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

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locomotor efficiency and decreases strain energy transmitted to bones (Pollock 1991 ). Adult tyrannosaurids were notably large, ranging from 2 to perhaps 8 tonnes (Paul 1988; cross scaling of measurements from fragmentary metatarsals: UCMP V91181). Extrapolating Rom research on living animals, it is reasonable to expect that certain characteristics of the limbs of large extinct animals will be suggestive of agility and high relative speed. In addition to a musculoskeletal configuration that promotes high out velocity at joints (Hildebrand 1988), it can also be predicted that the morphology of ligaments and bones will interact to promote effective dissipation and distribution of footfall forces. Because the arctometatarsus is proportionally long compared with primitive theropod metapodia, it would be fruitfui to compare other aspects of its morphology with extant systems that promote rapid progression. The imbricacy of the arctometatarsus in distal cross section (Holtz 1994a; this study) resembles the interdigitating wedge arrangement of horse carpals (Deane and Davies 1995). This suggests that interelement dynamics may be analogous in these phylogenetically disparate structures. Ligaments that connect horse carpals store and retum elastic strain energy (Figure 3.1), which may reduce the potential for injury to the wrist when the animal is mnning (Deane and Davies 1995). By compan'ng the results of detailed analysis of interrnetatarsal movement in tyrannosaurids with dowmented intercarpal kinematics of horse wrists, I tested the following hypothesis: Tyrannosaurid metatarsals and their ligaments
dynamically transmitted Iommotor forces in a manner similar to that seen in the equine carpus. Inference from comparative anatomy enables testing of the main assumption of this comparison, that ligaments sirnilar in composition and properties to those of modem animals were present between theropod metatarsals. Ligaments connect bones at syndesmotic joints in living vertebrates (Hildebrand 1988), and relevant articular surfaces on the metatarsals of large theropods should display the rugose scamng or discrete facets indicative of ligaments in living animals. However, the presence of ligaments cannot be declaimed a prion; ligament attachments may be confused with tendinous insertions, and aRicular cartilage rnay wver closely articulating joint surfaces and resuk in sculpturing of the bone surface. I therefore refer to potential soft tissue attachment sites on bone as osteological correlates, or simply conelates. There are three major correlates of arthrotogical soft tissue. Surfaces associated with articular cartilage are usually smooth, slightly raised, and ofteri occur at weight bearing hinge joints, bathed in synovial fluid within a joint capsule (Hidebrand 1988). Ligament or tendon fixation sites rnay be recognized on metatarsals by two pnmary correlates. Rugosity marks the location of Sharpey's fibers, su bperïosteal mineralized wllagen fibers wntinuous with fibers of the attaching ligament (Woo et al. 1987). Ligaments and tendons may also attach to bone by so-called direct insertions, through a gradient of ligament, fibrocartilage, mineratized fibrocartilage, and bone. Direct insertions are indicated on a bone
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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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Page 84 and 85: In addition, a hook like proximal a
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Page 88 and 89: 1s the MT 111 shape segregation fun
Page 90 and 91: tyrannosaurids, ornithomimid, and t
Page 92 and 93: proximal constriction, white EImisa
Page 95: Figure 2.2. Tyrannosaurid, trwdonti
Page 99: Figure 2.4. Dromaeosaurid and carno
Page 102 and 103: - Anterior (flexor) surface medial
Page 105 and 106: Figure 2.7. Left MT III of Tyrannos
Page 107 and 108: Figure 2.8. Left MT III of Gorgosau
Page 109 and 110: Figure 2.9. Left MT III of an omith
Page 111 and 112: Figure 2.10. Left metatarsus of Tmd
Page 113 and 114: Figure 2.1 1. Left MT III of Elmisa
Page 115 and 116: Figure 2.12. Right MT III of Deinon
Page 117 and 118: Figure 2.13. Left MT III of Allosau
Page 119 and 120: Figure 2.14. Left metatarsus of Sin
Page 121 and 122: Figure 2.15. Plots of MT Ill specir
Page 123: Figure 2.16. Plot of third metatars
Page 126 and 127: tv . ... . Ri. . . .
Page 129 and 130: Figure 2.19. Plot of third metatars
Page 131: CHAPTER 3: Tensile keystone model o
Page 135 and 136: Table 3.1 . Metatarsi examined in t
Page 137 and 138: interrnetatarsal dynamics in these
Page 139 and 140: compelled analysis of movement evid
Page 141 and 142: Once prepared, TMP 94.1 2.602 and i
Page 143 and 144: Daspletosaurus tomsus (MOR 590), an
Page 145 and 146: 2. Freedom of movement inferred fro
Page 147 and 148: Table 3.2. Surface areas of intemet
Page 149 and 150: phylogenetic bracketing Wtmer 1995)
Page 151 and 152: 4) Forces from anterior displacerne
Page 153 and 154: e expected for tyrannosaurid interm
Page 155 and 156: The preceding discussion derives fr
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Page 159: Figure 3.1. Schematic representatio
Page 163: Figure 3.3. CT reconstruction of ri
Page 167: 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
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Figure 3.13. Ligament contribution
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Figure 3.15. Anatorny of the equid
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Figure 3.16. Cornparison of loading
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(nodes), and solving stresslstrain
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The practicality of the finite elem
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collective density of trabeculae (K
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A FORCE INPUTS AND MATERIAL PROPERT
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Alexander et al. (1 979) rewrded a
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The preceding loading regime entait
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detemined by using the photographic
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2. Preparation of data for contour
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. - Elastic modulus (GP4 Ex EY Ez P
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conversion. Volume-to-nuages facili
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maintained. An angle of 2 degrees p
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of MT III (Figure 4.4, pinpointed b
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anterodorsal rotation of the distal
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allowed to stretch slightly under t
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Figure 4.1. Initial loads and bound
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Figure 4.3. Finite element mesh of
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Figure 4.5. Strain distribution in
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CHAPTER 5: Mechanical and evolution
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y distal intermetatarsal ligaments,
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Probable fundion of the tyrannosaur
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Ornitholestes, and camosaurs) incre
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morphology matches in degree the co
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discussion elucidates the evolution
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face of the skull met part of the m
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Alternately, a morphological novelt
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outgroup to ail theropods, and the
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convincingly evident in figures of
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parsimonious scenarios are outlined
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dromaeosaurids], and a dade compris
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arctometatarsalian forrns. The conv
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Proximal intermetatarsal ligaments
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witti a surfeit of potentially supp
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Figure 5.2. 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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