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

Page 69 and 70: outlined. This includes description

Page 71 and 72: eneath it while also extending post

Page 73 and 74: y a pair of parallel grooves (Fig.

Page 75 and 76: the skull (Fig. 2.1). The length of

Page 77 and 78: margin clearly had a posteriorly pr

Page 79 and 80: groove measures about 0.29 mm in di

Page 81 and 82: 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

Page 87 and 88: Plesiadapis tricuspidens MNHN CR 12

Page 89 and 90: is roughly 2.8 mm long. Medial to t

Page 91 and 92: is not convoluted like many other s

Page 93 and 94: 110) and the only squamosal/alisphe

Page 95 and 96: two regions for the internal jugula

Page 97 and 98: ventral to the sinuous suture (132)

Page 99 and 100: provides measurements of these and

Page 101 and 102: of the promontorium of the Pellouin

Page 103 and 104: seen on the HRxCT scan is expressed

Page 105 and 106: 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

Page 123 and 124: Table 2.3a. Petrosal features of pl

Page 125 and 126: Table 2.4. List of cranial measurem

Page 127 and 128: Shape variables (Table 2.6) ac/GM -

Page 129 and 130: Table 2.5. continued Specimen MNHN

Page 131 and 132: Appendix Table 2.1. Specimens scann

Page 133 and 134: Fig. 2.39 - bs, Ptr, rtp, s1, s2 Fi

Page 135 and 136: Figure 2.1. UALVP 46685 Pronothodec







Page 149 and 150: Figure 2.8. USNM 309902 Nannodectes

Page 151 and 152: Figure 2.9. USNM 309902 Nannodectes

Page 153 and 154: Figure 2.10. USNM 309902 Nannodecte



Page 159 and 160: Figure 2.15. AMNH 17388 Nannodectes

Page 161 and 162: Figure 2.16. AMNH 17388 Nannodectes



Page 167 and 168: Figure 2.22. MNHN CR 965, Plesiadap




Page 175 and 176: Figure 2.26. Pellouin skull Plesiad




Page 183 and 184: Figure 2.30. MaPhQ 33y Adapis paris


Page 187 and 188: Figure 2.32. SBU MRd-12 Sciurus car

Page 189 and 190: Figure 2.33. UMMZ 58983 Tupaia glis

Page 191 and 192: Figure 2.34. Boyer coll. Marmota mo

Page 193 and 194: Figure 2.35. UMMZ TS13 Lagostomus m

Page 195 and 196: Figure 2.36. AMNH 41527 Lagostomus

Page 197 and 198: Figure 2.37. AMNH 185638 Indri indr

Page 199 and 200: Figure 2.38. USNM 482353 Ignacius c

Page 201 and 202: Figure 2.39. UM 108207 Acidomomys h

Page 203 and 204: Figure 2.40. Reconstruction of ples

Page 205 and 206: CHAPTER 3: DESCRIPTION OF THE FIRST

Page 207 and 208: these species. Changing ecological

Page 209 and 210: 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

Page 249 and 250: Figure 3.1. Cranium of Plesiadapis

Page 251 and 252: Figure 3.3. Right maxillary teeth (




Page 259 and 260: Figure 3.8. Fragment from right nuc

Page 261 and 262: Figure 3.9. Right promontorium of P


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

Page 273 and 274: Institutional and collections abbre

Page 275 and 276: CaL - capitulum (of humerus) antero

Page 277 and 278: HSV - head shape variable = ln(DEW/

Page 279 and 280: 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:

Page 339 and 340: 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 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 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 475 and 476: Figure 4.10. Surface reconstruction



Page 481 and 482: Figure 4.14. Plesiadapis cookei or


Page 485 and 486: Figure 4.17. Plot of principal coor






Page 497: Figure 4.27. Plesiadapis cookei (UM

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 512 and 513: Figure 4.40. Stereophotographic vie



Page 518 and 519: Figure 4.46. 490

Page 520 and 521: Figure 4.47 492


Page 524 and 525: Figure 4.50. 496


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

cookei

tricuspidens

plesiadapis

mnhn

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proximal

foramen

medial

lateral

facet

boyer

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