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Ornitholestes, and camosaurs) increased ligament cross section, and also prevented proximal anteroposterior displacement of this element. Peculiarities of MT III morphology invoke more specific hypothses for several taxa. These hypotheses largely trace the succession of descriptions presented in Chapter 2. Taxa with particularly noteworthy features are discussed, in wmparison with tyrannosaurids and other forms. 1 ) cf. Omithomimidae and Troodon formosus. These MT III specimens are dassically arctometatarsalian. Proximally some degree of parasagittal rotation of this element may have been possible in either forrn (Wilson and Currie 1985), but because the animals were probably low in mass relative to the cross section of ligaments, little displacement was Iikely (Pollock 1991 ). The distal morphology of the ornithomirnid MT III (Figure 2.9) was so closely akin to that of tyrannosaurids that fundion in this region was probably similar to that of the larger forms, although scaling variance must be considered. The modest size of the specirnen measured in this study indicates an omithomimid of relatively low mass. Presurnably this mass imposed lower elastic displacements of distal ligaments than those likely in adult tyrannosaurids (Pollock 1991). This proportionality of ligament strain probably applied to adults of both clades; adult ornithomirnids were generally smaller than adult tyrannosaurids. Small ligament elastic displacements in large omithomimids were probably paralleled in very young tyrannosaurids, whose mas, limb proportions, and presumed running performance were very similar to those of adult omithomimids (Cume 2000).
The precise scaling of metatarsal ligament cross sections between omithomimids, tyrannosaurids, and troodontids has yet to be investigated. The morphology of buttressing surfaces in the T don metatarsus (Figure 2.10) was the mirror image of the condition in omithomimids and tyrannosaurids, in which footfall energies were transferred from MT III to MT II. Instead, with Troodon long axis compressive forces on MT III would be transferred mainly to MT IV. Its buttressing surface for MT III was more acutely angled to the substrate than that of MT II, and would have ken loaded by the component of the ground- reaction force parallel to the long axis of the metatarsus. In contrast, the MT II- MT III contact is in a parasagittal plane. In Traodon, MT II is shorter than MT IV, and bears a retractile ungual phalanx presumably held clear of the ground; it is unlikely that the second pedal digit contnbuted to weight bearing. 2) Oviraptorosauria: Hmisaurus sp. MT III of this specimen shows proximal constriction on its anterior surface, but plantar constriction is restrided to the distal portion of the metatarsal (Figure 2.1 1). As in other elmisaurids (Currie and Russell 1988), the metatarsals are also fused proximally. Compressive energy would transfer from the distal third metatarsal to the proximal ankylosis of the bones, and would not be transmitted by ligaments to the astragalar condyles, as was likely in tyrannosaurids (Chapter 3, Holtz 19944. 3) Deinonychus anfinhopus. MT III of Deinonychus displays a sligM plantar angulation on its media1 surface, and a distally extensive articular facet for MT II (Figure 2.12). Neither
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THE UNIVERSITY OF CALGARY Functiona
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Many figures in this thesis use col
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ACKNOWLEûGEM ENTS Many friends and
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Approval page Acknowledgements Tabi
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CONCLUDING COMMENTS Figures for Cha
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CHAPTER 5: Mechanical and phylogene
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LIST OF FIGURES Figure 1 .l. Phylog
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Figure 3.6. Osteological correlates
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AMNH GI HMN IVPP MOR NMC OMNH PIN P
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and constructional morphology revea
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1985, Bryant and Seymour 1990). If
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surfaces of a Tyrannosaurus rex spe
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3. Troodontidae (Figure 1.2): Trwdo
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Hypothesized functions of the tyran
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1. Atomization (Chapters 2 and 3).
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Figure 1.1. P hylogenetic diagram o
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Figure 1.2. Phylogenetic diagram of
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Figure 1 .S. Skeletal restorations
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did not quantify the degree of prox
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qualitative debate over arctometata
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This hypothesis focuses on tyrannos
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Table 2.1. Theropod and Plafeosauru
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specimens in the figures reflects a
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were not attempted. While this limi
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Chapter 3). 1 now address variation
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- Ç6'P 1 68'€tr 1 L'OC 99'66 L'9
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Table 2.3: Log-transformed values f
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experiences a sharp laterally indin
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ecause it can be cursorily dassifie
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arctometatarsalian specimens, the l
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) Shaf€ and region of proximal ar
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a) Ginglymus. Figure 2.14 shows the
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) Shaft and region of proximal arti
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measurements of width pV2S%TU-DE an
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Table 2.4b Variancelmeasurement eac
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the largest sum of linear measureme
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Table 2.5: Specimen statistics Herr
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and Sinraptor dongi specimens. With
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In addition, a hook like proximal a
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oth physical narrowing and elongati
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1s the MT 111 shape segregation fun
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tyrannosaurids, ornithomimid, and t
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proximal constriction, white EImisa
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Figure 2.2. Tyrannosaurid, trwdonti
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Figure 2.4. Dromaeosaurid and carno
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- Anterior (flexor) surface medial
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Figure 2.7. Left MT III of Tyrannos
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Figure 2.8. Left MT III of Gorgosau
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Figure 2.9. Left MT III of an omith
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Figure 2.10. Left metatarsus of Tmd
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Figure 2.1 1. Left MT III of Elmisa
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Figure 2.12. Right MT III of Deinon
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Figure 2.13. Left MT III of Allosau
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Figure 2.14. Left metatarsus of Sin
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Figure 2.15. Plots of MT Ill specir
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Figure 2.16. Plot of third metatars
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tv . ... . Ri. . . .
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Figure 2.19. Plot of third metatars
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CHAPTER 3: Tensile keystone model o
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dynamically transmitted Iommotor fo
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Table 3.1 . Metatarsi examined in t
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interrnetatarsal dynamics in these
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compelled analysis of movement evid
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Once prepared, TMP 94.1 2.602 and i
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Daspletosaurus tomsus (MOR 590), an
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2. Freedom of movement inferred fro
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Table 3.2. Surface areas of intemet
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phylogenetic bracketing Wtmer 1995)
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4) Forces from anterior displacerne
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e expected for tyrannosaurid interm
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The preceding discussion derives fr
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weapons sparring), and the arctomet
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Figure 3.1. Schematic representatio
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Figure 3.3. CT reconstruction of ri
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Figure 3.5. MT II (left element) an
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Figure 3.1 1 ae. The metatarsal rec
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Figure 3.13. Ligament contribution
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Page 202 and 203: A FORCE INPUTS AND MATERIAL PROPERT
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Page 232: Figure 4.5. Strain distribution in
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Page 238 and 239: y distal intermetatarsal ligaments,
Page 240 and 241: Probable fundion of the tyrannosaur
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Page 262 and 263: Proximal intermetatarsal ligaments
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Page 267: Figure 5.2. Phylogeny of the Therop
Page 278 and 279: Bock W.J. and von Wahlert G. Evolut
Page 280 and 281: Cume P.J. 1997. Theropoda. In: Fari
Page 282 and 283: Grenard S. 1991. Handbook of Alliga
Page 284 and 285: Liem K.F. and Osse J.W.M. 1975. Bio
Page 286 and 287: Richtsmeier J.T. and Cheverud J.M.
Page 288 and 289: Woo S., Maynard J., Butler O. 1987.
Page 290 and 291: Different variables in a PCA will h
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