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his 1959 work. Figures 13-19 of Russell (1964) have been frequently relied upon as a reference for the cranial anatomy of P. tricuspidens (Gingerich, 1976; Bloch and Silcox, 2006). Russell (1964) was the only researcher to document the sutural and foraminal patterns in P. tricuspidens with original illustrations, although he provided no photographs. Due to the diagramatic form of the illustrations and controversy concerning sutural patters that followed, the lack of photographic evidence has become problematic. In fact, none of the cranial sutures, or foramina of the orbital region, have ever been adequately photographed. In Russell’s (1964) figures 14 and 19, the orbitotemporal region is reconstructed as preserving a human- or tupaiid treeshrew-like pattern of foramina for cranial nerves with separate foramina for ophthalmic (t.d.r.) and maxillary (t.r.) branches of the trigeminal nerve. Although Russell also documented most major sutures in his figures 14 and 19, the alisphenoid/orbitosphenoid suture and the posterior termination of the orbitosphenoid/frontal suture were not illustrated, or discussed. Russell (1964) changed some of the interpretations he made in his original note in 1959. For instance, he reconsidered the stapedial artery to have been reduced relative to the promontorial branch, or absent. Szalay (1971) concluded that the skull of P. tricuspidens had been incorrectly reconstructed in lateral view by Simons (1960) and Russell (1964), in which the tips of the upper and lower incisors were shown to meet one another while the cheek teeth were occluded. Szalay further suggested that the premaxillae did not contact the frontal and provided a photographic dorsal view of the skull (fig. 2) showing transverse cracks (as interpreted by other authors) in the premaxillae looking very much like symmetrical sutures, intersecting the nasals before reaching the maxillae. 20
Gingerich (1971) rebutted Szalay (1971), pointing out that Simons (1960) actually reconstructed the skull as Szalay (1971) claimed to have “re-reconstructed” it. Gingerich further suggested that Szalay (1971) had no new evidence to support his reinterpretation of the premaxilla as lacking frontal contact. Szalay (1972) described a skull of a paromomyid plesiadapiform, Phenacolemur jepseni (AMNH 48005), and compared it to P. tricuspidens specimens MNHN CR 125 and MNHN CR 7377, as well as to skulls of extant and fossil euprimates. This represents the initial publication of the latter P. tricuspidens specimen. Although the photographs he provided of P. tricuspidens MNHN CR 125 are of high quality (Szalay, 1972: figs. 5, 6), they reveal little about the carotid plexus pathway. His descriptions, however, are consistent with those of Russell (1964). Szalay (1972: figs. 7-9) showed different stereophotographic views of MNHN CR 7377, an isolated squamosal glenoid, petrosal, and ectotympanic. A trough-like remnant of the carotid canal on this specimen is labeled with an arrow on figure 8, which effectively illustrates the carotid canal morphology and a groove extending anteriorly from it onto the promontorium. Gingerich (1975) announced a new specimen of P. tricuspidens from Berru (the Pellouin skull). He provided a stereophotograph of its right ear (p.112, fig. 2) and suggested that its expanded external auditory meatus links it with tarsiiform euprimates. Gingerich (1976) newly described a frontal fragment (YPM-PU 24618) from the Berru locality, and an edentulous splanchnocranium (YPM-PU 19642) of P. anceps from 7-up Butte, a late Paleocene locality in the Medicine Rocks area, Montana. The descriptions are brief and the section on cranial anatomy serves more as an addendum and review than as a comprehensive reassessment of basicranial evidence. Gingerich 21
Page 1 and 2: NEW CRANIAL AND POSTCRANIAL REMAINS
Page 3 and 4: Stony Brook University The Graduate
Page 5 and 6: postcranial anatomy more accurately
Page 7 and 8: Table of Contents List of Figures .
Page 9 and 10: Zygomatic. . . . . . . . . . . . .
Page 11 and 12: Zygomatic. . . . . . . . . . . . .
Page 13 and 14: Humerus . . . . . . . . . . . . . .
Page 15 and 16: Description . . . . . . . . . . . .
Page 17 and 18: Description . . . . . . . . . . . .
Page 19 and 20: Craniodental material of Plesiadapi
Page 21 and 22: Figure 2.25. MNHN CR 965 Plesiadapi
Page 23 and 24: Figure 4.24. UM 87990 Plesiadapis c
Page 25 and 26: Table List Chapter 2 Table 2.1. Num
Page 27 and 28: Table 4.30. Caudal vertebrae measur
Page 29 and 30: CHAPTER 1: INTRODUCTION Plesiadapif
Page 31 and 32: primates, sharing many dental featu
Page 33 and 34: were acquired. Specifically, the
Page 35 and 36: of the plesiadapiform skeleton. Ext
Page 37 and 38: Gervais, M.P., 1877. Enumération d
Page 39 and 40: Szalay, F.S., 1972. Cranial morphol
Page 41 and 42: CHAPTER 2: A REEVALUATION OF CRANIA
Page 43 and 44: Information on the cranium of basal
Page 45 and 46: petrosal bulla predicts that a sutu
Page 47: History of descriptive study of ple
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 and 98: 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
Page 137 and 138:
Page 139 and 140:
Page 141 and 142:
Page 143 and 144:
Page 145 and 146:
Page 147 and 148:
Page 149 and 150:
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 157 and 158:
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Figure 2.15. AMNH 17388 Nannodectes
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Figure 2.16. AMNH 17388 Nannodectes
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Figure 2.18. MNHN CR 125 Plesiadapi
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Figure 2.22. MNHN CR 965, Plesiadap
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Page 171 and 172:
Page 173 and 174:
Page 175 and 176:
Figure 2.26. Pellouin skull Plesiad
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Page 179 and 180:
Page 181 and 182:
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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
Page 385 and 386:
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,
Page 415 and 416:
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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Page 451 and 452:
Page 453 and 454:
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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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Page 479 and 480:
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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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Page 491 and 492:
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Page 495 and 496:
Page 497:
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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
Page 514 and 515:
Page 516 and 517:
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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
Page 538 and 539:
RESULTS Phylogenetic reconstruction
Page 540 and 541:
Optimization of postcranial traits
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Therefore, character optimization r
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carpolestid bulla is not split into
Page 546 and 547:
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
Page 554 and 555:
Novacek, M.J., 1986. The skull of l
Page 556 and 557:
TABLES Table 5.1. Dental characters
Page 558 and 559:
Table 5.2. Dental character matrix.
Page 560 and 561:
asisphenoid and basioccipital bones
Page 562 and 563:
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
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
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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