Boyer diss 2009 1046..

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INTRODUCTION Bloch et al. (2007) analyzed the relationships among plesiadapiforms using a mixture of species, genus, and family level taxa. They used a family level operational taxonomic unit (O.T.U.) to represent plesiadapids and based their coding primarily on the literature available for the skull and skeleton of Plesiadapis tricuspidens (e.g., Russell, 1964; Szalay et al., 1975; Gingerich, 1976), a humerus of Plesiadapis walbeckensis (Szalay and Dagosto, 1980), and the postcranium of Nannodectes (Simpson, 1935; Beard, 1989), as well as on direct observations of the skeleton of Plesiadapis cookei (UM 87990). When different reference species showed variance in character states, Bloch et al. (2007) represented the Plesiadapidae as polymorphic. New information on plesiadapid skeletons presented in chapters 2-4 allows a more accurate coding of the “Plesiadapidae” in the Bloch et al. (2007) matrix. In this study, the use of polymorphic codings for the “Plesiadapidae” is avoided as much as possible: a “composite” Plesiadapidae O.T.U. is generated by optimizing states of characters onto the ancestral node of a newly generated species level tree. The over arching hypotheses to be tested with the new skeletal material are that (1) plesiadapids are close relatives of carpolestids and (2) these taxa plus saxonellids and Chronolestes simul are the sister taxon to Euprimates. The former hypothesis is an apparently fragile aspect of recent phylogenetic analyses (Bloch et al., 2007). While a carpolestid-plesiadapid clade has been favored by specialists familiar with dental morphology (Simpson, 1937; Szalay, 1968; Gingerich, 1976; Fox, 1993; Van Valen, 500
1994), only one (Bloch et al., 2007) of several (i.e., also Bloch and <strong>Boyer</strong>, 2002; Bloch and Silcox, 2006) recently published cladistic analyses has recovered unambiguous support for such a grouping. Bloch and Silcox (2006) described known crania of Carpolestes simpsoni. They showed the nasals to have a narrow caudal extent, the frontal and maxilla to have a sutural contact in the orbit, and described the presence of a foramen rotundum. Additionally, they documented the carpolestid basicranium to have the following features: a posteromedially positioned posterior carotid foramen; a large internal carotid canal (proportionally and absolutely larger than that of even the largest plesiadapids: see tables 3.3; 5.7); well developed stapedial and promontorial grooves on the promontorium; a promontorial groove that does not clearly lead to any septa; and a bone that may be either a dividing wall within a two-chambered auditory bulla or an inconspicuously preserved ectotympanic. The cochlea of Carpolestes has since been measured and shown to be unusually short (Table 2.3) (Coleman and <strong>Boyer</strong>, 2008). In contrast, as demonstrated in chapters 2 and 3, most plesadapids have nasals that are mediolaterally broad at their caudal extent, the nature of suturing in the orbital mosaic is ambiguous, and there is no foramen rotundum. Furthermore, virtually all known plesiadapids have a laterally positioned posterior carotid foramen, an internal carotid plexus foramen of small diameter, no expression of a stapedial system, a narrow promontory groove that leads to an anterior septum (s1), no evidence of multi-chambered bullae, no solid evidence of an ectotympanic bone that is distinct from the bullar wall, and a relatively long cochlea (Table 2.3). However, in Chapter 2, two plesiadapids were argued to have a non-tubular ectotympanic, and Plesiadapis tricuspidens was shown to 501
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NEW CRANIAL AND POSTCRANIAL REMAINS
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Stony Brook University The Graduate
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postcranial anatomy more accurately
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Table of Contents List of Figures .
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Zygomatic. . . . . . . . . . . . .
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Zygomatic. . . . . . . . . . . . .
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Humerus . . . . . . . . . . . . . .
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Description . . . . . . . . . . . .
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Description . . . . . . . . . . . .
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Craniodental material of Plesiadapi
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Figure 2.25. MNHN CR 965 Plesiadapi
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Figure 4.24. UM 87990 Plesiadapis c
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Table List Chapter 2 Table 2.1. Num
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Table 4.30. Caudal vertebrae measur
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CHAPTER 1: INTRODUCTION Plesiadapif
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primates, sharing many dental featu
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were acquired. Specifically, the
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of the plesiadapiform skeleton. Ext
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Gervais, M.P., 1877. Enumération d
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Szalay, F.S., 1972. Cranial morphol
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CHAPTER 2: A REEVALUATION OF CRANIA
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Information on the cranium of basal
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petrosal bulla predicts that a sutu
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History of descriptive study of ple
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Gingerich (1971) rebutted Szalay (1
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9c, Gingerich (1976) labeled a groo
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portion of the bulla medial to the
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Bloch and Silcox (2006) described t
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MATERIALS AND METHODS Material exam
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whitening, dark and light areas on
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SYSTEMATIC PALEONTOLOGY Class MAMMA
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efore meeting a large, anteroposter
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The canine is a simple, single-root
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existence and/or nature of contacts
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outlined. This includes description
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eneath it while also extending post
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y a pair of parallel grooves (Fig.
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the skull (Fig. 2.1). The length of
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margin clearly had a posteriorly pr
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groove measures about 0.29 mm in di
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2.13: 50). The suture with the supr
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preserved (Fig. 2.15: 55). In fact,
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the posterior septum, a deeply inci
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Plesiadapis tricuspidens MNHN CR 12
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is roughly 2.8 mm long. Medial to t
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is not convoluted like many other s
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110) and the only squamosal/alisphe
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two regions for the internal jugula
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ventral to the sinuous suture (132)
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provides measurements of these and
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of the promontorium of the Pellouin
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seen on the HRxCT scan is expressed
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primates, as well as treeshrews and
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canaliculus are present on the sept
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it is missing from the other side o
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observations and interpretations ma
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REFERENCES Beard, K.C., 1993. Phylo
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Russell, D.E., 1959. Le crâne de P
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TABLES Table 2.1. Numerical list of
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81 - Occipital/petrosal suture (Fig
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Table 2.2. Abbreviations for crania
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Table 2.3a. Petrosal features of pl
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Table 2.4. List of cranial measurem
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Shape variables (Table 2.6) ac/GM -
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Table 2.5. continued Specimen MNHN
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Appendix Table 2.1. Specimens scann
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Fig. 2.39 - bs, Ptr, rtp, s1, s2 Fi
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Figure 2.1. UALVP 46685 Pronothodec
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Figure 2.8. USNM 309902 Nannodectes
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Figure 2.9. USNM 309902 Nannodectes
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Figure 2.10. USNM 309902 Nannodecte
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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
Page 477 and 478: Figure 4.12. Surface reconstruction
Page 481 and 482: Figure 4.14. Plesiadapis cookei or
Page 485 and 486: Figure 4.17. Plot of principal coor
Page 487 and 488: Figure 4.19. Plesiadapis cookei (UM
Page 497: Figure 4.27. Plesiadapis cookei (UM
Page 500 and 501: Figure 4.30 472
Page 502 and 503: Figure 4.31. Measurements of astrag
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Page 506 and 507: Figure 4.34. Measurements of calcan
Page 512 and 513: Figure 4.40. Stereophotographic vie
Page 518 and 519: Figure 4.46. 490
Page 520 and 521: Figure 4.47 492
Page 524 and 525: Figure 4.50. 496
Page 530 and 531: have a lacrimal bone that retains i
Page 532 and 533: Institutional abbreviations AMNH, A
Page 534 and 535: level cladogram. A total of 33 cran
Page 536 and 537: plesiadapiform Ignacius graybullian
Page 538 and 539: RESULTS Phylogenetic reconstruction
Page 540 and 541: Optimization of postcranial traits
Page 542 and 543: Therefore, character optimization r
Page 544 and 545: carpolestid bulla is not split into
Page 546 and 547: 2008). I therefore changed the codi
Page 548 and 549: Re-coding and optimization of crani
Page 550 and 551: 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
Page 564 and 565: 158 (p50). Metatarsal I facet on en
Page 566 and 567: Table 5.4C. Postcranial characters
Page 568 and 569: Table 5.7. Posterior carotid forame
Page 570 and 571: Figure 5.2. 542
Page 572 and 573: Figure 5.3 544
Page 574 and 575: Figure 5.4. Plot of posterior carot
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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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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