Boyer diss 2009 1046..

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DISCUSSION AND CONCLUSIONS Morphology of P. cookei The skeleton of P. cookei UM 87990 is significant in that it provides a more complete view of several regions of the anatomy than previously available for any plesiadapid. Bones preserved in UM 87990 that were previously unknown for any species of Plesiadapis include the pollical metacarpal, scaphoid, lunate, pisiform, trapezoid, trapezium, ectocuneiform, hallucal metatarsal and much of the vertebral column. Bones previously only known from fragmentary elements for Plesiadapis include the innominate and tibia. The new morphology has allowed some significant progress in understanding the articular relationships of wrist bones, metacarpals, and metatarsals. It must be acknowledged, however, that due to problems with associations and the persistent lack of an actual complete wrist of a single individual there is still a large amount of uncertainty regarding the reconstructions of the wrist and hand presented in the descriptions. Phylogenetic significance.—Several new observations with possible phylogenetic significance came to light. These will be discussed in detail with regard to specific character matrices in Chapter 5. A reconstruction of the wrist showing that the lunate intervened between the scaphoid and triquetrum refutes the reconstruction by Beard (1989, 1993a) who suggested that the scaphoid and triquetrum articulated with one another. However, Stafford and Thorington (1998) have already showed that, if plesiadapiforms were in fact characterized by the configuration proposed by Beard (1989, 1993a), this would be an 380
autapomorphy, because it appears that the lunate ossification center in Cynocephalus intervenes between the triquetrum and scaphoid before fusing to the scaphoid. From a phenetic perspective, the fact that plesiadapids have humeri that are morphologically more similar to those of primates than to those of other euarchontans could reflect a special relationship of plesiadapids (and other plesiadapiforms) to euprimates. However, this signal is not paralleled by similar analyses of the morphology of the astragalus, calcaneum, vertebrae, or body segments. Functional significance.—The opportunity to consider the functional significance of mobility at several joints of the limbs at a time is a major benefit provided by such a complete skeleton of a single individual. Analyses presented in the Results section reveal that P. cookei probably utilized postures in which the arm was typically extended or flexed in the sagittal plane to some degree, but not substantially abducted or laterally rotated (i.e., the distal end of the humerus pointed caudally or ventrally, but not substantially laterally). As discerned in previous studies of P. tricuspidens (Szalay et al., 1975), it seems that the elbow was typically flexed from a fully extended posture by around 90º or more. The forearm would have been abducted by ~30º, relative to the anteroposterior axis of the humerus. The manus was typically partly supinated and slightly dorsiflexed, although it had the capacity for a large range of dorsiflexion. The mechanical capacity for such a large range of dorsiflexion of the hand (and digits) is most likely important for pronograde quadrupedalism, and possibly descents of large diameter vertical substrates (Jenkins, 1974). A typical hind limb posture would have been characterized by a femur that was abducted, flexed, and laterally rotated relative to the innominate. The knee would have 381
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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
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Figure 3.14. A, Plot of relief inde
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CHAPTER 4: THE FIRST KNOWN SKELETON
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among plesiadapiforms (e.g., Szalay
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Institutional and collections abbre
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CaL - capitulum (of humerus) antero
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HSV - head shape variable = ln(DEW/
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MSD - mid-shaft dorsoventral or ant
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Ry - ray (as in “digit ray”) S-
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History of descriptive study of the
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illustrations of this material, exc
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astragalus and calcaneum was highly
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discussion of the femur indicates t
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supinator crests. He also noted tha
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that it may not even be an archonta
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unstudied material. Specifically, h
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5321), some metapodials (MNHN R 529
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Gingerich and Gunnell (1992) publis
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prehensility they provide, is an in
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euarchontans (Fig. 1.1). Their anal
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for comparison. These include isola
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plesiadapid samples have the same m
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Organization of results Each bone i
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Bloch and Boyer (2002) and N. inter
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clavicle reflects some basic aspect
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Humerus Description.—The right an
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epicondyle actually projects somewh
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cookei is absolutely longer than an
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tuberosity. This crest probably del
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olecranon process to estimate its t
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distinct, convex distal radial face
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of the midcarpal joint), and its pr
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(there is no evidence for more than
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matches the opposing facet on the t
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mobility at the trapezoid-trapezium
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Function.—The three proximal carp
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the bone presently being described:
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the “set 2” MC II is a larger,
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differs from MC II and III in havin
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even more pronounced. The distal en
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etween the distal carpals and the
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have stouter shaft diameters for th
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difference makes them more like kno
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antipronograde clinging postures, o
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foramina, and faces slightly proxim
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spine at the superior tip of the il
Page 357 and 358: the thigh (Gambaryan, 1974). The ha
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Page 363 and 364: The anteromedial side of the tibial
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Page 371 and 372: Function.—The functional features
Page 373 and 374: flexor fibularis groove surface. Ex
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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
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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
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Page 403 and 404: preserved. The ribs are slender and
Page 405 and 406: Carpolestes simpsoni (UM 101963) an
Page 407: third metacarpal, similar to arbore
Page 411 and 412: Jenkins (1974) found that Tupaia gl
Page 413 and 414: Nannodectes and other plesiadapids,
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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
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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
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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
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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
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Coleman, M. N. and Boyer, D.M. 2008
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Godinot, M., Beard, K.C., 1991. Fos
Page 592 and 593:
MacPhee, R.D.E., 1981. Auditory Reg
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Savage, D.E., Russell, D.E., 1983.
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Szalay, F.S., Drawhorn, G., 1980. E
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