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

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the animals ran on uneven ground. This is an intnguing possibility, although Carpenter (1997) did not offer a detailed morphological analysis. Hypotheses of biological role or function of the arctometatarsus must derive from thorough assessment of the observed inter-relatedness of its parts. Therefore, this thesis attempts a large-sale analysis of arctometatarsus function and its selective implications, through study of basic morphology, systematic variation of the arctometatarsalian pes, and possible connectivities (Bock 1989) between this organ system's constituent structures. Elicitation and testing of hypotheses: atomization, synthesis, and extrapolation The salient features of the arctometatarsus are the proximal and plantar constnction of MT Ill. These observations alone suggest a general prirnary hypothesis and nuIl hypothesis: H(a): MT III constnction imparted functional differences between the arctometatarsus and other theropod metatarsus morphologies. H(o): The constricted MT Ill of the arctometatarsus functioned identically to unconstricted theropod third metatarsais. Superficially these hypotheses appear overly broad. However, generating hypotheses of specific function or performance advantage requires more comprehensive observation. M en these focused hypotheses are supported or falsified, the validity of the more general hypothesis can be upheld or discounted. This thesis seeks out and tests more specific hypotheses of arctometatarsus function through the following methods:
1. Atomization (Chapters 2 and 3). The tyrannosaurid arctometatarsus is chosen as a basis for comparisan with other morphologies. The third metatarsals of theropods are described in detail, and quantitative variation in MT II1 morphology is assessed through a Principal Components Analysis of shape. lndividual bones and soft structure correlates are assessed for the tyrannosaurid arctometatarsus. 2. Synthesis (Chapters 3 and 4). These chapters address possible integrated functions for the ligaments and bones of the arctometatarsus. The extent and orientation of soft tissue correlates in the metatarsus of tyrannosaurids are wmpared with those of the carnosaur Allosaurus fragilis, which lacks a constricted third metatarsal. Range of movement in the metatarsi of several tyrannosaurids is tested through computed tornographic (CT) scans, and physical manipulation of casts. This kinernatic and morpholagical data is synthesized into a model of arctometatarsus function in tyrannosaurids, which is compared with the function of an extant analog, the quine wrist The hypothesized model of tyrannosaurid metatarsus function, and the energy transference hypothesis (Holtz 19944, are tested with finite element stress analysis. The wmputer model integrates the suggested kinematics of the qualitative hypotheses, and assumptions of material properties based on measurements of modem bones. 3. Extrapolation (Chapter 5). The preceding results for tyrannosaurid metatarsals are applied comparatively to potential functions in other theropods, in a revisiting of variation revealed in Chapter 2 Metatarsa1 morphology is placed in
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 20 and 21: and constructional morphology revea
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: Hypothesized functions of the tyran
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
Page 70 and 71: a) Ginglymus. Figure 2.14 shows the
Page 72 and 73: ) Shaft and region of proximal arti
Page 74 and 75: measurements of width pV2S%TU-DE an
Page 76 and 77: Table 2.4b Variancelmeasurement eac
Page 78 and 79: 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
Page 232:
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
Page 271:
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.
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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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