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

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morphology matches in degree the condition in tyrannosaurids. These morphologies probably correlate with a strong MT II-MT III articulation in life. Deinonychus has a large, trenchant ungual phalanx on digit II. The strong interrnetatarsal articulation may have resisted torsional forces when the animal employed this weapon during predation (Ostrom 1969). 4) Carnosauria: Sinraptor dong? Unlike in the Allosaurus specimens, the Sinmptor MT III (Figure 2.14) shows some plantar angulation along its articular surface with MT II. As with Deinonychus, there is also a more extensive distal articular facet with MT II. This indicates that the MT II-MT III articulation was potentially stronger in Sinraptor than in Allosaurus. Interestingly, recent examination of MT III of a unnarned giant South Arnerican carnosaur (Carcharodontosauridae, n. gen., n. spec: Plaza Huincul specimen;TMP loan) reveals a discrete area of distal rugosity on the distal articular surface with MT II. MT II of this specirnen is unavailable, so it cannot be deterrnined if it bears a rugosîty corresponding to that on MT III. However, if the roughened surface on MT III of this animal is a ligament scar and not a pathology, these morphologies indicate a diversity of MT II-MT III articulation mechanisms and strength among large camosaurs. The functional implications of this diversity are undear. Sinraptor and large Allosaurus specimens were of similar mass (Paul 1988, 1997), wtiile the carcharodontosaurid was much larger (Currie: pers. comm. 2000). Speculations on whether stronger articulations correlateci with size or activity level are not
tenable from the available evidence. However, graaIity of the MT III of Sinraptor (UUVP10600) may indicate immaturity (Cum'e: pers. comm. 2000). Theropod metatarsals scale negatively with femur Iength during ontogeny (Cum'e 2000). An arctometatarsus occurs in coelurosaurs of a broad range of adult sizes (Figure 1.5, Chapter 1; Holtz 1994a,b), induding tyrannosaurids comparable in size to giant camosaurs. Therefore aspects of their morphology other than size must be considered in examination of the arctometatarsus' biological role. The following section explores the possible phytogenetic and selective ramifications of the arctometatarsus, with the avemding caveat that hypotheses of biological role are diffïcult to test adequately. EVOLUTIONARY CONTUCT FOR <strong>THE</strong> ARCTOMETATARSUS This thesis tests the hypothesis that the tyrannosaurid arctometatarsus irnparted benefits ta foot resilience and strength. These advantages have been mechanically conoborated, but their oflgin and utility require the contexts of phylogeny and historical aptation (traditionally adaptation). In the following section, I consider the arctometatarsus from an evolutionary perspective. The discussion begins with an explanation of aptational teminology. With this background in place, I then use recently published cladograms as baseline distributions of relevant charaders and taxa. From there I proceed into more complex evolutionary hypotheses (scenarios) based upon the pertinent cladistic hypotheses and the morphologicat evidence presented in this thesis. This
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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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Figure 3.15. Anatorny of the equid
Page 193: Figure 3.16. Cornparison of loading
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Page 198 and 199: The practicality of the finite elem
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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
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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 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
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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
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Page 258 and 259: dromaeosaurids], and a dade compris
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Page 262 and 263: Proximal intermetatarsal ligaments
Page 264 and 265: witti a surfeit of potentially supp
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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