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convincingly evident in figures of Allosaurus fiagilis elements (Madsen 1976). These metacarpals articulate tightly with one another, much like the proximal portions of theropod metatarsals. While the manus and pes of theropods are disjunct functionally (Gatesey and Middleton 1997), developmental conespondence predicts that if osteological indications of interrnetatarsal ligaments are present, correlates of intermetacarpal ligaments woiild not be surprising. However, identification of correlates as ligament scars on either manus or pes is uncertain. Arguments for iteratively associated intrametapodial ligaments are therefore circular, without independent evidence for ligaments in either metapodium. By phylogenetic inference (Bryant and Russell 1992) or bracketing (Bryant and Russell 1992, Witmer 1995), we can deduce the presence of interrnetatarsal ligaments in taxa intervening between carnosaurs and tyrannosaurs, and perhaps in Mesozoic theropods prirnitively (Figs. 5.1 and 5.2). The distribution of osteological ligament correlates in both phylogenies indicates that welurosaurs inheflted them from the common ancestor of al1 saurischians, or from that of tyrannosaurs and carnosaurs (Figs. 5.1 and 5.2). The postulate emerges that proximal ligaments were a necessary step before the acquisition of distal ligaments, which would bind the metatarsals where their distal plantar angulation occurs. Osteological carrelates in the arctometatarsus indicate a disjunct ontogeny for ligaments, in which strong ligaments developed proximally and distally, but not in the intermediate region. Further testing will reveai whether or not this apparent ligament ontogeny of the arctometatarsus
was the default developmental pattern for coelurosaurs, or how men it became expressed or lost in various taxa. How offen did the arctometatarsus evolve? The presented phylogenies conflict in regard ta how often the arctometatarsus emerged, and in the distribution and inclusiveness of arctometatarsalian clades. The simplest hypothesis of character evolution usually attrads provisional acceptance, because it entails the fewest evolutionary steps (acquisitions and reversais). However, the most parsimonious scenario is not necessarily correct. The broad distribution of the arctometatarsus in competing phylogenies suggests that the simplest and other scenarios be tested with independent character evidence. The lsvels of parsimony of the Sereno (1 999) and Holtz (2000) phylogenies cannot be directiy compared. The most econornical resulting cladograms for each analysis are internally consistent with their respective data sets, and are the most parsimonious solutions given the taxa and characters included in that study. Nevertheless, if one analysis resuits in fewer steps for the emergence of a given character, its results may be interpreted provisionally as the sirnpler evolutionary explanation for that character. The arctometatarsus is the relevant subject of character evolution here, and several patterns of loss and emergence were deduced from its occurrence on both phylogenies. These patterns divulge which phylogeny presents the simplest hypothesis of arctometatarsus evolution. Because the simplest explanation does not definitively refi ect ttie true evolutionary pattern, at minimum the first two most
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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
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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
Page 204 and 205: Alexander et al. (1 979) rewrded a
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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
Page 260 and 261: arctometatarsalian forrns. The conv
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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