THE UNIVERSITY OF CALGARY Eric Snively A ... - Ohio University

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ABST RACT The arctometatarsus is a metapodial structure exclusive to Cret- PC~OUS melurosaurian dinosaurs. The central third metatarsal (MT III) of the arctometatarsus is proximally constricted between the second and fourth (MT II and MT IV), MT III is also triangular in distal cross section, with the apex towards the plantar surface. Descriptive morphology and Principal Components Analysis (PCA) segregate the arctometatarsus from other morphologies, and suggest hypotheses of function for tyrannosaurid metatarsi and similar forms. Through CT analysis of metatarsal shape, physical manipulation of casts, assessrnent of osteological correlates of ligaments, and camparison with the equid carpus, two hypotheses are evaluated for the tyrannosaurid arctometatarsus. First, ligament anatomy provides a rnechanism for a previously proposed hypothesis, that axial Iocomotor energies were transferred from MT III to the outer elements. Second, the tensional keystone hypothesis holds that upon angled footfalls during linear locomotion, distal intermetatarsal figaments would prevent anterodorsôl displacement of MT III, and unrfy the metapodium. Finite element analysis of strain energy in Gorgosaurus libratus metatarsals strongly supports the energy transference hypothesis, and indiredy supports the tensional keystone hypothesis. From the perspective of this evaluation of the tyrannosaurid arctometatarsus, functional hypotheses are proposed for the metatarsi of other theropods. Mapping the arctometatarsus ont0 two phylogenies of theropods suggests a tentative sœnario for the structure's evolution and biological rote. iii
Page 1 and 2: THE UNIVERSITY OF CALGARY Functiona
Page 3: Many figures in this thesis use col
Page 7 and 8: my splendid students provided neces
Page 9 and 10: General approach and hypotheses MAT
Page 11 and 12: CHAPTER 4: Finite elernent mode1 of
Page 13 and 14: Table 2.1. Theropod and Plateosauru
Page 15 and 16: Figure 2.1 1. Notable features of l
Page 17 and 18: metatarsus. Figure 4.4. Simplified
Page 19 and 20: CHAPTER 1: INTRODUCTION Animais as
Page 21 and 22: Completing the system in fossil ani
Page 23 and 24: The arctometatarsus as an example o
Page 25 and 26: Figure 1.1 shows a phylogeny of the
Page 27: coelurosaurs with an enlarged trenc
Page 30 and 31: the animals ran on uneven ground. T
Page 32 and 33: an evolutionary wntext, in an effor
Page 34 and 35: Camosauria 3 4, Ornitholestes 4 Dro
Page 36 and 37: Prosauropoda Hemrasaurus Elap hrosa
Page 38 and 39: L Prosauropoda Henerasaunrs Dromaeo
Page 40 and 41: Dromaeosaurida Aves - Therizhosauri
Page 43 and 44: CHAPTER 2: Descriptive and quantita
Page 45 and 46: definition of the arctometatarsus h
Page 47 and 48: 1995), can informatively map specim
Page 49 and 50: about clustering by metatarsal morp
Page 51 and 52: Specimenl Taxon (arctometatanus*) S
Page 53 and 54: c) Region of proximal articulation
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measurements are shown in Table 2.3
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Table 2.2. Measurements of theropod
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Table 2.3. Log-transforms of measur
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The distal intermetatarsal articula
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III, which has a pronounced medial
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c) Region of proximal articulation.
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depression. The entire ginglymus is
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contours seen in Ornithalestes. The
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has a very shallow U-shape. The sha
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PC3 1.026%. Other wmponents contrib
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Table 2.4. Loading, variance, and c
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positively with relative metatarsal
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Table 2.5. Statistics for specimens
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Troodon, with the most gracile meta
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metatarsals, including those of tyr
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have PC2 values above 0.22, substan
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qualitative rnetatarsal data for He
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and detailed hypotheses for the tyr
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a) There is no substitute for morph
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Figure 2.1. Specimens from figures
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Figure 2.3. Oviraptorosaur specimen
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Figure 2.5. Relative directional an
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Figure 2.6. Template for PCA measur
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egion of proximal articulations med
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proximal hook: lateral defiection p
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medial deflection proximal edge of
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vertical MT II articulation - proxi
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proximal edge- of gingly I - nus pr
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egion of proximal articulations pro
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eg ion of proximal proximal expansi
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egion of proximal articulations - r
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Component I Score
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Figure 2.17. Plot of third metatars
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Figure 2.1 8. Plot of third metatar
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Component 1 Score
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locomotor efficiency and decreases
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surface that is smooth and slightly
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Taxon (arctometatarsus*) Specimen n
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proximal end. This was repeated wit
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evealed through observation and mea
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The metal frame caused diffraction
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RESULTS lntemetatarsal movement 1.
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tyrannosaurids (Figure 3.5), but ar
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Prox.: MT III-II:II P~ox.: MT III-I
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Kinematic mode1 of the tyrannosauri
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part of MT III and its ligaments as
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Tensionat keystone dynarnics may ex
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Chapter 2 aIso outlines metatarsus
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of other theropods, and yet allowed
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Figure 3.2. Freedom of intermetatar
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Figure 3.4. Freedom of intermetatar
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Figure 3.6. Osteological correlates
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Figure 3.8. Osteological correlates
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Figure 3.10. Step sequence of Gorgo
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Figure 3.12. CT reconstructions of
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Figure 3.14. Torsional loading tran
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Lu- dorsal 81igarnents
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CHAPTER 4: Finite element model of
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forces are applied to nodes on the
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Material and Ioading regimes of fhe
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Two analyses were fun in order to t
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Dunng Iinear progression when an an
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This quantity approximates the vert
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to its right foot to prepare for a
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elements in the finite element rnes
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Table 4.1. Material properties of b
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Photoshop 3 for Macintosh, to ensur
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this program. NUAGES typically foun
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1) Informative strain results versu
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of strain energy evident in Figure
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Specifically, osteological correlat
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custiioned the rnetatarsals against
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Figure 4.2. Initial loads and bound
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Figure 4.4. Simplified representati
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Figure 4.6. Strain distribution in
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The analyses of the Gorgosaurus Iib
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individual organs with ideally effi
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These considerations of ligament mo
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The precise scaling of metatarsal l
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tenable from the available evidence
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equivalent horsepower, 15 economy c
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Other aptations refiect multiple fu
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This and other systernatically info
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(Russell 1975, McGregor 2000). Meta
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was the default developmental patte
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Arnong dromaeosaurids and birds, th
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hypothesis. Fossil evidence of basa
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suggestive of grasping ability. Neo
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available evidence without propriet
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Figure 5.1. Phylogeny of the Therop
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LITERTURE ClTED Alexander R.M. 1977
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Carter D.R., Fyhrie, D.P., and Whal
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Galton P.M. 1982. Elaphrosaurus, an
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Hutchinson J.R., Stidham T.A., Sniv
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Novas F.E. 1993. New information on
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Stein B.R. and Casinos 1997. What i
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APPENDIX Principal Components Analy
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highest degree, and so on. Specimen
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THE UNIVERSITY OF CALGARY Eric Snively A ... - Ohio University

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