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

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face of the skull met part of the medially expanding lower jaw, The bones formed a stout new joint that braœs the jaw articulation when the skimmer spears a fish (Bock 1959). The sequence of functional acquisition, determined through character analysis (Bock 1965) and ideally through fossil evidence, allows us to classify the relevant aptations. The new function of bracing the jaw is an exaptation derived from Iateral extension of the basitemporal processes, which was not originally involved with a second jaw joint. The original function of increased muscle attachment is still in effect, and thus falls under the category of adaptation proper (Gould and Vrba 1982). 00th functions (termed postadaptation by Bock 1959) are amenable to further refinement. Epistemology and evolution of aptations With this background of terminology, further examples illustrate the evolution of exaptive functional complexes. A simple instance involves one aspect of the origin of flight in birds. The range of motion evident in the forelimb joints of small theropod dinosaurs indicates these animals could sweep their claws down and forward; one possible utility of this action was to stfike and grasp prey. Gauthier and Padian (1985) propose that the same motion was incorporated into the downward component of the flightstroke in birds, the descendants of earlier predaceous theropods. If a predation strike was the primary selective benefit of forelimb kinernatics in the ancestors of birds, the avian downward fiightstroke may be interpreted as an exaptation derived from an anteceding function.
Other aptations refiect multiple fundons that potentially stem from a single, broad morphological innovation. African cichlid fishes have developed a second set of jaws within their pharynx, exapted from tooth bearing pharyngobranchials and ceratobranchials, and associated branchiorneric musculature and innervation (Liern 1973). These cichlids can transport and process food within their throats, and their primary jaws have been freed for wildly diverse specializations in food collection. Liem (1 973) and Lem and Osse (1 975) term such fortuitous structures key innovations, because they allow extensive exploitation of new adaptive zones. Other key innovations enabte less dramatic diversification than that of cichlids, but can still trigger elaboration and multiple functions when selective forces are brought to bear. The strength of the rostrurn of synapsids increased substantially with the development of a full secondary palate (Thomason and Russell 1986). This resistance to bending and torsion enabled a diversification of mechanisms for continuous and energetic mastication in mammals (Thomason and Russell 1986). Caniniform and canine teeth in mammals and other synapsids have been expressed as saber-like modifications in the gorgonopsian Inosfancivia, herbivorous pantodonts, marsupial carnivores, and in saber-toothed cats and their nimravid relatives (Carroll 1.990). Lateral body foIds in gekkonid lizards, initially involved in fat storage, have become modified for crypsis and parachuting (Russell 1979a). These variegated fundions in geckos are not mutually exclusive, and their selective expression depended upon the interplay of behavior, morphology, and environment (Russell 1979a).
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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
Page 198 and 199: The practicality of the finite elem
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Page 202 and 203: A FORCE INPUTS AND MATERIAL PROPERT
Page 204 and 205: Alexander et al. (1 979) rewrded a
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Page 210 and 211: 2. Preparation of data for contour
Page 212 and 213: . - Elastic modulus (GP4 Ex EY Ez P
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Page 228: Figure 4.3. Finite element mesh of
Page 232: Figure 4.5. Strain distribution in
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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 258 and 259: dromaeosaurids], and a dade compris
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Page 262 and 263: Proximal intermetatarsal ligaments
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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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