Boyer diss 2009 1046..

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een flexed, and the leg slightly abducted. The foot would have been dorsiflexed and slightly inverted with the capacity for a large range of inversion. Abducted limbs result in a posture with sprawled hands and feet. Such a posture would have made P. cookei adept at navigating relatively large diameter substrates. A physiologically supinated hand and inverted foot would have accommodated cylindrical substrates well (i.e., tree trunks and branches). Jenkins (1974) presents data on Tupaia glis showing it to adopt such supinated hand and inverted foot postures during quadrupedal locomotion on relatively narrow branches. The nature of mobility, in the hind limb especially, would have been useful in ascending vertical substrates, because the hind limb has much of its mobility concentrated in an anterposteriorly extending plane with a mediolateral, rather than dorsoventral orientation. Thus, extension of the hind limb would not tend to push the body farther away from the substrate. The femur of Plesiadapis extends and flexes in the transverse plane. Because the knee would have been laterally rotated during much of the support phase of locomotion, flexion and extension of the knee also occurred mainly in a transverse plane. Finally, as explained in the Results section, although flexion and extension of the astragalotibial joint is limited, abduction-adduction movements and inversion-eversion movements are emphasized. It is important to keep the body close to the substrate during vertical climbing because the moment created by gravity on body mass with respect to a vertical substrate increases when the center of mass moves away from the substrate. To be clear, what suggests orthograde postures and locomotion in Plesiadapis is the unusual degree to which its limbs appear to have been sprawled. For instance, 382
Jenkins (1974) found that Tupaia glis utilizes sprawled limb postures and relies on abduction-adduction (instead of flexion and extension) movements in its hind limbs to a significant degree: Tupaia glis is a pronograde scansorialist, not an orthograde arborealist. However, the hind limb of Tupaia does not resemble that of P. cookei very much in features that would seem to have promoted transverse plane mobility in P. cookei. Furthermore, the hind limb postures that Jenkins (1974) illustrates for Tupaia still show the femur to be more sagittally oriented and mobile than suggested in the reconstructions presented above for P. cookei. The hind limb of the treeshrew Ptilocercus does resemble that of P. cookei in many respects (Sargis, 2002b). Ptilocercus is arboreally committed, engaging in vertical clinging and climbing behaviors (Le Gros Clark, 1926). So it seems reasonable to conclude that Ptilocercus limits its movements to a transverse plane even more than does Tupaia (although no data are available), that this helps Ptilocercus locomote in arboreal settings, and that P. cookei moved more like Ptilocercus than Tupaia. The reduced ability for plantarflexion in plesiadapids relative to treeshrews and many primates (see above) begs the question of how vertical descent postures could have been accomplished in plesiadapiforms. In fact, Beard (1989) suggested plesiadapiforms were not effective at descending vertical supports, and suggested gliding behaviors might have represented an alternative means of descending from tree canopies. However, assembling the hind limb of P. cookei with an abducted, extended femur, a knee flexed at 90º, an astragalotibial joint plantarflexed by 90º, and a fully inverted astragalocalcalcaneal joint (Fig. 4.51) shows an approximation of a limb posture that could be used for vertical descents. It should be noted that further inversion of the foot 383
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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
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the thigh (Gambaryan, 1974). The ha
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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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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: autapomorphy, because it appears th
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
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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
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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