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

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did not quantify the degree of proximal constridion of MT III that diierentiates an arctometatarsus from other morphologies. A theropod MT Ill with any degree of proximal constriction, such as that displayed by Allosaurus (Figure 2.1 : e), wuld qualify as arctometatarsalian if the definition is interpreted with some latitude (for example, Hutchinson and Padian 1997). This unquantified morphological definition has become confiated with the hypothesis of a polyphyletic origin of the structure, given the controversial constituency of Holtz's Arctometatarsalia (Tyrannosauridae + Bullatosaura; Holîz 1994b, Hutchinson and Padian 1997, Sereno 1 999). Sereno challenged the monophyly of the Arctometatarsalia (Hoitz 1994b) at the 1994 Society of Vertebrate Paleontology meeting, in part because Deinonychus and Allosaurus appear to display some proximal narrowing of MT 111. Most parties to this controversy initially overlooked Holtz's (1 994a) character of plantar constriction of MT III, which separates the arctometatarsus from other theropod morphologies. Thus, Holtz's (1 994a,b) characterization of MT III morphology that typified the Arctometatarsalia was a three dimensional assessment, and accounted for more than a simple narrowing of the proximal end of this elernent. Sereno (1999) redressed this imbalance by incorparating plantar constriction and further qualitative character states into a phylogenetic analysis. This systematic reassessment of the Theropoda (Sereno 1999) dismpted the monophyly of HolKs (1994b, 1996) Arctometatarsalia. Regardless of the ultimate phytogenetic distribution of the arctometatarsus, the ambiguity surrounding its morphology invites further elucidation. The
definition of the arctometatarsus has so far relied upon comparative qualitative study, which has spurred quantitative investigation of biomechanics (Holtz 1994a) and phylogeny (Holtz 1994b, 1996, Sereno 1999). A thorough assessment of theropod MT III diversity must precede efforts to place the arctometatarsus in a systematic and fundional frarnework, and thus to explicate its roles in theropod evolution and ecology. 60th descriptive and quantitative techniques are useful in addressing these issues. Description provides a salutary prerequisite to mathematical inquiries into morphological diversity. White quantitative analysis is ostensibly a more objective starting point, a grounding in qualitative data is necessary for assessing previous morphological perceptions, and for interpretation of statistical results (Pimentel 1979). Statistical methods must yield wholly to morphological description wtien sample size is very low (often the case with palaeontological specimens: Kemp 1999). Description and observation are the primary methods for arriving at phyiogenetic charaders when morphology is the only criterion available (Grande and Bernis 1998). ln addition, subtleties of morphology may be missed when a worker chooses [andmarks for morphometric study (see conclusions to this chapter for examples). Therefore, this chapter proceeds from descriptive to quantitative assessment of theropod third metatarsals. The development and province of MT Ill affected that of both adjacent weight bearing metatarsals (Holtz 1994a). Morphological segregation of MT III by description and statistical clustering has the potential to elucidate variation in the entire pes of a wide array of taxa. Because the initial
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 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 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
Page 80 and 81: Table 2.5: Specimen statistics Herr
Page 82 and 83: and Sinraptor dongi specimens. With
Page 84 and 85: In addition, a hook like proximal a
Page 86 and 87: oth physical narrowing and elongati
Page 88 and 89: 1s the MT 111 shape segregation fun
Page 90 and 91: tyrannosaurids, ornithomimid, and t
Page 92 and 93: proximal constriction, white EImisa
Page 95:
Figure 2.2. Tyrannosaurid, trwdonti
Page 99:
Figure 2.4. Dromaeosaurid and carno
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- Anterior (flexor) surface medial
Page 105 and 106:
Figure 2.7. Left MT III of Tyrannos
Page 107 and 108:
Figure 2.8. Left MT III of Gorgosau
Page 109 and 110:
Figure 2.9. Left MT III of an omith
Page 111 and 112:
Figure 2.10. Left metatarsus of Tmd
Page 113 and 114:
Figure 2.1 1. Left MT III of Elmisa
Page 115 and 116:
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
Page 131 and 132:
CHAPTER 3: Tensile keystone model o
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dynamically transmitted Iommotor fo
Page 135 and 136:
Table 3.1 . Metatarsi examined in t
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interrnetatarsal dynamics in these
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compelled analysis of movement evid
Page 141 and 142:
Once prepared, TMP 94.1 2.602 and i
Page 143 and 144:
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
Page 157 and 158:
weapons sparring), and the arctomet
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Figure 3.1. Schematic representatio
Page 163:
Figure 3.3. CT reconstruction of ri
Page 167:
Figure 3.5. MT II (left element) an
Page 171:
Figure 3.7. Osteological correlates
Page 175:
Figure 3.9. Osteological correlates
Page 179:
Figure 3.1 1 ae. The metatarsal rec
Page 187:
Figure 3.13. Ligament contribution
Page 191 and 192:
Figure 3.15. Anatorny of the equid
Page 193:
Figure 3.16. Cornparison of loading
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(nodes), and solving stresslstrain
Page 198 and 199:
The practicality of the finite elem
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collective density of trabeculae (K
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
Page 208 and 209:
detemined by using the photographic
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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maintained. An angle of 2 degrees p
Page 218 and 219:
of MT III (Figure 4.4, pinpointed b
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anterodorsal rotation of the distal
Page 222 and 223:
allowed to stretch slightly under t
Page 224:
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
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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morphology matches in degree the co
Page 246 and 247:
discussion elucidates the evolution
Page 248 and 249:
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
Page 256 and 257:
parsimonious scenarios are outlined
Page 258 and 259:
dromaeosaurids], and a dade compris
Page 260 and 261:
arctometatarsalian forrns. The conv
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Proximal intermetatarsal ligaments
Page 264 and 265:
witti a surfeit of potentially supp
Page 267:
Figure 5.2. Phylogeny of the Therop
Page 271:
Page 275:
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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