Roger Cuffey, pers. consult.).

There is little evidence of interspecies interactions in the fossil remains. A possible exception may be two occurrences of Cyclonema bilix lata (Fig. 20B), a probably coprophagous gastropod often found attached to the tegmen of crinoids, including

Glyptocrinus (Ausich, 1999; Meyer and Davis, 2009; Steve Felton, pers. consult.). These

Cyclonema are not in positions that associate them with any particular crinoid, and it is impossible to say if they were alive or dead when the obrution event occurred. It may be noted, however, that they are in a better state of preservation than most other incidental elements, and show little evidence of corrasion. As their preservation mirrors that of some

83 of the more intact crinoid fossils, it could be suggested that they were alive within the crinoid colony prior to final burial.

There are multiple occurrences of four brachiopod genera: Hebertella, Dalmanella

(Onniella), and Rafinesquina with encrusted inarticulate Orbiculoidea (Brenda Hunda, pers. consult.; Fig. 20C-E). Calcite crusts of Rhodophyta or red algae appear twice (Fig. 22B). Only the mollusk Ambonychia robusta, preserved as an articulated internal mold, is probably preserved in its semi-infaunal life position within the crinoid colony (Fig. 22A).

Thin sectioning of the mudstones revealed four Skolithos-type burrows, all between

1.5 and 3 cm, in close proximity to degraded crinoid elements (Fig. 22C). In general, the

Droser-Bottjer ichnofabric index, which grades the amount of trace fossil activity on a 1-6 scale, would put the Maysville slab at "2," the next-to-lowest grading, representing

"discrete, isolated trace fossils (with) up to 10% of the original bedding disturbed" (Droser and Bottjer, 1986). As previously stated, little or no scavenging is apparent in the final obrution (Category 1) specimens.

All the preceding epibenthic and infaunal forms have been previously reported from mid-Cincinnatian horizons (Ausich, 1992; Meyer and Davis, 2009; Roger Cuffey, pers. consult.), and are associated either with the deep subtidal environment of the crinoid community or the shallow subtidal up-slope of the site. The single exception is the monoplacaphoran Cyrtolites minor (Fig. 20F), which occurs twice. Cyrtolites is known from the Kope to the Whitewater formations, but C. minor has not previously been reported in the Fairview; while its mode of life is still under investigation, it is usually associated with offshore environments, adjacent to but deeper than that of the Maysville Lagerstätte (Dzik,

1980; Steve Felton, pers. consult.).

A B

C D

E F

FIGURE 20 — Other elements from the slab. Note that as slab is prepared from underneath, these are the undersides of the specimens as preserved. A) moult remains of Isotelus. B) Cyclonema bilix lata. C) Dalmanella (Onniella). D) Hebertella. E) Rafinesquina with encrusted inarticulate Orbiculoidea. F) Cyrtolites minor.

85 FIGURE 21 — Trepostome bryozoans from the slab. Atactoporella schucherti, Batostoma implicatum (or possibly Batostoma prosseri ), Batostomella gracilis, Parvohallopora ramosa, and Parvohallopora subplana

86 A B

Skolithos-type burrow C crinoid remains

FIGURE 22 — Other elements from the slab. A) Ambonychia robusta. B) Rhodophyta. C) thin section revealing Skolithos-type burrow in vicinity of crinoid remains.

87 Geochemistry

X-ray fluorescent spectrometry (XRF) scans for major elements reveal the slab to be comprised of a fairly homogenous mixed siliciclastic/lime mud, which is particularly iron- rich. Scanned thin sections containing poorly-preserved crinoid elements show the presence of phosphorous "halos" around some crinoid remains, consistent with what can occur when decay products leak from a carcass into the surrounding matrix (Martin et al.,

1985; Allison, 1988; Lin et al., 2008; Barry Maynard, pers. consult.; Fig. 23). Datillo et al.

(2013) found similar phosphorous deposits around Cincinnatian crinoids to decrease in concentration away from the lumen, likely because indigenous microorganisms were concentrated in and around that area; microbial processes are known to strongly influence the way phosphorous concentrates and is distributed (Orr, 2000; Brasier, 2010). The presence of these halos also supports the argument for anoxic conditions, as it is anaerobic decomposition which liberates these dissolved phosphates from the organic matter, and in cases of more decay and/or higher phosphorous concentrations, can lead to the creation of phosphatic nodules or phosphatization as a primary mode of preservation, although neither occur in this case (Brett and Baird, 1986).

The scans also reveal sulfur nodules scattered about the matrix. These nodules were neither seen nor felt during preparation with hand tools, a process that makes differences in texture and the consistency of the matrix obvious. Because the sulfur concentrations are only revealed by the scans, they are thought to be nodules, the result of replacement, rather than concretions, which form by mineral precipitation around a nucleus and are generally harder/denser than the surrounding matrix. Elemental sulfur deposition is associated with the activity of sulfate-reducing bacteria (Maynard, 1980; Sageman et al., 1991; Zierenberg,

88 1994), and experimental results obtained by Vasconceles et al. (1994) on sulfur nodules in deep-water mudstones confirmed post-depositional microbial involvement in the formation of such nodules.

The sediment-water interface is where a mixture of solid sediment and interstitial water meet and are separated from an overlying body of water. Concentrations of manganese at the deposit’s upper surface are consistent with evidence for this interface.

The decay of organic matter beneath the sediment surface decreases the redox potential of oxidized manganese, causing it to destabilize. The resulting reductive dissolution increases the porewater concentrations of the manganese and associated elements, such as iron. If there is a high enough concentration gradient, the ions will diffuse upward towards the surface (Fig. 24). The manganese revealed at the deposit's upper surface by the XRF is, then, a good indicator of sediment stopping (Lerman, 1978; Sundby et al., 1986;

Pakhomova et al., 2007; Higgins and Schrag, 2010; Barry Maynard, pers. consult.).

Most of the fossil crinoids preserved gray, similar to the mudstones, but several are unusually dark (Fig. 25). These specimens do not seem to be in unique positions regarding the other fossils or within the matrix, nor is their skeletal preservation remarkable. There are several possible explanations.

Iron oxides and other inorganic pigments are a common cause of coloration in fossils, which makes this a plausible explanation, considering the iron-rich mudstones of the slab. The dark coloration may also be the remnants of organic material extruded from the fossil elements during decomposition, similar to what is seen as "smudges" in some elements of the Burgess Shale (Whittington, 1971; Lin et al., 2007). Similarly, the color could be the carbon residue of decaying plant matter ("The Holdfast Forest," this

89 FIGURE 23 — XRF scan revealing sulfur, calcium, iron, and phosphorous as major elements of the slab. Bottom center is a crinoid fossil.

90 document). A third explanation entertains the idea that the unusual coloration was the crinoids' own. The range of crinoid fossil coloration across the Paleozoic, often in abutting specimens within a single bed, has long been a puzzle (Blumer, 1965; Ausich et al., 1999a;

O'Malley et al., 2008). Recent work by O'Malley et al. (2009, 2013) points to the unique crinoid morphology to explain why the animals' original coloration may be preserved: crinoid ossicles are each a single calcite crystal, with roughly half the plate volume being calcite and half porosities filled with soft tissue. Some of the earliest diagenetic processes after death involve the occlusion of those porosities by precipitated calcite cement. Organic molecules containing pigments thus may be trapped inside calcite crystals or what have been called "crystal caskets" (Dickson, 2001), geochemically stable structures resistant to leaching and other diagenetic processes. While there is evidence that original taxon- specific pigments may be preserved in this way, it is still unclear if this process could occur selectively within a single bed containing a single taxon.

FIGURE 24  Extracted spectra from XRF, with enlargement for manganese (Mn). Note Mn concentration at upper surface.

FIGURE 25 — Crinoid stem fragment with dark pigmentation.

93 Mechanism for Obrution

The crinoid fossils exhibit some current alignment, although it is weak. Their positions and orientation are, for the most part, reminiscent of silk scarves dropped in a mild breeze; their settling positions, particularly the random splaying of the arms, suggest that the smothering sediment came primarily as a rain of fines. This is consistent with the work of a lofting turbidity current, which can have several modes of generation.

Turbidity currents — turbulent, fast-moving, sediment-laden currents propelled downslope by gravity — are often initiated in relatively shallow-water shelf settings by some catastrophic event and move down-gradient and out across the deeper-water horizons (Brett and Seilacher, 1991; Wright et al., 2001; Potter et al., 2005; Stanley, 2009).

Their genesis may be in what Gladstone and Pritchard (2010) called a quasi-steady flow, such as that created by river discharge, or it can be surge-like, or generated rapidly, as by a storm event. These are typically described as "auto-suspending flows in that the turbulence generated by the flow itself is sufficient to maintain the sediment in suspension"

(Traykovski, 2000). Once mobilized downslope, the suspended sediment serves as a horizontal pressure gradient, and the current continues to flow, driven by the sediment which renders it of higher density than the surrounding fluid (Middleton, 1993; Potter et al., 2005; Gladstone and Pritchard, 2010). Eventually the slope decreases, the turbulent energy wanes, and the turbidity current begins to shed sediments, beginning with the heaviest and coarsest. Sands, then silts, then clays are lost in succession over what can be a very wide area: using Stokes Law, Potter et al. (2005) calculated that a grain of fine sand

100 microns in diameter would fall 140 times faster than an 8-micron silt grain and 6,600 times faster than a particle 1 micron in diameter (the mudstone particles of the Lagerstätte

94 would be between 3.90625 and 6.25 microns). Such fall velocities translate to long lateral transit times, and thus the great lateral continuity of muds in general. These characteristics are, however, greatly affected by the density of both the flow itself and the ambient fluid into which it flows (Wright et al., 2001).

The interstitial fluid within which the sediment is suspended is generally of lower density than the ambient fluid, due to the normal stratification of the ocean and the interstitial fluid's origin in shallower, hence warmer, fresher, lower-density, water. The ocean is stratified along both thermal and salinity gradients, although the thermal gradients have a more powerful effect on fluid density (Traykovski, 2000; Wright et al.,

2001; Gladstone and Pritchard, 2010). This is true even where the temperature change is slight: the surface of the modern Atlantic Ocean, for example, has an average density of

~1024 kg per m3. The density increases to about 1028 kg per m3 at depths greater than

4000 meters, which is an increase of less than 0.5% but is still significant enough to have a profound effect on fundamental oceanic processes, such as vertical mixing and circulation.

The areas where the density gradient is highest are pycnoclines, thin but effective barriers between the turbulent mixed waters above and the calmer, colder waters below, and which strongly influence the vertical profile of the ocean in regards to the movement of heat, salt, and nutrients. Permanent pycnoclines at about 500 and 1000 meter depths exist in modern ocean basins, but seasonal ones can form in waters as shallow as 20 meters or less (Franks and Franks, 2009). It can reasonably be surmised that similar conditions existed during the

Paleozoic, whether or not ocean densities were different from those in modern seas.

What becomes of the suspended sediment in the flow when it reaches a change in the density gradient depends on the flow's initial bulk density (Lowe, 1982; Mulder et al.,

95 2003). A hypopycnal flow is one in which the turbidity current is less dense than the water it is flowing into, due to the relative lightness of the interstitial fluid and a low and/or exceedingly fine sediment load; the result would be for the current to spread out, plume- like, over the ocean surface and subsequently shed its fine sediments near the shoreline.

Another possibility is a hyperpycnal flow, wherein the flow is denser than the water it flows into by virtue of the sediment it carries; in this case the composition of the flow, as regards the relative bulk densities of the sediment and the interstitial fluid, determine what next occurs. If the interstitial fluid is similar in density to the surrounding water, or if it becomes so through mixing in the early life of the current, the heavier sediments will prevail and the flow will become negatively buoyant, sinking to the ocean floor and becoming a typical "ground-hugging" turbidity current. Another scenario exists wherein the interstitial fluid is still less dense than the surrounding water even upon reaching the ocean floor. As viscous effects decrease the energy of the flow, the heavier sediments settle out first; bulk flow density decreases until the flow reverses in buoyancy, with some sediments still in suspension. What remains of the flow, that is, the light water and the finer sediment, is now lighter than the ambient water and will loft upwards and spread out as a plume or cloud (Fig. 26). The sediment-laden cloud will rise to a depth of neutral buoyancy: depending on the composition of the cloud, this could be the ocean surface in shallower water or a pycnocline at intermediate or greater depth. It has been suggested that this is a more common phenomenon than the fossil record would seem to indicate, and is even the most logical explanation for deposition from the distal end of a turbidity current

(Stanley, 1983; Stow and Wetzel, 1990; Mulder et al., 2003; Potter et al., 2005; Franks and

Franks, 2009).

96 Lofting, then, effectively separates heavier, coarser sediments from finer ones. Still,

Gladstone and Pritchard (2010) found in laboratory modeling experiments that such lofting turbidity currents could contain between a tenth and a quarter of their initial sediment load after lofting. This would suggest that significant amounts of very fine sediment could be transported in this manner. When a turbidity current loses enough of its sediment to reverse buoyancy, the flow decelerates rapidly. The ensuing turbulence causes massive and sudden dumping of the last of the coarsest sediments. As the cloud lofts vertically, it remains stationary horizontally, and increasingly finer sediments rain out of it as it accelerates upward to the point of neutral buoyancy. Once the cloud reaches the surface barrier, be it the ocean surface or a pycnocline, it intrudes. Finally, the cloud of the finest sediments spreads horizontally along the cline and will be carried onwards by the current, resulting in a lateral flow without great speed or force. It travels to the point where its energy is depleted and/or it encounters increased friction, and it fails.

Where the last and finest of sediments from the cloud rain down is a fall-out deposit that may be called a hemiturbidite, after Stow and Wetzel (1990), who first described such a phenomenon as a fine, muddy deposit of a slightly different origin: in their estimation, the distal end of a ground-hugging turbidity current would loft upwards to neutral density and then discharge the last of its remaining suspension into the water column where it would float as a suspension cloud hundreds or thousand of meters above the ocean floor. The suspended matter would subsequently settle to the ocean floor over a long period of time, long enough to create ample opportunities for bioturbators to penetrate throughout the deposit, as they observed.

The Maysville deposit was neither laid down in the offshore environment Stow and

Crinoid communit y on rise

FIGURE 26  Schematic of lofting turbidity current into distal environment encountering the crinoid community on the rise. Not to scale. Modified from Potter, et al. in Mud and Mudstones: Introduction and Overview, 2005, and Wetzel and Meyer, 2006.

98 Wetzel described, nor is it heavily bioturbated. It does, however, share some characteristics with their hemiturbidite, which is described as "partly turbiditic and partly hemipelagic" in character: the Maysville slab has the sharp, irregular base of a turbidite deposit, but unlike such a deposit, such structures as erosive scours which might indicate moving water are not seen, nor does it grade upward (Fagerstrom, 1964); rather, it shows the unstructured, unlaminated fabric of a hemipelagic deposit. It is composed of the finest sediments of dominantly turbiditic material, indicating that it is a product of the last material deposited from a turbidity current. But the positions of the crinoids suggest the sediments settled gently, without strong directional flow; likewise, a hemiturbidite is deposited from a stationary or slow-moving suspension cloud, the product a dying turbidity current. Yet unlike a "classic" hemiturbidite, the Maysville Lagerstätte shows evidence for rapid, short- term deposition. The sediment of the Maysville slab, therefore, while not possessing all the defining characteristics of a hemiturbidite, does share enough to put it in a similar category: the product of a rapidly-deposited yet soft rain of very fine sediment with low- velocity horizontal trend, as has been observed in Recent settings (Traykovski et al., 2000;

Wright et al., 2001; Wetzel and Meyer, 2006).

99 Sediment Capture

The plant matter and its resident crinoids formed a dense aggregation which may have had a baffling-and-binding effect on the slow-flowing sediments that ultimately formed the

Lagerstätte. To reiterate, a sediment plume of fines extended laterally along a pycnocline, intruding into distal regions and the crinoids' habitat on the rise. It is possible that the subsequent decelerating flow velocity coupled with the additional friction of the elevated relief of the crinoid community reduced the flow competence enough to result in sediment deposition. This suggests a feedback process, a hypothesis supported by the tapering off of both mudstones and crinoids at what appears to be the outer edge of the deposit, implying that deposition did not occur outside the colony. Friction would have been lower outside the crinoid habitat, and sediments not trapped by the community may have been carried away by residual storm currents (Wetzel and Meyer, 2006). The waning storm overhead could have created low-energy orbital currents within the water column, stirring small pockets of the freshly-deposited light muds and entombing the crinoids into their current orientation patterns. The obrution both killed the crinoids by asphyxiation and preserved them.

The seeming contradiction between "low-energy" and "rapid" deposition is seen in the preservation. The crinoids gently-draped positions suggest low-energy deposition, while those in starburst, pressed to the substrate while in feeding position, suggest rapid.

Also, while deposition was swift enough to trap many of the crinoids in life position, it was not so fast nor laid down so thickly as to prevent the escape of any other mobile, benthic fauna which may have been at the site. It is also, however, possible that the sheer density of crinoids and their anchor-sites precluded much or any other fauna from living within

100 the crinoid habitat, and that the assemblage was always essentially monospecific.

Finally, the mudstones are consistent for grain size and color and were certainly winnowed from the same proximal site of origin, but it's not impossible that at least some of the muds were there prior to final obrution. Muds from the same source may have washed in earlier and the dense crinoid community would have trapped them, too; many of the crinoids would have been elevated just far enough above the substrate to survive. The oscillatory currents which slightly stirred the settling sediments could have mixed the newly arrived and recently-arrived standing muds, eliminating internal lamination. It's very likely that the event which killed some crinoids prior to obrution and the final obrution which buried living crinoids was generated by the same massive and lengthy storm.

101 Paleoautecology

Paleoecology, the reconstruction of an ancient ecosystem, involves first extracting data from the fossils. There are numerous taphonomic processes that must be considered when interpreting fossil remains. The information derived from such considerations includes the reconstruction of the paleoenvironment and therefore, by extension, the fossilized animal's relationship to it in life. Paleoecologists, therefore, must be taphonomists first, as post- mortem history is all that remains for the reconstruction and understanding of a long-dead animal. Crinoids lend themselves especially well to this pursuit, retaining as they do evidence of their feeding, attachment, and other behaviors on their exoskeletons

(Lawrence, 1968; Brett et al., 2008).

Yet the use of modern analogs in determining the paleoecology of a Paleozoic community has its limitations. Paleozoic shallow-water communities were actually very different from modern ones. Whereas modern soft-substrate shallow-water communities are largely made up of infaunal suspension-feeding mollusks and annelid worms, the environmental counterpart in the Paleozoic was dominated by epifaunal suspension feeders such as bryozoans, brachiopods, and crinoids. There are no modern shallow-water settings that consist solely of crinoids, stalked or otherwise (Ausich, 1980).

Interpreting the paleoautecology of a site such as the Maysville can nevertheless be very enlightening as to the animals preserved there, provided the site is considered on its own terms. These terms are based on established models for identifying types of fossil aggregations, behaviors and other strategies, and, finally, the cautious use of modern analogs in order to obtain as complete a picture as possible of life within the crinoid community.

102

B C

I J D

E F

H G

FIGURE 27 — Construction of G. decadactylus A) Free arms B) Tegmen (hidden within arms) C) Fixed arms (brachials) D) Secundibrachs E) Radials (in black) F) Basals G) Distal coil H) Stem I) Interrradials (in blue) J) Calyx K) Crown (calyx + arms). Modified from Hess, H., Ausich W.I., Brett, C.E., and Simms, M.J. in Fossil Crinoids, 1999.

103 Morphology

Implications for paleobathymetry

The morphology of Glyptocrinus decadactylus was adapted to its particular mode of life with respect to the depth of the waters it inhabited, its feeding habits, and its survival strategies — in short, the morphology is the first key to reconstructing the life habits of the animal.

Glyptocrinus was one of the more robust crinoids of the Cincinnatian, due largely to its rigidly articulated theca (consisting of the calyx and the tegmen) incorporating fixed brachials, interradials, and the plates of the calyx (Fig. 27). The plates are highly ornamented with angular ridges radiating from the center of each, bracing the theca further. The heavy, rigid tegmen is likewise strongly cemented to the calyx (Nichols, 1967;

Simms, 1999; Ausich 1980, 2002; Meyer et al., 2002). This is a distinguishing feature of the subclass Camerata, a subclass of Crinoidea that also includes Cladida, Disparida, and

Flexibilia. There are both dicyclic and monocyclic groups within Camerata (which also appear in other subclasses), distinguished by the occurrence or otherwise, respectively, of infrabasal plates along with the radial and basal plates common to both groups. Of the two prevalent orders within Camerata, Monobathrida and Diplobathrida, it is Monobathrida —

Camerata without infrabasals — that appears the more common (McFarlan, 1931; Lane,

1963; Ausich et al., 1999; Ausich, 2002). This grouping includes Glyptocrinus.

Water velocity is an important factor in the distribution of suspension feeders

(Meyer, 1973; Messing et al., 1990). Larger, more robust crinoid forms are associated with higher-energy settings, typically shallower and more wave-swept. As previously stated, G. decadactylus occurs in facies interpreted as having been deposited between fair-weather

104 and storm base in a frequently stormy region, and so was capable of withstanding fairly high-energy currents. Its comparative toughness is made apparent by its preservation: monobathrid camerate crinoids are found to be the most resistant to disarticulation in comparison to other crinoid morphotypes, flexibles in particular (Meyer, 1971; Meyer et al

1989; Brett and Seilacher, 1991; Ausich and Sevastopulo 1994; Brett et al., 1997; Ausich,

1998, 2001; Gahn and Baumiller, 2004).

"High-energy" current is, of course, a relative term in regards to crinoid habitats.

Paleozoic crinoids are generally interpreted as low-energy rheophiles, analogous to modern stalked crinoids (Nichols, 1962; Hess et al., 1999). These support themselves above the substrate by means of an anchored stem and arrange their arms in parabolic filtration fan position perpendicular to the current, with the arms deflected into the current and the oral surface facing downcurrent. Suspended food particles are captured by the fan, secured by mucus, and passed by tube feet along the ambulacral grooves to the mouth (Nichols,

1962; Ausich, 1980, 1996; Hess et al., 1999). While the feeding behavior of G. decadactylus was likely very similar to that of its modern stalked counterparts, morphology of the feeding structures further indicates its higher-energy habits.

Aerosol filtration theory states that crinoids with densely-branched filtration fans capture food particles most efficiently at higher-velocity current flows, and lower-density branched fans are most efficient in slower currents (Meyer, 1979; Kammer and Ausich,

1987; Baumiller, 1993; Brower, 1994; Holterhoff, 1997; Meyer et al., 2007). There is a minimum and maximum velocity at which efficiency falls off again, however, and therefore every type of filtration fan has a water velocity range at which it captures food particles most effectively. Furthermore, the particles, once caught, must be moved along ambulacral

105 grooves which themselves vary in width. The width of the ambulacral groove limits the size of the food particle that can be ingested by the crinoid (Ausich, 1980; Brower, 2004, 2006): consequently, between the density of the filtration fan and the ambulacral groove width, the feeding niche of a crinoid might be thought fairly established (but see "The

Community," this document). In general, low-density fans and wider ambulacral grooves characterize deep-water crinoids, where slower, gentler currents prevail; the wider grooves allow them to consume a wider range of food particle sizes in an environment where food may be scarcer. By contrast, crinoids with high-density fans, such as those created by glyptocrinids' 20 well-pinnulated arms, are generally found to have narrower ambulacral grooves, particularly when compared to disparids (Ausich, 1980; Brower, 1994,

2006; Meyer et al, 2002). Narrower food grooves serve to narrow glyptocrinids' food options by particle size, although this would be less of an issue in a higher-energy environment where not only is a wider range of food passing through the crinoids' habitat, but flow rates are powerful enough to drive the food-bearing fluid through the crinoids' dense fans (Brett et al., 2008).

Baumiller (1993) hypothesized that camerate crinoids would have been specialized to environments with current velocities greater than 4 to 8.5 cm/second, whereas most cladids, disparids, and flexibles would have done better at lower water velocities, between

1.5 to 3 cm/second. Studies of Recent crinoids yield similar results regarding morphotype, with crinoids bearing closely-spaced pinnules and tube feet favoring more exposed and elevated feeding positions than those with more widely-spaced tube feet, even when both types occur at similar depths (Meyer, 1979). Nichols (1962) reiterates Clark's (1915) findings that the number of arms in Recent crinoids correspond to depth but also to

106 temperature, with "shallow warm" species averaging 40 arms, "deep cold" averaging 10 arms, and "intermediate moderate" between 20 and 30 arms.

Meyer et al. (2002), in surveying six Paleozoic species from the Cincinnati Arch region, further found smaller, more delicate crinoid morphotypes typically occurring in deeper, calmer waters, while larger, more robust crinoids were associated with shallower, more energetic environments. Stem length, for example, was positively correlated with depth, with shorter stems possibly securing the animal more effectively so as to increase its resistance to the wear-and-tear of current buffeting. Glyptocrinus, one of the largest and hardiest of the dataset, also had the shortest stalk; this is supported by measurements from the Maysville site which found the slab crinoids limited to stalks 20 cm or shorter. These measurements contrast with, for example, the often 1-meter stalks observed in the more delicate disparid crinoids and the 40-cm stalks of the more robust disparids from the same region at different depths (Meyer et al., 2002).

Implications for preservation position

Crinoid pentameral symmetry is expressed as five rays or plates that lead into arms. In G. decadactylus the rays branch twice on the calyx, with two secundibrachs in each ray and fixed tertibrachs passing into free arms. There are 20 uniserial arms, meaning that a single row of brachials runs along each arm (Nichols, 1962; Ausich, 1996).

Taphonomic evidence, i.e., the crinoids’ preserved arm positions, strongly suggests that glyptocrinids possessed physical control limited to only the most proximal portion of the free arms. This resulted in a distinctive trauma position, the afore-mentioned modified shaving brush, in which the arms are seen to be drawn inward in the defensive manner

107 typical of Recent crinoids until a point just distal to where the arms become free, from which the arms are splayed. The implication, then, is that physical control was lacking from this point altogether, which agrees with the generally accepted view that muscled arms in crinoids evolved in the Devonian (Ausich and Baumiller, 1993, 1998). This is further supported by the lack of a fulcral ridge on the articular facet, implying limited arm flexibility and control (Baumiller et al., 2008).

It is unclear how the crinoids functioned in life without the fully-muscled arms that are a feature of all extant crinoids. Ligamentary connective tissue alone would normally have provided little or no contractile ability. Despite suggestions that the crinoids could have relied largely on external forces such as gravity or fluid drag to assume feeding or defensive postures, this method alone is very unlikely, as most of the slab crinoids had clearly achieved a defensive position, at least to the best of their ability. Other proposed substitutions for muscles in arms include a hydrostatic skeleton based on a water vascular system, unusual ligaments in which elasticity/viscosity could be controlled, or ligaments with weak contractile abilities by cytoplasmic filaments (Baumiller et al., 2008).

Unfortunately, none of these features are preservable. Some validity to the suggestion of limited ligamentary control may, however, be made on the grounds that the crinoids that died just prior to the final obrution event (those of Categories 3 and 4, specifically) had remained in trauma position despite showing evidence of some decay, particularly in the arms. The slower decay rate of ligamentary tissue over muscular may explain the persistence of the modified shaving brush, even as the arms disintegrated.

None of the proposed mechanisms for arm control would be capable of a very speedy response. Likewise, the response time implied by muscular control limited to the

108 base of the arms would likely be sluggish in comparison to that of fully muscled arms, rather like closing a door by applying pressure near the hinge. It seems, in fact, that the crinoids were responding to events neither strongly nor swiftly. The crinoids preserved in starburst position were almost certainly amongst the living when obrution occurred, judging by their preservation. Therefore, assuming they were capable of responding to the onslaught of sediment, it must further be assumed that they went down too rapidly to affect a response. Yet other evidence is for fairly gentle deposition rather than a high-speed onslaught, even when taking oscillating currents into consideration.

The starburst position seen on the slab is almost entirely aboral-up. This is note- worthy in light of the assertion that pre-Devonian crinoids would be unlikely to preserve in oral-up position, for reasons referring again to the limited muscles in said crinoids' arms.

Recent stalked crinoids are known to be capable of crawling, which they do oral surface up, in the manner of Recent comatulids, or with the crown parallel to the substrate (Messing et al., 1988; Baumiller et al., 2008). The same limited musculature that prevented Ordovician crinoids from achieving a full shaving brush defensive position would have likewise made crawling impossible. Baumiller et al. (2008) tellingly found a strong correlation between pre-Devonian crinoids' apparent inability to crawl and a paucity of specimens preserved in oral-up position.

Escape through mobility was therefore an impossibility for glyptocrinids. And while

Recent crinoids, especially the vigorous comatulids, are able to respond rapidly to changes in condition, it appears that this was not the case for Ordovician crinoids. Slow to respond and unable to escape, glyptocrinids might more potentially be found in aboral-up starburst position, which is what Baumiller et al. (2008) did find: of 35 starburst specimens on M-

109 slab, 33 were aboral-up. Figure 28 shows the process by which this occurs: a glyptocrinid is subject to drag on its filtration fan or to sediment weight and is forced downstream and pressed oral surface downward to the substrate.

Implications for attachment strategy

It has already been noted that the stem of Glyptocrinus was short in relation to other

Paleozoic crinoids of the region, which is attributed to an adaptation for withstanding the higher current velocities of shallower water. A filtration fan will not function properly, however, if the animal is not firmly anchored in place, making the crinoid unsuited for an eleutherozoic existence, nor would it likely benefit the crinoid to be buffeted about in the current.

Baumiller et al. (2008) noted that the lack of musculature necessary for crawling in

Ordovician crinoids was paired with a lack of any means to reattach to the substrate. As the distal coils grew in position, their initial site of attachment would necessarily have been a permanent one. This disadvantage, however, was probably outweighed by the wider range of attachment-site possibilities afforded by a distal coil. Guensburg (1992) found encrusting crinoids in the Middle Ordovician of Tennessee to be limited to hardgrounds that accommodated their holdfasts, while coil-attached crinoids colonized any site that supported their food-gathering needs. This often included living as commensals on the encrusting crinoids. Distal coils were, then, found to be strongly associated with opportunistic species, which are also usually associated with accelerated growth rates and briefer life spans, perhaps nullifying the need to disattach and reattach. Distal coils were

110 also found to be often paired with dense filtration fans, which suggests that such a fan is more efficient for food-gathering than aerosol filtration theory would allow (Guensburg,

1992).

The question of autotomy

Autotomy, wherein crinoids disattach parts of their bodies, generally their crowns, in response to stress, is usually attributed to a defensive response to imminent predation.

Autotomized stalked crinoids have been found in obrution deposits, which implies that the response was extended to include any manner of stress; it's possible that escape capacity was enhanced without the weight of the stem. Yet while the Maysville crinoids may have suffered the stress of predation, and certainly did obrution, it is unlikely that they were capable of autotomizing their crowns in response to it (Baumiller and Gahn, 2004).

It is very difficult to state with real certainty whether autotomy actually occurred in many cases. This is all the more true for Ordovician crinoids, which lacked the particular skeletal morphology indicative of the ability to autotomize, which doesn't appear until the

Mississippian, and the arrangement of ligaments necessary to the task are unfortunately not preserved, if they were ever present. It is possible that the ability to autotomize would evolve only later, as predation pressure increased (Baumiller and Ausich, 1992; Thomka,

2010). It is further possible that points of weakness along the stem made it more likely to break at certain points, as previously stated. There is, however, no compelling indication of crown autotomy on the slab, and where very well-preserved crowns exist without a stem, there is almost always a break in the matrix to explain it (Fig. 29). Furthermore, the ability to drop the stem might be useless for a crinoid incapable of making good its escape or reestablishing itself upon having done so.

111

FIGURE 28 — Schematic representing response of stalked crinoid with non-muscular arm articulations to increased current. Drawing after Baumiller, et al., in Echinoderm Paleontology, 2008.

112

FIGURE 29 — Crinoid crowns that appear to have separated from their stems during collection, rather than by autotomy.

113 The Community

All fossil assemblages were once, whole or in part, living communities. Well-preserved, autochthonous fossil communities are the most highly prized for paleoecologicial studies: the so-called snapshot of life covers, to the best ability that taphonomy can offer, every aspect of the community concept. The concept itself requires definition.

Fagerstrom (1964), building on categories previously established by Boucot (1953) and others, defined a fossil assemblage as a group of fossils from the same restricted stratigraphic interval and geographic locality. It's a very general definition, containing four broad categories, two of which are termed "assemblages" and two of which are called

"communities." The first, a transported assemblage, is one in which all or most of the specimens were carried into the burial site from more than one contemporaneous community. The second, a mixed fossil assemblage, contains specimens from either one or more than one community; the mixing occurs through some of the specimens being transported and some not; moreover, the assemblage may contain specimens eroded from pre-existing rock.

A fossil community, by contrast, would be defined as an assemblage in which all the organisms were ecologically compatible and living as a single community at the same time.

Preserved communities are usually the result of a catastrophic event resulting in mass mortality, as the Maysville Lagerstätte is interpreted to be. Not all communities are complete: a residual fossil community has undergone a certain amount of preburial alteration as to the numbers and membership of the community; winnowing may remove certain groups from the community and/or physical conditions may favor or disfavor the preservation of other groups. A community proper, however, is complete or nearly so, and

114 further narrowing of the definition yields the fossil census community, wherein all specimens are contemporaneous. The Maysville Lagerstätte conforms to the narrowest definition offered, and the rarest in occurrence: the fossil census population, in which the specimens are not only a complete representation of a contemporaneous community, but are all the same species. The existence of possibly escaped fauna is impossible to prove either way, but sheer crinoid density points to there not being much living space to spare.

An average of 230 individuals per square meter is very high density for crinoids.

Considering that the fossil remains are in situ, Kidwell et al. (1986) distinguished between two types of high-density fossil aggregations: those produced by extrinsic factors, such as the shell pits of predator discards, and intrinsic biogenic concentrations, which would describe the Maysville site. This second type is typically an autochthonous or parautochthonous concentration representing density resulting from the organisms' own intrinsic behavior. Gregarious species typically accumulate for three reasons, two of which may be recorded in the Maysville Lagerstätte (the third, consisting of dense aggregations associated with spawning, feeding, or moulting, is not applicable in the present study).

While all three categories are meant to be distinct, it is suggested that there could be some overlap.

1. The preferential colonization by larvae of sites with abundant adults: Initially free-swimming, crinoid larvae settle after a few days onto whatever substrate is most suitable to their particular morphology; most attach themselves to a hard substrate by an adhesive adaptation on their ventral surface. Glyptocrinid larvae would have settled preferentially on something branchlike and graspable — a stalk or another crinoid — and would, once established at such a site, begin to grow the stem with permanently coiled

115 holdfast around the object. The location is further selected for its suitability to the particular crinoid's feeding strategies and general hardiness in a current. The Maysville site must have been an especially suitable location for glyptocrinids on all factors, and the resulting dense monospecific assemblage further strongly suggests autochthony, as such assemblages do (Meyer, 1997; Brett and Allison, 1998; Thomka, 2010).

The spatial "patchiness" of the crinoid fossils is likewise typical of larval settling, and the effect is similar to what is seen in living stalked crinoid assemblages, if more dense

(Thomka, 2010). Trawling surveys (Meyer et al., 1978) and submersible observations

(Neumann et al., 1977; Messing et al., 1990) found Recent isocrinids living in relatively dense groupings (several specimens per m) and scattered ones (one or two specimens per several m), but all groupings shared a patchy distribution similar to what is seen in

Maysville.

2. Single colonization events by an r-selected or opportunistic species: The great quantity of larvae that settled at the Maysville site is characteristic of opportunistic species, which are both highly reproductive and excel in their powers of dispersal so as to extend their habitat (Whitlach and Zajac, 1985; Brett et al., 2008). Opportunists are typically less highly-specialized and so are less restricted by resource availability, unlike equilibrium species which maintain relatively stable population levels in a physiologically-relegated zone. This characterization of Glyptocrinus as an opportunistic generalist seems to disagree with earlier assertions relating to aerosol filtration theory. However, Holterhoff (1997) demonstrated aerosol filtration theory to be imperfect regarding distribution of taxa in the

Paleozoic in certain instances, one being that, of all taxa sampled, the most widely distributed and abundant were found to be two especially dense fan crinoids. This

116 indicates that while aerosol filtration theory is generally applicable, Paleozoic crinoids with denser fans were actually more niche-flexible than it would suggest.

Typical opportunists are known for their ability to withstand more marginal conditions, and the large population of the slab implies that the glyptocrinids were thriving in the wave and current-dominated habitat. Indeed, opportunists are known for generating explosive population bursts in young, physically-stressful environments; these environments were often ephemeral, arising sporadically between destructive events

(Levinton, 1970; Brett et al., 2008). The resulting high-density, monospecific clusters of animals were themselves often young, and short-lived as well. It is worth noting that the green algae that is the proposed anchorhold for the glyptocrinids is itself considered an opportunistic species.

It is suggested that some overlap between the two preceding categories is possible and occurred in the Maysville Lagerstätte: explosive colonization by an opportunistic species, as per the second category, yet transpiring over more than one generation, as per the first. The use of some crinoids as anchorholds for others, combined with a range of body sizes, indicates more than one colonization event in close succession ("Community

Strategy," this document).

117 The Holdfast Forest

Glyptocrinid holdfasts consisted of a clinging whorl, and were neither weighty nor buried

(Seilacher and MacClintock, 2005). Yet despite their weak attachment capacity, the

Maysville Lagerstätte crinoids had been anchored to something sturdy enough to secure them in the relatively energetic environment. Their anchors are no longer in evidence, exceptions being the animals wrapped around each other or attached to a bryozoan (Fig.

30). Knowing that crinoids' coils were not prehensile but permanent, and that these were of a remarkably consistent diameter, would seem to indicate that all but the exceptions were anchored to a very similar structure, one which also positioned them above the substrate. Seaweeds, a macrophytic green algae, are the most likely anchorhold (Fig. 31).

That there is no evidence for anchoring seaweeds is much to be expected.

Ordovician plant matter is very rarely preserved. Few occurrences, however, do exist throughout the Cincinnatian, most commonly in the shallows, where photosynthesis potential was enhanced (Meyer and Davis, 2009). Cyanophyta in the form of algal mats has been found, as have several species of the calcareous green algae group Dasycladaceae,

Rhodophyta or coralline red algae, and microscopic phytoplankton, including acritarchs

(Cross et al., 1996). The preservation potential of all of these is greater than that of non- calcareous green algae, however, including the largest green algae, the seaweeds. Unlike

Rhodophyta, which occurs twice on the slab, seaweed does not produce preservable calcite crusts or any other hard parts. All the same, carbonaceous compression films of branching macrophytic algae have been recovered from Ordovician rocks in the Cincinnati area. Cross et al. (1996) presumed them to be more numerous, but allowed that they are nondescript enough to be ignored by most collectors.

118 Protein sequence analysis by Heckman et al. (2001) indicate green algae and major lineages of fungi to have been present as long as 1000 mya, and phosphatized lichen fossils have been found in the Doushantou Formation dating between 551 and 600 mya (Yuan et al., 2005). Also, there is ample microfossil evidence of plant matter in dispersed spores on land; while accounts differ, the oldest uncontroversial record of such spores is Mid-

Ordovician (Steemans et al., 2001; Parnell and Foster, 2012). Based on these findings, it is generally accepted that green algae were the dominant plants of the Ordovician, as well as the ancestral type for the earliest land plants.

These were non-vascular, frondose marine plants, and likely grew in small, dense patches, as their descendants are prone to do today. A "forest" of such plants would present a good anchor-hold prospect for opportunistic glyptocrinid larvae.

The seaweeds would serve as a buffer against energetic water motion, which may have been of some importance considering the crinoids' weak attachment ability coupled with a feeding morphology and crown adapted toward higher-energy currents. The seaweeds would aid in binding any asphyxiating muddy sediment that may have been in the crinoid habitat, keeping the sediment at bay and out of crinoid ambulacral grooves. This would clearly benefit the crinoids, although, as previously stated, the additional friction created by the plants could have also added to the habitat's undoing, inducing or facilitating the inducement of sediment deposition. Finally, the presence of plant matter may imply elevated conditions for photosynthesis, and would likely indicate a well-oxygenated setting

(Meyer and Davis, 2009).

Certainly the crinoids were anchored to something in life and during the event that finally buried them. Furthermore, other faunal elements that were not living amongst the

119 crinoids ended up being buried with them; these elements likely washed in earlier but were stopped from progressing further than the crinoids' habitat, as if trapped there.

Similarly, loose crinoid elements, stem fragments in particular, appear to have been held fast within the crinoid/seaweed forest, accumulating despite the currents.

A cohesive microbial mat as anchoring plant matter was considered and rejected as very unlikely, because such a structure is associated with sulfur-fixing bacteria that forms over decaying matter, and could probably only form under very quiet, disoxic conditions

(Brett and Baird, 1986). Nor does the slab display the spherical structures or the "microbial cloak" associated with a microbial mat (Meyer and Milsom, 2001). The black patina of a

"cloak" could, however, be an explanation for the dark coloration of some of the crinoid fossils. Decaying plant matter, whether or not in "cloak" form, can contribute a dark lamination to fossils, similar to what Meyer and Milsom (2001) observed in the field and

Meyer and Oji (1992) induced in laboratory experiments.

120

FIGURE 30 — Examples of crinoid stems anchored to other crinoids.

121

FIGURE 31 — Artist's rendition of the Ordovician seafloor with an emphasis on seaweeds, from geology.wisc.edu.

122 Community Strategy

Communities of opportunistic species are frequently short-lived. Although an aggregation of opportunists is almost ephemeral by definition, there still appears to have been more than one occurrence of mass larval settling at the Maysville site. Two factors could have made the crinoids' dense community livable for a longer span of time: attachment strategy and the size of the crinoids themselves.

The number of complete/near complete fossils combined with the assemblage of ossicles implies numbers of crinoids that initially appear too dense to survive were they all alive at the same time. Yet with evidence for a single burial event, no evidence of transportation, and no prominent background assemblage, it's very likely that the fossil density of the slab represents the population density in life. Tiering is suggested as a strategy to cope with the problem of overcrowding.

Tiering, a community structure in which organisms are distributed vertically in space, is a common solution to the problem of resource allocation (Ausich, 1980; Ausich and Bottjer, 1982). As passive suspension feeders, crinoids need clear access to horizontally flowing currents in order to collect food. In stalked crinoids this is achieved largely through the stem, which otherwise serves no purpose (Hess et al., 2003): the adhesive ventral surface of crinoid larvae means that a stem is not needed for attachment, but instead develops in order to elevate the animal off the substrate, the better to take advantage of food-bearing currents and likely to avoid ambulacral-clogging sediments. The stem does, then, afford crinoids with the potential to take advantage of multiple levels above a substrate (Bottjer and Ausich, 1986; Guensberg, 1992).

Niche differentiation is a term generally applied to different guilds within a single

123 community (Ausich, 1980; Watkins, 1991; Holterhoff, 1997), wherein a guild is defined as a group of species that exploit the same resources in basically the same way (Root, 1967).

Lane (1963) considered first-order niche differentiation as that practiced by benthic taxa in their chosen height above the sea floor; in Paleozoic crinoids this is coupled with stem length (Guensburg, 1992). Likewise, Recent stalked crinoids of different guilds are reported arranged in parallel rows, to the extent that "they appear planted" on lithoherms (Messing et al., 1990), apparently for the purpose of avoiding overlapping filtration fans (Neumann et al., 1977; Hagdorn, 1999; Hess, 1999). But while interspecific tiering is well-documented

(Lane, 1963; Ausich, 1980; Ausich and Bottjer, 1982; Bottjer and Ausich, 1986; Watkins,

1991; Guensburg, 1992; Taylor and Brett, 1996), intraspecific tiering as a niche strategy, though less often seen, is also not unknown (Taylor and Brett, 1996; Hagdorn, 1999).

Brett et al. (2008) found calyx height to have a relatively strong correlation with stem length "but with considerable scatter." It is suggested that this scatter was actually functional, and served the purpose of differentiating the height elevation of the crinoids above the substrate. The seaweed itself likely possessed some variation in height, and the crinoids could have enhanced this by settling at different points along the seaweed's length, but further deviations from the height "norm" would have given these acknowledged opportunists an added advantage for colonization. Adaptability is yet another hallmark of opportunistic species. Having established an attachment site, glyptocrinids grew the stem distal coil first, then added length by adding columnals up the stem at the calyx (Hess et al.,

2003). Therefore, the crinoids may have been exploiting "niches within niches:" growing their stem out to the point of best advantage whether by adding height or even extending laterally along the substrate (Holterhoff, 1997; Brett et al., 2008), with the best feeding

124 positions likely going to the earliest-established individuals. The "considerable scatter" of earlier measurements can go to extremes at times, with even the smallest crowns displaying inordinately long stems and very large crowns having relatively stumpy ones

(Fig. 32).

The suggestion of "earliest-established individuals" leads to the issue of one or more than one mass settling event. To all appearances, the crinoid community was multi- generational. Primarily, some crinoids are wrapped around others, indicating that the anchoring individuals were already there when the newer larvae settled. This is not uncommonly seen in gregarious echinoderms, and would also indicate a short larval period, as is typical of many opportunists (Guensburg, 1992; Hagdorn, 1999; Hess, 1999).

Furthermore, the crinoids are not all the same size. Crown height, though undersized overall, is varied enough to indicate a variety of age groups.

So as is usually true, the community probably persisted beyond the original colonizers, and what was likely a prime, if rather energetic, location attracted and held several spatfalls. But it is also true that the crinoids are small, by even local standards. As previously established, the Maysville crinoids are roughly 39% smaller than other

Glyptocrinus decadactylus collected from the type-Cincinnatian.

The small size of the Maysville crinoids would qualify them as what Gould (1977) called "proportional dwarves," characterized by adult features paired with small adult size relative to related specimens. Causes are unclear. It has been observed that very small crinoid taxa are common in offshore facies in the North American midcontinent, and this has been interpreted as being indicative of colder water (Pabian and Strimple, 1979;

Heckel and Pabian, 1981). Conversely, large, robust crinoids were typically found in coeval

125 nearshore habitats of warmer waters. The Maysville site, interpreted as being in the transition zone between near and offshore, may have been unusually cold, although the warm climate and the fact that the community was living within the photic zone would suggest otherwise. Another possibility is a hypothesis put forth by Holterhoff (1997) regarding undersized crinoids among the Copan fauna, which he attributed to oxygen stress. Yet this seems even less a potential single cause than temperature: the high population density of the crinoids and the likely dense stand of plant matter they occupied would indicate well-oxygenated waters. The community was close enough to storm wave base and subject to robust enough currents to supply ample oxygen through that dissolved at the air-water interface, and photosynthetic processes by the algae would supplement oxygen supplies further. If the water were, in fact, very cold, this would only increase oxygen levels, as oxygen dissolves more readily in colder water.

"Dwarfed" fossil assemblages are not common in the fossil record. Cloud (1948) nevertheless created categories to describe the three types of dwarf assemblages observed amongst brachiopods, those being (1) dwarfing due to physical retardation, including the above listed causes, (2) assemblages segregated by currents, and (3) assemblages of immature specimens. The second option would require that the crinoid community was transported, which it was not. The third is unlikely, as the crinoids are at least superficially physically mature (Brower, 1974 and 1994; Clark, 2012); and this would not account for the observed size diversity. Option one, physical retardation by stress, is the best explanation (Fagerstrom, 1964).

Barring the possibility of the Maysville crinoids being a new subspecies (for which there is no evidence either way), the community was stressed, whether by very vigorous

126 currents, cold water, overcrowding, or a combination of the three. Repeated, highly successful colonization events at the same site argue against the habitat being severely stressed, at least not beyond what the hardy glyptocrinids were capable of tolerating.

Therefore, it is most likely that stress by overcrowding was the greatest factor in the dwarfism, which may have reached a point where, while conditions were otherwise conducive to larval settling, new attachment sites were lacking. This led the crinoids to settle atop established crinoids when necessary and grow their stems out to a length that best exposed the feeding fan to the current. Under these circumstances, smaller size may have been an advantage or even a necessity. The crinoids were likely short-lived and fast- growing but they were established long enough to reach full maturity, though they never reached full size.

It is important to note that the disarticulated ossicles in various stages of decay are equal in size to those of the best-preserved specimens, indicating that there is not a background assemblage of perhaps older, larger specimens.

127

FIGURE 32 — Disparity in stem length between similarly-sized crinoids. A) Short stem. B) Long stem. The size of the distal coils is a good indication of scale.

128

Comparison with other Lagerstätten

The Maysville slab can be best compared to two particular types of obrution deposit, described by Brett and Seilacher (1991) as the Gmünd and the Crawfordsville. The Gmünd consists of well-preserved fossils, primarily crinoids, entombed in thin shale layers buried along the top contacts of skeletal limestone. The limestone is typically a hardground resulting from sediment starvation, and the bedding layer a storm-generated rapid influx of siliciclastic mud. This type of deposit is often associated with transgressions and deepening water, although this is not the case for the Maysville deposit (Ausich et al., 1999b).

The second type is named for the Mississippian Edwardsville Formation near

Crawfordsville, Indiana. The Crawfordsville accounts for many, possibly most, Paleozoic

Lagerstätten, and like in Maysville, is thought to represent an ephemeral assemblage.

Characterized by primarily fine-grained mud and siltstone bedding planes overlying skeletal debris, these mainly differ from the Gmünd in that they are overlain by the same skeletal debris as underlies them, or something very similar. This obrution type is interpreted as opportunistic colonization of a seafloor during a brief period of sediment starvation: the assemblage is buried by mud within a generation or two (Ausich et al.,

1999b).

The relationship between the preceding two types is obvious, and one may grade into the other. As the rocks that originally overlaid the Maysville deposit are lost to erosion and weathering, one of the defining characteristics of the Crawfordsville-type deposit is unfortunately lacking, although it otherwise falls into this category. The following examples of obrution also fall along the spectrum of the two types described.

129 Catastrophic storm deposition is a hallmark of the type-Cincinnatian, and is the usual source of obrution. It is therefore not surprising to find that these conditions generated another excellent Glyptocrinus Lagerstätte in the Stamping Ground Member, not far from Maysville. Cincinnatian crinoid assemblages tend toward low-diversity (Brett et al., 2008), and more than five species in a deposit is very rare; therefore it is not unusual for assemblages to be monospecific (Ausich, 1999a). Also like the Maysville deposit, the

Stamping Ground deposit is interpreted as resulting from rapid opportunistic colonization.

Up to 100 fossils were found within 15 to 20 cm of skeletal grainstone covering roughly two m in the upper part of the Lexington Limestone at Georgetown, Kentucky. The preservation is very good, although not as well articulated as the specimens of the

Maysville deposit, displaying perhaps the effects of slightly more decay. Like the Maysville deposit, the preservation is attributed largely to the fact that these crinoids are preserved in situ; however, unlike the Maysville, these crinoids are stacked in layers about 5 cm thick and separated by thin muds. This is thought to represent multiple deposits, although still within a single obrution event. The crinoids are slightly larger than those at Maysville, averaging between 14 and 16 mm in crown height (Brett et al., 2008). The crinoids' habitat is thought to have been just below normal wave base, and, like the Maysville crinoids, sheltered in some way from normal wave action and minor storms. Their generally short stems would have aided in stabilization, as would the density of the community (Brett et al.,

2008).

The intervening mud layers of the Stamping Ground deposit could make it more difficult to identify as belonging to a single obrution event. Conversely, some deposits that appear initially to be single event may turn out to belong to several. One such is the Copan

130 Lagerstätte from the Upper Pennsylvanian (Missourian) Barnsdall Formation near Copan,

Washington County, northeastern Oklahoma. Like the Maysville, this is an episode of rapid burial by distal storm event, but close examination by Thomka (2010) determined low sedimentation rates between multiple events to be responsible for only the appearance of a single event. In fact the deposit represents stacked intervals of many crinoid communities that became condensed into 50 cm of what is predominantly mudstone. Within many layers, each representing a single event, crinoid remains show evidence of some specimens having died in the obrution while others appear to have died a short time beforehand, in a manner thought similar to what is seen in Maysville. The diversity of the Copan deposit is, however, the greatest difference between the two: even accounting for the fact that the low-energy, offshore muddy shelf that created the Copan deposit is the area where

Paleozoic crinoids were typically most diverse, the diversity of the Copan, at 44 genera and

50 species, is impressive (Thomka, 2010).

Less diversity is seen in a well-known assemblage from the Lower Devonian

Manlius/Coeymans Formation of Central New York, discovered in the late 1800's (Brett,

1999). Crinoids are not common in most of the Manlius, and seven species total have been found; a very notable exception is Lasiocrinus occurring in dense clumps. One famous high- density slab was excavated and delivered to the Yale Peabody Museum early in the last century measuring 2.1 by 1.8 meters. It contained four species of crinoids encased in a light grey limestone. The environment is interpreted as shallow subtidal, not as deep as the

Maysville site, which makes it more probable that the crinoids were transported by higher energy currents, which appears to be the case. Nonetheless, the preservation is very good, likely aided by the fact that most of the crinoids are the more robust camerates. Some, like

131 Lasiocrinus, are also considered opportunistic species (Brett, 1999).

A more richly-varied fauna is seen in the Middle Ordovician Trenton Group, consisting of thinly-bedded grey limestones containing over 75 species of echinoderm, brachiopod, bryozoan, and trilobite (Brett, 1999). The Rust Member, outcropping near

Trenton Falls, New York, is the most productive for crinoids. Seventeen species have been identified, primarily disparids such as Cincinnaticrinus and Ectenocrinus. Glyptocrinus is less common, with the exception of a single bed with conditions reminiscent of the

Maysville site: a thin layer of storm-deposited lime mud holds many glyptocrinids in starburst position, with the aboral surfaces in the expected upward orientation. As previously stated, this is diagnostic for very rapid burial in Ordovician stalked crinoids, and many communities within the Trenton Group, crinoidal and otherwise, show signs of being buried rapidly while alive or shortly thereafter. The Rust Member appears deposited mostly in a middle shelf environment, indicating water depths similar to those of the

Maysville site. Therefore the Rust Member crinoids are considered a stressed, ephemeral community of opportunists from a transitional environment between a shallow, stormy shelf and the deep offshore.

Whereas all the preceding Lagerstätten were Paleozoic, a certain deposit from the

Triassic Muschelkalk of Central Europe also suggests comparison with the Maysville slab.

The Crailsheim and Neckarwestheim Encrinus Lagerstätten were formed on bioherms situated along a carbonate ramp in the Crailsheim Member, and burial is interpreted as resulting from bottom backflows of mud avalanches down the ramp (Hagdorn, 1999). The bioherms are made primarily of false oysters and encrusted with worm tubes and forams.

One large bioherm complex contains over 120 specimens of Encrinus, which, like the

132 Maysville crinoids, were living between normal and storm wave base. This assemblage is of interest because, like the Maysville crinoids, the Crailsheim Encrinus apparently employed intraspecific tiering as a niche strategy. Positions suggest a pioneer settling, after which increased sedimentation made more height above the substrate a necessity to avoid smothering muds. Subsequent spatfalls of larvae then tended to settle on the basal stems of adults. The colony ultimately consisted of juveniles at lower levels and adults up to 160 cm above the sea floor. Encrinus, like Glyptocrinus, could not reattach its stem if dislodged, and so this arrangement was apparently working for successful long-term settlement (Hagdorn,

1999).

133 Conclusions

The Maysville Lagerstätte is interpreted as an autochthonous community of Glyptocrinus decadactylus entombed in a classic "smothered bottom" obrution deposit with low directional flow, not atypical where the deposition site is more distal to the shoreline. The crinoid community had colonized a rise slightly elevated above the deep subtidal regions, high energy enough to support glyptocrinid feeding habits, yet sheltered enough to accommodate their weak attachment capacity. The crinoids' distal coils were wrapped around what was probably seaweed, which would have anchored the animals and elevated them above the substrate. Further elevation by column growth into tiers and the retention of sub-adult size into maturity were employed in order to avoid overcrowding and the resulting overlapping filtration fans.

Turbidity currents began with a storm event proximal to the coastline. Conditions likely associated with the storm effected some changes within the crinoid habitat, resulting in some early mortality, followed by scavenging. Finally, the storm caused denser, sediment-charged water to be moved down-slope by gravity where the heavier particles settled out, resulting in a bulk density decrease and reversing the flow's buoyancy. The resulting sediment plume of fines extended laterally along a pycnocline, intruding into distal regions and the crinoids' habitat on the rise. The eventual decelerating flow velocity, in combination with the friction of the seaweed/crinoid stand on the rise, reduced the flow competence enough to induce sediment deposition. The crinoid community, then, facilitated its own burial, and the Lagerstätte represents the result of a feedback process.

Scavenging essentially ceased with the final burial event.

134 G. decadactylus is typically found in the deep subtidal regions, although "deep" in the context of the shallow sea represented by the Cincinnatian is still likely within the photic zone. All the same, aerosol filtration theory would put them morphologically best-suited for shallower, more energetic waters. It appears that glyptocrinids were actually generalists, adapting to a wider range of conditions than are typical of crinoids. Generalists are typically opportunists, and evidence suggests that this best describes the glyptocrinids: the robust calyx could withstand strong currents, the distal coil allowed for a multitude of attachment possibilities, and they were by all appearances highly reproductive. One spatfall after another settled on the marginal environment of the rise, and when conditions became too crowded, the crinoids settled atop one another and stymied their growth, while still growing their columns to compensatory lengths.

A population estimate of over 400 Glyptocrinus is conservative, as it takes no account of the many stem fragments of varying lengths. The crinoid community maintained itself despite its high density, apparently thriving before the obrution that ended it.

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144 Appendix 1: Measurements

calyx stem coil stem calyx slab height condition length diameter length 2 height J 9.4 2 13.6 NA NA NA J NA NA 82.5 2.43 82.5 NA J NA 2 NA NA NA NA J 13.7 2 NA NA NA NA J 9.5 3 NA NA NA NA J 9.3 3 NA NA NA NA J NA 3 NA NA NA NA J NA 4 NA NA NA NA J 11.3 2 20.4 NA NA NA J NA NA 52.2 5.8 52.2 NA J NA NA 22.3 4 22.3 NA J NA NA 62.1 3.3 62.1 NA J NA 5 NA NA NA NA G 16.5 4 NA NA NA NA F NA 4 NA NA NA NA B NA 2 NA NA NA NA B 15.3 3 NA NA NA NA B 8.4 4 NA NA NA NA B NA 5 NA NA NA NA B 6.7 2 NA NA NA NA B NA 4 8.2 NA NA NA B NA 4 NA NA NA NA B NA NA 5 2.1 5 NA B NA 4 NA NA NA NA B NA 4 NA NA NA NA B 10.7 2 NA NA NA NA B 8 3 142 NA NA NA B 11.3 1 104 2.4 104 11.3 B 9.2 3 54.5 NA NA NA B 6.9 2 34.5 NA NA NA B 7.5 1 43.2 closed 43.2 7.5 B NA 3 NA NA NA NA B NA NA 96 4.5 96 NA B NA 4 NA NA NA NA B NA 3 NA NA NA NA B NA 4 20 NA NA NA B 12 3 NA NA NA NA D NA 5 NA NA NA NA D NA 6 NA NA NA NA D NA 4 NA NA NA NA D 13.7 2 NA NA NA NA D 11.6 2 NA NA NA NA D 6.9 2 21.5 NA NA NA D NA 4 NA NA NA NA A 11.5 3 14.6 NA NA NA

145 A 13.9 3 32.3 NA NA NA A NA 4 22.6 NA NA NA A 13.6 4 66.4 NA NA NA A 12 2 71.5 NA NA NA A NA 4 41.8 NA NA NA A NA 4 60.3 2.5 60.3 NA A NA NA 6.5 3.7 6.5 NA A 10.5 3 6.8 NA NA NA A NA NA 26 3.1 26 NA A NA NA 32 NA NA NA A NA 4 16.2 NA NA NA A NA 4 18.9 NA NA NA A 4.2 3 192 NA NA NA A 12.8 4 70 NA NA NA A 17.1 3 33.3 NA NA NA A NA 4 36.4 NA NA NA A 6.6 3 20.3 NA NA NA C NA 1 46.4 0.9 46.4 NA C 6.5 2 24.2 NA NA NA C 11 3 93.6 3.8 93.6 11 C NA 4 12.8 NA NA NA C NA 5 NA NA NA NA C NA 4 NA NA NA NA C NA 4 6.1 NA NA NA C NA 3 22.3 NA NA NA C NA 4 NA NA NA NA C NA 3 26 NA NA NA C NA 4 22.3 NA NA NA C NA 4 9.3 NA NA NA I 8.1 2 20.7 NA NA NA I 9.1 2 7.7 NA NA NA I NA 2 55 NA NA NA I 10.4 2 10 NA NA NA I 16.3 2 9.5 NA NA NA I NA NA NA NA NA NA I 11.4 2 13.2 NA NA NA I NA NA 8.3 3.4 8.3 NA I NA NA 14 3.6 14 NA I 12.5 2 70.5 NA NA NA I NA 2 24.7 NA NA NA I NA NA 106 2.6 106 NA I 10 2 202 NA NA NA I 11.8 2 NA NA NA NA I NA 3 NA NA NA NA I NA 4 NA NA NA NA I NA 3 NA NA NA NA I NA NA 83.8 7.6 83.8 NA I NA 3 36.6 NA NA NA I NA NA 82 6.6 82 NA

146 I NA 3 26 NA NA NA I 9.5 3 41.7 NA NA NA I NA 6 NA NA NA NA I NA 2 NA NA NA NA I NA 6 NA NA NA NA I NA 5 NA NA NA NA I 11.5 1 135.6 7.3 135.6 11.5 I 13.2 2 28.4 NA NA NA I NA NA 46 5.2 46 NA I 12.9 2 5.1 NA NA NA I NA 6 NA NA NA NA I 7.3 3 25.3 NA NA NA I 10.7 1 116 3.6 116 10.7 I 7.7 2 6 NA NA NA I NA 5 NA NA NA NA I NA 6 NA NA NA NA I 8.1 1 85 6.8 85 8.1 I NA NA 120.2 5.4 120.2 NA I NA NA 100 4.6 100 NA I NA NA 80 4.3 80 NA I NA NA 81 2.1 81 NA I NA 5 NA NA NA NA I NA 6 NA NA NA NA I NA 4 NA NA NA NA I 9.5 2 49.9 NA NA NA I 8.9 1 74.1 4.8 74.1 8.9 I 8.1 2 50.5 NA NA NA I NA NA 26 2.9 26 NA I 6.3 1 58.8 2.8 58.8 6.3 I NA NA 38 5.8 38 NA I NA 4 34.8 NA NA NA I NA 4 9.8 NA NA NA I NA 4 58.6 NA NA NA I 12.8 1 116 4.2 116 12.8 I NA 5 NA NA NA NA I 9.8 1 65.5 7.7 65.5 9.8 I NA 4 NA NA NA NA I NA NA 142 5.1 142 NA I 10.1 1 76.4 2.9 76.4 10.1 I NA 3 66 NA NA NA I NA NA 120 5.8 120 NA I 9.4 1 64.2 1.6 64.2 9.4 I NA 4 122 6.1 122 NA I 4.9 2 71.1 NA NA NA I 8.1 3 11.8 NA NA NA I NA NA 136 5.8 136 NA I NA 4 NA NA NA NA I NA NA 89.8 4.8 89.8 NA I NA 4 27.6 NA NA NA

147 I 6.2 2 20.9 NA NA NA E NA 3 12.7 NA NA NA E NA 4 20.8 NA NA NA E NA 3 45.7 NA NA NA E NA NA 66 2.5 66 NA E NA NA 52.6 5.5 52.6 NA E 8.9 1 89.8 4.3 89.8 8.9 E NA 4 52 NA NA NA E NA NA 56 7.7 56 NA E 12.3 2 44.4 NA NA NA E NA 4 112 8.1 112 NA E NA 4 29.2 NA NA NA E NA 4 7.6 NA NA NA E NA 4 26.1 NA NA NA E NA 4 34.1 NA NA NA E 8 2 50.4 NA NA NA E 13.2 3 NA NA NA NA E NA 4 30.2 NA NA NA E NA NA 105.2 4.2 105.2 NA E NA 4 120 NA NA NA E NA NA 44 3.2 44 NA E NA NA NA NA NA NA E NA 4 47.2 NA NA NA E 12.6 3 26.9 NA NA NA E NA 3 17.3 NA NA NA E 10 2 NA NA NA NA E 8.4 3 9 NA NA NA E NA NA 43 3.9 43 NA E NA 3 73.3 NA NA NA E NA NA 78 4.9 78 NA E NA NA NA NA NA NA E NA NA 76 4.1 76 NA E NA NA 35 3.4 35 NA E 10.6 3 22.1 NA NA NA E 8.8 2 3.4 NA NA NA E 8.4 2 26 NA NA NA E NA 2 NA NA NA NA E 17.4 3 NA NA NA NA E 10.4 2 3.3 NA NA NA T NA NA 107 6.3 107 NA T 11.3 3 10.3 NA NA NA T NA 4 62.7 NA NA NA T NA NA 60 7.4 60 NA T NA 4 71.4 NA NA NA T NA 4 NA NA NA NA T NA NA NA NA NA NA T NA NA 110 6.2 110 NA T 10 2 32.6 NA NA NA T NA 4 NA NA NA NA

148 T 8.3 2 15.2 NA NA NA K NA 2 NA NA NA NA K NA 3 NA NA NA NA K NA 4 80 NA NA NA K 11 2 86 NA NA NA K NA 2 50 NA NA NA K NA 4 NA NA NA NA K NA 5 NA NA NA NA K NA 5 NA NA NA NA K 9.5 3 37 NA NA NA K NA NA 60 3.4 60 NA K 8.8 2 4.6 NA NA NA K 8.5 3 13.1 NA NA NA K 9.8 3 NA NA NA NA K NA NA 45 4.8 45 NA K NA 3 NA NA NA NA K 12.6 3 NA NA NA NA K 12.2 2 4 NA NA NA K 10.4 2 NA NA NA NA K 9.4 2 48 NA NA NA K NA NA NA NA NA NA K NA NA NA NA NA NA K NA 4 NA NA NA NA K 10.1 3 NA NA NA NA K 4.2 1 61 4.5 61 4.2 K NA NA 58 3.4 58 NA K 8.7 2 62 NA NA NA K NA NA 70 4.4 70 NA K 11.5 3 65.1 NA NA NA K NA NA 30 2.7 30 NA K 11.2 2 107 NA NA NA K 9.1 2 6.9 NA NA NA K 10 NA 11.3 NA NA NA K NA NA NA NA NA NA K NA NA NA NA NA NA K NA 4 25.3 NA NA NA K NA 4 57 NA NA NA K NA NA NA NA NA NA K NA NA 56 5.2 56 NA K NA NA 50 3.2 50 NA K 10.3 2 68.5 NA NA NA K NA NA 97 NA NA NA K 9.1 3 36.6 NA NA NA K NA NA 100 1.6 100 NA K NA 4 12.1 NA NA NA K NA 5 NA NA NA NA K NA NA NA NA NA NA K 7.4 3 46.5 3.2 46.5 7.4 K NA 4 NA NA NA NA

149 K NA NA 6.4 NA NA NA K 9.9 3 8.4 NA NA NA K NA NA 53.3 6.71 53.3 NA K NA 4 46 NA NA NA K 6.1 3 24.3 NA NA NA K 6.2 3 34.8 NA NA NA K NA 4 7.8 NA NA NA K 10.7 3 NA NA NA NA K NA 3 46 NA NA NA K 9.1 2 39.9 NA NA NA K NA NA 12.1 NA NA NA K NA NA NA NA NA NA K NA NA NA NA NA NA c 10.6 M 11.4 M 12.1 M 9.5 M 11.8 M 8.2 M 13.3 M 13.3 M 12.9 M 11.9 M 10.8 M 8.1 M 15.6 M 8.6 M 12.3 M 13.4 M 7.6 M 12.4 M 7 M 14.6 M 7.2 M 11.2 M 15.3 M 9.7 M 16 M 12.2 M 15.5 M 7.9 M 9.3 M 12 M 10 M 9.5 M 9.6 M 9.3 M 14.7

150 Appendix 2: Orientation data from entire slab

Calyces Round to Set 360 to Slab orientation Calc Round "10" deg 0 deg J 263 26.3 26 260 260 j 208 20.8 21 210 210 j 90 9 9 90 90 j 340 34 34 340 340 j 360 36 36 360 0 j 332 33.2 33 330 330 j 13 1.3 1 10 10 I 85 8.5 9 90 90 i 210 21 21 210 210 i 176 17.6 18 180 180 i 153 15.3 15 150 150 i 85 8.5 9 90 90 i 303 30.3 30 300 300 i 217 21.7 22 220 220 i 165 16.5 17 170 170 i 323 32.3 32 320 320 i 12 1.2 1 10 10 i 333 33.3 33 330 330 i 10 1 1 10 10 i 74 7.4 7 70 70 i 113 11.3 11 110 110 i 33 3.3 3 30 30 i 116 11.6 12 120 120 i 170 17 17 170 170 i 20 2 2 20 20 i 350 35 35 350 350 i 293 29.3 29 290 290 i 5 0.5 1 10 10 i 358 35.8 36 360 0 i 214 21.4 21 210 210 i 348 34.8 35 350 350 i 193 19.3 19 190 190 i 160 16 16 160 160 i 78 7.8 8 80 80 i 340 34 34 340 340 i 150 15 15 150 150 i 145 14.5 15 150 150 D 180 18 18 180 180 d 158 15.8 16 160 160 d 190 19 19 190 190 d 240 24 24 240 240 H 45 4.5 5 50 50 H 160 16 16 160 160 H 118 11.8 12 120 120 H 343 34.3 34 340 340

151 H 40 4 4 40 40 H 130 13 13 130 130 H 190 19 19 190 190 H 320 32 32 320 320 H 187 18.7 19 190 190 H 113 11.3 11 110 110 H 139 13.9 14 140 140 H 83 8.3 8 80 80 H 165 16.5 17 170 170 H 115 11.5 12 120 120 H 310 31 31 310 310 H 360 36 36 360 0 H 10 1 1 10 10 H 80 8 8 80 80 H 160 16 16 160 160 H 20 2 2 20 20 H 344 34.4 34 340 340 H 166 16.6 17 170 170 H 153 15.3 15 150 150 H 300 30 30 300 300 H 150 15 15 150 150 H 86 8.6 9 90 90 H 310 31 31 310 310 H 336 33.6 34 340 340 H 112 11.2 11 110 110 H 155 15.5 16 160 160 H 260 26 26 260 260 H 350 35 35 350 350 H 120 12 12 120 120 H 120 12 12 120 120 H 75 7.5 8 80 80 H 354 35.4 35 350 350 H 85 8.5 9 90 90 H 11 1.1 1 10 10 K 267 26.7 27 270 270 K 170 17 17 170 170 K 240 24 24 240 240 K 178 17.8 18 180 180 K 138 13.8 14 140 140 K 320 32 32 320 320 K 342 34.2 34 340 340 K 255 25.5 26 260 260 K 210 21 21 210 210 K 160 16 16 160 160 K 80 8 8 80 80 K 105 10.5 11 110 110 K 238 23.8 24 240 240 K 33 3.3 3 30 30 K 50 5 5 50 50

152 K 113 11.3 11 110 110 K 180 18 18 180 180 K 210 21 21 210 210 K 170 17 17 170 170 K 137 13.7 14 140 140 K 180 18 18 180 180 K 10 1 1 10 10 K 13 1.3 1 10 10 K 120 12 12 120 120 K 250 25 25 250 250 K 30 3 3 30 30 K 340 34 34 340 340 K 50 5 5 50 50 K 120 12 12 120 120 K 143 14.3 14 140 140 K 180 18 18 180 180 C 293 29.3 29 290 290 C 185 18.5 19 190 190 C 30 3 3 30 30 C 320 32 32 320 320 C 227 22.7 23 230 230 C 200 20 20 200 200 C 220 22 22 220 220 C 180 18 18 180 180 C 40 4 4 40 40 C 215 21.5 22 220 220 A 112 11.2 11 110 110 A 250 25 25 250 250 A 195 19.5 20 200 200 A 235 23.5 24 240 240 A 198 19.8 20 200 200 A 210 21 21 210 210 A 187 18.7 19 190 190 A 230 23 23 230 230 A 152 15.2 15 150 150 A 60 6 6 60 60 A 180 18 18 180 180 A 105 10.5 11 110 110 A 333 33.3 33 330 330 A 335 33.5 34 340 340 A 208 20.8 21 210 210 A 220 22 22 220 220 A 352 35.2 35 350 350 B 68 6.8 7 70 70 B 10 1 1 10 10 B 360 36 36 360 0 B 275 27.5 28 280 280 B 25 2.5 3 30 30 B 340 34 34 340 340

153 B 248 24.8 25 250 250 B 110 11 11 110 110 B 190 19 19 190 190 B 300 30 30 300 300 B 210 21 21 210 210 B 297 29.7 30 300 300 B 350 35 35 350 350 B 330 33 33 330 330 M 230 23 23 230 230 M 305 30.5 31 310 310 M 95 9.5 10 100 100 M 250 25 25 250 250 M 18 1.8 2 20 20 M 20 2 2 20 20 M 10 1 1 10 10 M 55 5.5 6 60 60 M 220 22 22 220 220 M 300 30 30 300 300 M 10 1 1 10 10 M 68 6.8 7 70 70 M 310 31 31 310 310 M 173 17.3 17 170 170 M 290 29 29 290 290 M 162 16.2 16 160 160 M 180 18 18 180 180 M 330 33 33 330 330 M 150 15 15 150 150 M 360 36 36 360 0 M 153 15.3 15 150 150 M 165 16.5 17 170 170 M 130 13 13 130 130 M 350 35 35 350 350 M 312 31.2 31 310 310 M 350 35 35 350 350 M 160 16 16 160 160 M 363 36.3 36 360 0 M 220 22 22 220 220 M 273 27.3 27 270 270 M 83 8.3 8 80 80 M 353 35.3 35 350 350 M 335 33.5 34 340 340 M 345 34.5 35 350 350 M 270 27 27 270 270 M 195 19.5 20 200 200 M 116 11.6 12 120 120 M 92 9.2 9 90 90 M 338 33.8 34 340 340 M 160 16 16 160 160 M 355 35.5 36 360 0

154 M 70 7 7 70 70 M 270 27 27 270 270 M 40 4 4 40 40 M 353 35.3 35 350 350 M 200 20 20 200 200 M 190 19 19 190 190 M 283 28.3 28 280 280 M 255 25.5 26 260 260 M 138 13.8 14 140 140 M 304 30.4 30 300 300 M 345 34.5 35 350 350 M 60 6 6 60 60 M 50 5 5 50 50 M 260 26 26 260 260 M 50 5 5 50 50 M 110 11 11 110 110 M 60 6 6 60 60 M 313 31.3 31 310 310 M 222 22.2 22 220 220 M 180 18 18 180 180 M 190 19 19 190 190 M 230 23 23 230 230 M 306 30.6 31 310 310 M 12 1.2 1 10 10 M 150 15 15 150 150 M 312 31.2 31 310 310 M 320 32 32 320 320 M 20 2 2 20 20 M 190 19 19 190 190 M 263 26.3 26 260 260 M 140 14 14 140 140 M 228 22.8 23 230 230 M 195 19.5 20 200 200 M 305 30.5 31 310 310 M 304 30.4 30 300 300 M 312 31.2 31 310 310 M 165 16.5 17 170 170 M 70 7 7 70 70 M 15 1.5 2 20 20 M 190 19 19 190 190 M 340 34 34 340 340 M 10 1 1 10 10 M 130 13 13 130 130 M 275 27.5 28 280 280 M 160 16 16 160 160 M 195 19.5 20 200 200 M 160 16 16 160 160 M 295 29.5 30 300 300 M 113 11.3 11 110 110

155 M 340 34 34 340 340 M 32 3.2 3 30 30 M 190 19 19 190 190 M 350 35 35 350 350 M 305 30.5 31 310 310 M 235 23.5 24 240 240 M 40 4 4 40 40 M 250 25 25 250 250 M 210 21 21 210 210 M 68 6.8 7 70 70 M 310 31 31 310 310 M 240 24 24 240 240 M 80 8 8 80 80 M 190 19 19 190 190 M 313 31.3 31 310 310 M 40 4 4 40 40 M 320 32 32 320 320 M 105 10.5 11 110 110 M 170 17 17 170 170 M 293 29.3 29 290 290 M 220 22 22 220 220 M 170 17 17 170 170 M 320 32 32 320 320 M 355 35.5 36 360 0 M 350 35 35 350 350 M 72 7.2 7 70 70 M 190 19 19 190 190 M 240 24 24 240 240 M 20 2 2 20 20 M 190 19 19 190 190 M 100 10 10 100 100 M 230 23 23 230 230 M 150 15 15 150 150 M 340 34 34 340 340 M 142 14.2 14 140 140 M 70 7 7 70 70 M 170 17 17 170 170 M 340 34 34 340 340 M 200 20 20 200 200 M 90 9 9 90 90 M 303 30.3 30 300 300 M 330 33 33 330 330 M 320 32 32 320 320 M 320 32 32 320 320 M 240 24 24 240 240 M 354 35.4 35 350 350 M 180 18 18 180 180 M 210 21 21 210 210 M 192 19.2 19 190 190

156 M 310 31 31 310 310 M 352 35.2 35 350 350 M 360 36 36 360 0 M 237 23.7 24 240 240 M 190 19 19 190 190 M 270 27 27 270 270 M 352 35.2 35 350 350 M 242 24.2 24 240 240 M 160 16 16 160 160 M 170 17 17 170 170 M 130 13 13 130 130 M 73 7.3 7 70 70 M 310 31 31 310 310 M 350 35 35 350 350 M 140 14 14 140 140 M 10 1 1 10 10

157 Appendix 3: Orientation for North and South areas

Calyx Round or to "10" Set 360 Quadrant Group Orientation stem Calc Round deg to 0 deg 1 NORTH 85 c 8.5 9 90 90 1 NORTH 13 c 1.3 1 10 10 1 NORTH 176 c 17.6 18 180 180 1 NORTH 85 c 8.5 9 90 90 1 NORTH 303 c 30.3 30 300 300 1 NORTH 153 c 15.3 15 150 150 1 NORTH 12 c 1.2 1 10 10 1 NORTH 10 c 1 1 10 10 1 NORTH 210 s 21 21 210 210 1 NORTH 320 s 32 32 320 320 1 NORTH 160 s 16 16 160 160 1 NORTH 335 s 33.5 34 340 340 1 NORTH 5 s 0.5 1 10 10 1 NORTH 240 s 24 24 240 240 1 NORTH 190 s 19 19 190 190 2 NORTH 165 c 16.5 17 170 170 2 NORTH 208 c 20.8 21 210 210 2 NORTH 217 c 21.7 22 220 220 2 NORTH 323 c 32.3 32 320 320 2 NORTH 74 c 7.4 7 70 70 2 NORTH 240 c 24 24 240 240 2 NORTH 230 s 23 23 230 230 2 NORTH 180 s 18 18 180 180 2 NORTH 357 s 35.7 36 360 0 3 NORTH 338 c 33.8 34 340 340 3 NORTH 195 c 19.5 20 200 200 3 NORTH 160 c 16 16 160 160 3 NORTH 355 c 35.5 36 360 0 3 NORTH 263 c 26.3 26 260 260 3 NORTH 90 c 9 9 90 90 3 NORTH 340 c 34 34 340 340 3 NORTH 340 c 34 34 340 340 3 NORTH 360 c 36 36 360 0 3 NORTH 332 c 33.2 33 330 330 3 NORTH 13 c 1.3 1 10 10 3 NORTH 263 s 26.3 26 260 260 3 NORTH 360 s 36 36 360 0 3 NORTH 360 s 36 36 360 0 4 NORTH 230 c 23 23 230 230 4 NORTH 160 c 16 16 160 160 4 NORTH 95 c 9.5 10 100 100 4 NORTH 305 c 30.5 31 310 310 4 NORTH 355 c 35.5 36 360 0 4 NORTH 116 c 11.6 12 120 120

158 4 NORTH 250 c 25 25 250 250 4 NORTH 338 c 33.8 34 340 340 4 NORTH 96 c 9.6 10 100 100 4 NORTH 10 c 1 1 10 10 4 NORTH 270 c 27 27 270 270 4 NORTH 10 c 1 1 10 10 4 NORTH 18 c 1.8 2 20 20 4 NORTH 88 s 8.8 9 90 90 4 NORTH 90 s 9 9 90 90 5 NORTH 210 c 21 21 210 210 5 NORTH 68 c 6.8 7 70 70 5 NORTH 30 c 3 3 30 30 5 NORTH 310 c 31 31 310 310 6 NORTH 190 c 19 19 190 190 6 NORTH 210 c 21 21 210 210 6 NORTH 180 c 18 18 180 180 6 NORTH 192 c 19.2 19 190 190 6 NORTH 240 c 24 24 240 240 6 NORTH 173 c 17.3 17 170 170 6 NORTH 290 c 29 29 290 290 6 NORTH 360 c 36 36 360 0 6 NORTH 343 s 34.3 34 340 340 6 NORTH 180 s 18 18 180 180 6 NORTH 278 s 27.8 28 280 280 7 NORTH 188 c 18.8 19 190 190 7 NORTH 330 c 33 33 330 330 7 NORTH 160 s 16 16 160 160 7 NORTH 130 s 13 13 130 130 7 NORTH 162 s 16.2 16 160 160 7 NORTH 157 s 15.7 16 160 160 8 NORTH 112 c 11.2 11 110 110 8 NORTH 250 c 25 25 250 250 8 NORTH 195 c 19.5 20 200 200 8 NORTH 197 s 19.7 20 200 200 8 NORTH 50 s 5 5 50 50 8 NORTH 362 s 36.2 36 360 0 8 NORTH 150 s 15 15 150 150 9 NORTH 10 c 1 1 10 10 9 NORTH 240 c 24 24 240 240 9 NORTH 20 c 2 2 20 20 9 NORTH 180 c 18 18 180 180 9 NORTH 333 c 33.3 33 330 330 9 NORTH 170 c 17 17 170 170 9 NORTH 158 c 15.8 16 160 160 9 NORTH 10 c 1 1 10 10 9 NORTH 190 c 19 19 190 190 9 NORTH 113 c 11.3 11 110 110 9 NORTH 293 c 29.3 29 290 290 9 NORTH 68 s 6.8 7 70 70

159 9 NORTH 30 s 3 3 30 30 9 NORTH 200 s 20 20 200 200 9 NORTH 138 s 13.8 14 140 140 9 NORTH 190 s 19 19 190 190 9 NORTH 90 s 9 9 90 90 9 NORTH 312 s 31.2 31 310 310 10 NORTH 10 c 1 1 10 10 10 NORTH 33 c 3.3 3 30 30 10 NORTH 116 c 11.6 12 120 120 10 NORTH 350 c 35 35 350 350 10 NORTH 358 c 35.8 36 360 0 10 NORTH 340 c 34 34 340 340 10 NORTH 214 c 21.4 21 210 210 10 NORTH 183 s 18.3 18 180 180 10 NORTH 140 s 14 14 140 140 10 NORTH 40 s 4 4 40 40 10 NORTH 330 s 33 33 330 330 11 NORTH 270 c 27 27 270 270 11 NORTH 345 c 34.5 35 350 350 11 NORTH 138 c 13.8 14 140 140 11 NORTH 255 c 25.5 26 260 260 11 NORTH 40 c 4 4 40 40 11 NORTH 70 c 7 7 70 70 11 NORTH 283 c 28.3 28 280 280 11 NORTH 58 s 5.8 6 60 60 12 NORTH 190 c 19 19 190 190 12 NORTH 200 c 20 20 200 200 12 NORTH 363 c 36.3 36 360 0 12 NORTH 113 c 11.3 11 110 110 12 NORTH 110 c 11 11 110 110 12 NORTH 162 s 16.2 16 160 160 12 NORTH 170 s 17 17 170 170 12 NORTH 251 s 25.1 25 250 250 12 NORTH 244 s 24.4 24 240 240 13 NORTH 235 c 23.5 24 240 240 13 NORTH 305 c 30.5 31 310 310 13 NORTH 240 c 24 24 240 240 13 NORTH 180 c 18 18 180 180 13 NORTH 130 s 13 13 130 130 14 NORTH 303 c 30.3 30 300 300 14 NORTH 325 c 32.5 33 330 330 14 NORTH 364 c 36.4 36 360 0 14 NORTH 330 c 33 33 330 330 14 NORTH 90 c 9 9 90 90 14 NORTH 320 c 32 32 320 320 14 NORTH 352 c 35.2 35 350 350 14 NORTH 310 c 31 31 310 310 14 NORTH 340 c 34 34 340 340 14 NORTH 90 c 9 9 90 90

160 14 NORTH 170 c 17 17 170 170 14 NORTH 190 s 19 19 190 190 15 NORTH 237 c 23.7 24 240 240 15 NORTH 270 c 27 27 270 270 15 NORTH 360 c 36 36 360 0 15 NORTH 352 c 35.2 35 350 350 15 NORTH 153 c 15.3 15 150 150 16 NORTH 187 c 18.7 19 190 190 16 NORTH 235 c 23.5 24 240 240 16 NORTH 198 c 19.8 20 200 200 16 NORTH 210 c 21 21 210 210 16 NORTH 60 c 6 6 60 60 16 NORTH 152 c 15.2 15 150 150 16 NORTH 60 c 6 6 60 60 16 NORTH 230 c 23 23 230 230 16 NORTH 180 c 18 18 180 180 16 NORTH 105 c 10.5 11 110 110 16 NORTH 352 c 35.2 35 350 350 16 NORTH 333 c 33.3 33 330 330 16 NORTH 240 s 24 24 240 240 16 NORTH 279 s 27.9 28 280 280 16 NORTH 150 s 15 15 150 150 16 NORTH 320 s 32 32 320 320 16 NORTH 323 s 32.3 32 320 320 17 NORTH 360 c 36 36 360 0 17 NORTH 300 c 30 30 300 300 17 NORTH 275 c 27.5 28 280 280 17 NORTH 340 c 34 34 340 340 17 NORTH 110 c 11 11 110 110 17 NORTH 248 c 24.8 25 250 250 17 NORTH 25 c 2.5 3 30 30 17 NORTH 348 c 34.8 35 350 350 17 NORTH 210 c 21 21 210 210 17 NORTH 190 c 19 19 190 190 17 NORTH 350 c 35 35 350 350 17 NORTH 330 c 33 33 330 330 17 NORTH 143 c 14.3 14 140 140 17 NORTH 80 s 8 8 80 80 17 NORTH 313 s 31.3 31 310 310 17 NORTH 8 s 0.8 1 10 10 18 NORTH 78 c 7.8 8 80 80 18 NORTH 160 c 16 16 160 160 18 NORTH 150 c 15 15 150 150 18 NORTH 80 s 8 8 80 80 18 NORTH 80 s 8 8 80 80 18 NORTH 97 s 9.7 10 100 100 18 NORTH 194 s 19.4 19 190 190 18 NORTH 30 s 3 3 30 30 18 NORTH 135 s 13.5 14 140 140

161 18 NORTH 145 c 14.5 15 150 150 18 NORTH 353 c 35.3 35 350 350 18 NORTH 110 c 11 11 110 110 18 NORTH 90 s 9 9 90 90 18 NORTH 38 s 3.8 4 40 40 19 NORTH 50 c 5 5 50 50 19 NORTH 260 c 26 26 260 260 19 NORTH 190 c 19 19 190 190 19 NORTH 263 c 26.3 26 260 260 19 NORTH 335 c 33.5 34 340 340 19 NORTH 312 c 31.2 31 310 310 19 NORTH 220 c 22 22 220 220 19 NORTH 150 c 15 15 150 150 19 NORTH 60 c 6 6 60 60 19 NORTH 180 c 18 18 180 180 19 NORTH 38 c 3.8 4 40 40 19 NORTH 20 c 2 2 20 20 19 NORTH 140 s 14 14 140 140 19 NORTH 130 s 13 13 130 130 20 NORTH 15 c 1.5 2 20 20 20 NORTH 70 c 7 7 70 70 20 NORTH 165 c 16.5 17 170 170 20 NORTH 312 c 31.2 31 310 310 20 NORTH 60 c 6 6 60 60 20 NORTH 195 c 19.5 20 200 200 20 NORTH 10 c 1 1 10 10 20 NORTH 304 s 30.4 30 300 300 20 NORTH 195 s 19.5 20 200 200 21 NORTH 350 c 35 35 350 350 21 NORTH 313 c 31.3 31 310 310 21 NORTH 190 c 19 19 190 190 21 NORTH 40 c 4 4 40 40 21 NORTH 293 c 29.3 29 290 290 21 NORTH 320 c 32 32 320 320 21 NORTH 105 c 10.5 11 110 110 21 NORTH 170 c 17 17 170 170 21 NORTH 190 s 19 19 190 190 21 NORTH 41 s 4.1 4 40 40 21 NORTH 350 s 35 35 350 350 21 NORTH 350 s 35 35 350 350 22 NORTH 150 c 15 15 150 150 22 NORTH 350 c 35 35 350 350 22 NORTH 340 c 34 34 340 340 22 NORTH 247 c 24.7 25 250 250 22 NORTH 230 c 23 23 230 230 22 NORTH 142 c 14.2 14 140 140 22 NORTH 190 c 19 19 190 190 22 NORTH 73 c 7.3 7 70 70 23 NORTH 160 s 16 16 160 160

162 23 NORTH 165 s 16.5 17 170 170 23 NORTH 140 s 14 14 140 140 24 SOUTH 220 c 22 22 220 220 24 SOUTH 333 c 33.3 33 330 330 24 SOUTH 293 c 29.3 29 290 290 24 SOUTH 208 c 20.8 21 210 210 24 SOUTH 335 c 33.5 34 340 340 24 SOUTH 40 c 4 4 40 40 24 SOUTH 40 c 4 4 40 40 24 SOUTH 285 s 28.5 29 290 290 24 SOUTH 155 s 15.5 16 160 160 24 SOUTH 115 s 11.5 12 120 120 24 SOUTH 335 s 33.5 34 340 340 24 SOUTH 217 s 21.7 22 220 220 24 SOUTH 340 s 34 34 340 340 24 SOUTH 178 s 17.8 18 180 180 24 SOUTH 217 s 21.7 22 220 220 24 SOUTH 180 s 18 18 180 180 25 SOUTH 297 c 29.7 30 300 300 25 SOUTH 40 c 4 4 40 40 25 SOUTH 45 c 4.5 5 50 50 25 SOUTH 130 c 13 13 130 130 25 SOUTH 118 c 11.8 12 120 120 25 SOUTH 190 c 19 19 190 190 25 SOUTH 160 c 16 16 160 160 25 SOUTH 320 c 32 32 320 320 25 SOUTH 113 c 11.3 11 110 110 25 SOUTH 160 s 16 16 160 160 25 SOUTH 190 s 19 19 190 190 25 SOUTH 155 s 15.5 16 160 160 25 SOUTH 320 s 32 32 320 320 25 SOUTH 60 s 6 6 60 60 25 SOUTH 138 s 13.8 14 140 140 26 SOUTH 343 c 34.3 34 340 340 26 SOUTH 187 c 18.7 19 190 190 26 SOUTH 165 c 16.5 17 170 170 26 SOUTH 130 s 13 13 130 130 26 SOUTH 80 s 8 8 80 80 27 SOUTH 222 c 22.2 22 220 220 27 SOUTH 190 c 19 19 190 190 27 SOUTH 273 c 27.3 27 270 270 27 SOUTH 230 c 23 23 230 230 28 SOUTH 306 c 30.6 31 310 310 28 SOUTH 195 c 19.5 20 200 200 28 SOUTH 295 c 29.5 30 300 300 28 SOUTH 32 c 3.2 3 30 30 29 SOUTH 72 c 7.2 7 70 70 29 SOUTH 350 c 35 35 350 350 29 SOUTH 345 c 34.5 35 350 350

163 29 SOUTH 170 c 17 17 170 170 29 SOUTH 190 c 19 19 190 190 29 SOUTH 60 s 6 6 60 60 29 SOUTH 8 s 0.8 1 10 10 29 SOUTH 40 s 4 4 40 40 29 SOUTH 30 s 3 3 30 30 30 SOUTH 30 c 3 3 30 30 30 SOUTH 140 c 14 14 140 140 30 SOUTH 10 c 1 1 10 10 30 SOUTH 310 c 31 31 310 310 30 SOUTH 240 c 24 24 240 240 30 SOUTH 350 c 35 35 350 350 31 SOUTH 185 c 18.5 19 190 190 31 SOUTH 30 c 3 3 30 30 31 SOUTH 320 c 32 32 320 320 31 SOUTH 30 c 3 3 30 30 31 SOUTH 180 c 18 18 180 180 31 SOUTH 227 c 22.7 23 230 230 31 SOUTH 260 c 26 26 260 260 31 SOUTH 215 c 21.5 22 220 220 31 SOUTH 220 c 22 22 220 220 31 SOUTH 62 s 6.2 6 60 60 31 SOUTH 160 s 16 16 160 160 31 SOUTH 150 s 15 15 150 150 31 SOUTH 246 s 24.6 25 250 250 31 SOUTH 66 s 6.6 7 70 70 31 SOUTH 242 s 24.2 24 240 240 31 SOUTH 310 s 31 31 310 310 32 SOUTH 10 c 1 1 10 10 32 SOUTH 139 c 13.9 14 140 140 32 SOUTH 83 c 8.3 8 80 80 32 SOUTH 80 c 8 8 80 80 32 SOUTH 360 c 36 36 360 0 32 SOUTH 153 c 15.3 15 150 150 32 SOUTH 153 c 15.3 15 150 150 32 SOUTH 166 c 16.6 17 170 170 32 SOUTH 310 c 31 31 310 310 32 SOUTH 115 c 11.5 12 120 120 32 SOUTH 300 c 30 30 300 300 32 SOUTH 20 c 2 2 20 20 32 SOUTH 344 c 34.4 34 340 340 32 SOUTH 130 c 13 13 130 130 32 SOUTH 336 c 33.6 34 340 340 32 SOUTH 160 s 16 16 160 160 32 SOUTH 232 s 23.2 23 230 230 32 SOUTH 190 s 19 19 190 190 32 SOUTH 220 s 22 22 220 220 32 SOUTH 200 s 20 20 200 200 32 SOUTH 180 s 18 18 180 180

164 33 SOUTH 160 c 16 16 160 160 33 SOUTH 310 c 31 31 310 310 33 SOUTH 86 c 8.6 9 90 90 33 SOUTH 350 c 35 35 350 350 33 SOUTH 320 s 32 32 320 320 33 SOUTH 33 s 3.3 3 30 30 33 SOUTH 182 s 18.2 18 180 180 34 SOUTH 120 c 12 12 120 120 34 SOUTH 120 c 12 12 120 120 34 SOUTH 190 c 19 19 190 190 34 SOUTH 170 s 17 17 170 170 34 SOUTH 160 s 16 16 160 160 34 SOUTH 140 s 14 14 140 140 34 SOUTH 62 s 6.2 6 60 60 34 SOUTH 243 s 24.3 24 240 240 34 SOUTH 320 s 32 32 320 320 34 SOUTH 200 s 20 20 200 200 35 SOUTH 155 c 15.5 16 160 160 35 SOUTH 83 c 8.3 8 80 80 35 SOUTH 75 c 7.5 8 80 80 35 SOUTH 354 c 35.4 35 350 350 35 SOUTH 11 c 1.1 1 10 10 35 SOUTH 350 c 35 35 350 350 35 SOUTH 170 s 17 17 170 170 35 SOUTH 247 s 24.7 25 250 250 35 SOUTH 142 s 14.2 14 140 140 35 SOUTH 180 s 18 18 180 180 36 SOUTH 267 c 26.7 27 270 270 36 SOUTH 240 c 24 24 240 240 36 SOUTH 170 c 17 17 170 170 36 SOUTH 210 c 21 21 210 210 36 SOUTH 320 c 32 32 320 320 36 SOUTH 138 c 13.8 14 140 140 36 SOUTH 190 s 19 19 190 190 36 SOUTH 10 s 1 1 10 10 36 SOUTH 185 s 18.5 19 190 190 36 SOUTH 170 s 17 17 170 170 37 SOUTH 180 c 18 18 180 180 37 SOUTH 350 c 35 35 350 350 37 SOUTH 143 c 14.3 14 140 140 37 SOUTH 160 c 16 16 160 160 37 SOUTH 33 c 3.3 3 30 30 37 SOUTH 250 c 25 25 250 250 37 SOUTH 120 c 12 12 120 120 37 SOUTH 140 s 14 14 140 140 37 SOUTH 180 s 18 18 180 180 37 SOUTH 170 s 17 17 170 170 37 SOUTH 202 s 20.2 20 200 200 37 SOUTH 160 s 16 16 160 160

165 37 SOUTH 200 s 20 20 200 200 37 SOUTH 160 s 16 16 160 160 37 SOUTH 160 s 16 16 160 160 37 SOUTH 260 s 26 26 260 260 37 SOUTH 355 s 35.5 36 360 0 37 SOUTH 55 s 5.5 6 60 60 37 SOUTH 22 s 2.2 2 20 20 37 SOUTH 190 s 19 19 190 190 37 SOUTH 10 s 1 1 10 10 37 SOUTH 40 s 4 4 40 40

166 Appendix 4: Orientation for East and West areas

Calyx Round or to "10" Set 360 Quadrant Group Orientation stem Calc Round deg to 0 deg 1 West 85 c 8.5 9 90 90 1 West 13 c 1.3 1 10 10 1 West 176 c 17.6 18 180 180 1 West 85 c 8.5 9 90 90 1 West 303 c 30.3 30 300 300 1 West 153 c 15.3 15 150 150 1 West 12 c 1.2 1 10 10 1 West 10 c 1 1 10 10 1 West 210 s 21 21 210 210 1 West 320 s 32 32 320 320 1 West 160 s 16 16 160 160 1 West 335 s 33.5 34 340 340 1 West 5 s 0.5 1 10 10 1 West 240 s 24 24 240 240 1 West 190 s 19 19 190 190 2 West 165 c 16.5 17 170 170 2 West 208 c 20.8 21 210 210 2 West 217 c 21.7 22 220 220 2 West 323 c 32.3 32 320 320 2 West 74 c 7.4 7 70 70 2 West 240 c 24 24 240 240 2 West 230 s 23 23 230 230 2 West 180 s 18 18 180 180 2 West 357 s 35.7 36 360 0 3 West 338 c 33.8 34 340 340 3 West 195 c 19.5 20 200 200 3 West 160 c 16 16 160 160 3 West 355 c 35.5 36 360 0 3 West 263 c 26.3 26 260 260 3 West 90 c 9 9 90 90 3 West 340 c 34 34 340 340 3 West 340 c 34 34 340 340 3 West 360 c 36 36 360 0 3 West 332 c 33.2 33 330 330 3 West 13 c 1.3 1 10 10 3 West 263 s 26.3 26 260 260 3 West 360 s 36 36 360 0 3 West 360 s 36 36 360 0 4 East 230 c 23 23 230 230 4 East 160 c 16 16 160 160 4 East 95 c 9.5 10 100 100 4 East 305 c 30.5 31 310 310 4 East 355 c 35.5 36 360 0 4 East 116 c 11.6 12 120 120

167 4 East 250 c 25 25 250 250 4 East 338 c 33.8 34 340 340 4 East 96 c 9.6 10 100 100 4 East 10 c 1 1 10 10 4 East 270 c 27 27 270 270 4 East 10 c 1 1 10 10 4 East 18 c 1.8 2 20 20 4 East 88 s 8.8 9 90 90 4 East 90 s 9 9 90 90 5 East 210 c 21 21 210 210 5 East 68 c 6.8 7 70 70 5 East 30 c 3 3 30 30 5 East 310 c 31 31 310 310 6 East 190 c 19 19 190 190 6 East 210 c 21 21 210 210 6 East 180 c 18 18 180 180 6 East 192 c 19.2 19 190 190 6 East 240 c 24 24 240 240 6 East 173 c 17.3 17 170 170 6 East 290 c 29 29 290 290 6 East 360 c 36 36 360 0 6 East 343 s 34.3 34 340 340 6 East 180 s 18 18 180 180 6 East 278 s 27.8 28 280 280 7 East 188 c 18.8 19 190 190 7 East 330 c 33 33 330 330 7 East 160 s 16 16 160 160 7 East 130 s 13 13 130 130 7 East 162 s 16.2 16 160 160 7 East 157 s 15.7 16 160 160 8 West 112 c 11.2 11 110 110 8 West 250 c 25 25 250 250 8 West 195 c 19.5 20 200 200 8 West 197 s 19.7 20 200 200 8 West 50 s 5 5 50 50 8 West 362 s 36.2 36 360 0 8 West 150 s 15 15 150 150 9 West 10 c 1 1 10 10 9 West 240 c 24 24 240 240 9 West 20 c 2 2 20 20 9 West 180 c 18 18 180 180 9 West 333 c 33.3 33 330 330 9 West 170 c 17 17 170 170 9 West 158 c 15.8 16 160 160 9 West 10 c 1 1 10 10 9 West 190 c 19 19 190 190 9 West 113 c 11.3 11 110 110 9 West 293 c 29.3 29 290 290 9 West 68 s 6.8 7 70 70

168 9 West 30 s 3 3 30 30 9 West 200 s 20 20 200 200 9 West 138 s 13.8 14 140 140 9 West 190 s 19 19 190 190 9 West 90 s 9 9 90 90 9 West 312 s 31.2 31 310 310 10 West 10 c 1 1 10 10 10 West 33 c 3.3 3 30 30 10 West 116 c 11.6 12 120 120 10 West 350 c 35 35 350 350 10 West 358 c 35.8 36 360 0 10 West 340 c 34 34 340 340 10 West 214 c 21.4 21 210 210 10 West 183 s 18.3 18 180 180 10 West 140 s 14 14 140 140 10 West 40 s 4 4 40 40 10 West 330 s 33 33 330 330 11 West 270 c 27 27 270 270 11 West 345 c 34.5 35 350 350 11 West 138 c 13.8 14 140 140 11 West 255 c 25.5 26 260 260 11 West 40 c 4 4 40 40 11 West 70 c 7 7 70 70 11 West 283 c 28.3 28 280 280 11 West 58 s 5.8 6 60 60 12 East 190 c 19 19 190 190 12 East 200 c 20 20 200 200 12 East 363 c 36.3 36 360 0 12 East 113 c 11.3 11 110 110 12 East 110 c 11 11 110 110 12 East 162 s 16.2 16 160 160 12 East 170 s 17 17 170 170 12 East 251 s 25.1 25 250 250 12 East 244 s 24.4 24 240 240 13 East 235 c 23.5 24 240 240 13 East 305 c 30.5 31 310 310 13 East 240 c 24 24 240 240 13 East 180 c 18 18 180 180 13 East 130 s 13 13 130 130 14 East 303 c 30.3 30 300 300 14 East 325 c 32.5 33 330 330 14 East 364 c 36.4 36 360 0 14 East 330 c 33 33 330 330 14 East 90 c 9 9 90 90 14 East 320 c 32 32 320 320 14 East 352 c 35.2 35 350 350 14 East 310 c 31 31 310 310 14 East 340 c 34 34 340 340 14 East 90 c 9 9 90 90

169 14 East 170 c 17 17 170 170 14 East 190 s 19 19 190 190 15 East 237 c 23.7 24 240 240 15 East 270 c 27 27 270 270 15 East 360 c 36 36 360 0 15 East 352 c 35.2 35 350 350 15 East 153 c 15.3 15 150 150 16 West 187 c 18.7 19 190 190 16 West 235 c 23.5 24 240 240 16 West 198 c 19.8 20 200 200 16 West 210 c 21 21 210 210 16 West 60 c 6 6 60 60 16 West 152 c 15.2 15 150 150 16 West 60 c 6 6 60 60 16 West 230 c 23 23 230 230 16 West 180 c 18 18 180 180 16 West 105 c 10.5 11 110 110 16 West 352 c 35.2 35 350 350 16 West 333 c 33.3 33 330 330 16 West 240 s 24 24 240 240 16 West 279 s 27.9 28 280 280 16 West 150 s 15 15 150 150 16 West 320 s 32 32 320 320 16 West 323 s 32.3 32 320 320 17 West 360 c 36 36 360 0 17 West 300 c 30 30 300 300 17 West 275 c 27.5 28 280 280 17 West 340 c 34 34 340 340 17 West 110 c 11 11 110 110 17 West 248 c 24.8 25 250 250 17 West 25 c 2.5 3 30 30 17 West 348 c 34.8 35 350 350 17 West 210 c 21 21 210 210 17 West 190 c 19 19 190 190 17 West 350 c 35 35 350 350 17 West 330 c 33 33 330 330 17 West 143 c 14.3 14 140 140 17 West 80 s 8 8 80 80 17 West 313 s 31.3 31 310 310 17 West 8 s 0.8 1 10 10 18 West 78 c 7.8 8 80 80 18 West 160 c 16 16 160 160 18 West 150 c 15 15 150 150 18 West 80 s 8 8 80 80 18 West 80 s 8 8 80 80 18 West 97 s 9.7 10 100 100 18 West 194 s 19.4 19 190 190 18 West 30 s 3 3 30 30 18 West 135 s 13.5 14 140 140

170 18 West 145 c 14.5 15 150 150 18 West 353 c 35.3 35 350 350 18 West 110 c 11 11 110 110 18 West 90 s 9 9 90 90 18 West 38 s 3.8 4 40 40 19 West 50 c 5 5 50 50 19 West 260 c 26 26 260 260 19 West 190 c 19 19 190 190 19 West 263 c 26.3 26 260 260 19 West 335 c 33.5 34 340 340 19 West 312 c 31.2 31 310 310 19 West 220 c 22 22 220 220 19 West 150 c 15 15 150 150 19 West 60 c 6 6 60 60 19 West 180 c 18 18 180 180 19 West 38 c 3.8 4 40 40 19 West 20 c 2 2 20 20 19 West 140 s 14 14 140 140 19 West 130 s 13 13 130 130 20 East 15 c 1.5 2 20 20 20 East 70 c 7 7 70 70 20 East 165 c 16.5 17 170 170 20 East 312 c 31.2 31 310 310 20 East 60 c 6 6 60 60 20 East 195 c 19.5 20 200 200 20 East 10 c 1 1 10 10 20 East 304 s 30.4 30 300 300 20 East 195 s 19.5 20 200 200 21 East 350 c 35 35 350 350 21 East 313 c 31.3 31 310 310 21 East 190 c 19 19 190 190 21 East 40 c 4 4 40 40 21 East 293 c 29.3 29 290 290 21 East 320 c 32 32 320 320 21 East 105 c 10.5 11 110 110 21 East 170 c 17 17 170 170 21 East 190 s 19 19 190 190 21 East 41 s 4.1 4 40 40 21 East 350 s 35 35 350 350 21 East 350 s 35 35 350 350 22 East 150 c 15 15 150 150 22 East 350 c 35 35 350 350 22 East 340 c 34 34 340 340 22 East 247 c 24.7 25 250 250 22 East 230 c 23 23 230 230 22 East 142 c 14.2 14 140 140 22 East 190 c 19 19 190 190 22 East 73 c 7.3 7 70 70 23 East 160 s 16 16 160 160

171 23 East 165 s 16.5 17 170 170 23 East 140 s 14 14 140 140 24 West 220 c 22 22 220 220 24 West 333 c 33.3 33 330 330 24 West 293 c 29.3 29 290 290 24 West 208 c 20.8 21 210 210 24 West 335 c 33.5 34 340 340 24 West 40 c 4 4 40 40 24 West 40 c 4 4 40 40 24 West 285 s 28.5 29 290 290 24 West 155 s 15.5 16 160 160 24 West 115 s 11.5 12 120 120 24 West 335 s 33.5 34 340 340 24 West 217 s 21.7 22 220 220 24 West 340 s 34 34 340 340 24 West 178 s 17.8 18 180 180 24 West 217 s 21.7 22 220 220 24 West 180 s 18 18 180 180 25 West 297 c 29.7 30 300 300 25 West 40 c 4 4 40 40 25 West 45 c 4.5 5 50 50 25 West 130 c 13 13 130 130 25 West 118 c 11.8 12 120 120 25 West 190 c 19 19 190 190 25 West 160 c 16 16 160 160 25 West 320 c 32 32 320 320 25 West 113 c 11.3 11 110 110 25 West 160 s 16 16 160 160 25 West 190 s 19 19 190 190 25 West 155 s 15.5 16 160 160 25 West 320 s 32 32 320 320 25 West 60 s 6 6 60 60 25 West 138 s 13.8 14 140 140 26 West 343 c 34.3 34 340 340 26 West 187 c 18.7 19 190 190 26 West 165 c 16.5 17 170 170 26 West 130 s 13 13 130 130 26 West 80 s 8 8 80 80 27 West 222 c 22.2 22 220 220 27 West 190 c 19 19 190 190 27 West 273 c 27.3 27 270 270 27 West 230 c 23 23 230 230 28 East 306 c 30.6 31 310 310 28 East 195 c 19.5 20 200 200 28 East 295 c 29.5 30 300 300 28 East 32 c 3.2 3 30 30 29 East 72 c 7.2 7 70 70 29 East 350 c 35 35 350 350 29 East 345 c 34.5 35 350 350

172 29 East 170 c 17 17 170 170 29 East 190 c 19 19 190 190 29 East 60 s 6 6 60 60 29 East 8 s 0.8 1 10 10 29 East 40 s 4 4 40 40 29 East 30 s 3 3 30 30 30 East 30 c 3 3 30 30 30 East 140 c 14 14 140 140 30 East 10 c 1 1 10 10 30 East 310 c 31 31 310 310 30 East 240 c 24 24 240 240 30 East 350 c 35 35 350 350 31 West 185 c 18.5 19 190 190 31 West 30 c 3 3 30 30 31 West 320 c 32 32 320 320 31 West 30 c 3 3 30 30 31 West 180 c 18 18 180 180 31 West 227 c 22.7 23 230 230 31 West 260 c 26 26 260 260 31 West 215 c 21.5 22 220 220 31 West 220 c 22 22 220 220 31 West 62 s 6.2 6 60 60 31 West 160 s 16 16 160 160 31 West 150 s 15 15 150 150 31 West 246 s 24.6 25 250 250 31 West 66 s 6.6 7 70 70 31 West 242 s 24.2 24 240 240 31 West 310 s 31 31 310 310 32 West 10 c 1 1 10 10 32 West 139 c 13.9 14 140 140 32 West 83 c 8.3 8 80 80 32 West 80 c 8 8 80 80 32 West 360 c 36 36 360 0 32 West 153 c 15.3 15 150 150 32 West 153 c 15.3 15 150 150 32 West 166 c 16.6 17 170 170 32 West 310 c 31 31 310 310 32 West 115 c 11.5 12 120 120 32 West 300 c 30 30 300 300 32 West 20 c 2 2 20 20 32 West 344 c 34.4 34 340 340 32 West 130 c 13 13 130 130 32 West 336 c 33.6 34 340 340 32 West 160 s 16 16 160 160 32 West 232 s 23.2 23 230 230 32 West 190 s 19 19 190 190 32 West 220 s 22 22 220 220 32 West 200 s 20 20 200 200 32 West 180 s 18 18 180 180

173 33 West 160 c 16 16 160 160 33 West 310 c 31 31 310 310 33 West 86 c 8.6 9 90 90 33 West 350 c 35 35 350 350 33 West 320 s 32 32 320 320 33 West 33 s 3.3 3 30 30 33 West 182 s 18.2 18 180 180 34 West 120 c 12 12 120 120 34 West 120 c 12 12 120 120 34 West 190 c 19 19 190 190 34 West 170 s 17 17 170 170 34 West 160 s 16 16 160 160 34 West 140 s 14 14 140 140 34 West 62 s 6.2 6 60 60 34 West 243 s 24.3 24 240 240 34 West 320 s 32 32 320 320 34 West 200 s 20 20 200 200 35 West 155 c 15.5 16 160 160 35 West 83 c 8.3 8 80 80 35 West 75 c 7.5 8 80 80 35 West 354 c 35.4 35 350 350 35 West 11 c 1.1 1 10 10 35 West 350 c 35 35 350 350 35 West 170 s 17 17 170 170 35 West 247 s 24.7 25 250 250 35 West 142 s 14.2 14 140 140 35 West 180 s 18 18 180 180 36 West 267 c 26.7 27 270 270 36 West 240 c 24 24 240 240 36 West 170 c 17 17 170 170 36 West 210 c 21 21 210 210 36 West 320 c 32 32 320 320 36 West 138 c 13.8 14 140 140 36 West 190 s 19 19 190 190 36 West 10 s 1 1 10 10 36 West 185 s 18.5 19 190 190 36 West 170 s 17 17 170 170 37 West 180 c 18 18 180 180 37 West 350 c 35 35 350 350 37 West 143 c 14.3 14 140 140 37 West 160 c 16 16 160 160 37 West 33 c 3.3 3 30 30 37 West 250 c 25 25 250 250 37 West 120 c 12 12 120 120 37 West 140 s 14 14 140 140 37 West 180 s 18 18 180 180 37 West 170 s 17 17 170 170 37 West 202 s 20.2 20 200 200 37 West 160 s 16 16 160 160

174 37 West 200 s 20 20 200 200 37 West 160 s 16 16 160 160 37 West 160 s 16 16 160 160 37 West 260 s 26 26 260 260 37 West 355 s 35.5 36 360 0 37 West 55 s 5.5 6 60 60 37 West 22 s 2.2 2 20 20 37 West 190 s 19 19 190 190 37 West 10 s 1 1 10 10 37 West 40 s 4 4 40 40

175 Appendix 5: Orientation for Sections A, B, C ("Quads")

Calyx or Round to Set 360 Quadrant Group Orientation stem Calc Round "10" deg to 0 deg 9 A 10 c 1 1 10 10 9 A 240 c 24 24 240 240 9 A 20 c 2 2 20 20 9 A 180 c 18 18 180 180 9 A 333 c 33.3 33 330 330 9 A 170 c 17 17 170 170 9 A 158 c 15.8 16 160 160 9 A 10 c 1 1 10 10 9 A 190 c 19 19 190 190 9 A 113 c 11.3 11 110 110 9 A 293 c 29.3 29 290 290 9 A 68 s 6.8 7 70 70 9 A 30 s 3 3 30 30 9 A 200 s 20 20 200 200 9 A 138 s 13.8 14 140 140 9 A 190 s 19 19 190 190 9 A 90 s 9 9 90 90 9 A 312 s 31.2 31 310 310 10 A 10 c 1 1 10 10 10 A 33 c 3.3 3 30 30 10 A 116 c 11.6 12 120 120 10 A 350 c 35 35 350 350 10 A 358 c 35.8 36 360 0 10 A 340 c 34 34 340 340 10 A 214 c 21.4 21 210 210 10 A 183 s 18.3 18 180 180 10 A 140 s 14 14 140 140 10 A 40 s 4 4 40 40 10 A 330 s 33 33 330 330 11 B 270 c 27 27 270 270 11 B 345 c 34.5 35 350 350 11 B 138 c 13.8 14 140 140 11 B 255 c 25.5 26 260 260 11 B 40 c 4 4 40 40 11 B 70 c 7 7 70 70 11 B 283 c 28.3 28 280 280 11 B 58 s 5.8 6 60 60 12 B 190 c 19 19 190 190 12 B 200 c 20 20 200 200 12 B 363 c 36.3 36 360 0 12 B 113 c 11.3 11 110 110 12 B 110 c 11 11 110 110 12 B 162 s 16.2 16 160 160 12 B 170 s 17 17 170 170 12 B 251 s 25.1 25 250 250

176 12 B 244 s 24.4 24 240 240 13 C 235 c 23.5 24 240 240 13 C 305 c 30.5 31 310 310 13 C 240 c 24 24 240 240 13 C 180 c 18 18 180 180 13 C 130 s 13 13 130 130 14 C 303 c 30.3 30 300 300 14 C 325 c 32.5 33 330 330 14 C 364 c 36.4 36 360 0 14 C 330 c 33 33 330 330 14 C 90 c 9 9 90 90 14 C 320 c 32 32 320 320 14 C 352 c 35.2 35 350 350 14 C 310 c 31 31 310 310 14 C 340 c 34 34 340 340 14 C 90 c 9 9 90 90 14 C 170 c 17 17 170 170 14 C 190 s 19 19 190 190 17 A 360 c 36 36 360 0 17 A 300 c 30 30 300 300 17 A 275 c 27.5 28 280 280 17 A 340 c 34 34 340 340 17 A 110 c 11 11 110 110 17 A 248 c 24.8 25 250 250 17 A 25 c 2.5 3 30 30 17 A 348 c 34.8 35 350 350 17 A 210 c 21 21 210 210 17 A 190 c 19 19 190 190 17 A 350 c 35 35 350 350 17 A 330 c 33 33 330 330 17 A 143 c 14.3 14 140 140 17 A 80 s 8 8 80 80 17 A 313 s 31.3 31 310 310 17 A 8 s 0.8 1 10 10 18 A 78 c 7.8 8 80 80 18 A 160 c 16 16 160 160 18 A 150 c 15 15 150 150 18 A 80 s 8 8 80 80 18 A 80 s 8 8 80 80 18 A 97 s 9.7 10 100 100 18 A 194 s 19.4 19 190 190 18 A 30 s 3 3 30 30 18 A 135 s 13.5 14 140 140 18 A 145 c 14.5 15 150 150 18 A 353 c 35.3 35 350 350 18 A 110 c 11 11 110 110 18 A 90 s 9 9 90 90 18 A 38 s 3.8 4 40 40 19 B 50 c 5 5 50 50

177 19 B 260 c 26 26 260 260 19 B 190 c 19 19 190 190 19 B 263 c 26.3 26 260 260 19 B 335 c 33.5 34 340 340 19 B 312 c 31.2 31 310 310 19 B 220 c 22 22 220 220 19 B 150 c 15 15 150 150 19 B 60 c 6 6 60 60 19 B 180 c 18 18 180 180 19 B 38 c 3.8 4 40 40 19 B 20 c 2 2 20 20 19 B 140 s 14 14 140 140 19 B 130 s 13 13 130 130 20 B 15 c 1.5 2 20 20 20 B 70 c 7 7 70 70 20 B 165 c 16.5 17 170 170 20 B 312 c 31.2 31 310 310 20 B 60 c 6 6 60 60 20 B 195 c 19.5 20 200 200 20 B 10 c 1 1 10 10 20 B 304 s 30.4 30 300 300 20 B 195 s 19.5 20 200 200 21 C 350 c 35 35 350 350 21 C 313 c 31.3 31 310 310 21 C 190 c 19 19 190 190 21 C 40 c 4 4 40 40 21 C 293 c 29.3 29 290 290 21 C 320 c 32 32 320 320 21 C 105 c 10.5 11 110 110 21 C 170 c 17 17 170 170 21 C 190 s 19 19 190 190 21 C 41 s 4.1 4 40 40 21 C 350 s 35 35 350 350 21 C 350 s 35 35 350 350 22 C 150 c 15 15 150 150 22 C 350 c 35 35 350 350 22 C 340 c 34 34 340 340 22 C 247 c 24.7 25 250 250 22 C 230 c 23 23 230 230 22 C 142 c 14.2 14 140 140 22 C 190 c 19 19 190 190 22 C 73 c 7.3 7 70 70

178 Appendix 6: Distal coil diameter

coil slab diameter J 2.43 J 5.8 J 4 J 3.3 B 2.1 B 2.4 B 0 B 4.5 A 2.5 A 3.7 A 3.1 C 0.9 C 3.8 I 3.4 I 3.6 I 2.6 I 7.6 I 6.6 I 7.3 I 5.2 I 3.6 I 6.8 I 5.4 I 4.6 I 4.3 I 2.1 I 4.8 I 2.9 I 2.8 I 5.8 I 4.2 I 7.7 I 5.1 I 2.9 I 5.8 I 1.6 I 6.1 I 5.8 I 4.8 E 2.5 E 5.5 E 4.3 E 7.7 E 8.1 E 4.2

179 E 3.2 E 3.9 E 4.9 E 4.1 E 3.4 T 6.3 T 7.4 T 6.2 K 3.4 K 4.8 K 4.5 K 3.4 K 4.4 K 2.7 K 5.2 K 3.2 K 1.6 K 3.2 K 6.71 Average 4.3241 Standard Deviation 1.7868

180