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

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allowed to stretch slightly under the bending loads, stresses on the bone will decrease. In addition, elastic deformation and remil of tendons would decrease the rate of strain on the bone; unlike tendons and ligaments, bone is less brittle under longer periods of loading. A longer collective loading period on tyrannosaurid hind limb elements, including soft and hard tissues, may have allowed their bones to withstand torque imposed by lower duty factors than nomially expected for animals of their great sire. While these relationships hold for mammals over wide range of body mass (Pollock 1991), benefits of elastic storage have yet to be quantified in tyrannosaurids. Based on regression equations from quadrupedal mammals (Pollock 1991 ), (1 1) ~=0.14m'.~~ U=elastic strain energy storage, m=body mass, the M. gastrocnemius tendon in G. librafus would be expected to store and release at least 910 Joules. Because tendons of M. flexor digitonim longus also span the mesotarsal joint, and because G.libratus is a biped, the maximum capacity for elastic energy storage of al1 tendons around the joint was almost certainly higher. The cross section of these tendons in ostriches is 1.95 times that predided for an antelope carrying the same amount of mass on its back legs (Alexander et al. 1979), and the tendons are much longer than those of ungulates. If the same were true for G. libratus versus a quadrupedal mammal of its mass, its mesotarsal tendons would potentially be quite efficient at elasüc starage. The tendons thus may have
custiioned the rnetatarsals against bending force incurred after moderately long suspended phases. CONCLUDING COMMENTS AND FUTURE DIRECTIONS While elastic storage by tendons and ligaments may have enhanced the locomotor performance of tyrannosaurids, the preceding considerations do not support the 70 kmihr speeds postulated by Bakker (1986) and Paul (1 988). These speeds would require lower duty factors or higher stride frequencies than are probably realistic for such long-legged, heavy animals (Christiansen 2000). However, the elastic properties of ligaments and tendons should be incorporated into future studies, including investigations expanding upon the finite element analyses presented in this chapter. Without consideration of elastic wnnective tissues, attempts to set rigorous upper and lower bounds on tyrannosaurid capabilities will be inadequate.
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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
Page 171: Figure 3.7. Osteological correlates
Page 175: Figure 3.9. Osteological correlates
Page 179: Figure 3.1 1 ae. The metatarsal rec
Page 187: Figure 3.13. Ligament contribution
Page 191 and 192: Figure 3.15. Anatorny of the equid
Page 193: Figure 3.16. Cornparison of loading
Page 196 and 197: (nodes), and solving stresslstrain
Page 198 and 199: The practicality of the finite elem
Page 200 and 201: collective density of trabeculae (K
Page 202 and 203: A FORCE INPUTS AND MATERIAL PROPERT
Page 204 and 205: Alexander et al. (1 979) rewrded a
Page 206 and 207: The preceding loading regime entait
Page 208 and 209: detemined by using the photographic
Page 210 and 211: 2. Preparation of data for contour
Page 212 and 213: . - Elastic modulus (GP4 Ex EY Ez P
Page 214 and 215: conversion. Volume-to-nuages facili
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Page 218 and 219: of MT III (Figure 4.4, pinpointed b
Page 220 and 221: anterodorsal rotation of the distal
Page 224: Figure 4.1. Initial loads and bound
Page 228: Figure 4.3. Finite element mesh of
Page 232: Figure 4.5. Strain distribution in
Page 236 and 237: CHAPTER 5: Mechanical and evolution
Page 238 and 239: y distal intermetatarsal ligaments,
Page 240 and 241: Probable fundion of the tyrannosaur
Page 242 and 243: Ornitholestes, and camosaurs) incre
Page 244 and 245: morphology matches in degree the co
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Page 248 and 249: face of the skull met part of the m
Page 250 and 251: Alternately, a morphological novelt
Page 252 and 253: outgroup to ail theropods, and the
Page 254 and 255: convincingly evident in figures of
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Page 258 and 259: dromaeosaurids], and a dade compris
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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 271: Figure 5.4. Phylogeny of the Therop
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Figure 5.6. Phylogeny of the Therop
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Bock W.J. and von Wahlert G. Evolut
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Cume P.J. 1997. Theropoda. In: Fari
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Grenard S. 1991. Handbook of Alliga
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Liem K.F. and Osse J.W.M. 1975. Bio
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Richtsmeier J.T. and Cheverud J.M.
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Woo S., Maynard J., Butler O. 1987.
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Different variables in a PCA will h
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THE UNIVERSITY OF CALGARY Eric Snively A ... - Ohio University

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