Boyer diss 2009 1046..

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Chapter 4: The first known skeleton of Plesiadapis cookei and basic features of the skeleton of Plesiadapidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242 Major objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 Anatomical terminology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 Institutional and collections abbreviations . . . . . . . . . . . . . . . . . . . . . . 245 Generic abbreviations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 Other abbreviations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 History of descriptive study of the plesiadapid postcranium . . . . . . . . 245 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276 Materials examined. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276 Methods of examination and documentation . . . . . . . . . . . . . . . . . . . . 277 Measurements and analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278 Univariate comparisons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278 Multivariate comparisons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279 Body mass estimation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280 Intrinsic hand proportions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Organization of results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Systematic Paleontology (Results). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 Type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 Referred specimen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 Ontogenetic state of UM 87990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 Clavicle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 Scapula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286 xi
Humerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290 Radius. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294 Ulna. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297 Scaphoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299 Lunate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300 Triquetrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 Pisiform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303 Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303 Trapezium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304 xii
Page 1 and 2: NEW CRANIAL AND POSTCRANIAL REMAINS
Page 3 and 4: Stony Brook University The Graduate
Page 5 and 6: postcranial anatomy more accurately
Page 7 and 8: Table of Contents List of Figures .
Page 9 and 10: Zygomatic. . . . . . . . . . . . .
Page 11: Zygomatic. . . . . . . . . . . . .
Page 15 and 16: Description . . . . . . . . . . . .
Page 17 and 18: Description . . . . . . . . . . . .
Page 19 and 20: Craniodental material of Plesiadapi
Page 21 and 22: Figure 2.25. MNHN CR 965 Plesiadapi
Page 23 and 24: Figure 4.24. UM 87990 Plesiadapis c
Page 25 and 26: Table List Chapter 2 Table 2.1. Num
Page 27 and 28: Table 4.30. Caudal vertebrae measur
Page 29 and 30: CHAPTER 1: INTRODUCTION Plesiadapif
Page 31 and 32: primates, sharing many dental featu
Page 33 and 34: were acquired. Specifically, the
Page 35 and 36: of the plesiadapiform skeleton. Ext
Page 37 and 38: Gervais, M.P., 1877. Enumération d
Page 39 and 40: Szalay, F.S., 1972. Cranial morphol
Page 41 and 42: CHAPTER 2: A REEVALUATION OF CRANIA
Page 43 and 44: Information on the cranium of basal
Page 45 and 46: petrosal bulla predicts that a sutu
Page 47 and 48: History of descriptive study of ple
Page 49 and 50: Gingerich (1971) rebutted Szalay (1
Page 51 and 52: 9c, Gingerich (1976) labeled a groo
Page 53 and 54: portion of the bulla medial to the
Page 55 and 56: Bloch and Silcox (2006) described t
Page 57 and 58: MATERIALS AND METHODS Material exam
Page 59 and 60: whitening, dark and light areas on
Page 61 and 62: SYSTEMATIC PALEONTOLOGY Class MAMMA
Page 63 and 64:
efore meeting a large, anteroposter
Page 65 and 66:
The canine is a simple, single-root
Page 67 and 68:
existence and/or nature of contacts
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outlined. This includes description
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eneath it while also extending post
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y a pair of parallel grooves (Fig.
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the skull (Fig. 2.1). The length of
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margin clearly had a posteriorly pr
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groove measures about 0.29 mm in di
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2.13: 50). The suture with the supr
Page 83 and 84:
preserved (Fig. 2.15: 55). In fact,
Page 85 and 86:
the posterior septum, a deeply inci
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Plesiadapis tricuspidens MNHN CR 12
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is roughly 2.8 mm long. Medial to t
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is not convoluted like many other s
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110) and the only squamosal/alisphe
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two regions for the internal jugula
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ventral to the sinuous suture (132)
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provides measurements of these and
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of the promontorium of the Pellouin
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seen on the HRxCT scan is expressed
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primates, as well as treeshrews and
Page 107 and 108:
canaliculus are present on the sept
Page 109 and 110:
it is missing from the other side o
Page 111 and 112:
observations and interpretations ma
Page 113 and 114:
REFERENCES Beard, K.C., 1993. Phylo
Page 115 and 116:
Russell, D.E., 1959. Le crâne de P
Page 117 and 118:
TABLES Table 2.1. Numerical list of
Page 119 and 120:
81 - Occipital/petrosal suture (Fig
Page 121 and 122:
Table 2.2. Abbreviations for crania
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Table 2.3a. Petrosal features of pl
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Table 2.4. List of cranial measurem
Page 127 and 128:
Shape variables (Table 2.6) ac/GM -
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Table 2.5. continued Specimen MNHN
Page 131 and 132:
Appendix Table 2.1. Specimens scann
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Fig. 2.39 - bs, Ptr, rtp, s1, s2 Fi
Page 135 and 136:
Figure 2.1. UALVP 46685 Pronothodec
Page 137 and 138:
Page 139 and 140:
Page 141 and 142:
Page 143 and 144:
Page 145 and 146:
Page 147 and 148:
Page 149 and 150:
Figure 2.8. USNM 309902 Nannodectes
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Figure 2.9. USNM 309902 Nannodectes
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Figure 2.10. USNM 309902 Nannodecte
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Page 157 and 158:
Page 159 and 160:
Figure 2.15. AMNH 17388 Nannodectes
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Figure 2.16. AMNH 17388 Nannodectes
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Figure 2.18. MNHN CR 125 Plesiadapi
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Page 167 and 168:
Figure 2.22. MNHN CR 965, Plesiadap
Page 169 and 170:
Page 171 and 172:
Page 173 and 174:
Page 175 and 176:
Figure 2.26. Pellouin skull Plesiad
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Page 179 and 180:
Page 181 and 182:
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Figure 2.30. MaPhQ 33y Adapis paris
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Figure 2.32. SBU MRd-12 Sciurus car
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Figure 2.33. UMMZ 58983 Tupaia glis
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Figure 2.34. Boyer coll. Marmota mo
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Figure 2.35. UMMZ TS13 Lagostomus m
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Figure 2.36. AMNH 41527 Lagostomus
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Figure 2.37. AMNH 185638 Indri indr
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Figure 2.38. USNM 482353 Ignacius c
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Figure 2.39. UM 108207 Acidomomys h
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Figure 2.40. Reconstruction of ples
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CHAPTER 3: DESCRIPTION OF THE FIRST
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these species. Changing ecological
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Institutional abbreviations AMNH, A
Page 211 and 212:
Methods of examination and document
Page 213 and 214:
SYSTEMATIC PALEONTOLOGY Class MAMMA
Page 215 and 216:
Premaxilla and premaxillary dentiti
Page 217 and 218:
nerve and vessels in life (Fig. 3.5
Page 219 and 220:
identifiable. No ethmoid foramina c
Page 221 and 222:
process is quite large, projecting
Page 223 and 224:
vestibuli. This groove’s point of
Page 225 and 226:
9: 40). The right side reveals an a
Page 227 and 228:
e seen as a wedge-shaped, rugose de
Page 229 and 230:
process appears as solid bone. Admi
Page 231 and 232:
16) for P. tricuspidens and Rose (1
Page 233 and 234:
DENTAL FUNCTIONAL MORPHOLOGY OF P.
Page 235 and 236:
Lower premolar molarization As indi
Page 237 and 238:
SUMMARY AND CONCLUSION The skull of
Page 239 and 240:
REFERENCES Bloch, J.I., Boyer, D.M.
Page 241 and 242:
TABLES Table 3.1. List of anatomica
Page 243 and 244:
Table 3.2. Anatomical abbreviations
Page 245 and 246:
Table 3.3. Size comparison among pl
Page 247 and 248:
Table 3.4 continued. European plesi
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Figure 3.1. Cranium of Plesiadapis
Page 251 and 252:
Figure 3.3. Right maxillary teeth (
Page 253 and 254:
Page 255 and 256:
Page 257 and 258:
Page 259 and 260:
Figure 3.8. Fragment from right nuc
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Figure 3.9. Right promontorium of P
Page 263 and 264:
Page 265 and 266:
Figure 3.12. Right dentary of Plesi
Page 267 and 268:
Figure 3.14. A, Plot of relief inde
Page 269 and 270:
CHAPTER 4: THE FIRST KNOWN SKELETON
Page 271 and 272:
among plesiadapiforms (e.g., Szalay
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Institutional and collections abbre
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CaL - capitulum (of humerus) antero
Page 277 and 278:
HSV - head shape variable = ln(DEW/
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MSD - mid-shaft dorsoventral or ant
Page 281 and 282:
Ry - ray (as in “digit ray”) S-
Page 283 and 284:
History of descriptive study of the
Page 285 and 286:
illustrations of this material, exc
Page 287 and 288:
astragalus and calcaneum was highly
Page 289 and 290:
discussion of the femur indicates t
Page 291 and 292:
supinator crests. He also noted tha
Page 293 and 294:
that it may not even be an archonta
Page 295 and 296:
unstudied material. Specifically, h
Page 297 and 298:
5321), some metapodials (MNHN R 529
Page 299 and 300:
Gingerich and Gunnell (1992) publis
Page 301 and 302:
prehensility they provide, is an in
Page 303 and 304:
euarchontans (Fig. 1.1). Their anal
Page 305 and 306:
for comparison. These include isola
Page 307 and 308:
plesiadapid samples have the same m
Page 309 and 310:
Organization of results Each bone i
Page 311 and 312:
Bloch and Boyer (2002) and N. inter
Page 313 and 314:
clavicle reflects some basic aspect
Page 315 and 316:
Humerus Description.—The right an
Page 317 and 318:
epicondyle actually projects somewh
Page 319 and 320:
cookei is absolutely longer than an
Page 321 and 322:
tuberosity. This crest probably del
Page 323 and 324:
olecranon process to estimate its t
Page 325 and 326:
distinct, convex distal radial face
Page 327 and 328:
of the midcarpal joint), and its pr
Page 329 and 330:
(there is no evidence for more than
Page 331 and 332:
matches the opposing facet on the t
Page 333 and 334:
mobility at the trapezoid-trapezium
Page 335 and 336:
Function.—The three proximal carp
Page 337 and 338:
the bone presently being described:
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the “set 2” MC II is a larger,
Page 341 and 342:
differs from MC II and III in havin
Page 343 and 344:
even more pronounced. The distal en
Page 345 and 346:
etween the distal carpals and the
Page 347 and 348:
have stouter shaft diameters for th
Page 349 and 350:
difference makes them more like kno
Page 351 and 352:
antipronograde clinging postures, o
Page 353 and 354:
foramina, and faces slightly proxim
Page 355 and 356:
spine at the superior tip of the il
Page 357 and 358:
the thigh (Gambaryan, 1974). The ha
Page 359 and 360:
The femoral shaft is smooth, lackin
Page 361 and 362:
lacking a lateral extension of its
Page 363 and 364:
The anteromedial side of the tibial
Page 365 and 366:
fibular notch and the strong crest
Page 367 and 368:
to the peroneal surface. The perone
Page 369 and 370:
could even be described as having t
Page 371 and 372:
Function.—The functional features
Page 373 and 374:
flexor fibularis groove surface. Ex
Page 375 and 376:
tubercle, which is centrally locate
Page 377 and 378:
Cuboid Description.—The right cub
Page 379 and 380:
Ectocuneiform Description.—A left
Page 381 and 382:
Metatarsals Hallucal metatarsal des
Page 383 and 384:
articulation with the entocuneiform
Page 385 and 386:
medial side facet on MT IV from a v
Page 387 and 388:
Vertebral column Vertebral column d
Page 389 and 390:
measurements, see caption of Fig. 4
Page 391 and 392:
transverse processes, and the poste
Page 393 and 394:
taxa appear to have slightly more p
Page 395 and 396:
dorsoventrally than craniocaudally.
Page 397 and 398:
The zygapophyses increase in size b
Page 399 and 400:
vertebrae. It is also similar to th
Page 401 and 402:
identifications have been reversed.
Page 403 and 404:
preserved. The ribs are slender and
Page 405 and 406:
Carpolestes simpsoni (UM 101963) an
Page 407 and 408:
third metacarpal, similar to arbore
Page 409 and 410:
autapomorphy, because it appears th
Page 411 and 412:
Jenkins (1974) found that Tupaia gl
Page 413 and 414:
Nannodectes and other plesiadapids,
Page 415 and 416:
are consistent with more frequent u
Page 417 and 418:
differences generated from informat
Page 419 and 420:
Perry, M. Silcox, R. Secord and man
Page 421 and 422:
ubriventer: implications for the fu
Page 423 and 424:
Linnaeus, C., 1758. Systema naturae
Page 425 and 426:
Stern, J.T.,Jr., 1988. Essentials o
Page 427 and 428:
Table 4.3A. Measurements of the pro
Page 429 and 430:
Table 4.3C. Comparative shape varia
Page 431 and 432:
Table 4.5A. Measurements and shape
Page 433 and 434:
Table 4.10. Measurements and shape
Page 435 and 436:
Table 4.13. Measurements of plesiad
Page 437 and 438:
Page 439 and 440:
Table 4.17B. Measurements of the di
Page 441 and 442:
Table 4.18B. Measurements of the di
Page 443 and 444:
Table 4.20B. Measurements 11-18 of
Page 445 and 446:
Table 4.21A. Measurements 1-10 and
Page 447 and 448:
Table 4.23. Measurements and shape
Page 449 and 450:
Page 451 and 452:
Page 453 and 454:
Page 455 and 456:
Table 4.35. Body segment lengths (m
Page 457 and 458:
Table 4.37A. Parameters for Gingeri
Page 459 and 460:
Table 4.38C. Summary of plesiadapid
Page 461 and 462:
Appendix Table 4.1B. Plesiadapis co
Page 463 and 464:
Appendix Table 4.3. Other plesiadap
Page 465 and 466:
Figure 4.2. Plesiadapis cookei (UM
Page 467 and 468:
Figure 4.4. Plot of principal coord
Page 469 and 470:
Figure 4.6. Surface reconstructions
Page 471 and 472:
Page 473 and 474:
Page 475 and 476:
Figure 4.10. Surface reconstruction
Page 477 and 478:
Page 479 and 480:
Page 481 and 482:
Figure 4.14. Plesiadapis cookei or
Page 483 and 484:
Page 485 and 486:
Figure 4.17. Plot of principal coor
Page 487 and 488:
Page 489 and 490:
Page 491 and 492:
Page 493 and 494:
Page 495 and 496:
Page 497:
Page 500 and 501:
Figure 4.30 472
Page 502 and 503:
Figure 4.31. Measurements of astrag
Page 504 and 505:
Figure 4.33 476
Page 506 and 507:
Figure 4.34. Measurements of calcan
Page 508 and 509:
Page 510 and 511:
Page 512 and 513:
Figure 4.40. Stereophotographic vie
Page 514 and 515:
Page 516 and 517:
Page 518 and 519:
Figure 4.46. 490
Page 520 and 521:
Figure 4.47 492
Page 522 and 523:
Page 524 and 525:
Figure 4.50. 496
Page 526 and 527:
Page 528 and 529:
INTRODUCTION Bloch et al. (2007) an
Page 530 and 531:
have a lacrimal bone that retains i
Page 532 and 533:
Institutional abbreviations AMNH, A
Page 534 and 535:
level cladogram. A total of 33 cran
Page 536 and 537:
plesiadapiform Ignacius graybullian
Page 538 and 539:
RESULTS Phylogenetic reconstruction
Page 540 and 541:
Optimization of postcranial traits
Page 542 and 543:
Therefore, character optimization r
Page 544 and 545:
carpolestid bulla is not split into
Page 546 and 547:
2008). I therefore changed the codi
Page 548 and 549:
Re-coding and optimization of crani
Page 550 and 551:
and paromomyids. This, however, is
Page 552 and 553:
REFERENCES Beard, K.C., 1989. Postc
Page 554 and 555:
Novacek, M.J., 1986. The skull of l
Page 556 and 557:
TABLES Table 5.1. Dental characters
Page 558 and 559:
Table 5.2. Dental character matrix.
Page 560 and 561:
asisphenoid and basioccipital bones
Page 562 and 563:
111 (p3). Deltopectoral crest of hu
Page 564 and 565:
158 (p50). Metatarsal I facet on en
Page 566 and 567:
Table 5.4C. Postcranial characters
Page 568 and 569:
Table 5.7. Posterior carotid forame
Page 570 and 571:
Figure 5.2. 542
Page 572 and 573:
Figure 5.3 544
Page 574 and 575:
Figure 5.4. Plot of posterior carot
Page 576 and 577:
ostral end of the nasals) and prema
Page 578 and 579:
its body size would fit predictions
Page 580 and 581:
vertically-to-caudally projecting t
Page 582 and 583:
More generally speaking, this disse
Page 584 and 585:
Figure 6.1. CT reconstruction of Pl
Page 586 and 587:
BIBLIOGRAPHY Alexander, R.M., Jayes
Page 588 and 589:
Coleman, M. N. and Boyer, D.M. 2008
Page 590 and 591:
Godinot, M., Beard, K.C., 1991. Fos
Page 592 and 593:
MacPhee, R.D.E., 1981. Auditory Reg
Page 594 and 595:
Savage, D.E., Russell, D.E., 1983.
Page 596 and 597:
Szalay, F.S., Drawhorn, G., 1980. E
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