UNIVERSITY OF CINCINNATI
Date:______Jan 26, 2009
I, ______,Donald Anton Esker hereby submit this work as part of the requirements for the degree of: Master of Science in: Geology
It is entitled: An Analysis of the Morrison Formation’s Terrestrial Faunal Diversity Across Disparate Environments of Deposition, Including the Aaron Scott Site Dinosaur Quarry in Central Utah
This work and its defense approved by:
Chair: ______Dr. Glenn Storrs ______Dr. Arnold Miller ______Dr. Carton Brett ______Dr. David Meyer ______
An Analysis of the Morrison Formation’s Terrestrial Faunal Diversity
Across Disparate Environments of Deposition,
Including the Aaron Scott Site Dinosaur Quarry in Central Utah
A thesis submitted to the
Division of Research and Advanced Studies
Of the University of Cincinnati
in partial fulfillment of the
requirements for the degree of
MASTER OF SCIENCE
In the Department of Geology
of the College of Arts and Sciences
2009
by
Donald A Esker
B.S., Marietta College, 2004
Committee Chair: Dr. Glenn W. Storrs i Esker 1/26/2009
ABSTRACT -- The Aaron Scott Site dinosaur quarry (Quarry) in the Morrison Formation of Utah offers a unique view of Late Jurassic patterns of terrestrial diversity. The Quarry represents a rare perennial lacustrine environment of deposition, preserving a diverse population of large and small vertebrate and invertebrate fauna.
The null hypotheses state that patterns of diversity at the Quarry do not differ significantly from those found at ephemeral lacustrine sites elsewhere in the Morrison, and an even spread of terrestrial diversity across the Late Jurassic Morrison Basin.
While evidence has revealed a similarity between the Quarry and ephemeral lacustrine sites, multivariate analysis reveals distinct patterns in terrestrial diversity of the Morrison Formation, most prominently, a division between wetland and dry land taxa, and between ornithischian and sauropod dominated environments. Unusual patterns found among several taxa pairs hint that two (or more) Morrison genera may be sexual dimorphs or organisms at different stages in their ontogeny. While the Quarry itself may not be wholly unique, the Morrison was far more complex than traditionally portrayed.
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Table of Contents
ABSTRACT ...... II INTRODUCTION ...... 1
BROADER SCIENTIFIC IMPACTS ...... 2 BACKGROUND ...... 5
PALEONTOLOGY ...... 5 STRATIGRAPHY ...... 6 QUARRY HISTORY AND ACTIVITY ...... 11
EXCAVATION TECHNIQUES ...... 11 VERTEBRATE PREPARATION TECHNIQUES ...... 14 QUARRY POTENTIAL ...... 15 A BRIEF HISTORY OF MORRIS ON FORMATION RESEARCH ...... 17
OVERVIEW OF THE MORRISON FORMATION ...... 19 LITERATURE REVIEWED ...... 21 The Implications of a Dry Climate for the Paleoecology of the Morrison Formation Engelmann, Chure and Fiorillo 2004 ...... 21 Regional Paleohydrologic and Paleoclimatic Settings of Wetland/Lacustrine Depositional Systems in the Morrison Formation, Western Interior, USA. Dunagan and Turner, 2004 ...... 23 Jurassic “Savannah” – Plant Taphonomy and Climate of the Morrison Formation Parrish, T., Peterson, F., and Tu rner, C. 2004...... 25 Reconstruction of the Upper Jurassic Morrison Formation extinct ecosystem – a synthesis Turner, C., and Pete rson, F. 2004...... 28 Vertebrate Biostratigraphy of the Morrison Formation Near Cañon City, Colorado, Carpenter, K. 1998...... 33 Vertebrate Microfossil Sites and Their Contribution to Studies of Paleoecology. Brinkman, D., Russell, A., and Peng, J. 2005...... 36 Documentation ...... 37 Bonebed diversity: ...... 37 Matrix ...... 38
FAUNAL LIST ...... 40 STATISTICAL ANALYSIS ...... 52
NAVIGATING MATRICES ...... 53 MATRIX DATA TRANSFORMATIONS ...... 54 LOG TRANSFORMATION ...... 54 PERCENT TRANSFORMATION ...... 55 SIMILARITY COEFFICIENTS ...... 56 DENDROGRAMS WPGMA VS. UPGMA ...... 57 ANALYSIS OF THE UPPER JURASSIC MORRISON FORMATION ...... 59 THE MATRIX ...... 59 DATA TRANSFORMATIONS ...... 61 SIMILARITY COEFFICIENTS ...... 61 DENDROGRAMS ...... 62 CONCLUSIONS ...... 69
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POLAR ORDINATION ...... 70
DESCRIPTION OF TECHNIQUE ...... 70 PROJECTIONS ...... 70 CHOOSING ENDPOINTS ...... 72 INTERPRETATION ...... 73 TRANSFORMATIONS ...... 76 MORRISON ORDINATIONS ...... 76 PRINCIPAL COMPONENTS ANALYSIS OF THE MORRISON ...... 85
ANALYSIS OF THE MORRISON FORMATION ...... 85 PRINCIPAL COMPONENTS ANALYSIS -DESCRIPTION OF TECHNIQUE ...... 85 SUMMARY ...... 95
FAUNAL ANALYSIS AND PALEOENVIRONMENT OF THE AARON SCOTT SITE ...... 95 MULTIVARIATE ANALYSIS OF MORRISON FAUNA ...... 96 CONCLUSION AND FUTURE RESEARCH ...... 99 BIBLIOGRAPHY ...... 101 APPENDIX A – CINCINNATI MUSEUM CENTER – AARON SCOTT SITE SPECIMENS ...... 108 APPENDIX B – AARON SCOTT SITE FIELD NOTES & SPECIMEN IDENTIFICATION ...... 111 APPENDIX C – PALEOBIOLOGY DATABASE (PBDB) MORRISON SITE TERRESTRIAL TAXA ...... 131 APPENDIX D – ANNOTATED OCCURRENCE MATRIX OF THE MORRISON FORMATION ...... 132
Table of Figures
Figure 1: Simplified Stratigraphy of the Morrison Formation – Source: Kowallis et al. 1998. ... 7 Figure 2: Location of Aaron Scott Site – Source: Microsoft Virtual Earth ...... 12 Figure 3: View of the Aaron Scott Site ...... 12 Figure 4: Extent of the Morrison Formation. Source: Google Earth ...... 18 Figure 5: (From Carpenter, 1998) Two way Dendrogram of the top 10 most diverse vertebrate sites, and the 12 most common taxa...... 35 Figure 6: Ventral view of Glyptops neural bone, field # ASS 149...... 41 Figure 7: Lateral view of Opisthias right maxilla VP 8586...... 43 Figure 8: View of several goniopholid teeth, CMC VP # 8554...... 44 Figure 9: Anterior Allosaurus tooth in labial view...... 45 Figure 10: Coelurus 3rd metatarsal in lateral view...... 46 Figure 11: Dorsal view of Diplodocus chevron CMC VP 7675...... 48 Figure 12: Lingual view of left Dryosaurus dentary field number ASS 133...... 50 Figure 13: Left lateral view of Stegosaurus mid-dorsal vertebra CMC VP 8551...... 51 Figure 14: Simple two-dimensional matrix X...... 53 Figure 15: Matrix X, a simple 4 x 5 matrix...... 54 Figure 16: Transformed matrix...... 55 Figure 17: A 4 x 5 matrix...... 55 Figure 18: Percent transformed matrix...... 56 Figure 19: Dice matrix generated by the data in Figure 15...... 56 Figure 20: Similarity matrix from typical Dice matrix – Step 1...... 57
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Figure 21: Similarity matrix – Step 2...... 57 Figure 22: Similarity matrix – Step 3...... 57 Figure 23: Similarity matrix – Step 4...... 58 Figure 24: Similarity matrix – Step 5...... 58 Figure 25: A dendrogram for the data in Figure 20...... 58 Figure 26: Matrix of the Morrison...... 60 Figure 27: Q-mode dendrogram of all sites and taxa (inset enlarged) ...... 62 Figure 28: Q-Mode dendrogram of all taxa...... 64 Figure 29: Two way dendrogram (all non-empty sites and taxa, truncated) with two or more occurrences ...... 65 Figure 30: Top 22 Sites, Top 20 Taxa Q-mode...... 66 Figure 31: Top 20 Genera, Top 22 Sites R-mode...... 67 Figure 32: Top 22 Sites, Top 20 Genera 2-Way...... 69 Figure 33: A small, Q-mode matrix...... 70 Figure 34: Sites in two- dimensional space...... 71 Figure 35: Sites in one-dimensional space...... 71 Figure 36: Non-perpendicular axes...... 72 Figure 37: Outlier-based axes...... 73
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Introduction
The Aaron Scott Site Dinosaur Quarry (Quarry) is a major new Jurassic vertebrate locality in the Morrison Formation of Utah that provides a strong educational and outreach resource for undergraduate institutions in Ohio and Kentucky. Since its discovery in 2001, the
Quarry has gained importance in its paleontology and stratigraphic context, due in part to the richness and diversity of its taxa. The Quarry provides abundant and robust paleoecological, taphonomic, paleolimnologic, fluviodeltaic sedimentological, sequence stratigraphic, and paleoclimatic data on the Morrison Formation’s depositional setting. Given the unique depositional environment at the Quarry, it would be reasonable to suppose that fauna there was distinct from the rest of the Morrison. This paper tests that hypothesis, examining the Quarry’s faunal assemblage in the context of the Morrison Formation as a whole. In doing so, this paper also tests the long-held hypothesis that the ancient Morrison Basin was environmentally and ecologically homogenous.
A collaborative team of paleontologists and stratigraphers from Cincinnati Museum
Center, the University of Cincinnati, Marietta College, and Northern Kentucky University continues research at the Quarry that includes vertebrate excavation, preparation, and study of the fossils, and stratigraphic, sedimentologic and geochemical studies of the enclosing package of sediments. The team consists of faculty with experience in paleontology and stratigraphy who lead a diverse group of undergraduates, graduate students and volunteers.
The Quarry is of significance in large part because the abundant accumulation of vertebrate fossils – particularly “microvertebrates” – provides insights about its depositional setting and faunal history within the Morrison Formation. Traditionally known as a river and
1 Esker 1/26/2009 floodplain environment, the Morrison Formation is also widely known for its rich dinosaur accumulations (Turner and Peterson, 2004). Large dinosaur fossils in the Quarry include a variety of sauropod bones and isolated elements of Allosaurus, but this site also abounds in microvertebrates, including several sphenodontid mandibles.
Discoveries in the Quarry hint at the potential to produce Jurassic mammal fossils, although to date, none has been found. Very little is known of Jurassic mammals due to their rarity and the paucity of fossils in the Morrison Formation, but the high frequency of microvertebrates at this site is indicative of a significant potential to produce mammalian and associated fauna. In order to fully understand the environment in which these animals lived and died, the enclosing strata are being analyzed, stratigraphically and geochemically.
Marietta College and Northern Kentucky University are undergraduate institutions that use the Quarry in their respective public outreach programs, making this site of great value for basic research and science education by involving undergraduate students in the research process
(field work, lab work, and presentation of the results).
Broader Scientific Impacts Ongoing work has demonstrated that the sediments enclosing the Quarry assemblage are the consequence of fluvio-deltaic processes resulting in infilling of a lake basin (Jeffery Bertog, and Bishop, 2005). The strata display sequence stratigraphic relationships caused by interactions between sediment supply and fluctuations in lake water volume. The fluctuations in water volume are interpreted to have resulted from severe drought conditions that would have driven animals to congregate around large, permanent lakes, thus resulting in the rich accumulation at the Quarry. These interpretations are possible because of the excellent exposure of the bone beds and the enclosing sediments that help put the deposit into a more profound depositional context.
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The Morrison Formation is traditionally considered a meandering river floodplain environment (Turner and Peterson, 2004). This formation is known for its dinosaur population and is the most dinosaur-rich formation in the United States (Turner and Peterson, 1999). The
Quarry contains fragments of typical Jurassic dinosaurs – especially Diplodocus, Barosaurus,
Stegosaurus and Apatosaurus as well as Allosaurus – but among these typical, Morrison
Formation dinosaurs are numerous (but less well known) microvertebrates. The abundance of these microvertebrates is a strong indicator that the Quarry may also contain early mammal remains (Bertog, et al., 2005). While Jurassic mammals have been found in the Morrison
Formation, their numbers have been relatively few; thus very little is known about these animals.
Continued research at this locality may yet reveal important information about population size, diversity and development during this early period in mammalian history.
In order to understand the preservation of these microvertebrates and the environment where these animals lived, research at the Quarry includes sedimentologic and taphonomic analysis. Detailed stratigraphy is being measured and analyzed (Jeffery, Bertog, and Bishop,
2005) to delineate the Quarry within a broader context. On a finer scale, as fossils are excavated, they are mapped using traditional grid methods and a Nokia laser transit. The resulting distribution grid is used to create a 3-dimensional map of the Quarry. Bone orientation is also measured to discern preferred orientation that would be suggestive of reworking of the bones by current (Bertog et al., 2005). Bone orientation and distribution relative to the stratigraphic relationships of the enclosing strata, can provide insights into where and why they were preserved (Jeffery, Bertog, and Bishop, 2005; Bertog et al., 2005).
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The factors responsible for the distribution and concentration of rich terrestrial vertebrate fossil deposits are fundamentally tied to the biology of the organisms and to the climate and landscape of their environment. The death assemblage and the sediments supply proxy evidence for biology, climate, and landscape as recorded in the relationships between the taphonomy and the stratigraphic architecture of the accumulation. Distinctive fossil localities afford a well exposed view of stratal relationships of the bone beds and their enclosing sediments and thus contribute compelling and substantial evidence about their paleoecology, paleogeography, and paleoclimate. While skeletal relationships of fluvial bone bed accumulations have been well documented, lacustrine bone beds that display sequence stratigraphic relationships are rare or possibly, unrecognized. Thus, the Quarry offers a distinctively promising opportunity to study these relationships.
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Background
Paleontology The site was discovered in 2001, within the lower portion of the Brushy Basin Member of the Upper Jurassic Morrison Formation and contains a rich assemblage of vertebrates and invertebrates (Jeffery, Bertog, and Bishop, 2004; Jeffery, Bertog, and Bishop, 2005; Bertog et al., 2005). The Quarry is located in Emery County, Utah, southeast of the town of Ferron.
The Morrison Formation crops out within the Western Interior region of the United States and Canada and extends south to central New Mexico (Turner and Peterson, 2004). The exposures of east-central Utah lie within the Colorado Plateau region, where it generally consists of flat-lying beds of sedimentary rock including variegated siltstones, shales, and sandstones
(Craig, 1955). The Quarry is located on the western margin of the San Rafael Swell, a NE- trending broad, asymmetrical Laramide upwarp roughly 75 miles long and 30 miles wide with a steeply dipping eastern margin and a gently dipping western margin (Lawton et al., 1983, Stokes,
1986). The landscape at the Quarry consists of numerous cuestas, dipping westward on the order of 11 degrees. It is difficult to differentiate what proportion of the dip is depositional, but most is probably tectonic.
Hundreds of bones from the Quarry have been collected with preliminary identification of vertebrate taxa including fragments of a sauropod skull (possibly Barosaurus), articulated sauropod vertebrae and other bones from one or more individuals, bones of at least two
Allosaurus individuals (based on size classes of pedal phalanges, one individual is a very small specimen, less than 1 meter tall), numerous teeth – including a broken-off tooth penetrating a bone, numerous bones belonging to Stegosaurus, crocodiles, turtles, and small ornithopods
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(possibly Dryosaurus), a large collection of as yet unidentified small bones and teeth, a partial skull and several lower jaws of sphenodontids.
The bones are in poor to good condition, black, and fragmentary to complete. The bone horizon consists of calcite-cemented, very light gray lithoclastic silty sandstone overlain by a medium gray, heavily bioturbated silty mudstone with abundant bivalves of the freshwater genus
Unio.
The Morrison Formation contains an extremely well known assemblage of Jurassic vertebrates that has been extensively studied since the late 1800’s (Foster, 2003). Recent individual articles and article collections provide prolific detail and summary of the sedimentology, stratigraphy, paleontology, and history of this vast dinosaur resource (Carpenter et al. 1998; Gillette, 1999; Foster, 2003; Turner and Peterson, 2004). Numerous studies focus on dinosaur assemblages within individual quarry localities (e.g., Bilbey, 1998; Richmond and
Morris, 1998; Gates, 2005) or summarize published quarry data from the rich Morrison literature
(Foster, 2003). Turner and Peterson, (2004) introduced a series of papers, each of which is a synthesis of the “state of the paleoecology of the Morrison” in terms of the sub-disciplines of sedimentology and paleontology.
Stratigraphy The Morrison Formation in east-central Utah has been subdivided into three members: the Tidwell, Salt Wash, and Brushy Basin. These units are described in great detail by numerous authors (Dodson et al., 1980; Peterson and Turner-Peterson, 1987) and in the research notes and references on the nearby Cleveland Lloyd Dinosaur Quarry (Bilbey, 1998; Gates, 2005). The
Tidwell Member contains mudstones, limestones, and evaporites that are interpreted as mudflat,
6 Esker 1/26/2009 playa, and sabkha environments and is the transition from the marine and marginal marine sediments of the underlying Summerville Formation (Turner and Peterson, 2004). The Salt
Wash Member consists primarily of coarse fluvial sands and lake and floodplain mudstones
(Peterson and Turner-Peterson, 1987; Gates, 2005). The Brushy Basin Member near the Quarry consists of a lower gray lake unit that interfingers with the Salt Wash Member and an upper red, gray, and green banded unit (Figure 1).
Figure 1: Simplified Stratigraphy of the Morrison Formation – Source: Kowallis et al. 1998.
At the Quarry, the apparent thickness of the interval between the Summerville Formation and the upper Brushy Basin Member of the Morrison (that is, the thickness of the lower Brushy
Basin Member) may be 30-50 m, but this estimate is problematic because the beds of the
Tidwell, Salt Wash, and lower Brushy Basin members are westward off-lapping wedges that crop out in a broad, north-south belt (Bertog and Jeffery, 2005). The lower Brushy Basin
Member consists primarily of gray mudstones with interbedded, thin limestone and discontinuous sands.
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The upper Brushy Basin Member is up to 175 m thick and predominantly contains red, green, and gray mudstones giving it distinctive horizontal banding with minor discontinuous epsilon cross-bedded sand lenses. This unit has largely been interpreted as a meandering river and floodplain deposit (Gates, 2005). The distinctive red coloring of the upper unit is primarily due to the abundant paleosols contained therein (Demko et al., 2004). The Quarry lies in the upper portions of the gray mudstones of the lower Brushy Basin Member that is interpreted to have been laid down in a lacustrine depositional environment.
The Morrison ecosystem has been interpreted as a seasonally arid plain, dominated by meandering river systems and ephemeral lakes (Dodson et al., 1980; Parrish, Peterson, and
Turner,, 2004, Turner and Peterson, 2004; Gates, 2005). Turner and Fishman (1991) cite the presence of a large, alkaline, saline lake that covered a large portion of the Colorado Plateau, primarily east of the Four Corners area during late Kimmeridgian time. More recent interpretations cite evidence that the large lake may have been more of a wetland/lacustrine complex (Dunagan and Turner, 2004).
Lake sediments in the area of the Quarry are far to the west of the large lake cited by
Turner and Fishman. They lie directly above and interfinger with the middle Kimmeridgian Salt
Wash Member (Turner and Peterson, 2004). The lake sediments enclosing the Quarry lie within the depositional area of the Tidwell Member in east-central Utah, interpreted as a small
“embayment” in the Jurassic Epeiric Seaway during the early Kimmeridgian by Turner and
Peterson (2004). Once this early Kimmeridgian sea withdrew, the “embayment” could have become a lake basin that progressively freshened and filled during the middle Kimmeridgian.
Stratal geometries, cyclical stacking patterns and detailed taphonomic analysis of the
Quarry in the lower Brushy Basin Member of the Morrison Formation can be used to infer
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sequence stratigraphic relationships and reveal that sediments were deposited during the
progressive fill of a large lake, accumulating within a prograding fluvio-deltaic lacustrine system
(Jeffery, Bertog, and Bishop, 2005; Bertog et al., 2006; Bertog et al., 2005). Sequence
stratigraphic relationships have been well documented in numerous studies dealing with aspects
of lacustrine and related strata in terms of petroleum source rock deposition and stratal
geometries of progressive lake basin fill (e.g., Keighley et al., 2003; Carroll and Bohacs, 2001;
Oviatt, McCoy, and Nash, 1994; Dam and Surlyk, 1992).
Observations from both stratigraphic and taphonomic analyses in this study support the
hypothesis that the deposit is within lake-margin sands, probably at a beach or distributary bar
and that variations in water volume within the lake caused the changes in base level (Jeffery,
Bertog, and Bishop, 2005). The variations in base level are responsible for the preservation of an
assemblage that is interpreted to have accumulated as the result of a brief lowering and
subsequent rise of lake level during the late stages of an overall fall in base level (late highstand),
but prior to the formation of the next distinct cycle boundary.
The brief lowering in base level is believed to have been the result of temporary drought
conditions that eliminated smaller watering holes. The presence of the perennial lake resulted in
a high rate of dinosaur and other vertebrate traffic at the margins of the largest regional body of
water1. This situation likely lasted for the duration of the drought. Such brief fallen
parasequences or drought-forced regression episodes are conducive to preservation of rich
assemblages because of the high animal traffic and the rapid submergence and burial at the end
of each drought. Evidence for this can be seen in the varying quality of preservation of fossils at
1 Further evidence for the Quarry representing a drought forced lake level drop is the absence of obligate aquatic taxa from within the dinosaur bearing layer. It seems likely that investigating the beds above and below the bonebed will yield substantial fish and amphibian