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

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and constructional morphology reveal developmental and material capacities and constraints on structure. Phylogenetic analysis allows testing of evolutionary hypothesis that anse from apparent rnorphological similanty. Functional analysis entails the atomization and subsequent integration of biological subsystems in order to understand a more complex system. Finaliy, deductive methods involve quantitative modeling, which gives rise to predictions about how natural systems operate. These methods are readily applied to extant organisms. Assessing functional morphology of extinct animals presents obvious constraints on some of the preceding approaches. Ontogenetic study is limited to evidence from hard tissues, and low sample sizes curtail developmental investigation of large fossil vertebrates. Constructional morphology of fcssil hard tissues is readily amenable to study, but that of soft tissues requires inference of their presence and assumptions about their composition. This caution also overiies functional analysis of organs that indude unpreserved components. Fossil systems can be modeled deductively, but falsifying such a model is even more difiicult than with studies of extant organisms (Lauder 1995). Of these approaches, only phylogenetic analysis of fossil organisms attains kill reciprocity with neontology. Phylogenetic study of extant biota requires a palaeontological perspective for falsification (Gauthier et al. 1988) The common tnpwire for many of these approaches is the required inference of unpreserved structures. This impediment is surrnountable to some degree through careful examination of the fossil evidence, using principles of comparative anatomy.
Completing the system in fossil animals: inference of unpreserved structures Two fruitful approaches facilitate inference of unpreserved soft tissues in vertebrat5 fossils. 60th methods require anatomical comparisons, and assessments of correlates to soft tissue occumng on bones of extant vertebrates. One method involves inference from phylogeny, and the other approach entails extrapolation from histologieal and kinctional relationships beiween soft and hard tissues. Recently the method of phylogenetic inference has been formalized (Bryant and Russell 1992, Witmer 1995), invoking cornparison with living or well- preserved extinct relatives of the fossil taon. Phylogenetic inference mandates detailed anatomical examination of modem forms, and the framework of explicit cladistic hypotheses. Soft tissue organs in fossil taxa are robustly inferable when derived and primitive modem outgroups (the extant phylogenetic bracket) possess osteological correlates to the structure also present in the extinct clade (Witmer 1995). For example, heads of extant dinosaurs (birds) and the nearest modem relatives of dinosaurs (mcodilians) have pneumatic diverticula or air sacs in the region anterior to the eyes, associated with fenestration or foramina in the skull bones. Extinct dinosaurs had these openings in their skulls; hence it is quite likely that these dinosaurs also had cranial air sacs (Witmer 1997). Phylogenetic inference becomes Iess certain in taxa without the both elements of the extant bracket (Bryant and Russell 19Ç2, Witrner 1995), or if correlates are ambiguous in any of the examined taxa (Nicholls and Russell
Page 1 and 2: THE UNIVERSITY OF CALGARY Functiona
Page 3: Many figures in this thesis use col
Page 6 and 7: ACKNOWLEûGEM ENTS Many friends and
Page 8 and 9: Approval page Acknowledgements Tabi
Page 10 and 11: CONCLUDING COMMENTS Figures for Cha
Page 12 and 13: CHAPTER 5: Mechanical and phylogene
Page 14 and 15: LIST OF FIGURES Figure 1 .l. Phylog
Page 16 and 17: Figure 3.6. Osteological correlates
Page 18 and 19: AMNH GI HMN IVPP MOR NMC OMNH PIN P
Page 22 and 23: 1985, Bryant and Seymour 1990). If
Page 24 and 25: surfaces of a Tyrannosaurus rex spe
Page 26 and 27: 3. Troodontidae (Figure 1.2): Trwdo
Page 29 and 30: Hypothesized functions of the tyran
Page 31 and 32: 1. Atomization (Chapters 2 and 3).
Page 33 and 34: Figure 1.1. P hylogenetic diagram o
Page 35 and 36: Figure 1.2. Phylogenetic diagram of
Page 41: Figure 1 .S. Skeletal restorations
Page 44 and 45: did not quantify the degree of prox
Page 46 and 47: qualitative debate over arctometata
Page 48 and 49: This hypothesis focuses on tyrannos
Page 50 and 51: Table 2.1. Theropod and Plafeosauru
Page 52 and 53: specimens in the figures reflects a
Page 54 and 55: were not attempted. While this limi
Page 56 and 57: Chapter 3). 1 now address variation
Page 58 and 59: - Ç6'P 1 68'€tr 1 L'OC 99'66 L'9
Page 60 and 61: Table 2.3: Log-transformed values f
Page 62 and 63: experiences a sharp laterally indin
Page 64 and 65: ecause it can be cursorily dassifie
Page 66 and 67: arctometatarsalian specimens, the l
Page 68 and 69: ) 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 3.7. Osteological correlates
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Figure 3.9. Osteological correlates
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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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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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Page 275:
Page 278 and 279:
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.
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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