Boyer diss 2009 1046..

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Specimens included for direct comparison and contrast include those of the plesiadapiforms Carpolestes simpsoni (UM 101963 and USNM 482354), Ignacius clarksforkensis (UM 82616), and Ignacius graybullianus (USNM 421608 and USNM 482353); the fossil euprimates Adapis parisiensis (MaPhQ 33y) and Leptadapis leenhardti (YPM-PU 11481); and the extant rodents Lagostomus maximus (AMNH 41527, 41522), Dipodomys heermani swarthi (AMNH 124181, 39836, 121077), and Sciurus carolinensis (SBU MRd-12). Methods of examination and documentation Specimens were studied under a binocular light microscope. Sutures and cranial foramina were identified by comparison to extant and fossil skulls. Morphology was photo-documented using a Nikon Coolpix 4500 camera mounted on a copy stand or tripod. For more detailed morphology, I used a continuously-calibrated (for pixel scale) digital camera mounted on a steREO Discovery V12 Zeiss microscope with a 0.63x objective lens and 10x ocular lens, motorized focus and zoom, and capacity for reflected and transmitted illumination of objects in the object field (maximum = 36.5mm) via two Zeiss LCD 2500 light boxes. Measurements were taken from the resulting photographs (on structures such as cranial foramina) to the nearest hundredth of a millimeter using the measurement software Axiovision 4.4. In some cases, for specimens that could not be imaged with the Zeiss microscope, camera lucida drawings of minute morphological features were made and measured. Prior to photography for documentation purposes, specimens were whitened with ammonium chloride salt or magnesium powder to remove tonal contrasts due to the mottled coloration of the fossil or glare off the surface. After 30
whitening, dark and light areas on a specimen correspond predominantly to shadows and highlights, respectively, and reveal the specimen’s shape more effectively. All externally visible morphological structures pertinent to description are labeled on the figures. Bones are identified with abbreviations (see Table 2.2). Other features are labeled with numbers (Table 2.1). Italicized numbers following figure citations in the description section correspond to labels on the morphology depicted in some or all of the cited figures. High resolution x-ray computed tomography (HRxCT) data were acquired from the Center for Quantitative Imaging of Pennsylvania State University, the High- Resolution X-ray Computed Tomography Facility of the University of Texas at Austin, and the Center for Biotechnology of Stony Brook University on most specimens (data available upon request). These data were visualized with the software Amira 4.1.2-1 and Image J and assisted in description of internal morphology. HRxCT data were particularly important for verifying identifications of various foramina. Measurements Four measurements were taken on the petrosal bone (Table 2.3) and 61 measurements were taken on various aspects of the cranium (Table 2.4-5) using digital calipers, digital photographs, camera lucida drawings and skeletal reconstructions from scan imagery. These measurements were used to quantitatively compare specimens considered in this study. 31
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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
Page 7 and 8: Table of Contents List of Figures .
Page 9 and 10: Zygomatic. . . . . . . . . . . . .
Page 11 and 12: Zygomatic. . . . . . . . . . . . .
Page 13 and 14: Humerus . . . . . . . . . . . . . .
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: MATERIALS AND METHODS Material exam
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
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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 2.18. MNHN CR 125 Plesiadapi
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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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The femoral shaft is smooth, lackin
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lacking a lateral extension of its
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The anteromedial side of the tibial
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fibular notch and the strong crest
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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
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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
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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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