Boyer diss 2009 1046..

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similarly present in the euprimate Indri (Fig. 2.37), this could also explain the “bilaminar” appearance of its medial process. Given the similarity between the medial tympanic processes of paromomyids, Indri, and plesiadapids, the more detailed interpretation of paromomyids can be applied to plesiadapids as well. Thus the apparent sutures relating to the medial tympanic process of some of the Pronothodectes specimens and the Pellouin skull likely were formed in a way similar to those of the Ignacius and Acidomomys specimens. That is, these sutures are likely to be so-called “petroso-petrosal” sutures that have been observed in Tarsius (MacPhee and Cartmill, 1986). It is explained that these sutures occur as a result of relatively rapid growth of the tympanic processes of the petrosal. SUMMARY AND CONCLUSIONS Description and analysis of new plesiadapid cranial specimens revealed the Plesiadapidae to be more diverse than previously recognized. Some features thought to be autapomorphic for the clade are revealed to be less developed in species other than P. tricuspidens. Specifically, the nasal bones are broader, the premaxillae are narrower and the external auditory meati are shorter in non-P. tricuspidens plesiadapids. Furthermore, previous conceptions of the clade’s basicranial anatomy were revised. Contrary to some previous interpretations, there is strong morphological evidence that plesiadapids have a posterolaterally positioned posterior carotid foramen; a consistent intratympanic route for the internal carotid plexus; a combined superior orbital fissure and foramen rotundum (i.e., a sphenorbital fissure); and a petrosally derived auditory bulla. These new 82
observations and interpretations may bear on previously proposed phylogenetic relationships among plesiadapids, other plesiadapiforms and euprimates. ACKNOWLEDGMENTS I am particularly grateful to R. C. Fox and C. S. Scott for allowing me to lead the description and analysis of the UALVP Pronothodectes gaoi specimens. A. Walker and T. Ryan provided HRxCT scans of this specimen and others at the Center for Quantitative Imaging, Pennsylvania State University. M. Colbert and T. Rowe provided HRxCT scans of USNM 309902 at the High-Resolution X-ray CT Facility of the University of Texas at Austin. S. Judex and C. Rubin provided HRxCT scans at the Center for Biotechnology of the Department of Biomedical Engineering at Stony Brook University. M. Godinot, P. Tassy, and C. Sagne of MNHN, and M. Pellouin provided access to important comparative specimens of Plesiadapis tricuspidens. R. Emry of the USNM granted access to and study of USNM 309902, Nannodectes intermedius. M. Jin of the AMNH granted access to and study of AMNH 17388, Nannodectes gidleyi. M. Coleman provided measurements of cochlea length in Table 2.3. J. Scott of the University of Alberta, Department of Biological Sciences helped acquire scanning electron microscope images of UALVP 46685, 46687, and 49105. J. Wible of the Carnegie Museum of Natural History assisted in interpreting soft anatomical correlates of the osteology of the tympanic region. This research was further enhanced by discussions with J. Bloch, P. Gingerich, M. Godinot, J. Perry, M. Silcox, and many other researchers. D. Krause, M. O’Leary, J. Fleagle, W. Jungers, and P. Gingerich read and enhanced 83
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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
Page 59 and 60: whitening, dark and light areas on
Page 61 and 62: SYSTEMATIC PALEONTOLOGY Class MAMMA
Page 63 and 64: efore meeting a large, anteroposter
Page 65 and 66: The canine is a simple, single-root
Page 67 and 68: existence and/or nature of contacts
Page 69 and 70: outlined. This includes description
Page 71 and 72: eneath it while also extending post
Page 73 and 74: y a pair of parallel grooves (Fig.
Page 75 and 76: the skull (Fig. 2.1). The length of
Page 77 and 78: margin clearly had a posteriorly pr
Page 79 and 80: groove measures about 0.29 mm in di
Page 81 and 82: 2.13: 50). The suture with the supr
Page 83 and 84: preserved (Fig. 2.15: 55). In fact,
Page 85 and 86: the posterior septum, a deeply inci
Page 87 and 88: Plesiadapis tricuspidens MNHN CR 12
Page 89 and 90: is roughly 2.8 mm long. Medial to t
Page 91 and 92: is not convoluted like many other s
Page 93 and 94: 110) and the only squamosal/alisphe
Page 95 and 96: two regions for the internal jugula
Page 97 and 98: ventral to the sinuous suture (132)
Page 99 and 100: provides measurements of these and
Page 101 and 102: of the promontorium of the Pellouin
Page 103 and 104: seen on the HRxCT scan is expressed
Page 105 and 106: primates, as well as treeshrews and
Page 107 and 108: canaliculus are present on the sept
Page 109: it is missing from the other side o
Page 113 and 114: REFERENCES Beard, K.C., 1993. Phylo
Page 115 and 116: Russell, D.E., 1959. Le crâne de P
Page 117 and 118: TABLES Table 2.1. Numerical list of
Page 119 and 120: 81 - Occipital/petrosal suture (Fig
Page 121 and 122: Table 2.2. Abbreviations for crania
Page 123 and 124: Table 2.3a. Petrosal features of pl
Page 125 and 126: Table 2.4. List of cranial measurem
Page 127 and 128: Shape variables (Table 2.6) ac/GM -
Page 129 and 130: Table 2.5. continued Specimen MNHN
Page 131 and 132: Appendix Table 2.1. Specimens scann
Page 133 and 134: Fig. 2.39 - bs, Ptr, rtp, s1, s2 Fi
Page 135 and 136: Figure 2.1. UALVP 46685 Pronothodec
Page 149 and 150: Figure 2.8. USNM 309902 Nannodectes
Page 151 and 152: Figure 2.9. USNM 309902 Nannodectes
Page 153 and 154: Figure 2.10. USNM 309902 Nannodecte
Page 159 and 160: Figure 2.15. AMNH 17388 Nannodectes
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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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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
Page 477 and 478:
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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
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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
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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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