Boyer diss 2009 1046..

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MATERIALS AND METHODS Material examined All known specimens referable to the Plesiadapidae and representing a major or critical portion of the cranium were examined in this study except for MNHN CR 7377, an isolated squamosal glenoid, petrosal, and ectotympanic of Plesiadapis tricuspidens from the Berru locality near Reims, France, which could not be located. Previously unpublished, or largely undescribed, specimens studied include all cranial material referable to a single specimen of Plesiadapis cookei. UM 87990, the primary focus of this report, is a skull and skeleton discovered in 1987 at University of Michigan locality SC-117 in strata of the Fort Union Formation, dated to the middle Clarkforkian (late Paleocene biochron Cf-2) in the Clarks Fork Basin, Wyoming. It was found preserved in a freshwater limestone nodule and was prepared using acid reduction techniques described in Bloch and <strong>Boyer</strong> (2001). P. cookei is compared to other plesiadapids including Pronothodectes gaoi (UALVP 46685, 46687 and 49105), Nannodectes intermedius (USNM 309902), N. gidleyi (AMNH 17388), other species of Plesiadapis, mainly P. tricuspidens (MNHN CR 125, 126, 965, 4306; and the Pellouin skull), and the fragmentary, edentulous rostrum of P. anceps (YPM-PU 19642). Other specimens included for comparison are isolated petrosals of P. tricuspidens (MNHN BR 17414-19, 1371). For the most part, the comparative sample of plesiadapids utilized here represents the study sample of Chapter 2. 182
Methods of examination and documentation Methods of microscopic examination, photographic documentation and measurement follow Chapter 2. All externally visible morphological structures pertinent to the description are labeled on the figures. Bones are identified with abbreviations (see Table 3.2). Other features are labeled with numbers (Table 3.1). Italicized numbers following figure citations in the description section correspond to labels on the morphology depicted in some or all of the cited figures. Table 3.1 is a list of features corresponding to the labels in the figures. High resolution x-ray computed tomography (HRxCT) data were acquired from Pennsylvania State University (data available upon request) for UM 87990. The original scan consisted of 1435 slices spaced at 0.0614 mm, each slice having a pixel resolution of 0.0527 mm. During the scan, the specimen was embedded in florist foam to hold it in place (see Appendix Table 2.1). Additionally, the florist foam was saturated with water, which constituted a wedge to help improve contrast between matrix and bone. These data were visualized with the software Amira 4.1.2-1 and Image J and assisted in description of internal morphology. HRxCT data was particularly important for verifying identifications of various foramina. Measurements and analysis Various measurements were taken using digital calipers, digital photographs, camera lucida drawings and bone reconstructions from scan imagery (Tables 2.3-5). Measurements of foramina are accurate to at least a twentieth of a millimeter. Other 183
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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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Page 159 and 160: Figure 2.15. AMNH 17388 Nannodectes
Page 161 and 162: Figure 2.16. AMNH 17388 Nannodectes
Page 163 and 164: Figure 2.18. MNHN CR 125 Plesiadapi
Page 167 and 168: Figure 2.22. MNHN CR 965, Plesiadap
Page 175 and 176: Figure 2.26. Pellouin skull Plesiad
Page 183 and 184: Figure 2.30. MaPhQ 33y Adapis paris
Page 187 and 188: Figure 2.32. SBU MRd-12 Sciurus car
Page 189 and 190: Figure 2.33. UMMZ 58983 Tupaia glis
Page 191 and 192: Figure 2.34. Boyer coll. Marmota mo
Page 193 and 194: Figure 2.35. UMMZ TS13 Lagostomus m
Page 195 and 196: Figure 2.36. AMNH 41527 Lagostomus
Page 197 and 198: Figure 2.37. AMNH 185638 Indri indr
Page 199 and 200: Figure 2.38. USNM 482353 Ignacius c
Page 201 and 202: Figure 2.39. UM 108207 Acidomomys h
Page 203 and 204: Figure 2.40. Reconstruction of ples
Page 205 and 206: CHAPTER 3: DESCRIPTION OF THE FIRST
Page 207 and 208: these species. Changing ecological
Page 209: Institutional abbreviations AMNH, A
Page 213 and 214: SYSTEMATIC PALEONTOLOGY Class MAMMA
Page 215 and 216: Premaxilla and premaxillary dentiti
Page 217 and 218: nerve and vessels in life (Fig. 3.5
Page 219 and 220: identifiable. No ethmoid foramina c
Page 221 and 222: process is quite large, projecting
Page 223 and 224: vestibuli. This groove’s point of
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Page 227 and 228: e seen as a wedge-shaped, rugose de
Page 229 and 230: process appears as solid bone. Admi
Page 231 and 232: 16) for P. tricuspidens and Rose (1
Page 233 and 234: DENTAL FUNCTIONAL MORPHOLOGY OF P.
Page 235 and 236: Lower premolar molarization As indi
Page 237 and 238: SUMMARY AND CONCLUSION The skull of
Page 239 and 240: REFERENCES Bloch, J.I., Boyer, D.M.
Page 241 and 242: TABLES Table 3.1. List of anatomica
Page 243 and 244: Table 3.2. Anatomical abbreviations
Page 245 and 246: Table 3.3. Size comparison among pl
Page 247 and 248: Table 3.4 continued. European plesi
Page 249 and 250: Figure 3.1. Cranium of Plesiadapis
Page 251 and 252: Figure 3.3. Right maxillary teeth (
Page 259 and 260: Figure 3.8. Fragment from right nuc
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Figure 3.9. Right promontorium of P
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Figure 3.10. Cranium of Plesiadapis
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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
Page 588 and 589:
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.
Page 596 and 597:
Szalay, F.S., Drawhorn, G., 1980. E
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