Boyer diss 2009 1046..

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to refine views of the evolutionary origin of Euprimates (Szalay, 1968; Szalay and Dagosto, 1980; Bloch and Boyer, 2003; Kirk et al., 2003). This additional documentation must begin with descriptions of new specimens of poorly represented species. In this dissertation I aim to provide comprehensive descriptions and character analyses of newly discovered cranial and postcranial material referable to the Plesiadapidae, the core family of plesiadapiforms. I also address previous phylogenetic hypotheses for the Plesiadapidae and consider some functional morphological questions that have implications for the degree to which plesiadapids occupied a “primate-like” ecological niche. Specifically, new information on plesiadapid crania and postcrania can help address the hypothesis that the specialized grasping hallux of Euprimates evolved in (and was inherited from) a “plesiadapiform” ancestor that lacked euprimate visual system features, such as a postorbital bar and convergent orbits on the cranium (Bloch and Boyer, 2002; Bloch et al., 2007; Sargis et al., 2007). If cladistic analyses incorporating new cranial and postcranial information on plesiadapids overturn the phylogenetic hypothesis that plesiadapids and carpolestids are sister taxa, and/or members of the sister taxon to euprimates (Bloch et al., 2007), this could show that characters relating to the hallucal grasp complex of the carpolestid plesiadapiform Carpolestes simpsoni (Bloch and Boyer, 2002) were not inherited from a common ancestor with Euprimates. This result, in turn, could have implications for theories on the original adaptive significance of euprimate features (e.g., Sussman, 1991; Cartmill, 1992; Bloch et al., 2007). Different ideas regarding the adaptive significance of euprimate features have different predictions regarding the evolutionary sequence in which features of interest 4
were acquired. Specifically, the “angiosperm exploitation hypothesis” of R. Sussman (Sussman and Raven, 1978; Sussman, 1991) argues that features reflecting the ability to locomote in trees (like relatively long fingers and toes, and a divergent, opposable hallux with a nail) originally facilitated access to angiosperm resources (flowers, nectar, pollen, fruits, seeds, etc.) in a fine branch niche, making grasping features of primary importance. Sussman believed that teeth should and do show increasing specialization towards frugivory through early primate evolution, as elegantly argued by Szalay (1968). Visual system features are predicted to appear at the same time as or after grasping features, but there is no specific point in primate-euprimate evolution when visual system features are seen as integral or critical in Sussman’s hypothesis. Instead, the visual system features are thought to be further improvements for a life spent in the trees searching out angiosperm products, and are described as an eventual consequence of coevolution between angiosperms, primates and other vertebrates that utilize angiosperm resources in an arboreal milieu (e.g., bats and various birds). The view of early primateeuprimate evolution presented by phylogenetic hypotheses that are supported by recent cladistic analyses corroborates the angiosperm exploitation hypothesis (Bloch et al. 2007), but refutes the “visual predation hypothesis” of Cartmill (1972). The visual predation hypothesis suggests that the synchronized evolution of euprimate pedal grasping and visual features allowed euprimates to visually locate and manually apprehend insects in the shrub-layer of forests at night. Decoupled evolution of grasping and visual features, and the precedence of grasping features in the course of evolution leading up to euprimates, is inconsistent with this hypothesis. 5
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: primates, sharing many dental featu
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 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
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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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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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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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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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Figure 4.14. Plesiadapis cookei or
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Figure 4.17. Plot of principal coor
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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
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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
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
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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
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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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
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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