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specimen from the Bison Basin Saddle locality (AMNH 92011: p. 9, fig. 6), which they identified on the basis of its similarity to craniodentally associated elements from the Berru locality, one of which was also figured (p. 10, fig. 7). The authors did not provide a number for the Berru specimen, but Beard (1989) later recognized it as pertaining to MNHN R 416. Szalay and Dagosto considered the distinctive features of the entocuneiform to be its large plantar-projecting process and a facet for MT I that is mediolaterally broad, relatively flat, and saddle-shaped. They viewed the bone as morphologically similar to that of the extant primitive treeshrew Ptilocercus lowii. Because Ptilocercus is capable of grasping with a divergent hallux that exhibits mobility at the metatarsal-entocuneiform joint, Szalay and Dagosto suggested that the same activities could be inferred for plesiadapids. However, no first metatarsals had yet been described for any plesiadapids. Gunnell (1989: p. 41-48) discussed the astragalus, calcaneum, and cuboid of Plesiadapis. His figures 16 and 17 (p. 45) compare the astragalus and calcaneum to that of Marmota. Gunnell acknowledged that the astragalus and calcaneum indicate a high degree of mobility, as argued by Szalay and Decker (1974). However, he brought up the terrestriality hypothesis of (Gingerich, 1976) and noted that mobility of the ankle does not necessarily argue for an arboreal habitus, as indicated by the mobile foot of terrestrial Marmota and as convincingly argued by Jenkins (1974). However, he stated that an ongoing study by him and Gingerich indicated an arboreal habitus for the skeleton of P. cookei. Beard (1989) brought studies of plesiadapiform postcranial anatomy several steps forward with an excellent dissertation, wherein he described and illustrated previously 266
unstudied material. Specifically, he described and figured a skeleton of Nannodectes intermedius (USNM 442229: p. 20-89, figs. 1-16) including the following bones: left scapula, right humerus, right ulna, left radius, right scaphoid, right lunate, left pisifom, right capitate, both hamates, left MC I, right MC II, right MC III, four proximal manual phalanges, four intermediate manual phalanges, several distal manual phalanges, left tibia, left cuboid, left hallucal metatarsal, four non-hallucal metatarsal fragments and two pedal intermediate and two pedal distal phalanges. All of this material plus additional elements associated with this specimen (but not described by Beard) were observed, measured, and in some cases HRxCT-scanned in the course of this study. Furthermore, Beard described isolated elements from Cedar Point Quarry attributed to P. rex (p. 89- 101, figs. 17, 18) including YPM-PU 23976, a proximal humeral fragment (not figured); YPM-PU 23975, a distal tibial fragment; YPM-PU 23977 and UM 94816, right and left astragali, respectively. Of these bones, I was only able to directly study UM 94816. Beard (1989) additionally described a number of previously unstudied bones of P. tricuspidens, relying on Russell’s (1964) documentation of their association to a partial skeleton mentioned above (p. 101-132, figs. 19-21). The bones newly described by Beard included the following: MNHN R 415, a right cuboid; MNHN R 416, a left entocuneiform; MNHN R 5295, a right MT III; MNHN R 5296, a pedal intemediate phalanx; MNHN R 5297, a manual proximal phalanx; MNHN R 5298, a left MT IV; MNHN R 5300, a right MT III; MNHN 5301 and 5303, manual proximal phalanges; MNHN R 5305, a left MC II; MNHN R 5306, a distal epiphysis of a right MT I; MNHN R 5309, a pedal distal phalanx; MNHN R 5310, a fragment of a pedal distal phalanx; MNHN R 5312, a manual intermediate phalanx; MNHN R 5313, a pedal distal phalanx; 267
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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 2.2. UALVP 46685 Pronothodec
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Figure 2.3. UALVP 46685 Pronothodec
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Figure 2.4. UALVP 46685 Pronothodec
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Figure 2.5. UALVP 46685 Pronothodec
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Figure 2.6. UALVP 49105 Pronothodec
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Figure 2.7. UALVP 46687 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 2.11. USNM 309902 Nannodecte
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Figure 2.13. 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 2.20. MNHN CR 125 Plesiadapi
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Figure 2.22. MNHN CR 965, Plesiadap
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Figure 2.23. MNHN CR 965 Plesiadapi
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Figure 2.24. MNHN CR 965 Plesiadapi
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Figure 2.25. MNHN CR 965, Plesiadap
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Figure 2.26. Pellouin skull Plesiad
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Figure 2.27. Pellouin skull Plesiad
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Figure 2.28. Pellouin skull Plesiad
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Figure 2.29. Pellouin skull Plesiad
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Figure 2.30. MaPhQ 33y Adapis paris
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Figure 2.31. MNHN CR 126, Plesiadap
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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 3.4. Cranium of Plesiadapis
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Figure 3.5. Cranium of Plesiadapis
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Figure 3.6. Cranium of Plesiadapis
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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 3.10. 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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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 4.15. 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 4.26. Measurements of plesiad
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Table 4.28. Measurements of plesiad
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Table 4.32. Measurements of plesiad
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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 4.8. Surface reconstructions
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Figure 4.9. Surface reconstructions
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Figure 4.10. Surface reconstruction
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Figure 4.12. Surface reconstruction
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Figure 4.13. Surface reconstruction
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Figure 4.14. Plesiadapis cookei or
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Figure 4.15. Surface reconstruction
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Figure 4.17. Plot of principal coor
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Figure 4.19. Plesiadapis cookei (UM
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Figure 4.21. Plesiadapis cookei (UM
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Figure 4.23. Plesiadapis cookei (UM
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Figure 4.24. Plesiadapis cookei (UM
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Figure 4.25. Surface reconstruction
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Figure 4.27. Plesiadapis cookei (UM
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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 4.36. Plesiadapis cookei (UM
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Figure 4.38. Plesiadapis cookei (UM
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Figure 4.40. Stereophotographic vie
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Figure 4.42. Plesiadapis cookei (UM
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Figure 4.44. Plesiadapis cookei (UM
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Figure 4.46. 490
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Figure 4.47 492
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Figure 4.48. Plesiadapis cookei (UM
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Figure 4.50. 496
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Figure 4.51. Surface reconstruction
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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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Page 554 and 555:
Novacek, M.J., 1986. The skull of l
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TABLES Table 5.1. Dental characters
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Page 558 and 559:
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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Page 576 and 577:
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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Page 582 and 583:
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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Page 588 and 589:
Coleman, M. N. and Boyer, D.M. 2008
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Page 590 and 591:
Godinot, M., Beard, K.C., 1991. Fos
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Page 592 and 593:
MacPhee, R.D.E., 1981. Auditory Reg
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Page 594 and 595:
Savage, D.E., Russell, D.E., 1983.
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Page 596 and 597:
Szalay, F.S., Drawhorn, G., 1980. E
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