Boyer diss 2009 1046..

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distal end implies that the forearms were typically somewhat abducted during pronograde postures (Fig. 4.3). Comparison.—A dataset from Szalay and Dagosto (1980) was modified to compare the distal end of the humerus of P. cookei to those of other plesiadapids and mammals (mainly other euarchontans) (Table 4.3C, Fig. 4.4). Six size-standardized variables were analyzed with principal coordinates analysis (PCA). P. cookei is revealed to be most similar in its joint morphology to that of other plesiadapids, as well as adapid and omomyid euprimates. On the other hand, primitive eutherian mammals and nonprimate euarchontans are well-separated from P. cookei. Comparing P. cookei only to other plesiadapids in a more focused, if qualitative fashion, reveals it to differ from smaller forms including Pr. gaoi, N. intermedius, and N. gidleyi in having much greater lateral deviation of its distal end (Table 4.3A: LD). P. tricuspidens, P. remensis, and Pl. daubrei appear to have the distal end of the humerus oriented like that of P. cookei. While all known plesiadapids have a prominent medial keel on the ulnar trochlea of the humerus, P. cookei differs from Pr. gaoi, P. rex, P. tricuspidens and Pl. daubrei in lacking a lateral keel on its trochlea. This feature is present in many euprimates, but is also found in the primitive arboreal treeshrew Ptilocercus lowii. In taxa possessing a lateral keel, the trochlea is more easily distinguished from the zona conoidea lateral to it. The specimens of Nannodectes are like P. cookei in lacking the lateral keel. P. cookei further differs from P. rex alone in the presence of a dorsoepitrochlear fossa (it is retained in all other plesiadapid specimens). Finally, P. cookei appears to differ from at least P. tricuspidens in having a more gracile humerus (Table 4.3A: note higher SSV). Furthermore, while the left humerus of P. 290
cookei is absolutely longer than any complete elements known for P. tricuspidens, almost all of the fragmentary humeri of P. tricuspidens have greater midshaft diameters than those of P. cookei, further implying that the humerus is a more gracile bone in UM 87990 than in any known individual of P. tricuspidens. Radius Description.—The left radius and the distal epiphysis of the right radius are preserved. However, the two elements have been adhered to one another with paste epoxy as if they originally belonged to the same bone (Fig. 4.5A). A medical CT scan image was manipulated to estimate the morphology of the actual element. The diaphysis was separated from the surface reconstruction, inverted and repositioned on the epiphysis (Fig. 4.6). In the following description, the terms “ulnar” and “radial” are used in place of lateral and medial. The “ulnar side” of the radius is that which contacts the ulna at the distal radioulnar joint, and to which the interosseous membrane attaches. The radial side is the opposite side (i.e., the side that the pollex is on; also equivalent to the pre-axial side). Overall, the radius is a slender element with a slight dorsoulnar convexity to its shaft. The radioulnar axis of the distal end is rotated dorsoulnarly by ~40° relative to the proximal end (radial head). That is, when the proximal end of the radius is articulated with the humerus in a fully pronated position, the manus would have been supinated by ~40° (full supination is equal to 180°) (e.g., Fig. 4.3). Furthermore, the manipulated CT image reveals that the distal articular surface faced ventrally and ulnarly. 291
Page 1 and 2:
NEW CRANIAL AND POSTCRANIAL REMAINS
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Stony Brook University The Graduate
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postcranial anatomy more accurately
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Table of Contents List of Figures .
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Zygomatic. . . . . . . . . . . . .
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Zygomatic. . . . . . . . . . . . .
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Humerus . . . . . . . . . . . . . .
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Description . . . . . . . . . . . .
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Description . . . . . . . . . . . .
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Craniodental material of Plesiadapi
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Figure 2.25. MNHN CR 965 Plesiadapi
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Figure 4.24. UM 87990 Plesiadapis c
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Table List Chapter 2 Table 2.1. Num
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Table 4.30. Caudal vertebrae measur
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CHAPTER 1: INTRODUCTION Plesiadapif
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primates, sharing many dental featu
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were acquired. Specifically, the
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of the plesiadapiform skeleton. Ext
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Gervais, M.P., 1877. Enumération d
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Szalay, F.S., 1972. Cranial morphol
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CHAPTER 2: A REEVALUATION OF CRANIA
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Information on the cranium of basal
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petrosal bulla predicts that a sutu
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History of descriptive study of ple
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Gingerich (1971) rebutted Szalay (1
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9c, Gingerich (1976) labeled a groo
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portion of the bulla medial to the
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Bloch and Silcox (2006) described t
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MATERIALS AND METHODS Material exam
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whitening, dark and light areas on
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SYSTEMATIC PALEONTOLOGY Class MAMMA
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efore meeting a large, anteroposter
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The canine is a simple, single-root
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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
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preserved (Fig. 2.15: 55). In fact,
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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
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canaliculus are present on the sept
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it is missing from the other side o
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observations and interpretations ma
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REFERENCES Beard, K.C., 1993. Phylo
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Russell, D.E., 1959. Le crâne de P
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TABLES Table 2.1. Numerical list of
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81 - Occipital/petrosal suture (Fig
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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
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Shape variables (Table 2.6) ac/GM -
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Table 2.5. continued Specimen MNHN
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Appendix Table 2.1. Specimens scann
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Fig. 2.39 - bs, Ptr, rtp, s1, s2 Fi
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Figure 2.1. UALVP 46685 Pronothodec
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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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Figure 2.15. AMNH 17388 Nannodectes
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Figure 2.16. AMNH 17388 Nannodectes
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Figure 2.22. MNHN CR 965, Plesiadap
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Figure 2.26. Pellouin skull Plesiad
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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
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Methods of examination and document
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SYSTEMATIC PALEONTOLOGY Class MAMMA
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Premaxilla and premaxillary dentiti
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nerve and vessels in life (Fig. 3.5
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identifiable. No ethmoid foramina c
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process is quite large, projecting
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vestibuli. This groove’s point of
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9: 40). The right side reveals an a
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e seen as a wedge-shaped, rugose de
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process appears as solid bone. Admi
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16) for P. tricuspidens and Rose (1
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DENTAL FUNCTIONAL MORPHOLOGY OF P.
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Lower premolar molarization As indi
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SUMMARY AND CONCLUSION The skull of
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REFERENCES Bloch, J.I., Boyer, D.M.
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TABLES Table 3.1. List of anatomica
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Table 3.2. Anatomical abbreviations
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Table 3.3. Size comparison among pl
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Table 3.4 continued. European plesi
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Figure 3.1. Cranium of Plesiadapis
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Figure 3.3. Right maxillary teeth (
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Figure 3.8. Fragment from right nuc
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Figure 3.9. Right promontorium of P
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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
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Page 315 and 316: Humerus Description.—The right an
Page 317: epicondyle actually projects somewh
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
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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
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could even be described as having t
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Function.—The functional features
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flexor fibularis groove surface. Ex
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tubercle, which is centrally locate
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Cuboid Description.—The right cub
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Ectocuneiform Description.—A left
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Metatarsals Hallucal metatarsal des
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articulation with the entocuneiform
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medial side facet on MT IV from a v
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Vertebral column Vertebral column d
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measurements, see caption of Fig. 4
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transverse processes, and the poste
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taxa appear to have slightly more p
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dorsoventrally than craniocaudally.
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The zygapophyses increase in size b
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vertebrae. It is also similar to th
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identifications have been reversed.
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preserved. The ribs are slender and
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Carpolestes simpsoni (UM 101963) an
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third metacarpal, similar to arbore
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autapomorphy, because it appears th
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Jenkins (1974) found that Tupaia gl
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Nannodectes and other plesiadapids,
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are consistent with more frequent u
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differences generated from informat
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Perry, M. Silcox, R. Secord and man
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ubriventer: implications for the fu
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Linnaeus, C., 1758. Systema naturae
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Stern, J.T.,Jr., 1988. Essentials o
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Table 4.3A. Measurements of the pro
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Table 4.3C. Comparative shape varia
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Table 4.5A. Measurements and shape
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Table 4.10. Measurements and shape
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Table 4.13. Measurements of plesiad
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Table 4.17B. Measurements of the di
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Table 4.18B. Measurements of the di
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Table 4.20B. Measurements 11-18 of
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Table 4.21A. Measurements 1-10 and
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Table 4.23. Measurements and shape
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Table 4.35. Body segment lengths (m
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Table 4.37A. Parameters for Gingeri
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Table 4.38C. Summary of plesiadapid
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Appendix Table 4.1B. Plesiadapis co
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Appendix Table 4.3. Other plesiadap
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Figure 4.2. Plesiadapis cookei (UM
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Figure 4.4. Plot of principal coord
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Figure 4.6. Surface reconstructions
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Figure 4.10. Surface reconstruction
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Figure 4.14. Plesiadapis cookei or
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Figure 4.17. Plot of principal coor
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Figure 4.30 472
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Figure 4.31. Measurements of astrag
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Figure 4.33 476
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Figure 4.34. Measurements of calcan
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Figure 4.40. Stereophotographic vie
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Figure 4.46. 490
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Figure 4.47 492
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Figure 4.50. 496
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INTRODUCTION Bloch et al. (2007) an
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have a lacrimal bone that retains i
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Institutional abbreviations AMNH, A
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level cladogram. A total of 33 cran
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plesiadapiform Ignacius graybullian
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RESULTS Phylogenetic reconstruction
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Optimization of postcranial traits
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Therefore, character optimization r
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carpolestid bulla is not split into
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2008). I therefore changed the codi
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Re-coding and optimization of crani
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and paromomyids. This, however, is
Page 552 and 553:
REFERENCES Beard, K.C., 1989. Postc
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Novacek, M.J., 1986. The skull of l
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TABLES Table 5.1. Dental characters
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Table 5.2. Dental character matrix.
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asisphenoid and basioccipital bones
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111 (p3). Deltopectoral crest of hu
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158 (p50). Metatarsal I facet on en
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Table 5.4C. Postcranial characters
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Table 5.7. Posterior carotid forame
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Figure 5.2. 542
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Figure 5.3 544
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Figure 5.4. Plot of posterior carot
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ostral end of the nasals) and prema
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its body size would fit predictions
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vertically-to-caudally projecting t
Page 582 and 583:
More generally speaking, this disse
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Figure 6.1. CT reconstruction of Pl
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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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