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dromaeosaurids], and a dade comprising ornithomirnids and Mononychus. The potential for arctometatarsus development would have first arisen in the common ancestor of Mononychus and tyrannosaurids (Figure 5.6). if his was the initial condition, the structure was wnvergently expressed in four instances: the clade comprising Mononychus, omithornirnosaurs, and their common ancestor, and separately in the dades Umisaunrs, Troodontidae, and Tyrannosauridae. If the arctometatarsus itselfwas present in the common ancestor of tyrannosaurids and Mononycbus, and passed on to the descendents of that ancestor, the phylogeny implicates four losses. Theriùnosaurs, Nomingia, dromaeosaurids, and Aves wouId have lost the structure. The association of Mononychus with omithomimids minimires the permutations involved in the occurrence of an arctometatarsus in that taon (see the preceding discussion). An alternative hypothesis, suggested by both phylogenies, is that the arctometatarsus is a synapomorphy of the clade comprising the common ancestor of Tyrannosauridae and Aves and ail its descendents (Tyrônnoraptora: Sereno 1999). Losses of the morphology would have occurred within the Eumaniraptora (dromaeosaurids and birds: Holtz: 2000), and therizinosaur- oviraptorosaur (Holtz 2000) or therizinosaur-omithomimosaur (Sereno 1999) clades. This would require only one gain and four losses on one phylogeny (Sereno 1999), and minimally one gain and three losses on the other (HoItz 2000). The universal Iack of an arctometatarsus in adult birds, the most speciose and accessible coelurosaurian clade, may be a spurious bias against this
hypothesis. Fossil evidence of basal tyrannoraptorans (sensu Sereno 1999) will potentially corroborate or falsify the proposal of a single arctometatarsus origin. Overall, the distribution mapped onto Holtz's topology (Holtz 2000; Figure 5.3) is more parsimonious, as it requires only tfiree gains in one scenano, or one gain and three losses, of this complex kinctional suite. More crucially, however, both phylogenies indicate multiple independent ongins andlor losses. The implications of this convergence must be discussed in the context of selective pressures leading to fixation of the arctometatarsus once it emerged. The following section explores this selective aspect of arctometatarsus evolution What selective factors contributed to arctometatarsus evolution? The hypothesis tested here is that the arctometatarsus was an innovation initially involved in prey capture. The long legs of theropods have been considered adaptations enhanang predatory behavior (Gauthier 1986), and with equal validity as an adaptation associateci with increases in home range size (Carrano 1998). Tensile keystone dynamics imply the capacity for increased agility in arctometatarsalians, as presented abuve in Chapter 3. Modem animals employ agility in order to procure prey or to estxtpe predators, and also for intraspecific combat. Potential modem behavioral analogs suggest parallel hypotheses of biological role in theropods. Although observational corroboration is impossible, hypotheses of the biological role of potential agility are subject ta testing by falsification. We can falsify the prey capture hypothesis if herbivory was the primary habit of
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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
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Alexander et al. (1 979) rewrded a
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The preceding loading regime entait
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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 260 and 261: arctometatarsalian forrns. The conv
Page 262 and 263: Proximal intermetatarsal ligaments
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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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THE UNIVERSITY OF CALGARY Eric Snively A ... - Ohio University

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