An Actualistic, CT-Based Study in Taphonomy

Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy

A dissertation presented to

the faculty of

the College of Arts and Sciences of Ohio University

In partial fulfillment

of the requirements for the degree

Doctor of Philosophy

Joseph C. Daniel

August 2012

This dissertation titled

Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy

JOSEPH C. DANIEL

has been approved for

the Department of Biological Sciences

and the College of Arts and Sciences by

Lawrence M. Witmer

Professor of Biomedical Sciences

Benjamin M. Ogles

Dean, College of Arts and Sciences 3

ABSTRACT

DANIEL, JOSEPH C., Ph.D., August 2012, Biological Sciences

Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy

Director of Dissertation: Lawrence M. Witmer

Reconstruction of extinct animals is hampered by lack of soft-tissue preservation.

To test the hypothesis that sediment interacts with soft tissue during burial in predictable ways that may be of use in reconstructions, 29 ostrich heads in fresh, desiccated, rotten, or cleaned conditions were buried in two flumes; a short, deep-water flume with slow- moving water and a shallow-water flume with fast-moving water, designed to emulate common burial conditions in alluvial systems. After an initial CT scan, two heads from each setup (16 total) were reburied for seven months and then rescanned. Sediment was examined to confirm patterns detected by CT. Primary void space was retained in all conditions, especially within the tympanic recesses, increasing after prolonged burial, allowing crude estimates of burial conditions to be predicted. Sediment drapes overlay fresh and rotten heads in the shallow flume and to a lesser extent desiccated heads, but only rotten heads developed any drapes in the deep flume. Decompositional outgassing only partially disrupted external patterns in long-term-burial scans. Most internal soft- tissue patterns were obliterated by decomposition, although rostral conchae and auditory canals were still visible. Fat readily became adipocere, which is highly resistant to decay and can potentially be preserved as sediment traces. The best patterns were preserved in rotten or desiccated heads. These data indicate sediment patterns may be preserved in fossils that may reflect soft tissue and that may help to refine interpretations of fossils, 4 although not enough for detailed information for reconstructions. Void space origins and taphonomic implications for fossil preservation may be determined. Reassessments of published fossils include the preserved intestinal track of Scipionyx as a potential impacted bowel, preservation of a dinosaur “heart” is contraindicated, and possible conchae within Panoplosaurus. The presence of feathers on Sinornithosaurus and

Sinosauropteryx is supported, but not strongly, and the chance of preserved internal organs in dinosaur mummies is low, although epidermal tissues and peripheral structures are high. Questions remain why some fossils show exceptional preservation while others in similar situations do not.

Approved: ______

Lawrence M. Witmer

Professor of Biomedical Sciences

This work is dedicated to my wife Katharyn, without whose unflagging faith in me— even

when I didn’t—and support this dissertation would never have been completed; and my

children, Lorelai and Griffin, who provided the inspiration, hugs, and smiles to get me through. I would also like to dedicate this dissertation to my mother, for her gift of books

and summers in the library; and my father, for seeing that I spent my youth in college,

but did not live to see me finish.

Today there remain but a few small areas on the world's map unmarked by explorers'

trails. Human courage and endurance have conquered the Poles; the secrets of the

tropical jungles have been revealed. The highest mountains of the earth have heard the

voice of man. But this does not mean that the youth of the future has no new worlds to

vanquish. It means only that the explorer must change his methods.

—Roy Chapman Andrews, 1926

ACKNOWLEDGMENTS

I am indebted to my advisor, Lawrence Witmer, for his guidance and aid throughout the project on both the science and the sociopolitical process of the career. I appreciate his patience with my idiosyncrasies and the roller-coaster life experiences as I dealt with the births, deaths, and other life experiences, both expected and unexpected, during my time at Ohio University. Thanks also to my supportive and unusually patient committee members, Patrick O’Connor, Susan Williams, and Greg Nadon. Thanks for the help and advice provided by Nancy Stevens and Don Miles and other faculty members that provided assistance. I would also like to thank Willem Roosenberg for serving on my comprehensive exam committee and, when asked later why he avoided certain topics, answering, “Because I already knew you knew all that.”

I thank Dr. Gar Rothwell (OU Environmental & Plant Biology) for advice, assistance, and access to his thin-sectioning equipment and masonry saw and Dr. Mark

Stoy of the Ohio Coal Research Center for advice on and use of the Malvern Mastersizer

2000. Special thanks go to Ryan Ridgely (graphics guru), and Casey Holliday, Tobin

Hieronymus, Dave Dufeau, and Justin Tickhill of the Witmer lab for assistance in many aspects of the project. Thanks go to Angie Nielson for knowing everything about how to navigate the OU bureaucracy and getting things done.

This work would not have been possible without access to the extensive collection of CT scans made available to me by Lawrence Witmer, for which I am very grateful. For loan of specimens for CT scanning and/or for providing existing CT datasets, I thank

Mark Norell & Carl Mehling (American Museum); Michael Ryan, Bruce Latimer, & the 7 late Mike Williams (Cleveland Museum); and Matt Lamanna (Carnegie Museum).

Thanks to Heather Rockhold, RT(CT), O’Bleness Memorial Hospital (Athens, OH), Ron

Beshears (NASA Marshall Space Flight Center), and Tim Rowe & Rich Ketcham

(UTCT, Austin) for CT scanning.

Support for this work was provided by grants from Sigma Xi, the Jurassic

Foundation, the Department of Biological Sciences and the Graduate Student Senate at

Ohio University, as well as National Science Foundation awards IBN-9601174, IBN-

0343744, IOB-0517257 to LMW.

Finally, I would like to thank my family for all their help. My wife, Katharyn, provided support and encouragement when I hit the inevitable moments of despair. My kids, Lorelai and Griffin, provided the joy and inspiration to see it through. I am indebted to my in-laws, John and Shannon Chamberlin for making all of this even possible and encouraging me—and so many others— to “follow your bliss.”

TABLE OF CONTENTS

Page

Abstract ...... 3 Acknowledgments...... 6 List of Tables ...... 9 List of Figures ...... 10 Chapter 1: Void space within buried ostrich heads and its implications for the interpretation of fossil patterns: an experimental, actualistic, CT-based study in taphonomy...... 13 Chapter 2: Sediment/carcass interactions at the integument and their implications for fossil interpretation: A CT-based actualistic taphonomy study ...... 53 Chapter 3: The utility of CT-based, taphonomic study of sediment/carcass interactions for soft-tissue reconstruction of extinct organisms...... 95 Chapter 4: New interpretations of fossils based on CT-based, actualistic taphonomy studies ...... 158 References ...... 209 Appendix A: Grain-size estimation techniques ...... 230 Appendix B: Grain-size estimation in this study ...... 234 Appendix C: Grain-size data ...... 240 Appendix D: Thigh circumference estimation ...... 343

LIST OF TABLES

Page

Table 1: Ohio University Vertebrate Collections (OUVC) specimens of ostrich used in this study, arranged by treatment condition for short-term burial set ...... 51

Table 2: Standard flume sediment mixture ...... 51

Table 3: Statistical summary of void space comparisons ...... 52

Table 4: Grain-size categories for sediment distribution estimates...... 94

Table 5: MANOVA results using full data set from both flumes ...... 94

Table 6: Endpoints for original void space within carcasses...... 207

Table 7: Institutional abbreviations ...... 208

LIST OF FIGURES

Page

Figure 1: Anatomy of the head of Struthio camelus ...... …42

Figure 2: Flumes used to bury ostrich heads ...... …43

Figure 3: Potential fluvial depositional environments emulated by this study ...... 44

Figure 4: Methods diagram ...... 45

Figure 5: Void space measured in buried ostrich heads ...... 46

Figure 6: Void space within heads by group ...... 47

Figure 7: Reconstructions of void space seen in short-term burial CT scans ...... 48

Figure 8: Reconstructions of void space seen in long-term-burial CT scans ...... 49

Figure 9: CT images showing disruption haloes of gas bubbles around orbits and nasals . ………………………………………………………………………………………….50

Figure 10: Flumes used to bury ostrich heads ...... 85

Figure 11: Line probe analysis example for OUVC 10500 ...... 86

Figure 12: CT scans of sediment blocks taken from the deep flume and immediately frozen...... 87

Figure 13: Position of sloughed integument on a rotten head buried in the shallow flume (OUVC 10475)...... 88

Figure 14: CT scans of sediment blocks taken from the shallow flume and immediately frozen ...... 90

Figure 15: CT scans of sediment blocks taken from the deep flume after seven months.91

Figure 16: CT scans of sediment blocks taken from the shallow flume after seven months ...... 93

Figure 17: Simplified chart of taphonomic factors and their interrelationships ...... 134

Figure 18: Visual confirmation of CT scan sediment patterns ...... 135 11

Figure 19: CT density variations can distinguish sediment grain-size parameters ...... 137

Figure 20: Sediment patterns in fresh heads buried in the deep flume ...... 138

Figure 21: Short-term sediment patterns in desiccated heads buried in the deep flume.141

Figure 22: Sediment patterns within rotten heads buried in the deep flume ...... 143

Figure 23: Sediment patterns of clean skulls ...... 144

Figure 24: Sediment patterns within fresh heads buried in the shallow flume ...... 147

Figure 25: Variation of short-term sediment patterns seen in CT slices of desiccated heads buried in the shallow flume...... 149

Figure 26: Short-term sediment patterns seen in CT slices of rotten heads buried in the shallow flume ...... 151

Figure 27: Variation in desiccated heads buried in the deep flume after seven months seen in CT slices...... 152

Figure 28: CT slices from desiccated heads buried in the shallow flume after seven months ...... 1534

Figure 29: Pneumatic sinuses that collected sediment ...... 156

Figure 30: Axial slices through the nasal passage of OUVC 10475, rotting head in shallow flume, showing preservation of conchae in sediment ...... 157

Figure 31: Hypothetial spectrum of preservability for different tissues ...... 194

Figure 32: Void space in Tyrannosaurus rex (FMNH PR 2081)...... 195

Figure 33: Braincase of Allosaurus (UMNH VP 18050)...... 196

Figure 34: Metallic infills within the premaxilla of Allosaurus (UMNH VP 18046). .197

Figure 35: CT scan of Nanotyrannus (CMNH 7541)...... 198

Figure 36: CT of Hypacrosaurus premaxilla (ROM 702)...... 199

Figure 37: Tyrannosaurus rex (AMNH 5027) as figured by Osborn 1912 ...... 199

Figure 38: A: CT reconstruction of a juvenile Tarbosaurus bataar MPC-D 107/7. B: Ikechosaurus (adapted from Liu and Wang 2008)...... 200

Figure 39: Adipocere formed on buried, rotting ostrich head...... 201

Figure 40: Possible conchal sediment traces in CT reconstructions of Panoplosaurus (ROM 1215) ...... 202

Figure 41: Monjurosuchus. Photo: IVPP modified from Liu and Wang (2008)...... 203

Figure 42: Psittacosaurus bed buried in lahar deposit (modified from Qi, et al. 2007). ………………………………………………………………………………………...204

Figure 43: People preserved as casts by ash fall at Pompeii...... 205

Figure 44: A child’s skeleton preserved by pyroclastic density flows at Herculaneum …………………………………………………………………………………………206

CHAPTER 1: VOID SPACE WITHIN BURIED OSTRICH HEADS AND ITS

IMPLICATIONS FOR THE INTERPRETATION OF FOSSIL PATTERNS: AN

EXPERIMENTAL, ACTUALISTIC, CT-BASED STUDY IN TAPHONOMY

Abstract

As part of an experimental, actualistic study on sediment/carcass interactions and their implications for fossil interpretation, patterns of void spaces within buried heads were investigated. Using two flumes designed to emulate deep, slow-moving water and shallow, fast-moving water, 29ostrich heads in four conditions (fresh, rotting, desiccated, clean skull) were buried. The sediment blocks containing the heads were then subjected to computed tomographic (CT) scanning. Sixteen heads were then reburied and CT- scanned after seven months. A substantial amount of void space was retained in all conditions, which was contributed to by gases released by decomposition, particularly in the fresh and rotting heads. Bony recesses were largely gas-filled even in clean skulls.

Sediment collapse after soft-tissue decomposition was limited, most notably affecting the narial region and, to a lesser extent, the orbits. Sediment collapse created a halo effect of disrupted sediment around the collapsed regions. These experiments indicate that void spaces in fossils are likely primary and not due to mineral dissolution, and these spaces might contain trapped gaseous content from the time of burial. Specific patterns of void spaces may also be useful in evaluating taphonomic conditions during burial and therefore provide additional insight into local ecology.

Introduction

When most researchers think of air spaces within fossils, they are usually trying to reconstruct air ways within the nasal cavities or pneumatic sinuses within the bones (e.g.,

Bruner and Manzi, 2002; Witmer et al., 2008). These spaces can be important for aiding interpretations of lung design and thermoregulatory physiology (e.g., Ruben et al., 1999;

O’Connor and Claessens, 2005; Sereno et al., 2008). The other main use of air spaces or, more properly, voids is in providing easily distinguished boundaries in computed tomographic (CT) slices for the actual anatomy of interest, because the radiographic density of air is radically different from that of bone and rock and so usually forms a clear edge on CT scans (Tate and Cann, 1982; Zollikofer and Ponce De León, 2005;

Dierick et al., 2007). Generally, however, voids within fossils are simply acknowledged as being present (e.g., Rühli et al., 2004; Straight et al., 2009) or, more often, ignored

(e.g., Brochu, 2003). Martill et al. (2000) may be unique in attempting to interpret a void space outside of a bony recess anatomically, speculating that the void space represented a postpubic air sac.

There have been no published studies examining voids within vertebrate fossils.

Thus, little is known about their preservation in the fossil record. What patterns they might exhibit in terms of size and location within individual fossils has not been studied.

Their mode of origin is equally unknown. It is possible that they formed from secondary dissolution of previously deposited material, or they could be primary in origin, existing since the time of burial. 15

Such studies have not been done because the techniques have been present until relatively recently. A study of void spaces in vertebrates requires a readily accessible means of noninvasive, nondestructive three-dimensional imaging. Voids in fossils could be studied by serial grinding and other physical methods, but these methods are impractical for any sort of reasonable sample size due to the time-consuming and destructive nature of the methods. Better visualization techniques are now possible, with

CT scanning being the tool of choice (Zollikofer and Ponce De León, 2005). An understanding of the presence of voids in fossils begins with actualistic, experimental taphonomic studies utilizing CT and requiring specialized, dedicated apparatus. It also requires software capable of mapping the patterns three-dimensionally.

Even with the recent advent of these tools and their availability to researchers, voids have not been examined in detail. Research thus far has been focused on the remains themselves, with imaging software being used to extricate fossils from the surrounding matrix. Little attention has been paid to the rock matrix itself and what information it may hold about the fossil it contains. As a result, the implications of void spaces within fossils are unclear. They have only recently begun to be studied in forensic postmortem studies (Thali et al., 2003; Jackowski et al., 2007; Heng et al., 2008;

O’Donnell and Woodward, 2008). Specific to paleontology, void spaces may have implications for preservation potential both during and after fossilization. The trapped gases inside the voids may also have potential for geochemical analyses, dependent on the nature of their formation, development, and preservational potential. 16

This article explores the origin and pattern of void spaces using an experimental, actualistic taphonomic approach, using CT to map void spaces within buried heads under a variety of conditions. Void spaces were examined to determine their relationship to the amount and condition of soft tissue present during burial. The overall amount of void space and the patterns relating to their placement were examined to determine possible taphonomic filters related to burial and to explore what implications they may have for fossil interpretation.

Materials and Methods

Struthio camelus (ostrich) heads were provided by a commercial dealer. The animals had been euthanized by electrical stunning to the head, with minimal to no damage to most specimens, after which the heads were removed and frozen. All heads were essentially the same size and age (young adults). Struthio was chosen for its large size, increasing the potential for sediment to infiltrate the head as well as making any patterns easier to observe. Due to the relative reduction of bone in Struthio heads, soft tissue influences compartmentation in Struthio heads more so than in mammals, and thus soft tissue should have a correspondingly large effect on water flow and sediment patterns. Struthio heads are also exceptionally pneumatic (Fig. 1), with extensive sinuses in both the soft tissue and bone, providing many potential areas for sediment to be deposited (Witmer and Ridgely, 2008).

To examine how various states of decomposition affected sediment patterns, 29 heads were buried in four “state” conditions: fresh, rotten, desiccated, and clean (Table 17

1). The fresh set was buried immediately after thawing. The rotten set was allowed to decompose in a water tank for nine days before burial. The desiccated set was placed in a fume hood under a sun lamp at ~45°C for 7–10 days until there was no outward evidence of water (although this did not preclude trace water inside the skull). The clean set was scavenged by dermestid beetles until all available soft tissue was removed (ligamentous connections between the bones were still intact).

Heads in all four states were buried individually in two flumes built in the lab for this project (Fig. 2). One flume was designed to emulate burial in slow-moving deep water, such as at the entrance of a pool. The flume was a 60 cm cube, in which water flowed from a trough at the top of one side so that it flowed along the entire edge. The outlet was at the top of the far wall. The water was recirculated through ¾” (~19 mm) pipes with a 90 gal/h (~340 l/h) aquarium pump. The second flume was designed to emulate burial on a sandbar deposit of a river (Fig. 3). It was 240 cm in length, with the water outlet at the bottom and recirculated with a 2400 gal/h (~9085 l/h) pump through

1.5” (~38 mm) pipes so that water level was kept shallow and fast-moving. All water used in the experiment was tap water at room temperature (in this case, 28°C). A 4”

(~102 mm) barrier was placed just before the outlet so that sediment collected on the heads, which were placed just before the barrier. All burials used a consistent heterogeneous mixture of sediment ranging from clay to 2 mm sand, with fine to medium sand dominating (Table 2). Sediment was derived from a mixture of Quikrete All-

Purpose Sand, Lowe’s Play Sand, and clay-rich sediment from Good Earth Garden

Center. All sediment was sieved to eliminate particles larger than 2 mm and regularly 18 analyzed by sieving to maintain consistency. The experiment was stopped when the composition of Quikrete changed beyond the parameters of the previous sediment.

All heads were placed into the flumes dorsal-side up and facing upstream. Initial tests with a fresh head and two models indicated this was the most hydrodynamically stable position for the head (sans neck). Heads positioned ventral-side up facing upstream tended to flip backwards and those facing downstream tended to roll, except under very low flow conditions. In some conditions of high velocity but low water-depth, those heads that were dorsal-side up and facing upstream sometimes flipped backwards, as well, but, once moving, never settled on a stable position. Many of the heads could not be sunk because of their inherent buoyancy, so, to maintain consistency, all heads were held in place by a rubber band. Although this means of restraint was imposed of necessity and for reasons of experimental standardization, it replicates natural conditions whereby real heads would not commonly be so mobile, being attached to bodies and/or being restrained by natural debris (e.g., branches).

The initial burial took one to two days, after which the heads were removed from the flumes by pushing a plastic box through the sediment around the head and removing the sediment block en masse with as little disruption of the surrounding sediment as possible. The blocks were then promptly frozen for subsequent CT scanning (described below). Some of these blocks were not subjected to further burial, and this set is referred to here as the “Short-Term Burial” set. However, to examine the effects of further decomposition after burial, two blocks from each set were stored at room temperature for seven months, comprising the “Long-Term Burial” set. The blocks were placed in four 19 boxes and packed with sediment. The boxes were then covered to prevent scavenging and to retain moisture. A small hole was placed approximately 1 cm from the bottom to allow water to pass through the sediment. To keep the sediment wet, sufficient water was added twice a week until water remained pooled on the surface and was added shortly before termination of the experiment. Sediment was added as needed to ensure the heads remained completely buried. After seven months, the blocks were removed and frozen.

The frozen blocks were CT scanned at O’Bleness Memorial Hospital, Athens,

Ohio, using a GE LightSpeed Ultra Multislice scanner with the extended Hounsfield option at a slice thickness of 1.25 mm, 120 kV, 350 mA and a bow-tie head filter. All heads were scanned after initial removal from the flumes. The heads kept for long-term burial were CT scanned again after the end of the seven months. After scanning, the blocks were then sliced into cross-sections approximately 2–3 cm thick (Fig. 4). Limited scanner time prevented acquiring pretreatment scans for all the heads, so three additional fresh heads were CT scanned without burial as controls to provide data on the normal void space in live ostrich heads for comparison.

CT scans were analyzed using the software program Amira (ver. 3.1, 4.1, and

5.0). Radiodensity values typically formed a trimodal distribution, with peaks for void space, soft tissue, and sediment. Radiodensity is measured in CT scans using Hounsfield

Units (HU), with air at standard temperature and pressure defined as -1000 HU and distilled water as 0 HU. Bone typically fell into the upper soft-tissue peak. To measure void space volumes within the heads, each head was outlined and a threshold value set to map the density values corresponding to void space. Because measured volume is 20 affected by the threshold value, the threshold value for all scans was set at -530

Hounsfield units (HU). This value was determined by recording the range of minimal density values between the air and soft-tissue peaks for each scan, and then taking the value that fell into the range of the majority of heads. Only four scans had values falling outside -530 HU. For those, the most extreme value of the minimal range between void space and soft tissue that was closest to -530 HU was used.

All skulls used in the study are archived in the Ohio University Vertebrate

Collection (OUVC) and are available from Lawrence Witmer, as are the CT-scan data generated during the study.

Anatomical abbreviations used in figures: airway = main nasal airway (respiratory region of the nasal cavity); antorb = antorbital sinus; art = articular sinus (tympanic sinus); cho = choanae; con = conchal spaces in the airway; ctym = caudal tympanic sinus; dtym = dorsal tympanic sinus; endo = endocranial cavity; eth = ethmoidal portion of fronto-ethmoidal sinus; fmag = foramen magnum; fr = frontal portion of fronto- ethmoidal sinus; ics = inferior conjunctival sac; lac = lacrimal sinus proper (from suborbital sinus); mes = mesethmoidal portion of fronto-ethmoidal sinus; nar = narial opening; olf = olfactory region of the nasal cavity; oral = oral cavity; orbit = orbit or eye socket; palp = palpebral fissure; para = sinus within parasphenoid rostrum (from rostral tympanic sinus); premaxilla = premaxilla; pter = pterygoid sinus (from suborbital sinus); quad = quadrate sinus (tympanic sinus); ros = rostral tympanic sinus; rtym = rostral tympanic sinus; sub = suborbital sinus (from antorbital sinus); tymp = main middle ear cavity and paratympanic sinuses. 21

Results

Short-term burial set

An analysis of the CT scan data showed that void spaces were present in all of the heads. Although there was no significant difference overall between the deep and shallow flumes, the effects due to decompositional state both within and between the two flow regimes varied considerably, with the heads in the shallow flume showing much more variability between decompositional states than those in the deep flume (Figs. 5, 6).

Despite having no soft tissue, even the clean skulls retained void spaces within the paratympanic and caudal fronto-ethmoidal sinuses (terminology following Witmer, 1990;

Witmer and Ridgely, 2008). Whereas the fresh heads buried in the shallow flume had the most void space on a consistent basis, the rotten heads buried in either flume and the desiccated heads buried in the shallow flume retained extensive void space as long as the orbits remained sealed against the intrusion of sediment.

Fresh heads

Deep flume.—In the deep flume, the fresh heads preserved void space in the rostral oral cavity and, to a lesser extent, in the nasal cavity, chiefly in the olfactory region (Fig. 7). Both the paratympanic and paranasal sinuses retained considerable quantities of void space. Substantial amounts of void space were retained in the fronto- ethmoidal sinuses. The antorbital sinus usually contained some void space, but the suborbital sinuses only had void space in one head, except for the lacrimal sac, which 22 was typically gas-filled. The mesethmoid did not retain void space, probably due to the opening lying at the top of the sinus, although there were typically a few small gas bubbles trapped in the soft tissue adjacent to the mesethmoid. The dorsal tympanic diverticula retained void space in all cases, with void space usually extending through the caudal tympanic, articular, and quadrate diverticula. The rostral tympanic diverticula within the parasphenoid rostrum retained void space in one head, but were almost completely fluid-filled in the others, and the rest of the sphenoid region was fluid-filled in all of the heads. Small amounts of void space were also trapped under the eyelids.

Shallow flume.—In the shallow flume, fresh heads retained only minor void space in the oral cavity, but retained large quantities within the nasal cavity, with the olfactory and most of the respiratory regions being gas-filled. The paranasal and paratympanic sinuses were mostly gas-filled. The fronto-ethmoidal and antorbital sinuses were gas- filled, although the suborbital sinuses were not. Unlike the deep flume, the mesethmoid was gas-filled, as were the rostral tympanic diverticula and almost all the rest of the paratympanic sinuses. Void space was also retained under the eyelids and within the orbit.

Desiccated heads

Deep flume.—The desiccated heads buried within the deep flume did not retain much void space within the oral or nasal cavity proper. The antorbital and suborbital sinuses were mostly fluid-filled. The caudal fronto-ethmoidal sinuses were gas-filled to varying extents, but always contained void space in the caudal-most regions. The 23 paratympanic sinuses always contained void space, chiefly in the dorsal and caudal tympanic diverticula, with a variable amount in the quadrate and articular diverticula. The orbits often held considerable void space, originating either from replacement of fluid in the eye with gas during desiccation or from the rupture of the suborbital and antorbital sinuses, which then became trapped in the dorsal orbital concavity. The endocranial cavities were mostly fluid-filled, but did retain up to one third of its volume in void space. Other soft-tissue regions also showed numerous gas bubbles, particularly in the major jaw muscles.

Shallow flume.—The desiccated heads buried in the shallow flume had a bimodal distribution, with two heads only slightly more void space than clean skulls, retaining around 12 cc, and the other two heads retaining the most void space of any of the heads in the short-term burial group at approximately 140 cc. The head with the most void space had an almost 16-fold increase over the head with the smallest amount. The difference between the two can be attributed mostly to the integrity of the eyelid, which, when intact, allowed the head to retain most of its void space, but once compromised, allowed most of the void space in the head to be replaced by water.

In the two desiccated heads buried in the shallow flume (Ohio University

Vertebrate Collections [OUVC] 10465 and 10466) that retained little void space, the void space that was retained was primarily in the fronto-ethmoidal sinuses, extending in some cases into the lacrimal sacs, and the dorsal paratympanic diverticula. The other paranasal sinuses were primarily fluid-filled, although gas did collect in the dorsal-most portion of the mesethmoid, extending into the olfactory recess of the nasal cavity. Of the remaining 24 paratympanic diverticula, the caudal tympanic diverticula were partially gas-filled, and the quadrate diverticula contained major quantities of void space, whereas the articular diverticula were primarily fluid-filled, as were the rostral tympanic diverticula.

In the two desiccated heads buried in the shallow flume (OUVC 10510 and

10511) that retained a lot of void space, the eyelid maintained a successful seal against gas loss, which is the likely explanation for substantial gas retention. The nasal cavities were almost completely gas-filled, along with all the paranasal sinuses, including the antorbital and suborbital sinuses. The paratympanic sinuses were likewise gas-filled, with void space filling the dorsal tympanic, caudal tympanic, articular, and quadrate diverticula and most of the rostral tympanic diverticula. The orbits were mostly gas- filled, with void space within and around the eye, both medially and laterally, and with void space under the eyelids. The endocranial cavities were mostly gas-filled. The oral cavities, however, were mostly fluid-filled, except for minor bubbles in the rostral tip of the beak and under the tongue.

Rotten heads

Deep flume.—The rotten heads had the most void space of any decompositional state within the deep flume. The four heads also showed a strongly bimodal distribution, with two heads retaining almost 60 cc of void space, whereas the other two heads retained twice as much. In all of the heads, extensive small bubbles appeared throughout the soft tissue, giving it a riddled, spongy appearance. It appears that even though the 25 heads underwent the same amount of time for decomposition, the rates of decomposition differed in areas.

In two of the rotten heads buried in the deep flume (OUVC 10472 and 10473), the nasal cavities were mostly fluid-filled, except for a very thin layer of void space along the top of the nasal cavity in the respiratory region and a small pocket of void space above the ossified portion of the mesethmoid in the olfactory recess. In addition, there were several small bubbles within the soft tissue lining the cavities, with water filling the antorbital and suborbital sinuses. The mesethmoid itself was partially gas-filled, but the void space was not coalesced into a single space, but rather riddled throughout the mesethmoid. Gas filled the caudal portions of the fronto-ethmoidal sinuses, but did not extend into the rostral portions; the lacrimal sacs were fluid-filled. The air within the oral cavity was also dispersed into small bubbles rather than being coalesced to any great degree. Of the paratympanic sinuses, the dorsal tympanic diverticula were gas-filled, but the caudal tympanic diverticula were mostly fluid-filled. The parasphenoid rostrum demonstrated a similar pattern as the mesethmoid, having a riddled appearance. The articular and quadrate diverticula were partially gas-filled, although the articular tended to have more void space than the quadrate diverticula. There were also sizable gas bubbles within the joint capsule itself. The orbit contained gas bubbles mostly between the eye and the medial fat layer within the orbit. Water did not appear to enter the endocranial cavity, but the brain tissues were shrunken. Gas filled 25–35% of the endocranial cavity and gas bubbles were dispersed within the brain itself. 26

In the other two rotten heads in the deep flume (OUVC 10499 and 10500), the nasal cavities were mostly gas-filled, even within the respiratory regions. The paranasal sinuses were almost completely gas-filled, including the lacrimal sacs. The paratympanic sinuses were often fluid-filled, however. The dorsal tympanic diverticula were partially gas-filled, but the caudal tympanic diverticula were mostly fluid-filled, as were the quadrate and articular diverticula. The parasphenoid rostrum was partially or mostly fluid-filled. Nevertheless, there were gas bubbles present in all diverticula. Gas bubbles were also present in the basisphenoid and within the joint capsules. The oral cavities retained significant quantities of void space, with gas bubbles extending into the trachea as well (while much of the gas within the oral cavity is likely original air, the tracheal gas is likely trapped decompositional gases). The orbits also retained significant gas pockets, both between the eye and fat layers within the orbit and in the eye itself. The endocranial cavities were almost completely gas-filled, with little remaining of the brain.

Shallow flume.—The rotten heads buried in the shallow flume had a likewise strongly bimodal distribution, with two heads retaining no more void space than clean skulls and two heads retaining as much or more than the fresh heads. Here again the difference was due to eyelid integrity. In the heads with little void space, most of the void space was retained in the fronto-ethmoidal and dorsal paratympanic sinuses. The lacrimal sacs retained some void space, but the remaining paranasal sinuses were fluid-filled, as were the nasal passages other than a small gas pocket in the olfactory region. In the paratympanic sinuses, the quadrate and articular diverticula were partially gas-filled, but the caudal and rostral diverticula were almost completely fluid-filled. A few small 27 bubbles were interspersed throughout the soft tissue, and a little gas was trapped in the rostral portion of the oral cavity.

The fronto-ethmoidal sinuses in the two rotten heads buried in the shallow flume were almost completely gas-filled, with gas extending into the lacrimal sacs and the dorsal paratympanic sinuses. The other major gas pockets were in the orbits, which contained void space within the eyeballs, antorbital sinus, under the eyelid, and between the eyeball and the medial fat layer. The mesethmoid and suborbital sinuses were mostly fluid-filled. The caudal and rostral tympanic diverticula were mostly fluid-filled, but the articular and quadrate diverticula retained void space. The endocranial cavity was partially gas-filled and the remaining soft tissue had numerous gas bubbles dispersed throughout, but the nasal and oral cavities were for the most part fluid-filled.

Clean skulls

Deep flume.—The clean skulls buried in the deep flume were not completely fluid-filled, as might be expected, although they retained significantly less void space than those with soft tissue. Void space was retained mostly in the dorsal-most fronto- ethmoidal sinuses, followed by the dorsal tympanic diverticula. The lacrimal sacs often contained void space, as did the quadrate and articular diverticula, although the amount varied considerably within and between skulls. The caudal tympanic diverticula were often mostly fluid-filled. The mesethmoid and parasphenoid rostrum were usually completely fluid-filled, as was the endocranial cavity. The pterygoid and jugal 28 intraosseous recesses retained traces of void space, as did the rostral-most mandible and premaxilla.

Shallow flume.—The clean skulls buried in the shallow flume were similar to the heads buried in the deep flume in terms of quantity and mostly in pattern as well. Most void space was in the dorsal portions of the fronto-ethmoidal sinuses, extending down in some instances into the lacrimal sacs. The dorsal tympanic diverticula also retained significant gas pockets. The articular diverticula retained sizable gas pockets, although the quadrate diverticula were mostly fluid-filled. The caudal tympanic diverticula had small bubbles of gas, but were mostly fluid-filled. Gas pockets were retained in the rostral tympanic diverticula, although the parasphenoid rostrum itself was usually completely fluid-filled. The mesethmoid and endocranial cavities were completely fluid- filled. Small gas bubbles appeared in the rostral-most dentary and premaxilla, as well as small bubbles trapped within the oral cavity. The most notable difference seen in the clean skulls buried in the shallow flume as compared to the deep flume was the presence of gas pockets in the rostral tympanic diverticula caudal to the parasphenoid rostrum, which was not observed in the skulls from the deep flume.

Long-term burial set

The patterns that developed after seven months of continued decomposition became much simpler and differences between flumes and decompositional states became less well defined, with the exception of the clean skulls, which already displayed similar patterns of void space retention (Fig. 8). All heads, including the clean skulls, 29 contained substantially higher volumes of void space. The patterns can essentially be summarized as whatever soft tissue was present decayed and was replaced in large part not by sediment or water influx, but by void space, most likely a result of decompositional gases, as evidenced by the fact that the void space in all the heads buried with soft tissue was higher after the long-term burial than that seen in the unburied control heads (Figs. 5, 6).

The highest amount of void space was seen in the fresh heads buried in the shallow flume and one rotten head buried in the deep flume. All three of these heads left an essentially head-shaped void within the sediment. Virtually all the sinuses located within bone were gas-filled, as were the endocranial cavities. Almost the entire extent of the nasal cavities were gas-filled, as well as the orbits. The oral cavities were primarily fluid-filled, but even here gas occupied the medial palatal surface down to the rostral end of the beak.

Scans of the remaining heads with soft tissue revealed greatly increased quantities of gas over the short-term-burial scans. With the exception of the suborbital sinuses, the paranasal sinuses were completely gas-filled. The paratympanic sinuses were also gas- filled, with the exception of portions of the rostral tympanic diverticula. The orbits were in all cases mostly gas-filled. The olfactory regions of the nasal cavities were gas-filled, although the respiratory regions were typically sediment or fluid-filled, as were the oral cavities, with the exception of a medial palatal gas layer, although the sediment was typically riddled with small gas bubbles. 30

The clean skulls of both the deep and shallow flumes developed similar patterns.

The caudal fronto-ethmoidal sinuses were gas-filled, extending rostrally to varying extents. The other intraosseous paranasal sinuses were filled with either fluid or sediment.

The dorsal tympanic, quadrate, and articular diverticula of the paratympanic sinuses were all gas-filled, although curiously the caudal tympanic diverticulum was primarily fluid- filled. The amount of gas within the rostral tympanic diverticula varied considerably between skulls. Small pockets of gas were also present under the rostral tips of both the dentary and premaxilla.

Statistical Analyses

The amount of void space was analyzed via MANOVA after log-transforming the measurements to normalize the data, followed by individual ANOVAs and Tukey-

Kramer Multiple Comparisons to analyze specific comparisons. Results were verified using a three-way ANOVA and Kruskal-Wallis nonparametric ANOVA. ProStat version

4.83 and NCSS 2004 were used for all analyses. As a whole, state of decomposition and time (short-term-burial scans versus long-term-burial scans taken seven months later) showed highly significant variations (Table 3). No significant variation was seen in the type of flume, nor were there any significant interaction effects. These results did not change if the variables were analyzed separately.

In most cases, Tukey-Kramer Multiple Comparisons tests indicated little difference in void space retention within the heads buried with soft tissue attached, whether it was fresh, dried, or rotten, but the clean skulls retained significantly less void 31 space than the other states. The short-term-burial scans in the deep flume and the long- term-burial scans in the shallow flume revealed slightly more detailed differences. In the short-term-burial deep-flume data, clean skulls showed significantly less void space than the other groups. Fresh heads were not significantly different from any other group other than the clean skulls. However, whereas the desiccated and rotten heads were not significantly different from the fresh, they were significantly different from each other, with the rotten heads retaining more void spaces than the desiccated heads. In the long- term-burial shallow-flume data, clean skulls were again different from the other groups, but here, the fresh heads were also significantly different from any other group, showing the largest retention of void space of any group. The desiccated and rotten groups, unlike the short-term-burial deep-flume data, showed no significant difference in the long-term- burial shallow-flume data.

The only individual ANOVA that indicated a different result from that of the

MANOVA was the short-term-burial shallow-flume data, which did not indicate a significant difference between decompositional states. This lack of statistical significance is due to the bimodal patterns in the desiccated and rotten heads. In both situations, four heads were used. In each case, two heads retained considerable quantities of void space

(more than that seen in the fresh heads), and the other two had very little void space, similar in level to the clean skulls. The difference related to the integrity of the orbits. If the orbits maintained a seal, gasses built up. Once the eyelids degraded sufficiently to allow gas to escape, however, virtually all the gasses not trapped within bony pneumatic sinuses exited the skull. 32

Exterior Patterns

Exterior to the heads, patterns of gas bubbles created a halo around the heads related to sediment collapse into the void spaces created by soft-tissue decay (Fig. 9). Gas bubbles 1–2 mm in diameter were dispersed through the sediment, extending 2–4 cm dorsolaterally from the orbits and nasal cavities. The amount and extent of the gas bubbles were loosely associated with the amount of soft tissue in the heads at the time of burial. The fresh and rotten heads had the most extensive haloes. The desiccated heads demonstrated less extensive haloes, and no haloes appeared on the clean skulls. Heads buried in the deep and shallow flumes exhibited similar patterns.

Discussion

The results of this study indicate that substantial amounts of gas remain in carcasses after burial, which increases during further decomposition and may persist for at least seven months. The origins of these void spaces are due in some part to experimental design, but that is unlikely to be the entirety of the explanation, which may explain in part why many fossils are preserved in the manner they are.

It could be argued that the void space can simply be attributed to drainage from the blocks before they were frozen. Despite concerted effort to rapidly extract and freeze the blocks, drainage almost certainly occurred to some extent, but drainage alone cannot fully account for the patterns observed. If drainage were the cause, one would expect all of the blocks to have responded in a similar fashion. There were no large sedimentologic 33 differences between the blocks containing heads with large void spaces and those with little void space, so there is little reason to expect that some blocks would be significantly more drained than others. There is good evidence to suggest that drainage of the blocks contributed little artifact to the observed patterns. For example, many of the blocks, particularly those in the short-term-burial scans, retained water pooled on the surface.

Several heads also retained substantial amounts of water, in many cases exhibiting regions containing sediment, water, and void space in three distinct layers. For the long- term-burial study, water was added regularly to maintain water flow. Each block received on average 65 L of water, yet less than 4 L drained over the course of the seven months, indicating that, despite being covered, much more water evaporated than drained. Even then, when the blocks were removed from the storage containers, water was pooled at the surface and the surrounding sediment was heavily water-saturated. Thus, drainage of water can be discounted as a confounding factor to interpretation of the data.

The largest contributor to the void space observed in the short-term-burial scans was most likely natural air trapped during burial. Struthio heads contain extensive pneumatic sinuses, residing in both bone and soft tissue. Despite attempts to let as much air escape before being placed on the flume bottom, some air was unable to escape.

Because of the small exits and torturous routes, the paratympanic sinuses are particularly difficult to fill with fluid, resulting in the high degree of retained air in these sinuses.

Gaseous decay byproducts likely explained much of the void spaces in the long- term-burial scans and the short-term scans of the rotten heads in the deep flume, and, to a lesser extent, in the desiccated heads. Decomposition creates considerable gas 34 byproducts, including carbon dioxide, hydrogen, methane, hydrogen sulfide, and ammonia, as well as several volatile organic compounds, such as cadaverine and putrescine (Sakata et al., 1980; Gill-King, 1997; Jackowski et al., 2007). Decay presumably would have been initiated during the early stages of desiccation, but would have slowed or stopped toward the end; nevertheless, gas byproducts would have been produced during the early stages, which may have become trapped within the head depending on the integrity of the integument. Decay of ligamentous tissues and collagen within the bone would likely have supplied suitable decay products for the increase in gases seen even within the clean skulls and abundant material was available in the other heads.

Bone and soft tissue appeared to be a sufficiently effective barrier to gas and water diffusion to prevent air trapped within the heads from escaping into the surrounding water, even after almost two days underwater. Whereas one might have assumed that the fresh heads would have retained more primary air due to intact soft tissue, this proved not to be the case in the deep flume, in which the fresh heads retained less than either the desiccated or rotten heads. Desiccation appeared to have caused little damage to the integrity of the skin while at the same time trapping more gas within the desiccated tissues. The gas in this case is likely a combination of primary air and trapped decompositional gasses. The skin of the rotten heads was loose, but intact for the most part, and decomposition created additional gas that remained trapped within the tissues, likely mixing with primary air within the original pneumatic spaces as well. Heads buried in the shallow flume underwent similar decompositional patterns, but the end result 35 differed due to the increased water flow, which caused small differences in decomposition to have a greater effect. In both the desiccated and rotten heads, gas built up as long as the eyelids remained intact. The water flow in the shallow flume washed the eyes, reducing the integrity of the decaying skin. When the eyelids became compromised, virtually all the gas in the head was replaced by water flowing directly into the head.

Thus, a fossil containing large void spaces within the orbits may be interpreted as having been buried with intact eyelids.

The decay process itself may have provided another factor in the retention of gases. Microbial biofilms can greatly impede diffusion of liquids and gases and are known to create bubbles of gas within the biofilms itself as metabolic byproducts

(Stewart, 2003; Noffke et al., 2008). Microbial mats can make gas domes several centimeters across even without the aid of a nutrient-rich carcass (Noffke et al., 2008).

Certainly all the heads in the long-term-burial trials created abundant biofilms during the decomposition of soft tissues. The rotten heads in the short-term-burial trials had thick biofilms covering them when placed in the flumes. Even though decay had slowed on the desiccated heads by the time they were placed into the flume, decay had been initiated during desiccation and would have developed biofilms at that time which would have been quickly rejuvenated when placed in the water for burial in the flumes.

In addition to creating much of the gases filling the heads, the microbial biofilms also likely directly aided in limiting sediment influx into the head. The mucilaginous secretions forming the biofilms are known to play a major role in the cohesiveness of sediments in both marine (Black, 2002) and riverine sediments (Gerbersdorf et al., 2008) 36 and play important roles in preserving a variety of traces, such as ripples and tracks

(Sutherland et al., 1998; Noffke et al., 2001; Friend et al., 2008).

Microbial biofilms may have facilitated the development of the haloes as well, and would be crucial for the possibility of their persistence. Creation of the haloes was created in large part by collapse of sediment into the void spaces left by decaying tissue.

However, it is likely that the driving cause of the sediment collapse was the gas buildup created by the very decay that created the void. Gas released from microbial biofilms has been proposed as a mechanism for creating fluid escape tubes in sediment by increasing pore pressure within the sediment (Murphy and Sumner, 2008). In these cases, fluid is squeezed out vertically due to surrounding pressure. In this study, the void space would have created a weak wall that would be more likely to collapse inward rather than create a tube upwards due to the large size of the void space within the head and the skull forming a bony roof preventing a vertical collapse of sediment. Without the presence of that weaker wall, it is likely the biostabilization created by the biofilms (Paterson, 1994;

Noffke and Paterson, 2008) would have continued to support the void spaces.

The haloes could potentially be preserved by the microbial biofilms. In addition to their role as biostabilizers, microbes are known to precipitate a wide variety of minerals

(Lowenstam and Weiner, 1989; Simkiss and Wilbur, 1989), particularly in association with decaying organisms (Allison and Briggs, 1991; Wilby et al., 1996; Sagemann et al.,

1999), and can do so within minutes to days of beginning decomposition under the proper circumstances (Hirschler et al., 1990; Martill and Harper, 1990; Dunn et al., 1997). If the gas bubbles forming the haloes were mineralized quickly, the haloes could persist as 37 small mineralized pellets surrounding the orbits and nasal cavities, similar to structures known as bird’s eye structures found in some carbonate rocks, also thought to be formed by gas bubbles (Shinn, 1968). They could be distinguished from unassociated grains by being concentrated around the middle of the skull, and would most likely consist of calcite or apatite.

The presence of large void spaces being created as the result of soft-tissue decay and the failure of sediment to readily displace the decomposition gases likely affects preservation patterns and could explain the condition of many vertebrate fossil heads.

Without sediment filling the voids, a carcass will continue to decay until it eventually collapses of its own accord if not earlier due to the weight of overlying sediment. This explanation has been proposed for the condition of several human skeletons in Egyptian mummies and a York cemetery (Brothwell, 1987) and likely occurred during the formation of flattened but otherwise exquisite fossils, such as many that have come out of the Messel oil shales, such as Darwinius (Franzen et al., 2009) or Liaoning Province in

China, such as Microrapter (Xu et al., 2000) and many others. These fossils are justifiably famous for their soft-tissue preservation surrounding the skeletons, but the skeletons themselves are generally compressed.

Fossils preserved with compression by decay can be distinguished from those being crushed or diagenetically compressed by the condition of the bones. Compression by decay-induced collapse will leave the bones jumbled but in fair condition, unless crushed or altered later. Crush compression will leave the bones in more of a life position, but will greatly damage the bones. Diagenetic compression may plastically warp 38 the bones and possibly pull them apart, but it will leave them in articulation and intact.

Bones broken in the process will remain relatively in position. Of course, these preservational styles are not mutually exclusive and could all happen to a greater or lesser extent to the same fossil. A carcass could begin undergoing decay collapse and then become crushed in the later stages, followed by further diagenetic alteration at any point afterwards. An example of this is the American Museum of Natural History’s

Tyrannosaurus rex (AMNH 5027; Osborn, 1912), which began disarticulating due to decay as shown by the right ectopterygoid being in the left antorbital fenestra, indicating additionally that the carcass lay on its left side for an extended time. After it was buried, the skull suffered additional diagenetic alteration that slightly warped the skull, although the warping could also have occurred early due to tissue deformation simply due to the weight of its skull.

The data presented here indicate that the voids in most fossils, especially in the pneumatic sinuses are primary in origin and are not due to secondary dissolution. This potentially means that some of them might be amenable to gaseous isotope studies, which could serve as an additional and independent source of information to that obtain from other sources, such as teeth, paleosols, etc. If the voids could be sealed— or even potentially filled— by early apatite mineralization, exceptionally well preserved voids may retain traces of the gases at the time of burial. Amber had been thought to be impermeable in some studies (Berner and Landis, 1987, 1988), but its impermeability is disputed by others (Beck, 1988; Cerling, 1989). If this were true, it would make any preservation of ancient air in these sinuses seem unlikely. However, gas and fluid 39 inclusions in mineral crystals are usually thought to be more reliable (Goldstein, 2001;

Lowenstein et al., 2001). If this were true, then these voids may indeed prove to be useful sources for such studies.

Few fossils are found with conspicuously large void spaces such as those seen here. In many instances, this can be explained by compaction, crushing the skull as previously discussed. However, it is likely that bioturbation is a more common process, unless the carcass is buried beyond the bioturbation zone very quickly. Most sediments are rife with macro-organisms that will quickly scavenge the carcass, bringing sediment into the carcass along with them as they break up the soft tissue and the protective microbial mats. An excellent example of this phenomenon is the Thuliadanta presented by Colbert and Eberle (2007).

These data indicate that there is potential information in not only the bones but also in the sediment and voids in and around the fossils. Therefore, the analysis of well- preserved fossils should include an examination of the sediment matrix around the fossil before freeing it from its confines in the field and when we get it back to the lab, taking careful note of the sediment adjacent to and within the fossil as well. We are just beginning to elucidate the information we may be able to retrieve from the stone, but only if we are not in too much of a rush to clean the fossil first, thereby throwing out a potentially useful source of information.

Conclusions

In conclusion, it is apparent that void spaces can be utilized in the interpretation of fossils. Substantial gas remains trapped in heads after burial, which increases during decomposition. Only the most easily accessible pneumatic recesses could be expected to collect sediment during burial, such as were the mesethmoid recesses in the ostrich sample. Other recesses are likely to remain free of sediment unless the skull is cleaned before burial and/or broken before or during burial. The sediment surrounding fossil skulls in which void spaces are found (e.g., within the endocranial cavity, orbits, or nasal cavity) should be carefully examined for soft-tissue traces as these void spaces likely indicate that the head was buried with substantial soft tissue surrounding the skull.

During excavation in the field, sediment around a skull should be examined for small mineral pellets around the orbit and nasal cavities, which would also indicate the presence of substantial soft tissue around the skull during burial.

Further experimental work needs to be done to examine the effects of different positions of the head and possible tumbling during burial, as this may change specific patterns. The response of void spaces to compression also needs to be examined and should go a long ways towards explaining the condition of many fossils. Burial in heavily mineral-rich waters also needs to be studied, due to its potential to create rapid mineralization on the carcass, which should enhance preservation. Understanding these factors and their relationships will provide increased information on early burial conditions and may point to predictions of rock units that are more likely to preserve exceptional fossils. 41

Figure 1. Anatomy of the head of Struthio camelus, labeling structures discussed in the text. Nasal airway is in yellow, with the olfactory region in red. Paranasal sinuses that communicate with nasal passage are in purple, blue, and bright green. Tympanic sinuses within bony pneumatic recesses are in darker green. Endocast of endocranial cavity is in light purple. Modified from Witmer and Ridgely (2008). 43

Figure 2. Flumes used to bury ostrich heads. A) Deep flume schematic. B) Shallow flume schematic, side view. C) Shallow flume schematic, top view. D) Photograph of deep flume. E) Photograph of shallow flume.

Figure 3. Potential fluvial depositional environments emulated by this study, as indicated by the area encompassed by the rectangle. The deep flume is representative of conditions involving a drop in water velocity and/or deeper water, such as might be seen when a river enters a lake or pool. The shallow flume is representative of a point bar. Both situations are particularly applicable in flood stage. Adapted from Tarbuck et al. (2005).

Figure 4. Methods diagram. All CT datasets were analyzed using Amira, and sediment blocks were sliced for visual confirmation. Sixteen sediment blocks were reburied for seven months and rescanned before slicing, providing two sets of data for these blocks, a short-term burial set and a long-term burial set. 46

Figure 5. Void space measured in buried ostrich heads. Error bars are standard error. CON = unburied, control ostrich heads. Remaining conditions as indicated on x axis. First letter: Flume type, D = Deep, S = Shallow; Second letter: Burial time, S = Short- term, L = Long-term; Third letter: Decompositional state, F = Fresh, D = Desiccated, R = Rotten, C = Clean skull. Void space increases in all conditions from short- to long-term, going from less than control levels in the short-term burials to more than control levels in the long-term burials, with the exception of the clean skulls, which had a less pronounced increase. Shallow burials showed more variability than deep burials initially, but less in the long-term burials. Fresh heads had significantly more void space in the long-term shallow burial than did either the desiccated or rotten heads, although in the deep burials, rotten heads had significantly more than either fresh or desiccated in both the short-term and long-term burials. 47

250

200

150

Void Space (cc) 100

0 CON Deep Shallow Short Long Fresh Dry Rot Clean Flume Type Burial Time Tissue Condition at Burial

Figure 6. Void space within heads by group. Data from all heads were pooled and plotted. Error bars indicate standard error. Burial time shows significant difference (p<0.01), as do clean skulls compared to other conditions, but no difference is seen in overall flume type or between tissue conditions which still retained soft tissue at time of burial.

Figure 7. Reconstructions of void space seen in short-term-burial CT scans. For each set, the top image is a right lateral view, and the bottom image is dorsal view. A) Shallow flume, fresh head; B) Deep flume, clean skull; C) Shallow flume, desiccated head; D) Deep flume, rotten head.

Figure 8. Reconstructions of void space seen in long-term-burial CT scans. The top images are right lateral views, and the bottom images are dorsal views. A) Shallow flume, fresh head; B) Deep flume, clean skull.

Figure 9. CT images showing disruption haloes of gas bubbles around orbits and nasals. A) Horizontal section (rostral is to the right), B) Axial (cross) section, C) Sagittal section showing location of horizontal and axial sections through head. 51

Table 1. Ohio University Vertebrate Collections (OUVC) specimens of ostrich used in this study, arranged by treatment condition for short-term burial set. Bold numbers indicate those reburied and subsequently rescanned for long-term burial set.

Burial Condition Tissue condition Deep Flume Shallow Flume Control Fresh 10467, 10468, 10471, 10476, 10634, 10635, 10470 10477 10636 Desiccated 10457, 10458, 10465, 10466, 10459, 10460 10510, 10511 Rotten 10472, 10473, 10474, 10475, 10499, 10500 10501, 10502 Clean skull 10454, 10455, 10451, 10453, 10497, 10498 10456

Table 2. Standard flume sediment mixture. Due to variability in source material, the precise mixture could not be maintained precisely, but varied no more than 10% for any size category. Variations greatly favored slight reductions in the coarsest fraction in favor of finer fractions.

Wentworth size Grain size (mm) Phi % (by wt.) class 1–2 -1–0 Very coarse sand 9 0.5–1 0–1 Coarse sand 22 0.25–0.5 1 – Medium sand 33 2 0.125–0.25 2–3 Fine sand 22 0.063–0.125 3–4 Very fine sand 7 <0.063 >4 Silt and clay 5

Table 3. Statistical summary of void space comparisons. N indicates number of CT scans in each comparison. Wilk’s lambda values reported for MANOVA. Hotelling-Lawley Trace, Roy’s Largest Root, and Pillai’s Trace were also calculated, but resulted in identical values. For Tukey-Kramer Multiple Comparisons, those enclosed by parentheses or brackets are not significantly different, those outside are significantly different. P values considered significant at 0.05 level.

N Df F value P value MANOVA, full data set 45 Depth 1 0.12 0.73686 Time 1 56.49 <0.0001 (Depth)(Time) 1 0.10 0.75186 State 3 12.99 <0.0001 (Depth)(State) 3 1.26 0.30160 (Time)(State) 3 0.24 0.86478 (Depth)(Time)(State) 3 0.38 0.76557

ANOVA, Long-term data only 16 Depth 1 0.00 0.96388 State 3 53.30 0.00001 (Depth)(State) 3 1.48 0.29265

ANOVA, Short-term data only 29 Depth 1 0.22 0.64204 State 3 8.71 0.00060 (Depth)(State) 3 1.53 0.23591

Individual ANOVAs for State Short-term, Deep flume 15 3 29.34 0.00001 Short-term, Shallow flume 14 3 1.22 0.35186 Long-term, Deep flume 8 3 13.07 0.01556 Long-term, Shallow flume 8 3 121.99 0.00022

Tukey-Kramer Multiple Comparisons N Group Differences Short-term, Deep flume 15 ([FD])(C), ([FR])(C) Short-term, Shallow flume 14 (FDRC) Long-term, Deep flume 16 (FDR)(C) Long-term, Shallow flume 8 (F)(DR)(C) 53

CHAPTER 2: SEDIMENT/CARCASS INTERACTIONS AT THE INTEGUMENT

AND THEIR IMPLICATIONS FOR FOSSIL INTERPRETATION: A CT-BASED

ACTUALISTIC TAPHONOMY STUDY

Abstract

The type of integument and its various appendages greatly influence many aspects of an animal’s life, functions, and evolutionary history. But because soft tissue rarely preserves in the fossil record, reconstructing the appearance of fossil animals draws largely on knowledge of extant relatives and comparatively rare exceptionally-preserved specimens. CT scans of 30 ostrich heads with differing amounts of soft tissue (fresh, desiccated, rotting, and cleaned), buried in two flumes emulating common burial conditions (deep and slow flow vs. shallow and fast flow), were examined immediately post-burial and after seven months to determine if sediment patterns could be useful in elucidating integument characteristics, thereby increasing potential information derived from fossils. Interpretations based on CT data were verified by correlating sediment data from examination of 460 points within the sediment surrounding the heads to density measurements from the CT scans. These data then served as a guide for interpreting CT scans based on an additional 2489 points of sediment data. Heads buried in the deep flume showed little to no patterns deriving from the integument, although there was a slight enrichment of fine sediment near the rotting heads. The heads buried in the shallow flume showed more variation, with drapes of fine sediment associated with the fresh and rotting heads, as well as, to a lesser extent, the desiccated heads. Clean skulls in the 54 shallow flume exhibited no sediment drape, thus confirming that the drapes were induced by the intact integument. Sediment patterns in heads subjected to long-term burial were disrupted in some areas by decompositional outgassing. Nevertheless, sufficient sediment integrity was retained around those areas with little soft tissue between the bone and integument to identify patterns seen in the short-term burial scans. Some patterns seen in the short-term burial scans were even enhanced after the long-term burial in some shallowly buried heads due to the migration of fine sediment towards the head. CT scans were therefore effective in elucidating these sediment patterns in the extant ostrich sample, suggesting that CT scanning likewise can provide some clues towards both integument and burial conditions in fossils.

Introduction

The integument, including the skin and the structures within or overlying it, is diverse in both scope and function. Integumentary appendages can take the form of scales, fur, feathers, or keratinous sheaths forming, in part or in whole, horns, claws, nails, beaks, and tooth-like structures. They can be as unnoticeable as fine hairs on a baby or as flamboyant as a peacock’s tail. The integument itself can form diverse soft tissue structures, such as wattles, combs, snoods, or even trunks.

Integument and the various integumentary appendages play numerous roles that profoundly influence—or are influenced by—the animal’s physiology, behavior, ecology, and evolution. Feathers, hair, and scales all play a role in thermoregulation, protection against the elements, fluid retention, camouflage, display, body contouring, and either 55 sound production or alteration (e.g., Maderson, 1972, 2003; Morbeck, 1979; Ortolani,

1998). Specialized hairs are used for mechanoreception (e.g., Carvell and Simons, 1990).

Feathers and scales are also used for locomotion, albeit in vastly different ways (Dial,

2003; Homberger and deSilva, 2003; Hu et al., 2009). Modified keratinous sheaths are used for a variety of purposes, including digging, predation, or defense (Hildebrand,

1988). Rhampotheca are modified for highly diverse uses (Proctor and Lynch, 1993). But perhaps most obviously, integumentary variation is used in species or sexual communication (e.g., Fitzpatrick, 1998; Moller and Cuervo, 1998; Caro, 2005).

Reconstructing the evolutionary history of integumentary appendages is a required step in sorting out the adaptive responses to selective pressures driving the evolution of the integument and the animal as a whole and can even affect our understanding of the evolution of large clades (Maderson, 2003; Wu et al., 2004; Hill,

2005; Sire et al., 2009). However, few fossils preserve soft tissue of any kind, leaving large gaps in our knowledge, requiring interpolations across huge evolutionary distances and in many cases extrapolation beyond what our data support. As a result, work has been done almost exclusively in the extant realm, with minor input from the fossil realm.

Nevertheless, when the fossils are included, they can radically change our perceptions of evolution within important groups (Donoghue et al., 1989; Eernise and Kluge, 1993).

One need only look at the change in perception of theropod dinosaurs from scaly reptiles to feathered bird antecedents and the early evolution of mammals to see the effect of the fortuitous fossil find. Feathers, or feather antecedents, once thought to be exclusive to birds, have now been found in hundreds of specimens of theropod dinosaurs (e.g., 56

Chuong et al., 2003; Norell and Xu, 2005; Prum, 2005; Hu et al., 2009; Xu et al., 2009), and other feather-like structures have even been found in ornithischians (Mayr et al.,

2002; Zheng et al., 2009), making the picture of feather evolution more interesting

(Witmer, 2009). Similarly, the discovery of a fossil mammaliaform, Castorocauda, with soft tissue preserved (Ji et al., 2006) demonstrated that fur evolved prior to the first appearance of Mammalia, as well as shifting the first evidence of aquatic specializations in mammaliforms 110 million years deeper in time (Martin, 2006).

Specific and often apomorphic integumentary structures are little known in extinct organisms, with the exceptions of those integrated with bone. The bony cores of casques on hornbills (Bucerotidae) preserve, but generally not the keratinous appendages that extend and form most casques. Likewise, feathery crests or the skin folds and thickenings that form wattles, combs, caruncles, and throat pouches (Stettenheim, 2000) are almost never preserved. Without the preservation of soft tissue, determining whether integumentary appendages were present can be difficult. The basal ceratopsid

Psittacosaurus would never have been reconstructed with bristles on its tail if they had not been preserved (Mayr et al., 2002), and it is doubtful that Archaeopteryx would have been reconstructed with feathers if it had not been found with them preserved

(Wellnhofer, 2009).

This chapter examines the hypothesis that some insight into the presence of integumentary appendages is gained by looking at sediment traces left behind after the decay of soft tissue by using an actualistic, experimental taphonomic approach, using computed tomography (CT scanning) to map external sediment patterns around buried 57 heads. Sediment patterns developing around buried heads were examined to determine their relationship to the condition of soft tissue present during burial. If sediment patterns indeed relate to the presence of soft tissue after further decay, this could greatly aid our ability to reconstruct extinct organisms. Whereas it would be unlikely to provide detailed information without good soft-tissue preservation, it may provide clues to the presence of soft tissue that extends beyond the confines of the skeleton.

Materials and Methods

Specific details concerning the general experimental design are presented in

Chapter 1. Briefly, 30 Struthio (ostrich) heads were buried in two flumes (Fig. 10) designed to emulate salient aspects of two common burial conditions for carcasses in terrestrial freshwater ecosystems: (1) burial in deep, slow-moving water, such as is found in the upstream end of a pool or just downstream of a blockage, and (2) fast, shallow- moving water, such as is found in a sandbar type of deposit. The heads were buried in four different levels of decomposition: fresh heads, desiccated heads, rotten heads (nine days in room temperature water), and clean skulls. After burial, they were removed and

CT-scanned using a medical CT scanner at O’Bleness Memorial Hospital, in Athens, OH.

A subset was then reburied in a shallow container that allowed for slow percolation of tap water through the blocks and excluded scavengers for seven months and then rescanned.

Struthio was chosen for their large size, which would make resultant sediment patterns easier to evaluate, and for their availability. The decompositional levels covered the range of common burial states for carcasses. All skulls used in the study are archived in the 58

Ohio University Vertebrate Collection (OUVC) and are available from Lawrence

Witmer, as are the CT-scan data generated during the study.

For this study, it was necessary to determine if sedimentological data could be correlated with density measurements determined by the CT scans. Use of CT scans have been used by other workers (e.g., Vinegar and Wellington, 1987) for geological studies to study textural relationships, fluid flow, and void space, although these studies were done with high-energy CT scans or with metallic grains that enhanced contrast (Kyle and

Ketcham, 2003; Hersum et al., 2005; Ketcham and Iturrino, 2005; Nettles and McSween,

2006). However, medical scanners have been successfully used to study marine sediments with minimal contrast (Orsi et al., 1994, 1999). Preliminary experiments using the GE LightSpeed 16 CT scanner at O’Bleness Memorial Hospital in Athens, OH, indicated that such scanning could be done sufficiently well to elucidate overall sediment patterns, if not precise sediment parameter characteristics (Daniel and Witmer, 2005).

Correlation of sediment parameters with CT scan data was further evaluated on the sediment in which the heads were buried. After the sediment blocks were CT scanned, they were sliced into approximately 2–3 cm sections. Initially, the plan was to collect several sediment samples from the sections, allowing grain-size distributions to be precisely measured. The data were lost because of a software error in the sediment analyzer, coincident with an equipment failure, and it was impossible to retrieve more samples. Instead, we estimated grain-size distributions from detailed photographs of the sections. This method was limited in resolution compared to direct measurement of sediment distributions for several reasons: (1) determination from photographs is 59 restricted by the resolution of the image, preventing fine grains from being measured; (2) image clarity, which in many cases was limited by frost forming on the grains of the frozen sections during photography; and (3) precision of measurement, which must be balanced between accuracy for a few grains or accepting less precision with the ability to examine more samples (see Appendix A for more details). Despite these caveats, estimation from photographs, such as that done here, although less accurate, is more reflective of what would be available to a researcher examining a fossil.

We determined grain-size distributions in three ways. (1) The dominant grain size of the area of interest was estimated by counting the number of grains in one centimeter transects and categorized according to size class of the Udden-Wentworth scale (Table

4). (2) We then used the scale bar in the photographs to measure the largest grains and the smallest grains to determine the range of grain sizes, again categorized by size class. (3)

We then estimated relative fines contribution into the following categories: none observable, moderate, or considerable. Relative fines contribution was estimated for two reasons. First, the volumetric contribution by coarse particles so overwhelms the contribution by fine-grained particles that a few coarse particles can contribute as much volume as fine-grained particles that number orders of magnitude more than coarse particles. Second, clay particles have been shown to be important for exceptional preservation and are attracted by organic matter (Martin et al., 2005). Additionally, clays help to stabilize organic material for thousands of years, making preservation much more likely (Wattel-Koekkoek et al., 2003). See Appendix A for more details of grain-size measurement in general and Appendix B for details specific to this study. Using these 60 parameters, we were able to determine distributions for 2949 areas on 10 blocks

(Appendix C). Due to the increased gas pockets permeating the blocks that were scanned after seven months of decomposition, only the blocks that were frozen immediately after burial and sectioned shortly after CT scanning were used for grain-size estimation.

Blocks containing a fresh head, desiccated head, and a clean skull from both the deep and shallow flume were used, as were rotten heads from each flume. Both rotten heads were used because the rotten heads had more intra-group variation in sediment patterns than the other heads.

Of the 2949 regions for which grain-size parameters were estimated, 460 were matched with Hounsfield density values taken from the CT scans and measured using the line probe measurement tool (although in a few cases, a spline probe was used) in the software program Amira 5.1, utilizing 500 readings along the transect line (Fig. 11). The regions for grain-size distribution varied from regions as small as 2 mm in thickness to 20 mm, depending on the variation within the sediment. Regions in which density readings were taken varied from 8–20 mm. The smallest areas were excluded from the density measurements, as were those that had an abnormally high level of CT artifact, such as evident beam hardening or edge effects. The remaining regions for which grain-size parameters were estimated were used to confirm CT-based grain-size estimates for sediment pattern interpretations along with direct comparisons with slices from the sediment blocks.

Results

Reliability of Sediment Parameter Estimations

Statistical tests determined that density variations as measured from CT scans were useful as a crudely reliable relative indicator of dominant grain size and range, provided sufficient samples are taken, but less so for relative fine-sediment contribution.

Regions dominated by the coarsest fraction were clearly differentiated from those dominated by the finest fractions, as were the most heterogeneous regions from the most homogeneous samples. Samples that fell in between were not clearly distinguishable from either extreme. This result was likely due at least in part to the low resolution of the estimates for the sediment distribution parameters. With finer detail in measuring sediment distributions, determinations from density variations would likely improve.

Nevertheless, when used as a relative scale within a specimen, this method was sufficiently reliable to determine sediment patterns. It was not sufficient, however, to give precise measurements of sediment distributions that could be compared with specimens from different preservational features.

To make comparisons between the sediment parameters and the density values, the 500 density values measured for each corresponding sediment sample needed to be compressed into one or a few variables that both adequately described the population and could be statistically compared to the sediment parameters. Typically, this is done using average values, but this technique loses the complexity of the distribution. Thus, the sediment parameters were compared with the main statistical distribution parameters, or

“moments,” those being mean, standard deviation, skewness, and kurtosis (Zar, 2009). 62

Mean and standard deviation proved most relevant, with generally lesser contributions from skewness. Kurtosis rarely had a significant effect. All the following statistical procedures were run in SAS 9.2

A MANOVA was run to test the hypothesis that the categories of sediment parameters were not significantly different based on density measures (Table 5). Mean density proved significant for both dominant grain size and grain-size range (p<0.0001), but not for fines determination (p=0.4726). But whereas the MANOVA indicated that the categories for dominant grain size and range had statistically distinguishable density distributions, a series of exploratory graphs and regressions indicated a generally poor predictive capacity in most circumstances. Mean density, when the shallow flume data were separated from the deep flume data, proved the most reliable, achieving an r2 value of 0.5788 for dominant grain size and 0.5908 for grain-size range when regressed

(Appendix C).

A series of discriminant function analyses were performed to more fully test the predictability of the categories for dominant grain size and range. Because density values had little predictive power for relative fines contribution, other than in the samples with the lowest means, discriminant function analysis was not performed for this parameter.

Mean density and standard deviation proved the most informative population parameters for dominant grain size. Density proved to be a poor predictor, with a 58.9% error rate, but the error was not equal across all categories, with size categories 1 and 6 having 33.3% and 39.6% respectively, whereas category 4 had a 72.6% error rate. This is compared to an error rate based on a random distribution of 83%. Estimates from the 63 deep flume were less accurate than from the shallow flume (62.0% error rate vs. 49.3%, respectively). When considered by head condition, predictions of dominant grain size by density in the fresh heads were the most accurate (41.0% error rate) and the rotten heads were the least accurate (54%). Relaxation of the categories, which is defined as counting the readings correct if it was within one category above or below, dropped the error rate to only 15.2% overall, with category 1 having only a 4.8% error and category 4 having an error rate of 21.4% compared to an expected error rate of 63.9%. Thus, density values can be used as general indicators of dominant grain size, provided multiple readings are taken and absolute estimates of grain size are not required.

Grain-size range mirrored dominant grain size in that mean density and standard deviation were the dominant parameters and that base predictive capability was poor, with an overall error rate of 42.6%, compared to an expected random error rate of 80%.

Thus, while performing much better than random, it was not highly predictive. Here again, the extreme ranges were better predicted than the middle ranges. Readings in the upper ranges from the shallow flume were more reliable than the deep flume (e.g., 20.8% error for category 6 in the shallow flume compared to 47.9% error for the deep flume) and the fresh heads were markedly more reliable than the clean skulls (overall 35.6% versus 55.1%). Using the relaxed categories, the error rate dropped to an overall error rate of 12.0%, with category 2 and 6 achieving a 6.2% and 12.5%, respectively, and with category 4 having 19.0%, compared to a random error rate of 60% for categories 2 and 6 and 40% for category 4. Relaxing the categories only marginally improved the predictive capability for the middle categories, whereas it greatly improved the predictive ability of 64 the extreme ranges. Thus, grain-size range could be predicted on a relative scale between homogeneous sediments and very heterogeneous sediments with multiple readings, as long as precise measurements were not required.

Sediment Patterns

Collectively, the burial conditions showed a large effect on sediment patterns, with the shallow flume being much more diverse in both sediment composition and pattern. Blocks from the shallow flume typically had a lower layer consisting of mud, with coarse material overlying it, itself being highly variable in its composition from place to place, whereas blocks from the deep flume were dominated by relatively unpatterned sediments throughout.

Short-Term-Burial Sediment Patterns

Three fresh heads were buried in the deep flume (Ohio University Vertebrate

Collections [OUVC] 10467, 10468, 10470). Sediment patterns rostral to the heads were relatively homogeneous, with stratification indistinct (Fig. 12). The sediment itself was highly heterogeneous, consisting primarily of mid-range sediment but covering the entire range. In one head, as the sediment began to cover the bill, the sediment began to shift toward finer particles dorsally and larger particles laterally. The shift to finer sediment formed a distinct layer running caudally along the base of the head; forming a thick wedge near the head thinning quickly laterally. Above this layer was typically a very coarse layer that was also rich in fine sediment, itself overlain by layers with less fine 65 material. In the other two heads, no distinct layer was deposited, although there was a wedge of less coarse sediment at the eye level and in the occipital region. A diffuse sediment layer, containing more coarse sediment than seen in the dorsally adjacent sediment layers, covered all the heads from the nasal apertures to the apex of the heads.

Feathers above the surface did not seem to have a consistent effect on the sediments, in some places seeming to have more fines than adjacent sediment, in other places having more coarse material. In one head, feathers did appear to have aided in trapping a coarser fraction than that seen adjacent to them. The feathers surrounding the ear canals contained sediment coarser than that seen either within the ear canal itself or just outside of it. The wedge of finer sediments behind the occipital region intersected the ear canals, providing the sediment adjacent to the coarser sediment in the feathers. In general, however, the sediment did not substantially change as it encountered the head in any way that might be clearly indicative of the underlying tissue.

The four rotten heads buried in the deep flume (OUVC 10472, 10473, 10499,

10500) shared the initial sediment patterns seen in the blocks with fresh heads (Fig. 12C,

D). The rest of the block showed very little modification of the sediment. The sediments did not change markedly toward the head. There seemed to have been a slight decrease in the coarsest material near the heads, possibly a small enrichment of fines, but little else.

Dorsally, there was no distinct layer, but there apparently was an indistinct layer which exhibited a small decrease in coarse particles and possibly increased fines. It extended up to 1 cm above the apex of the head, but faded even more into obscurity 1–2 cm above the rostral part of the mesethmoid. The thin layer of coarse material over the dorsal surface of 66 the rostral portion of the skull seen in the fresh heads was only partially seen here in small patches, but was very thin and intermittent. No wedge or layer of finer sediment was present caudal to the heads as was seen in the fresh heads.

The skin around the rotten heads tended to slide off the head in a way not generally seen in the other conditions. The skin formed a flat layer around the base of the head with the feathers generally aligned with the head, the rachises pointing downstream

(Fig. 13). Unlike the surrounding sediment which was dominated by coarse sand, the sediment most closely associated with the sloughed-off skin, and feathers tended to be dominated by fine sand, although it retained the high levels of heterogeneity seen in the surrounding sediments.

The four desiccated heads buried in the deep flume (OUVC 10457, 10458, 10459,

10460) showed little to no sediment patterns. Any influence of the soft tissue on what few sediment patterns were seen is questionable (Fig. 12E, F). There was enrichment of fine grains in the bottom-most layer of sediment, but the layer was diffuse and not clearly localized to the heads and so did not appear to be related to any factor specific to the heads. There was no clear change in the sediment directly overlying any of the heads.

Two of the heads had an indistinct layer no more than 1–2 mm thick overlying the top of the head that appeared to have been enriched in coarse material, giving the appearance of the feathers having simply inhibited the grains from rolling off the head, as the feathers sticking farther out into the sediment do not appear to have otherwise affected the sediment around them. Places where the eyes had sunken into the orbits had finer sediments in the dorsal portions of the orbits than in the ventral portions, but there were 67 otherwise no observable patterns that relate to the heads. One head had a layer of sediment appearing to lack much coarse material around the orbits and ear canals that faded caudally, and photos indicated a possible increase in coarse particles in the feathers around the ear canals, but this was difficult to make out and not seen in the CT scans.

There was no wedge of finer sediment close to the heads as was seen in the fresh and rotten heads.

The four clean skulls (OUVC 10454, 10455, 10497, 10498) in the deep flume showed the least amount of sediment patterns and were more similar to the desiccated heads than the fresh or clean heads (Fig. 12G, H). No evidence of interaction between the skull and the sediment was evident, except for a reduction in coarse material caudally in the occipital region, most likely due to coarser material being trapped within the skull initially, leaving the finer sediments to more easily fill in behind the skull. No trace of a layer associated with the dorsal surface of the skull was present.

In contrast to the deep flume, all of the heads within the shallow flume showed much more complex sediment patterns and more interactions between the heads and the sediment (Fig. 14). All three fresh heads buried in the shallow flume (OUVC 10471,

10476, 10477) exhibited a distinct layer of mud and fine sand, beginning from the near the tip of the bill to the nares and extending to just caudal to the apex of the head, reaching a maximal thickness of 11–12 mm near the lacrimals. This layer paralleled the surface of the head, forming a crosscutting relationship with the layering seen in the surrounding sediment. This layer was possibly related to a layer of mud and fine sands that began as a thin, indistinct layer rostral to the heads, but became thicker and much 68 more distinct around the heads. Caudal to the head, the mud layer reduced in thickness and the grain size generally shifted to slightly larger grains, becoming dominated more by fine sand than mud. In one head, the mud layer faded almost completely in sections caudal to the head.

The sediment patterns relating to the integument were suggestive, but not conclusive (Fig. 14). The layer overlying the heads was itself overlain by a very thin, intermittent layer of coarse material. Laterally, the variation of the sediment adjacent to the integument was inconsistent. Sediment in contact with the heads varied from being coarser than the sediment farther away, exhibiting no changes, or was finer than the distal sediments. One head appeared to have a very thin, patchy, layer adjacent to the skin consisting of enriched mud and coarse sand, but this layer was only apparent when viewing the CT data axially and was not apparent when viewing the data horizontally or sagittally. Most areas seemed to be enriched in muds, but the extent varied considerably.

The eyelashes were either encased within mud and fine sand, despite being otherwise surrounded by coarse sand, or showed no observable change.

A Chi square analysis of 57 matched pairs of observed sediment samples was done to elucidate trends in grain size variation related to the integument. For each pair of samples, one point lay adjacent to the integument, whereas the other point lay slightly farther away. Dominant grain size only showed a small trend towards smaller grain size adjacent to the skin (λ=2.00, p=0.37), but the range showed a clear trend towards reduced ranges, that is, more homogeneity (λ=5.16, p=0.08). Relative fines contribution significantly increased (λ=35.37, p<0.0001). This analysis did not distinguish between 69 the left and right sides of the head. Considering that the heads had a clear mud and fine sand layer on the dorsal surface, it is likely that, if separated out, the lateral sides would display no clear trend in dominant grain size and only a marginal range reduction.

The four rotten heads buried in the shallow flume (OUVC 10474, 10475, 10501,

10502) showed a distinct layer of fine sediment on the dorsal surface of the head similar to that seen on the fresh heads. However, the increase in the relative fines fraction and loss of coarse sediment near the head was much more prominent here. Rostral to the head, there was only an indistinct, intermittent layer showing a relative lack of coarse material, but it quickly became a thick, muddy layer engulfing the mandible, with a further fine sandy layer covering the lower half of the head. This muddy layer formed a prominent wedge caudal to the head, becoming patchy and coarser distally. The feathers were generally covered in mud, except for the distal ends of the longer feathers. A thin, coarse layer of sediment covers one head caudal to the apex and the layer of fine sediment, but this appeared to be due to the head intersecting a downstream sloping layer unrelated to the head itself. This layer was actually deflected over the head by the layer of fine sediment that runs parallel to the head surface.

A Chi square analysis of 89 pair-matched sediment samples indicated much stronger correlations than in the fresh heads buried in the shallow flume. Dominant grain size, range, and relative fines contribution either did not noticeably change or shifted to finer-grained sediments (λ2=14.18, 15.12, 139.3, respectively; p=0.0008, 0.0005,

<<0.0001, respectively). Whereas there were some cases in which the dominant grain size increased and/or the range widened, there were no cases in which the relative fines 70 contribution decreased. In every case, the mud and fine sand fraction either stayed the same or increased.

Four desiccated heads were buried in the shallow flume (OUVC 10465, 10466,

10510, 10511). They all showed a distinct layer of fine sediment overlying the skull, but it was not as prominent as those above the fresh and rotten heads, except for one head

(Fig. 14E,F). The layer began around the nares and terminated near the apex of the head, but did not always pass the apex and was not as thick or distinct as seen in the fresh or rotten heads. There was a ventral layer of fine sediment here as in the others, but it did not extend over the bill. Thus, there was a clear break between the layer of fine sediment on the skull and that covering the base of the block. Moreover, the muddy layer at the base of the block was generally thinner than those seen in the fresh and rotten heads and did not develop into a thick, distinct layer until the sediment reached the main portion of the head with the orbits and braincase, after which it decreased in both width and distinctiveness as the sediments began to blend more with the surrounding sediment.

Laterally, the sediment around the bill did not change significantly. However, a very thin layer of fine sediment appeared to have been associated with the integument in most parts of the caudal portion of the cranium, particularly the feathers surrounding the ear canals.

The sediment patterns surrounding the three clean skulls buried in the shallow flume (OUVC 10451, 10453, 10456) bore a marked contrast to those around the other heads buried in that flume (Fig. 14G, H). There were no distinct layers of fine sediment overlying the skulls. The only place in which fine sediment did collect preferentially was over the craniofacial hinge, which exhibited a sudden change in angle of the surface 71 encountered by the sediment. The ventral mud and fine sand layer was also present here, but it was generally not as pronounced laterally from the skull as seen in the other heads

(although one skull did exhibit a thick mud layer in one area that had a depression in the floor of the block). The lateral sides of the mandibles had finer material adjacent to them than more distally, with mud dominating medially. As the skulls gained in height, the interaction with the sediment decreased such that the braincase did not appear to have any interaction with the sediment that altered the sediment composition.

Long-Term-Burial Sedimentation Patterns

Because fossils represent burial under extended periods, usually well after all the soft tissue has rotted away, sediment blocks containing two heads from all four decompositional conditions in both burial regimes were reburied for seven months to determine how the sediment patterns may have changed. In general, factoring out the disruption of the sediment by the escape of decompositional gases, the basic sedimentation patterns did not change markedly.

The sediment surrounding the fresh heads buried in the deep flume (OUVC

10467, 10468) were substantially disrupted by the infiltration of copious amounts of what were presumably decompositional gases (Fig. 15). Nevertheless, the patterns within the sediments—such as they were—were not substantially altered. The wedges of less coarse material were still observable around the heads, and no other new patterns were found.

The two desiccated heads from the deep flume (OUVC 10459, 10460) showed very different patterns. One head showed virtually no change in sediment. There were a 72 few lenses of gas or fluid between sediment layers, but they did not disrupt the sediment patterns; they were much like intrusive micro-sills in geology. The decompositional gases seemed to have been largely contained within the head itself, with little escaping into the surrounding sediment. The other head, however, showed extensive disruptions around the orbits and bill where sediments collapsed into void spaces within the head. Nevertheless, as the sediment did not show significant patterns relating to the head to begin with, there was little, if anything, to alter.

Both rotten heads buried in the deep flume (OUVC 10472, 10473) showed considerable void spaces after seven months, with extensive infiltration of gases into the adjacent sediment, particularly around the orbits and nares. However, the extent to which it affected the surrounding sediment varied. In one, only the sediment immediately adjacent to the head was affected, and here, the sediment was only expanded by the addition of gases, not substantially disrupted. In the other head, collapse into portions of the head and more extensive gas infiltration into the sediment obliterated any patterns that may have been present laterally to the head and above the nares. In both cases, gas formed sills between sediment layers as seen in the other heads. Nevertheless, the sediment dorsal to both heads remained unaffected and sediment patterns were unchanged.

The long term scans of the clean skulls in the deep flume (OUVC 10454, 10455) appeared virtually identical to the initial scans. Some of the water within the skull had been replaced by gas and a few small gas lenses appeared in the sediment, but nothing affected the structure of the sediment in any meaningful fashion. 73

The prolonged burial actually enhanced some of the patterns around the fresh heads buried in the shallow flume (OUVC 10476, 10477; Fig. 16). Areas of sediment dominated by coarse grains appeared to have lost some of the fine sediment, whereas the layers of sediment near the head that were dominated by fines appear to have gained material. This phenomenon was particularly noticeable around the orbits in one head in which the shape of the eyes remained intact, which had the effect of enhancing contrast in areas. However, in areas in which the sediment collapsed into void spaces within the head created by the decay of soft tissue, patterns were obliterated, such as occurred in one head in which sediment collapsed into the orbit and oral cavity. Elsewhere, places in which the skull supported soft tissue retained the sediment patterns around them.

Scans of the desiccated heads buried in the shallow flume after prolonged burial

(OUVC 10465, 10466) exhibited some of the enhancement seen in the fresh heads (Fig.

16). The adipocere that formed within and on the heads could be clearly seen in the CT scans, as could the feathers around the head and residual brain matter that had not yet decomposed. The rest of the soft tissue had completely decomposed, including the ligaments connecting the bones. Nevertheless, the sediment patterns formed during burial were still intact everywhere that had not suffered catastrophic sediment collapse. Fine- grained sediment marked locations previously occupied by soft tissue around the cranium and mandible and was more clearly marked in the same manner as that seen in the fresh heads.

Both rotten heads buried in the shallow flume (OUVC 10474, 10475) showed similar patterns in the long-term burial scans. The sediment adjacent to the integument 74 directly dorsal to the head was unperturbed. The sediment contacting the lateral aspects were extensively infiltrated by gas and fluids that obliterated any patterns directly adjacent to the integument. The lateral extent of the infiltration was limited, however, to a relatively narrow zone compared to the fresh head. Feathers that remained after seven months formed a thin layer adjacent to the bone that was as visible on the long term scans as they were on the initial scans and in some areas were more visible.

The clean skulls buried in the shallow flume (OUVC 10451, 10453) were very similar to those buried in the deep flume in that virtually no change took place between the initial scans and the long-term burial scans. Here again, a small amount of void space opened up in the bones. There was also a small level of fine-sediment transfer away from regions near the dorsal sections of the block that were dominated by very coarse sediment to the ventral sections of the block.

Discussion

Grain-Size Analysis

The use of CT for grain-size analysis has potential as a viable tool. Here, differences were limited to determining gross patterns in sediment with distinctive separations in either dominant grain size or heterogeneity. Much of the limitations in the method, however, are likely due to the imprecision of measuring sediment distributions.

More accurate and precise measurements would almost certainly have increased correlations with density readings. 75

High-resolution CT scans have been used to determine grain sizes in absolute terms (Kyle and Ketcham, 2003; Ketcham and Iturrino, 2005), and medical scanners have been used to determine relative differences in grain size (Orsi et al., 1994, 1999), so this technique is not new. The level of verification presented here, however, has not been done previously. Sediment data from numerous locations within ten of the sediment blocks allowed us to accurately interpret the sediment patterns found within the other blocks despite the relatively low resolution of our CT data.

Even though further refinement and more precision in sediment distribution estimates may allow the precision of this technique to be greatly improved, each situation will still need to be individually assessed. This study did not touch upon differences caused by variations in cementation or elemental composition of sediment, both of which could be affected by organic interactions with the sediment, which would affect density values. Additionally, varying levels of compaction during the early lithification stages of the carcass and the surrounding sediments would affect packing of the sediments as well as influencing the degree of cementation by controlling pore space and water flux. Thus, local depositional conditions must be taken into account for proper interpretation. he converse may also be true. The condition of the fossil and the sediment patterns around the fossil may help determine the conditions under which the sediments were deposited.

While there does exist the danger of circularity, iterative data processing of the fossil and the sedimentary matrix may increase the precision of the interpretations of both by allowing contextualization of data for which interpretation is ambiguous. 76

To be of use in other contexts, a detailed physical examination of the sedimentologic characteristics of the matrix around a fossil should be undertaken to precisely determine grain-size distributions, composition, and cementation. These characteristics can then be used to accurately assess the CT data from the fossil and other fossils from the same area, allowing a detailed understanding of the sediment and the capacity to map sediment characteristics from the CT data. Knowledge of the matrix characteristics can be of particular use in identifying small patterns that may be more readily detected in CT-scan data than by visual examination, as well as in determining patterns in areas inaccessible to visual assessment without removing the very material to be examined. The CT data also provides a three-dimensional permanent record of the sediment patterns that can be examined even after the specimen has been fully prepared.

Currently, few institutions regularly CT scan fossils before preparation. Hopefully, as scanning becomes more available and the utility to which it can be used becomes better appreciated, more institutions will acquire such facilities, either separately or in conjunction with partner institutions.

Sediment Patterns

Variation in Burial Environment

The results of this experiment indicate that there are recognizable differences in the sediment patterns created by different burial regimes and, to some extent, differences in the presence of soft tissue at the time of burial. The level of detail, however, is poor, and the meanings of the specific patterns are difficult to elucidate conclusively. 77

Nevertheless, there is justification for the expectation that the patterns are real and potentially useful.

The heads in the deep flume were all surrounded by sediment that, as a whole, had a fairly homogeneous texture but was highly heterogeneous in composition, whereas the sediment in the shallow flume had a highly variable texture, reflecting the fact that many areas had restricted grain-size variation. This difference was entirely controlled by the mode of burial. The deep flume had sediment entering a pool of slow-moving water, which caused the larger, and thus heavier, particles to sink to the bottom quickly while the fine sediment settled out more slowly. The resulting sediment accumulation was an initial pile of very coarse sediment covered by sediment that contained particles of all sizes. As more sediment continued to enter the system, the coarsest grains settled out concurrently with finer sediments that had been introduced earlier. In contrast, the first particles to reach the end of the shallow flume were the finest grains, with the coarsest grains following up much more slowly. In addition, the embayment at the end of the flume caused the grains that were not small enough to be carried in suspension to settle out of the flow as the water eddied around the end of the flume. This created an initial layer of fine grained sediment that was then overlain by coarser material. With these initial patterns in mind, we may now turn our attention to the patterns that vary from this template to elucidate the organically derived sediment patterns.

Variation in Carcass Condition

Heads buried in either a fresh or rotting state in the deep flume altered the general sediment patterns only minimally, if at all, with a wedge of material enriched in fines laterally and an inconsistent very thin layer enriched in coarse grains dorsally. The lateral wedge of sediment enriched in fine-grained sediment extended all the way across the block, despite being thickest adjacent to the head, indicating normal sediment stratification. However, it did not extend rostral to the heads, arguing for a specific head effect on the sediment. One possibility is that a layer of sediment finer in composition than earlier sediment developed on the leading edge of the sediment advance, which was subsequently buried by coarser sediment. The other possibility is that organics leached into the surrounding water from the head, attracting clay particles. Because the water was relatively still, fine-grained sediment built up in the water near the head, becoming incorporated into the sediment as it was deposited. The thin layer of coarse material on the dorsal surface is likely due to entrapment of grains by the feathers, preventing them from sliding off the surface. This interpretation is supported by the lack of evidence of these patterns surrounding the desiccated heads and none around the clean skulls. The only pattern exhibited by the clean skulls was that of finer sediment caudal to the occiput.

This can be explained by the coarser particles being trapped more easily by the bones of the skull, allowing the smaller grains to more quickly infill the area behind the head.

In the shallow flume, the most predominant pattern is a distinct, drape-like layer of sediment overlying the heads with soft tissue. The drapes were very distinct in the fresh and rotten heads, and, while they were still present in the desiccated heads, they 79 were less distinct. The drapes could simply be a result of the same settling patterns that resulted in the fine-grained layer found ventrally in all the blocks, but the fact that the layers are thin and indistinct rostral to the heads argues against this explanation. One could also argue that the layer started at the embayment and progressed upstream as more sediment was added, only getting intertongued with the coarser sediment around the heads. Although possible, this explanation requires special pleading for the timing of the sedimentation. More importantly, it is contradicted by the clean skulls, which show no evidence of such drapes. If the soft tissue were not involved in forming this layer, one would expect the clean skulls to exhibit a comparable drape, which is not the case.

The differences in the drapes may be best explained by the varying states of the integument and soft tissue. The fresh heads exhibit a distinct layer of fine-grained sediment which may have occurred due to trapping of sediment by the feathers. The first sediments encountered by the feathers were the smallest grains and thus the first to be deposited. They were also preferentially deposited by attraction of clays to organics which allowed a layer to build up on the surface in advance of the sediment that rose up around the head. As the collection of clay built up, this would have allowed the collection of larger silt and fine sand grains. The top of the head provided a stable platform for the entrapment and collection of grains. The steeper inclines laterally would have prevented a significant buildup of sediment, except at the base. The rotten heads exhibited a thicker layer in general than the fresh heads which may be because the rotten heads quickly dispersed abundant organic material into the water, causing the sediments around the head to turn black in some areas. These dispersed organics likely enhanced the sediment 80 collection on and near the head. The desiccated heads, on the other hand, had a smaller degree of soft tissue and what was there was tightly adhered to the skull and presumably less effective at sediment entrapment. The tissue would have been rehydrated in the water, but it would take time for the tissue to fully absorb the water, by which time the burial process would have already started. Additionally, even after hydration, the desiccation process irreversibly altered the composition, cohesion, and amount of soft tissue. The clean skulls, of course, had no soft tissue and so were buried simply as a result of sediment deposition covering the skull. The surface did not attract sediments, so the finer particles simply washed over and rolled off the head until such time as the sediment built up around the skull, as it would any obstruction to flow.

The rotten heads exhibited sloughed off skin forming a flat layer around the base of the head. Because this was not generally seen in the other heads, the appearance of sloughed skin near the carcass would seem to indicate that the carcass was already in a state of decomposition during burial and thus there was some period of time between death and burial. This also indicates a potential confounding issue with soft-tissue reconstruction in that position of the skin is not necessarily a good indicator of live placement and must be taken into account during fossil reconstruction. This is a phenomenon understood by many workers (e.g., Xu et al., 2001), although it is not often mentioned in descriptions of skeletons surrounded by soft-tissue traces (e.g., Lindgren et al., 2010) or carbon traces which are often considered to mark body outlines (Franzen et al., 2009). It is essential that position of the integument in relation to the carcass, 81 depositional environment, and the potential for slippage such as that seen here be taken into account when interpreting traces of soft-tissue remains.

The variation of sediment patterns between heads in the shallow flume and the deep flume indicate that such sediment interactions may provide additional information on environmental factors, in addition to tissue content. The drapes and sediment wedges that formed around the heads gave a strong indication of flow patterns. Relative velocity could be inferred from the grain-size distributions along the surface of the head and the interactions between the sediment and the carcass. Of potentially more use, however, is as an indicator of flow direction. Flow direction in the specific region around the head could be determined more easily and precisely than flow velocity. These parameters have great utility in determining small scale environments as an iterative tool for refining interpretations. Whereas these interactions would not be of great use by themselves, they provide additional independent measures, as well as conceptual context in support of interpretations of other data.

Substrate composition will affect sediment/carcass interactions which will in turn affect preservation potential. Most Lagerstätten, such as the Burgess or Messel Shales, have a high proportion of clay and silt, thus enhancing preservation. Particles of this size are attracted to organic matter and allow a high resolution in detail. Additionally, they generally inhibit water flow and thus serve to slow decay and the spread of decompositional fluids. Sandy substrates, however, have higher porosities and water flow, thereby enhancing decomposition over that of finer-grained sediments. Interactions such as the drapes seen here may then aid preservation by reducing the sediment size 82 nearest to the carcass. Such protection may aid in protecting microbes involved in decomposition, providing a structural foundation for forming molds of the surface or otherwise protecting the soft tissue from complete destruction. Microbial presence on the surface may also enhance the formation of the drape, which may explain the increased drape formation on rotting carcasses. These types of interactions are likely to form at least part of the explanation for the preservation seen in such specimens as the

Edmontosaurus MRF-03 (Manning et al., 2009) and the brachylophosaur “Leonardo”

(Murphy et al., 2009).

Conclusions

Sediment patterns surrounding skulls can be utilized in the interpretation of fossils. Water flow direction is clearly interpretable by examination of sediment patterns around the skull. Information about the amount and condition of soft tissue around the skull at the time of burial also can be obtained. The presence of finer-grained sediment drapes over the head indicates both a moderately fast water flow rate and the presence of soft tissue, with the most distinct drapes occurring with fresher tissues. Intact feathers or other integumentary appendages will increase the thickness and distinctiveness of the drape. Tissue actively undergoing decomposition at the time of burial creates a thicker, more diffuse drape with relatively more fine-grained material as compared to both the surrounding sediment and heads with fresher soft tissue. Additionally, the amount of skin sloughing off rotting carcasses can create a tissue halo around the carcass that can superficially appear as a midline structure for carcasses residing on their side and thus 83 makes argument by location problematic without corroborating evidence and a comprehensive view of the entire carcass.

This work points the way toward identifying the presence of epidermal appendages before more solid evidence, such as the clear preservation of definitive structures. It will not determine what exactly the structures are. For an example, the origin and distribution of feathers is a hotly debated topic (Feduccia et al., 2005; Xu,

Zheng and You, 2009; Dhouailly, 2009; Witmer, 2009; Lingham-Soliar, 2009, 2011;

Buchwitz and Voigt, 2012), partially due to taphonomic difficulties in determining precisely what type of structure one is studying (Benton et al., 2008; Foth, 2012).

Bristle-like or feathery structures have been found on such disparate species as pterosaurs

(Bakhurina and Unwin, 1995; Lu, 2002; Wang et al., 2002), ornithischians (Mayr et al.,

2002; Zheng et al. 2009), and a variety of nonavian theropods (Norell and Xu, 2005;

Witmer, 2009; Xu et al., 2011, 2012). The relationship between these structures is unclear. Genetic studies have been ambiguous, because in modern species, such as turkeys, we see both feathers and unrelated bristles, with the genes controlling their development widespread throughout Archosauria (Sawyer et al., 2003, 2005). Whereas sedimentologic data cannot provide direct support for homology, the data can provide indirect support (or lack thereof) by reducing discontinuities in our phylogenetic knowledge about the presence or absence of epidermal appendages. At the very least, sedimentologic data will allow more refined reconstructions of soft tissue, providing support for more than conservative estimates of simple skin stretched over a bony frame even when soft-tissue preservation is not immediately obvious. These data will strengthen 84 and provide context for ambiguous data, improving interpretations for not only soft-tissue reconstructions, but also local taphonomic conditions and environmental parameters. 85

Figure 10. Flumes used to bury ostrich heads. A) Deep flume schematic. B) Shallow flume schematic, side view. C) Shallow flume schematic, top view. D) Photograph of deep flume. E) Photograph of shallow flume. 86

Figure 11. Line probe analysis example for OUVC 10500. A) Sediment slice through back of head. B) CT scan image corresponding to sediment slice using Amira 5.3. Orange line indicates location of line probe analysis for density measurement. C) Output plot of 500 readings along line probe, measured in Hounsfield units. Mean and standard deviation of data for this line probe is also given.

Figure 12. CT scans of sediment blocks taken from the deep flume and immediately frozen. No distinct sediment drape appears on any head, although an indistinct layer may be seen on the rotten head (red arrows). Column one: Sagittal slice images. Column two: 88 axial slice images through orbit. A, B: fresh heads; C, D: rotten heads; E, F: desiccated heads; G, H: clean skulls.

Figure 13. Position of sloughed integument on a rotten head buried in the shallow flume (OUVC 10475). A: CT slice, B: slice through sediment block. C, D: enlarged sections denoted by red rectangles in A and B. 89

Figure 14. CT scans of sediment blocks taken from the shallow flume and immediately frozen. Red arrows indicate sediment drape distinct to the ostrich heads. Column one: parasagittal slice images. Column two: axial slice images through the orbit (H is through the braincase). A, B: fresh heads; C , D: rotten heads; E, F: desiccated heads; G, H: clean skulls. 91

Figure 15. CT scans of sediment blocks taken from the deep flume after seven months. Decompositional gases separated sediment layers near head which may be mistaken for actual sediment differences (black arrows). Indistinct sediment drape seen in the initial 92 scan is still present (red arrows). Column one: parasagittal slice images. Column two: axial slice images through the orbit (H is through the braincase). A, B: fresh heads; C, D: rotten heads; E, F: desiccated heads; G, H: clean skulls. 93

Figure 16. CT scans of sediment blocks taken from the shallow flume after seven months. Red arrows indicate sediment drape distinct to the heads. White arrows indicate adipocere. Column one: parasagittal slice images. Column two: axial slice images 94 through the orbit (H is through the braincase). A, B: fresh heads; C, D: rotten heads; E, F: desiccated heads; G, H: clean skulls.

Table 4. Grain-size categories for sediment distribution estimates.

Category Grain Size (mm) Phi

1 ≤0.062 4+

2 0.062–0.125 3–4

3 0.125–0.25 2–3

4 0.5–0.25 1–2

5 0.5–1.0 0–1

6 1.0–2.0 -1–0

Table 5. MANOVA results using full data set from both flumes. Only Wilk's lambda is reported, but Pillai's and Hotelling-Lawley's Trace and Roy's Greatest Root gave similar answers.

Wilk’s λ F Value Num DF Dem DF Pr > F

Grain Size 0.7744 5.92 20 1476.8 <0.0001

Range 0.8782 3.69 16 1360.1 <0.0001

Fines 0.9831 0.95 8 890 0.4726

CHAPTER 3: THE UTILITY OF CT-BASED, TAPHONOMIC STUDY OF

SEDIMENT/CARCASS INTERACTIONS FOR SOFT-TISSUE RECONSTRUCTION

OF EXTINCT ORGANISMS

Abstract

Reconstruction of extinct organisms is complicated by the lack of soft-part preservation in fossils. Finding additional data sources that expand our knowledge of soft tissues could greatly improve our understanding of past life. To test the hypothesis that sediment interacts with soft tissue during burial in predictable ways that may be of use in the reconstruction of fossil animals, 30 ostrich heads ranging from fresh heads to clean skulls were buried in two flumes: a short, deep flume with slow-moving water and a shallow flume with faster-moving water, each designed to emulate common burial situations in alluvial systems. After the heads were buried, the sediment blocks containing the heads were CT scanned, after which 16 heads were reburied for seven months and then rescanned. Sediment was mostly excluded from heads containing soft tissue in the deep flume until rot or desiccation created substantial openings, although rostral conchae were usually discernible in heads with soft tissue, as were, to a lesser extent, external auditory canals. Sediment more clearly delineated nasal and oral cavities in the shallow flume, as well as increased clarity in the auditory canals. Paranasal and paratympanic sinus recesses were generally clear of sediment, except in the clean skulls, which had sediment partially fill the mesethmoid and parasphenoid rostrum. The endocranial cavity was only partially filled in even the clean skulls. The best sediment 96 signals within the skulls appeared to be in the rotten or desiccated heads, although most of the patterns were destroyed with further decay. However, traces of rostral conchae were still evident within the sediment in some cases even after complete decay. The amount of disruption within the sediment may aid in determining the amount of soft tissue around a fossil when buried and peripheral soft-tissue structures may be interpretable, but interior soft tissue patterns seem out of reach, as decay will obliterate sediment patterns.

Introduction

Reconstructing the soft tissues of extinct animals is problematic, because with rare exceptions (e.g., Schweitzer et al., 2005), soft tissues, unlike bone, typically do not preserve as anything more than the occasional skin or feather impression (e.g., Mayr et al., 2002). Yet many of our inferences concerning the animal, its behavior, interactions, and function within an ecological context, are predicated on inaccurate reconstruction of its soft tissues (Bryant and Russell, 1992; Witmer, 1995). Any errors made in soft-tissue reconstruction are amplified as one attempts to make inferences beyond that which is provided from the primary osteological data (Witmer, 1995). Therefore, inferences made in reconstructing soft tissues should be based on as much sound data as possible.

Soft tissues rarely fossilize and therefore their reconstructions are normally based on modern comparisons. Strong phylogenetic controls (Witmer, 1995) for comparisons provide confidence in the soft-tissue inferences, but as the comparisons become weaker so do the reconstructions (Bryant and Russell, 1992; Witmer 1995). This approach has 97 yielded considerable success when dealing with tissue that directly affects the bone in predictable ways, forming osteological correlates (e.g., Witmer et al., 2003; O’Connor and Claessens, 2005; Hurum et al., 2006; Hieronymus and Witmer, 2010). Other methods have used biomechanical models to inform and constrain reconstructions (e.g.,

Hutchinson, 2004; Hutchinson and Gatesy, 2005). Inferences are based on clues obtained from the bone, and thus the methodology is limited to the tissues that make diagnosable changes to the bone itself. A largely unexplored avenue that may increase the amount of available soft tissue information is the examination of the sediment surrounding the bones. The construction of the bones and the soft tissue surrounding them may affect the way sediment collects around them during burial.

Due to its complexity, the head is of prime importance in many animals, containing a wealth of phylogenetic, anatomical, and physiological data. It is also a potentially useful region for sediment analysis, because it has multiple passages through which sediment may infiltrate (e.g., mouth, nose, eyes, ears). Variations within and between the passages may alter sediment composition (i.e., grain size) by acting as filters.

For instance, one may expect coarser grain sizes to fill the oral cavity due to its size and progressively finer grains within the ear canals and the passages connecting the ears and throat. Soft tissue may alter sediment transport in ways that leave traces within the deposit. For example, nasal conchae are rarely preserved due to their fragile structure.

Nevertheless, they may alter sediment flow into the head during burial and thereby leave an identifiable pattern. Regional boundaries may also be preserved by varying sediment type due to different preservational capacity of tissues. Differences in collagen and lipid 98 content will cause skin, muscle, and fat to decompose at different rates, allowing sediment to infill these areas at different times.

Head construction is also a factor in sediment deposition around an animal. The lower the percentage of bone forming the skull, the more regions within the head will be defined by soft tissue. For instance, the nasal passage in mammals is clearly defined by bone, but not in birds and most other dinosaurs. The extent of the brain, particularly the olfactory bulb, is also less defined in dinosaurs than either birds or mammals. The size and shape of these and other spaces within the skull has generated much debate among paleontologists (e.g., Ruben et al., 1996; Witmer and Sampson, 1999; Brochu, 2000;

Witmer and Ridgely, 2009). Residual traces within the sediment may provide insight to these questions.

It may seem counterintuitive that degrading tissues may leave sediment traces, but the very act of degradation may aid in preserving sediment differences. Whereas the ability of microbial biofilms to preserve impressions by precipitating minerals that stabilize the sediment is well known (e.g., Davis and Briggs, 1995; Gehling, 1999), microbes may also precipitate minerals within decaying tissues themselves to stabilize the tissues (Briggs, 2003a; Carpenter, 2005; Daniel and Chin, 2010). Microbial ability to precipitate carbonates, for instance, is so proficient that it can be harnessed for building materials (Castanier et al., 2000). In addition to mineral precipitation, bacterial degradation can fix clay minerals present in the sediment into the organic tissue thereby preserving the tissue in a clay mold (Martin et al., 2004).

On a broader level, a study such as this that explores sediment/carcass interactions 99 may help to identify taphonomic patterns. Even if specific soft tissues cannot be inferred, it may help to identify how much tissue was left on the carcass during burial, which in turn will provide insights into the condition of the carcass at the time of burial and by extension may give clues useful for interpreting behavioral and environmental parameters.

Taphonomic studies are inherently multidisciplinary because they integrate biology and geology, among other fields, but there are currently few studies that directly combine the two fields. The study of taphonomy is complicated by the numerous factors involved and the poorly understood interactions between them (Fig. 17). Taphonomic studies generally consist of three types: (1) biostratinomy, the study of the time between death and burial, including decay or destruction and post-mortem transport, and can generally be thought of as focusing on the biological aspects of individual deaths (e.g.,

Voorhies, 1969; Behrensmeyer, 1978; Sansom et al., 2010); (2) the study of environmental parameters and community compositions of fossil assemblages (e.g., Staff et al., 1986; Gates, 2005); and (3) diagenetic studies involving biogeochemical or mechanical changes post-burial (e.g., Kohn et al., 1999; Trueman and Tuross, 2002). The goal of taphonomic studies typically is to understand the biases in the fossil record to better determine what information is lost through destructive processes or to help define sedimentary environmental parameters.

There is currently no taphonomic method for the reconstruction of organismal morphology. A shift in viewpoint is needed, from a bones-to-biology/rock-to- environment dichotomy to a more multidisciplinary sediment-to-biology approach. It also 100 requires appropriate tools, which have not been available until recently. Utilizing this type of data requires the capability to map the sediment differences around a carcass in three dimensions. Advances in computed tomography (CT) and 3D visualization software allow such a study. However, a modern comparative dataset, which is not currently available, is also required to enable interpretations of patterns found within fossils. CT scans have revealed sediment differences around and within numerous fossils, yet without a basis of comparison, these differences are not interpretable. Such an approach has been demonstrated in plants, in which permineralized fruits and seeds can be clearly imaged with CT (Devore, 2005). It remains to be taken further and studied for vertebrate animals, in which the sediment may retain trace permineralized organics, but may also retain residual soft-tissue traces in the sediment type itself, either in grain size distribution or cementation differences.

Here we report a series of actualistic laboratory experiments designed to create a baseline for comparison with CT scans of fossils. Struthio (ostrich) heads were buried in a controlled laboratory environment so that variables involved in the burial process could be more rigorously controlled and recorded. These experiments allowed the interplay between soft tissue and sediment to be directly observed and measured.

The objective of the study was to determine the feasibility of utilizing sedimentologic data from CT scans to infer soft-tissue traces and organizational patterns that may be of use in the reconstruction of extinct animals. Our hypothesis that soft tissue within heads affects sediment influx in predictable ways has several corollaries that lead to several testable predictions. 101

1. Soft tissue surrounding and within the head affects sediment influx. Below some threshold, smaller points of entry into the head should show increased sorting with finer sediment. Deeper sediment penetration of the head should result in deposition of finer and better sorted sediment. The inner ear, for example, should show finer sediment than either the naso- or oropharynx. Soft-tissue projections, such as nasal conchae, may increase sorting, either by causing variations within the flow or by being replaced by finer sediment as they decay. Tissue replacement by sediment during decay may also involve finer sediment than that found within larger cavities and these sediments will be better cemented by the decay process. Tissues of different decompositional patterns may be detectable through changes in either sediment or diagenesis. For example, fatty areas may potentially cause different patterns than muscular areas as the fat will much more quickly be removed or altered to adipocere or “grave wax” that may protect areas from short-term decay processes.

2. State of decomposition affects sediment influx. The lesser the extent of decomposition, the more intact the soft tissue and the greater the effect soft tissue may have on sediment influx. Therefore, clean skulls should show the least sediment sorting and fewest patterns. The only patterns that develop should be due solely to skull construction, which is far less complex than a fleshed out head and of much less interest for reconstructions. Fresh heads are predicted to have the best sorting as they are the most complex organizationally and have the best potential for recruitment of microbial decay stabilizing effects. Rotted heads are predicted to have some sorting, but soft-tissue patterns should be less noticeable. The percentage of clay is also expected to be increased 102 throughout the rotted head, and unsorted sediment is expected to penetrate more deeply into the head than compared to fresh heads. Desiccated heads are expected to show the least evidence of sorting for heads with soft tissue, but are expected to develop a rind of cemented fine grained sediment close to the bone.

3. Water velocity and decompositional environment will interact in more complex ways than just determining available sediment grain size. Faster flowing water should cause greater penetration of larger sediment sizes into the head along with greater initial sorting. Soft-tissue organizational patterns will be more apparent compared to low-energy environments. Decay should cause patterns to become less apparent as decomposition opens the heads, causing disruptions of much of the initial sediment patterns. Slow moving water will result in less sediment sorting, but an increase in silt throughout the head. The environment with the best preservational potential is expected to be one that is fast enough to transport the sediment into the head but not fast enough to disrupt accumulated sediment patterns. Pattern resolution is higher with fine grained sediment, so the best velocities would entrain fine sands, but have little effect on coarse sand or higher.

4. Head construction affects sediment influx. This hypothesis was only tangentially tested here, as all work was done with Struthio heads, but predicts that places enclosed by bone within the head will show more sorting during decomposition than areas more open to the outside environment. The more influence the bone has on defining regions within the head, the less influence soft tissue will have on sediment patterns.

Whereas soft-tissue-derived sediment patterns may still be present, they will likely be 103 more difficult to distinguish from the signal created by the construction of the bony skull itself. By contrast, a more open head will be more likely to distinguish sediment patterns derived from soft tissue, because the signal from bone construction will be weaker. This hypothesis is best tested by comparing open-skull types to more closed-skull types, but in a more limited fashion can be tested by comparisons between bone-enclosed sinuses to open passages within the same head.

Materials and Methods

Specific details concerning the general experimental design were presented in chapter 1. Briefly, 30 Struthio (ostrich) heads were buried in two flumes designed to emulate salient aspects of burial in deep, slow-moving water—such as in the upstream end of a pool or just downstream of a blockage—and fast, shallow-moving water in a sand-bar type of deposit. Both are common burial conditions for carcasses in terrestrial freshwater ecosystems. The heads were buried in different states of decomposition covering the range of common burial conditions for carcasses: fresh, desiccated, rotten

(nine days in room-temperature water), and clean skulls. After burial, they were removed and CT scanned using a medical CT scanner at O’Bleness Hospital, in Athens, OH.

Sediment patterns were determined from the CT scans and verified by slicing the blocks into approximately 2 cm slices and visually confirming sediment patterns (Fig. 18). A subset of sediment blocks containing heads was then reburied in a shallow container that allowed for slow percolation of tap water through the blocks and excluded scavengers for seven months and then rescanned. Struthio was chosen for their large size, which would 104 make resultant sediment patterns easier to evaluate, and for their availability. All skulls used in the study are archived in the Ohio University Vertebrate Collection (OUVC) and are available from Lawrence Witmer, as are the CT-scan data generated during the study.

To determine sediment patterns using the CT scans, grain-size distribution estimates had to be correlated with the CT scans. The full methodology was described in chapter 2. Briefly, almost 3000 sediment distributions were estimated from high- resolution photographs of slices through ten of the sediment blocks containing buried heads, of which 460 were correlated with density measurements from the CT scans. This demonstrated the method was reliable for a low-level, comparative interpretation within blocks, but not sufficiently reliable to determine exactly what sediment distributions were present across the entire experiment. Thus, sediment parameters could be estimated accurately relative to other regions within the same block, but detailed descriptions of sediment parameters across multiple blocks was not possible. The remaining 2500 sediment estimates were used to refine high-resolution sediment patterns along with using the photographs of sediment block slices in conjunction with the CT scans to confirm sediment patterns (Fig. 19).

Results

Short-term sediment patterns

Deep flume

Fresh heads.—Of the three fresh heads (OUVC 10467, 10468, 10470) buried in the deep flume, only one had any sediment in the oral cavity, and that was minimal and 105 restricted to the rostral part of the bill in front of the nares. The infilling sediment was heterogeneous and differed from the surrounding sediment only in the loss of the coarsest grains. Sediment did not significantly enter the trachea from the cut end, nor did it show any gradient. Sediment was completely excluded from the endocranial cavity (Fig. 20A–

E).

Sediment penetrated the nasal cavity far enough to delineate the rostral-most nasal conchae, but did not continue significantly past the nasal aperture and did not appear to enter the paranasal sinuses. The sediment within the nasal cavity was heterogeneous, but enriched in fine sediment. The coarsest material remained adjacent to the aperture, but did not otherwise show a strong gradient once past the aperture. Sediment entering the external auditory meatuses filled the canal, showing fine sediment enrichment in association with the tympanum, which was inconsistently delineated. In OUVC 10467, one side was completely delineated by fine sediment, but was obscured by inadequate entry on the other. The coarsest grains were largely prevented from entering the canal, although whether that was due to the feathers surrounding the auditory canal or some other actor could not be determined.

The ventral conjunctival sac collected sediment in two of the heads. The grain size was fine to medium, with little coarse material, and was generally similar in texture to the sediment adjacent to the exterior, although the sediment filling the ventral conjunctival sac in one eye of OUVC 10470 showed clear separation with only mud and fine sand entering the palpebral fissure. Larger grains were probably prevented from 106 reaching the palpebral fissures by the eyelashes. Sediment entry was highly inconsistent, ranging from complete exclusion to clear delineation of the entire conjunctival sac.

Desiccated heads.—The sediment influx patterns present in the four desiccated heads buried in the deep flume (OUVC 10457–10460) were variable in extent and difficult to track through the head in some instances. The texture of the sediment was consistent throughout, and similar to the exterior sediment, with the common exception of a very thin fine-grained layer directly adjacent to soft tissue separating the tissue from sediment that was very heterogeneous texturally, but relatively homogeneous spatially

(Fig. 21).

Sediment influx into the oral cavity was greater than that seen in the fresh heads, but was variable in extent and entry pathway. In OUVC 10457, sediment entered the left side through the bill ventral to the antorbital sinus. OUVC 10459 and 10460 had significant sediment filling much of the right portion of the oral cavity, appearing to have entered through the bill rostrally. Small quantities of sediment in the oral cavity of

OUVC 10458 entered rostrally through the bill. The origin of the small amount of sediment caudally is unclear as it was not connected to any other sediment, but due to its proximity to the choanae is likely to have entered via the nasal cavity. Sediment was mostly excluded from the cut end of the trachea from all the heads, with the exception of

OUVC 10458, although even here the sediment did not approach the oral cavity.

Sediment influx into the nasal cavity extended further in some of the desiccated heads than in the fresh heads. The respiratory conchae were completely encompassed by sediment, which extended into the antorbital sinuses in OUVC 10457 and 10458, but was 107 limited to only the rostralmost portions of the respiratory conchae and nasal cavity in

OUVC 10459 and 10460. Sediment entered the right antorbital and suborbital sinuses in

OUVC 10457 and 10458, but appeared to have entered via the orbit, the floor of which was covered in sediment (Fig. 21). OUVC 10459 had sediment on the right orbital floor, as well, but the sediment did not extend into the paranasal sinuses. Sediment was largely excluded from the left orbits, and no sediment entered either orbit in OUVC 10460.

Sediment generally entered the external auditory meatuses to partially delineate the tympanum (if intact), but in OUVC 10459 and 10460, the tympana were ruptured, allowing access into the middle ear. No sediment was found in any of the paratympanic sinuses. Despite the presence of feathers surrounding and partially covering the meatus, sediment texture was not noticeably altered other than a thin layer of fines adjacent to the interior-most walls.

Rotten heads.—The sediment that entered the four rotten heads buried in the deep flume (OUVC 10472, 10473, 10499, 10500) was similar in texture to that found in the fresh and desiccated heads, i.e., little alteration between external and internal sediment, except for a thin layer of fine sediment adjacent to some soft-tissue barriers (Fig. 22). The entry paths in these four examples were different in that sediment was mostly prevented from entering the oral cavity through the bill. Instead, considerable sediment entered the oral cavity through disruption of the soft palate. Sediment entered downward through the nasal cavity or through the orbit, which was opened up by the decay of the soft tissues within the orbit and the epithelia of the suborbital and antorbital paranasal sinuses. These 108 tissues did not decay in OUVC 10499 and 10500, resulting in an oral cavity without sediment.

Sediment entered the nares to a variable extent, but in all cases, delineated at least the rostral end of the respiratory conchae. The sediment differed here from the surrounding sediment, being noticeably skewed towards finer grain sizes. The degree to which the sediment shifted toward finer grain sizes varied; it was barely noticeable in

OUVC 10500 but clearly differentiated in 10473.

Entry into the orbits was also highly variable. Decomposition in OUVC 10499 did not open the orbits significantly, resulting in sediment reaching only as far as shrinkage of the eyeball allowed. The eyeballs remained intact in OUVC 10500, but they shrank enough for sediment to enter laterally around the eye and enter the suborbital and antorbital paranasal sinuses through epithelia disrupted by decay. The left orbits of

OUVC 10472 and 10473 were opened by decay, allowing sediment to fill the ventrolateral portions of the orbit and partially extending into the suborbital and antorbital sinuses. The sediment in the antorbital sinus of OUVC 10473 was enriched in fine sediment, but the sediment otherwise did not show significant differentiation.

The external auditory meatuses were often completely blocked, with little to no sediment entry into even the lateral portions. When sediment did enter, it was of limited extent. Only one canal was filled with sediment and demonstrated a ruptured tympanum, although sediment did not extend into the paratympanic sinuses.

Unlike the fresh and desiccated heads, there was some entry into the endocranial cavity in the rotten heads, with sediment filling the floor of the cavity in OUVC 10500, 109 but not in the other heads. This variation demonstrates a key characteristic of the rotten heads. Sediment entry at the time of burial was strongly dependent on the specific characteristics of the level of decomposition within the head, which was highly variable, not only between heads having undergone identical decomposition regimes, but between different areas of the head itself.

Clean skulls.—Four clean skulls were buried in the deep flume (OUVC 10454,

10455, 10497, 10498). The only soft tissue remaining on these skulls was minor ligamentous cartilage. As a result, sediment was able to freely enter most spaces normally filled by soft tissue (Fig. 23A, B). However, the bony recesses normally housing paranasal and paratympanic sinuses remained largely devoid of sediment. In three of four skulls, the mesethmoid was partially filled with fine sand and mud, which entered chiefly through the caudodorsal opening by the olfactory foramina, with sediment contributions from both the endocranial cavity and the orbit. The paratympanic sinuses within the basicranium and caudoventral portion of the parasphenoid rostrum collected minor amounts of fine-grained sediment in OUVC 10454. Other than these cases, all other sinuses were empty of sediment, with OUVC 10497 having almost no sediment in any sinus. In all the skulls, the endocranial cavity only partially filled with sediment showing a coarse, heterogeneous core and becoming finer and more homogeneous the farther away from the foramen magnum until it merged with sediment entering from the olfactory foramina.

Sediment patterns exterior to the bone were largely unaffected by the skull, flowing without disruption until directly adjacent to the bone. Laminae of fine sediment 110 were present adjacent to the ventral surfaces of some bones with a large, relatively flat surface, such as the frontals forming the dorsal wall of the orbits. Thin bands of fine sediments were also often present medial to large bones around the basicranium and suspensorium, such as the articular bones and mandibles. It appeared the larger bones served as a temporary barrier to coarser sediment, allowing finer sediment to collect before being mixed with coarser material that collected on the rostrolateral surfaces.

Shallow flume

Fresh heads.—The three fresh heads buried in the shallow flume (OUVC 10471,

10476, 10477) showed considerable infiltration of sediment, as well as gradients not seen in the deep flume (Fig. 24A–F). In OUVC 10477, only minimal amounts of sediment entered the oral cavity via the tip of the bill, but in OUVC 10471 and 10477, the oral cavity was filled with sediment clearly delineating the tongue. The sediment was coarse and heterogeneous alongside the labial margins, but became more homogeneous lingually. The sediment also decreased in grain size toward the trachea, with fine-grained sediment extending down the trachea. It is possible that sediment entered via the cut end of the trachea and flowed upwards, but overall, the pattern is consistent with most of the sediment pattern flowing from oral cavity to trachea. No sediment appeared to enter the esophagus in any of the heads, which is to be expected as the esophagus is typically flattened when not in use, which would prevent entry unless more force was applied than was present in the flume. 111

In OUVC 10471 and 10476, the entire floor of the nasal cavity was covered with sediment, making even the caudal respiratory conchae clearly visible in CT scans. The sediment extended through the choanae into the oral cavity, although sediment did not appear to enter the paranasal sinuses. Sediment did not reach the olfactory conchae.

Consistent with the lack of sediment entering the mouth, OUVC 10477 showed limited entry of sediment into the nasal cavity, extending only as far as the palatal maxillary processes. The sediment was much finer than the majority of sediment overlying the head, but was derived from a thin layer of fine mud to medium sand that separated the bill from the overlying coarser, heterogeneous sediment. Thus, even though some sorting did occur, the degree of sorting was less than it appeared due to the influx of the fine sediment layer.

Sediment was excluded from the orbits of OUVC 10477, and fine to medium sand was seen behind the right ventral palpebral of OUVC 10471, but more sediment was seen in OUVC 10476. On the left side, coarse, heterogeneous sediment entered the fissure, but did not significantly extend under the palpebrals. On the right side however, fine-grained sediment passed through the palpebral fissure and entered both dorsal and ventral conjunctival sacs, delineating the external boundaries of the eye.

Unlike the deep flume, in which coarse, heterogeneous, undifferentiated sediment filled the external auditory meatuses, all the fresh heads in the shallow flume contained fine-grained and more homogeneous sediment than that surrounding the head. This seemed to be linked to the same processes that led to the thin, fine-grained layer present on some dorsal aspects of the head and the finer-grained sediment commonly found 112 surrounding the ventral-most sections of the head. However, the external auditory meatuses only directly intersect one of these layers in OUVC 10476 and were separated by coarse, heterogeneous sediment covering the external aspect of the meatuses in the other two heads.

Desiccated heads.—The four desiccated heads buried in the shallow flume

(OUVC 10465, 10466, 10510, 10511) showed considerable differences depending on how much damage the desiccation did to the epithelia, particularly in the palate, but there were some similarities (Fig. 25). The rostral respiratory conchae were clearly delineated, as were the external auditory meatuses for the most part. The oral cavity was more clearly delineated here than in any other condition. However, little to no sediment entered the endocranial cavities or any of the bony recesses, limited to only minor fine-grained sediment in the basalmost basicranial paratympanic sinuses.

Desiccation damaged the palate of OUVC 10465, particularly near the left choana, causing the nasal and oral cavities to merge in this region. The sediment in the oral cavity was finer grained than the surrounding sediment near the tip of the bill.

Increased sediment input along the bill under the nasal passages caused the sediment to become coarser and more heterogeneous, although the sediment was still finer grained than the encompassing sediment outside the bill. As in the fresh heads, the tongue was clearly delineated. Rostrally, the nasal passages were well defined by sediment, with the respiratory conchae easily visible within the sediment, which was distinctly finer grained than the exterior sediment, although not as fine-grained as the sediment within the oral cavity. Caudally, the oral and nasal cavities merged through the damaged palate and the 113 sediments mixed along with possible input from the cut end of the trachea. The trachea here was not as clear as in the fresh heads, being compressed and shrunken. As a result, there was a clear trace of sediment throughout the oral and ventral nasal cavities into the oropharynx, but the precise positions of the soft tissue were not as clear. Likewise, there was clearly sediment within the areas that contain the antorbital and suborbital sinuses, but the sinuses were not distinguishable as they appear to have ruptured. The sediment in these regions appeared to have come from the oral and nasal cavities, as the orbits themselves looked intact. One eye was partially filled with fine grained sediment, while the other was filled with sediment that was similar to the external sediment. In neither instance was the eye filled to the point of distinguishing any soft tissue markers other than a partially decayed eye. The external auditory meatuses were filled with fine- to medium-grained sediment. In the right ear, fine-grained sediment entered the paratympanic sinuses, partially filling the diverticula within the basicranium most proximal to the tympanum.

The oral cavity in OUVC 10466 was filled with relatively fine-grained sediment, with limited coarse sand input. The tongue was well delineated rostrally, but was not as clear caudally. The integument under the oral cavity was ruptured just rostral to the right orbit, so beyond this point, the interior and exterior sediment mixed. The floors of both nasal passages were filled with sediment, but one contained coarse and heterogeneous sediment, whereas the other was filled with fine-grained sediment that progressively fined caudally. Fine-grained sediment from the nasal passages reached the suborbital and pterygoid sinuses partially filling the ventral sections. The left orbit had coarse, 114 heterogeneous sediment surrounding the eye rostrally, but the eye itself was partially filled with fine-grained sediment. Coarse, heterogeneous sediment filled the auditory canals superficially, but fined inwards to the end of the canals. A small amount of fine- grained material entered the braincase, but the braincase was mostly filled with soft tissue and water.

OUVC 10510 and 10511 were similar to each other, with only minor differences between them. The oral cavity was filled with sediment clearly finer than the exterior sediment, with the coarsest grains limited mainly to small areas around the margins. The entire oral cavity in OUVC 10510 could be traced through the oropharynx, although with the collapse of the trachea and the input from the nasal passages caudally, the individual regions became difficult to distinguish. The nasal cavities were initially filled with coarse, heterogeneous sediment, but this quickly transitioned to fine to medium sediment filling the floors of the passages, continuing into the antorbital and suborbital sinuses, although the sinuses were not separable. The orbits were predominantly void space and fatty tissue, although some fine to medium sediment filled portions of the ventral-most sections in OUVC 10510. OUVC 10511 showed a similar pattern, but with coarser sediment in the left orbit. Sediment was by and large excluded from the orbits by tissue that retained an exterior connectivity even though shrunken and perforated inside.

Sediment was almost completely excluded from the endocranial cavity. The external auditory meatuses were filled almost completely with coarse, heterogeneous sediment, although the proximal margins were finer grained in OUVC 10510 and somewhat more fine-grained sediment near the tympanum in OUVC 10511. Fine-grained sediment also 115 infiltrated the basicranial paratympanic recesses, extending into the caudalmost section of the parasphenoidal rostrum in OUVC 10510, although there was no clear evidence of any sediment in the paratympanic sinuses for OUVC 10511.

Rotten heads.—Four rotting heads (OUVC 10474, 10475, 10501, 10502) were buried in the shallow flume. Sediment filtered through the nasal cavities farther than in other conditions, fining caudodorsally, although sediment influx elsewhere was generally inhibited. Respiratory conchae were clearly visible, with olfactory conchae visible occasionally. Otherwise, sediment influx was very limited or excluded from other routes of possible entry (Fig. 26A–E).

Whereas overall sediment influx and patterns were generally consistent between heads, the extent and specific patterns showed variability between heads. Sediment within the nasal passages of OUVC 10475 filled the nasal passages, with some fine- grained sediment reaching the olfactory conchae as well as the antorbital and suborbital sinuses. Sediment within OUVC 10501 filled only the rostral nasal passages with very little fine-grained sediment extending farther along the floor of the nasal passages.

OUVC 10474 and 10502 were intermediate, both showing sediment in a caudally fining pattern covering the floor of the nasal passages and antorbital sinuses. The degree of fining was reduced, because all four heads were surrounded by a sometimes diffuse layer of fine to medium grained sediment that was thickest by the head and thinned laterally.

Little sediment entered the endocranial cavity or external auditory canals.

Sediment was completely excluded from the endocranial cavity of OUVC 10502, and

OUVC 10474 and 10475 limited influx to a small amount of coarse, heterogeneous 116 sediment in the area adjacent to the foramen magnum. Only in OUVC 10501 did sediment collect to any real extent, filling the floor of the endocranial cavity, fining rostrodorsally. Even here, sediment was limited to the height of the foramen magnum, with most of the endocranial cavity filled by soft tissue, water, and gas. The external auditory canals were at best only partially filled with sediment, with grain size usually fining inwards.

Sediment did not reach into most of the bony recesses. The mesethmoid sinus contained fine grained sediment in OUVC 10475. In OUVC 10501 and 10502, minor amounts of sediment may have infiltrated the basalmost paratympanic recesses, although its identification and how it may have gotten there is unclear. Some sediment may have entered via the external auditory canals, which were subsequently blocked by a shift in soft tissue either during or post burial. Fine sediment may also have entered the paratympanic sinuses via the pharynx by way of the nasal cavities, although this was not traceable. No bony recesses contained any sediment in OUVC 10474.

The soft tissue prevented entry of much sediment into the orbits. Sediment collected in the lower conjunctival recess of OUVC 10475 and 10501. The left eye of

OUVC 10502 ruptured, allowing fine- to medium-grained sediment to fill the lower third of the orbit, merging with sediment from the nasal cavity in the suborbital and pterygoid sinuses. The right eye of OUVC 10502 partially collapsed, allowing sediment to enter the suborbital sinus, but the sediment influx was not as great as in the left eye and did not extend as far. 117

Sediment was almost completely excluded from the oral cavity, unlike heads in other conditions. Small amounts of sediment entered the rostralmost portion of the cavity, but did not extend past the beak tip. In OUVC 10474 and 10475, decay ruptured the right antorbital sinus, allowing a limited spread of coarse sediment to pass through the antorbital sinus into the oral cavity at that point, but sediment did not enter from the margins as seen in other situations.

Clean skulls.—Three clean skulls were buried in the shallow flume (OUVC

10451, 10453, 10456). As with the skulls in the deep flume, all soft tissue other than minor amounts of ligamentous cartilage had been removed. As a result, sediment was able to freely enter most open areas of the skull, although sediment did not enter most of the bony recesses (Fig. 23 C, D). Fine-grained sediment entered the mesethmoid and the basal paratympanic recesses as well as the caudal portion of the parasphenoid rostrum, but did not spread beyond these sections. The endocranial cavities were only partially filled with fine-grained sediment, coarser material limited to the peripheries near the foramen magnum and olfactory tract foramina. Fine-grained sediment entered the endocranial cavities mostly from the rostral direction through the olfactory tract foramina, followed by a coarser layer.

Unlike in the deep flume, sediment around the clean skulls in the shallow flume showed considerable differences inside and outside the skull. Sediment along the ventral margins between the mandibles consisted mostly of finer-grained sediments, with coarse sediment mostly exterior to the mandible. The fine-grained layer was overlain by a coarse, heterogeneous layer. The orbits were filled with sediment similar to that outside 118 the skull. The sediment around the medial skull bones varied between the skulls. Other than a very thin, fine, intermittent layer of sediment overlying the skull followed by an equally thin coarse layer covering the cranium, the sediment around OUVC 10451 was little affected by the skull. The basicranium and parasphenoidal rostrum in OUVC 10456 was surrounded by fine grained sediment, coarsening dorsolaterally, the effect of the skull decreasing dorsally such that the top half of the skull showed little interaction with the sediment. OUVC 10453, however, showed a fine-grained layer covering the full extent of the caudal right orbit, with a diffuse fine-grained layer around the nasals and other bones to some extent. The left side, however, showed little interaction, with the coarse, heterogeneous sediment from the exterior extending to the mesethmoid without disruption.

Long-term sediment patterns

Two blocks containing heads from each decompositional state in both flumes were reburied for seven months and then CT-scanned again. By the end of this time, almost all soft tissue had decomposed. Feathers surrounding the head remained, as did adipocere formed from fat and, surprisingly, brain matter with a mucoid appearance also remained in some heads. All traces of other soft tissue, including the cartilage and ligaments between bones had decayed, causing the skulls to completely disarticulate upon removal from the sediment. Much of the soft tissue was not replaced by sediment, leaving large void spaces. Whereas many of the patterns seen in the initial scans did not hold up over time, there were some notable exceptions. Several heads preserved traces of the 119 rostral respiratory conchae, as well as soft tissue aspects of the external auditory meatuses.

Deep flume

Fresh heads.—The soft tissue of the two fresh heads buried in the deep flume retained for long-term burial (OUVC 10467, 10468) was almost completely decomposed, leaving only bone, feathers, and an abundant amount of adipocere within the orbit, as well as an unexpectedly large amount of mucoid brain matter, which pooled in the ventral third of the endocranial cavity. The ligaments were almost completely decomposed, leaving the bones held in place solely by interdigitating bony processes and the sediment surrounding the skulls.

Sediment collapsed into the oral and nasal cavities, but large void spaces remained (Fig. 20F, G). Many of the previous sediment patterns were lost as coarse, heterogeneous sediment entered the oral and nasal cavities dorsally. Some trace of rostral nasal conchae may have been retained in OUVC 10468, but it was ambiguous. Sediment entry into the orbits was variable, ranging from a complete void to more than half of the orbit and suborbital sinus domain filled with coarse, heterogeneous sediment, limited in medial spread by adipocere. Decomposition widened the external auditory canal, but basic placement and external appearance was still evident, if not as cleanly demarcated as before. Sediment did not enter the endocranial cavity, nor any other bony recess for the pneumatic sinuses. 120

Desiccated heads.—The two desiccated heads buried in the deep flume (OUVC

10459, 10460) were similar to the fresh heads, but showed less disruption from their initial state overall (Fig. 27). With the exception of one ventral conjunctival sac, the orbits were devoid of sediment. Disruption was primarily limited to the oral and nasal cavities. Unlike the fresh heads, the oral and nasal cavities are at least partially distinct rostrally, with most disruption through the antorbital sinuses. Decomposition was more extensive than in the fresh heads, with no residual brain matter, but adipocere formation was extensive.

Despite the loss of all internal soft tissue other than adipocere, respiratory conchae in the rostral nasal passage were still evident within the sediment as thin lines of fine sediment and residual organic traces, as well as some traces of the external integument overlying the antorbital sinuses. The sediment within the oral and nasal cavities was coarse and heterogeneous, much like the initial condition, with the exception of numerous gas bubbles introduced during decomposition. As in the fresh heads, the sediment filled the external auditory canals, but here, the sediment was more restricted in extent and did not pass into the paratympanic recesses, nor did sediment enter the endocranial cavity or the paranasal recesses.

Rotten heads.—The two rotten heads buried in the deep flume retained for long- term burial (OUVC 10472, 10473) showed the preservation of rostral respiratory conchae in one head (OUVC 10472), but not the other (Fig. 22F–I). Overall, sediment influx was not substantially changed in extent, with the exception of increased sediment in the oral cavity of OUVC 10473 due to the decay of the ventral integument, although sediment 121 was disrupted by numerous gas bubbles. Sediment influx was limited by the presence of extensive adipocere deposits, feathers, and organic residuals which aided in stabilization of sediment. Sediment within the nasal and oral cavities of OUVC 10473 did not preserve structures such as conchae due to sediment disruption. Structures were preserved within

OUVC 10472, but distortion caused by decay and sediment movement made identification dubious at best.

Clean skulls.—The clean skulls buried in the deep flume (OUVC 10454, 10455) showed no significant change in the long-term-burial scans other than minor compaction of sediment (Fig. 23E, F). No noticeable additional sediment entered the skulls. The clean skulls buried in the shallow flume (OUVC 10451, 10453) showed a similar pattern.

The largest effect from the compaction was to reduce water content and bring sediment patterns into starker contrast.

Shallow flume

Fresh heads.—The fresh heads buried in the shallow flume retained for long-term burial (OUVC 10476, 10477) retained patterns within the rostral section of the nasal and oral cavities, preserving rostral respiratory conchae traces within the sediment patterns

(Fig. 24G–I). Whereas abundant adipocere and feathers remained, all other soft tissue, including cartilage, had decayed, leaving residual organic carbon stains within the sediment and/or were replaced by clay. Conchal traces were visible in OUVC 10477, but were less evident in OUVC 10476 due to extensive sediment influx and disruption of sediment patterns within the caudal nasal and oral cavities of OUVC 10476. Decay of 122 soft tissue within OUVC 10477 left a void within the sediment that resisted collapse and entry of sediment. The creation of this void led to the obliteration of the sediment traces within the caudal nasal passages created by the respiratory and olfactory conchae as sediment fell to the bottom of the void space. While there was sufficient adipocere within the orbits to preserve sediment patterns, no sediment entered the orbits initially, and none entered even after the eyes decayed. Sediment within the external auditory canals remained unchanged from the initial scans and proceeded no farther into the skull. None of the bony recesses received any more sediment than what they received during the initial burial.

Desiccated heads.—The two desiccated heads buried in the shallow flume retained for long-term burial (OUVC 10465, 10466) were grossly similar in pattern to the short-term-burial scans and that of the fresh heads in that overall patterns were unchanged, but details were reduced or eliminated (Fig. 28). Although large void spaces remained, the sediment was so disrupted from the buildup of decompositional gases and sediment influx that little in the way of soft-tissue patterning could be observed.

Separation between sediment within the oral cavity and rostral nasal cavity was retained rostrally, although sediment collapse into the nasal cavity eliminated clear indications of nasal conchae. Abundant adipocere prevented much sediment from entering the orbits, but substantial sediment did enter the antorbital and suborbital sinuses dorsolaterally. The external auditory meatuses retained what sediment was present but did not gain any more and lost coherency. Unexpectedly, mucoid brain tissue remained, which prevented sediment from entering the cavity. Other than adipocere and feathers, all other soft tissue 123 had decayed. The brain tissue had not been converted to adipocere, but retained a wet, slimy consistency. No new sediment entered other bony recesses.

Rotten heads.—The rotten heads buried in the shallow flume retained for long- term burial (OUVC 10474, 10475) showed surprisingly little change when compared to the fresh or desiccated heads (Fig. 26F–I). The soft tissue decayed almost completely away, leaving only feathers, adipocere, and a small amount of brain matter, along with extensive organic residue throughout the sediment. However, sediment collapse was almost completely absent, leaving an ostrich-head-shaped void space in the sediment.

Most of the sediment within the nasal passages collapsed to the floor of the void space, but some traces of the pattern remained. The rostral respiratory conchae in OUVC 10475 were still visible in the sediment despite the cartilage having decayed. Sediment that had previously entered the mesethmoid sinus settled to the floor of the sinus, leaving the sinus only partially filled. Elsewhere, the sinuses that had been devoid of sediment remained void. The external auditory meatuses did not fill any further than the partial infilling they demonstrated initially, but that partial filling was still evident.

Discussion

Soft tissue surrounding and within the head affects sediment influx

Soft tissue had a strong effect on sediment patterns within the head which could in some cases be correlated with specific soft-tissue structures. In all cases, soft tissue had pronounced inhibition of sediment influx even well after the organic tissue had decayed.

Sediment patterns in both grain size and overall patterns were always distinguishable 124 from the clean skulls. Sediment was excluded from many areas even well after the decay of the tissue. Sediment patterns in the sediment that did enter the remaining skull were also disrupted, not only by the flow of sediment itself, but by the accumulation of decompositional gas creating numerous gas pockets within the sediment. External sediment patterns have been dealt with elsewhere (Chapter 2).Thus, it is worthwhile to more closely examine the internal patterns here.

The preservation of sediment patterns over the long term requires either the sediment to remain undisturbed or that the sediment become lithified before a disturbance occurs, preserving the sediment patterns in a durable matrix. In this experiment, the sediment surrounding all heads buried with substantial soft tissue―those being fresh and rotten heads and to a lesser extent the desiccated heads―accumulated considerable organic residue as a black stain in the general region of the head in the long-term burials.

This material could support a rich bacterial population. Previous studies have shown the role played by bacteria to precipitate minerals helping to stabilize sediments (e.g., Noffke

2010), precipitate minerals within bone (e.g., Grupe and Piepenbrink, 1989; Trueman and

Tuross, 2002) and soft tissue (e.g., Briggs and Kear, 1994a; Wilby and Briggs, 1997), and binding sediment to bone and soft tissue (Martin et al., 2004; Carpenter, 2005). No mineral traces were directly observed during this study, but they were not the focus of this study, and the water used was municipal tap water and not enriched in minerals, which would have inhibited mineralization. Nevertheless, it is likely that some degree of mineral precipitation contributed to the stabilization of the voids seen in this study and the preservation of some of the resulting patterns. Daniel and Chin (2010) found full 125 permineralization of bone as early as six weeks post burial using calcium carbonate saturated waters. Therefore, it is possible that in more mineral-rich waters, more sediment patterns could have been preserved more clearly.

The most likely types of soft tissue to preserve traces within sediment are ones that consist of refractory materials, such as feathers or scales, or adipose tissue, which may turn into adipocere, but their preservation in place may depend upon the more labile tissues. In this experiment, feathers that remained in place were coated with thick organic residue that permeated the sediment, whereas the feathers separated from the head were generally clear of extraneous organics. The organic residue may therefore have been instrumental in helping to stabilize the sediment.

Adipocere may leave behind traces, as well. Adipocere is exceptionally resistant to decay and can last hundreds of years (Fiedler et al., 2009), far beyond estimates needed for soft-tissue preservation via microbial stabilization of both soft tissue and surrounding sediment (Briggs and Kear, 1994a; Dunn et al, 1997; Noffke and Paterson, 2008; Daniel and Chin, 2010). With the attraction of organics to clay particles seen here and elsewhere

(Martin et al., 2004), it is plausible that adipocere formed during burial could be replaced by clay over a long period, allowing possible recognition of patterns within fossils. The chief difficulty here would be distinguishing adipocere residuals from the fine sediment that naturally lines bones in virtually all depositional conditions. One possibility lies in the observation that orbital adipocere tended to partially collapse, forming a deformed cup. Naturally-occurring fine-sediment layers would be coincident with bone surface patterns. Layers of fine sediment replacing adipocere would appear similar along a 126 dorsally buried surface, but may be partially detached from a ventrally located surface, much like a retina detaching from an eyeball. The amount of fine sediment may also be expected to be greater when replacing adipocere than that seen by simple deposition along the borders of a clean skull.

An intriguing possibility for this work is helping to elucidate the visual system of dinosaurs and other extinct animals. Several papers have estimated visual acuity in animals such as ichthyosaurs (Motani et al., 1999), tyrannosaurs (Stevens, 2006), and the circadian behavioral patterns of a variety of dinosaurs, both avian and nonavian (Hall,

2008; Schmitz and Motani, 2011a). However, these measurements were based on scleral rings to estimate the diameter of the cornea and estimates of the overall size of the eyeball, the accuracy of which has been debated (Hall, 2009; Hall et al., 2011; Schmitz and Motani, 2011b). In hominids, it is relatively easy to estimate the size of the eye, as they have more closed eye sockets. It is considerably more difficult to do so in other animals, however, because most animals have open sockets which do not mark a distinct boundary for the back of the orbit. Birds and other reptiles in particular have large adipose layers behind the eyeball. Without knowing how much adipose lies behind the eye, there is substantial latitude in possible estimates. Our work has shown that this adipose layer can be preserved as adipocere, which lasted far longer than other soft tissues. Moreover, this layer, while partially collapsing in our studies, was a reliable indicator of the previous adipose layer. Because of adipocere’s exceptional resistance to decay, it is possible that sediment markers for adipose could be found even when no other 127 soft-tissue preservation has been noted, allowing for better estimation of eyeball size and the resultant behavioral implications to be more firmly drawn.

State of decomposition affects sediment influx

Decompositional state greatly affected sediment influx patterns. Although these differences decreased over the long term as all heads approached the same state of decay, some patterns remained. Soft tissue, as expected, inhibited sediment influx, although the degree to which it did so varied considerably.

Although structural considerations would indicate that fresh heads would allow greater opportunity for sediment infiltration to preserve soft-tissue patterns, this proved not to be the case. The amount of soft tissue prevented sufficient sediment from entering the head while the tissue was still present. Thus, only peripheral structures were preserved. Rotten and desiccated heads allowed more sediment to enter the head and resulted in comparatively better access to sediment. Moreover, the rotten heads allowed greater attraction and adherence of sediment. Unfortunately, the advanced decay and desiccation destroyed many structures before sediment could interact with them. Rotten tissue would sometimes shift, causing blockage of pathways that would ordinarily have been accessible. Thus, preservation potential proved dependent on specific and random microenvironment parameters. Thus, the best preservation potential occurred with suboptimal soft-tissue conditions, greatly (and ironically) limiting preservational capacity. Preservation of deep tissue, such as internal organs thus appears almost impossible. Even if a carcass were opened up so that sediment could gain direct access to 128 internal organs, decay would prevent anything more than the most superficial of impressions. Internal preservations of muscles or intestines would be implausible.

Water velocity and decompositional environment interactions

Burial environmental effects matched predictions in general. Deep, slow moving water resulted in little sediment influx into any aspect of the head other than the most peripheral, and little to no sorting occurred. Moreover, the difference between decompositional states in the deep flume was minimal, other than that between soft-tissue present versus clean skull. The sorting that did occur produced a limited fining upwards pattern consistent with normal settling patterns. In contrast, shallow burial in faster moving water produced much more infiltration of sediment into the head as well as some degree of sorting, with more dramatic differences between decompositional states.

The results were strongly influenced by the specifics of the burial regime and so the applicability of these patterns to real world conditions requires caveats. The deep flume conditions could be representative for conditions in which the flow regime experienced a sudden widening and deepening of the channel, such as floodwaters into an oxbow lake or at the base of a steep slope, in which finer or less dense particles remain in the water column longer than larger and denser particles. However, the heads were not buried in a long-running system, but one set up for test purposes. It is likely that a real- world system would show effects not seen in the tests. Nevertheless, test conditions should have simulated a burial that might occur during a storm event that increased sediment influx for a brief period of time to a sufficient degree that broad scale patterns 129 should be similar. Once the storm was over, sediment deposition should return to normal, indicated by a sharp reduction in grain size, which should be apparent in the sediment and could thus be taken into account. The number of parameters in the shallow system is much more complex and varied, which greatly expands the realm of possibilities and thus has fewer restrictions that can be placed upon interpretations.

One such parameter that would affect results is placement of the head. Because the head was separated from the neck at the base of the skull, hydrodynamic effects caused the head to be most stable in the dorsal up, rostrum upstream position. However, if any part of the neck remained attached, positioning would be very different, which would greatly alter results, particularly in the faster moving streams. Nevertheless, it is a common occurrence for the head to disarticulate from the body early during the decomposition process at the base of the skull (Pfeiffer and Stevenson, 1998) and, given the number of headless fossil skeletons and isolated skulls, is likely to hold true in deep time for a variety of animals. Also, a head separated from the neck would have a tendency to float, which would greatly alter burial patterns. However, while these alterations would affect specific placements of sediment within the skulls, it is unlikely that the major patterns noted here would be altered beyond logical extensions and considerations of the specific burial regime and carcass placement.

Head construction affects sediment influx

This study did not directly evaluate head construction in a rigorous manner, but some conclusions can be drawn from the data that should be applicable to other situations 130 by looking at sediment within the bony recesses and endocranial cavity. Sediment was almost always excluded from bony recesses that housed pneumatic sinuses (Fig. 29).

Clean skulls collected sediment within the mesethmoid sinuses and the basicranial paratympanic sinuses. Fresh heads never collected sediment in any sinus under either burial regime. No condition allowed any sinus to be fully filled. The endocranial cavity had similar results, with even the clean skulls leaving a substantial portion of the endocranial cavities empty.

One can conclude from this that sinuses within fossils that are filled with sediment were filled after the soft tissue had decayed, and most likely after fracture or disarticulation. The specific results here are influenced by the specific burial conditions, altering them would give different results. For instance, turning the skull over could allow the endocranial cavity to fill. However, the architecture of the sinuses would seem to preclude a significant amount of sediment influx so long as the skull is intact.

Utility in Interpretations

Several patterns noted here may be of interest to paleontologists for additional interpretations of fossils and their burial conditions. Sediment patterns within a skull will provide information on the carcass at the time of burial simply by the broad scale patterns within the skeleton, because the patterns within the skulls with soft tissue are inherently different than skulls buried after being scavenged. Thus, if a skeleton was found in any sort of graded or layered sediment, one could easily and quickly determine how much soft tissue was present on the body at the time of burial simply by looking at the amount 131 of disruption within the sediment patterns. Sediment patterns within the confines of a skull that replaced soft tissue would be expected to be less apparent, with the sediment overall being more homogeneous—as compared to the external graded or layered sediment—due to disruptions caused by decompositional gases. Alternatively, these pockets may also have been filled in by mineral precipitation. These filled gas pockets should be recognizable as numerous nodules of similar composition to each other, but not of the surrounding sediment.

Whereas internal soft-tissue domains were generally out of reach of sediment that may have preserved them, peripheral structures were not. Rostral conchae were frequently preserved in sediment patterns (Fig. 30). Such structures have been used to determine metabolic capacities of animals (e.g., Ruben et al., 1996; Ruben and Jones,

2000). In mammals, the turbinates that support conchae are bony, but still preserve poorly due to their very fine structure. In most other animals, these turbinates are cartilaginous and do not preserve at all. In birds (e.g. Struthio), the turbinates are cartilaginous and were decayed, but still left traces within the sediment, traces that would assuredly have been prepared away unless someone were specifically looking for them. Other areas of interest include placement of nares, which has also been a subject of debate in dinosaurs

(Witmer, 2001), a not inconsequential topic as nares placement affects the size of the nasal passage and thus the potential space for conchae. Determining nasal chamber size in mammals is difficult despite having well defined bony parameters. Making such a determination in archosaurs is considerably more difficult due to the presence of the antorbital sinus and soft-tissue boundaries of the nasal chamber. Other peripheral 132 structures that could be elucidated are extent of the external auditory meatuses, as well as fleshy crests or other structures which depend on extraordinary preservation to be found, such as the crests of the pterosaur Nyctosaurus (Bennett, 2003).

Conclusions

The hypothesis that soft tissue affects sediment influx into the head was corroborated, but the ability to determine anatomical information was restricted to features near the body’s surface, such as the rostral nasal cavity, orbits, ear canals, and oral cavity. Decompositional state did affect sediment influx, but not as predicted. Rotten heads allowed greater influx of sediment and preserved the best sediment traces of soft tissue, but decomposition limited preservation of internal structures. Faster flowing water allowed better sediment entry overall and was the only flow state to get sediment into any pneumatic sinus or to get any substantial sediment in the oral cavity. Sediment sorting was only observed in the faster flowing water. Rostral nasal conchae were preserved as sediment traces in most conditions in both flumes. Ear canals were best preserved in fresh heads. The production of adipocere and its resilience to decay has potential to preserve traces of the shape of the eye and the brain even if initial sediment input during burial is excluded. There are definite limits to what sediment patterns can tell us about the soft- tissue anatomy of extinct organisms, but there are some important areas where they can tell us valuable information. Sediment patterns will not give us information on the interior of a heart or the musculature attached to a liver, but it may tell us the difference 133 between closely related species or give us clues to metabolism not recorded in the skeleton.

Many aspects of the effects of burial on soft-tissue preservation need to be examined further. Position of the carcass and potential movement during burial will affect what patterns are seen. Burial in mineral-laden waters could potentially increase preservation of more tissue, and water chemistry in general could have a large impact on preservation potential. Compression during burial could affect sediment flow into the head and the preservation of both the skull and the sediment patterns in and around it after burial. Temperature has a strong influence on decay rates and could potentially affect sediment patterns.

Sediment patterns typically are not studied in detail before the fossils are cleaned.

As a result, we may be destroying information that we can use. Much of this information can be inferred from close relatives or by extraordinary fossils. Yet inference from close relatives will miss the unusual structures. Extraordinary fossils with spectacular preservation are just that, extraordinary and rare. The sediment patterns may provide information for less than obviously exceptionally preserved specimens and thereby turn what may look to be an above average fossil into an extraordinary one.

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Figure 17. Simplified chart of taphonomic factors and their interrelationships. The burial environment includes factors specific to the water and sediment during and post-burial. Exposure time includes factors pre-burial factors and factors non-specific to the burial environment that are time-specific, i.e. are most relevant to the possibility of surviving the short-term decompositional stages, whereas the water and sediment factors may have influence throughout the life of the potential fossil. Specific factors (i.e., not the main groups) indicated in bold were examined in this study. Note grain size was examined in detail, but as a consequence of the other factors, not as a particular factor in itself. 135

Figure 18.Visual confirmation of CT scan sediment patterns. A) axial CT slice through nasal passage of OUVC 10458, a desiccated head buried in the deep flume. B) physical slice through sediment block in same position. C) fresh head preburial, red line indicates placement of slices shown in A and B. D and E are expanded views of the areas indicated by the black boxes in A and B, respectively. The small amount of sediment covering the floor of the nasal passage is clearly indicated in the CT scan (red arrows), despite being relatively inconspicuous in the physical slice (black arrows).

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Figure 19. CT density variations can distinguish sediment grain-size parameters. A) Axial CT slice through nasal passage of OUVC 10471 (fresh head, shallow flume). B) Slice through sediment block containing the same head. The arrows indicate locations sediment parameters were estimated. The numbers correspond to the sample reported in Chapter 2, Appendix 3a. C) Graph of 500 density values (in Hounsfield units) for each sediment sample. The numbers in parentheses are the parameters estimated from high-resolution photographs (see chapter 2, the first digit indicates most dominant grain size, second number indicates range, third number indicates clay contribution). Coarser and more heterogeneous sediments had on average higher density than finer sediments, as well as having higher variability. Fine grained, homogeneous sediments with high clay contributions had lower density and less variation.

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Figure 20. Sediment patterns in fresh heads buried in the deep flume. A–C) 3D reconstructions of sediment patterns formed during initial burial of in OUVC 10467 139 mapped via the Amira software. Density varies from very low (red) to very high (orange). The brownish color of the sediment is due to the high heterogeneity of densities within the sediment. A) Lateral view. B) Frontal view. C) Dorsal view. The soft tissue is ghosted to provide localization reference. D) Sagittal CT slices of OUVC 10467. E) Axial CT slice. Orange line indicates position of the adjacent slice. Note the respiratory conchae plainly visible in the sediment (white arrows). F, G) CT slices from the same head after seven months additional burial showing collapse of sediment into ventral portion of head.

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Figure 21. Short-term sediment patterns in desiccated heads buried in the deep flume. A– C) 3D reconstructions of sediment in OUVC 10457 using Amira software. Soft tissue is shown in ghosted green to denote placement of sediment displayed on a red to yellow scale, with red being low density and yellow being high density. A) Lateral view. B) Frontal view. C) Dorsal view. D) composite CT image showing horizontal sections at two different levels, left is caudal, showing external auditory meatuses containing sediment (red arrows), right is rostral, showing conchae within sediment (white arrows). Orange lines denote position of sagittal and axial views shown in E–G. E) Sagittal view showing sediment in nasal passage. Orange lines denote position of coronal views. Respiratory conchae can be seen in all images but F (white arrows). 142

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Figure 22. Sediment patterns within rotten heads buried in the deep flume. A–C) 3D reconstructions of sediment in OUVC 10472 using Amira software. Soft tissue is shown in ghosted green to denote placement of sediment displayed on a red to yellow scale, with red being low density and yellow being high density. A) Lateral view. B) Frontal view. C) Dorsal view. D, E) Sagittal and axial CT slices, respectively. Orange lines denote placement of each view compared to the other. F–I) CT slices of head after seven-months additional burial. F) horizontal slice, orange lines denote placement of views in G–I. White arrows show respiratory conchae preserved within sediment. Red arrows indicate adipocere preserving fat tissue.

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Figure 23. Sediment patterns of clean skulls. Left: sagittal CT slices. Right: axial slices through orbits showing mesethmoid (white arrows) and parasphenoid rostrum (red 145 arrows) pneumatic recesses, the only pneumatic recesses sediment typically entered. A, B) Deep flume, OUVC 10454. C, D) Deep flume after seven months. E, F) Shallow flume, OUVC 10451. G, I) Shallow flume after seven months. Other than minor sediment settling and replacement of water with decompositional gases, there is no substantial change between short and long-term sediment patterns. 146

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Figure 24. Sediment patterns within fresh heads buried in the shallow flume. A–C) 3D reconstructions of sediment after short-term burial in OUVC 10471 using Amira software. Soft tissue is shown in ghosted green to denote placement of sediment displayed on a red to yellow scale, with red being low density and yellow being high density. A) Lateral view. B) Frontal view. C) Dorsal view. D) Head reconstruction, dorsal view, showing placement of sagittal and axial CT slices shown in E and F, respectively. G–I) CT slices of head after seven months additional burial. Orange lines in sagittal slice (G) show placement of axial slices (H, I). This set shows the most extensive sediment pattern within the oral cavity, as well as preserving rostral conchae in the nasal cavity, although these patterns did not survive after decomposition of the soft tissue. 148

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Figure 25. Variation of short-term sediment patterns seen in CT slices of desiccated heads buried in the shallow flume. A–D) OUVC 10465. A) Sagittal slice.B-D) Axial slices. E– H) OUVC 10466. E) Sagittal slice. F–H) Axial slices. I–L) OUVC 10510. I) Sagittal slice. J-L) Axial slices. M–O) Horizontal slices through OUVC 10465, 10466, and 10510, respectively. Orange lines through sagittal slices denote placement of horizontal slices. Orange lines on horizontal slices show placement of sagittal and axial slices. Despite overall variation, all heads preserved rostral conchae traces (white arrows).

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Figure 26. Short-term sediment patterns seen in CT slices of rotten heads buried in the shallow flume. A–E) OUVC 10475. A) Horizontal slice. B) Sagittal slice. C–E) Axial slices. Orange lines on A denote placement of B–E. Orange line on sagittal slice denote placement of A. F–I) OUVC 10475 after seven months after all soft tissue but feathers and adipocere have decayed away. F) Horizontal slice. G) Sagittal slice. H, I) Axial slices. Orange lines on F denote placement of G–I. Sediment traces of conchae were preserved in long-term burial (white arrows) and orbital fat was preserved as adipocere (red arrows).

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Figure 27. Variation in desiccated heads buried in the deep flume after seven months seen in CT slices. A–C, E) OUVC 10459. D, F–H) OUVC 10460. Orange lines in horizontal slices shown in C and D denote position of sagittal and axial slices in A, B, and E–H. 153

Despite similar conditions, the amount of disruption is considerably different, allowing conchae (white arrows) to be preserved in OUVC 10459, but not in OUVC10460, for instance. Adipocere was preserved in both, although it was better preserved in OUVC 10459 (red arrows).

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Figure 28. CT slices from desiccated heads buried in the shallow flume after seven months. No soft tissue remained other than feathers, adipocere (red arrows), or brain matter. A–G) OUVC 10466. Orange line on sagittal slice in A denotes position of horizontal slice in G. Orange lines in G denote positions of sagittal slice in A and axial slices in B–F. H) OUVC 10465 axial slice through rostral nasal cavity. Traces of rostral respiratory conchae were equivocal in OUVC 10466, but were more evident in OUVC 10465 (white arrows). OUVC 10465 was otherwise similar to OUVC 10466.

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Figure 29. Pneumatic sinuses that collected sediment. A) Horizontal slice showing locations of axial and sagittal sections shown in B–E. Image is from skull shown in B and D, but locations are approximately correct for C and E, as well. B) Axial slice, rotten head, shallow flume. C) Axial slice, dried skull, shallow flume. D) Sagittal slice, rotten head, shallow flume, same head as B. E) Sagittal slice, clean skull, deep flume. Arrows in B, D, and E point to sediment in mesethmoid sinuses, arrow in C points to sediment in parasphenoid rostrum. 157

Figure 30. Axial slices through the nasal passage of OUVC 10475, rotting head in shallow flume, showing preservation of conchae in sediment. Arrows indicate conchae. A) CT scan after initial burial. B) CT scan after seven months. C) Photograph of sediment block slice. All organic tissues except feathers, bone, and adipocere have completely decayed in B and C. The conchae were preserved as clay suspended in water surrounded by coarser sediment most clearly seen in the left (from viewer’s perspective) nasal passage.

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CHAPTER 4: NEW INTERPRETATIONS OF FOSSILS BASED ON CT-BASED,

ACTUALISTIC TAPHONOMY STUDIES

Abstract

Interpretation and reconstruction of fossils have typically rested on homology, analogy, and the rare exceptional fossil. Recent taphonomic studies, however, have provided a new way of looking at both the fossilization process and the fossils themselves. Although not providing detailed information on reconstructions of animals, they do help to refine interpretations and point the way to further explorations of usually ignored data. The origin of void spaces within fossils can be interpreted, providing information on burial conditions. These studies indicate, for example, that the preserved intestinal tract in the small theropod Scipionyx is unlikely to have been preserved from the exterior and is potentially a unique case of a fossilized impacted bowel. Study of decay patterns makes the controversial claims of a preserved dinosaur “heart” increasingly unlikely, although conchae within Panoplosaurus are possible. Integument slippage during decay can be taken into account to determine how much skin is preserved in a given fossil, as well as demonstrating the limits of interpretability in specimens such as Sinornithosaurus and Sinosauropteryx, which, while supporting the presence of feather-like integumentary appendages, are not as clear-cut as supporters of either side of the debate makes it out to be. There also seems to be less chance of finding internally preserved organs in the hadrosaur mummies “Leonardo” and “Dakota” than might be hoped considering their unusually high levels of preservation. Tissues around the 159 periphery, such as in the nose, ears, mouth, and eyes are more amenable to preservation, so study (and preparation) should be especially careful in these areas. Taphonomic questions remain as to why some fossils show exceptional preservation while others in similar situations, or those situations that should be optimal for soft-tissue preservation, do not.

Introduction

Fossil interpretation is typically done in several ways, which can be broadly grouped into biological and geophysical methods. Biological methods are mostly morphological studies using comparative methods that have become increasingly guided and constrained by use of the Extant Phylogenetic Bracket approach (Witmer, 1995) which emphasizes homology but also is supplemented by comparative analogy studies when the region of interest involves character novelty or lack of clarity in homology, phylogeny, or environment. Our understanding of such diverse structures as brains and ears, (e.g., Kundrát, 2007; Zhou et al., 2007; Witmer and Ridgely, 2008; Witmer et al.,

2008) as well as structures that often leave little or no physical trace, such as nostril position, skin, or appendices (Witmer, 2001; Hieronymus et al., 2009; Smith et al., 2009) and behaviors implicated by these structures (e.g., Witmer et al., 2003; Witmer and

Ridgely, 2009).

Numerous taphonomic studies have contributed by analyzing (1) disarticulation, transport, and preservation potential of skeletal elements; (2) environmental parameters that aid in determining ecological and behavioral characters; and (3) mode of 160 preservation (for reviews see Lyman [1994] and Martin [1999]). However, detailed studies using taphonomic methods to determine constraints of morphological interpretation or possibly even enhance reconstructive interpretations are more limited and are almost completely limited to studies of invertebrates (e.g., Briggs and Kear,

1993; Norris, 1989; Plotnick, 1986,) or primitive chordates (Briggs and Kear, 1994b;

Sansom et al., 2010). Recent studies on ostrich heads are beginning to help elucidate the interaction of soft tissue with sediment during burial (Chapters 1–3) and the relative chances of preservation for various tissues (Fig. 31). These studies help constrain how far interpretation can be taken as well as enhancing some interpretations when soft tissue is not directly preserved. This chapter will examine several fossils that have been previously reported and discuss the findings in light of the new taphonomic interpretive possibilities. What follows are a series of case studies. As such, there are no methods to report, other than the use of the software program Amira 5.3, which was used for visualizing three dimensional relationships.

Preservation of Internal Soft-Tissue Structures

Almost all soft tissues preserved as fossils are derived from “external” tissues, meaning soft tissues near the surface of the body. Internally located tissues (e.g., internal organs) are exceedingly rare, and the few that have been found are generally hotly debated (e.g,. Fisher et al., 2000; Rowe et al., 2001; Cleland et al., 2011). Gut contents, referring to partially digested matter, are more common and less ambiguous than preserved soft tissue of the actual animal, but, as gut contents tell us about what the 161 animal ate, they do little to aid in the reconstruction of the animal itself. Gut content preservation is sometimes associated with preservation of other tissues, but a full description of gut-content taphonomy is beyond the scope of this chapter. Preservation of external integumentary structures, while rare, is relatively straightforward and fairly easily understood. The integument (as well as any epidermal appendages) has direct contact with the environment, which allows impressions to be made in the surrounding sediment. This location also provides access to both endogenous and exogenous bacteria that may induce rapid permineralization of the tissue (Briggs and Kear, 1993; Briggs,

2003a, b). These bacteria also have access to exogenous sources of nutrients which may aid the induction of mineralization. Epidermal tissues are keratinized to varying extents, making them much more durable than other tissues, increasing their resistance to degradation and allowing them to persist for long periods, thus enhancing their chances of preservation as fossils (Schopf, 1975; Briggs, 1999).

By contrast, internally located soft tissues are mostly protected from the environment. Sediment impressions will not be formed, except with tissues exposed to the environment due to rupture of the integument. The tissues contain a ready source of lysosomal enzymes which initiate autolytic degradation (Gill-King, 1997). Endogenous bacteria within the gut will also initiate decay of the carcass from within unless the temperature of the body is swiftly reduced. Most internal soft tissues decay quickly at room temperature (Mann et al., 1990; Megyesi, et al., 2005), even when shallowly buried and protected from scavengers (see Chapter 3) unless the whole organism is small enough to completely mineralize,such as embryos (Xiao and Knoll, 1999; Donaghue and 162

Dong, 2005; Yin, Z.J. and Zhu, M.Y., 2012), bacteria (Bailey et al., 2006), or sporocysts

(Huldtgren, T. et al., 2011) or be entombed in amber, which has its own preservational issues (Beck, 1988; Cerling, 1989).

Decay of internal structures as seen in experimental taphonomic work (Chapters

1, 3) has several ramifications for fossil preservation and interpretation. The interpretation of void spaces within fossils—regarding their origins, soft-tissue interpretation, and consequences for preservation—may now be more fully delineated.

We also may be able to say something about the initial taphonomic conditions of the carcass based on originally air-filled cavities that are no longer empty space and provide more parameters for characterizing exceptional finds.

Void Spaces

The amount of endogenous air space within the sinuses and airway of vertebrate heads, coupled with the rapid decay of internal soft tissue, creates abundant void spaces within rapidly buried carcasses. Their ultimate manifestation within a fossil can be categorized into four main types (Table 6).

Type 1 void space.—The first type is a primary void, meaning the void existed in the living animal and was never filled in. In my actualistic experiments on ostriches, sediment rarely entered most sinuses to any great degree as long as the skull was intact

(Chapters 1, 3). Even if the skull had been cleaned prior to burial, sediment did not enter most of the paratympanic recesses, as well as the portions of the paranasal pneumatic recesses distal to the recess openings. Therefore, type 1 voids are the most likely 163 explanation for void spaces within many fossil bones, such as those seen in

Tyrannosaurus (FMNH PR 2081, Fig. 32). In addition to the air trapped within sinuses, decay adds to the air volume by the release of decompositional gases such as methane and carbon dioxide.

Type 2 void space.—Voids can be filled in secondarily through mineral precipitation, either through microbially mediated precipitation (Carpenter, 2005; Daniel and Chin, 2010) or through inorganic precipitation (Trueman and Tuross, 2002). Infills are most likely to be either a calcium carbonate or calcium phosphate, such as Allosaurus

(UMNH VP 18050, Fig. 33) and in the London specimen of Archaeopteryx (BMNH

37001; Davis, 1996), or iron in the form of iron sulfides or iron oxides. Carbonate infills may be difficult to distinguish via computed tomographic (CT) scans, as the density of the infill would be similar to that of the surrounding bone. Iron, however, would be much more easily seen and mapped (Allosaurus, UMNH VP 18046; Fig. 34), and typically are the cause of a great deal of CT scan artifacts due to their density.

The high-density nodules in the Allosaurus premaxilla shown in Figure 34 look to be deposited mainly within the pulp cavity of the teeth and the alveolar and trabecular portions of the premaxilla. These are likely mineral infillings of voids created by decomposing soft tissue. One might expect to see void spaces created by decompositional gases, but these would not be discernible from original air except in the case of voids outside the bone, which would be created by decomposition (air in pneumatic sinuses would either be replaced by fluid or be mixed with decompositional gases, the spaces themselves potentially distorted beyond recognition by decomposition for those sinuses 164 solely bounded by soft tissue). These voids would likely be preserved as a collection of nodules concentrated around the orbital and nasal cavities as these areas are most likely to both collect gases and serve as points of entry for sediment, the interaction of which is necessary for the preservation of gas bubbles discernible as originating from decomposition (Chapter 1).

Type 3 void space.—Based on examination of a large diversity of CT-scan datasets, most fossils are filled with sediment and do not exhibit large numbers of void spaces. This third type of void spaces is indicative of several possibilities that may be distinguishable from each other, but not mutually exclusive. First, the bones would need to be buried after thorough cleaning by scavengers or simple long term exposure to the elements, allowing sediment direct access to the bone creating greater influx of sediment.

This process will not completely fill the pneumatic sinuses or even the braincase, much less introduce sediment into marrow cavities. The processes leading to cleaned bones are often damaging to the bones themselves, however. Long-term exposure weakens and cracks bone, and scavengers cause both mechanical and chemical damage to bones.

These processes can open up cavities within the bone allowing influx of sediment.

Mechanical damage during the burial itself may also do so.

Soft-sediment flow can bring sediment further into the skull with less damage.

Bioturbation, such as that noted by Colbert and Eberle (2007) in an Eocene skull of the tapiroid Thuliadanta, may bring sediment into the interior recesses. Provided the carcass is not buried too quickly and remains within the zone of bioturbation (Aller, 1982; Davies et al., 1989), burrowing scavengers may carry sediment deeper into the carcass as a result 165 of sediment disturbance, although this method is limited to the size of the burrower.

Unless the burrowers are small enough to fit inside a sinus or a portion thereof, they will not be effective sediment distributors. Compression early in the burial process while the sediment is still loose and capable of flow during deformation can carry sediment deep within the sinuses, although it is unknown how far and under what viscosities and pressure this method is effective before either inhibiting flow or causing overt damage to the bone. Void spaces filled in this manner will show a homogeneous mixture of sediment, although the finer the sediment, the more efficient this process should be.

The holotype skull of Nanotyrannus (CMNH 7541; Gilmore, 1946; Bakker, 1988;

Witmer and Ridgely, 2009, 2010; note: there is controversy as to whether this taxon is valid or the specimen pertains to a juvenile of Tyrannosaurus rex, but that debate is peripheral to the discussion here, and we use holotypic name for convenience) appears to be an excellent example of soft-sediment entry after decay (Fig. 35). The skull was buried in fine-grained sediment of the Hell Creek Formation. For the most part, it is in excellent shape with sediment partially infilling many pneumatic recesses, particularly in the basicranium. CT scans revealed infillings that are less dense than the surrounding matrix.

This can be representative of either finer-grained or less compact sediment. The surrounding matrix is fine-grained itself, so the distinction is hardly important, but does indicate that the sediment was still fluid at the time of entry into the recesses. The skull was found without any associated postcrania. Actualistic taphonomic studies (Davis,

1996; Brand et al., 2003; see Chapters 1–3; but see Bickart, 1984) have indicated that, although the skulls of mammals and birds (but not amphibians and lizards) disarticulate 166 from the body fairly early, the skull as a whole is more resistant to disarticulation than is the atlanto-occipital joint, so it is expected that the Nanotyrannus skull had already undergone considerable decay before being buried, at which time, the soft mud could then enter the recesses.

A fourth type of sediment infilling is indicative of slow, pulsed influx of small amounts of sediment. In this case, the sediment-to-water ratio is low, and the carcass gets buried over a longer period of time, resulting in a layered appearance to the sediment in the cavities, as well as a greater likelihood of the bone suffering more damage due to the time exposed on the surface. The premaxilla of the lambeosaurine hadrosaur

Hypacrosaurus (ROM 702) in Figure 36 illustrates this type of infilling with sediment stratified horizontally like bedding planes, rather than circumferentially as one might expect in concretionary deposition. In addition to indicating taphonomic condition of the hypacrosaur, it provides spatial orientation of the premaxilla during burial as well as data on the flow regime. Sinuses that are filled more rapidly in one major pulse would contain sediment that is more mixed and consistent, and more texturally similar to the surrounding matrix, such as that seen in the maxillaries and left jugal of Rhabdognathus

(CNRST-SUNY 190; Brochu et al., 2002). Our CT scan data of that specimen not only show recesses filled through breaks in the bone, but also primary void space around the less damaged braincase. Another example of this is Deinonychus (OMNH 50268), which shows some air-filled recesses (primary voids), but sediment in the larger ones. The sediment is heterogeneous, but clearly entered post-scavenging. The skull is incomplete and likely scavenged or otherwise damaged before burial. 167

Type 4 void space.—Type 4 void spaces are those in which voids have been previously infilled by mineralization and then secondarily dissolved out. These are expected to be rare and may be only theoretical. Any situation that would dissolve the mineral infillings would likely also act on the surrounding bone. Thus, for these to occur, the mineral infilling the cavities would have to have a lower threshold of dissolution than the surrounding bone. Otherwise, the bone would dissolve preferentially as it would be in more contact with the groundwater due to the bone surrounding the void space. Not only would it come into contact with the groundwater first, but it would have a greater surface to volume ratio than the mineral within the cavities making it much more likely to dissolve.

Void Spaces: Structural Consequences

Carcasses buried with significant amounts of soft tissue will form large void spaces as the internal tissues decay and drain away. Such large void spaces are structurally unstable and will usually either collapse or get filled in. How this happens will depend on the conditions of the carcass during sediment influx—which itself is partially dependent on the size and type of carcass—and the parameters describing the sediment, including size distributions of grains, composition, and degree of water saturation, as well as depth. Studies to examine this in detail are few and do not address most aspects, but some aspects can be examined in the context of fossil preservation.

There are three main possibilities that can result in sediment filling in large cavities in a skull. (1) The skull will stay in its original configuration as the sediment fills 168 it in, which is obviously the preferred condition for paleontologists but is the rarest of the three possibilities. Far more likely is that portions of the skull will at least partially collapse, which leads to the other two possibilities: (2) the skull may fall apart as the ligaments connecting the bones decay, or (3) the skull may simply be crushed by the overlying sediment.

These possibilities are not mutually exclusive, and they may all appear in the same skull, controlled primarily by the skull’s construction. As demonstrated by taphonomic studies (Trapani, 1998; Brand et al., 2003; Chapter 3), the neurocranium, particularly the basicranium, is the most durable section of the skull. Thus, it is more likely to remain preserved in its original configuration than other parts of the skull. The bones in the rostrum are much more likely to separate and break than the relatively compact braincase, because, being longer and thinner, they are more fragile. The bones of the rostrum are also more commonly joined by overlapping or grooved contacts compared to the interlocking sutures of the cranium. When the connective tissue between the bones decays, the rostral bones are therefore much more likely to disarticulate. This explains why it is not uncommon to find skulls in which the neurocranial bones of the braincase are intact and articulated, whereas the rostral bones are disarticulated, with the generally lighter bones in the rostrum, such as the palatines, crushed and the usually stouter rostral bones, such as the premaxilla, intact.

Understanding these possibilities is not entirely academic. Being able to tell the difference between these taphonomic possibilities may be of interest during reconstruction of the skull when attempting to determine the type and amount of 169 diagenetic alteration that needs to be considered. Skulls that began to disarticulate before being crushed may give deceptive skull reconstructions if this situation is not noted.

Bones connected by overlapping sliding joints, as is common in the rostral bones, can sometimes be difficult to place accurately. Warping (plastic deformation) of the bones post-burial may also be influenced by the type of collapse undergone by the skull.

Additionally, distinguishing pre- and post-burial breakage could have taphonomic implications for paleoecological interpretations.

The classic Tyrannosaurus rex skull (AMNH 5027; Fig. 37) described by Osborn

(1912) illustrates some aspects of void space and decay. This skull is in excellent condition, showing little more than slight deformation of the skull, indicating that the skull was likely at least partially buried while retaining some soft tissue. Before sediment filled in the skull, however, sufficient decay occurred to allow the right ectopterygoid to disarticulate, falling into the left antorbital fenestra. The placement of this bone and the warping of the skull also indicate the head lay on its left side during burial. A juvenile

Tarbosaurus skull described by Tsuihiji et al. (2011) shows further developments along this line. As preserved (Fig. 38A), this skull is mediolaterally compressed, but is in relatively good condition. The bones are not crushed for the most part. Instead, they appear to have disarticulated before sediment influx and compression occurred, allowing the bones to shift to some degree, thereby preserving much of their integrity. They are not out of position, indicating that the skull was still articulated during burial, thereby indicating the presence of soft tissue during burial. In contrast, the skull of Ikechosaurus gaoi (Fig. 38B) is simply crushed and broken, apparently collapsing due to compression 170 of the sediments. The Sinornithosaurus specimens reported by Kobayashi and Lu (2003) are also described as being crushed, a common feature of many fossils from Liaoning.

The well-articulated specimens and the crushed nature of the specimens suggest burial occurred either at or shortly after death, the fragility of these skulls contributing to their fracturing before disarticulation. Smaller skulls with thinner bones are more likely to be broken than larger, more solid skulls, which could resist fracturing and may be more likely to disarticulate when compressed by soft sediment.

Internal Soft-Tissue Preservation Examples

Although soft tissue typically decays, leaving a void in its place, there are some examples in the literature of specimens that appear to have preserved significant quantities of soft tissue. In light of the recent taphonomic work on soft-tissue preservation, it is worth examining them with this new perspective. These examples include a fish brain, a dinosaur with preserved abdominal organs, and a dinosaur heart.

Fish Brain.—Pradel (2007) reported a 300 million year old chimaeroid fish with a phosphatized structure that they refer to as a “purported” brain, leaving open the possibility that it could be something else. The structure is preserved in three dimensions and lies on the floor of the braincase. It is less than two millimeters thick. Due to the position and thinness of the material, their assessment that it is indeed phosphatized residual brain tissue is likely correct. The polymerization of lipids within the brain can preserve brain matter for a considerable amount of time under the right circumstances

(Gill-King, 1997; Gupta and Briggs, 2011). The small size of the material is probably 171 what allowed its preservation. They are also likely correct in their assertion that this preservation resulted from microbially mediated phosphatization in anoxic, phosphate- enriched waters enhanced by the fall of pH caused by the carbon dioxide produced during decay and the release of fatty acids from the lipids present in the brain tissue.

Scipionyx.—One of the most remarkably preserved fossils is that of Scipionyx, reported by Dal Sasso and Signore (1998), which preserved muscles within the pectoral region and tail, possible liver traces, tracheal rings, and most remarkably, a complete intestine. The specimen is flattened, but the intestine is preserved in three dimensions.

Additionally, the intestine is preserved as an endocast, showing striations interpreted as transverse folds of the gut (Dal Sasso and Signore, 1998; Halls, 2003). Interestingly, no epidermal remains are present, except for claw sheaths (Dal Sasso and Maganuco (2011).

Whereas it is difficult to imagine the taphonomic situation that would cause this, it is also difficult to imagine that it could be otherwise as the preservation is so spectacular. Intestinal tissue is one of the first tissues to decay due to the digestive enzymes and endogenous microbial fauna of the gut (Gill-King, 1997). Dal Sasso and

Signore (1998) reported that the intestines did not themselves preserve, but what was within them. However, Dal Sasso and Maganuco (2011) reported that further work found traces of intestinal tissue preserved. It also seems impossible that the intestines could have been sediment-filled during burial. Holtz et al. (2004) suggested that what was preserved was not tissue preservation of the dinosaur itself, but preservation of what would have been fecal material within the intestine, a cololite. As a carnivore, the meat would have provided a nutrient supply rich in potassium, iron, and calcium, all of which 172 would have supported mineralization by endogenous bacteria (Bradley, 1946; Hollocher et al., 2001; Chin, 2002; Hollocher et al., 2005). A fair question then would be why so much of the intestine was preserved in such a manner. It may be that less intestine is preserved than thought. Dal Sasso and Signore (1998) did comment on the short length of the intestinal tract, although one would expect a short tract in a carnivorous juvenile. It is also possible this animal was suffering from an impacted bowel, possibly due to the ingestion of sand, as occasionally is seen in horses and dogs (Udenberg, 1979; Papzoglou et al., 2004), which would explain the sand within the gut as well as demonstrating behaviors not normally capable of being preserved in fossils. Dal Sasso and Maganuco

(2011) call the suggestion of a cololite as being simplistic, but it seems clear from their description that most of the preservation is indeed a cololite with the addition of some tissue preservation as well.

The “Dinosaur Heart”.—Fisher et al. (2000) made quite a stir when they published CT scans of a Thescelosaurus nicknamed “Willo” with a purported three- dimensionally preserved heart, complete with ventricles and an aorta. Almost immediately, others disputed the finding, with Rowe et al. (2001) concluding it was nothing more than a concretion. Whereas it is fair to wonder whether Rowe et al. may have set an unreasonably high bar for the level of detail required to consider it a heart, on the basis of the experimental taphonomy results presented elsewhere (Chapters 1, 3), they were likely correct in their opinion that the “heart” is a concretion. The lack of sediment entry into the interior of the heads as reported in chapters 1 and 3, along with the decay of internal tissues preventing the preservation of sediment patterns that would support the 173 preservation of a heart make that interpretation problematic. The taphonomic work discussed here therefore supports the histochemical and CT reexamination of the “heart” by Cleland et al., (2011). Nevertheless, it is certainly possible that the concretion was formed from the iron-rich residues that would be expected from the heart muscle, even if it did not preserve intact.

The “heart” as reported by Fisher et al. (2000) had ventricles filled with iron-free quartz silt. It is difficult to postulate any scenario that would allow sediment to fill the ventricles without destroying the heart in the process. Sediment would not have been able to enter the heart unless the ventricles were open to the environment, in which case one would have to wonder how the heart could have been present to begin with. In accordance with the decay patterns reported in chapter 3 and as argued previously

(Chapters 1 and 3; Gill-King, 1997), the soft tissue of the heart would have decayed long before it could have been mineralized to that extent, as it is highly nutrient-rich and is readily consumed by decomposers, much earlier than the cartilaginous tendons that were not preserved elsewhere on the carcass. Mineralization of that much non-keratinous soft tissue has never been demonstrated. The authors supposition that saponification may have allowed preservation is inadequate. Saponification is a good explanation for adipose tissue (Fig. 39), which preserves well and in proper conditions can last for hundreds of years. But muscle tissue decays rapidly, leaving few traces, even in situations in which saponification completely preserves the adipose tissue. The only other tissue that might be preserved through saponification would be the brain, due to its high level of lipids

(Gill-King, 1997). While not preserved structurally, brain tissue was preserved in a few 174 of the ostrich heads studied in the actualistic taphonomy studies presented previously, although not in the same form as adipose tissue. The brain tissue did not form adipocere, but rather a gelatinous sludge (Chapters 1, 3).

One complicating factor making extrapolation from the taphonomic studies problematic is that the work done on the mineralization of soft tissue (Briggs, 2003a;

LaBarbera and Kowaleski, 2004) has never dealt with temperature differences. It is possible that a sharp and extreme drop in core temperature may have preserved the tissue long enough for mineralization. However, that still leaves open the question of how the sediment entered the purported ventricles as well as how such a drop in temperature could have occurred to an animal living in the Hell Creek Formation environs, which are thought to have been rather warm (Lehman, 1987; Nudds and Selden, 2008).

Orbital fat.—A possible example of soft tissue not previously reported is that of

Diplodocus (CMNH 3452). This specimen shows a vague and faint partial cup shape within the left orbit that may be indicative of residual adipose around the eye. If it is, the adipose cup has partially deformed and is not complete, but this is to be expected

(Chapter 3). Only in the best preserved specimens do the adipose cups faithfully mark the boundaries of the eyeball. The rest of the skull does not have any other noticeable organic signatures. The sediment within the skull is fairly homogeneous, but appears to have inorganic, heterogeneous packing, limiting any possible organic signature, as does the CT scan beam hardening artifacts present and the limits of the CT scan resolution itself. 175

Respiratory conchae.—The existence of respiratory conchae in dinosaurs has been much debated. In extant species, they play an arguably important role as a physiological regulating mechanism for endothermic animals by conditioning the air and serving as heat exchangers to moderate respiratory evaporative water loss, and thus their presence or absence may thereby provide important clues to dinosaurian thermoregulation. Unfortunately, mammals are the only animals that routinely form bony turbinates within their respiratory conchae (although some birds form bony olfactory turbinates). Ruben et al. (1996) asserted that dinosaurs did not have respiratory conchae.

However, problems with the accuracy of some of the reconstructions used in their study and their small sample size call into question their results. Nevertheless, little definitive evidence for them has been published, although indications that they existed have been presented (Witmer and Sampson, 1999; Witmer and Ridgely, 2010). Therefore, further study of the narial anatomy of two ankylosaurids published by Witmer and Ridgely

(2008) could shed light on this debate.

The rock matrix found within Panoplosaurus (ROM 1215; Fig. 40) is coarser rostrally within the nasal passage than elsewhere in the skull, as one might expect. It appears that the head was initially buried upside down with at least some soft tissue still present. The coarse, heterogeneous material situated dorsally in the nasal passage indicates the orientation and the degree of penetration of sediment during the initial burial, the remainder of the skull being filled in with finer sediment that infiltrated over time as tissue degraded and fine material was either brought in suspension and settled out or via bioturbation. There are several small areas of less dense material within the caudal 176 section of the rostral loop of the nasal chamber that suggest the presence of soft tissue affecting sediment deposition within the skull. These may indicate possible simple cartilaginous rostral nasal conchae along the lines of those present in the ostrich rostrum.

A difficulty in this interpretation is that the regions are situated dorsally in the chamber, when one might expect them to be ventrally located if the heads were buried upside down. However, these areas are also places in which mucus would collect in an overturned head. Any collected mucus would have blocked sediment from filling those regions and would preferentially be replaced by finer sediment as it decayed. There is a bleb of less dense material within the braincase that may indicate these blebs are artifactual, but may also indicate the presence of residual brain tissue similar to the fish described above. There is not enough to see any features, but its position at the top of the braincase, which would have been the lowermost portion during burial, supports the possibility of residual tissue. Additionally, the trabecular bone within the skull contains void space which is likely primary (type I) and high density nodules, which are likely mineral infilling (type 2).

Euoplocephalus (AMNH 5405) has much more complex nasal passages than does

Panoplosaurus. There is no evidence here of anything resembling rostral conchae, although the olfactory conchae seem evident (Witmer and Ridgely, 2008). Here again, the sediment patterns indicate that the head was preserved upside down, with the right side tilted upward. If these interpretations are correct, and Panoplosaurus had rostral conchae whereas Euoplocephalus did not, this may suggest a possible explanation for the unusually tortuous nasal passage in Euoplocephalus. It is conceivable that nodosaurids 177 developed simple conchae to expand the surface area of the nasal passage, but the ankylosaurids took an alternate route by lengthening the route itself. These interpretations are admittedly speculative, but they are consistent with the data.

Preservation of External Soft-Tissue Structures

In contrast to internal soft tissues, external tissues are relatively common and much less ambiguous. Many dinosaur skin impressions have been found (reviewed by

Carpenter, 2007), as well as fish scales and skin from a variety of other animals. Some of the most spectacular have come from the Liaoning province in China and have revolutionized our understanding of the evolution and anatomy of theropod dinosaurs, as well as greatly increasing our knowledge of other archosaurs, squamates, mammals, and many others (Zhou et al., 2003; Benton et al., 2008). The Messel shales of Germany are another locality famous for its exceptional preservation (Franzen, 1985; Davis and

Briggs, 1995; Franzen et al. 2009). External tissue preservation is relatively easy due to the more durable nature of the tissues and their close contact with the external environment and preservational mechanisms, as discussed earlier. Even so, many aspects are not well understood and have caused interpretations of even the best preserved fossils to be debated. The discussion here will highlight a few illustrative examples and discuss taphonomic aspects of some much-publicized and, in some cases, highly controversial fossils, as well as point out areas that need more study.

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Monjurosuchus and Integument Slippage

A specimen of the Cretaceous choristodere Monjurosuchus splendens exquisitely illustrated by the IVPP in Liu and Wang (2008) shows preserved integument over much of the caudal portion of the specimen (Fig. 41). In particular, the right femur is framed on both sides by well-preserved skin. The cranial portion shows relatively large scales, whereas the caudal section shows a proximal section of comparatively small scales and a distal portion of very small scales.

The skin preservation here appears to show the full extent of the integument around the leg, showing both dorsal and ventral integument. How much is actually present, however, is unclear. Integument sometimes slides off of the carcass during decomposition as intact pieces preserving what appears to be complete skin (Chapter 2).

Without careful examination, this sloughed-off skin could appear to be either a frill around the body or could make the outline of the carcass appear much bigger than it actually is. In specimens that are exceptionally well preserved, determining whether the skin is one layer or multiple layers, indicating a folded and compressed section, is possible. Most times though, this sort of determination cannot be done.

Extant comparisons may be of value here. A preliminary study of extant lizards has indicated a relationship between the length of the femur and the cross-sectional area of the thigh (Appendix D). Extant lizards are not closely related to choristoderes such as

Monjurosuchus, but they are the closest living relative having the same overall morphology. Using this relationship, the Monjurosuchus fossil is shown to have approximately half the amount of integument what one would expect for an animal of this 179 size, indicating that the internal soft tissue decayed and the surrounding integument was simply flattened, albeit rolled to one side exposing portions of the dorsal and ventral sides.

Another specimen of Monjurasuchus illustrated by Gao et al. (2000) is reported to have preserved integument showing a webbed pes. However, while the specimen as illustrated does indeed show clearly preserved integument as well as sedimentary traces of claws beyond the tips of the exposed phalanges, the integument is not clearly indicative of webbing. It may well be, but it is also consistent with slippage of unwebbed phalangeal integument. Reconstructions of a semi-aquatic lifestyle for this animal based on this feature are therefore equivocal. This specimen, if not already prepared more fully, seems to be a good candidate for CT scanning, which may potentially allow better three- dimensional relationships of the bones to the preserved integument.

Feather Controversies

Few things have stirred up more controversy and acrimonious debate within paleontology than has the origin of birds. A key character in the debate has been the presence of feathers or featherlike structures found in association with several dinosaur skeletons. Among these, probably no specimens have been more debated than those of

Sinornithosaurus and Sinosauropteryx. Both have been described as having filamentous integumentary appendages described as either ‘protofeathers’ or degraded collagen fibers.

Taphonomy has played an important role in the debate, although the subjectivity of interpretations has made it difficult to come to firm conclusions. The specifics of feather 180 degradation and its effects on preservation are also not well understood (Benton et al.

2008; Foth, 2012), which adds another complication to taphonomic interpretation.

Sinornithosaurus—Sinornithosaurus was first described by Xu et al. (1999) as having filaments widely covering the body, but not in their original position “owing to posthumous displacement” (p. 263). They go on to say that their “broad distribution” made them unlikely to be dermal collagenous fibers, as had been suggested by skeptics

(see below). Xu et al. (2001, p. 201) argued that the fibers “are unlikely to be collagenous dermal fibers or musculoskeletal structures because they are clearly preserved as external integumentary structures.” They concluded that the tufts on the fossil are feathers because they remained as clumps of strands despite being dissociated from the body and in varying orientation.

Sinosauropteryx—Due to its basal position within Coelurosauria, the debate around

Sinosauropteryx is even more highly charged. Chen et al. (1998) described the dorsal fibers as integumentary structures similar to plumulaceous feathers. They also described them as probably being all over the body as there are patches of fibers surrounding the skull, not just dorsally, as well as a small patch over the left ribs and patches along the left (lateral) side of the tail. Chen et al. (1998) and Currie and Chen (2001) claimed that the clumped nature of the feathers is an artifact due to the uneven separation plane between the part and counterpart (NIGP 127587 and NGMC 2123). They also stated that the structures were “soft and pliable,” noting the variable orientation and sinuosity of the strands. 181

Dissenting opinion— Dissenting analyses of the structures have been presented by

Lingham-Soliar (2003a, b), Feduccia et al. (2005), and Lingham-Soliar et al. (2007). The authors asserted that the structures were dermal collagen fibers, with the neck and tail fibers forming a stiff frill along the spine, rather than a whole body covering. After comparing fossil evidence with decaying skin from a dolphin, a white shark, a chicken, and a small variety of reptiles, they concluded that the strands could be explained as decomposing collagen fibers, forming the purported structures through a variety of possible taphonomic pathways.

The dispute involves three lines of evidence, fiber appearance, randomness of fiber orientation, and location of the fibers in terms of external versus internal position, distribution over the body, and distance from the skeleton. Of the three, fiber appearance is perhaps the most straightforward. While the collagenous decay products shown by

Lingham-Soliar (2003a, b), Feduccia et al. (2005), and Lingham-Soliar et al. (2007) do look superficially similar to the strands on Sinornithosaurus, they are also an order of magnitude smaller, arguing against the collagen hypothesis. It should be noted that the size of the similarly-appearing strands given by Currie and Chen (2001) for

Sinosauropteryx (>0.3 mm to <0.1 mm) are within the range for collagen fiber bundles

(<0.05 mm to >1 mm, according to Feduccia et al., 2005). The strands are far larger than collagen fibers, which are between 0.001 and 0.012 mm (Burns and Cave, 2007). This range is slightly smaller than the 0.004–0.02 mm given by Feduccia et al. (2005), but the latter source supplied no reference for this figure. 182

Interpreting the randomness of fiber orientation has been more difficult. Feduccia et al. (2005) argued that the strands on Sinosauropteryx and Sinornithosaurus are more random than attributed by Currie and Chen (2001) and, combined with erosional issues, makes identifying them as feathers unjustified, whereas Lingham-Soliar et al. (2007) argued that they are unlikely to be protofeathers because they are not random enough.

Notably, the patterning of the strands is not completely random. The tufts that are detached from the carcass show patterning expected for keratinous feathery structures.

Currie and Chen (2001) stated that the fibers in the integumentary corona overwhelmingly have the orientation of originating near the body and flowing out and caudally, which is consistent with a feather explanation but not consistent with either a collagenous-dermal-fiber- or frill explanation or a sloughed-skin explanation.

The argument over fiber orientation is of limited value without detailed tapho- environment information, because without such information, taphonomic considerations preclude a clear expectation of orientation. Fiber orientation regardless of origin can be either parallel or random. A parallel orientation would likely be natural. Whereas a very low current may realign the fibers into a parallel orientation, current sufficient to move them would much more likely simply float them away. Random orientation, on the other hand, could occur from either an initial unstructured pattern or as random placement due to chance during decomposition. The argument by Lingham-Soliar et al. (2007) for a random position for protofeathers would be valid for a carcass decaying on a dry substrate. However, on a wet substrate, one would expect an incompletely ordered arrangement, because the feathers by the body would be stuck in place, fixed in almost a 183 natural position, whereas feathers not affixed to the substrate would either drift away as decomposition released them or fall exterior to the immediate corona (Bickart, 1984).

This pattern is indeed consistent with the fossils.

There is no reason to expect dermal fibers to have any particular pattern close to the carcass. Dermal fibers may or may not be parallel and ordered as Lingham-Soliar has shown (Lingham-Soliar, 2003a, b; Feduccia et al., 2005; Lingham-Soliar et al., 2007).

Parallel strands would not be expected to have any particular orientation relative to the carcass. One would also expect, as they have illustrated, clear crosscutting relationships.

This would make them difficult to distinguish from very small feathers or hair-like structures on sloughed off skin, which would have a similarly random orientation with respect to the carcass, but does provide a method for distinguishing them from feathers preserved in place. Feduccia et al. (2005) illustrated this sort of random relationship for

Sinosauropteryx, but the preservation in those areas illustrated is unclear and the interpretations are debatable.

Location of the fibers has perhaps been argued the most. Xu et al. (2001) and

Norell and Xu (2005) claimed that the fibers were clearly external integumentary structures based on their location on the fossil. It is fair to ask how these fibers, which are not in their original position, are clearly external structures. Norell and Xu (2005) ruled out fibers specifically around the head as being collagenous because, in their view, collagenous fibers would have been tightly bound to the head. However, Feduccia et al.

(2005) disputed this view, correctly stating that it should not be unusual for rotting tissue to accumulate around the periphery. The suppositions by Feduccia et al. (2005) about 184 flayed skin or a frill being possible explanations are plausible in this context. Skin that has been sloughed off from the surface of a carcass and deposited around the periphery can contain both external (i.e., epidermal) integumentary and dermal collagenous structures. Whereas it is most common for the epidermis to slough off leaving the dermis behind, whole sections of skin can also easily come off, particularly if decay has begun before burial (Chapter 2). Xu et al. (2001, p. 201) made this point themselves:

“independent hair-like filaments from independent integumental structures might remain associated if they were attached to a single piece of skin that sloughed off the body during decomposition, but the varying orientation of the dissociated overlying

Sinornithosaurus appendages argues directly against this interpretation.“

Lingham-Soliar et al. (2007) reported similar structures as being located deeper within the body. Unfortunately, while examination of the actual specimen was not possible, the images and locations described in their paper are unconvincing that they are indeed within the body. The locations that they note are all above the vertebrae. Whereas it is possible they were indeed deeper within the body, they could also be external structures compressed onto the skeleton during decay. There is no way to know for sure and are just as likely to be epidermal in origin because fibers from both the dermis and epidermis would be located in the same place on the skeleton after degradation of remaining tissue, provided no significant transport occurred.

The distribution of the fibers around the body has been a major point of contention. Geist et al. (1997) claimed that the fibers are strictly along the midline. This claim is repeated by Feduccia et al. (2005), who interpreted the structures as a sagittal 185 crest or frill. However, both Xu et al. (1997) and Currie and Chen (2001) reported preservation of fibers laterally, not just along the midline, supporting the interpretation of a covering over the entire body. Additionally, Currie and Chen (2001) described the strands as being piled thickly, indicating there were multiple strands overlying each other, which is consistent with the strands being an entire body covering, although it does not rule out a frill or collagenous fibers.

Finally, the distance of the fibers from the skeleton has been debated. Currie and

Chen (2001) stated that in regions with large amounts of skin and muscle, the integument would have degraded along with the underlying soft tissue, presumably leaving the feathers and accounting for the distance from the bone. Feduccia et al. (2005) stated that

Currie and Chen neglected the toughness of skin. Taphonomic work (Chapters 2, 3; Gill-

King, 1997) has demonstrated the ability of skin to last longer than the underlying tissue, due to the more durable nature of collagen. Nevertheless, collagen is far more susceptible to decay than keratinous tissue, so feathers remain long after collagen (at least that not locked away inside bone) is completely degraded (Chapter 3). In fact, sagittal preservation of feathers at approximately the correct distance from the bones with no other soft tissue preserved is precisely what one would expect to find of an animal covered in feathers whose carcass decomposed within wet sediment (Bickart, 1984;

Chapter 2). Avian carcasses tend to become “glued” to wet sediment, such that they do not move from the position in which they came to rest, even when exposed to significant wind or wave action. Decomposition can remove all trace of everything but the bones and 186 feathers within a few days, but the feathers may remain in position for extended periods allowing substantial time for them to fossilize.

This pattern is much more consistent with what we observe in the fossils than is preservation of a collagenous frill or dermal collagen fibers. In the latter case, no gap should be present. A frill would be located along the midline and thus next to the spine along the entire section, preserving clear, crosscutting relationships of the fibers. Dermal collagen fibers would not be separated by any distance from the skeleton because they would simply collapse onto the skeleton and would not pile up around the edges with the fibers flowing out and caudally while leaving a gap between fibers and skeleton, as seen in the fossils.

These statements do not conclusively support either feathers or dermal fibers, but they do state the limitations of interpretations based on position. In general, determination of integumentary type based on location is fraught with uncertainty at best and is untenable when it is apparent that displacement has occurred. However, considering the preservational advantage of keratinous structures over collagen, the more likely but certainly not conclusive interpretation is that the fossils indeed preserve feather-like structures. The partial disarticulation of the bones of the skull (Chen et al., 1998), as opposed to simple breakage, argues for the degradation of any exposed collagen (Chapter

3) and further supports the keratinous feather hypothesis.

Padian et al. (2001, p. 121) argued that the integumentary structures of NGMC

2123 are feathers and stated, “The view that these structures are collagenous fibers from the body midline is simply indefensible.” While they were correct that the midline 187 statement is indefensible, they ruled out collagenous fibers apparently solely based on placement (stating that preservation does not allow for anatomical detail), which is itself an indefensible statement. In this, I am in agreement with Feduccia et al. (2005) that “due regard is required with respect to changes, …that occur in soft tissue with the onset of death and degradation.” Although the data may be set in stone, the interpretations are not, so we are forced to weigh the evidence in total. Since none of the evidence is conclusive, we have to gauge the whole picture in context and cannot afford to be too reductionist in our thinking. In this case, it appears that the debate has polarized interpretations beyond the data, with the pro-feathers camp being too accepting of the structures as feathers and not sufficiently examining the taphonomic possibilities, whereas the anti-feather camp has attempted to introduced taphonomic rigor into the discussion, but have set the bar for acceptance of feathers so high that it is impossible to achieve.

The hadrosaur “mummies” “Leonardo” and “Dakota”

Two hadrosaur mummies that exhibit excellent skin preservation have drawn a great deal of attention, and the taphonomic processes leading to their preservation and interpretation have been much discussed in the popular press, if not in academic journals.

In Montana, a specimen of Brachylophosaurus canadensis (JRF 115H), colloquially named “Leonardo,” was found in 2000 in the Judith River Formation. The other specimen, MRF-03, an Edmontosaurus nicknamed “Dakota,” was found in the Hell

Creek Formation of North Dakota. Both are from the Upper Cretaceous and are noted for unusual skin preservation that is at an apparently lifelike distance from the bone. Neither 188 one has been fully examined yet, as work is in progress, which provides an unparalleled opportunity to examine preservation in detail throughout the preparation process.

MRF-03 was preserved through what appears to be rapid burial in a sandy river channel with rapid carbonate precipitation due to bacterial methanogenesis. The soft tissue of MRF-03 wa s preserved in sediment that is much finer grained than the surrounding matrix, with the skin replaced with clay minerals and siderite (Manning et al., 2009). This is in agreement with taphonomic work that has indicated the conjunction of microbial mineral precipitation and clay adherence (e.g., Martin et al., 2004) as well as broader-scale sediment interactions (Chapters 2, 3). This fossil is therefore a very likely candidate for finding traces of soft tissue near body orifices. The head of this animal is a prime candidate for detailed CT analysis before preparation of the bones. Attempts have been made to scan the whole body block, but this proved unsuccessful (Manning, 2008).

However, if the head is separated from the body, CT scanning might be more successful, and this find could be potentially of great importance.

One caveat that should be kept in mind with this fossil, as with all fossils with presumed soft tissue preservation, is that even in the best of circumstances, the soft tissue will not be perfectly in place. Manning (2008) stated that the successful CT scan of the tail indicated that the tail was much broader than previously thought. This may be true, but it is also very possibly a preservational artifact. The tail would almost certainly have been laterally compressed and dorsoventrally broadened during decomposition and this possibility must be accounted for before proper reconstruction can be made. 189

Less taphonomic work has been published on JRF-115, but some aspects may be tentatively discussed. The fossil was found with skin from the right shoulder in a

“natural” position, although the remaining skin was compressed to the bone. There was also a “vertical comb or frill” (Manning, 2008) preserved along the spine. Initially, in addition to skin, Murphy had reported (Halls, 2003) that he had found muscle, internal organs, stomach contents, a neck pouch, and a tongue. However, in the published description of the fossil, he only discussed the skin preservation (Murphy et al., 2007).

He interpreted the taphonomy as the animal having been mummified during a drought and then buried in a river after the drought broke (Halls, 2003). However, if this were true, it is extremely unlikely that any internal tissue would have been left behind in any recognizable form as the endogenous bacteria would have decomposed the internal tissue.

The skin may have been dried in the sun and served as protection for the bacteria to continue decomposition of the internal tissue (Galloway, 1997; Chapters1–3).

A question not answered by Murphy et al. (2007) is how the sand managed to fill the interior while at the same time preserving the skin in place. One would presume that the skin would collapse after the soft tissue degraded, and the skeleton should have disarticulated to some degree. The key here is likely the burial of the right shoulder underneath the carcass. As Murphy et al. (2007) noted, the remainder of the carcass was more exposed to decay processes and not as well preserved. Thus, the internal tissues would have decayed faster than those in the lower section. With a carcass this large, sediment would have collapsed into the carcass before decomposition was complete, eventually burying the underlying skin from the inside while partially stabilizing the 190 skeleton in place. The overlying skin would collapse onto the bones as the soft tissue decayed in more typical dinosaur “mummy” fashion. Meanwhile, the sediment surrounding the skin underneath would be stabilized by mineral precipitation caused by bacterial decay forming the skin impressions that were found. If this hypothesis were true, internal soft tissue will not be found other than perhaps plant material from gut contents, which themselves would be somewhat displaced and would be located on the lower (relative to the position at burial) side of the abdomen.

Questions

There are, of course, those finds that create more questions because of what they do not have than what they do. One case in point is the collection of psittacosaurs reported by Qi et al. (2007). This bonebed contains exquisitely preserved, well-articulated complete skeletons preserved in what they refer to as a lahar deposit, a volcanic mud/debris flow (Fig. 42). The sediment is very poorly sorted and shows no bioturbation and should be a prime candidate for soft-tissue preservation, yet no trace of soft tissue has been found. External tissue preservation could be explained by the flow being simply too thick and quickly deposited to be sorted, much like some turbidite flows (Lowe, 1997;

Major, 1997) and like that seen in previously supported taphonomic studies (Chapter 2).

However, one would expect some heterogeneity within the skeletons as the soft tissue decayed and was replaced by collapsing sediment. The lack of sorting suggests the burial depth was considerable, causing increased pressure which may have squeezed out any gaseous traces before any significant lithification occurred, although the depth could not 191 have been too great as the skeletons were in pristine condition. The three-dimensionality of the skeletons suggests a rapid decay of soft tissue and displacement of gas by still-fluid sediment. Nevertheless, the sediment within the skulls has not been examined for internal soft-tissue traces and would seem to be potentially fruitful. The heterogeneity of the sediment and its fluid nature during burial are optimal for grain sorting due to soft-tissue interactions to occur within the head. It is possible that the sediment had too high of a density to enter the carcass through small openings, but this seems contraindicated by the other taphonomic indicators.

Wang and Zhou (2008) described the preservation of the Jehol Biota as being similar to Pompeii. However, in the case of the Jehol Biota, the animals died in a lake and floated briefly before sinking to bottom and being covered in ash. All of these fossils are preserved as flattened fossils, unlike the three-dimensional corpses of people found at

Pompeii. However, many of the Jehol specimens preserve at least integumentary soft tissue. Yet the psittacosaurs described by Qi et al. (2007) are three-dimensional and preserve no obvious soft tissue. The psittacosaurs all died quickly in a nonbioturbated muddy debris flow, so the preservation of some soft tissue might be expected. They were from the Yixian Formation as were the fossils described in the Jehol Biota. However, the psittacosaurs were from the Lujiatun Beds, known for fully articulated three-dimensional specimens (e.g., Mei long reported by Xu and Norell, 2004), but not soft tissue, unlike the flattened fossils of the Jianshangou and Dawangzhangzi beds which did preserve soft tissues. Why the skeletons were preserved in excellent shape without soft tissue in one 192 place, but in another were less well preserved but retained soft-tissue traces warrants explanation.

More detailed studies of Pompeii and Herculaneum have found important differences in preservation, which explain the preservation in Pompeii and Herculaneum as well as the psittacosaurs. It has been widely reported that Pompeii and Herculaneum were buried under lahar deposits with massive pumice and ash falls. However, these deposits were not formed from a single event (Luongo et al. 2003). Instead of one event, there were multiple events with two distinct types. The first type involved a massive ash and pumice fall, which blanketed the cities. This event caused the exquisite casts seen in

Pompeii (Fig. 43). The second type consisted of pyroclastic density flows, which caused articulated skeletons in life poses, but no soft tissue whatsoever (Fig. 44), just like that seen in the psittacosaurs reported by Qi et al. (2007).

The pyroclastic density flows were formed from ash, pumice, mud, and superheated gasses reaching a temperature of approximately 500 °C. This extreme heat vaporized the soft tissue instantaneously. However, that vaporization also cooled the flow at the same time. This cooling prevented serious damage to the skeleton as well as causing the ash cloud to deflate and solidify around the skeleton within seconds. Further temperature drops and ash fall over the next half hour or so solidified the layer and preserved the skeletons in place (Mastroloenzo et al. 2001).

It is apparent from these descriptions that the preservation of the Jehol Biota can be likened to the eruption of Vesuvius and the resultant destruction —and preservation— of Pompeii and Herculaneum. The layers preserving the exquisitely preserved fossils with 193 soft tissue were a result of ash falls, whereas the skeletons of the psittacosaurs and others like them were the victims of the pyroclastic density flows.

Conclusions

The identification of soft tissues in fossils without obvious signs of soft tissue preservation can greatly expand our knowledge of extinct animals and broaden our understanding of evolution within known clades. While it may seem trivial to say that advanced theropods had feathers, it is not trivial to identify when they developed, nor is it necessarily valid to interpolate traits within a clade. For instance, Göhlich and Chiappe

(2006) described Juravenator as a basal coelurosaur, noting its importance in understanding feather evolution due its phylogenetic position within feathered theropods despite preserving a scaly integument. Butler and Upchurch (2007) placed it as a basal maniraptoran, making Juravenator even more interesting. Pelecanimimus (LH 7777), as reported by Feduccia et al. (2005) is similarly interesting as a scaly ornithomimid within a feathered clade. These finds, as well as the heterodontosaurid Tianyulong with feather- like integumentary appendages (Zheng et al., 2009) further complicate our understanding of feather evolution even assuming the feather interpretations are correct. Body coverings, such as feathers, are evolutionarily labile structures in their type, form, and distribution over the body, to say nothing of more exclusive traits, such as the combs, wattles, and other integumentary appendages ornamenting a wide variety of birds and animals as a whole. Enhancing our understanding of the taphonomic processes enhances our understanding of the animals themselves and their evolutionary trajectories. 194

Figure 31. Hypothetical spectrum of preservability for different tissues. Scale is relative and nonlinear. 195

Figure 32. Void space in Tyrannosaurus rex (FMNH PR 2081). A: Volume rendering of CT scan of skull. B: Position of CT slices shown in C and D. C: Axial CT slice through skull. D: Horizontal CT slice. White arrows indicate primary void space.

196

Figure 33. A: Braincase of Allosaurus (UMNH VP 18050). Red rectangle indicates area shown in B. White arrow indicates mineral infilling pneumatic sinus. Scale of rectangle ~1 cm in length. 197

Figure 34. Metallic infills within the premaxilla of Allosaurus (UMNH VP 18046). A, B, E: Amira reconstructions from CT data, infill in green, bone translucent grey, views 198 lateral, rostral, ventral. C, D: CT slices through premaxilla, sagittal and axial, respectively.

Figure 35. CT scan of Nanotyrannus (CMNH 7541). orange lines on center image demonstrate location of slice images in skull. Arrows indicate less dense matrix within pneumatic sinuses.

199

Figure 36. CT of Hypacrosaurus premaxilla (ROM 702). A: Sagittal slice. Orange line indicates position of axial slice shown in B. Sediment filling sinus shows pulsed layering indicating that the premaxilla was buried lateral side down.

Figure 37. Tyrannosaurus rex (AMNH 5027) as figured by Osborn 1912. Arrow indicates right ectopterygoid in left antorbital fenestra. 200

Figure 38. A: CT reconstruction of a juvenile Tarbosaurus bataar MPC-D 107/7. B: Ikechosaurus (adapted from Liu and Wang 2008).

201

Figure 39. Adipocere formed on buried, rotting ostrich head. A: axial CT slice. B: sliced ostrich head within sediment block. C: ostrich basicranium with adipocere removed from sediment. 202

Figure 40. Possible conchal sediment traces in CT reconstructions of Panoplosaurus (ROM 1215). Yellow nasal passage (adapted from Witmer and Ridgely 2008) overlies ghosted skull. Segmented sediment traces overlie nasal passage. Tubular sediment traces lying within wide final chamber in front of choanae aligned with air flow as shown in Witmer and Ridgely (2008). A: Right lateral view. B: Oblique left view. C: Dorsal view. D. Ventral view. 203

Figure 41. Monjurosuchus. Photo: IVPP modified from Liu and Wang (2008). 204

Figure 42. Psittacosaurus bed buried in lahar deposit (modified from Qi, et al. 2007).

205

Figure 43. People preserved as casts by ash fall at Pompeii (adapted from Luongo et al. 2003).

206

Figure 44. A child’s skeleton preserved by pyroclastic density flows at Herculaneum (from Mastrolorenzo et al., 2001).

207

Table 6. Endpoints for original void space within carcasses

Type Interpretation Example

1 Primary voids Original space Tyrannosaurus FMNH PR 2081 2 Secondarily Original space mineral infill Allosaurus mineralized UMNH 18046 3 Sediment infilled Cleaned before burial, fractured prior or during burial Fractured prior to or during burial Deinonychus OMNH 50268 3a

Bioturbation, limited access Thuliadonta (see text) 3b

Infiltration via sediment flow forming Nanotyrannus homogeneous fill CMNH 7541 3c

Slow pulses of sediment forming layered Hypacrosaurus patterns ROM 702 3d

4 Secondary voids Type 2 showing dissolution

208

Table 7. Institution abbreviations

Abbreviation Institution Location AMNH American Museum of Natural History New York, New York BMNH Natural History Museum London, United Kingdom CMNH Cleveland Museum of Natural History Cleveland, Ohio CNRST-SUNY Centre Nationale de la Recherche Paris, France; Stony Scientifique et Technologique-Stony Brook Brook, New York University FMNH Field Museum of Natural History Chicago, Illinois IVPP Institute of Vertebrate Paleontology and Beijing, China Paleoanthropology JRF Judith River Foundation, Great Plains Malta, Montana Dinosaur Museum LH Las Hoyas Collection, Museo de Cuenca Cuenca, Spain MRF Marmarth Research Foundation Marmarth, North Dakota NGMC National Geological Museum of China Beijing, China NIGP Nanjing Institute of Geology and Nanjing, China Paleontology OMNH Oklahoma Museum of Natural History, Norman, Oklahoma Oklahoma University ROM Royal Ontario Museum Toronto, Ontario, Canada UMNH University of Utah Museum of Natural Salt Lake City, Utah History 209

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APPENDIX A: GRAIN-SIZE ESTIMATION TECHNIQUES

There are currently four generally accepted methods for estimating sediment grain size. The most common method is by filtering the sediment through a series of sieves, the standard being those certified by the American Society for Testing and Materials

(ASTM). This method however, only records the smallest diameter of the grain. A spindle-shaped grain will be measured as a much smaller grain than a spherical grain with the same surface area. Another problem with sieving is that clay and silt particles cannot be effectively measured, and a different technique using settling velocities is utilized.

This method, referred to as the pipette method (there is also a variant called the hydrometer method, but it uses the same principles and assumptions), uses Stokes’ Law, which states that the larger the particle, the more friction must be overcome for the particle to move through the liquid, with the effect that larger particles sink slower than smaller particles in a liquid of a given viscosity. This statement assumes that all the particles are of equal shape (spherical) and density. A small, dense particle will settle faster than a large, light particle, but will be recorded as a larger particle. Both of these methods report results in terms of mass, assuming that density is not a significant variable

(Folk, 1980; Lewis, 1984).

The third method uses laser diffraction systems that measure light scatter off particles. This method is more direct in that it removes the problem of density. Results can also be reported in terms of both volume and the number of counts. The diameter reported with this method is not the minimum, but rather a random diameter, depending on the orientation of the particle as it passes through the light beam. It assumes a 231 spherical shape, and so a particle with a length several times longer than its width will be reported as a range of sizes. The method also depends on the particles passing through the light beam individually. If the stream is too fast, multiple particles will be measured together as one particle, greatly increasing the size of the reported particle. Clay particles are particularly susceptible to adhering to each other. Several methods are utilized to limit this behavior, such as increased air pressure or a liquid suspension with the addition of anti-flocculants, but the effectiveness of these methods varies greatly by sediment type and can run the risk of breaking particles, thereby counting them as smaller than they actually were. An additional problem with this method is that it assumes equal reflectance for all particles. Particles of different colors or crystal patterns will be measured as different sizes. In effect, this method assumes that all particles are simply size variations of the same material. Finally, it is very difficult to compare numerical reports with volumetric reports as the volume of particles increases greatly as size increases. For example, one spherical grain that is 1 mm in diameter will count the same as 1,000,000

1µm particles. Thus, a single large grain will overwhelm any smaller particles in a volumetric measurement, but will be lost in a numerical measurement (Konert and

Vandenberghe, 1997; Kippax, 2005).

The third method is a photographic measurement technique in which the sediment is spread out and each particle is physically measured (for detailed review and methods, see Francus, 2005). This method has traditionally been done by hand, thereby greatly limiting the utility of this technique, but recently some machines have entered the market to automate this procedure, making it much more viable, although it is still much more 232 time-consuming than the other techniques. This technique combines the benefits and precision of the laser diffraction techniques with the ability to measure several other grain shape parameters at the same time. But like the other methods, this technique is not without its flaws. Particles are measured while at rest on a surface, so the diameter is measured in the most stable position. A plate will therefore be measured as having the same volume as a sphere, despite its having a small fraction of the mass or volume of the sphere. Various methods have been proposed to deal with this problem, but all assume the grains are of a common type and are thus inappropriate for heterogeneous mixtures, which defines virtually all known natural sediments. The biggest problem with this method, however, is that a minimum of 200–300 grains are required to be measured for every sample to be meaningful. This means that the number of samples that one can measure is extremely limited, making any statistical comparison requiring large sample numbers impossible.

All of these methods assume the grains can be separated. If the sediment is indurated, none of these methods will work. However, the photographic method can still provide a rough estimate of grain-size distributions, but will perforce not have the accuracy and precision of the methods utilizing separated grains. So, in cases of indurated sediment, it is the only available method that will give results, however limited they may be. Each method has its advantages and disadvantages. Unfortunately, none of the methods are comparable with each other as they will all give different answers to some extent. Thus, the relative shape of the distributions maybe be roughly compared, but one should not measure some sediment with one technique and use it in an analysis with 233 sediment measured with a different technique, nor even really directly compare analyses.

These techniques all have their place, but should only be used as a way of comparing sediments within the same study and should not be used to make precise comparisons with other studies.

234

APPENDIX B: GRAIN-SIZE ESTIMATION IN THIS STUDY

The initial plan was to analyze all the sediment samples with a laser diffraction system. Unfortunately, after processing the samples, it was discovered that a malfunction in the machine caused all the data to be unreliable. However, high-resolution photographs were taken of all slices of all sediment blocks. It was decided to estimate grain-size distributions of several regions on each slice for ten of the blocks. Estimates were taken for one block containing a fresh head, desiccated head, and a clean skull for both flumes.

Both blocks from each flume containing rotten heads were examined, because the rotten heads had demonstrated more variability in sediment patterns than had the other conditions. Only the short-term burial blocks were used for sediment parameter estimation. Long-term burial blocks, those that had been reburied for seven months after being initially CT-scanned, were not used for sediment parameter estimation because the overall exterior patterns were not changed considerably except in those areas that had collapsed into the void spaces that formed within the decomposing heads. Additionally, the sediment was riddled with gas pockets, which would have made density measures of the sediment without gas impractical.

Due to the time constraints imposed by this method, we elected to categorize the distributions rather than attempt to obtain closely accurate measurements for a few hundred grains for each sample. Although limiting the resolution of the data, this approach allowed a far greater number of samples to be measured as well as allowing measurement of samples too small to be counted by the higher-resolution method.

Moreover, the quality of the photographs did not allow higher quality estimations in 235 many cases, making higher-resolution studies moot in any case. Grain-size categories followed the commonly accepted classification in the Udden-Wentworth scale in Table 1.

Table 1. Grain-size categories for sediment distribution estimates.

Sediment Estimates

Category Grain Size (mm) Phi Size Class

1 <0.062 4+ silt/clay

2 0.062–0.125 3–4 very fine sand

3 0.125–0.25 2–3 fine sand

4 0.5–0.25 1–2 medium sand

5 0.5–1.0 0–1 coarse sand

6 1.0–2.0 -1–0 very coarse sand

Grain-size distributions were measured according to three parameters: dominant grain size, range, and relative fines (silt/clay) contribution. For each sampled region, a representative sample of grains was categorized by comparing a one-millimeter length as determined by the in-photo scale bar to the individual grains. Grains were measured along their longest axis for approximately spherical grains and an average between the longest and shortest axes for grains that were substantially nonspherical. The number of grains varied according to the size and heterogeneity of the sampled region. The smaller 236 and more homogeneous the sampled region, the fewer grains that were required for determination.

Determination of dominant grain size involves two parameters: absolute

(volumetric) and relative (numeric) abundance. Because of the immense volumetric differences between grain categories (one 2 mm grain = 8x109 μm grains), using either parameter separately provides a skewed interpretation of the actual distribution. Sieving techniques utilize solely mass measurements, which are assumed to be essentially identical to volumetric measurements, although this is not the case in many circumstances. Characterization of fines utilizing settling velocities also uses mass measurements, with the same inherent problems. Laser diffraction techniques utilize the numeric distributions, which can then be converted into volumetric distributions without regard for mass differences, making this a preferred method, but has the limitations discussed above. For this study, dominance was given to the category with the highest volumetric percentage, except in cases in which that volumetric dominance was achieved by a small number of very large particles in a region otherwise dominated by a much smaller category. In contrast, range was much more straightforward, involving determination of the largest and smallest grains and including all categories in between.

Thus, a region that contained both categories 1 and 6 was given a range of 6, a region with the largest grain size in category 5 and the smallest in category 2 was given a range of 4. Relative fines contribution was determined on a relative three-point scale: none observable, noticeable (“some” or “moderate”), and substantial (“lots”) contribution. This was initially counted on a scale of 0–2, with 0 being none observable. However, for 237 statistical analyses, this was converted to a 1–3 scale to avoid zeroes in the data set, which could potentially pose problems with some analyses.

This method proved to have several limitations regarding the precision and reproducibility of the estimates, although these limitations were alleviated somewhat by substantial sample size and did not unduly overwhelm the needed level of accuracy. The most obvious limitation was that of photographic resolution. In most cases, it was possible to distinguish grain sizes down to coarse silt, leaving finer silt and clay as a pasty texture. Larger particles could be difficult to measure accurately many times due to a lack of clarity in the boundaries between grains. This proved to be an issue when the grain was close to the border between size categories. Grains close to the borders of categories added an additional complication. A sample of grains that were on average 0.9 mm in diameter was counted as category 5. They were comparatively closer to category 6 than they were category 4, yet data analyses would count them as equidistant. This problem, however, pertains whenever one attempts to separate continuous data into arbitrary bins and is not unique to this study.

Another limitation was clarity of the image due to frost. The sediment slices were frozen when photographed and in some instances, frost appeared on the slices during photography. The ice contained within and on the slices also began to melt during the process. The addition of ice and water limited clarity in some photographs and may have altered the apparent grain-size distribution, particularly in the fines category.

Finally, there was the standard subjectivity of the person making the determinations. One place this subjectivity played a role was in determinations of the 238 relative fines contributions, where it proved difficult to reliably make determinations between categories 2 and 3. If the fines were the dominant fraction, it was of course simple. Where it proved difficult was when there were regions within the same slice that had multiple clearly different levels of fines contribution. For instance, in some cases, there were clear areas in which the fines were present in low amounts adjacent to another region which had a much larger fines contribution, but contained within that region were small layers of sediment in which the fines were the dominant size class. Another place of acute subjectivity was when a region with a small dominant size fraction, but with a few grains of a much larger size fraction graded into a region dominated by that much larger size fraction. For example, a region that would be categorized as having a dominant size category of 2, but with a range of 6 due to a small amount of fines and a few grains of category 6, in some instances graded into a region dominated by the highest size fraction. The determination of when it switched from 2, 6, 1 to 6, 6, 1 could depend on subtle differences that nevertheless radically altered the characterization of the sediment without necessarily requiring any transitional sediments with intervening size categories.

Multiple checks were put in place to address these difficulties. Human subjectivity was partially controlled by only one person making all the determinations.

Additionally, approximately 10% of the determinations were redone on different days for comparison and this resulted in over a 90% success rate. Those that were found different were repeated and revised if needed. Samples chosen for density comparisons were those with a suitably large area and away from edges if possible to avoid obvious CT artifacts, 239 such as beam hardening and edge effects caused by a sharp delineation between two regions of highly different densities. Situations with the fines as mentioned above with multiple levels of fines contribution were able to be delineated when combined with the dominant size fraction. For instance, the region with high fines contribution might be described as 4, 6, 2, whereas the thin layer in which the fines dominated might be 1, 3, 2.

This illustrates the fact that all three parameters are needed to adequately describe the sediment. Any one parameter misses vital information about the nature of the sediment.

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APPENDIX C: GRAIN-SIZE DATA

Appendix C.1. Sediment sample characterizations

Head Dom. Flume Condition Block slice sample Size range fines Shallow Clean 10456 2 1 2 3 2 Shallow Clean 10456 2 2 3 4 2 Shallow Clean 10456 2 3 2 3 2 Shallow Clean 10456 2 4 6 6 2 Shallow Clean 10456 2 5 6 6 2 Shallow Clean 10456 2 6 6 6 1 Shallow Clean 10456 2 7 5 6 2 Shallow Clean 10456 2 8 5 6 2 Shallow Clean 10456 2 9 4 6 2 Shallow Clean 10456 2 10 3 6 1 Shallow Clean 10456 2 11 3 5 2 Shallow Clean 10456 2 12 3 5 1 Shallow Clean 10456 2 13 4 6 1 Shallow Clean 10456 2 14 5 6 1 Shallow Clean 10456 2 15 6 6 1 Shallow Clean 10456 3 1 2 3 2 Shallow Clean 10456 3 2 1 3 2 Shallow Clean 10456 3 3 2 3 2 Shallow Clean 10456 3 4 2 3 1 Shallow Clean 10456 3 5 1 3 2 Shallow Clean 10456 3 6 3 4 2 Shallow Clean 10456 3 7 2 3 2 Shallow Clean 10456 3 8 5 6 2 Shallow Clean 10456 3 9 6 6 2 Shallow Clean 10456 3 10 6 6 1 Shallow Clean 10456 3 11 3 5 1 Shallow Clean 10456 3 12 3 6 1 Shallow Clean 10456 3 13 6 6 1 Shallow Clean 10456 3 14 5 6 2 Shallow Clean 10456 3 15 2 3 2 Shallow Clean 10456 3 16 6 6 1 Shallow Clean 10456 3 17 6 6 2 Shallow Clean 10456 3 18 4 6 1 Shallow Clean 10456 3 19 3 5 1 Shallow Clean 10456 3 20 5 6 1 241

Shallow Clean 10456 3 21 6 6 2 Shallow Clean 10456 3 22 6 6 2 Shallow Clean 10456 3 23 2 3 2 Shallow Clean 10456 3 24 3 4 1 Shallow Clean 10456 3 25 2 3 2 Shallow Clean 10456 3 26 2 3 2 Shallow Clean 10456 3 27 5 6 2 Shallow Clean 10456 3 28 5 6 2 Shallow Clean 10456 4 1 2 3 2 Shallow Clean 10456 4 2 2 3 2 Shallow Clean 10456 4 3 2 3 2 Shallow Clean 10456 4 4 2 3 2 Shallow Clean 10456 4 5 2 3 2 Shallow Clean 10456 4 6 2 3 2 Shallow Clean 10456 4 7 5 6 2 Shallow Clean 10456 4 8 5 6 2 Shallow Clean 10456 4 9 5 6 1 Shallow Clean 10456 4 10 3 5 1 Shallow Clean 10456 4 11 3 5 2 Shallow Clean 10456 4 12 4 5 2 Shallow Clean 10456 4 13 5 6 2 Shallow Clean 10456 4 14 6 6 2 Shallow Clean 10456 4 15 5 6 2 Shallow Clean 10456 4 16 5 6 2 Shallow Clean 10456 4 17 3 5 2 Shallow Clean 10456 4 18 2 3 2 Shallow Clean 10456 4 19 5 6 2 Shallow Clean 10456 4 20 3 6 2 Shallow Clean 10456 4 21 2 4 2 Shallow Clean 10456 4 22 6 6 1 Shallow Clean 10456 4 23 4 5 1 Shallow Clean 10456 4 24 6 4 0 Shallow Clean 10456 4 25 6 6 1 Shallow Clean 10456 4 26 5 6 2 Shallow Clean 10456 4 27 5 6 1 Shallow Clean 10456 4 28 3 5 2 Shallow Clean 10456 4 29 4 5 2 Shallow Clean 10456 4 30 4 5 2 Shallow Clean 10456 4 31 5 6 2 Shallow Clean 10456 4 32 5 6 2 Shallow Clean 10456 4 33 5 6 2 242

Shallow Clean 10456 4 34 6 6 2 Shallow Clean 10456 4 35 4 5 2 Shallow Clean 10456 4 36 5 6 2 Shallow Clean 10456 4 37 5 6 1 Shallow Clean 10456 4 38 5 6 2 Shallow Clean 10456 4 39 6 6 1 Shallow Clean 10456 4 40 6 6 2 Shallow Clean 10456 4 41 6 6 1 Shallow Clean 10456 4 42 4 6 1 Shallow Clean 10456 5 1 6 6 2 Shallow Clean 10456 5 2 5 6 2 Shallow Clean 10456 5 3 5 6 2 Shallow Clean 10456 5 4 2 3 2 Shallow Clean 10456 5 5 2 3 2 Shallow Clean 10456 5 6 4 6 2 Shallow Clean 10456 5 7 4 5 2 Shallow Clean 10456 5 8 5 6 1 Shallow Clean 10456 5 9 5 6 1 Shallow Clean 10456 5 10 6 6 1 Shallow Clean 10456 5 11 5 6 1 Shallow Clean 10456 5 12 5 6 1 Shallow Clean 10456 5 13 6 6 1 Shallow Clean 10456 5 14 6 6 2 Shallow Clean 10456 5 15 5 6 1 Shallow Clean 10456 5 16 4 6 1 Shallow Clean 10456 5 17 4 6 2 Shallow Clean 10456 5 18 5 6 2 Shallow Clean 10456 5 19 4 6 2 Shallow Clean 10456 5 20 5 6 1 Shallow Clean 10456 5 21 5 6 1 Shallow Clean 10456 5 22 6 6 2 Shallow Clean 10456 5 23 6 6 2 Shallow Clean 10456 5 24 6 6 2 Shallow Clean 10456 5 25 1 2 2 Shallow Clean 10456 5 26 1 2 2 Shallow Clean 10456 5 27 1 2 2 Shallow Clean 10456 5 28 5 6 2 Shallow Clean 10456 5 29 2 3 2 Shallow Clean 10456 5 30 5 6 1 Shallow Clean 10456 5 31 6 6 1 Shallow Clean 10456 5 32 5 6 1 243

Shallow Clean 10456 5 33 6 6 1 Shallow Clean 10456 5 34 3 5 2 Shallow Clean 10456 5 35 2 3 2 Shallow Clean 10456 6 1 1 3 2 Shallow Clean 10456 6 2 1 2 2 Shallow Clean 10456 6 3 1 2 2 Shallow Clean 10456 6 4 1 2 2 Shallow Clean 10456 6 5 1 3 2 Shallow Clean 10456 6 6 5 6 1 Shallow Clean 10456 6 7 5 6 1 Shallow Clean 10456 6 8 6 6 2 Shallow Clean 10456 6 9 5 6 1 Shallow Clean 10456 6 10 4 5 2 Shallow Clean 10456 6 11 4 6 2 Shallow Clean 10456 6 12 6 6 1 Shallow Clean 10456 6 13 6 6 2 Shallow Clean 10456 6 14 2 4 2 Shallow Clean 10456 6 15 1 2 2 Shallow Clean 10456 6 16 4 6 2 Shallow Clean 10456 6 17 4 5 2 Shallow Clean 10456 6 18 4 5 1 Shallow Clean 10456 6 19 5 6 1 Shallow Clean 10456 6 20 4 5 1 Shallow Clean 10456 6 21 6 6 2 Shallow Clean 10456 6 22 5 6 1 Shallow Clean 10456 6 23 6 6 1 Shallow Clean 10456 6 24 5 6 1 Shallow Clean 10456 6 25 5 6 2 Shallow Clean 10456 6 26 1 3 2 Shallow Clean 10456 7 1 2 3 2 Shallow Clean 10456 7 2 1 2 2 Shallow Clean 10456 7 3 1 3 2 Shallow Clean 10456 7 4 4 6 2 Shallow Clean 10456 7 5 5 6 2 Shallow Clean 10456 7 6 2 3 2 Shallow Clean 10456 7 7 3 6 2 Shallow Clean 10456 7 8 4 6 2 Shallow Clean 10456 7 9 5 6 2 Shallow Clean 10456 7 10 3 5 2 Shallow Clean 10456 7 11 3 4 2 Shallow Clean 10456 7 12 4 5 2 244

Shallow Clean 10456 7 13 5 6 2 Shallow Clean 10456 7 14 5 6 2 Shallow Clean 10456 7 15 6 6 2 Shallow Clean 10456 7 16 5 6 2 Shallow Clean 10456 7 17 4 5 2 Shallow Clean 10456 7 18 6 6 2 Shallow Clean 10456 7 19 4 5 2 Shallow Clean 10456 7 20 5 6 2 Shallow Clean 10456 7 21 5 6 2 Shallow Clean 10456 7 22 5 6 2 Shallow Clean 10456 7 23 1 2 2 Shallow Clean 10456 7 24 5 6 2 Shallow Clean 10456 7 25 5 6 2 Shallow Clean 10456 7 26 5 6 2 Shallow Clean 10456 7 27 2 3 2 Shallow Clean 10456 7 28 2 3 2 Shallow Clean 10456 7 29 2 3 2 Shallow Clean 10456 7 30 1 2 2 Shallow Clean 10456 7 31 1 2 2 Shallow Clean 10456 7 32 2 3 2 Shallow Clean 10456 8 1 1 2 2 Shallow Clean 10456 8 2 6 6 2 Shallow Clean 10456 8 3 5 6 2 Shallow Clean 10456 8 4 6 6 2 Shallow Clean 10456 8 5 5 6 2 Shallow Clean 10456 8 6 6 6 1 Shallow Clean 10456 8 7 3 5 1 Shallow Clean 10456 8 8 3 5 2 Shallow Clean 10456 8 9 2 3 2 Shallow Clean 10456 8 10 3 5 2 Shallow Clean 10456 8 11 5 6 1 Shallow Clean 10456 8 12 5 6 2 Shallow Clean 10456 8 13 3 4 2 Shallow Clean 10456 8 14 3 4 2 Shallow Clean 10456 8 15 1 2 2 Shallow Clean 10456 8 16 5 6 2 Shallow Clean 10456 8 17 6 6 2 Shallow Clean 10456 8 18 1 2 2 Shallow Clean 10456 8 19 6 6 2 Shallow Clean 10456 8 20 2 5 2 Shallow Clean 10456 8 21 1 2 2 245

Shallow Clean 10456 8 22 5 6 2 Shallow Clean 10456 8 23 1 2 2 Shallow Clean 10456 8 24 1 2 2 Shallow Clean 10456 8 25 1 2 2 Shallow Clean 10456 8 26 1 2 2 Shallow Clean 10456 8 27 1 2 2 Shallow Clean 10456 8 28 2 3 2 Shallow Clean 10456 8 29 5 6 2 Shallow Clean 10456 8 30 3 5 2 Shallow Clean 10456 8 31 5 6 2 Shallow Clean 10456 9 1 2 3 2 Shallow Clean 10456 9 2 1 2 2 Shallow Clean 10456 9 3 2 3 2 Shallow Clean 10456 9 4 1 2 2 Shallow Clean 10456 9 5 2 3 2 Shallow Clean 10456 9 6 1 2 2 Shallow Clean 10456 9 7 2 3 2 Shallow Clean 10456 9 8 6 6 2 Shallow Clean 10456 9 9 4 6 2 Shallow Clean 10456 9 10 5 6 2 Shallow Clean 10456 9 11 5 6 2 Shallow Clean 10456 9 12 4 5 2 Shallow Clean 10456 9 13 3 4 2 Shallow Clean 10456 9 14 1 3 2 Shallow Clean 10456 9 15 6 6 2 Shallow Clean 10456 9 16 5 6 2 Shallow Clean 10456 9 17 5 6 2 Shallow Clean 10456 9 18 5 6 2 Shallow Clean 10456 9 19 3 5 2 Shallow Clean 10456 9 20 4 5 2 Shallow Clean 10456 9 21 3 5 2 Shallow Clean 10456 9 22 3 5 1 Shallow Clean 10456 9 23 3 5 2 Shallow Clean 10456 9 24 3 4 2 Shallow Clean 10456 9 25 4 5 1 Shallow Clean 10456 9 26 4 6 1 Shallow Clean 10456 9 27 5 6 2 Shallow Clean 10456 9 28 3 5 2 Shallow Clean 10456 9 29 3 5 2 Shallow Clean 10456 9 30 4 6 2 Shallow Clean 10456 9 31 5 6 2 246

Shallow Clean 10456 9 32 2 3 2 Shallow Clean 10456 9 33 4 6 2 Shallow Clean 10456 9 34 4 6 2 Shallow Clean 10456 9 35 2 3 2 Shallow Clean 10456 9 36 4 5 2 Shallow Clean 10456 9 37 5 6 2 Shallow Clean 10456 9 38 5 6 2 Shallow Clean 10456 9 39 4 5 2 Shallow Clean 10456 9 40 3 5 2 Shallow Clean 10456 9 41 4 5 2 Shallow Clean 10456 9 42 5 6 2 Shallow Clean 10456 9 43 4 5 2 Shallow Clean 10456 9 44 4 5 2 Shallow Clean 10456 9 45 3 4 2 Shallow Clean 10456 9 46 4 5 2 Shallow Clean 10456 9 47 4 5 1 Shallow Clean 10456 9 48 6 6 2 Shallow Clean 10456 10 1 2 3 2 Shallow Clean 10456 10 2 2 3 2 Shallow Clean 10456 10 3 1 2 2 Shallow Clean 10456 10 4 1 2 2 Shallow Clean 10456 10 5 3 3 2 Shallow Clean 10456 10 6 1 2 2 Shallow Clean 10456 10 7 2 3 2 Shallow Clean 10456 10 8 1 2 2 Shallow Clean 10456 10 9 1 2 2 Shallow Clean 10456 10 10 5 6 2 Shallow Clean 10456 10 11 6 6 2 Shallow Clean 10456 10 12 4 6 2 Shallow Clean 10456 10 13 4 5 2 Shallow Clean 10456 10 14 3 5 2 Shallow Clean 10456 10 15 4 5 2 Shallow Clean 10456 10 16 4 5 1 Shallow Clean 10456 10 17 3 5 1 Shallow Clean 10456 10 18 5 6 2 Shallow Clean 10456 10 19 3 5 1 Shallow Clean 10456 10 20 4 5 2 Shallow Clean 10456 10 21 4 6 2 Shallow Clean 10456 10 22 3 5 2 Shallow Clean 10456 10 23 4 5 2 Shallow Clean 10456 10 24 4 6 2 247

Shallow Clean 10456 10 25 6 6 2 Shallow Clean 10456 10 26 6 6 2 Shallow Clean 10456 11 1 1 2 2 Shallow Clean 10456 11 2 2 3 2 Shallow Clean 10456 11 3 5 6 2 Shallow Clean 10456 11 4 4 5 2 Shallow Clean 10456 11 5 6 6 2 Deep Desiccated 10457 1 1 3 6 1 Deep Desiccated 10457 1 2 3 5 1 Deep Desiccated 10457 1 3 5 6 1 Deep Desiccated 10457 1 4 5 6 1 Deep Desiccated 10457 2 1 4 6 1 Deep Desiccated 10457 2 2 5 6 1 Deep Desiccated 10457 2 3 4 6 1 Deep Desiccated 10457 2 4 2 5 2 Deep Desiccated 10457 2 5 3 6 1 Deep Desiccated 10457 2 6 2 5 2 Deep Desiccated 10457 2 7 4 6 1 Deep Desiccated 10457 3 1 3 6 1 Deep Desiccated 10457 3 2 3 4 1 Deep Desiccated 10457 3 3 2 3 2 Deep Desiccated 10457 3 4 2 3 2 Deep Desiccated 10457 3 5 3 4 1 Deep Desiccated 10457 3 6 3 5 2 Deep Desiccated 10457 3 7 3 6 1 Deep Desiccated 10457 3 8 3 5 2 Deep Desiccated 10457 3 9 3 5 1 Deep Desiccated 10457 3 10 3 6 1 Deep Desiccated 10457 3 11 3 5 1 Deep Desiccated 10457 3 12 3 4 1 Deep Desiccated 10457 3 13 3 4 2 Deep Desiccated 10457 3 14 2 3 2 Deep Desiccated 10457 3 15 3 5 2 Deep Desiccated 10457 3 16 5 6 1 Deep Desiccated 10457 3 17 3 6 1 Deep Desiccated 10457 3 18 3 6 1 Deep Desiccated 10457 3 19 3 6 2 Deep Desiccated 10457 3 20 3 6 1 Deep Desiccated 10457 3 21 3 6 1 Deep Desiccated 10457 3 22 3 6 1 Deep Desiccated 10457 3 23 2 5 2 248

Deep Desiccated 10457 3 24 3 6 2 Deep Desiccated 10457 3 25 3 5 1 Deep Desiccated 10457 3 26 5 6 1 Deep Desiccated 10457 3 27 4 6 1 Deep Desiccated 10457 3 28 4 6 2 Deep Desiccated 10457 3 29 5 6 1 Deep Desiccated 10457 3 30 3 6 1 Deep Desiccated 10457 4 1 3 6 1 Deep Desiccated 10457 4 2 3 5 1 Deep Desiccated 10457 4 3 3 6 1 Deep Desiccated 10457 4 4 5 6 1 Deep Desiccated 10457 4 5 4 5 2 Deep Desiccated 10457 4 6 4 6 2 Deep Desiccated 10457 4 7 3 5 2 Deep Desiccated 10457 4 8 3 5 2 Deep Desiccated 10457 4 9 3 5 1 Deep Desiccated 10457 4 10 3 4 2 Deep Desiccated 10457 4 11 3 5 2 Deep Desiccated 10457 4 12 3 4 2 Deep Desiccated 10457 4 13 3 4 1 Deep Desiccated 10457 4 14 3 4 1 Deep Desiccated 10457 4 15 3 5 1 Deep Desiccated 10457 4 16 3 5 1 Deep Desiccated 10457 4 17 3 5 1 Deep Desiccated 10457 4 18 3 5 2 Deep Desiccated 10457 4 19 3 4 1 Deep Desiccated 10457 4 20 3 5 1 Deep Desiccated 10457 4 21 3 4 1 Deep Desiccated 10457 4 22 3 6 1 Deep Desiccated 10457 4 23 3 4 2 Deep Desiccated 10457 4 24 3 4 2 Deep Desiccated 10457 4 25 5 6 2 Deep Desiccated 10457 4 26 3 5 2 Deep Desiccated 10457 4 27 3 5 1 Deep Desiccated 10457 4 28 3 4 1 Deep Desiccated 10457 4 29 3 4 1 Deep Desiccated 10457 4 30 3 5 1 Deep Desiccated 10457 4 31 3 4 2 Deep Desiccated 10457 4 32 2 5 2 Deep Desiccated 10457 4 33 3 5 2 Deep Desiccated 10457 4 34 4 6 2 249

Deep Desiccated 10457 4 35 5 6 1 Deep Desiccated 10457 4 36 4 5 2 Deep Desiccated 10457 4 37 5 6 2 Deep Desiccated 10457 4 38 3 5 2 Deep Desiccated 10457 4 39 3 5 2 Deep Desiccated 10457 4 40 5 6 1 Deep Desiccated 10457 4 41 4 5 1 Deep Desiccated 10457 4 42 4 6 1 Deep Desiccated 10457 4 43 4 6 1 Deep Desiccated 10457 4 44 5 6 1 Deep Desiccated 10457 4 45 5 6 1 Deep Desiccated 10457 4 46 3 6 1 Deep Desiccated 10457 4 47 4 5 0 Deep Desiccated 10457 4 48 4 6 1 Deep Desiccated 10457 4 49 3 6 1 Deep Desiccated 10457 4 50 4 6 1 Deep Desiccated 10457 4 51 3 4 1 Deep Desiccated 10457 5 1 4 6 2 Deep Desiccated 10457 5 2 4 6 1 Deep Desiccated 10457 5 3 4 6 1 Deep Desiccated 10457 5 4 3 5 2 Deep Desiccated 10457 5 5 5 6 1 Deep Desiccated 10457 5 6 5 6 2 Deep Desiccated 10457 5 7 3 5 1 Deep Desiccated 10457 5 8 4 6 1 Deep Desiccated 10457 5 9 3 6 2 Deep Desiccated 10457 5 10 4 6 1 Deep Desiccated 10457 5 11 4 6 1 Deep Desiccated 10457 5 12 na na na Deep Desiccated 10457 5 13 3 6 2 Deep Desiccated 10457 5 14 3 5 2 Deep Desiccated 10457 5 15 5 6 2 Deep Desiccated 10457 5 16 3 5 2 Deep Desiccated 10457 5 17 3 5 1 Deep Desiccated 10457 5 18 3 5 1 Deep Desiccated 10457 5 19 4 5 2 Deep Desiccated 10457 5 20 5 6 1 Deep Desiccated 10457 5 21 4 6 1 Deep Desiccated 10457 5 22 3 4 2 Deep Desiccated 10457 5 23 5 6 1 Deep Desiccated 10457 5 24 3 5 2 250

Deep Desiccated 10457 5 25 3 5 1 Deep Desiccated 10457 5 26 3 6 2 Deep Desiccated 10457 5 27 3 5 2 Deep Desiccated 10457 5 28 3 5 1 Deep Desiccated 10457 5 29 3 6 1 Deep Desiccated 10457 5 30 3 6 2 Deep Desiccated 10457 5 31 5 6 1 Deep Desiccated 10457 5 32 5 6 1 Deep Desiccated 10457 5 33 3 6 1 Deep Desiccated 10457 5 34 5 6 1 Deep Desiccated 10457 5 35 3 6 2 Deep Desiccated 10457 5 36 5 6 1 Deep Desiccated 10457 5 37 3 5 2 Deep Desiccated 10457 5 38 3 6 1 Deep Desiccated 10457 5 39 5 6 1 Deep Desiccated 10457 5 40 5 6 2 Deep Desiccated 10457 5 41 5 6 1 Deep Desiccated 10457 5 42 5 6 1 Deep Desiccated 10457 5 43 5 6 2 Deep Desiccated 10457 5 44 2 4 2 Deep Desiccated 10457 5 45 2 3 2 Deep Desiccated 10457 5 46 3 4 2 Deep Desiccated 10457 5 47 4 6 2 Deep Desiccated 10457 5 48 5 6 2 Deep Desiccated 10457 5 49 4 6 1 Deep Desiccated 10457 5 50 4 6 2 Deep Desiccated 10457 5 51 4 6 2 Deep Desiccated 10457 5 52 4 6 2 Deep Desiccated 10457 5 53 6 6 2 Deep Desiccated 10457 5 54 4 6 2 Deep Desiccated 10457 5 55 4 6 1 Deep Desiccated 10457 5 56 3 5 2 Deep Desiccated 10457 5 57 3 5 2 Deep Desiccated 10457 5 58 3 5 2 Deep Desiccated 10457 5 59 3 5 2 Deep Desiccated 10457 5 60 3 5 2 Deep Desiccated 10457 5 61 3 5 1 Deep Desiccated 10457 5 62 3 5 1 Deep Desiccated 10457 5 63 3 6 2 Deep Desiccated 10457 5 64 4 6 1 Deep Desiccated 10457 5 65 4 6 2 251

Deep Desiccated 10457 5 66 4 6 2 Deep Desiccated 10457 5 67 4 6 1 Deep Desiccated 10457 5 68 3 4 2 Deep Desiccated 10457 6 1 5 6 1 Deep Desiccated 10457 6 2 4 6 1 Deep Desiccated 10457 6 3 3 4 1 Deep Desiccated 10457 6 4 4 6 1 Deep Desiccated 10457 6 5 4 6 1 Deep Desiccated 10457 6 6 5 6 1 Deep Desiccated 10457 6 7 3 5 1 Deep Desiccated 10457 6 8 5 6 1 Deep Desiccated 10457 6 9 3 4 1 Deep Desiccated 10457 6 10 3 5 2 Deep Desiccated 10457 6 11 3 5 1 Deep Desiccated 10457 6 12 3 4 2 Deep Desiccated 10457 6 13 5 6 1 Deep Desiccated 10457 6 14 4 6 1 Deep Desiccated 10457 6 15 4 6 1 Deep Desiccated 10457 6 16 3 4 2 Deep Desiccated 10457 6 17 3 5 2 Deep Desiccated 10457 6 18 3 5 1 Deep Desiccated 10457 6 19 4 6 2 Deep Desiccated 10457 6 20 3 4 2 Deep Desiccated 10457 6 21 3 5 2 Deep Desiccated 10457 6 22 3 6 2 Deep Desiccated 10457 6 23 3 6 2 Deep Desiccated 10457 6 24 3 5 2 Deep Desiccated 10457 6 25 3 6 1 Deep Desiccated 10457 6 26 3 6 1 Deep Desiccated 10457 6 27 4 6 1 Deep Desiccated 10457 6 28 4 6 1 Deep Desiccated 10457 6 29 5 6 1 Deep Desiccated 10457 6 30 4 6 1 Deep Desiccated 10457 6 31 4 6 2 Deep Desiccated 10457 6 32 4 6 1 Deep Desiccated 10457 6 33 4 6 1 Deep Desiccated 10457 6 34 4 5 0 Deep Desiccated 10457 6 35 3 6 2 Deep Desiccated 10457 6 36 3 5 1 Deep Desiccated 10457 6 37 3 4 2 Deep Desiccated 10457 7 1 5 6 1 252

Deep Desiccated 10457 7 2 5 6 1 Deep Desiccated 10457 7 3 4 6 1 Deep Desiccated 10457 7 4 4 6 1 Deep Desiccated 10457 7 5 3 5 1 Deep Desiccated 10457 7 6 4 6 1 Deep Desiccated 10457 7 7 3 5 1 Deep Desiccated 10457 7 8 4 5 1 Deep Desiccated 10457 7 9 3 5 1 Deep Desiccated 10457 7 10 3 4 1 Deep Desiccated 10457 7 11 3 4 2 Deep Desiccated 10457 7 12 3 5 1 Deep Desiccated 10457 7 13 3 4 1 Deep Desiccated 10457 7 14 3 4 1 Deep Desiccated 10457 7 15 4 6 1 Deep Desiccated 10457 7 16 4 6 1 Deep Desiccated 10457 7 17 3 5 2 Deep Desiccated 10457 7 18 4 6 1 Deep Desiccated 10457 7 19 3 5 1 Deep Desiccated 10457 7 20 3 5 1 Deep Desiccated 10457 7 21 5 6 1 Deep Desiccated 10457 7 22 3 5 1 Deep Desiccated 10457 7 23 3 5 1 Deep Desiccated 10457 7 24 4 6 1 Deep Desiccated 10457 7 25 3 6 2 Deep Desiccated 10457 7 26 4 6 2 Deep Desiccated 10457 8 1 5 6 1 Deep Desiccated 10457 8 2 4 6 2 Deep Desiccated 10457 8 3 6 6 1 Deep Desiccated 10457 8 4 5 6 1 Deep Desiccated 10457 8 5 3 5 1 Deep Desiccated 10457 8 6 3 5 0 Deep Desiccated 10457 8 7 4 6 1 Deep Desiccated 10457 8 8 4 6 1 Deep Desiccated 10457 8 9 5 6 1 Deep Desiccated 10457 8 10 3 4 1 Deep Desiccated 10457 8 11 4 6 1 Deep Desiccated 10457 8 12 3 4 1 Deep Desiccated 10457 8 13 4 6 1 Deep Desiccated 10457 8 14 5 6 2 Deep Desiccated 10457 8 15 4 6 2 Deep Desiccated 10457 8 16 3 5 1 253

Deep Desiccated 10457 9 1 4 6 1 Deep Desiccated 10457 9 2 3 6 1 Deep Desiccated 10457 9 3 4 6 1 Deep Desiccated 10457 9 4 3 6 1 Deep Desiccated 10457 9 5 4 6 1 Deep Fresh 10470 1 1 5 6 1 Deep Fresh 10470 1 2 3 4 2 Deep Fresh 10470 1 3 3 4 1 Deep Fresh 10470 1 4 3 4 2 Deep Fresh 10470 1 5 5 6 1 Deep Fresh 10470 1 6 4 6 1 Deep Fresh 10470 1 7 5 6 2 Deep Fresh 10470 1 8 6 6 1 Deep Fresh 10470 1 9 5 6 1 Deep Fresh 10470 1 10 6 6 1 Deep Fresh 10470 2 1 4 6 2 Deep Fresh 10470 2 2 3 4 2 Deep Fresh 10470 2 3 3 4 1 Deep Fresh 10470 2 4 3 5 2 Deep Fresh 10470 2 5 3 5 2 Deep Fresh 10470 2 6 3 4 2 Deep Fresh 10470 2 7 3 5 2 Deep Fresh 10470 2 8 4 6 1 Deep Fresh 10470 2 9 4 6 1 Deep Fresh 10470 2 10 3 5 2 Deep Fresh 10470 2 11 3 5 1 Deep Fresh 10470 2 12 3 5 2 Deep Fresh 10470 2 13 4 6 1 Deep Fresh 10470 2 14 3 6 1 Deep Fresh 10470 2 15 4 6 1 Deep Fresh 10470 2 16 3 6 1 Deep Fresh 10470 2 17 3 5 2 Deep Fresh 10470 2 18 4 5 2 Deep Fresh 10470 2 19 4 6 1 Deep Fresh 10470 2 20 4 5 2 Deep Fresh 10470 2 21 3 6 2 Deep Fresh 10470 2 22 3 4 2 Deep Fresh 10470 2 23 3 6 2 Deep Fresh 10470 2 24 3 5 2 Deep Fresh 10470 2 25 3 6 2 Deep Fresh 10470 2 26 4 6 1 254

Deep Fresh 10470 2 27 5 6 1 Deep Fresh 10470 3 1 4 6 1 Deep Fresh 10470 3 2 4 6 2 Deep Fresh 10470 3 3 5 6 2 Deep Fresh 10470 3 4 4 6 2 Deep Fresh 10470 3 5 4 6 2 Deep Fresh 10470 3 6 4 6 1 Deep Fresh 10470 3 7 3 4 2 Deep Fresh 10470 3 8 3 4 2 Deep Fresh 10470 3 9 4 6 2 Deep Fresh 10470 3 10 3 4 2 Deep Fresh 10470 3 11 6 6 2 Deep Fresh 10470 3 12 6 6 1 Deep Fresh 10470 3 13 4 6 1 Deep Fresh 10470 3 14 5 6 1 Deep Fresh 10470 3 15 4 6 1 Deep Fresh 10470 3 16 4 6 2 Deep Fresh 10470 3 17 3 5 2 Deep Fresh 10470 3 18 3 5 2 Deep Fresh 10470 3 19 4 6 2 Deep Fresh 10470 3 20 4 6 2 Deep Fresh 10470 3 21 4 6 1 Deep Fresh 10470 3 22 4 6 1 Deep Fresh 10470 3 23 5 6 1 Deep Fresh 10470 3 24 6 6 1 Deep Fresh 10470 3 25 6 6 1 Deep Fresh 10470 3 26 3 4 2 Deep Fresh 10470 3 27 5 6 1 Deep Fresh 10470 3 28 3 4 2 Deep Fresh 10470 3 29 3 4 1 Deep Fresh 10470 3 30 3 4 2 Deep Fresh 10470 4 1 4 6 1 Deep Fresh 10470 4 2 4 6 2 Deep Fresh 10470 4 3 3 5 2 Deep Fresh 10470 4 4 4 6 1 Deep Fresh 10470 4 5 3 4 2 Deep Fresh 10470 4 6 3 4 2 Deep Fresh 10470 4 7 3 4 1 Deep Fresh 10470 4 8 3 4 2 Deep Fresh 10470 4 9 na na na Deep Fresh 10470 4 10 4 6 2 255

Deep Fresh 10470 4 11 6 6 2 Deep Fresh 10470 4 12 5 6 2 Deep Fresh 10470 4 13 4 6 1 Deep Fresh 10470 4 14 4 6 1 Deep Fresh 10470 4 15 6 6 2 Deep Fresh 10470 4 16 3 4 2 Deep Fresh 10470 4 17 5 6 2 Deep Fresh 10470 4 18 6 6 2 Deep Fresh 10470 4 19 4 6 2 Deep Fresh 10470 4 20 4 6 1 Deep Fresh 10470 4 21 na na na Deep Fresh 10470 4 22 4 6 1 Deep Fresh 10470 4 23 5 6 2 Deep Fresh 10470 4 24 4 6 1 Deep Fresh 10470 4 25 3 4 1 Deep Fresh 10470 4 26 2 3 2 Deep Fresh 10470 4 27 2 5 2 Deep Fresh 10470 4 28 3 5 1 Deep Fresh 10470 4 29 5 6 1 Deep Fresh 10470 4 30 5 6 1 Deep Fresh 10470 4 31 3 4 2 Deep Fresh 10470 4 32 3 4 2 Deep Fresh 10470 4 33 3 5 2 Deep Fresh 10470 4 34 3 6 2 Deep Fresh 10470 5 1 4 6 1 Deep Fresh 10470 5 2 3 4 2 Deep Fresh 10470 5 3 4 5 2 Deep Fresh 10470 5 4 4 5 2 Deep Fresh 10470 5 5 3 4 2 Deep Fresh 10470 5 6 4 5 2 Deep Fresh 10470 5 7 5 6 1 Deep Fresh 10470 5 8 3 4 2 Deep Fresh 10470 5 9 5 6 2 Deep Fresh 10470 5 10 4 6 1 Deep Fresh 10470 5 11 2 3 2 Deep Fresh 10470 5 12 2 3 2 Deep Fresh 10470 5 13 3 4 2 Deep Fresh 10470 5 14 6 6 2 Deep Fresh 10470 5 15 5 6 2 Deep Fresh 10470 5 16 5 6 2 Deep Fresh 10470 5 17 5 6 2 256

Deep Fresh 10470 5 18 5 6 1 Deep Fresh 10470 5 19 6 6 1 Deep Fresh 10470 5 20 4 6 1 Deep Fresh 10470 5 21 5 6 2 Deep Fresh 10470 5 22 3 4 2 Deep Fresh 10470 5 23 4 6 1 Deep Fresh 10470 5 24 4 6 1 Deep Fresh 10470 5 25 3 6 2 Deep Fresh 10470 5 26 5 6 2 Deep Fresh 10470 5 27 3 4 1 Deep Fresh 10470 5 28 3 4 2 Deep Fresh 10470 5 29 4 6 1 Deep Fresh 10470 5 30 Na Na na Deep Fresh 10470 5 31 5 6 2 Deep Fresh 10470 5 32 2 3 1 Deep Fresh 10470 5 33 2 3 2 Deep Fresh 10470 5 34 3 5 1 Deep Fresh 10470 5 35 3 4 1 Deep Fresh 10470 5 36 4 6 1 Deep Fresh 10470 5 37 4 6 1 Deep Fresh 10470 5 38 3 5 2 Deep Fresh 10470 5 39 3 5 1 Deep Fresh 10470 5 40 3 5 2 Deep Fresh 10470 5 41 4 6 1 Deep Fresh 10470 5 42 5 6 1 Deep Fresh 10470 5 43 3 6 1 Deep Fresh 10470 6 1 3 6 1 Deep Fresh 10470 6 2 4 6 1 Deep Fresh 10470 6 3 4 5 2 Deep Fresh 10470 6 4 4 6 2 Deep Fresh 10470 6 5 3 4 2 Deep Fresh 10470 6 6 3 5 1 Deep Fresh 10470 6 7 5 6 1 Deep Fresh 10470 6 8 4 6 1 Deep Fresh 10470 6 9 3 4 2 Deep Fresh 10470 6 10 3 5 1 Deep Fresh 10470 6 11 5 6 2 Deep Fresh 10470 6 12 3 5 1 Deep Fresh 10470 6 13 3 4 2 Deep Fresh 10470 6 14 4 6 1 Deep Fresh 10470 6 15 5 6 2 257

Deep Fresh 10470 6 16 6 6 1 Deep Fresh 10470 6 17 3 4 2 Deep Fresh 10470 6 18 5 6 1 Deep Fresh 10470 6 19 3 5 2 Deep Fresh 10470 6 20 5 6 2 Deep Fresh 10470 6 21 5 6 2 Deep Fresh 10470 6 22 5 6 2 Deep Fresh 10470 6 23 5 6 2 Deep Fresh 10470 6 24 3 5 2 Deep Fresh 10470 6 25 4 6 1 Deep Fresh 10470 6 26 3 4 2 Deep Fresh 10470 6 27 3 4 2 Deep Fresh 10470 6 28 5 6 1 Deep Fresh 10470 6 29 5 6 2 Deep Fresh 10470 6 30 5 6 2 Deep Fresh 10470 6 31 3 4 2 Deep Fresh 10470 6 32 4 5 2 Deep Fresh 10470 6 33 3 5 1 Deep Fresh 10470 6 34 4 5 1 Deep Fresh 10470 6 35 3 5 2 Deep Fresh 10470 6 36 5 6 1 Deep Fresh 10470 6 37 2 2 2 Deep Fresh 10470 6 38 6 6 2 Deep Fresh 10470 6 39 5 6 1 Deep Fresh 10470 7 1 5 6 1 Deep Fresh 10470 7 2 5 6 1 Deep Fresh 10470 7 3 4 6 1 Deep Fresh 10470 7 4 4 6 1 Deep Fresh 10470 7 5 5 6 1 Deep Fresh 10470 7 6 5 6 1 Deep Fresh 10470 7 7 4 6 1 Deep Fresh 10470 7 8 5 6 1 Deep Fresh 10470 7 9 3 5 2 Deep Fresh 10470 7 10 4 6 2 Deep Fresh 10470 7 11 3 5 2 Deep Fresh 10470 7 12 5 6 2 Deep Fresh 10470 7 13 3 4 2 Deep Fresh 10470 7 14 3 4 2 Deep Fresh 10470 7 15 4 6 1 Deep Fresh 10470 7 16 3 4 2 Deep Fresh 10470 7 17 3 3 2 258

Deep Fresh 10470 7 18 3 5 1 Deep Fresh 10470 7 19 2 3 2 Deep Fresh 10470 7 20 3 4 2 Deep Fresh 10470 7 21 3 4 2 Deep Fresh 10470 7 22 2 3 2 Deep Fresh 10470 7 23 1 3 2 Deep Fresh 10470 7 24 3 4 2 Deep Fresh 10470 7 25 3 4 2 Deep Fresh 10470 7 26 3 4 2 Deep Fresh 10470 7 27 3 5 2 Deep Fresh 10470 7 28 1 3 2 Deep Fresh 10470 7 29 3 4 2 Deep Fresh 10470 7 30 3 6 1 Deep Fresh 10470 7 31 5 6 1 Deep Fresh 10470 7 32 5 6 1 Deep Fresh 10470 7 33 4 6 1 Deep Fresh 10470 7 34 3 5 2 Deep Fresh 10470 7 35 3 5 2 Deep Fresh 10470 7 36 3 5 1 Deep Fresh 10470 7 37 5 6 2 Deep Fresh 10470 7 38 6 6 1 Deep Fresh 10470 7 39 5 6 2 Deep Fresh 10470 7 40 5 6 1 Deep Fresh 10470 7 41 3 4 2 Deep Fresh 10470 7 42 3 4 2 Deep Fresh 10470 7 43 3 5 2 Deep Fresh 10470 7 44 3 4 2 Deep Fresh 10470 7 45 4 5 1 Deep Fresh 10470 7 46 5 6 1 Deep Fresh 10470 8 1 4 6 1 Deep Fresh 10470 8 2 5 6 1 Deep Fresh 10470 8 3 4 6 1 Deep Fresh 10470 8 4 3 5 1 Deep Fresh 10470 8 5 2 4 2 Deep Fresh 10470 8 6 2 5 2 Deep Fresh 10470 8 7 3 4 2 Deep Fresh 10470 8 8 2 4 2 Deep Fresh 10470 8 9 5 6 1 Deep Fresh 10470 8 10 4 6 1 Deep Fresh 10470 8 11 3 6 2 Deep Fresh 10470 8 12 3 4 2 259

Deep Fresh 10470 8 13 3 5 1 Deep Fresh 10470 8 14 3 4 1 Deep Fresh 10470 8 15 5 6 1 Deep Fresh 10470 8 16 4 6 1 Deep Fresh 10470 9 1 4 6 1 Deep Fresh 10470 9 2 4 6 1 Deep Fresh 10470 9 3 5 6 1 Deep Fresh 10470 9 4 4 5 1 Deep Fresh 10470 9 5 3 5 1 Deep Fresh 10470 9 6 4 6 1 Deep Fresh 10470 9 7 3 4 1 Deep Fresh 10470 9 8 5 6 1 Shallow Fresh 10471 1 1 1 2 2 Shallow Fresh 10471 1 2 2 3 2 Shallow Fresh 10471 1 3 2 3 2 Shallow Fresh 10471 1 4 3 3 1 Shallow Fresh 10471 1 5 2 3 2 Shallow Fresh 10471 1 6 1 2 2 Shallow Fresh 10471 1 7 5 6 1 Shallow Fresh 10471 1 8 4 5 1 Shallow Fresh 10471 1 9 6 6 1 Shallow Fresh 10471 1 10 5 6 1 Shallow Fresh 10471 1 11 3 4 1 Shallow Fresh 10471 1 12 6 5 0 Shallow Fresh 10471 1 13 4 5 1 Shallow Fresh 10471 1 14 3 5 1 Shallow Fresh 10471 1 15 4 6 1 Shallow Fresh 10471 1 16 3 4 1 Shallow Fresh 10471 1 17 3 5 1 Shallow Fresh 10471 1 18 5 6 1 Shallow Fresh 10471 2 1 1 2 2 Shallow Fresh 10471 2 2 2 3 2 Shallow Fresh 10471 2 3 2 3 2 Shallow Fresh 10471 2 4 2 6 2 Shallow Fresh 10471 2 5 6 6 2 Shallow Fresh 10471 2 6 5 6 1 Shallow Fresh 10471 2 7 3 5 1 Shallow Fresh 10471 2 8 4 5 1 Shallow Fresh 10471 2 9 5 6 2 Shallow Fresh 10471 2 10 1 3 2 Shallow Fresh 10471 2 11 2 3 2 260

Shallow Fresh 10471 2 12 3 3 2 Shallow Fresh 10471 2 13 5 6 2 Shallow Fresh 10471 2 14 5 6 2 Shallow Fresh 10471 2 15 3 5 2 Shallow Fresh 10471 2 16 4 6 1 Shallow Fresh 10471 2 17 1 3 2 Shallow Fresh 10471 2 18 3 6 1 Shallow Fresh 10471 2 19 3 4 1 Shallow Fresh 10471 2 20 1 2 2 Shallow Fresh 10471 2 21 3 6 1 Shallow Fresh 10471 2 22 3 5 1 Shallow Fresh 10471 2 23 3 5 1 Shallow Fresh 10471 2 24 3 5 2 Shallow Fresh 10471 2 25 3 5 2 Shallow Fresh 10471 2 26 4 6 2 Shallow Fresh 10471 2 27 5 6 1 Shallow Fresh 10471 2 28 3 5 2 Shallow Fresh 10471 2 29 4 6 2 Shallow Fresh 10471 2 30 2 3 2 Shallow Fresh 10471 2 31 3 3 2 Shallow Fresh 10471 2 32 2 4 2 Shallow Fresh 10471 2 33 3 4 2 Shallow Fresh 10471 2 34 2 3 2 Shallow Fresh 10471 2 35 5 6 1 Shallow Fresh 10471 2 36 5 6 2 Shallow Fresh 10471 2 37 1 2 2 Shallow Fresh 10471 2 38 2 3 2 Shallow Fresh 10471 2 39 1 2 2 Shallow Fresh 10471 2 40 2 2 2 Shallow Fresh 10471 3 1 1 2 2 Shallow Fresh 10471 3 2 2 3 2 Shallow Fresh 10471 3 3 6 6 2 Shallow Fresh 10471 3 4 5 6 1 Shallow Fresh 10471 3 5 6 6 1 Shallow Fresh 10471 3 6 6 6 1 Shallow Fresh 10471 3 7 3 6 1 Shallow Fresh 10471 3 8 3 6 1 Shallow Fresh 10471 3 9 1 2 2 Shallow Fresh 10471 3 10 6 6 2 Shallow Fresh 10471 3 11 6 6 2 Shallow Fresh 10471 3 12 3 5 2 261

Shallow Fresh 10471 3 13 5 6 2 Shallow Fresh 10471 3 14 4 4 1 Shallow Fresh 10471 3 15 5 6 2 Shallow Fresh 10471 3 16 3 3 0 Shallow Fresh 10471 3 17 4 6 1 Shallow Fresh 10471 3 18 6 6 1 Shallow Fresh 10471 3 19 5 6 1 Shallow Fresh 10471 3 20 4 6 2 Shallow Fresh 10471 3 21 6 6 1 Shallow Fresh 10471 3 22 5 6 1 Shallow Fresh 10471 3 23 5 6 2 Shallow Fresh 10471 3 24 4 6 2 Shallow Fresh 10471 3 25 3 5 2 Shallow Fresh 10471 3 26 5 6 2 Shallow Fresh 10471 3 27 6 6 1 Shallow Fresh 10471 3 28 4 5 2 Shallow Fresh 10471 3 29 4 6 2 Shallow Fresh 10471 3 30 3 4 2 Shallow Fresh 10471 3 31 1 3 2 Shallow Fresh 10471 4 1 1 3 2 Shallow Fresh 10471 4 2 3 6 2 Shallow Fresh 10471 4 3 3 5 2 Shallow Fresh 10471 4 4 3 6 2 Shallow Fresh 10471 4 5 4 6 2 Shallow Fresh 10471 4 6 6 6 2 Shallow Fresh 10471 4 7 4 5 2 Shallow Fresh 10471 4 8 5 6 2 Shallow Fresh 10471 4 9 6 6 2 Shallow Fresh 10471 4 10 3 5 2 Shallow Fresh 10471 4 11 1 3 2 Shallow Fresh 10471 4 12 1 6 2 Shallow Fresh 10471 4 13 3 6 2 Shallow Fresh 10471 4 14 3 5 2 Shallow Fresh 10471 4 15 3 5 1 Shallow Fresh 10471 4 16 6 6 1 Shallow Fresh 10471 4 17 3 5 2 Shallow Fresh 10471 4 18 6 6 2 Shallow Fresh 10471 4 19 2 6 2 Shallow Fresh 10471 4 20 4 6 2 Shallow Fresh 10471 4 21 5 6 2 Shallow Fresh 10471 4 22 1 3 2 262

Shallow Fresh 10471 4 23 3 4 1 Shallow Fresh 10471 4 24 1 3 2 Shallow Fresh 10471 4 25 3 4 1 Shallow Fresh 10471 4 26 1 3 2 Shallow Fresh 10471 4 27 6 6 2 Shallow Fresh 10471 4 28 3 5 1 Shallow Fresh 10471 4 29 3 4 2 Shallow Fresh 10471 4 30 6 6 2 Shallow Fresh 10471 4 31 6 6 1 Shallow Fresh 10471 4 32 6 6 2 Shallow Fresh 10471 4 33 5 6 2 Shallow Fresh 10471 4 34 4 6 2 Shallow Fresh 10471 4 35 4 6 2 Shallow Fresh 10471 4 36 5 6 2 Shallow Fresh 10471 4 37 4 5 1 Shallow Fresh 10471 4 38 2 5 2 Shallow Fresh 10471 4 39 1 3 2 Shallow Fresh 10471 4 40 5 6 2 Shallow Fresh 10471 4 41 5 6 2 Shallow Fresh 10471 4 42 5 6 2 Shallow Fresh 10471 4 43 4 6 2 Shallow Fresh 10471 4 44 5 6 1 Shallow Fresh 10471 4 45 1 3 2 Shallow Fresh 10471 4 46 1 3 2 Shallow Fresh 10471 4 47 3 3 1 Shallow Fresh 10471 4 48 3 4 1 Shallow Fresh 10471 4 49 2 4 2 Shallow Fresh 10471 4 50 2 3 2 Shallow Fresh 10471 4 51 1 2 2 Shallow Fresh 10471 4 52 2 3 2 Shallow Fresh 10471 5 1 1 3 2 Shallow Fresh 10471 5 2 1 3 2 Shallow Fresh 10471 5 3 2 3 2 Shallow Fresh 10471 5 4 3 4 1 Shallow Fresh 10471 5 5 2 3 2 Shallow Fresh 10471 5 6 2 3 1 Shallow Fresh 10471 5 7 2 3 2 Shallow Fresh 10471 5 8 6 6 2 Shallow Fresh 10471 5 9 4 6 2 Shallow Fresh 10471 5 10 6 6 2 Shallow Fresh 10471 5 11 6 6 2 263

Shallow Fresh 10471 5 12 1 3 2 Shallow Fresh 10471 5 13 2 3 2 Shallow Fresh 10471 5 14 3 4 2 Shallow Fresh 10471 5 15 3 5 1 Shallow Fresh 10471 5 16 3 6 2 Shallow Fresh 10471 5 17 3 4 2 Shallow Fresh 10471 5 18 3 4 1 Shallow Fresh 10471 5 19 3 6 2 Shallow Fresh 10471 5 20 4 6 1 Shallow Fresh 10471 5 21 6 5 0 Shallow Fresh 10471 5 22 6 6 1 Shallow Fresh 10471 5 23 3 6 1 Shallow Fresh 10471 5 24 3 6 1 Shallow Fresh 10471 5 25 3 5 1 Shallow Fresh 10471 5 26 3 3 0 Shallow Fresh 10471 5 27 3 5 1 Shallow Fresh 10471 5 28 6 6 1 Shallow Fresh 10471 5 29 4 6 1 Shallow Fresh 10471 5 30 4 6 1 Shallow Fresh 10471 5 31 4 6 2 Shallow Fresh 10471 5 32 6 6 1 Shallow Fresh 10471 5 33 6 6 2 Shallow Fresh 10471 5 34 1 4 2 Shallow Fresh 10471 5 35 1 3 2 Shallow Fresh 10471 5 36 3 4 1 Shallow Fresh 10471 5 37 6 6 1 Shallow Fresh 10471 5 38 1 4 2 Shallow Fresh 10471 5 39 6 6 1 Shallow Fresh 10471 5 40 4 5 2 Shallow Fresh 10471 5 41 5 6 2 Shallow Fresh 10471 5 42 4 5 1 Shallow Fresh 10471 5 43 3 5 2 Shallow Fresh 10471 5 44 3 4 1 Shallow Fresh 10471 5 45 5 6 2 Shallow Fresh 10471 5 46 4 6 2 Shallow Fresh 10471 5 47 5 6 2 Shallow Fresh 10471 5 48 4 5 2 Shallow Fresh 10471 5 49 5 6 2 Shallow Fresh 10471 5 50 4 5 2 Shallow Fresh 10471 5 51 5 6 2 Shallow Fresh 10471 5 52 6 6 2 264

Shallow Fresh 10471 5 53 2 6 2 Shallow Fresh 10471 5 54 3 4 2 Shallow Fresh 10471 5 55 2 3 2 Shallow Fresh 10471 5 56 1 3 2 Shallow Fresh 10471 5 57 1 3 2 Shallow Fresh 10471 5 58 3 4 1 Shallow Fresh 10471 5 59 2 4 2 Shallow Fresh 10471 5 60 4 6 2 Shallow Fresh 10471 5 61 5 6 1 Shallow Fresh 10471 5 62 4 6 1 Shallow Fresh 10471 5 63 4 6 1 Shallow Fresh 10471 5 64 5 6 2 Shallow Fresh 10471 5 65 5 6 2 Shallow Fresh 10471 5 66 2 4 2 Shallow Fresh 10471 5 67 2 3 2 Shallow Fresh 10471 5 68 1 3 2 Shallow Fresh 10471 5 69 6 6 1 Shallow Fresh 10471 5 70 6 6 1 Shallow Fresh 10471 5 71 3 5 1 Shallow Fresh 10471 6 1 1 4 2 Shallow Fresh 10471 6 2 1 3 2 Shallow Fresh 10471 6 3 2 5 2 Shallow Fresh 10471 6 4 3 4 1 Shallow Fresh 10471 6 5 2 4 2 Shallow Fresh 10471 6 6 5 6 2 Shallow Fresh 10471 6 7 4 6 2 Shallow Fresh 10471 6 8 5 6 2 Shallow Fresh 10471 6 9 3 4 2 Shallow Fresh 10471 6 10 1 3 2 Shallow Fresh 10471 6 11 1 4 2 Shallow Fresh 10471 6 12 1 6 2 Shallow Fresh 10471 6 13 1 4 2 Shallow Fresh 10471 6 14 1 2 2 Shallow Fresh 10471 6 15 1 3 2 Shallow Fresh 10471 6 16 1 3 2 Shallow Fresh 10471 6 17 6 6 2 Shallow Fresh 10471 6 18 5 6 2 Shallow Fresh 10471 6 19 4 6 2 Shallow Fresh 10471 6 20 6 6 2 Shallow Fresh 10471 6 21 6 6 1 Shallow Fresh 10471 6 22 3 5 2 265

Shallow Fresh 10471 6 23 3 6 1 Shallow Fresh 10471 6 24 5 6 1 Shallow Fresh 10471 6 25 4 6 1 Shallow Fresh 10471 6 26 4 6 1 Shallow Fresh 10471 6 27 4 6 2 Shallow Fresh 10471 6 28 4 6 1 Shallow Fresh 10471 6 29 3 4 2 Shallow Fresh 10471 6 30 1 2 2 Shallow Fresh 10471 6 31 6 6 1 Shallow Fresh 10471 6 32 3 4 1 Shallow Fresh 10471 6 33 3 5 1 Shallow Fresh 10471 6 34 6 6 1 Shallow Fresh 10471 6 35 5 6 1 Shallow Fresh 10471 6 36 5 6 2 Shallow Fresh 10471 6 37 3 4 2 Shallow Fresh 10471 6 38 5 6 1 Shallow Fresh 10471 6 39 6 6 2 Shallow Fresh 10471 6 40 1 6 2 Shallow Fresh 10471 6 41 5 6 2 Shallow Fresh 10471 6 42 5 6 2 Shallow Fresh 10471 6 43 5 6 2 Shallow Fresh 10471 6 44 4 6 2 Shallow Fresh 10471 6 45 4 6 2 Shallow Fresh 10471 6 46 6 6 2 Shallow Fresh 10471 6 47 2 4 2 Shallow Fresh 10471 6 48 3 4 1 Shallow Fresh 10471 6 49 6 6 2 Shallow Fresh 10471 6 50 3 6 1 Shallow Fresh 10471 6 51 6 6 2 Shallow Fresh 10471 6 52 2 3 2 Shallow Fresh 10471 6 53 1 3 2 Shallow Fresh 10471 6 54 1 2 2 Shallow Fresh 10471 6 55 1 3 2 Shallow Fresh 10471 7 1 2 3 2 Shallow Fresh 10471 7 2 1 2 2 Shallow Fresh 10471 7 3 1 2 2 Shallow Fresh 10471 7 4 2 3 2 Shallow Fresh 10471 7 5 1 3 2 Shallow Fresh 10471 7 6 6 6 2 Shallow Fresh 10471 7 7 6 6 2 Shallow Fresh 10471 7 8 1 2 2 266

Shallow Fresh 10471 7 9 5 6 1 Shallow Fresh 10471 7 10 1 3 2 Shallow Fresh 10471 7 11 6 6 2 Shallow Fresh 10471 7 12 6 6 2 Shallow Fresh 10471 7 13 5 6 2 Shallow Fresh 10471 7 14 6 6 2 Shallow Fresh 10471 7 15 6 6 2 Shallow Fresh 10471 7 16 3 4 1 Shallow Fresh 10471 7 17 3 4 1 Shallow Fresh 10471 7 18 5 6 1 Shallow Fresh 10471 7 19 5 6 2 Shallow Fresh 10471 7 20 3 5 1 Shallow Fresh 10471 7 21 6 6 1 Shallow Fresh 10471 8 1 1 3 2 Shallow Fresh 10471 8 2 3 6 2 Shallow Fresh 10471 8 3 6 6 2 Shallow Fresh 10471 8 4 6 6 2 Shallow Fresh 10471 8 5 5 6 2 Shallow Fresh 10471 8 6 6 6 2 Shallow Fresh 10471 8 7 4 6 1 Shallow Fresh 10471 8 8 4 6 1 Shallow Fresh 10471 8 9 3 5 0 Shallow Fresh 10471 8 10 3 5 0 Shallow Fresh 10471 8 11 3 4 0 Shallow Fresh 10471 8 12 6 6 1 Shallow Fresh 10471 8 13 5 6 2 Shallow Fresh 10471 8 14 5 6 1 Shallow Fresh 10471 8 15 6 6 1 Deep Clean 10498 1 1 3 5 1 Deep Clean 10498 1 2 3 6 1 Deep Clean 10498 1 3 3 5 1 Deep Clean 10498 1 4 3 6 1 Deep Clean 10498 1 5 2 3 2 Deep Clean 10498 1 6 3 5 1 Deep Clean 10498 1 7 3 6 1 Deep Clean 10498 1 8 3 6 1 Deep Clean 10498 1 9 4 6 2 Deep Clean 10498 1 10 3 6 2 Deep Clean 10498 1 11 5 6 1 Deep Clean 10498 1 12 4 6 1 Deep Clean 10498 1 13 4 5 1 267

Deep Clean 10498 1 14 4 6 2 Deep Clean 10498 1 15 4 6 2 Deep Clean 10498 1 16 2 3 2 Deep Clean 10498 2 1 1 3 2 Deep Clean 10498 2 2 3 5 1 Deep Clean 10498 2 3 3 5 1 Deep Clean 10498 2 4 1 3 2 Deep Clean 10498 2 5 3 5 2 Deep Clean 10498 2 6 3 5 1 Deep Clean 10498 2 7 1 3 2 Deep Clean 10498 2 8 3 5 1 Deep Clean 10498 2 9 4 6 2 Deep Clean 10498 2 10 4 6 2 Deep Clean 10498 3 1 3 6 2 Deep Clean 10498 3 2 4 6 2 Deep Clean 10498 3 3 3 5 2 Deep Clean 10498 3 4 4 6 2 Deep Clean 10498 3 5 4 6 2 Deep Clean 10498 3 6 4 6 2 Deep Clean 10498 3 7 5 6 1 Deep Clean 10498 3 8 5 6 1 Deep Clean 10498 3 9 3 6 2 Deep Clean 10498 3 10 4 6 1 Deep Clean 10498 3 11 3 5 2 Deep Clean 10498 3 12 4 6 2 Deep Clean 10498 3 13 5 6 1 Deep Clean 10498 3 14 3 5 1 Deep Clean 10498 3 15 5 6 2 Deep Clean 10498 3 16 4 6 2 Deep Clean 10498 3 17 2 5 2 Deep Clean 10498 3 18 3 6 2 Deep Clean 10498 3 19 2 3 2 Deep Clean 10498 3 20 3 5 1 Deep Clean 10498 3 21 4 6 1 Deep Clean 10498 3 22 4 6 2 Deep Clean 10498 3 23 3 6 1 Deep Clean 10498 3 24 3 6 1 Deep Clean 10498 3 25 5 6 2 Deep Clean 10498 3 26 4 5 2 Deep Clean 10498 3 27 4 6 2 Deep Clean 10498 3 28 2 4 2 268

Deep Clean 10498 3 29 3 5 2 Deep Clean 10498 3 30 4 6 2 Deep Clean 10498 3 31 3 5 2 Deep Clean 10498 3 32 3 5 2 Deep Clean 10498 3 33 4 6 1 Deep Clean 10498 3 34 2 5 2 Deep Clean 10498 3 35 2 4 2 Deep Clean 10498 3 36 2 5 2 Deep Clean 10498 3 37 5 6 1 Deep Clean 10498 3 38 3 4 2 Deep Clean 10498 3 39 2 3 2 Deep Clean 10498 3 40 3 6 2 Deep Clean 10498 3 41 5 6 2 Deep Clean 10498 3 42 6 6 1 Deep Clean 10498 3 43 5 6 2 Deep Clean 10498 3 44 5 6 2 Deep Clean 10498 3 45 5 6 2 Deep Clean 10498 3 46 4 6 2 Deep Clean 10498 3 47 5 6 2 Deep Clean 10498 4 1 5 6 1 Deep Clean 10498 4 2 4 6 1 Deep Clean 10498 4 3 5 6 1 Deep Clean 10498 4 4 5 6 1 Deep Clean 10498 4 5 5 6 2 Deep Clean 10498 4 6 5 6 2 Deep Clean 10498 4 7 2 5 2 Deep Clean 10498 4 8 1 3 2 Deep Clean 10498 4 9 5 6 1 Deep Clean 10498 4 10 5 6 2 Deep Clean 10498 4 11 5 6 1 Deep Clean 10498 4 12 5 6 2 Deep Clean 10498 4 13 3 6 2 Deep Clean 10498 4 14 3 6 2 Deep Clean 10498 4 15 4 6 1 Deep Clean 10498 4 16 3 5 2 Deep Clean 10498 4 17 2 5 2 Deep Clean 10498 4 18 3 5 2 Deep Clean 10498 4 19 5 6 2 Deep Clean 10498 4 20 2 3 2 Deep Clean 10498 4 21 5 6 2 Deep Clean 10498 4 22 5 6 2 269

Deep Clean 10498 4 23 5 6 2 Deep Clean 10498 4 24 5 6 2 Deep Clean 10498 4 25 5 6 2 Deep Clean 10498 4 26 5 6 2 Deep Clean 10498 4 27 4 6 2 Deep Clean 10498 4 28 3 5 2 Deep Clean 10498 4 29 4 5 1 Deep Clean 10498 4 30 4 6 2 Deep Clean 10498 4 31 5 6 1 Deep Clean 10498 4 32 4 6 2 Deep Clean 10498 4 33 3 6 2 Deep Clean 10498 4 34 4 6 1 Deep Clean 10498 4 35 4 6 2 Deep Clean 10498 4 36 5 6 2 Deep Clean 10498 4 37 4 6 2 Deep Clean 10498 4 38 3 5 2 Deep Clean 10498 4 39 4 6 2 Deep Clean 10498 4 40 3 5 2 Deep Clean 10498 4 41 5 6 2 Deep Clean 10498 4 42 4 6 2 Deep Clean 10498 4 43 3 6 2 Deep Clean 10498 4 44 2 4 2 Deep Clean 10498 4 45 4 6 2 Deep Clean 10498 4 46 5 6 2 Deep Clean 10498 4 47 3 5 2 Deep Clean 10498 4 48 2 4 2 Deep Clean 10498 4 49 3 6 2 Deep Clean 10498 4 50 3 6 2 Deep Clean 10498 4 51 4 6 2 Deep Clean 10498 4 52 3 4 2 Deep Clean 10498 4 53 5 6 2 Deep Clean 10498 4 54 3 5 1 Deep Clean 10498 4 55 3 4 1 Deep Clean 10498 4 56 3 5 2 Deep Clean 10498 4 57 5 6 2 Deep Clean 10498 4 58 5 6 2 Deep Clean 10498 4 59 3 6 2 Deep Clean 10498 4 60 5 6 2 Deep Clean 10498 4 61 4 6 2 Deep Clean 10498 4 62 4 6 1 Deep Clean 10498 4 63 5 6 1 270

Deep Clean 10498 4 64 5 6 2 Deep Clean 10498 4 65 4 6 2 Deep Clean 10498 4 66 5 6 1 Deep Clean 10498 4 67 3 6 1 Deep Clean 10498 4 68 5 6 1 Deep Clean 10498 4 69 3 6 1 Deep Clean 10498 4 70 4 6 1 Deep Clean 10498 4 71 4 6 1 Deep Clean 10498 4 72 4 6 1 Deep Clean 10498 4 73 5 6 1 Deep Clean 10498 5 1 3 6 1 Deep Clean 10498 5 2 3 6 1 Deep Clean 10498 5 3 5 6 1 Deep Clean 10498 5 4 5 6 1 Deep Clean 10498 5 5 5 6 1 Deep Clean 10498 5 6 3 5 2 Deep Clean 10498 5 7 3 4 1 Deep Clean 10498 5 8 3 6 2 Deep Clean 10498 5 9 2 4 2 Deep Clean 10498 5 10 5 6 2 Deep Clean 10498 5 11 5 6 1 Deep Clean 10498 5 12 4 6 1 Deep Clean 10498 5 13 3 6 1 Deep Clean 10498 5 14 5 6 1 Deep Clean 10498 5 15 5 6 1 Deep Clean 10498 5 16 5 6 1 Deep Clean 10498 5 17 3 6 1 Deep Clean 10498 5 18 3 5 2 Deep Clean 10498 5 19 3 5 1 Deep Clean 10498 5 20 3 6 1 Deep Clean 10498 5 21 3 6 1 Deep Clean 10498 5 22 3 6 1 Deep Clean 10498 5 23 2 5 2 Deep Clean 10498 5 24 3 6 1 Deep Clean 10498 5 25 3 6 1 Deep Clean 10498 5 26 2 5 2 Deep Clean 10498 5 27 3 6 1 Deep Clean 10498 5 28 3 5 2 Deep Clean 10498 5 29 3 6 1 Deep Clean 10498 5 30 4 6 1 Deep Clean 10498 5 31 3 6 1 271

Deep Clean 10498 5 32 3 6 1 Deep Clean 10498 5 33 3 6 1 Deep Clean 10498 5 34 2 6 2 Deep Clean 10498 5 35 3 6 1 Deep Clean 10498 5 36 2 4 2 Deep Clean 10498 5 37 4 6 1 Deep Clean 10498 5 38 3 6 1 Deep Clean 10498 5 39 3 6 1 Deep Clean 10498 5 40 4 6 2 Deep Clean 10498 5 41 4 6 1 Deep Clean 10498 5 42 4 6 2 Deep Clean 10498 5 43 4 6 1 Deep Clean 10498 5 44 5 6 1 Deep Clean 10498 5 45 4 6 2 Deep Clean 10498 5 46 4 6 1 Deep Clean 10498 5 47 3 6 1 Deep Clean 10498 5 48 3 6 1 Deep Clean 10498 5 49 5 6 1 Deep Clean 10498 5 50 3 6 1 Deep Clean 10498 5 51 3 6 1 Deep Clean 10498 5 52 3 6 1 Deep Clean 10498 5 53 3 4 1 Deep Clean 10498 5 54 2 4 2 Deep Clean 10498 5 55 2 4 2 Deep Clean 10498 5 56 1 2 2 Deep Clean 10498 5 57 3 3 2 Deep Clean 10498 5 58 3 6 1 Deep Clean 10498 5 59 4 5 1 Deep Clean 10498 5 60 5 6 2 Deep Clean 10498 5 61 3 6 1 Deep Clean 10498 5 62 5 6 1 Deep Clean 10498 5 63 4 5 2 Deep Clean 10498 5 64 3 5 2 Deep Clean 10498 5 64 3 5 2 Deep Clean 10498 5 65 3 5 1 Deep Clean 10498 5 66 3 6 1 Deep Clean 10498 6 1 6 6 1 Deep Clean 10498 6 2 5 6 1 Deep Clean 10498 6 3 4 6 1 Deep Clean 10498 6 4 5 6 2 Deep Clean 10498 6 5 5 6 2 272

Deep Clean 10498 6 6 3 4 2 Deep Clean 10498 6 7 4 6 1 Deep Clean 10498 6 8 5 6 2 Deep Clean 10498 6 9 3 5 2 Deep Clean 10498 6 10 3 5 2 Deep Clean 10498 6 11 5 6 2 Deep Clean 10498 6 12 4 5 1 Deep Clean 10498 6 13 2 2 2 Deep Clean 10498 6 14 4 6 2 Deep Clean 10498 6 15 6 6 2 Deep Clean 10498 6 16 4 6 2 Deep Clean 10498 6 17 5 6 2 Deep Clean 10498 6 18 4 6 2 Deep Clean 10498 6 19 2 5 2 Deep Clean 10498 6 20 4 5 2 Deep Clean 10498 6 21 3 5 2 Deep Clean 10498 6 22 3 5 2 Deep Clean 10498 6 23 4 6 2 Deep Clean 10498 6 24 4 6 2 Deep Clean 10498 6 25 5 6 2 Deep Clean 10498 6 26 4 6 2 Deep Clean 10498 6 27 5 6 2 Deep Clean 10498 6 28 5 6 2 Deep Clean 10498 6 29 5 6 2 Deep Clean 10498 6 30 4 6 2 Deep Clean 10498 6 31 3 6 2 Deep Clean 10498 6 32 5 6 2 Deep Clean 10498 6 33 4 6 2 Deep Clean 10498 6 34 5 6 2 Deep Clean 10498 6 35 5 6 2 Deep Clean 10498 6 36 5 6 2 Deep Clean 10498 6 37 4 6 2 Deep Clean 10498 6 38 5 6 2 Deep Clean 10498 6 39 3 6 2 Deep Clean 10498 6 40 5 6 2 Deep Clean 10498 6 41 5 6 2 Deep Clean 10498 7 1 5 6 1 Deep Clean 10498 7 2 4 6 1 Deep Clean 10498 7 3 3 6 2 Deep Clean 10498 7 4 5 6 1 Deep Clean 10498 7 5 5 6 2 273

Deep Clean 10498 7 6 3 6 2 Deep Clean 10498 7 7 3 5 2 Deep Clean 10498 7 8 5 6 1 Deep Clean 10498 7 9 5 6 1 Deep Clean 10498 7 10 5 6 1 Deep Clean 10498 7 11 6 6 1 Deep Clean 10498 7 12 3 4 2 Deep Clean 10498 7 13 5 6 1 Deep Clean 10498 7 14 6 6 1 Deep Clean 10498 7 15 5 6 1 Deep Clean 10498 7 16 5 6 1 Deep Clean 10498 7 17 5 6 1 Deep Clean 10498 7 18 5 6 2 Deep Clean 10498 7 19 5 6 1 Deep Clean 10498 7 20 5 6 2 Deep Clean 10498 7 21 5 6 2 Deep Clean 10498 7 22 3 6 2 Deep Clean 10498 7 23 5 6 2 Deep Clean 10498 7 24 3 6 2 Deep Clean 10498 7 25 4 6 2 Deep Clean 10498 7 26 3 6 2 Deep Clean 10498 7 27 3 6 1 Deep Clean 10498 7 28 3 4 1 Deep Clean 10498 7 29 3 5 2 Deep Clean 10498 7 30 2 3 2 Deep Clean 10498 7 31 3 4 1 Deep Clean 10498 7 32 5 6 1 Deep Clean 10498 7 33 2 6 2 Deep Clean 10498 7 34 3 6 2 Deep Clean 10498 7 35 3 6 2 Deep Clean 10498 7 36 3 6 2 Deep Clean 10498 7 37 3 6 2 Deep Clean 10498 7 38 5 6 2 Deep Clean 10498 7 39 3 6 1 Deep Clean 10498 7 40 3 4 1 Deep Clean 10498 7 41 5 6 1 Deep Clean 10498 7 42 3 5 2 Deep Clean 10498 7 43 5 6 1 Deep Clean 10498 7 44 6 6 1 Deep Clean 10498 7 45 6 6 1 Deep Clean 10498 7 46 5 6 1 274

Deep Clean 10498 7 47 5 6 1 Deep Clean 10498 7 48 4 6 1 Deep Clean 10498 7 49 2 6 2 Deep Clean 10498 7 50 3 6 2 Deep Clean 10498 7 51 3 6 1 Deep Clean 10498 7 52 3 6 1 Deep Clean 10498 7 53 3 6 1 Deep Clean 10498 7 54 5 6 1 Deep Clean 10498 7 55 5 6 1 Deep Clean 10498 7 56 5 6 2 Deep Clean 10498 8 1 5 6 1 Deep Clean 10498 8 2 3 6 1 Deep Clean 10498 8 3 3 5 1 Deep Clean 10498 8 4 4 6 1 Deep Clean 10498 8 5 2 5 2 Deep Clean 10498 8 6 3 6 1 Deep Clean 10498 8 7 3 6 1 Deep Clean 10498 8 8 5 5 1 Deep Clean 10498 8 9 3 3 2 Deep Clean 10498 8 10 3 6 2 Deep Clean 10498 8 11 4 6 2 Deep Clean 10498 8 12 3 6 1 Deep Clean 10498 8 13 3 4 2 Deep Clean 10498 8 14 3 6 2 Deep Clean 10498 8 15 4 5 1 Deep Clean 10498 8 16 5 6 1 Deep Clean 10498 8 17 4 5 1 Deep Clean 10498 8 18 3 6 2 Deep Clean 10498 8 19 3 6 2 Deep Clean 10498 8 20 4 6 2 Deep Clean 10498 8 21 3 5 1 Deep Clean 10498 8 22 4 6 2 Deep Clean 10498 9 1 5 6 1 Deep Clean 10498 9 2 4 6 1 Deep Clean 10498 9 3 3 4 2 Deep Clean 10498 9 4 4 6 2 Deep Clean 10498 10 1 4 6 2 Deep Clean 10498 10 2 3 5 1 Deep Clean 10498 10 3 4 6 1 Deep Clean 10498 10 4 5 6 2 Deep Rot 10499 1 1 5 6 1 275

Deep Rot 10499 1 2 6 6 2 Deep Rot 10499 1 3 3 6 2 Deep Rot 10499 1 4 3 6 1 Deep Rot 10499 1 5 3 6 1 Deep Rot 10499 1 6 3 6 2 Deep Rot 10499 1 7 3 5 1 Deep Rot 10499 1 8 3 6 2 Deep Rot 10499 1 9 3 6 2 Deep Rot 10499 1 10 3 5 2 Deep Rot 10499 1 11 3 5 2 Deep Rot 10499 1 12 3 6 2 Deep Rot 10499 1 13 4 6 1 Deep Rot 10499 1 14 3 4 2 Deep Rot 10499 1 15 4 6 1 Deep Rot 10499 2 1 3 6 1 Deep Rot 10499 2 2 3 5 1 Deep Rot 10499 2 3 3 6 2 Deep Rot 10499 2 4 3 5 1 Deep Rot 10499 2 5 3 5 1 Deep Rot 10499 2 6 3 5 2 Deep Rot 10499 2 7 3 5 2 Deep Rot 10499 2 8 3 6 2 Deep Rot 10499 2 9 3 5 2 Deep Rot 10499 2 10 3 6 1 Deep Rot 10499 2 11 3 5 1 Deep Rot 10499 2 12 3 5 2 Deep Rot 10499 2 13 4 6 1 Deep Rot 10499 3 1 3 6 1 Deep Rot 10499 3 2 3 4 0 Deep Rot 10499 3 3 3 6 1 Deep Rot 10499 3 4 3 5 1 Deep Rot 10499 3 5 3 6 1 Deep Rot 10499 3 6 3 4 2 Deep Rot 10499 3 7 3 6 2 Deep Rot 10499 3 8 3 5 2 Deep Rot 10499 3 9 3 6 2 Deep Rot 10499 3 10 3 6 1 Deep Rot 10499 3 11 3 4 0 Deep Rot 10499 3 12 3 6 1 Deep Rot 10499 3 13 3 6 1 Deep Rot 10499 3 14 3 5 1 276

Deep Rot 10499 3 15 3 4 0 Deep Rot 10499 3 16 3 6 2 Deep Rot 10499 3 17 3 6 1 Deep Rot 10499 3 18 3 6 2 Deep Rot 10499 3 19 2 5 1 Deep Rot 10499 3 20 2 5 1 Deep Rot 10499 3 21 2 4 2 Deep Rot 10499 4 1 3 6 2 Deep Rot 10499 4 2 2 6 1 Deep Rot 10499 4 3 3 5 2 Deep Rot 10499 4 4 3 5 1 Deep Rot 10499 4 5 3 5 1 Deep Rot 10499 4 6 5 6 2 Deep Rot 10499 4 7 3 6 1 Deep Rot 10499 4 8 3 6 2 Deep Rot 10499 4 9 5 6 2 Deep Rot 10499 4 10 5 6 2 Deep Rot 10499 4 11 5 6 1 Deep Rot 10499 4 12 3 6 1 Deep Rot 10499 4 13 3 5 2 Deep Rot 10499 4 14 3 5 2 Deep Rot 10499 4 15 3 6 1 Deep Rot 10499 4 16 3 6 2 Deep Rot 10499 4 17 3 6 2 Deep Rot 10499 4 18 3 6 2 Deep Rot 10499 4 19 3 6 1 Deep Rot 10499 4 20 3 6 2 Deep Rot 10499 4 21 3 5 2 Deep Rot 10499 4 22 3 5 2 Deep Rot 10499 4 23 3 6 2 Deep Rot 10499 4 24 3 6 1 Deep Rot 10499 4 25 3 6 2 Deep Rot 10499 4 26 3 4 1 Deep Rot 10499 4 27 4 6 1 Deep Rot 10499 4 28 3 5 2 Deep Rot 10499 4 29 1 4 2 Deep Rot 10499 4 30 4 6 1 Deep Rot 10499 4 31 4 6 2 Deep Rot 10499 5 1 6 6 1 Deep Rot 10499 5 2 6 6 2 Deep Rot 10499 5 3 3 6 2 277

Deep Rot 10499 5 4 6 6 2 Deep Rot 10499 5 5 6 6 1 Deep Rot 10499 5 6 4 6 1 Deep Rot 10499 5 7 6 6 1 Deep Rot 10499 5 8 3 6 2 Deep Rot 10499 5 9 3 6 1 Deep Rot 10499 5 10 3 6 1 Deep Rot 10499 5 11 3 5 1 Deep Rot 10499 5 12 4 6 1 Deep Rot 10499 5 13 3 6 1 Deep Rot 10499 5 14 3 6 2 Deep Rot 10499 5 15 3 6 1 Deep Rot 10499 5 16 3 6 2 Deep Rot 10499 5 17 3 6 2 Deep Rot 10499 5 18 3 4 1 Deep Rot 10499 5 19 6 6 2 Deep Rot 10499 5 20 3 6 2 Deep Rot 10499 5 21 6 6 1 Deep Rot 10499 5 22 6 6 2 Deep Rot 10499 5 23 6 6 2 Deep Rot 10499 5 24 3 6 2 Deep Rot 10499 5 25 3 5 2 Deep Rot 10499 5 26 3 6 2 Deep Rot 10499 5 27 3 6 1 Deep Rot 10499 5 28 3 5 2 Deep Rot 10499 5 29 3 5 1 Deep Rot 10499 5 30 3 6 1 Deep Rot 10499 5 31 3 5 2 Deep Rot 10499 5 32 3 6 2 Deep Rot 10499 5 33 3 6 1 Deep Rot 10499 5 34 3 6 2 Deep Rot 10499 5 35 3 6 2 Deep Rot 10499 5 36 3 6 2 Deep Rot 10499 6 1 3 6 2 Deep Rot 10499 6 2 3 6 1 Deep Rot 10499 6 3 2 5 2 Deep Rot 10499 6 4 3 6 1 Deep Rot 10499 6 5 3 6 1 Deep Rot 10499 6 6 5 6 2 Deep Rot 10499 6 7 6 6 2 Deep Rot 10499 6 8 4 6 2 278

Deep Rot 10499 6 9 5 6 2 Deep Rot 10499 6 10 4 6 1 Deep Rot 10499 6 11 3 5 1 Deep Rot 10499 6 12 3 6 2 Deep Rot 10499 6 13 4 6 2 Deep Rot 10499 6 14 5 6 2 Deep Rot 10499 6 15 6 6 2 Deep Rot 10499 6 16 3 5 1 Deep Rot 10499 6 17 2 3 2 Deep Rot 10499 6 18 3 5 1 Deep Rot 10499 6 19 6 6 1 Deep Rot 10499 6 20 4 6 2 Deep Rot 10499 6 21 6 6 2 Deep Rot 10499 6 22 3 5 1 Deep Rot 10499 6 23 3 4 1 Deep Rot 10499 6 24 3 6 2 Deep Rot 10499 6 25 3 5 1 Deep Rot 10499 6 26 3 6 2 Deep Rot 10499 6 27 3 5 1 Deep Rot 10499 6 28 5 6 2 Deep Rot 10499 6 29 6 6 2 Deep Rot 10499 7 1 5 6 2 Deep Rot 10499 7 2 5 6 1 Deep Rot 10499 7 3 3 6 1 Deep Rot 10499 7 4 3 6 2 Deep Rot 10499 7 5 3 6 1 Deep Rot 10499 7 6 6 6 1 Deep Rot 10499 7 7 6 6 1 Deep Rot 10499 7 8 4 6 2 Deep Rot 10499 7 9 3 6 2 Deep Rot 10499 7 10 2 6 2 Deep Rot 10499 7 11 3 5 1 Deep Rot 10499 7 12 4 6 1 Deep Rot 10499 7 13 3 5 2 Deep Rot 10499 7 14 6 6 1 Deep Rot 10499 7 15 4 6 1 Deep Rot 10499 7 16 6 6 1 Deep Rot 10499 7 17 3 6 1 Deep Rot 10499 7 18 5 6 1 Deep Rot 10499 8 1 6 6 1 Deep Rot 10499 8 2 5 6 2 279

Deep Rot 10499 8 3 6 6 1 Deep Rot 10499 8 4 3 5 2 Deep Rot 10499 8 5 5 6 1 Deep Rot 10499 8 6 2 5 2 Deep Rot 10499 8 7 3 5 2 Deep Rot 10499 8 8 5 6 1 Deep Rot 10499 8 9 3 5 2 Deep Rot 10499 8 10 5 6 1 Deep Rot 10499 8 11 2 5 2 Deep Rot 10499 8 12 3 5 2 Deep Rot 10499 8 13 6 6 1 Deep Rot 10499 8 14 3 6 2 Deep Rot 10499 8 15 6 6 1 Deep Rot 10499 8 16 5 6 1 Deep Rot 10499 8 17 5 6 1 Deep Rot 10499 8 18 4 6 1 Deep Rot 10499 8 19 3 6 1 Deep Rot 10499 8 20 6 6 1 Deep Rot 10499 8 21 5 6 1 Deep Rot 10499 9 1 5 6 2 Deep Rot 10499 9 2 5 6 1 Deep Rot 10499 9 3 3 5 2 Deep Rot 10499 9 4 5 6 1 Deep Rot 10499 9 5 3 5 2 Deep Rot 10499 9 6 5 6 1 Deep Rot 10499 9 7 6 6 2 Deep Rot 10499 9 8 5 6 2 Deep Rot 10499 9 9 6 6 2 Deep Rot 10499 9 10 6 6 1 Deep Rot 10500 1 1 6 6 2 Deep Rot 10500 1 2 5 6 2 Deep Rot 10500 1 3 6 6 2 Deep Rot 10500 1 4 3 5 2 Deep Rot 10500 1 5 3 5 2 Deep Rot 10500 1 6 2 3 2 Deep Rot 10500 1 7 3 5 2 Deep Rot 10500 1 8 6 6 2 Deep Rot 10500 1 9 3 5 2 Deep Rot 10500 1 10 6 6 2 Deep Rot 10500 1 11 5 6 2 Deep Rot 10500 2 1 5 6 2 280

Deep Rot 10500 2 2 5 6 2 Deep Rot 10500 2 3 6 6 2 Deep Rot 10500 2 4 5 6 2 Deep Rot 10500 2 5 6 6 2 Deep Rot 10500 2 6 6 6 2 Deep Rot 10500 2 7 3 4 2 Deep Rot 10500 2 8 3 6 2 Deep Rot 10500 2 9 3 4 2 Deep Rot 10500 2 10 3 6 2 Deep Rot 10500 2 11 6 6 1 Deep Rot 10500 2 12 5 6 2 Deep Rot 10500 2 13 4 6 2 Deep Rot 10500 2 14 6 6 2 Deep Rot 10500 2 15 3 6 2 Deep Rot 10500 2 16 3 5 2 Deep Rot 10500 2 17 4 6 2 Deep Rot 10500 2 18 3 5 2 Deep Rot 10500 2 19 3 6 2 Deep Rot 10500 2 20 3 5 2 Deep Rot 10500 2 21 3 6 2 Deep Rot 10500 2 22 5 6 2 Deep Rot 10500 3 1 6 6 2 Deep Rot 10500 3 2 6 6 2 Deep Rot 10500 3 3 5 6 2 Deep Rot 10500 3 4 6 6 2 Deep Rot 10500 3 5 5 6 2 Deep Rot 10500 3 6 4 6 2 Deep Rot 10500 3 7 5 6 2 Deep Rot 10500 3 8 5 5 2 Deep Rot 10500 3 9 6 6 2 Deep Rot 10500 3 10 5 5 2 Deep Rot 10500 3 11 3 5 2 Deep Rot 10500 3 12 3 6 2 Deep Rot 10500 3 13 5 6 2 Deep Rot 10500 3 14 4 6 2 Deep Rot 10500 3 15 3 6 2 Deep Rot 10500 3 16 3 5 2 Deep Rot 10500 3 17 6 6 2 Deep Rot 10500 3 18 6 6 2 Deep Rot 10500 3 19 5 6 2 Deep Rot 10500 3 20 5 6 2 281

Deep Rot 10500 3 21 5 6 2 Deep Rot 10500 3 22 5 6 2 Deep Rot 10500 3 23 5 6 2 Deep Rot 10500 3 24 3 5 2 Deep Rot 10500 3 25 3 6 2 Deep Rot 10500 3 26 5 6 2 Deep Rot 10500 3 27 2 3 1 Deep Rot 10500 3 28 3 6 2 Deep Rot 10500 4 1 4 6 2 Deep Rot 10500 4 2 6 6 2 Deep Rot 10500 4 3 5 6 2 Deep Rot 10500 4 4 6 6 1 Deep Rot 10500 4 5 3 6 2 Deep Rot 10500 4 6 6 6 2 Deep Rot 10500 4 7 5 6 2 Deep Rot 10500 4 8 3 6 2 Deep Rot 10500 4 9 2 4 2 Deep Rot 10500 4 10 3 5 1 Deep Rot 10500 4 11 4 6 2 Deep Rot 10500 4 12 3 6 1 Deep Rot 10500 4 13 6 6 2 Deep Rot 10500 4 14 3 6 2 Deep Rot 10500 4 15 5 6 2 Deep Rot 10500 4 16 4 6 1 Deep Rot 10500 4 17 5 6 1 Deep Rot 10500 4 18 6 6 1 Deep Rot 10500 4 19 5 6 2 Deep Rot 10500 4 20 3 6 2 Deep Rot 10500 4 21 5 6 2 Deep Rot 10500 4 22 6 6 2 Deep Rot 10500 4 23 4 6 2 Deep Rot 10500 5 1 3 6 2 Deep Rot 10500 5 2 6 6 2 Deep Rot 10500 5 3 5 6 2 Deep Rot 10500 5 4 6 6 2 Deep Rot 10500 5 5 3 6 2 Deep Rot 10500 5 6 4 6 2 Deep Rot 10500 5 7 3 4 2 Deep Rot 10500 5 8 3 6 2 Deep Rot 10500 5 9 3 4 2 Deep Rot 10500 5 10 5 6 2 282

Deep Rot 10500 5 11 4 6 2 Deep Rot 10500 5 12 6 6 2 Deep Rot 10500 5 13 3 5 2 Deep Rot 10500 5 14 5 6 2 Deep Rot 10500 5 15 6 6 1 Deep Rot 10500 5 16 5 6 1 Deep Rot 10500 5 17 5 6 1 Deep Rot 10500 5 18 3 6 2 Deep Rot 10500 5 19 3 6 2 Deep Rot 10500 5 20 4 6 2 Deep Rot 10500 5 21 5 6 2 Deep Rot 10500 5 22 6 6 2 Deep Rot 10500 5 23 3 4 2 Deep Rot 10500 5 24 5 6 2 Deep Rot 10500 5 25 5 6 2 Deep Rot 10500 5 26 6 6 2 Deep Rot 10500 5 27 6 6 2 Deep Rot 10500 5 28 6 6 2 Deep Rot 10500 5 29 6 6 2 Deep Rot 10500 5 30 3 6 2 Deep Rot 10500 5 31 3 5 2 Deep Rot 10500 5 32 5 6 2 Deep Rot 10500 5 33 6 6 2 Deep Rot 10500 6 1 5 6 2 Deep Rot 10500 6 2 6 6 2 Deep Rot 10500 6 3 6 6 2 Deep Rot 10500 6 4 5 6 2 Deep Rot 10500 6 5 5 6 2 Deep Rot 10500 6 6 5 6 1 Deep Rot 10500 6 7 4 5 2 Deep Rot 10500 6 8 3 5 2 Deep Rot 10500 6 9 5 6 2 Deep Rot 10500 6 10 5 6 2 Deep Rot 10500 6 11 5 6 2 Deep Rot 10500 6 12 5 6 2 Deep Rot 10500 6 13 4 6 2 Deep Rot 10500 6 14 5 6 2 Deep Rot 10500 6 15 5 6 2 Deep Rot 10500 6 16 3 6 2 Deep Rot 10500 6 17 6 6 2 Deep Rot 10500 6 18 5 6 2 283

Deep Rot 10500 6 19 5 6 2 Deep Rot 10500 6 20 5 6 2 Deep Rot 10500 6 21 5 6 1 Deep Rot 10500 6 22 6 6 2 Deep Rot 10500 6 23 1 4 2 Deep Rot 10500 7 1 5 6 2 Deep Rot 10500 7 2 2 3 2 Deep Rot 10500 7 3 4 6 2 Deep Rot 10500 7 4 5 6 2 Deep Rot 10500 7 5 2 6 2 Deep Rot 10500 7 6 3 6 2 Deep Rot 10500 7 7 2 6 2 Deep Rot 10500 7 8 5 6 1 Deep Rot 10500 7 9 2 3 2 Deep Rot 10500 7 10 3 6 1 Deep Rot 10500 7 11 4 6 1 Deep Rot 10500 7 12 3 4 1 Deep Rot 10500 7 13 2 4 2 Deep Rot 10500 7 14 3 4 2 Deep Rot 10500 7 15 5 6 2 Deep Rot 10500 7 16 4 6 1 Deep Rot 10500 8 1 5 6 2 Deep Rot 10500 8 2 5 6 2 Deep Rot 10500 8 3 6 6 1 Deep Rot 10500 8 4 5 6 1 Deep Rot 10500 8 5 6 6 1 Deep Rot 10500 8 6 5 6 1 Deep Rot 10500 8 7 5 6 1 Deep Rot 10500 8 8 6 6 2 Deep Rot 10500 8 9 3 6 1 Deep Rot 10500 8 10 2 5 1 Deep Rot 10500 8 11 2 3 1 Deep Rot 10500 8 12 3 6 1 Deep Rot 10500 8 13 3 6 1 Deep Rot 10500 8 14 5 6 2 Deep Rot 10500 8 15 5 6 2 Deep Rot 10500 8 16 6 6 1 Deep Rot 10500 9 1 5 6 2 Deep Rot 10500 9 2 6 6 2 Deep Rot 10500 9 3 2 6 2 Deep Rot 10500 9 4 4 6 2 284

Deep Rot 10500 9 5 5 6 2 Deep Rot 10500 9 6 6 6 1 Deep Rot 10500 9 7 6 6 1 Deep Rot 10500 9 8 5 6 1 Deep Rot 10500 9 9 5 6 2 Shallow Rot 10501 1 1 1 3 2 Shallow Rot 10501 1 2 1 2 2 Shallow Rot 10501 1 3 3 5 2 Shallow Rot 10501 1 4 5 6 2 Shallow Rot 10501 1 5 5 6 2 Shallow Rot 10501 1 6 6 6 2 Shallow Rot 10501 1 7 4 6 2 Shallow Rot 10501 1 8 2 4 2 Shallow Rot 10501 1 9 3 5 2 Shallow Rot 10501 1 10 3 5 1 Shallow Rot 10501 2 1 1 2 2 Shallow Rot 10501 2 2 2 3 2 Shallow Rot 10501 2 3 3 4 2 Shallow Rot 10501 2 4 1 2 2 Shallow Rot 10501 2 5 1 2 2 Shallow Rot 10501 2 6 5 6 2 Shallow Rot 10501 2 7 5 6 2 Shallow Rot 10501 2 8 4 6 2 Shallow Rot 10501 2 9 2 5 2 Shallow Rot 10501 2 10 4 6 2 Shallow Rot 10501 2 11 5 6 2 Shallow Rot 10501 2 12 5 5 2 Shallow Rot 10501 2 13 5 6 1 Shallow Rot 10501 2 14 4 6 1 Shallow Rot 10501 2 15 3 5 1 Shallow Rot 10501 2 16 5 6 1 Shallow Rot 10501 2 17 3 5 1 Shallow Rot 10501 2 18 5 6 1 Shallow Rot 10501 2 19 4 5 1 Shallow Rot 10501 2 20 5 6 1 Shallow Rot 10501 2 21 3 5 1 Shallow Rot 10501 2 22 5 6 1 Shallow Rot 10501 2 23 3 5 1 Shallow Rot 10501 2 24 6 6 2 Shallow Rot 10501 3 1 1 2 2 Shallow Rot 10501 3 2 1 3 2 285

Shallow Rot 10501 3 3 3 3 2 Shallow Rot 10501 3 4 1 2 2 Shallow Rot 10501 3 5 3 4 2 Shallow Rot 10501 3 6 2 3 2 Shallow Rot 10501 3 7 2 4 2 Shallow Rot 10501 3 8 5 6 2 Shallow Rot 10501 3 9 3 5 2 Shallow Rot 10501 3 10 2 4 2 Shallow Rot 10501 3 11 2 3 2 Shallow Rot 10501 3 12 1 2 2 Shallow Rot 10501 3 13 2 3 2 Shallow Rot 10501 3 14 1 3 2 Shallow Rot 10501 3 15 2 3 2 Shallow Rot 10501 3 16 5 6 2 Shallow Rot 10501 3 17 5 6 2 Shallow Rot 10501 3 18 5 6 2 Shallow Rot 10501 3 19 6 6 1 Shallow Rot 10501 3 20 4 6 1 Shallow Rot 10501 3 21 5 6 2 Shallow Rot 10501 3 22 6 6 2 Shallow Rot 10501 3 23 5 6 2 Shallow Rot 10501 3 24 6 6 2 Shallow Rot 10501 3 25 2 3 2 Shallow Rot 10501 3 26 2 5 2 Shallow Rot 10501 3 27 2 3 2 Shallow Rot 10501 3 28 3 4 2 Shallow Rot 10501 3 29 3 5 2 Shallow Rot 10501 3 30 5 6 2 Shallow Rot 10501 3 31 5 6 2 Shallow Rot 10501 3 32 1 2 2 Shallow Rot 10501 3 33 1 4 2 Shallow Rot 10501 3 34 2 4 2 Shallow Rot 10501 3 35 1 2 2 Shallow Rot 10501 4 1 1 2 2 Shallow Rot 10501 4 2 1 3 2 Shallow Rot 10501 4 3 2 3 2 Shallow Rot 10501 4 4 2 3 2 Shallow Rot 10501 4 5 3 3 2 Shallow Rot 10501 4 6 2 3 2 Shallow Rot 10501 4 7 1 3 2 Shallow Rot 10501 4 8 1 3 2 286

Shallow Rot 10501 4 9 1 3 2 Shallow Rot 10501 4 10 6 6 2 Shallow Rot 10501 4 11 5 6 1 Shallow Rot 10501 4 12 5 6 1 Shallow Rot 10501 4 13 5 6 1 Shallow Rot 10501 4 14 4 5 1 Shallow Rot 10501 4 15 1 2 2 Shallow Rot 10501 4 16 1 2 2 Shallow Rot 10501 4 17 5 6 1 Shallow Rot 10501 4 18 4 5 1 Shallow Rot 10501 4 19 4 6 1 Shallow Rot 10501 4 20 5 6 1 Shallow Rot 10501 4 21 5 6 2 Shallow Rot 10501 4 22 2 3 2 Shallow Rot 10501 4 23 2 3 2 Shallow Rot 10501 4 24 2 3 2 Shallow Rot 10501 4 25 1 3 2 Shallow Rot 10501 4 26 1 3 2 Shallow Rot 10501 4 27 3 3 2 Shallow Rot 10501 4 28 3 3 2 Shallow Rot 10501 4 29 1 2 2 Shallow Rot 10501 4 30 1 2 2 Shallow Rot 10501 4 31 2 3 2 Shallow Rot 10501 4 32 3 4 2 Shallow Rot 10501 5 1 1 2 2 Shallow Rot 10501 5 2 1 3 2 Shallow Rot 10501 5 3 2 3 2 Shallow Rot 10501 5 4 2 3 2 Shallow Rot 10501 5 5 1 2 2 Shallow Rot 10501 5 6 1 3 2 Shallow Rot 10501 5 7 1 2 2 Shallow Rot 10501 5 8 1 3 2 Shallow Rot 10501 5 9 1 3 2 Shallow Rot 10501 5 10 6 6 2 Shallow Rot 10501 5 11 5 6 2 Shallow Rot 10501 5 12 5 6 1 Shallow Rot 10501 5 13 5 6 2 Shallow Rot 10501 5 14 5 6 1 Shallow Rot 10501 5 15 6 6 1 Shallow Rot 10501 5 16 3 4 1 Shallow Rot 10501 5 17 3 4 2 287

Shallow Rot 10501 5 18 3 4 2 Shallow Rot 10501 5 19 4 6 2 Shallow Rot 10501 5 20 5 6 1 Shallow Rot 10501 5 21 3 5 1 Shallow Rot 10501 5 22 5 6 1 Shallow Rot 10501 5 23 4 6 1 Shallow Rot 10501 5 24 6 6 2 Shallow Rot 10501 5 25 5 6 1 Shallow Rot 10501 5 26 6 6 1 Shallow Rot 10501 5 27 6 6 1 Shallow Rot 10501 5 28 1 2 2 Shallow Rot 10501 5 29 3 4 1 Shallow Rot 10501 5 30 3 6 1 Shallow Rot 10501 5 31 5 6 1 Shallow Rot 10501 5 32 6 6 1 Shallow Rot 10501 5 33 3 5 1 Shallow Rot 10501 5 34 6 6 1 Shallow Rot 10501 5 35 5 6 1 Shallow Rot 10501 5 36 5 6 1 Shallow Rot 10501 5 37 3 5 1 Shallow Rot 10501 5 38 3 6 1 Shallow Rot 10501 5 39 3 6 1 Shallow Rot 10501 5 40 5 6 1 Shallow Rot 10501 5 41 5 6 2 Shallow Rot 10501 5 42 6 6 1 Shallow Rot 10501 5 43 5 6 1 Shallow Rot 10501 5 44 5 6 2 Shallow Rot 10501 5 45 4 5 2 Shallow Rot 10501 5 46 1 3 2 Shallow Rot 10501 5 47 1 3 2 Shallow Rot 10501 5 48 1 2 2 Shallow Rot 10501 5 49 1 2 2 Shallow Rot 10501 5 50 1 3 2 Shallow Rot 10501 5 51 1 3 2 Shallow Rot 10501 5 52 1 3 2 Shallow Rot 10501 5 53 1 2 2 Shallow Rot 10501 5 54 1 2 2 Shallow Rot 10501 6 1 2 4 2 Shallow Rot 10501 6 2 5 6 2 Shallow Rot 10501 6 3 2 3 2 Shallow Rot 10501 6 4 2 2 2 288

Shallow Rot 10501 6 5 1 2 2 Shallow Rot 10501 6 6 2 6 2 Shallow Rot 10501 6 7 1 2 2 Shallow Rot 10501 6 8 2 3 2 Shallow Rot 10501 6 9 3 4 2 Shallow Rot 10501 6 10 1 2 2 Shallow Rot 10501 6 11 3 4 2 Shallow Rot 10501 6 12 2 3 2 Shallow Rot 10501 6 13 3 4 2 Shallow Rot 10501 6 14 4 6 2 Shallow Rot 10501 6 15 3 4 2 Shallow Rot 10501 6 16 1 2 2 Shallow Rot 10501 6 17 2 2 2 Shallow Rot 10501 6 18 5 6 2 Shallow Rot 10501 6 19 4 6 2 Shallow Rot 10501 6 20 5 6 2 Shallow Rot 10501 6 21 4 6 2 Shallow Rot 10501 6 22 5 6 2 Shallow Rot 10501 6 23 5 6 2 Shallow Rot 10501 6 24 4 6 1 Shallow Rot 10501 6 25 3 5 1 Shallow Rot 10501 6 26 1 3 2 Shallow Rot 10501 6 27 5 6 2 Shallow Rot 10501 6 28 1 2 2 Shallow Rot 10501 6 29 5 6 2 Shallow Rot 10501 6 30 4 6 2 Shallow Rot 10501 6 31 5 6 2 Shallow Rot 10501 6 32 1 3 2 Shallow Rot 10501 6 33 1 3 2 Shallow Rot 10501 6 34 3 4 2 Shallow Rot 10501 6 35 6 6 2 Shallow Rot 10501 6 36 5 6 2 Shallow Rot 10501 6 37 4 5 2 Shallow Rot 10501 6 38 3 5 2 Shallow Rot 10501 6 39 3 6 2 Shallow Rot 10501 6 40 4 6 1 Shallow Rot 10501 6 41 3 5 2 Shallow Rot 10501 6 42 3 5 1 Shallow Rot 10501 6 43 5 6 1 Shallow Rot 10501 6 44 4 6 2 Shallow Rot 10501 6 45 5 6 1 289

Shallow Rot 10501 6 46 5 6 2 Shallow Rot 10501 6 47 6 6 1 Shallow Rot 10501 6 48 5 6 2 Shallow Rot 10501 6 49 4 6 2 Shallow Rot 10501 6 50 5 6 2 Shallow Rot 10501 6 51 6 6 1 Shallow Rot 10501 6 52 5 6 2 Shallow Rot 10501 6 53 6 6 2 Shallow Rot 10501 6 54 3 4 2 Shallow Rot 10501 6 55 3 5 2 Shallow Rot 10501 6 56 2 3 2 Shallow Rot 10501 6 57 5 6 2 Shallow Rot 10501 6 58 2 2 2 Shallow Rot 10501 6 59 2 3 2 Shallow Rot 10501 6 60 2 2 2 Shallow Rot 10501 6 61 1 2 2 Shallow Rot 10501 6 62 1 2 2 Shallow Rot 10501 6 63 1 2 2 Shallow Rot 10501 6 64 1 2 2 Shallow Rot 10501 6 65 1 2 2 Shallow Rot 10501 6 66 1 2 2 Shallow Rot 10501 6 67 2 4 2 Shallow Rot 10501 6 68 1 2 2 Shallow Rot 10501 6 69 1 2 2 Shallow Rot 10501 6 70 1 2 2 Shallow Rot 10501 6 71 1 2 2 Shallow Rot 10501 6 72 1 2 2 Shallow Rot 10501 6 73 1 2 2 Shallow Rot 10501 6 74 2 3 2 Shallow Rot 10501 6 75 1 3 2 Shallow Rot 10501 6 76 2 3 2 Shallow Rot 10501 6 77 2 5 2 Shallow Rot 10501 7 1 1 3 2 Shallow Rot 10501 7 2 2 3 2 Shallow Rot 10501 7 3 1 3 2 Shallow Rot 10501 7 4 2 3 2 Shallow Rot 10501 7 5 5 6 2 Shallow Rot 10501 7 6 2 3 2 Shallow Rot 10501 7 7 1 2 2 Shallow Rot 10501 7 8 2 2 2 Shallow Rot 10501 7 9 5 6 2 290

Shallow Rot 10501 7 10 5 6 2 Shallow Rot 10501 7 11 5 6 1 Shallow Rot 10501 7 12 5 6 1 Shallow Rot 10501 7 13 3 5 1 Shallow Rot 10501 7 14 2 4 2 Shallow Rot 10501 7 15 3 5 2 Shallow Rot 10501 7 16 1 3 2 Shallow Rot 10501 7 17 2 5 2 Shallow Rot 10501 7 18 3 5 1 Shallow Rot 10501 7 19 1 4 2 Shallow Rot 10501 7 20 3 5 1 Shallow Rot 10501 7 21 4 6 1 Shallow Rot 10501 7 22 4 6 1 Shallow Rot 10501 7 23 4 6 1 Shallow Rot 10501 7 24 3 5 2 Shallow Rot 10501 7 25 5 6 2 Shallow Rot 10501 7 26 4 5 2 Shallow Rot 10501 7 27 3 4 2 Shallow Rot 10501 7 28 5 6 2 Shallow Rot 10501 7 29 6 6 2 Shallow Rot 10501 7 30 4 5 2 Shallow Rot 10501 7 31 5 6 2 Shallow Rot 10501 7 32 6 6 2 Shallow Rot 10501 7 33 1 2 2 Shallow Rot 10501 7 34 5 6 2 Shallow Rot 10501 7 35 5 6 2 Shallow Rot 10501 7 36 5 6 2 Shallow Rot 10501 7 37 5 6 1 Shallow Rot 10501 7 38 6 6 1 Shallow Rot 10501 7 39 1 2 2 Shallow Rot 10501 7 40 1 3 2 Shallow Rot 10501 7 41 1 3 2 Shallow Rot 10501 7 42 1 2 2 Shallow Rot 10501 7 43 3 4 2 Shallow Rot 10501 7 44 2 3 2 Shallow Rot 10501 7 45 1 2 2 Shallow Rot 10501 7 46 1 2 2 Shallow Rot 10501 7 47 2 2 2 Shallow Rot 10501 7 48 2 2 2 Shallow Rot 10501 8 1 4 6 2 Shallow Rot 10501 8 2 2 3 2 291

Shallow Rot 10501 8 3 1 2 2 Shallow Rot 10501 8 4 3 3 2 Shallow Rot 10501 8 5 2 3 2 Shallow Rot 10501 8 6 3 4 2 Shallow Rot 10501 8 7 2 4 2 Shallow Rot 10501 8 8 3 5 2 Shallow Rot 10501 8 9 3 5 2 Shallow Rot 10501 8 10 5 6 1 Shallow Rot 10501 8 11 2 5 2 Shallow Rot 10501 8 12 1 4 2 Shallow Rot 10501 8 13 3 4 1 Shallow Rot 10501 8 14 3 4 1 Shallow Rot 10501 8 15 5 6 1 Shallow Rot 10501 8 16 4 6 1 Shallow Rot 10501 8 17 3 4 1 Shallow Rot 10501 8 18 4 6 1 Shallow Rot 10501 8 19 5 6 1 Shallow Rot 10501 8 20 3 4 2 Shallow Rot 10501 8 21 5 6 2 Shallow Rot 10501 8 22 5 6 2 Shallow Rot 10501 8 23 4 6 1 Shallow Rot 10501 8 24 3 4 2 Shallow Rot 10501 8 25 4 5 2 Shallow Rot 10501 8 26 3 4 2 Shallow Rot 10501 8 27 3 5 2 Shallow Rot 10501 8 28 3 6 2 Shallow Rot 10501 8 29 4 6 2 Shallow Rot 10501 8 30 4 5 2 Shallow Rot 10501 8 31 5 6 2 Shallow Rot 10501 8 32 3 6 2 Shallow Rot 10501 8 33 3 5 2 Shallow Rot 10501 8 34 3 5 2 Shallow Rot 10501 8 35 2 3 2 Shallow Rot 10501 8 36 2 3 2 Shallow Rot 10501 8 37 1 2 2 Shallow Rot 10501 8 38 1 2 2 Shallow Rot 10501 8 39 1 2 2 Shallow Rot 10501 8 40 1 2 2 Shallow Rot 10501 8 41 1 2 2 Shallow Rot 10501 8 42 4 6 2 Shallow Rot 10501 8 43 2 3 2 292

Shallow Rot 10501 9 1 5 6 2 Shallow Rot 10501 9 2 3 4 2 Shallow Rot 10501 9 3 2 3 2 Shallow Rot 10501 9 4 1 2 2 Shallow Rot 10501 9 5 2 3 2 Shallow Rot 10501 9 6 1 3 2 Shallow Rot 10501 9 7 3 3 2 Shallow Rot 10501 9 8 5 6 1 Shallow Rot 10501 9 9 2 3 2 Shallow Rot 10501 9 10 na na na Shallow Rot 10501 9 11 4 5 1 Shallow Rot 10501 9 12 3 4 1 Shallow Rot 10501 9 13 4 6 1 Shallow Rot 10501 9 14 3 5 2 Shallow Rot 10501 9 15 5 6 1 Shallow Rot 10501 9 16 3 4 0 Shallow Rot 10501 9 17 3 5 1 Shallow Rot 10501 9 18 4 5 1 Shallow Rot 10501 9 19 5 6 1 Shallow Rot 10501 9 20 5 6 1 Shallow Rot 10501 9 21 3 5 2 Shallow Rot 10501 9 22 3 4 1 Shallow Rot 10501 9 23 4 5 2 Shallow Rot 10501 9 24 5 6 1 Shallow Rot 10501 9 25 2 3 2 Shallow Rot 10501 9 26 3 3 2 Shallow Rot 10501 9 27 1 2 2 Shallow Rot 10501 9 28 2 3 1 Shallow Rot 10501 9 29 1 2 2 Shallow Rot 10501 9 30 3 4 2 Shallow Rot 10501 9 31 1 3 2 Shallow Rot 10501 9 32 3 4 2 Shallow Rot 10501 9 33 4 6 2 Shallow Rot 10501 9 34 3 4 1 Shallow Rot 10501 9 35 2 3 2 Shallow Rot 10501 9 36 4 6 2 Shallow Rot 10501 9 37 3 6 1 Shallow Rot 10502 1 1 6 6 2 Shallow Rot 10502 1 2 3 4 2 Shallow Rot 10502 1 3 4 5 2 Shallow Rot 10502 1 4 5 6 2 293

Shallow Rot 10502 1 5 2 3 2 Shallow Rot 10502 1 6 3 3 2 Shallow Rot 10502 1 7 5 6 2 Shallow Rot 10502 1 8 3 4 2 Shallow Rot 10502 1 9 3 3 2 Shallow Rot 10502 1 10 1 3 2 Shallow Rot 10502 1 11 2 3 2 Shallow Rot 10502 1 12 1 3 2 Shallow Rot 10502 1 13 2 3 2 Shallow Rot 10502 1 14 5 6 2 Shallow Rot 10502 1 15 2 3 2 Shallow Rot 10502 1 16 1 3 2 Shallow Rot 10502 1 17 2 3 2 Shallow Rot 10502 1 18 6 6 2 Shallow Rot 10502 1 19 5 6 2 Shallow Rot 10502 1 20 5 6 2 Shallow Rot 10502 1 21 6 6 2 Shallow Rot 10502 1 22 4 5 2 Shallow Rot 10502 1 23 3 5 2 Shallow Rot 10502 1 24 6 6 2 Shallow Rot 10502 1 25 5 6 2 Shallow Rot 10502 1 26 6 6 2 Shallow Rot 10502 1 27 6 6 2 Shallow Rot 10502 1 28 4 6 2 Shallow Rot 10502 1 29 5 6 2 Shallow Rot 10502 2 1 6 6 2 Shallow Rot 10502 2 2 4 6 2 Shallow Rot 10502 2 3 4 6 2 Shallow Rot 10502 2 4 5 6 2 Shallow Rot 10502 2 5 6 6 2 Shallow Rot 10502 2 6 5 6 2 Shallow Rot 10502 2 7 5 6 2 Shallow Rot 10502 2 8 3 5 2 Shallow Rot 10502 2 9 3 4 2 Shallow Rot 10502 2 10 5 6 2 Shallow Rot 10502 2 11 4 6 2 Shallow Rot 10502 2 12 3 6 2 Shallow Rot 10502 2 13 2 6 2 Shallow Rot 10502 2 14 2 3 2 Shallow Rot 10502 2 15 3 4 2 Shallow Rot 10502 2 16 3 4 2 294

Shallow Rot 10502 2 17 5 6 2 Shallow Rot 10502 2 18 3 4 2 Shallow Rot 10502 2 19 2 3 2 Shallow Rot 10502 2 20 6 6 2 Shallow Rot 10502 2 21 2 3 2 Shallow Rot 10502 2 22 2 5 2 Shallow Rot 10502 2 23 2 3 2 Shallow Rot 10502 2 24 2 3 2 Shallow Rot 10502 2 25 1 2 2 Shallow Rot 10502 2 26 2 5 2 Shallow Rot 10502 2 27 2 3 2 Shallow Rot 10502 2 28 2 6 2 Shallow Rot 10502 2 29 3 5 2 Shallow Rot 10502 2 30 2 6 2 Shallow Rot 10502 2 31 3 5 2 Shallow Rot 10502 2 32 2 4 2 Shallow Rot 10502 2 33 2 5 2 Shallow Rot 10502 2 34 3 5 2 Shallow Rot 10502 2 35 4 6 2 Shallow Rot 10502 2 36 5 6 2 Shallow Rot 10502 2 37 4 5 2 Shallow Rot 10502 2 38 5 6 2 Shallow Rot 10502 2 39 3 5 2 Shallow Rot 10502 2 40 5 6 2 Shallow Rot 10502 2 41 5 6 2 Shallow Rot 10502 2 42 5 6 2 Shallow Rot 10502 2 43 4 6 2 Shallow Rot 10502 2 44 5 6 2 Shallow Rot 10502 2 45 4 5 2 Shallow Rot 10502 2 46 4 5 2 Shallow Rot 10502 2 47 6 6 2 Shallow Rot 10502 2 48 5 6 2 Shallow Rot 10502 2 49 3 5 2 Shallow Rot 10502 2 50 4 5 2 Shallow Rot 10502 2 51 6 6 2 Shallow Rot 10502 2 52 5 5 2 Shallow Rot 10502 2 53 3 5 2 Shallow Rot 10502 2 54 3 6 2 Shallow Rot 10502 2 55 3 5 2 Shallow Rot 10502 2 56 3 5 2 Shallow Rot 10502 2 57 4 5 2 295

Shallow Rot 10502 2 58 3 4 2 Shallow Rot 10502 2 59 3 5 2 Shallow Rot 10502 2 60 3 4 2 Shallow Rot 10502 2 61 2 3 2 Shallow Rot 10502 2 62 2 3 2 Shallow Rot 10502 2 63 3 4 2 Shallow Rot 10502 2 64 2 3 2 Shallow Rot 10502 2 65 3 4 2 Shallow Rot 10502 2 66 3 4 2 Shallow Rot 10502 2 67 1 2 2 Shallow Rot 10502 2 68 3 4 2 Shallow Rot 10502 2 69 6 6 2 Shallow Rot 10502 2 70 2 3 2 Shallow Rot 10502 2 71 4 6 2 Shallow Rot 10502 2 72 3 5 2 Shallow Rot 10502 2 73 2 3 2 Shallow Rot 10502 2 74 3 5 2 Shallow Rot 10502 2 75 2 3 2 Shallow Rot 10502 2 76 1 3 2 Shallow Rot 10502 2 77 2 3 2 Shallow Rot 10502 2 78 3 4 2 Shallow Rot 10502 2 79 4 6 2 Shallow Rot 10502 2 80 5 6 2 Shallow Rot 10502 2 81 2 3 2 Shallow Rot 10502 2 82 3 5 2 Shallow Rot 10502 2 83 2 3 2 Shallow Rot 10502 2 84 2 3 2 Shallow Rot 10502 2 85 5 6 2 Shallow Rot 10502 2 86 4 6 2 Shallow Rot 10502 3 1 5 6 1 Shallow Rot 10502 3 2 5 6 1 Shallow Rot 10502 3 3 5 6 2 Shallow Rot 10502 3 4 3 4 2 Shallow Rot 10502 3 5 4 5 2 Shallow Rot 10502 3 6 2 3 2 Shallow Rot 10502 3 7 5 6 2 Shallow Rot 10502 3 8 3 4 2 Shallow Rot 10502 3 9 5 5 2 Shallow Rot 10502 3 10 3 5 2 Shallow Rot 10502 3 11 4 5 2 Shallow Rot 10502 3 12 5 5 2 296

Shallow Rot 10502 3 13 6 6 2 Shallow Rot 10502 3 14 4 6 2 Shallow Rot 10502 3 15 6 6 2 Shallow Rot 10502 3 16 6 6 2 Shallow Rot 10502 3 17 3 4 2 Shallow Rot 10502 3 18 4 5 2 Shallow Rot 10502 3 19 3 4 2 Shallow Rot 10502 3 20 3 4 2 Shallow Rot 10502 3 21 3 3 2 Shallow Rot 10502 3 22 1 3 2 Shallow Rot 10502 3 23 3 4 2 Shallow Rot 10502 3 24 4 5 2 Shallow Rot 10502 3 25 3 4 2 Shallow Rot 10502 3 26 2 3 2 Shallow Rot 10502 3 27 2 3 2 Shallow Rot 10502 3 28 3 4 2 Shallow Rot 10502 3 29 3 4 2 Shallow Rot 10502 3 30 2 3 2 Shallow Rot 10502 3 31 2 3 2 Shallow Rot 10502 3 32 2 3 2 Shallow Rot 10502 3 33 1 3 2 Shallow Rot 10502 3 34 5 6 2 Shallow Rot 10502 3 35 5 6 2 Shallow Rot 10502 3 36 5 6 2 Shallow Rot 10502 3 37 3 4 2 Shallow Rot 10502 3 38 3 4 2 Shallow Rot 10502 3 39 4 6 2 Shallow Rot 10502 3 40 5 6 2 Shallow Rot 10502 3 41 6 6 2 Shallow Rot 10502 3 42 3 4 2 Shallow Rot 10502 3 43 2 3 2 Shallow Rot 10502 3 44 3 4 2 Shallow Rot 10502 3 45 2 3 2 Shallow Rot 10502 3 46 3 5 2 Shallow Rot 10502 3 47 2 3 2 Shallow Rot 10502 3 48 3 4 2 Shallow Rot 10502 3 49 5 6 2 Shallow Rot 10502 3 50 4 5 2 Shallow Rot 10502 3 51 4 5 2 Shallow Rot 10502 3 52 5 6 2 Shallow Rot 10502 3 53 6 6 2 297

Shallow Rot 10502 3 54 3 4 2 Shallow Rot 10502 3 55 4 5 2 Shallow Rot 10502 3 56 3 4 2 Shallow Rot 10502 3 57 5 6 2 Shallow Rot 10502 3 58 5 6 1 Shallow Rot 10502 3 59 4 6 1 Shallow Rot 10502 3 60 4 6 1 Shallow Rot 10502 3 61 3 5 1 Shallow Rot 10502 3 62 4 5 2 Shallow Rot 10502 3 63 4 6 1 Shallow Rot 10502 3 64 4 5 1 Shallow Rot 10502 3 65 6 6 2 Shallow Rot 10502 3 66 3 4 2 Shallow Rot 10502 3 67 3 4 2 Shallow Rot 10502 3 68 2 4 2 Shallow Rot 10502 3 69 2 3 2 Shallow Rot 10502 3 70 2 3 2 Shallow Rot 10502 3 71 2 5 2 Shallow Rot 10502 3 72 3 4 2 Shallow Rot 10502 3 73 2 3 2 Shallow Rot 10502 3 74 1 2 2 Shallow Rot 10502 3 75 3 3 2 Shallow Rot 10502 4 1 5 6 2 Shallow Rot 10502 4 2 4 5 2 Shallow Rot 10502 4 3 5 6 2 Shallow Rot 10502 4 4 6 6 2 Shallow Rot 10502 4 5 3 4 2 Shallow Rot 10502 4 6 2 3 2 Shallow Rot 10502 4 7 4 5 2 Shallow Rot 10502 4 8 5 6 2 Shallow Rot 10502 4 9 6 6 2 Shallow Rot 10502 4 10 3 4 2 Shallow Rot 10502 4 11 2 3 2 Shallow Rot 10502 4 12 5 6 2 Shallow Rot 10502 4 13 6 6 2 Shallow Rot 10502 4 14 5 6 2 Shallow Rot 10502 4 15 6 6 2 Shallow Rot 10502 4 16 3 4 2 Shallow Rot 10502 4 17 2 2 2 Shallow Rot 10502 4 18 2 3 2 Shallow Rot 10502 4 19 3 4 2 298

Shallow Rot 10502 4 20 3 4 2 Shallow Rot 10502 4 21 2 3 2 Shallow Rot 10502 4 22 4 5 2 Shallow Rot 10502 4 23 5 6 2 Shallow Rot 10502 4 24 6 6 2 Shallow Rot 10502 4 25 3 4 2 Shallow Rot 10502 4 26 3 3 2 Shallow Rot 10502 4 27 6 6 2 Shallow Rot 10502 4 28 4 6 2 Shallow Rot 10502 4 29 3 5 2 Shallow Rot 10502 4 30 4 6 2 Shallow Rot 10502 4 31 6 6 1 Shallow Rot 10502 4 32 6 6 1 Shallow Rot 10502 4 33 6 6 2 Shallow Rot 10502 4 34 3 5 2 Shallow Rot 10502 4 35 3 6 2 Shallow Rot 10502 4 36 4 6 2 Shallow Rot 10502 4 37 2 2 2 Shallow Rot 10502 4 38 3 4 2 Shallow Rot 10502 4 39 6 6 2 Shallow Rot 10502 4 40 3 4 2 Shallow Rot 10502 4 41 1 2 2 Shallow Rot 10502 4 42 2 3 2 Shallow Rot 10502 4 43 3 3 2 Shallow Rot 10502 4 44 2 3 2 Shallow Rot 10502 4 45 3 4 2 Shallow Rot 10502 4 46 1 2 2 Shallow Rot 10502 4 47 3 4 2 Shallow Rot 10502 4 48 2 2 2 Shallow Rot 10502 4 49 2 2 2 Shallow Rot 10502 4 50 1 2 2 Shallow Rot 10502 4 51 1 2 2 Shallow Rot 10502 4 52 1 2 2 Shallow Rot 10502 4 53 1 2 2 Shallow Rot 10502 5 1 4 6 1 Shallow Rot 10502 5 2 6 6 2 Shallow Rot 10502 5 3 2 3 2 Shallow Rot 10502 5 4 5 6 2 Shallow Rot 10502 5 5 2 3 2 Shallow Rot 10502 5 6 4 6 2 Shallow Rot 10502 5 7 5 6 2 299

Shallow Rot 10502 5 8 4 5 2 Shallow Rot 10502 5 9 5 6 2 Shallow Rot 10502 5 10 5 6 2 Shallow Rot 10502 5 11 6 6 2 Shallow Rot 10502 5 12 2 3 2 Shallow Rot 10502 5 13 3 4 2 Shallow Rot 10502 5 14 2 3 2 Shallow Rot 10502 5 15 3 4 2 Shallow Rot 10502 5 16 3 4 2 Shallow Rot 10502 5 17 2 3 2 Shallow Rot 10502 5 18 1 3 2 Shallow Rot 10502 5 19 1 2 2 Shallow Rot 10502 5 20 3 4 2 Shallow Rot 10502 5 21 2 3 2 Shallow Rot 10502 5 22 2 3 2 Shallow Rot 10502 5 23 3 4 2 Shallow Rot 10502 5 24 5 6 2 Shallow Rot 10502 5 25 3 4 2 Shallow Rot 10502 5 26 2 3 2 Shallow Rot 10502 5 27 2 3 2 Shallow Rot 10502 5 28 1 2 2 Shallow Rot 10502 5 29 1 2 2 Shallow Rot 10502 5 30 2 3 2 Shallow Rot 10502 5 31 1 2 2 Shallow Rot 10502 5 32 5 6 2 Shallow Rot 10502 5 33 4 5 2 Shallow Rot 10502 5 34 4 6 2 Shallow Rot 10502 5 35 4 5 2 Shallow Rot 10502 5 36 5 6 2 Shallow Rot 10502 5 37 6 6 2 Shallow Rot 10502 5 38 3 4 2 Shallow Rot 10502 5 39 2 3 2 Shallow Rot 10502 5 40 3 3 2 Shallow Rot 10502 5 41 2 3 2 Shallow Rot 10502 5 42 2 2 2 Shallow Rot 10502 5 43 6 6 1 Shallow Rot 10502 5 44 3 5 2 Shallow Rot 10502 5 45 6 6 2 Shallow Rot 10502 5 46 4 5 2 Shallow Rot 10502 5 47 4 5 2 Shallow Rot 10502 5 48 5 6 2 300

Shallow Rot 10502 5 49 6 6 2 Shallow Rot 10502 5 50 6 6 2 Shallow Rot 10502 5 51 5 6 2 Shallow Rot 10502 5 52 6 6 2 Shallow Rot 10502 5 53 6 6 2 Shallow Rot 10502 5 54 6 6 2 Shallow Rot 10502 5 55 6 6 2 Shallow Rot 10502 5 56 5 6 2 Shallow Rot 10502 5 57 1 2 2 Shallow Rot 10502 5 58 3 4 2 Shallow Rot 10502 5 59 2 2 2 Shallow Rot 10502 5 60 2 3 2 Shallow Rot 10502 5 61 2 3 2 Shallow Rot 10502 5 62 1 2 2 Shallow Rot 10502 5 63 3 4 2 Shallow Rot 10502 5 64 3 4 2 Shallow Rot 10502 5 65 2 3 2 Shallow Rot 10502 5 66 2 3 2 Shallow Rot 10502 5 67 4 5 2 Shallow Rot 10502 5 68 4 6 2 Shallow Rot 10502 6 1 5 6 2 Shallow Rot 10502 6 2 2 3 2 Shallow Rot 10502 6 3 2 3 2 Shallow Rot 10502 6 4 3 4 2 Shallow Rot 10502 6 5 3 4 2 Shallow Rot 10502 6 6 2 4 2 Shallow Rot 10502 6 7 6 6 2 Shallow Rot 10502 6 8 6 6 2 Shallow Rot 10502 6 9 2 3 2 Shallow Rot 10502 6 10 3 4 2 Shallow Rot 10502 6 11 3 4 2 Shallow Rot 10502 6 12 2 3 2 Shallow Rot 10502 6 13 2 2 2 Shallow Rot 10502 6 14 2 3 2 Shallow Rot 10502 6 15 3 4 2 Shallow Rot 10502 6 16 2 3 2 Shallow Rot 10502 6 17 2 3 2 Shallow Rot 10502 6 18 4 6 2 Shallow Rot 10502 6 19 5 6 2 Shallow Rot 10502 6 20 6 6 2 Shallow Rot 10502 6 21 6 6 2 301

Shallow Rot 10502 6 22 6 6 2 Shallow Rot 10502 6 23 5 6 2 Shallow Rot 10502 6 24 4 5 2 Shallow Rot 10502 6 25 6 6 2 Shallow Rot 10502 6 26 5 6 1 Shallow Rot 10502 6 27 6 6 2 Shallow Rot 10502 6 28 2 3 2 Shallow Rot 10502 6 29 3 4 2 Shallow Rot 10502 6 30 2 2 2 Shallow Rot 10502 6 31 2 3 2 Shallow Rot 10502 6 32 5 6 2 Shallow Rot 10502 6 33 2 3 2 Shallow Rot 10502 6 34 2 6 2 Shallow Rot 10502 6 35 3 4 2 Shallow Rot 10502 6 36 2 6 2 Shallow Rot 10502 6 37 2 4 2 Shallow Rot 10502 6 38 3 5 2 Shallow Rot 10502 6 39 3 4 2 Shallow Rot 10502 6 40 2 3 2 Shallow Rot 10502 6 41 3 4 2 Shallow Rot 10502 6 42 1 3 2 Shallow Rot 10502 6 43 2 3 2 Shallow Rot 10502 6 44 3 3 2 Shallow Rot 10502 6 45 2 3 2 Shallow Rot 10502 6 46 2 3 2 Shallow Rot 10502 6 47 1 2 2 Shallow Rot 10502 6 48 3 5 2 Shallow Rot 10502 6 49 2 3 2 Shallow Rot 10502 7 1 5 6 2 Shallow Rot 10502 7 2 2 3 2 Shallow Rot 10502 7 3 3 4 2 Shallow Rot 10502 7 4 3 4 2 Shallow Rot 10502 7 5 2 3 2 Shallow Rot 10502 7 6 1 3 2 Shallow Rot 10502 7 7 2 3 2 Shallow Rot 10502 7 8 3 4 2 Shallow Rot 10502 7 9 5 6 2 Shallow Rot 10502 7 10 2 3 2 Shallow Rot 10502 7 11 2 4 2 Shallow Rot 10502 7 12 6 6 2 Shallow Rot 10502 7 13 2 3 2 302

Shallow Rot 10502 7 14 3 4 2 Shallow Rot 10502 7 15 5 6 2 Shallow Rot 10502 7 16 2 3 2 Shallow Rot 10502 7 17 2 4 2 Shallow Rot 10502 7 18 3 4 2 Shallow Rot 10502 7 19 4 6 2 Shallow Rot 10502 7 20 6 6 2 Shallow Rot 10502 7 21 3 4 2 Shallow Rot 10502 7 22 3 4 2 Shallow Rot 10502 7 23 4 5 2 Shallow Rot 10502 7 24 6 6 2 Shallow Rot 10502 7 25 6 6 2 Shallow Rot 10502 7 26 3 5 2 Shallow Rot 10502 7 27 3 5 2 Shallow Rot 10502 7 28 2 3 2 Shallow Rot 10502 7 29 2 3 2 Shallow Rot 10502 7 30 1 2 2 Shallow Rot 10502 7 31 2 3 2 Shallow Rot 10502 7 32 1 2 2 Shallow Rot 10502 7 33 2 3 2 Shallow Rot 10502 7 34 2 3 2 Shallow Rot 10502 7 35 1 2 2 Shallow Rot 10502 7 36 3 4 2 Shallow Rot 10502 7 37 2 3 2 Shallow Rot 10502 7 38 1 2 2 Shallow Rot 10502 7 39 2 3 2 Shallow Rot 10502 7 40 1 2 2 Shallow Rot 10502 8 1 6 6 2 Shallow Rot 10502 8 2 3 4 2 Shallow Rot 10502 8 3 2 3 2 Shallow Rot 10502 8 4 1 2 2 Shallow Rot 10502 8 5 2 3 2 Shallow Rot 10502 8 6 5 6 2 Shallow Rot 10502 8 7 6 6 2 Shallow Rot 10502 8 8 3 4 2 Shallow Rot 10502 8 9 2 3 2 Shallow Rot 10502 8 10 1 2 2 Shallow Rot 10502 8 11 4 6 2 Shallow Rot 10502 8 12 4 5 2 Shallow Rot 10502 8 13 4 5 2 Shallow Rot 10502 8 14 6 6 2 303

Shallow Rot 10502 8 15 4 5 2 Shallow Rot 10502 8 16 4 6 2 Shallow Rot 10502 8 17 5 6 2 Shallow Rot 10502 8 18 2 3 2 Shallow Rot 10502 8 19 4 5 2 Shallow Rot 10502 8 20 4 5 2 Shallow Rot 10502 8 21 5 6 2 Shallow Rot 10502 8 22 6 6 2 Shallow Rot 10502 8 23 6 6 2 Shallow Rot 10502 8 24 6 6 2 Shallow Rot 10502 8 25 2 3 2 Shallow Rot 10502 8 26 3 4 2 Shallow Rot 10502 8 27 3 6 2 Shallow Rot 10502 8 28 1 2 2 Shallow Rot 10502 8 29 2 3 2 Shallow Rot 10502 8 30 3 4 2 Shallow Rot 10502 8 31 5 6 2 Shallow Rot 10502 8 32 3 4 2 Shallow Rot 10502 8 33 1 2 2 Shallow Rot 10502 8 34 2 3 2 Shallow Rot 10502 8 35 5 6 2 Shallow Desiccated 10510 2 1 2 3 2 Shallow Desiccated 10510 2 2 5 6 2 Shallow Desiccated 10510 2 3 6 6 2 Shallow Desiccated 10510 2 4 5 6 2 Shallow Desiccated 10510 2 5 6 6 2 Shallow Desiccated 10510 2 6 5 6 2 Shallow Desiccated 10510 2 7 6 6 2 Shallow Desiccated 10510 2 8 4 6 2 Shallow Desiccated 10510 2 9 5 6 2 Shallow Desiccated 10510 3 1 2 3 2 Shallow Desiccated 10510 3 2 5 6 2 Shallow Desiccated 10510 3 3 6 6 2 Shallow Desiccated 10510 3 4 3 4 2 Shallow Desiccated 10510 3 5 6 6 2 Shallow Desiccated 10510 3 6 5 6 2 Shallow Desiccated 10510 3 7 5 6 2 Shallow Desiccated 10510 3 8 6 6 2 Shallow Desiccated 10510 3 9 2 3 2 Shallow Desiccated 10510 3 10 5 6 2 Shallow Desiccated 10510 3 11 6 6 2 304

Shallow Desiccated 10510 3 12 3 4 2 Shallow Desiccated 10510 3 13 5 6 2 Shallow Desiccated 10510 3 14 2 3 2 Shallow Desiccated 10510 3 15 2 4 2 Shallow Desiccated 10510 3 16 6 6 2 Shallow Desiccated 10510 3 17 5 6 2 Shallow Desiccated 10510 4 1 2 3 2 Shallow Desiccated 10510 4 2 6 6 2 Shallow Desiccated 10510 4 3 2 4 2 Shallow Desiccated 10510 4 4 6 6 2 Shallow Desiccated 10510 4 5 6 6 2 Shallow Desiccated 10510 4 6 5 6 2 Shallow Desiccated 10510 4 7 6 6 2 Shallow Desiccated 10510 4 8 5 6 2 Shallow Desiccated 10510 4 9 6 6 2 Shallow Desiccated 10510 4 10 2 3 2 Shallow Desiccated 10510 4 11 6 6 2 Shallow Desiccated 10510 4 12 5 6 2 Shallow Desiccated 10510 4 13 6 6 2 Shallow Desiccated 10510 4 14 5 6 2 Shallow Desiccated 10510 4 15 5 6 2 Shallow Desiccated 10510 4 16 2 3 2 Shallow Desiccated 10510 4 17 6 6 2 Shallow Desiccated 10510 4 18 1 2 2 Shallow Desiccated 10510 4 19 2 3 2 Shallow Desiccated 10510 4 20 3 4 2 Shallow Desiccated 10510 5 1 2 3 2 Shallow Desiccated 10510 5 2 5 6 2 Shallow Desiccated 10510 5 3 6 6 2 Shallow Desiccated 10510 5 4 5 6 2 Shallow Desiccated 10510 5 5 5 6 2 Shallow Desiccated 10510 5 6 2 2 2 Shallow Desiccated 10510 5 7 6 6 2 Shallow Desiccated 10510 5 8 5 6 2 Shallow Desiccated 10510 5 9 6 6 2 Shallow Desiccated 10510 5 10 4 6 1 Shallow Desiccated 10510 5 11 4 6 2 Shallow Desiccated 10510 5 12 5 6 2 Shallow Desiccated 10510 5 13 1 2 2 Shallow Desiccated 10510 5 14 3 3 2 Shallow Desiccated 10510 5 15 2 3 2 305

Shallow Desiccated 10510 5 16 1 2 2 Shallow Desiccated 10510 5 17 6 6 2 Shallow Desiccated 10510 5 18 6 6 2 Shallow Desiccated 10510 5 19 4 6 1 Shallow Desiccated 10510 5 20 4 6 2 Shallow Desiccated 10510 5 21 4 6 1 Shallow Desiccated 10510 5 22 6 6 2 Shallow Desiccated 10510 5 23 5 6 2 Shallow Desiccated 10510 5 24 5 6 2 Shallow Desiccated 10510 5 25 4 6 2 Shallow Desiccated 10510 5 26 5 6 2 Shallow Desiccated 10510 5 27 5 6 2 Shallow Desiccated 10510 5 28 4 6 2 Shallow Desiccated 10510 5 29 5 6 2 Shallow Desiccated 10510 5 30 5 6 2 Shallow Desiccated 10510 5 31 2 3 2 Shallow Desiccated 10510 5 32 6 6 2 Shallow Desiccated 10510 5 33 5 6 2 Shallow Desiccated 10510 5 34 2 3 2 Shallow Desiccated 10510 5 35 2 3 2 Shallow Desiccated 10510 5 36 1 2 2 Shallow Desiccated 10510 5 37 3 3 2 Shallow Desiccated 10510 5 38 1 2 2 Shallow Desiccated 10510 5 39 1 2 2 Shallow Desiccated 10510 5 40 1 2 2 Shallow Desiccated 10510 5 41 1 2 2 Shallow Desiccated 10510 5 42 2 3 2 Shallow Desiccated 10510 5 43 2 3 2 Shallow Desiccated 10510 5 44 6 6 2 Shallow Desiccated 10510 5 45 2 3 2 Shallow Desiccated 10510 5 46 1 2 2 Shallow Desiccated 10510 6 1 2 3 2 Shallow Desiccated 10510 6 2 6 6 2 Shallow Desiccated 10510 6 3 6 6 2 Shallow Desiccated 10510 6 4 5 6 2 Shallow Desiccated 10510 6 5 6 6 2 Shallow Desiccated 10510 6 6 6 6 2 Shallow Desiccated 10510 6 7 5 6 2 Shallow Desiccated 10510 6 8 6 6 2 Shallow Desiccated 10510 6 9 4 6 2 Shallow Desiccated 10510 6 10 4 6 2 306

Shallow Desiccated 10510 6 11 6 6 2 Shallow Desiccated 10510 6 12 4 6 2 Shallow Desiccated 10510 6 13 5 6 2 Shallow Desiccated 10510 6 14 4 5 2 Shallow Desiccated 10510 6 15 6 6 2 Shallow Desiccated 10510 6 16 5 6 2 Shallow Desiccated 10510 6 17 6 6 2 Shallow Desiccated 10510 6 18 1 2 2 Shallow Desiccated 10510 6 19 4 5 2 Shallow Desiccated 10510 6 20 3 5 1 Shallow Desiccated 10510 6 21 3 5 1 Shallow Desiccated 10510 6 22 4 6 2 Shallow Desiccated 10510 6 23 4 5 2 Shallow Desiccated 10510 6 24 5 6 2 Shallow Desiccated 10510 6 25 6 6 2 Shallow Desiccated 10510 6 26 4 6 2 Shallow Desiccated 10510 6 27 4 6 2 Shallow Desiccated 10510 6 28 5 6 2 Shallow Desiccated 10510 6 29 3 5 2 Shallow Desiccated 10510 6 30 4 6 2 Shallow Desiccated 10510 6 31 5 6 2 Shallow Desiccated 10510 6 32 4 6 2 Shallow Desiccated 10510 6 33 6 6 2 Shallow Desiccated 10510 6 34 5 6 2 Shallow Desiccated 10510 6 35 4 6 2 Shallow Desiccated 10510 6 36 4 6 2 Shallow Desiccated 10510 6 37 2 3 2 Shallow Desiccated 10510 6 38 6 6 2 Shallow Desiccated 10510 6 39 5 6 2 Shallow Desiccated 10510 6 40 2 3 2 Shallow Desiccated 10510 6 41 1 2 2 Shallow Desiccated 10510 6 42 1 2 2 Shallow Desiccated 10510 6 43 1 2 2 Shallow Desiccated 10510 6 44 2 2 2 Shallow Desiccated 10510 6 45 2 2 2 Shallow Desiccated 10510 6 46 1 3 2 Shallow Desiccated 10510 6 47 1 2 2 Shallow Desiccated 10510 6 48 2 2 2 Shallow Desiccated 10510 6 49 2 2 2 Shallow Desiccated 10510 6 50 2 3 2 Shallow Desiccated 10510 6 51 2 2 2 307

Shallow Desiccated 10510 6 52 1 2 2 Shallow Desiccated 10510 6 53 1 3 2 Shallow Desiccated 10510 6 54 1 2 2 Shallow Desiccated 10510 6 55 1 2 2 Shallow Desiccated 10510 6 56 1 2 2 Shallow Desiccated 10510 6 57 1 2 2 Shallow Desiccated 10510 7 1 1 4 2 Shallow Desiccated 10510 7 2 1 2 2 Shallow Desiccated 10510 7 3 2 3 2 Shallow Desiccated 10510 7 4 6 6 2 Shallow Desiccated 10510 7 5 6 6 2 Shallow Desiccated 10510 7 6 2 3 2 Shallow Desiccated 10510 7 7 6 6 2 Shallow Desiccated 10510 7 8 5 6 2 Shallow Desiccated 10510 7 9 5 6 2 Shallow Desiccated 10510 7 10 5 6 2 Shallow Desiccated 10510 7 11 6 6 2 Shallow Desiccated 10510 7 12 4 6 1 Shallow Desiccated 10510 7 13 6 6 1 Shallow Desiccated 10510 7 14 5 6 1 Shallow Desiccated 10510 7 15 5 6 2 Shallow Desiccated 10510 7 16 4 6 2 Shallow Desiccated 10510 7 17 5 6 2 Shallow Desiccated 10510 7 18 6 6 1 Shallow Desiccated 10510 7 19 6 6 2 Shallow Desiccated 10510 7 20 6 6 2 Shallow Desiccated 10510 7 21 4 6 2 Shallow Desiccated 10510 7 22 5 6 1 Shallow Desiccated 10510 7 23 4 6 1 Shallow Desiccated 10510 7 24 5 6 1 Shallow Desiccated 10510 7 25 4 6 1 Shallow Desiccated 10510 7 26 6 6 1 Shallow Desiccated 10510 7 27 6 6 1 Shallow Desiccated 10510 7 28 5 6 1 Shallow Desiccated 10510 7 29 6 6 2 Shallow Desiccated 10510 7 30 6 6 1 Shallow Desiccated 10510 7 31 5 6 1 Shallow Desiccated 10510 7 32 4 6 1 Shallow Desiccated 10510 7 33 5 6 1 Shallow Desiccated 10510 7 34 4 5 2 Shallow Desiccated 10510 7 35 5 6 2 308

Shallow Desiccated 10510 7 36 6 6 2 Shallow Desiccated 10510 7 37 2 3 2 Shallow Desiccated 10510 7 38 2 3 2 Shallow Desiccated 10510 7 39 6 6 2 Shallow Desiccated 10510 7 40 4 6 2 Shallow Desiccated 10510 7 41 3 5 2 Shallow Desiccated 10510 7 42 3 5 1 Shallow Desiccated 10510 7 43 3 5 1 Shallow Desiccated 10510 7 44 3 5 1 Shallow Desiccated 10510 7 45 4 5 1 Shallow Desiccated 10510 7 46 3 5 1 Shallow Desiccated 10510 7 47 4 5 2 Shallow Desiccated 10510 7 48 4 5 2 Shallow Desiccated 10510 7 49 5 6 2 Shallow Desiccated 10510 7 50 6 6 1 Shallow Desiccated 10510 7 51 6 6 2 Shallow Desiccated 10510 7 52 4 6 2 Shallow Desiccated 10510 7 53 5 6 2 Shallow Desiccated 10510 7 54 5 6 2 Shallow Desiccated 10510 7 55 5 6 2 Shallow Desiccated 10510 7 56 5 6 2 Shallow Desiccated 10510 7 57 5 6 2 Shallow Desiccated 10510 7 58 6 6 2 Shallow Desiccated 10510 7 59 2 6 2 Shallow Desiccated 10510 7 60 2 5 2 Shallow Desiccated 10510 7 61 3 6 1 Shallow Desiccated 10510 7 62 2 2 2 Shallow Desiccated 10510 7 63 2 3 2 Shallow Desiccated 10510 7 64 2 3 2 Shallow Desiccated 10510 7 65 1 3 2 Shallow Desiccated 10510 7 66 1 2 2 Shallow Desiccated 10510 7 67 1 3 2 Shallow Desiccated 10510 7 68 1 2 2 Shallow Desiccated 10510 7 69 2 3 2 Shallow Desiccated 10510 7 70 1 2 2 Shallow Desiccated 10510 7 71 1 3 2 Shallow Desiccated 10510 7 72 1 2 2 Shallow Desiccated 10510 7 73 1 2 2 Shallow Desiccated 10510 7 74 1 3 2 Shallow Desiccated 10510 7 75 1 2 2 Shallow Desiccated 10510 7 76 1 4 2 309

Shallow Desiccated 10510 7 77 1 3 2 Shallow Desiccated 10510 7 78 1 2 2 Shallow Desiccated 10510 7 79 1 2 2 Shallow Desiccated 10510 7 80 1 3 2 Shallow Desiccated 10510 7 81 6 6 2 Shallow Desiccated 10510 8 1 1 2 2 Shallow Desiccated 10510 8 2 2 3 2 Shallow Desiccated 10510 8 3 1 2 2 Shallow Desiccated 10510 8 4 2 3 2 Shallow Desiccated 10510 8 5 2 3 2 Shallow Desiccated 10510 8 6 1 3 2 Shallow Desiccated 10510 8 7 5 6 2 Shallow Desiccated 10510 8 8 5 6 2 Shallow Desiccated 10510 8 9 2 6 2 Shallow Desiccated 10510 8 10 2 3 2 Shallow Desiccated 10510 8 11 5 6 2 Shallow Desiccated 10510 8 12 6 6 2 Shallow Desiccated 10510 8 13 6 6 2 Shallow Desiccated 10510 8 14 6 6 2 Shallow Desiccated 10510 8 15 5 6 2 Shallow Desiccated 10510 8 16 5 6 2 Shallow Desiccated 10510 8 17 6 6 1 Shallow Desiccated 10510 8 18 5 6 2 Shallow Desiccated 10510 8 19 6 6 2 Shallow Desiccated 10510 8 20 5 6 2 Shallow Desiccated 10510 8 21 4 6 2 Shallow Desiccated 10510 8 22 5 6 2 Shallow Desiccated 10510 8 23 4 6 2 Shallow Desiccated 10510 8 24 4 6 2 Shallow Desiccated 10510 8 25 5 6 2 Shallow Desiccated 10510 8 26 5 6 2 Shallow Desiccated 10510 8 27 4 6 2 Shallow Desiccated 10510 8 28 6 6 1 Shallow Desiccated 10510 8 29 5 6 2 Shallow Desiccated 10510 8 30 4 5 2 Shallow Desiccated 10510 8 31 5 6 2 Shallow Desiccated 10510 8 32 5 6 2 Shallow Desiccated 10510 8 33 2 3 2 Shallow Desiccated 10510 8 34 1 2 2 Shallow Desiccated 10510 8 35 2 3 2 Shallow Desiccated 10510 8 36 3 4 1 310

Shallow Desiccated 10510 8 37 3 5 2 Shallow Desiccated 10510 8 38 4 6 1 Shallow Desiccated 10510 8 39 3 6 1 Shallow Desiccated 10510 8 40 4 6 1 Shallow Desiccated 10510 8 41 4 5 1 Shallow Desiccated 10510 8 42 4 6 1 Shallow Desiccated 10510 8 43 4 5 1 Shallow Desiccated 10510 8 44 5 6 1 Shallow Desiccated 10510 8 45 6 6 1 Shallow Desiccated 10510 8 46 5 6 1 Shallow Desiccated 10510 8 47 5 6 2 Shallow Desiccated 10510 8 48 6 6 2 Shallow Desiccated 10510 8 49 5 6 1 Shallow Desiccated 10510 8 50 5 6 1 Shallow Desiccated 10510 8 51 5 6 1 Shallow Desiccated 10510 8 52 5 6 2 Shallow Desiccated 10510 8 53 5 6 2 Shallow Desiccated 10510 8 54 6 6 2 Shallow Desiccated 10510 8 55 6 6 2 Shallow Desiccated 10510 8 56 6 6 2 Shallow Desiccated 10510 8 57 2 3 2 Shallow Desiccated 10510 8 58 2 3 2 Shallow Desiccated 10510 8 59 1 2 2 Shallow Desiccated 10510 8 60 2 3 2 Shallow Desiccated 10510 8 61 1 2 2 Shallow Desiccated 10510 8 62 2 5 2 Shallow Desiccated 10510 8 63 1 3 2 Shallow Desiccated 10510 8 64 1 2 2 Shallow Desiccated 10510 8 65 2 3 2 Shallow Desiccated 10510 8 66 1 2 2 Shallow Desiccated 10510 8 67 5 6 2 Shallow Desiccated 10510 8 68 2 3 2 Shallow Desiccated 10510 8 69 5 6 2 Shallow Desiccated 10510 8 69 5 6 2 Shallow Desiccated 10510 9 1 2 3 2 Shallow Desiccated 10510 9 2 1 2 2 Shallow Desiccated 10510 9 3 2 3 2 Shallow Desiccated 10510 9 4 6 6 2 Shallow Desiccated 10510 9 5 6 6 2 Shallow Desiccated 10510 9 6 5 6 2 Shallow Desiccated 10510 9 7 5 6 2 311

Shallow Desiccated 10510 9 8 5 6 2 Shallow Desiccated 10510 9 9 4 5 2 Shallow Desiccated 10510 9 10 5 6 2 Shallow Desiccated 10510 9 11 4 5 2 Shallow Desiccated 10510 9 12 4 5 2 Shallow Desiccated 10510 9 13 4 5 2 Shallow Desiccated 10510 9 14 3 5 1 Shallow Desiccated 10510 9 15 4 6 1 Shallow Desiccated 10510 9 16 4 6 1 Shallow Desiccated 10510 9 17 4 6 2 Shallow Desiccated 10510 9 18 5 6 1 Shallow Desiccated 10510 9 19 4 5 2 Shallow Desiccated 10510 9 20 3 5 2 Shallow Desiccated 10510 9 21 4 5 2 Shallow Desiccated 10510 9 22 4 5 2 Shallow Desiccated 10510 9 23 5 6 2 Shallow Desiccated 10510 9 24 5 6 2 Shallow Desiccated 10510 9 25 3 5 1 Shallow Desiccated 10510 9 26 3 5 2 Shallow Desiccated 10510 9 27 4 5 2 Shallow Desiccated 10510 9 28 5 6 2 Shallow Desiccated 10510 9 29 6 6 2 Shallow Desiccated 10510 9 30 5 6 2 Shallow Desiccated 10510 9 31 6 6 2 Shallow Desiccated 10510 9 32 6 6 2 Shallow Desiccated 10510 9 33 6 6 2 Shallow Desiccated 10510 9 34 6 6 2 Shallow Desiccated 10510 9 35 5 6 2 Shallow Desiccated 10510 9 36 5 6 2 Shallow Desiccated 10510 9 37 4 6 2 Shallow Desiccated 10510 9 38 5 6 2 Shallow Desiccated 10510 9 39 4 6 1 Shallow Desiccated 10510 9 40 4 6 2 Shallow Desiccated 10510 9 41 4 6 2 Shallow Desiccated 10510 9 42 6 6 2 Shallow Desiccated 10510 9 43 6 6 2 Shallow Desiccated 10510 10 1 4 6 2 Shallow Desiccated 10510 10 2 2 2 2 Shallow Desiccated 10510 10 3 2 3 2 Shallow Desiccated 10510 10 4 1 2 2 Shallow Desiccated 10510 10 5 6 6 2 312

Shallow Desiccated 10510 10 6 5 6 2 Shallow Desiccated 10510 10 7 5 6 2 Shallow Desiccated 10510 10 8 5 6 2 Shallow Desiccated 10510 10 9 4 5 2 Shallow Desiccated 10510 10 10 6 6 2 Shallow Desiccated 10510 10 11 5 6 2 Shallow Desiccated 10510 10 12 6 6 2 Shallow Desiccated 10510 10 13 6 6 1 Shallow Desiccated 10510 10 14 5 6 1

Appendix C.2. Summary Chart of Grain-size Data

Table C.2.1: Summary of samples by sediment rating.

Table C.2.2: Total number of samples per block.

Sediment rating: Grain-size parameters in the following order: dominant grain size, grain- size range, relative fines contribution.

# counted: Total number of that combination of sediment parameters counted. Blank cells indicate that no sediment sample matched that combination of parameters.

# density: Number of samples for that combination of sediment parameters that have corresponding density samples measured from CT scans using the lineprobe tool in the software program Amira 5.1.

%: percentage of total number of samples for that combination of sediment parameters that have corresponding density readings.

IP: Impossible combinations. For instance, it is impossible to have the size category 1 as the dominant grain size, yet have no observable fines contribution.

ID: Indistinguishable combinations. For instance, while it may be theoretically possible to have a sample consist solely of category 2, it would not be possible to distinguish it from one that also had any fines contribution and so would be categorized as 221 rather 313 than 210. Size category 2 was at the limit of resolution. As such, when it was the dominant size category, it was not possible to determine if fines were present or not and because it is exceedingly rare to have that category present without also having fines and requires hydrodynamic situations not found in this study, it was assumed that fines were present.

314

Table C.2.1. Summary of counts by sediment rating.

Sediment # # rating counted density % 110 IP 120 IP 130 IP 140 IP 150 IP 160 IP 210 ID 220 ID 230 ID 240 ID 250 ID 260 ID 310 320 330 2 0 0% 340 5 2 40% 350 3 1 33% 360 IP 410 420 430 440 450 2 1 50% 460 IP 510 520 530 540 550 560 IP 610 620 630 640 1 0 0% 650 2 1 50% 660 IP 111 IP 315

121 IP 131 IP 141 IP 151 IP 161 IP 211 IP 221 ID 231 5 0 0% 241 251 3 0 0% 261 1 0 0% 311 IP 321 IP 331 2 0 0% 341 62 9 15% 351 125 21 17% 361 121 25 21% 411 IP 421 IP 431 IP 441 1 0 0% 451 31 4 13% 461 173 38 22% 511 IP 521 IP 531 IP 541 IP 551 1 0 0% 561 220 39 18% 611 IP 621 IP 631 IP 641 IP 651 IP 661 99 22 22% 112 ID 122 161 28 17% 132 88 14 16% 142 12 0 0% 152 162 3 0 0% 316

212 IP 222 26 4 15% 232 247 39 16% 242 38 6 16% 252 31 4 13% 262 17 4 24% 312 IP 322 IP 332 23 2 9% 342 187 25 13% 352 183 24 13% 362 122 16 13% 412 IP 422 IP 432 IP 442 452 96 12 13% 462 189 29 15% 512 IP 522 IP 532 IP 542 IP 552 6 0 0% 562 417 60 14% 612 IP 622 IP 632 IP 642 IP 652 IP 662 244 30 12% Totals 2949 460 16%

317

Table C.2.2. Summary of samples per sediment block containing the specified head.

10456 56 10457 50 10470 43 10471 44 10498 41 10499 52 10500 41 10501 43 10502 45 10510 45 Total 460

318

Appendix C.3. Sediment samples showing grain-size parameters matched with the corresponding density measure.

Density was recorded along a line containing 500 readings. This chart indicates the mean density as measured in Hounsfield Units, standard deviation, skewness, and kurtosis of the density readings. For statistical comparison, the density parameters were standardized to a mean of zero and standard deviation and variance of one, which is listed in the Stmean (standardized mean), StSD (standardized standard deviation, Stskew

(standardized skewness), and StKurt (standardized kurtosis). The complete density readings are available upon request of the author.

Block Sl. Sam. Sed. mean Std.dev. Skew Kurt Stmean StSD StSkew StKurt rating 10456 2 2 3 4 3 1266 52 0.082 0.066 -0.947 -1.046 -0.099 0.565 10456 2 3 2 3 3 1147 47 -0.127 0.265 -1.498 -1.196 -0.591 0.873 10456 2 4 6 6 3 1630 111 0.555 -0.353 0.735 0.788 1.012 -0.088 10456 2 14 5 6 2 1724 106 0.437 -0.416 1.174 0.641 0.736 -0.186 10456 3 8 5 6 3 1717 128 0.026 -0.747 1.141 1.307 -0.232 -0.700 10456 3 11 3 5 2 1573 52 -0.145 -0.982 0.475 -1.025 -0.633 -1.067 10456 3 16 6 6 2 1759 170 0.261 0.643 1.336 2.610 0.321 1.462 10456 3 21 6 6 3 1637 157 0.090 -0.756 0.769 2.219 -0.079 -0.715 10456 3 23 2 3 3 1025 75 0.257 -0.900 -2.063 -0.321 0.312 -0.938 10456 4 5 2 3 3 860 93 0.466 -0.067 -2.828 0.246 0.804 0.357 10456 4 6 2 3 3 1234 61 0.204 -0.131 -1.096 -0.745 0.187 0.257 10456 4 7 5 6 3 1657 93 0.109 -0.881 0.863 0.224 -0.037 -0.909 10456 4 42 4 6 2 1655 79 0.213 0.262 0.854 -0.184 0.209 0.869 10456 5 6 4 6 3 1593 63 0.495 -0.289 0.569 -0.694 0.871 0.012 10456 5 7 4 5 3 1570 69 -0.074 -1.012 0.461 -0.503 -0.467 -1.113 10456 5 25 1 2 3 1068 66 -0.638 0.831 -1.862 -0.597 -1.793 1.755 10456 5 28 5 6 3 1497 133 0.085 -0.359 0.122 1.476 -0.092 -0.097 10456 5 33 6 6 2 1616 158 0.389 -0.177 0.674 2.230 0.622 0.186 10456 6 3 1 2 3 402 152 -0.306 -0.655 -4.946 2.071 -1.012 -0.558 10456 6 4 1 2 3 406 35 0.170 0.653 -4.928 -1.559 0.108 1.477 10456 6 5 1 3 3 1193 62 0.021 -0.218 -1.287 -0.726 -0.244 0.123 10456 6 11 4 6 3 1578 99 0.438 -0.466 0.497 0.416 0.737 -0.263 319

10456 6 12 6 6 2 1429 165 0.209 -0.098 -0.195 2.476 0.200 0.310 10456 6 13 6 6 3 1429 165 0.209 -0.098 -0.195 2.476 0.200 0.310 10456 6 14 2 4 3 1549 122 0.418 -0.595 0.361 1.140 0.691 -0.464 10456 6 25 5 6 3 1486 149 0.239 -0.233 0.071 1.962 0.271 0.099 10456 7 1 2 3 3 1263 45 -0.107 -0.813 -0.960 -1.243 -0.544 -0.803 10456 7 2 1 2 3 482 118 0.672 -0.279 -4.574 1.005 1.288 0.028 10456 7 3 1 3 3 617 145 0.279 -0.810 -3.952 1.858 0.365 -0.799 10456 7 4 4 6 3 1415 79 0.282 0.367 -0.257 -0.208 0.371 1.032 10456 7 5 5 6 3 1666 144 -0.415 0.464 0.905 1.821 -1.268 1.184 10456 7 6 2 3 3 1453 96 0.242 -0.073 -0.083 0.339 0.277 0.348 10456 7 19 4 5 3 1549 126 0.310 0.013 0.364 1.242 0.438 0.482 10456 7 30 1 2 3 966 56 -0.087 -0.330 -2.335 -0.914 -0.496 -0.051 10456 7 32 2 3 3 1277 56 0.462 -0.292 -0.897 -0.905 0.794 0.007 10456 8 1 1 2 3 1050 53 -0.256 -0.711 -1.948 -1.009 -0.895 -0.645 10456 8 7 3 5 2 1230 55 0.675 0.363 -1.115 -0.938 1.295 1.026 10456 8 8 3 5 3 1295 60 -0.395 -0.237 -0.814 -0.799 -1.222 0.093 10456 8 10 3 5 3 1483 88 0.286 0.383 0.056 0.080 0.380 1.057 10456 8 14 3 4 3 1411 46 -0.231 -0.767 -0.276 -1.216 -0.836 -0.732 10456 8 15 1 2 3 1013 65 -0.142 -0.875 -2.120 -0.635 -0.626 -0.900 10456 8 17 6 6 3 1486 74 -0.621 0.162 0.071 -0.344 -1.752 0.713 10456 8 30 3 5 3 1414 111 0.134 -0.696 -0.264 0.784 0.023 -0.621 10456 8 31 5 6 3 1624 92 0.316 -0.666 0.709 0.191 0.452 -0.574 10456 9 5 2 3 3 1312 94 -0.693 -0.526 -0.734 0.263 -1.921 -0.356 10456 9 6 1 2 3 1048 49 0.067 -0.616 -1.957 -1.134 -0.134 -0.497 10456 9 18 5 6 3 1595 130 0.535 0.417 0.575 1.368 0.966 1.110 10456 9 18 5 6 3 1535 116 -0.175 0.248 0.299 0.939 -0.704 0.847 10456 9 26 4 6 2 1480 75 0.375 -0.859 0.041 -0.329 0.589 -0.874 10456 9 28 3 5 3 1334 50 0.353 0.571 -0.631 -1.080 0.537 1.350 10456 9 38 5 6 3 1523 84 1.253 0.611 0.242 -0.046 2.655 1.413 10456 9 44 4 5 3 1465 79 0.011 -0.879 -0.027 -0.192 -0.266 -0.906 10456 10 2 2 3 3 1210 38 -0.231 -0.386 -1.206 -1.474 -0.836 -0.139 10456 10 3 1 2 3 881 66 0.124 -1.130 -2.731 -0.610 0.000 -1.296 10456 10 18 5 6 3 1587 109 0.524 0.033 0.540 0.744 0.941 0.514 10456 10 25 6 6 3 1423 154 0.774 0.294 -0.218 2.109 1.528 0.919 10457 1 1 3 6 2 1145 107 0.575 0.554 -1.509 0.677 1.059 1.323 10457 1 2 3 5 2 1580 88 0.124 -0.774 0.507 0.093 0.001 -0.742 10457 1 3 5 6 2 1617 134 0.435 -0.356 0.678 1.489 0.732 -0.092 10457 2 1 4 6 2 1505 90 0.243 -0.481 0.158 0.136 0.280 -0.287 10457 2 2 5 6 2 1572 89 0.330 -0.277 0.468 0.102 0.485 0.030 10457 2 3 4 6 2 1621 99 -0.017 -0.351 0.694 0.422 -0.333 -0.085 10457 2 6 2 5 3 1601 78 0.246 -0.007 0.602 -0.217 0.287 0.450 10457 2 7 4 6 2 1635 125 0.245 -0.237 0.759 1.239 0.285 0.093 320

10457 3 1 3 6 2 1593 88 0.371 -0.157 0.568 0.093 0.580 0.217 10457 3 2 3 4 2 1596 134 1.138 1.140 0.581 1.517 2.383 2.235 10457 3 3 2 3 3 1512 53 -0.010 -0.292 0.192 -0.989 -0.315 0.007 10457 3 10 3 6 2 1666 104 0.097 0.065 0.904 0.564 -0.064 0.563 10457 3 17 3 6 2 1460 86 -0.215 -0.297 -0.048 0.007 -0.797 -0.001 10457 3 23 2 5 3 1587 57 -0.008 -0.906 0.537 -0.890 -0.311 -0.948 10457 3 30 3 6 2 1619 127 0.549 -0.212 0.685 1.299 0.999 0.132 10457 4 4 5 6 2 1645 79 -0.161 -0.739 0.809 -0.213 -0.671 -0.688 10457 4 5 4 5 3 1599 71 0.085 -1.066 0.594 -0.440 -0.091 -1.197 10457 4 7 3 5 3 1610 52 -0.072 1.355 0.647 -1.040 -0.462 2.569 10457 4 19 3 4 2 1532 75 0.266 -0.288 0.284 -0.336 0.334 0.013 10457 4 20 3 5 2 1495 66 -0.085 -1.058 0.112 -0.594 -0.492 -1.184 10457 4 25 5 6 3 1515 69 -0.483 -0.047 0.203 -0.506 -1.428 0.388 10457 4 38 3 5 3 1568 50 0.132 -0.751 0.449 -1.108 0.019 -0.706 10457 4 45 5 6 2 1498 71 0.629 -0.212 0.124 -0.441 1.187 0.131 10457 5 8 4 6 2 1587 59 0.308 -0.508 0.540 -0.830 0.432 -0.329 10457 5 23 5 6 2 1630 87 -0.300 0.080 0.739 0.058 -0.996 0.585 10457 5 41 5 6 2 1590 89 0.527 0.116 0.554 0.112 0.948 0.642 10457 5 52 4 6 3 1496 77 -0.069 -0.910 0.116 -0.247 -0.453 -0.954 10457 5 57 3 5 3 1493 77 0.236 -0.943 0.104 -0.258 0.263 -1.005 10457 6 16 3 4 3 1533 52 -0.267 -0.341 0.290 -1.019 -0.919 -0.068 10457 6 17 3 5 3 1664 89 0.261 0.124 0.895 0.118 0.322 0.654 10457 6 19 4 6 3 1624 72 0.498 -0.113 0.711 -0.403 0.880 0.285 10457 6 24 3 5 3 1626 124 0.397 -0.689 0.720 1.189 0.642 -0.610 10457 6 28 4 6 2 1460 109 -0.192 -0.374 -0.049 0.720 -0.743 -0.120 10457 6 31 4 6 3 1678 97 0.770 0.580 0.959 0.369 1.518 1.364 10457 6 34 4 5 1 1668 110 0.775 0.430 0.916 0.760 1.530 1.131 10457 6 36 3 5 2 1655 91 -0.171 -0.850 0.852 0.170 -0.694 -0.860 10457 7 16 4 6 2 1684 100 -0.174 -0.136 0.987 0.445 -0.702 0.249 10457 7 17 3 5 3 1648 105 -0.058 -1.003 0.822 0.594 -0.429 -1.099 10457 7 21 5 6 2 1736 101 0.093 -0.561 1.229 0.474 -0.074 -0.411 10457 7 24 4 6 2 1637 103 0.344 -0.915 0.770 0.535 0.518 -0.961 10457 7 26 4 6 3 1605 105 0.377 -0.063 0.620 0.595 0.594 0.363 10457 8 8 4 6 2 1639 93 0.013 -0.494 0.778 0.224 -0.261 -0.307 10457 8 10 3 4 2 1472 64 -0.453 -0.094 0.004 -0.658 -1.356 0.316 10457 8 11 4 6 2 1630 51 0.085 -0.755 0.739 -1.063 -0.093 -0.714 10457 8 15 4 6 3 1526 96 0.570 -0.453 0.255 0.334 1.048 -0.243 10457 9 3 4 6 2 1639 87 -0.197 -0.603 0.780 0.049 -0.756 -0.476 10457 9 4 3 6 2 1751 133 -0.242 -0.663 1.297 1.473 -0.861 -0.570 10457 9 4 3 6 2 1724 106 -0.197 -0.031 1.175 0.648 -0.755 0.414 10457 9 5 4 6 2 1751 133 -0.242 -0.663 1.297 1.473 -0.861 -0.570 10457 9 5 4 6 2 1724 106 -0.197 -0.031 1.175 0.648 -0.755 0.414 321

10470 1 4 3 4 3 1667 139 0.193 -0.659 0.907 1.648 0.163 -0.563 10470 1 5 5 6 2 1660 142 0.157 -0.067 0.877 1.766 0.076 0.358 10470 1 6 4 6 2 1621 107 0.567 0.439 0.696 0.661 1.041 1.144 10470 1 6 4 6 2 1623 108 0.810 0.744 0.704 0.686 1.613 1.619 10470 1 10 6 6 2 1749 187 2.134 5.523 1.286 3.154 4.726 9.052 10470 2 1 4 6 3 1674 74 -0.086 0.170 0.940 -0.343 -0.494 0.726 10470 2 8 4 6 2 1641 140 0.042 -0.337 0.791 1.699 -0.194 -0.063 10470 2 9 4 6 2 1657 75 -0.297 0.219 0.864 -0.331 -0.991 0.802 10470 2 18 4 5 3 1415 62 -0.395 -0.395 -0.259 -0.732 -1.222 -0.154 10470 3 7 3 4 3 1478 60 -0.112 -0.809 0.035 -0.775 -0.555 -0.797 10470 3 8 3 4 3 1521 57 -0.884 -0.884 0.234 -0.875 -2.371 -0.914 10470 3 10 3 4 3 1494 54 -0.479 -0.479 0.108 -0.986 -1.418 -0.283 10470 3 11 6 6 3 1711 79 0.268 -0.302 1.112 -0.196 0.338 -0.008 10470 3 12 6 6 2 1499 186 0.004 -0.249 0.130 3.116 -0.282 0.075 10470 3 13 4 6 2 1508 130 0.919 0.333 0.174 1.364 1.870 0.979 10470 4 5 3 4 3 1439 52 -0.177 -0.955 -0.149 -1.045 -0.707 -1.024 10470 4 13 4 6 2 1468 63 0.353 -0.677 -0.011 -0.686 0.539 -0.592 10470 4 31 3 4 3 1465 64 -0.268 -0.212 -0.027 -0.668 -0.922 0.132 10470 5 9 5 6 3 1725 87 -0.315 0.187 1.179 0.044 -1.033 0.753 10470 5 29 4 6 2 1458 64 0.471 0.029 -0.060 -0.667 0.816 0.507 10470 5 35 3 4 2 1464 73 -0.104 0.320 -0.031 -0.389 -0.535 0.959 10470 5 37 4 6 2 1579 88 -0.053 0.164 0.502 0.073 -0.417 0.717 10470 5 39 3 5 2 1529 56 0.525 0.613 0.270 -0.921 0.942 1.415 10470 6 9 3 4 3 1452 66 0.032 0.032 -0.086 -0.594 -0.217 0.511 10470 6 10 3 5 2 1516 70 0.007 -0.622 0.211 -0.468 -0.275 -0.507 10470 6 11 5 6 3 1561 69 0.244 -0.743 0.419 -0.493 0.282 -0.694 10470 6 14 4 6 2 1528 76 0.214 -0.906 0.265 -0.289 0.211 -0.948 10470 6 34 4 5 2 1415 92 0.684 0.354 -0.256 0.196 1.316 1.013 10470 6 39 5 6 2 1668 75 0.086 -0.986 0.911 -0.336 -0.089 -1.072 10470 7 36 3 5 2 1612 65 -0.537 -0.793 0.654 -0.633 -1.554 -0.772 10470 7 42 3 4 3 1567 71 -0.702 -0.702 0.448 -0.430 -1.942 -0.630 10470 7 43 3 5 3 1545 73 -0.441 -0.042 0.344 -0.394 -1.329 0.397 10470 7 45 4 5 2 1605 70 0.246 -1.225 0.624 -0.483 0.286 -1.444 10470 7 46 5 6 2 1637 130 0.367 -0.346 0.771 1.371 0.570 -0.077 10470 8 8 2 4 3 1636 73 0.275 -0.411 0.765 -0.392 0.356 -0.178 10470 8 14 3 4 2 1560 56 -0.120 -1.229 0.414 -0.904 -0.575 -1.450 10470 8 15 5 6 2 1683 88 0.391 -0.233 0.983 0.080 0.628 0.099 10470 8 16 4 6 2 1627 75 0.209 -0.667 0.725 -0.333 0.200 -0.576 10470 9 2 4 6 2 1580 79 -0.243 0.152 0.504 -0.203 -0.862 0.699 10470 9 4 4 5 2 1569 66 -0.666 -0.666 0.455 -0.609 -1.857 -0.574 10470 9 5 3 5 2 1617 120 0.282 -0.375 0.676 1.074 0.371 -0.122 10470 9 7 3 4 2 1571 49 -0.232 -0.232 0.463 -1.123 -0.839 0.100 322

10470 9 8 5 6 2 1666 94 -0.178 -0.680 0.906 0.266 -0.711 -0.596 10471 1 5 2 3 3 1495 63 -0.546 -0.906 0.114 -0.696 -1.576 -0.947 10471 1 6 1 2 3 1037 146 -0.418 -0.679 -2.008 1.887 -1.274 -0.595 10471 1 9 6 6 2 1696 137 0.565 -0.521 1.042 1.601 1.037 -0.348 10471 1 11 3 4 2 1498 58 0.397 1.151 0.128 -0.852 0.643 2.251 10471 1 12 6 5 1 1522 171 0.372 -0.421 0.239 2.637 0.583 -0.193 10471 2 16 4 6 2 1552 66 0.000 -0.582 0.375 -0.603 -0.292 -0.443 10471 2 25 3 5 3 1385 67 0.413 -0.337 -0.398 -0.569 0.680 -0.063 10471 2 34 2 3 3 1382 20 -0.346 0.088 -0.412 -2.026 -1.107 0.598 10471 2 35 5 6 2 1448 95 0.410 -0.729 -0.103 0.299 0.671 -0.672 10471 2 36 5 6 3 1487 60 -0.096 -0.622 0.076 -0.788 -0.518 -0.506 10471 2 38 2 3 3 1150 47 -0.191 0.512 -1.485 -1.200 -0.742 1.258 10471 2 39 1 2 3 759 77 1.127 0.862 -3.295 -0.273 2.357 1.802 10471 2 40 2 2 3 864 65 -0.241 -0.634 -2.806 -0.633 -0.858 -0.524 10471 3 1 1 2 3 827 64 -0.137 -0.136 -2.978 -0.648 -0.613 0.250 10471 3 2 2 3 3 1250 70 0.407 -0.974 -1.019 -0.480 0.664 -1.053 10471 3 4 5 6 2 1489 130 0.308 -0.188 0.087 1.390 0.432 0.169 10471 3 7 3 6 2 1410 63 0.825 0.480 -0.280 -0.705 1.648 1.208 10471 3 9 1 2 3 1201 38 -0.515 -0.149 -1.248 -1.455 -1.504 0.230 10471 3 10 6 6 3 1603 153 0.851 0.408 0.613 2.080 1.710 1.096 10471 3 12 3 5 3 1373 41 -0.311 -0.569 -0.453 -1.382 -1.023 -0.424 10471 3 27 6 6 2 1423 127 0.113 -0.512 -0.218 1.288 -0.025 -0.336 10471 4 7 4 5 3 1333 52 0.183 -0.311 -0.636 -1.024 0.138 -0.022 10471 4 21 5 6 3 1483 161 0.348 -0.052 0.059 2.348 0.526 0.381 10471 4 29 3 4 3 1290 91 0.801 0.985 -0.836 0.169 1.591 1.993 10471 4 40 5 6 3 1474 71 0.812 0.778 0.016 -0.442 1.618 1.672 10471 5 32 6 6 2 1347 215 -0.467 -0.607 -0.572 4.012 -1.389 -0.482 10471 5 66 2 4 3 1390 48 0.161 -0.005 -0.371 -1.170 0.086 0.454 10471 5 67 2 3 3 1378 32 0.429 -0.844 -0.428 -1.652 0.718 -0.851 10471 5 71 3 5 2 1517 55 0.139 -0.662 0.214 -0.935 0.034 -0.568 10471 6 6 5 6 3 1571 183 0.448 -1.059 0.465 3.013 0.762 -1.185 10471 6 23 3 6 2 1492 56 0.175 -0.763 0.100 -0.919 0.120 -0.725 10471 6 33 3 5 2 1508 60 0.281 -0.805 0.173 -0.777 0.369 -0.791 10471 6 54 1 2 3 1191 55 0.275 -0.898 -1.292 -0.939 0.356 -0.936 10471 7 3 1 2 3 1132 82 0.237 -0.631 -1.566 -0.100 0.264 -0.520 10471 7 7 6 6 3 1699 90 -0.080 -0.580 1.056 0.146 -0.481 -0.441 10471 7 9 5 6 2 1642 106 -0.258 -0.798 0.791 0.633 -0.900 -0.780 10471 7 16 3 4 2 1119 67 -0.831 0.862 -1.626 -0.569 -2.247 1.802 10471 8 1 1 3 3 1176 80 0.469 0.486 -1.365 -0.165 0.812 1.217 10471 8 8 4 6 2 1587 80 0.815 2.082 0.538 -0.155 1.624 3.700 10471 8 9 3 5 1 1517 61 -0.125 -0.692 0.213 -0.747 -0.586 -0.614 10471 8 11 3 4 1 1431 79 -0.231 -0.316 -0.185 -0.198 -0.835 -0.031 323

10471 8 12 6 6 2 1614 166 0.020 -0.441 0.664 2.478 -0.245 -0.224 10471 8 13 5 6 3 1688 81 -0.224 -0.013 1.004 -0.144 -0.818 0.441 10471 8 14 5 6 2 1455 143 1.252 1.504 -0.071 1.782 2.652 2.801 10498 2 2 3 5 2 1239 70 -0.337 -0.709 -1.071 -0.463 -1.084 -0.641 10498 2 3 3 5 2 1465 65 0.456 -0.521 -0.026 -0.625 0.781 -0.349 10498 2 6 3 5 2 1603 80 1.025 0.087 0.614 -0.180 2.119 0.597 10498 2 10 4 6 3 1584 74 0.172 -1.262 0.524 -0.361 0.112 -1.502 10498 3 4 4 6 3 1559 49 0.046 -0.857 0.407 -1.120 -0.183 -0.871 10498 3 18 3 6 3 1459 97 0.783 -0.145 -0.055 0.371 1.549 0.236 10498 3 31 3 5 3 1598 48 0.132 -0.734 0.591 -1.150 0.018 -0.680 10498 3 32 3 5 3 1372 31 0.068 -0.817 -0.454 -1.687 -0.132 -0.810 10498 3 44 5 6 3 1641 78 -0.033 -0.236 0.787 -0.231 -0.371 0.095 10498 3 45 5 6 3 1613 76 -0.295 -0.127 0.657 -0.290 -0.984 0.264 10498 4 1 5 6 2 1490 70 0.243 -0.355 0.090 -0.489 0.280 -0.091 10498 4 20 2 3 3 1151 70 0.267 -0.442 -1.477 -0.484 0.335 -0.226 10498 4 21 5 6 3 1615 82 0.138 0.257 0.666 -0.118 0.032 0.861 10498 4 22 5 6 3 1509 71 -0.319 -0.622 0.178 -0.447 -1.043 -0.506 10498 5 1 3 6 2 1516 119 -0.133 -0.806 0.210 1.038 -0.604 -0.793 10498 5 16 5 6 2 1717 70 -0.110 -0.425 1.139 -0.482 -0.551 -0.199 10498 5 21 3 6 2 1465 113 -0.050 -1.240 -0.026 0.851 -0.410 -1.467 10498 5 26 2 5 3 1125 54 -0.001 -0.001 -1.599 -0.973 -0.295 0.460 10498 5 61 3 6 2 1604 83 -0.093 -0.976 0.619 -0.071 -0.510 -1.056 10498 6 13 2 2 3 1432 64 -0.003 -1.138 -0.178 -0.674 -0.299 -1.309 10498 6 14 4 6 3 1713 93 -0.359 -0.717 1.122 0.248 -1.137 -0.654 10498 6 40 5 6 3 1565 84 -0.093 -0.644 0.437 -0.040 -0.510 -0.540 10498 6 41 5 6 3 1689 81 -0.054 -0.737 1.009 -0.136 -0.419 -0.684 10498 7 36 3 6 3 1254 92 0.738 0.042 -1.002 0.215 1.443 0.527 10498 7 47 5 6 2 1684 81 -0.091 -0.304 0.986 -0.141 -0.505 -0.011 10498 7 49 2 6 3 1743 73 0.161 -0.907 1.260 -0.392 0.086 -0.949 10498 7 50 3 6 3 1682 73 -0.108 -0.540 0.979 -0.372 -0.545 -0.379 10498 7 54 5 6 2 1565 57 -0.198 -0.291 0.435 -0.876 -0.757 0.009 10498 8 1 5 6 2 1646 98 0.484 -0.510 0.811 0.391 0.845 -0.331 10498 8 5 2 5 3 1825 82 0.292 -0.539 1.639 -0.096 0.394 -0.378 10498 8 6 3 6 2 1661 76 -0.270 -1.036 0.880 -0.301 -0.928 -1.149 10498 8 11 4 6 3 1358 93 0.399 -0.011 -0.521 0.228 0.647 0.445 10498 8 19 3 6 3 1294 89 -0.187 -0.863 -0.819 0.102 -0.731 -0.881 10498 8 21 3 5 2 1582 70 0.039 -0.286 0.517 -0.463 -0.201 0.017 10498 9 1 5 6 2 1737 96 -0.129 0.221 1.233 0.317 -0.595 0.805 10498 9 2 4 6 2 1600 127 0.350 -0.571 0.601 1.300 0.531 -0.427 10498 9 4 4 6 3 1285 102 -0.116 -0.834 -0.857 0.510 -0.564 -0.836 10498 10 1 4 6 3 1453 85 0.456 -0.812 -0.083 0.001 0.779 -0.801 10498 10 2 3 5 2 1718 86 0.002 -0.869 1.145 0.016 -0.286 -0.891 324

10498 10 3 4 6 2 1645 129 0.152 -0.634 0.806 1.350 0.066 -0.525 10498 10 4 5 6 3 1223 88 0.516 -0.319 -1.148 0.075 0.922 -0.035 10499 1 1 5 6 2 1472 131 0.269 -0.322 0.006 1.416 0.341 -0.039 10499 1 4 3 6 2 1510 72 0.280 -0.675 0.184 -0.411 0.366 -0.588 10499 1 15 4 6 2 1144 113 0.193 -0.613 -1.510 0.850 0.162 -0.492 10499 2 1 3 6 2 1470 95 0.371 -0.769 -0.001 0.289 0.581 -0.735 10499 2 2 3 5 2 1500 42 -0.151 -0.326 0.134 -1.333 -0.646 -0.046 10499 2 4 3 5 2 1491 92 0.330 0.040 0.092 0.199 0.484 0.524 10499 2 6 3 5 3 1549 60 -0.419 -0.464 0.362 -0.783 -1.278 -0.260 10499 2 8 3 6 3 1658 50 -0.162 -0.658 0.869 -1.099 -0.672 -0.561 10499 2 13 4 6 2 1207 152 0.688 -0.792 -1.220 2.073 1.326 -0.770 10499 3 1 3 6 2 1526 120 0.665 -0.368 0.255 1.073 1.271 -0.110 10499 3 3 3 6 2 1511 93 -0.181 -0.600 0.186 0.238 -0.717 -0.472 10499 3 4 3 5 2 1373 67 -0.695 0.099 -0.454 -0.566 -1.925 0.615 10499 3 7 3 6 3 1643 72 0.006 -0.702 0.798 -0.420 -0.278 -0.630 10499 3 8 3 5 3 1366 39 -0.209 -0.516 -0.484 -1.433 -0.783 -0.342 10499 3 9 3 6 3 1439 44 0.242 -0.563 -0.147 -1.275 0.276 -0.414 10499 3 10 3 6 2 1446 51 0.528 0.143 -0.116 -1.073 0.951 0.683 10499 3 11 3 4 1 1357 32 0.030 -0.710 -0.527 -1.660 -0.221 -0.643 10499 3 13 3 6 2 1364 64 0.722 0.278 -0.495 -0.676 1.406 0.894 10499 4 1 3 6 3 1598 68 -0.294 -1.035 0.590 -0.550 -0.982 -1.148 10499 4 5 3 5 2 1364 56 0.174 -0.464 -0.493 -0.921 0.117 -0.260 10499 4 9 5 6 3 1410 111 0.630 -0.609 -0.280 0.790 1.189 -0.486 10499 4 16 3 6 3 1467 58 0.070 -0.369 -0.016 -0.834 -0.128 -0.113 10499 4 30 4 6 2 1349 49 0.176 0.199 -0.561 -1.133 0.121 0.770 10499 4 31 4 6 3 1529 65 0.004 -0.143 0.271 -0.620 -0.283 0.238 10499 5 1 6 6 2 1425 129 0.560 -0.773 -0.210 1.342 1.024 -0.741 10499 5 2 6 6 3 1352 116 0.857 0.493 -0.549 0.954 1.723 1.228 10499 5 14 3 6 3 1434 54 0.155 -0.281 -0.171 -0.965 0.073 0.025 10499 5 15 3 6 2 1447 70 0.175 -0.146 -0.110 -0.469 0.119 0.234 10499 5 34 3 6 3 1491 81 0.106 -1.124 0.093 -0.139 -0.043 -1.287 10499 6 14 5 6 3 1515 87 0.206 -0.506 0.205 0.050 0.192 -0.326 10499 6 26 3 6 3 1487 71 0.468 1.346 0.074 -0.452 0.808 2.555 10499 6 28 5 6 3 1560 100 0.165 -0.799 0.412 0.465 0.097 -0.782 10499 6 29 6 6 3 1430 103 0.267 -0.931 -0.187 0.543 0.335 -0.986 10499 7 1 5 6 3 1771 135 -0.376 -0.033 1.388 1.529 -1.177 0.410 10499 7 2 5 6 2 1701 81 0.224 -0.427 1.068 -0.133 0.235 -0.202 10499 7 2 5 6 2 1701 81 0.224 -0.427 1.068 -0.133 0.235 -0.202 10499 7 3 3 6 2 1615 77 0.048 -1.048 0.667 -0.248 -0.180 -1.168 10499 7 6 6 6 2 1546 88 0.974 1.441 0.350 0.085 1.999 2.702 10499 7 8 4 6 3 1430 114 0.923 -0.192 -0.190 0.895 1.879 0.164 10499 7 15 4 6 2 1553 114 -0.407 -0.719 0.379 0.890 -1.249 -0.656 325

10499 7 16 6 6 2 1450 114 1.349 1.574 -0.094 0.882 2.880 2.909 10499 7 16 6 6 2 1517 111 0.207 -0.726 0.214 0.794 0.195 -0.667 10499 8 1 6 6 2 1703 74 -0.220 -0.617 1.074 -0.359 -0.809 -0.499 10499 8 8 5 6 2 1640 89 0.443 -0.291 0.786 0.100 0.749 0.009 10499 8 10 5 6 2 1524 111 0.195 -0.779 0.246 0.792 0.168 -0.750 10499 8 15 6 6 2 1458 103 0.381 -0.675 -0.057 0.539 0.604 -0.588 10499 8 20 6 6 2 1580 80 0.277 -0.689 0.504 -0.181 0.360 -0.610 10499 8 21 5 6 2 1567 104 0.082 -0.762 0.446 0.588 -0.098 -0.723 10499 9 1 5 6 3 1823 127 0.176 -0.542 1.630 1.285 0.121 -0.382 10499 9 2 5 6 2 1656 115 0.060 -0.934 0.858 0.906 -0.151 -0.991 10499 9 4 5 6 2 1696 93 0.513 -0.477 1.045 0.223 0.914 -0.281 10499 9 6 5 6 2 1517 99 0.321 -0.257 0.217 0.417 0.464 0.062 10500 1 2 5 6 3 1203 108 0.972 1.846 -1.237 0.712 1.993 3.332 10500 1 4 3 5 3 1414 81 -0.209 -0.226 -0.262 -0.127 -0.783 0.111 10500 1 9 3 5 3 1635 73 0.055 -0.174 0.762 -0.373 -0.161 0.191 10500 1 10 6 6 3 1619 165 0.555 0.584 0.687 2.460 1.014 1.369 10500 2 2 5 6 3 1538 86 1.254 2.047 0.311 0.006 2.658 3.645 10500 2 5 6 6 3 1600 58 0.217 -0.042 0.598 -0.850 0.219 0.396 10500 2 10 3 6 3 1421 75 -0.492 0.355 -0.230 -0.331 -1.450 1.014 10500 2 11 6 6 2 1371 110 0.254 -0.039 -0.460 0.767 0.304 0.400 10500 2 12 5 6 3 1550 115 0.972 1.913 0.369 0.922 1.994 3.436 10500 2 19 3 6 3 1598 62 0.293 -0.363 0.588 -0.734 0.398 -0.102 10500 3 9 6 6 3 1494 63 0.095 0.286 0.108 -0.682 -0.069 0.907 10500 3 14 4 6 3 1429 81 0.359 -0.797 -0.193 -0.142 0.553 -0.778 10500 3 16 3 5 3 1462 63 0.634 -0.257 -0.038 -0.696 1.200 0.062 10500 3 17 6 6 3 1579 105 0.903 1.038 0.500 0.618 1.832 2.076 10500 4 11 4 6 3 1501 76 1.171 0.897 0.142 -0.305 2.462 1.856 10500 4 12 3 6 2 1364 56 0.172 -0.560 -0.492 -0.895 0.113 -0.410 10500 4 15 5 6 3 1615 63 0.154 -1.036 0.670 -0.706 0.071 -1.150 10500 4 23 4 6 3 1447 88 0.698 0.153 -0.112 0.083 1.349 0.700 10500 5 19 3 6 3 1563 81 0.179 -0.465 0.428 -0.129 0.128 -0.262 10500 5 20 4 6 3 1545 122 -0.641 -0.021 0.346 1.118 -1.800 0.429 10500 5 21 5 6 3 1436 88 0.570 0.486 -0.159 0.079 1.048 1.217 10500 5 27 6 6 3 1547 97 0.661 1.723 0.352 0.363 1.261 3.142 10500 6 10 5 6 3 1477 70 -0.150 0.736 0.028 -0.467 -0.645 1.606 10500 6 11 5 6 3 1646 99 0.910 0.648 0.813 0.424 1.847 1.469 10500 6 20 5 6 3 1619 83 0.563 -0.329 0.689 -0.082 1.032 -0.050 10500 6 21 5 6 2 1237 69 0.426 0.347 -1.081 -0.511 0.711 1.002 10500 7 3 4 6 3 1542 102 0.232 -0.080 0.330 0.522 0.254 0.337 10500 7 4 5 6 3 1627 69 -0.434 0.089 0.724 -0.493 -1.313 0.600 10500 7 5 2 6 3 1461 63 -1.066 -1.066 -0.045 -0.688 -2.799 -1.197 10500 7 6 3 6 3 1579 110 -0.146 -0.310 0.502 0.751 -0.634 -0.020 326

10500 7 7 2 6 3 1445 78 -0.595 -0.595 -0.118 -0.220 -1.691 -0.464 10500 7 8 5 6 2 1641 100 0.299 -0.127 0.789 0.456 0.412 0.263 10500 8 5 6 6 2 1855 147 0.386 -0.017 1.777 1.915 0.617 0.435 10500 8 7 5 6 2 1706 80 0.448 -0.091 1.091 -0.153 0.761 0.320 10500 8 8 6 6 3 1757 95 0.366 0.366 1.324 0.298 0.569 1.031 10500 8 9 3 6 2 1494 93 0.285 -0.640 0.109 0.233 0.377 -0.534 10500 8 12 3 6 2 1283 95 -0.198 -0.169 -0.867 0.298 -0.757 0.198 10500 8 13 3 6 2 1227 87 0.215 -0.472 -1.129 0.059 0.214 -0.272 10500 9 4 4 6 3 1779 103 0.297 -0.036 1.427 0.557 0.405 0.405 10500 9 5 5 6 3 1657 72 -0.118 -0.341 0.862 -0.420 -0.569 -0.069 10500 9 6 6 6 2 1506 99 0.465 -0.669 0.162 0.435 0.801 -0.579 10501 2 5 1 2 3 1049 61 0.171 -0.136 -1.951 -0.763 0.111 0.250 10501 2 6 5 6 3 1565 95 0.292 -0.516 0.435 0.304 0.394 -0.341 10501 2 8 4 6 3 1755 95 -0.113 0.053 1.316 0.287 -0.557 0.544 10501 2 14 4 6 2 1682 83 0.012 -0.149 0.980 -0.077 -0.264 0.230 10501 3 8 5 6 3 1675 118 0.824 0.276 0.948 1.015 1.646 0.891 10501 3 11 2 3 3 1407 64 0.269 -0.535 -0.295 -0.669 0.341 -0.370 10501 3 16 5 6 3 1787 112 0.221 -0.418 1.463 0.838 0.227 -0.189 10501 3 19 6 6 2 1700 93 0.390 -0.598 1.063 0.251 0.625 -0.468 10501 3 20 4 6 2 1701 104 -0.245 -0.240 1.065 0.574 -0.869 0.087 10501 3 23 5 6 3 1701 114 0.084 -0.461 1.065 0.895 -0.093 -0.255 10501 3 32 1 2 3 940 61 0.155 -0.496 -2.455 -0.752 0.072 -0.310 10501 4 5 3 3 3 1503 74 -0.958 -0.958 0.151 -0.338 -2.545 -1.029 10501 4 6 2 3 3 1394 89 0.230 -0.260 -0.353 0.109 0.248 0.057 10501 4 7 1 3 3 1376 75 -0.066 -0.595 -0.440 -0.314 -0.447 -0.465 10501 4 10 6 6 3 1813 85 -0.904 -0.904 1.585 -0.008 -2.417 -0.944 10501 4 13 5 6 2 1743 82 0.033 -0.691 1.259 -0.118 -0.215 -0.613 10501 4 14 4 5 2 1622 63 -0.402 -0.576 0.699 -0.685 -1.238 -0.434 10501 4 26 1 3 3 1165 43 0.090 -0.912 -1.416 -1.313 -0.079 -0.957 10501 5 2 1 3 3 1388 35 0.269 -0.829 -0.383 -1.556 0.341 -0.829 10501 5 3 2 3 3 1485 47 0.358 -0.225 0.066 -1.191 0.550 0.111 10501 5 11 5 6 3 1708 106 0.506 -0.594 1.097 0.645 0.897 -0.462 10501 5 14 5 6 2 1555 102 0.721 -0.165 0.392 0.528 1.403 0.205 10501 5 51 1 3 3 1216 51 -0.601 -0.382 -1.181 -1.055 -1.706 -0.133 10501 6 17 2 2 3 1414 75 -0.040 -1.371 -0.264 -0.325 -0.386 -1.671 10501 6 31 5 6 3 1530 120 -0.282 -0.772 0.274 1.075 -0.955 -0.739 10501 6 33 1 3 3 1420 83 0.011 0.071 -0.235 -0.076 -0.265 0.572 10501 6 65 1 2 3 1187 50 -0.070 -0.693 -1.311 -1.090 -0.458 -0.616 10501 7 4 2 3 3 1385 39 -0.139 -0.861 -0.396 -1.426 -0.619 -0.878 10501 7 16 1 3 3 1423 45 0.163 -0.958 -0.220 -1.240 0.091 -1.029 10501 7 27 3 4 3 1412 39 0.315 -0.566 -0.272 -1.425 0.448 -0.419 10501 7 36 5 6 3 1527 83 0.287 -0.712 0.261 -0.064 0.383 -0.645 327

10501 7 41 1 3 3 1199 89 -0.366 -0.936 -1.258 0.111 -1.154 -0.994 10501 7 42 1 2 3 1238 51 0.045 -1.026 -1.076 -1.050 -0.186 -1.135 10501 8 1 4 6 3 1499 86 -0.079 0.272 0.131 0.031 -0.478 0.885 10501 8 4 3 3 3 1544 102 -1.052 -1.052 0.340 0.509 -2.766 -1.175 10501 8 8 3 5 3 1572 76 0.368 -0.048 0.471 -0.295 0.572 0.387 10501 8 33 3 5 3 1556 91 0.181 -0.336 0.396 0.174 0.133 -0.062 10501 8 38 1 2 3 1172 115 -0.452 -1.026 -1.382 0.901 -1.355 -1.135 10501 9 1 5 6 3 1340 123 0.434 -0.624 -0.603 1.175 0.728 -0.509 10501 9 2 3 4 3 1469 76 0.577 -0.171 -0.006 -0.275 1.064 0.196 10501 9 4 1 2 3 1256 129 -0.255 -0.908 -0.992 1.335 -0.891 -0.950 10501 9 12 3 4 2 1500 90 0.294 -0.590 0.138 0.146 0.399 -0.456 10501 9 31 1 3 3 1403 87 0.023 -0.561 -0.313 0.035 -0.239 -0.412 10502 1 14 5 6 3 1363 94 0.566 0.463 -0.498 0.262 1.039 1.181 10502 1 14 5 6 3 1188 107 0.442 -0.240 -1.309 0.683 0.748 0.088 10502 1 15 2 3 3 1408 85 -0.498 -0.279 -0.289 -0.010 -1.462 0.028 10502 1 16 1 3 3 1107 147 0.071 -0.911 -1.683 1.918 -0.124 -0.955 10502 1 24 6 6 3 1840 130 -0.159 -0.482 1.709 1.378 -0.666 -0.287 10502 2 3 4 6 3 1511 68 -0.254 -0.402 0.189 -0.546 -0.889 -0.164 10502 2 16 3 4 3 1462 44 -0.393 -0.735 -0.040 -1.266 -1.215 -0.681 10502 2 21 2 3 3 1455 42 -0.519 -0.495 -0.074 -1.353 -1.512 -0.309 10502 2 45 4 5 3 1486 60 0.118 -0.464 0.070 -0.799 -0.015 -0.261 10502 2 65 3 4 3 1484 40 0.074 -0.624 0.062 -1.397 -0.117 -0.509 10502 3 2 5 6 2 1576 62 -0.270 -0.616 0.487 -0.739 -0.928 -0.497 10502 3 22 1 3 3 1162 23 -0.887 0.004 -1.428 -1.922 -2.378 0.468 10502 3 65 6 6 3 1399 133 -0.113 -0.439 -0.330 1.459 -0.558 -0.221 10502 3 72 3 4 3 1360 44 -0.149 -0.392 -0.512 -1.271 -0.641 -0.149 10502 3 73 2 3 3 1148 59 -0.336 -0.937 -1.495 -0.809 -1.082 -0.996 10502 4 20 3 4 3 1400 57 -0.009 -0.521 -0.328 -0.881 -0.313 -0.349 10502 4 21 2 3 3 1102 50 -0.575 0.143 -1.708 -1.081 -1.644 0.684 10502 4 27 6 6 3 1531 127 -0.274 -0.437 0.278 1.290 -0.935 -0.218 10502 4 31 6 6 2 1517 172 0.170 -0.518 0.215 2.670 0.107 -0.345 10502 4 44 2 3 3 1314 33 0.278 -0.843 -0.726 -1.621 0.362 -0.850 10502 4 45 3 4 3 1340 40 -0.159 -0.422 -0.605 -1.411 -0.667 -0.196 10502 4 46 1 2 3 1012 109 0.042 -0.767 -2.123 0.738 -0.194 -0.732 10502 5 15 3 4 3 1468 47 0.161 -0.709 -0.011 -1.180 0.086 -0.641 10502 5 19 1 2 3 1174 43 -0.441 -0.820 -1.373 -1.302 -1.329 -0.813 10502 5 43 6 6 2 1593 135 -0.004 -0.182 0.564 1.535 -0.301 0.178 10502 5 47 4 5 3 1503 90 0.151 -0.803 0.152 0.142 0.064 -0.787 10502 6 2 2 3 3 1074 53 0.362 -0.795 -1.834 -1.013 0.560 -0.775 10502 6 4 3 4 3 1460 51 -0.359 -0.844 -0.049 -1.070 -1.137 -0.851 10502 6 7 6 6 3 1733 81 0.396 -0.599 1.215 -0.139 0.639 -0.471 10502 6 24 4 5 3 1533 57 0.064 -0.524 0.287 -0.877 -0.142 -0.353 328

10502 6 36 2 6 3 1613 83 0.378 -0.535 0.661 -0.066 0.597 -0.371 10502 6 37 2 4 3 1580 51 0.504 -0.293 0.506 -1.049 0.893 0.005 10502 6 38 3 5 3 1409 49 0.168 -0.615 -0.286 -1.133 0.104 -0.496 10502 6 42 1 3 3 940 106 0.313 -0.691 -2.455 0.636 0.443 -0.613 10502 7 3 3 4 3 1489 47 -0.089 -0.392 0.083 -1.198 -0.500 -0.148 10502 7 24 6 6 3 1741 106 0.178 -0.677 1.253 0.633 0.126 -0.591 10502 7 26 3 5 3 1387 71 -0.435 0.716 -0.388 -0.434 -1.315 1.575 10502 7 36 3 4 3 1474 52 0.208 -0.019 0.018 -1.018 0.198 0.432 10502 7 37 2 3 3 1521 47 0.036 -0.496 0.231 -1.177 -0.206 -0.310 10502 7 38 1 2 3 1059 53 0.295 -0.443 -1.903 -1.013 0.402 -0.228 10502 8 7 6 6 3 1745 107 0.487 -0.418 1.268 0.666 0.854 -0.189 10502 8 8 3 4 3 1468 65 -0.354 -0.748 -0.012 -0.618 -1.125 -0.701 10502 8 9 2 3 3 1414 71 -0.399 -1.119 -0.264 -0.430 -1.231 -1.279 10502 8 10 1 2 3 1148 60 -0.056 -0.292 -1.493 -0.774 -0.424 0.007 10502 8 20 4 5 3 1665 61 -0.069 0.235 0.899 -0.749 -0.454 0.827 10510 2 1 2 3 3 1023 85 0.228 -1.058 -2.074 -0.008 0.243 -1.184 10510 2 4 5 6 3 1695 118 1.066 1.311 1.038 1.003 2.215 2.500 10510 2 5 6 6 3 1670 148 0.088 -0.308 0.921 1.930 -0.085 -0.017 10510 2 8 4 6 3 1637 85 0.605 0.077 0.769 0.001 1.132 0.581 10510 3 1 2 3 3 969 115 -0.018 -1.069 -2.323 0.900 -0.335 -1.202 10510 3 7 5 6 3 1646 106 0.029 -0.313 0.814 0.635 -0.224 -0.026 10510 3 12 3 4 3 1210 49 -0.553 -0.005 -1.208 -1.121 -1.593 0.454 10510 3 15 2 4 3 1089 63 0.404 -1.328 -1.768 -0.687 0.657 -1.605 10510 4 3 2 4 3 988 92 -0.857 -0.259 -2.232 0.209 -2.308 0.059 10510 4 8 5 6 3 1758 109 -0.129 -0.216 1.328 0.730 -0.596 0.126 10510 4 13 6 6 3 1679 95 0.819 2.044 0.966 0.303 1.633 3.641 10510 4 16 2 3 3 913 101 0.050 -0.514 -2.579 0.469 -0.173 -0.339 10510 5 1 2 3 3 991 50 0.496 -0.366 -2.218 -1.105 0.873 -0.109 10510 5 2 5 6 3 1691 103 0.534 -0.557 1.021 0.536 0.965 -0.405 10510 5 10 4 6 2 1631 75 -0.844 0.726 0.743 -0.317 -2.276 1.591 10510 5 28 4 6 3 1601 64 -0.196 -0.312 0.604 -0.651 -0.754 -0.024 10510 5 29 5 6 3 1460 118 0.652 0.115 -0.049 1.021 1.242 0.640 10510 6 3 6 6 3 1617 143 0.242 -0.965 0.678 1.793 0.277 -1.040 10510 6 19 4 5 3 1459 46 -0.233 -0.173 -0.052 -1.215 -0.840 0.192 10510 6 21 3 5 2 1319 52 -0.107 -0.493 -0.703 -1.024 -0.543 -0.305 10510 6 40 2 3 3 1344 37 -0.945 0.761 -0.587 -1.502 -2.513 1.646 10510 7 23 4 6 2 1468 50 0.100 0.643 -0.013 -1.102 -0.057 1.462 10510 7 34 4 5 3 1481 58 0.381 -0.347 0.049 -0.844 0.605 -0.079 10510 7 35 5 6 3 1493 76 0.107 -0.691 0.101 -0.278 -0.041 -0.614 10510 7 38 2 3 3 1491 49 -0.208 -0.818 0.094 -1.123 -0.781 -0.811 10510 7 39 6 6 3 1464 186 -0.654 -0.352 -0.033 3.099 -1.830 -0.087 10510 7 58 6 6 3 1665 61 -0.093 -0.805 0.898 -0.761 -0.511 -0.791 329

10510 7 62 2 2 3 1285 44 0.035 -1.239 -0.861 -1.283 -0.210 -1.466 10510 8 2 2 3 3 1250 49 -0.356 -0.757 -1.021 -1.136 -1.129 -0.716 10510 8 3 1 2 3 1029 119 0.805 -0.448 -2.045 1.052 1.601 -0.235 10510 8 21 4 6 3 1504 53 0.024 -1.021 0.154 -1.001 -0.235 -1.127 10510 8 50 5 6 2 1582 90 0.167 -0.661 0.517 0.152 0.100 -0.566 10510 8 52 5 6 3 1582 94 1.146 0.807 0.514 0.269 2.403 1.716 10510 8 57 2 3 3 1442 22 -0.045 -0.859 -0.134 -1.956 -0.398 -0.875 10510 9 1 2 3 3 1069 42 0.106 -1.109 -1.860 -1.341 -0.043 -1.264 10510 9 2 1 2 3 1137 33 0.273 -0.048 -1.545 -1.630 0.350 0.386 10510 9 3 2 3 3 1337 43 -0.535 -0.291 -0.618 -1.301 -1.549 0.009 10510 9 6 5 6 3 1680 65 -0.412 -1.025 0.968 -0.625 -1.260 -1.133 10510 9 15 4 6 2 1534 68 1.076 0.434 0.293 -0.533 2.238 1.136 10510 9 42 6 6 3 1714 136 0.526 -0.442 1.125 1.560 0.945 -0.225 10510 10 3 2 3 3 1325 44 -0.060 -0.871 -0.675 -1.290 -0.433 -0.894 10510 10 4 1 2 3 1255 67 -0.994 0.413 -0.997 -0.567 -2.630 1.104 10510 10 5 6 6 3 1707 159 0.547 0.191 1.093 2.263 0.994 0.758 10510 10 6 5 6 3 1175 116 1.119 1.215 -1.370 0.946 2.339 2.351 10510 10 7 5 6 3 1655 122 0.072 -0.636 0.855 1.120 -0.122 -0.528

330

Appendix C.4. Statistical Explanations

1. Assumption Testing

The standardized density values were checked for normality and outliers using the robustreg procedure in SAS 9.2 and as well as the programs NCSS and PAST. SAS found only one outlier in block 10498, slice 8, sample 5. However, when the residuals were compared to the calculated robust Mahalanobis distance, a recommended method for visually identifying outliers (Chen, 2002), it plotted with the other readings (Fig.

C.4.1). When looking at the data for that sample, we were unable to determine anything unusual about it. In contrast, the sample 10, from block 10470, slice 1, appeared noticeably different, with a robust Mahalanobis distance of 14.1, but which was not identified as an outlier by any test.

Figure C.4.1. Plot of sediment density residuals vs. Robust Mahalanobis Distance 331

A box plot of the individual variables indicated several potential outliers (Fig.

C.4.2). Block 10456, slice 6, samples 3 and 4, and slice 7, sample 2 all had means below

-4.5. Block 10456, slice 7, sample 3 had a mean of -3.95, the next lowest is -3.30. The lowest three all have grain-size parameters of 1 ,2, 3. The next two have 1, 3, 3 and 1, 2,

3, respectively, which corresponded to the general pattern of density correlations with grain-size parameters. The sample with the highest standard deviation was 10471, slice 5, sample 32, with an SD of 4.01 (the next highest SDs were 3.15 and 3.12, both from

10470, slices 1 and 3, samples 10 and 12, respectively). They both had grain-size parameters of 6, 6, 2, which again corresponded well with the overall pattern. The only outlier on skewness is 10470, slice 1, sample 10, with a skewness of 4.73 (the next highest is 2.88 from 10499, slice 7, sample 16. Both have grain-size parameters of 6, 6, 2.

This is the same one that is the biggest outlier for kurtosis, with a value of 9.05. The next highest value is 3.70, from 10471, slice 8, sample 8, with grain-size parameters of 4, 6, 2.

Thus, sample 10 from block 10470, slice 1, had the highest skewness and kurtosis by a considerable margin, the highest standard deviation, was among the highest means, and a substantially larger Mahalanobis distance than any other sample. It was, nevertheless, not calculated to be a multivariate outlier.

332

Box Plot

10.0

4.7 Amount -0.7

-6.0 Stmean StSD StSkewness StKurtosis Variables

Figure C.4.2. Box plot of density measures. Red indicates those samples three standard deviations from the univariate mean of the given variable.

The only outlier found by SAS was not an outlier by any single measure. A data screening by NCSS revealed 46 potential multivariate outliers. However, none of them matched any of the outliers of any single variable, nor did it count the outlier found by

SAS as an outlier. Thus, without any consensus from the statistical analyses and without any clear reason to eliminate any of the samples after examining the data for each of the points, it was decided to retain all the data for further analyses.

All of the variables failed normality tests in both programs. However, due to the sensitivity of these tests, large sample numbers often fail to pass these tests, even if they do have normal distributions. Additionally, most of the common statistical tests become 333 very robust to violations of normality with sample sizes greater than 50-100 (Hopkins,

2009). Thus, it is generally recommended that these types of data sets are not normalized.

Normalizing data can cause problems with interpretation of the results due to the alteration of the data, which may alter relationships between sample points. Of far more importance is equality of variance, which was made equal in all the variables by the standardization. Provided both normality and equality of variances are not violated, the tests are generally considered robust; with equal variances and large sample sizes, violations of normality are less worrisome (Quinn and O’Keough, 2002).

2. Hypothesis Testing: MANOVA

To test whether or not the categories of grain-size parameters were significantly different, a MANOVA was run using SAS. Table C.4.1 lists the results of the MANOVA.

The categories for dominant grain size and range showed clearly significant results for density parameters overall, but individually, only mean and standard deviation proved to be truly significant variables. Relative fines contribution categories, proved to be insufficiently separate to be distinguishable, except for standard deviation, which was the only variable able to distinguish the fines contribution. However, the degree to which the significance translates into reliable prediction was limited, as indicated by r2 values:

StMean – 0.53. StSD – 0.36, StSkewness – 0.10, and StKurtosis – 0.06. Thus, density mean and standard deviation seem to be important parameters, but for the most part, skewness and kurtosis can be dispensed with. Of the four MANOVA statistics used to determine significance, all four (Wilks’ lambda, Pillai’s Trace, Hotelling-Lawley’s Trace, and Roy’s Greatest Root), gave similar results, except in one case. For relative fines 334

contribution, Roy’s Greatest Root was 0.1131, as compared to the 0.47 reported by the

other three. While still not normally considered significant, this does indicate a trend that

may be more clearly detectable with refined practices.

Table C.4.1. Results of MANOVA on full data set. Wilk's Lambda is reported, although

similar results were obtained using Pillai's Trace and Roy's Greatest Root

Wilk’s λ F Value Num DF Dem DF Pr > F Grain Size 0.7744 5.92 20 1476.8 <0.0001 Range 0.8782 3.69 16 1360.1 <0.0001 Fines 0.9831 0.95 8 890 0.4726

DF Type III SS Mean Square F Value Pr > F StMean Dom_sedsize 5 9.73185071 1.94637014 4.08 0.0012 Sedrange 4 15.9320913 3.98302282 8.36 <.0001 Fines 2 0.20362469 0.10181235 0.21 0.8077 StSD Dom_sedsize 5 58.75392192 11.75078438 17.92 <.0001 Sedrange 4 9.21606673 2.30401668 3.51 0.0077 Fines 2 3.98672344 1.99336172 3.04 0.0488 StSkewness Dom_sedsize 5 7.33184772 1.46636954 1.58 0.1633 Sedrange 4 6.90988388 1.72747097 1.87 0.1154 Fines 2 0.83604843 0.41802422 0.45 0.637 StKurtosis Dom_sedsize 5 8.78306557 1.75661311 1.82 0.1082 Sedrange 4 1.19054696 0.29763674 0.31 0.8727 Fines 2 0.09269893 0.04634947 0.05 0.9532

335

3. Hypothesis Testing: Discriminant Function Analyses

The MANOVA indicated significance separation between the groups, indicating that, although they were distinguishable as a whole, the correlation was not great.

Because predictive capacity was the goal of this study, a discriminant function analysis was done for each grain-size parameter. In general, these confirmed the MANOVA results and allowed for more detailed examination.

a. Dominant Grain Size

As can be seen from the weightings, the mean and standard deviation of the density readings had the largest effects overall (Table C.4.2). Categories 1 and 5 were determined mostly by the mean density, whereas category 2 and 6 were influenced mainly by mean and standard deviation. Categories 3 and 4 were poorly constrained by any of the density variables. The variables do show a clear pattern between categories.

All four density parameters increase as one goes from fine-grained sediments to higher, indicating thesediment gets more dense and more heterogeneous as the grain size increases.

The pattern of the extremes performing differently carried on throughout the analyses. Despite the MANOVA indicating extremely significant differences between groups, their predictive value left much to be desired. Table C.4.3 lists the error rates after using the weightings to predict the grain sizes based on density values.

336

Table C.4.2. Discriminant function scores for sediment density parameters by grain-size

category.

Grain-size Categories Variable 1 2 3 4 5 6 Constant -3.73297 -1.04438 -0.13569 -0.15641 -0.4538 -1.52829 Stmean -3.81197 -1.54321 0.07318 0.77924 1.10384 1.21098 StSD -0.45138 -1.03183 -0.57945 0.009 0.55802 1.77743 StSkewness -0.51318 -0.22546 -0.12381 0.1506 0.29941 0.1432 StKurtosis -0.30647 -0.39976 -0.02849 0.06437 0.12504 0.40438

Table C.4.3. Predictive error rates by grain-size category.

1 2 3 4 5 6 Total

0.3333 0.6140 0.5840 0.7262 0.6768 0.3962 0.5551

SAS calculated the total error rate as simply an average of the categories with no regard to the amount of data within each category, i.e. it added the average error rates together and divided by the number of categories. A proper average error rate, dividing the total number incorrectly identified by the total number of samples, gives a total error rate of 0.5891. Random chance would have provided an error rate of 0.8333, so this is only marginally better than chance. There is much better differentiation at the two extremes, with little resolution in the middle ranges.

A close look at the distribution of the errors revealed that most errors were only one category off. Due to the inherent difficulties of gauging grain-size categories, it seemed reasonable to analyze the data using a rolling expanded bin system. By expanding 337 the results to include the bracketing grain sizes, allowing for a less precise measurement, results substantially improved (Table C.4.4). In this case, the scores for the groups on either side of the correct size category were combined, e.g. 1=1+2, 2=1+2+3, 3=2+3+4,

4=3+4+5, 5=4+5+6, and 6=5+6.

Table C.4.4. Predictive error rates by grain-size category using rolling expanded bins.

1 2 3 4 5 6 Total

0.0476 0.1228 0.1600 0.2143 0.1515 0.1509 0.1522

The error rate for a random sample in this case is 63.9%. Here, we see that it predicts very well if a sample is relatively finer-grained and fairly well if a sample is coarse-grained, with a little less reliability if it is medium-grained. As such, we can say that CT scan densities are suitable for broad, relative assessments, but not for precise measurements and multiple readings should be taken for reliable estimates.

Because the data consisted of sets composed of different burial situations and differing amounts of organics within the system (due to the varying amount of soft tissue present on the heads during burial), the possibility existed that these differences might appear in the reliability estimates of the density readings. Thus, separate analyses were done for each flume and head condition (Table C.4.5). For the different head conditions, data from both flumes were combined as the decreased size of the data sets made the 338 analyses less reliable, and it was deemed that fifty data points or less were too small to get a good estimate with six categories.

Table C.4.5. Predictive error rates by grain-size category, burial and tissue condition.

For ease of comparison, data for the full set is repeated here.

1 2 3 4 5 6 Total 0.3333 0.6140 0.5840 0.7262 0.6768 0.3962 0.5551 Shallow Flume 0.4048 0.6522 0.4750 0.5172 0.5682 0.3438 0.4935 Deep Flume (there were no category 1 data in the deep flume) ------0.3636 0.8000 0.8182 0.5455 0.5714 0.6197 Fresh Heads 0.2857 0.3750 0.2963 0.5789 0.6250 0.3000 0.4102 Rotten Heads 0.2381 0.5294 0.6226 0.7727 0.6250 0.4643 0.5420 Desiccated Heads 0.3333 0.4706 0.7727 0.6667 0.6316 0.2857 0.5268 Clean Skull 0.0909 0.4667 0.8261 0.6875 0.5833 0.2500 0.4841

The smaller groups in general improved performance at the ends by sacrificing reliability in the middle. Data from the shallow flume also appear more reliable than those from the deep flume. If the data from the head conditions were separated by flume, they would likely show a concomitant increase in reliability for the shallow flume data and decreased for the deep flume, particularly in the fresh heads and clean skulls. The poorer showing of the deep flume may be due to the fact that the sediment was less variable throughout than that seen in the shallow flume, which may have affected the reliability of the separations. 339

b. Grain-size Range

As was the case with the dominant grain size, mean density seems to have the greatest effect on determining range categories, followed by standard deviation, skewness, and kurtosis in that order, with kurtosis having increasingly less effect as range increases (Table C.4.6).

Table C.4.6. Sediment density parameters by range category.

Variable 2 3 4 5 6 Constant -4.56130 -1.47067 -0.36527 -0.14413 -0.33681 Stmean -4.28703 -2.15869 -0.38156 0.30739 0.95451 StSD -0.42910 -0.75739 -0.79667 -0.54132 0.47778 StSkewness -0.59494 -0.49591 -0.38966 0.02386 0.23216 StKurtosis -0.34635 -0.25242 0.14514 -0.07366 0.09080

Despite the MANOVA indicating extremely significant differences between groups, here again their predictive value left much to be desired, although it is better than dominant grain-size estimates. Table C.4.7 lists the error rates for each grain-size category after using the weightings to predict the grain sizes based on density values.

SAS calculated the total error rate as simply an average of the categories with no regard to the amount of data within each category; a more proper average error rate gives a total of 0.4261, actually improving the results. Random chance would have provided an error rate of 0.8000, so this is clearly better than chance. There is better differentiation at the two extremes, with little resolution in the middle ranges.

340

Table C.4.7. Predictive error rates by range category.

Category 2 3 4 5 6 Total

Error rate 0.3750 0.6545 0.4286 0.5882 0.3422 0.4777

However, if we expand the ranges as we did with the dominant grain-size estimates, we see that grain-size range using a broad stroke is well predicted, although again the middle range is less well predicted than the endpoints (Table C.4.8). Range seems to be slightly better predicted than dominant grain size, but CT density is still effective as only a crude, relative assessment tool.

Table C.4.8. Predictive error rates by range category using rolling expanded bins.

Category 2 3 4 5 6 Total

Error rate 0.0625 0.1272 0.1905 0.0735 0.1255 0.1196

If we separate out the data into groups, we find that, as in the dominant grain size, the capability of CT density readings to predict sediment grain-size range in the sediment blocks containing fresh heads and clean skulls showed a marginal improvement over the full set (Table C.4.9). Disregarding category 2, the shallow flume proved marginally more reliable in the higher ranges here as well. Category 2 should be discounted for the 341 deep flume as there was only one reading for the deep flume as compared to the 31 in the shallow flume. Likewise, there are only four readings for category 2 in the dry heads, making the error rate of little value. This may or may not be balanced by the high error rate in category 4 with only seven readings. The preference for improving the ends at the cost of the middle is still present, but not as apparent.

Table C.4.9. Predictive error rates by range category, burial and tissue condition.

2 3 4 5 6 Total Full set 0.3750 0.6545 0.4286 0.5882 0.3422 0.4777 Shallow flume 0.3871 0.6604 0.5600 0.5000 0.2083 0.4632 Deep flume 0.0000 0.5000 0.4706 0.7500 0.4790 0.4399 Fresh heads 0.2857 0.1667 0.4706 0.5625 0.2927 0.3556 Rotten heads 0.0909 0.6800 0.2667 0.6667 0.3304 0.4069 Dry heads 0.5000 0.5833 0.8571 0.4375 0.3750 0.5506 Clean skulls 0.2000 0.3333 0.6667 0.5556 0.2963 0.4104

Because of the differences between flumes, one may expect that error rates would be smaller if comparisons between tissue conditions were made within the individual flumes, and the calculated error rates bear this out (Table C.4.10). Due to the numbers being significantly smaller when broken down this far, we will only report the values for the expanded scale as discussed above and only total error rates.

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Table C.4.10. Predictive total error rates for range in each individual burial condition

using expanded rolling bins.

Shallow flume, fresh head: 0.0909 Deep flume, fresh head: 0.0930 (there are no category 2 or 3 data here) Shallow flume, rotten heads: 0.0795 Deep flume, rotten heads: 0.0323 (no cat.2 or 3 here) Shallow flume, dry heads: 0.0888 Deep flume, dry head: 0.1400 (no cat. 2 here) Shallow flume, clean skull: 0.0714 Deep flume, clean skull: 0.0976 (no cat. 4, only 1 value for cats. 2 and 3)

As can be seen, when painting with a broad brush, the CT density values have

utility in distinguishing grain-size range relative variation, particularly when one is

comparing within the same set of readings.

A Mann-Whitney-U Test was run using Pro-Stat, which confirmed that the

readings from the shallow and deep flume were significantly different in all three grain-

size parameters (Table C.4.11). Overall, sediment samples from the blocks taken from the

deep flume (N=227) were coarser and more heterogeneous, with less relative fines

contribution than those from the shallow flume (n=233).

Table C.4.11. Mann-Whitney U hypothesis significance test comparing sediment grain- size parameters between the deep and shallow flumes. Mean Ranks U P Deep - Shallow Dominant Grain Size 126.1 – 104.4 0.0002 <0.0001 Grains Size Range 138 – 92.53 0.0002 <<0.0001 Relative Fines Contribution 93.77 – 136.7 0.0002 <<0.0001

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APPENDIX D: THIGH CIRCUMFERENCE ESTIMATION

When examining a fossil with preserved skin, it is not always easy to know how much skin has been preserved and whether or not the skin preserved is dorsal, ventral, or both. To address this question, a pilot study was undertaken to determine the feasibility of using femoral length as a proxy to estimate thigh circumference in lizards. Skin coverage over the femur was estimated by taking measurements of 19 lizards preserved in formaldehyde held by the Museum of Discovery in Little Rock, AR, and linearly regressing thigh circumference with femoral length. Appendix Table 1 lists the species sampled. The amount of skin preserved around the left femur of Monjurosuchus splendens was then compared with these data. Monjurosuchus has been identified as a champsosaurid choristodere, semi-aquatic diapsid reptiles of uncertain relations. They have been considered eosuchians (Estes 1964), basal lepidosauromorphs (Erickson,

1972), basal diapsids (Benton 1985, Carroll 1988, Cvancara and Hoganson 1993), basal archosauromorphs (Gauthier et al. 1989, Peng et al. 2001), and even relatives of sauropterygians(Muller 2004). Nevertheless, Monjurosuchus is a small reptile with skin impressions and a body shape that have been compared to lizards (Liu and Wang, 2008), and so using lizards as a comparison seems appropriate, which is fortunate as lizards are much more numerous and available for study than are living crocodilians.

Dissections to determine exact measurements were not possible, so femoral lengths were measured by flexing the leg to make the distal end of the femur accessible with the skin pressed tightly to the joint. Congruent with the site of skin preservation on the Monjurosuchus specimen, only left thighs were measured. Measurements were taken 344 from the joint to the medial-most skin crease, which marked the pivot point of the joint and which palpation indicated overlay the proximal end of the femur. Measurements were taken to the nearest 0.05 mm, although realistically measurements could not be that precise due to not having clear access to the bone itself. Thigh circumference was measured by putting tape around the mid-point of the thigh, marking the circumference, removing the tape and measuring the distance on the flattened tape using the same calipers used to measure femoral length. Due to the error involved in measuring a pliant cylindrical object, each thigh was measured 4–6 times, and the results were averaged.

The precision of the measurements is limited, but as the goal is to obtain an estimate for general trends of skin surface to bone length to compare with unrelated but similar fossil reptiles, the limitations in precision make interpretations of fossils more robust, not less, as fossil measurements that fall outside the confidence intervals of the measurements are therefore more likely to be actual differences, not random or artifactual.

Monjurosuchus itself was unavailable for direct measurement, and so measurements of the skin impressions and femoral length were taken from the photograph in Liu and Wang (2008). The image is high quality and the skin impressions and femur are clearly visible. No scale bar is present in the image, so these measurements may not reflect the actual measurements of the animal, but since it is the ratio between the measurements that is important, not the precise lengths, this discrepancy is unimportant.

The lizards and the measurements are shown in Appendix Table 1. Due to the small number of lizards, the sample is not all-inclusive of lepidosaurs, but they do include 345 lizards of wide phylogenetic breadth, encompassing iguanids and two groups within

Scleroglossa, although the estimate of phylogenetic breadth depends very much on which phylogenetic analysis one uses though (Appendix Figs. 1, 2). There is considerable debate concerning squamate phylogeny. Morphological and combined analyses such as those by Lee (2005) show a fairly classic arrangement, with iguanids separated from other lizards and dibamids, amphisbaenians, and snakes being labile. Recent molecular phylogenies however, nest iguanids deeply within squamates, forming a clade within lizards along with the varanoids and snakes, with dibamids being an outgroup to everything else. Regardless of which phylogeny one chooses, however, the Iguanidae,

Teiidae, and Scincidae are highly divergent groups. In the classic sense, iguanids comprise a group separate from the scleoglossans, with the teiids and skinks forming separate groups within Scleroglossa. In the molecular phylogeny by Vidal and Hedges

(2005, 2009), iguanids are part of a large group referred to as Toxicofera, teiids are part of the Laterata, which together with the Toxicofera comprise the Episquamata, the sister clade to the Scinciformata. This subset of lizards does not encompass a large set of lizards, the Bifurcata and Dibamidae under the molecular phylogeny or the anguilloforms under the combined dataset by Lee (2005). However, the groups not included show virtually no overlap between Vidal and Hedges (2009) phylogeny and Lee’s (2005) phylogeny. Thus, regardless of which one is favored, the sampled species do not cover the whole phylogeny, although which part is left off is dependent on one’s point of view.

Needless to say, more sampling is needed to confirm the trend reported here. 346

The data were analyzed using PSI-Plot 9.01. The data showed a high degree of correlation, showing a Pearson correlation coefficient (r) of 0.95, fit to the line y=1.32x+0.08. With a femoral length of 69.1 mm, Monjurosuchus could be expected to have a thigh circumference of 91 mm. Instead, we see only 53.2 mm, indicating that despite having what appear to be both dorsal and ventral scales present, no dislocation of skin appears to have occurred. The amount of skin present is almost half the expected amount, meaning the thigh was simply compressed, although compressed in a non- vertical direction, allowing the skin to roll somewhat to show both sides of the thigh integument. Because of this, one might expect some shear deformation of the skeletal elements. The success of this project indicates that femoral length can be used to estimate thigh circumference, as long as the comparisons are being made on similar body types.

Being able to make such estimates is useful for determining the extent of skin preservation and thereby making inferences about placement of the skin on the body. The ability to accurately estimate these measurements will allow more accurate reconstruction of animals from their skeletons.

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Figure D.1. From Lee, 2005. Figure 7a. Combined dataset using ordered multistate characters. 348

Figure D.2. From Vidal and Hedges 2009

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Table D.1. Measurements of femoral length and thigh circumference. Standard deviation of thigh circumference measurements (Th.C st.dev.) and number of thigh measurements (n) also included. Species Femur length Thigh Circumference Th. C st. n (mm) (mm) dev. Iguanidae Basciliscus vittatus 45.20 52.1 1.95 4 Crotophytus collaris 17.35 22.1 0.24 4 Crotophytus collaris 25.95 30.5 1.33 4 Crotophytus collaris 27.30 25.0 1.32 4 Phrynosoma cornutum 14.20 17.0 0.98 4 Phrynosoma cornutum 19.40 24.4 0.80 5 Sauromalus obesus 40.20 58.3 1.12 4 Sauromalus obesus 45.20 59.2 1.39 4 Sceloporous graciosus 12.15 16.5 0.33 4 graciosus Sceloporous undulatus 13.85 17.7 0.61 4 Uta stansburiana 16.25 24.6 0.67 4 stejnegeri Iguanidae sp. 40.65 65.5 2.06 6

Scleroglossa Cnemidophorus tesselatus 16.95 23.3 0.50 4 Cnemidophorus tesselatus 18.40 22.0 0.93 4 Cnemidophorus tigris 12.50 14.2 0.74 4 tigris Cnemidophorus tigris 16.50 27.7 1.73 4 tigris Cnemidophorus tigris 25.05 28.3 0.34 4 tigris Eumeces laticeps 16.00 24.3 0.14 4 Eumeces laticeps 16.95 26.5 0.64 4

Fossil Choristodere Monjurasuchus splendens 69.10 53.2

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Figure D.3. Graph of thigh circumference to femoral length. Yellow dots are

Scleroglossa, red are Iguanidae, blue is Monjurosuchus.

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