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Plesiadapis tricuspidens MNHN BR 17414-19, 1371 These specimens represent isolated petrosals from Berru identified by M. Godinot. In all cases I assessed the morphology independently and confirmed the attribution to P. tricuspidens. The specimens have been measured from HRxCT scans generated at Pennsylvania State University at a resolution of 0.0500 mm for pixel dimensions, and 0.0581 mm for slice spacing. MNHN BR 17418 preserves a stapes that has fallen into the cochlea. This element was digitally extracted, measured, and helps confirm that P. tricuspidens did not have a functional stapedial artery. The area of its obturator foramen is small relative to its footplate length, which makes it like modern primates that lack stapedial arteries (Coleman and <strong>Boyer</strong>, 2008; in prep.). Table 2.3 is a codified description of these specimens and other petrosal specimens studied here. The presence/absence and measurements of various features is documented therein. None of these specimens reveals an apparent suture at the boundary between the medial tympanic process and the promontorium as in the case of the Pellouin skull. SELECT QUANTITATIVE DIFFERENCES BETWEEN P. TRICUSPIDENS AND OTHER PLESIADAPIDS As mentioned in the introduction, features thought to be distinctive for plesiadapids, as based on observations of P. tricuspidens, prior to this work, include a nasal bone that is narrow at its caudal extent; a premaxilla that has a broad contact with the frontal; and a tubular-shaped ectotympanic (e.g., Bloch et al., 2007). Table 2.5 70
provides measurements of these and other features. Table 2.6 provides natural log ratio variables quantifying their shape (see table 2.4 for descriptions of variables). Although sample sizes are too small for statistical confidence of any sort, it is interesting to note that P. tricuspidens has substantially proportionally narrower nasals (Table 2.6: N/GM), wider premaxillae (Table 2.6: N/Pmx, Pmx/GM), and a more tubular external auditory meatus (Table 2.6: EAM-S) than any of the other plesiadapids. Thus other plesiadapids are not as distinctive in these features as is P. tricuspidens. Another interesting difference between P. tricuspidens and other plesiadapids is its proportionally larger glenoid fossae (Table 2.6: Gld/GM). Finally, it appears that the largest plesiadapids (mainly P. tricuspidens) have the proportionally shortest cochleae (Table 2.6: Cl/GM), smallest petrosals (Table 2.6: Pcsa/GM) and smallest fenestra vestibulae (Table 2.6: ac/GM). NEW EVIDENCE BEARING ON THE COMPOSITION OF THE PLESIADAPID BULLA As mentioned in the introduction, the morphological prediction for a non-petrosal bulla is the presence of a suture separating the bulla from the pars cochlearis of the petrosal bone. However, whether and when this suture would have been obliterated by remodeling, and what the physical evidence for its presence should look like have not been outlined. A preliminary survey of mammals with basicranial forms similar to those of plesiadapids and basal euprimates includes several rodents, treeshrews, and two paromomyids. These taxa have large inflated bullar cavities with bony septa buttressing 71
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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
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 and 58: 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: ventral to the sinuous suture (132)
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 and 110: it is missing from the other side o
Page 111 and 112: observations and interpretations ma
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
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Figure 2.8. USNM 309902 Nannodectes
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Figure 2.9. USNM 309902 Nannodectes
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Figure 2.10. USNM 309902 Nannodecte
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Figure 2.15. AMNH 17388 Nannodectes
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Figure 2.16. AMNH 17388 Nannodectes
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Figure 2.22. MNHN CR 965, Plesiadap
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Figure 2.26. Pellouin skull Plesiad
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Figure 2.30. MaPhQ 33y Adapis paris
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Figure 2.32. SBU MRd-12 Sciurus car
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Figure 2.33. UMMZ 58983 Tupaia glis
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Figure 2.34. Boyer coll. Marmota mo
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Figure 2.35. UMMZ TS13 Lagostomus m
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Figure 2.36. AMNH 41527 Lagostomus
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Figure 2.37. AMNH 185638 Indri indr
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Figure 2.38. USNM 482353 Ignacius c
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Figure 2.39. UM 108207 Acidomomys h
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Figure 2.40. Reconstruction of ples
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CHAPTER 3: DESCRIPTION OF THE FIRST
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these species. Changing ecological
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Institutional abbreviations AMNH, A
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Methods of examination and document
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SYSTEMATIC PALEONTOLOGY Class MAMMA
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Premaxilla and premaxillary dentiti
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nerve and vessels in life (Fig. 3.5
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identifiable. No ethmoid foramina c
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process is quite large, projecting
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vestibuli. This groove’s point of
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9: 40). The right side reveals an a
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e seen as a wedge-shaped, rugose de
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process appears as solid bone. Admi
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16) for P. tricuspidens and Rose (1
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DENTAL FUNCTIONAL MORPHOLOGY OF P.
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Lower premolar molarization As indi
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SUMMARY AND CONCLUSION The skull of
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REFERENCES Bloch, J.I., Boyer, D.M.
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TABLES Table 3.1. List of anatomica
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Table 3.2. Anatomical abbreviations
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Table 3.3. Size comparison among pl
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Table 3.4 continued. European plesi
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Figure 3.1. Cranium of Plesiadapis
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Figure 3.3. Right maxillary teeth (
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Figure 3.8. Fragment from right nuc
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Figure 3.9. Right promontorium of P
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Figure 3.12. Right dentary of Plesi
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Figure 3.14. A, Plot of relief inde
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CHAPTER 4: THE FIRST KNOWN SKELETON
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among plesiadapiforms (e.g., Szalay
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Institutional and collections abbre
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CaL - capitulum (of humerus) antero
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HSV - head shape variable = ln(DEW/
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MSD - mid-shaft dorsoventral or ant
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Ry - ray (as in “digit ray”) S-
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History of descriptive study of the
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illustrations of this material, exc
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astragalus and calcaneum was highly
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discussion of the femur indicates t
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supinator crests. He also noted tha
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that it may not even be an archonta
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unstudied material. Specifically, h
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5321), some metapodials (MNHN R 529
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Gingerich and Gunnell (1992) publis
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prehensility they provide, is an in
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euarchontans (Fig. 1.1). Their anal
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for comparison. These include isola
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plesiadapid samples have the same m
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Organization of results Each bone i
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Bloch and Boyer (2002) and N. inter
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clavicle reflects some basic aspect
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Humerus Description.—The right an
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epicondyle actually projects somewh
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cookei is absolutely longer than an
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tuberosity. This crest probably del
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olecranon process to estimate its t
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distinct, convex distal radial face
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of the midcarpal joint), and its pr
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(there is no evidence for more than
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matches the opposing facet on the t
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mobility at the trapezoid-trapezium
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Function.—The three proximal carp
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the bone presently being described:
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the “set 2” MC II is a larger,
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differs from MC II and III in havin
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even more pronounced. The distal en
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etween the distal carpals and the
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have stouter shaft diameters for th
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difference makes them more like kno
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antipronograde clinging postures, o
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foramina, and faces slightly proxim
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spine at the superior tip of the il
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the thigh (Gambaryan, 1974). The ha
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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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