Boyer diss 2009 1046..

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Abstract of the Dissertation NEW CRANIAL AND POSTCRANIAL REMAINS OF LATE PALEOCENE PLESIADAPIDAE (“PLESIADAPIFORMES,” MAMMALIA) FROM NORTH AMERICA AND EUROPE: DESCRIPTION AND EVOLUTIONARY IMPLICATIONS by Douglas Martin Boyer Doctor of Philosophy in Anatomical Sciences Stony Brook University 2009 Plesiadapidae are a diverse monophyletic family of primate-like mammals that existed during the Paleogene in Europe and North America. They are thought to be nested within the Plesiadapoidea, which are in turn a member of a larger group of fossil mammals called the Plesiadapiformes. An understanding of plesiadapid evolutionary history is important for investigations into the origin of extant primates (including humans, apes, monkeys, lemurs, etc.) plus anatomically modern, extinct primates (euprimates). Specifically, this dissertation accumulates evidence bearing on the hypothesis that the ancestor of euprimates inherited a big toe that (1) is divergent from the lateral digits, (2) exhibits lateral torsion, and (3) has a nail, from its common ancestor with the Plesiadapoidea, in which these features were adaptations to foraging for angiosperm products in a fine branche niche. New information on plesiadapid skeletons may alter views on the phylogenetic relationship of various plesiadapiforms to euprimates. Altered phylogenetic arrangements of plesiadapids, other plesiadapiforms, and euprimates, are likely to indicate that the unusual features of the feet of euprimates were not inherited from plesiadapiforms and evolved at roughly the same time as forward facing orbits and other euprimate novelties. This would re-open the possibility that euprimate features are adaptations for a behavior besides foraging among small branches for angiosperm products. I describe the first known crania of Pronothodectes gaoi, a species of the most basal plesiadapid genus known; a skull of Nannodectes intermedius, the oldest known for a plesiadapid; and the first skull and postcranial skeleton of the large, late-occurring North American species Plesiadapis cookei. I provide new morphological observations for a cranial specimen of Nannodectes gidleyi and several specimens of Plesiadapis tricuspidens. These new data document basic characteristics of plesiadapid cranial and iii
postcranial anatomy more accurately and comprehensively than previously possible. I use quantitative comparative and cladistic methodologies to (1) reconstruct cranial characteristics for the ancestral node of the family Plesiadapidae, (2) generate hypotheses regarding phylogenetic relationships of various plesiadapid species to each other, and (3) test hypotheses regarding the phylogenetic relationship of Plesiadapidae and other plesiadapiforms to extant euarchontan mammals (Primates, Scandentia, and Dermoptera). I use quantitative comparative morphology to reconstruct functional and behavioral features of various plesiadapid species. The ancestral node of the family Plesiadapidae is reconstructed as having an enlarged premaxilla that contacts the frontal; a laterally-positioned, intratympanic, transpromontorial route for the internal carotid plexus; a non-functional internal carotid artery; a non-tubular external auditory meatus; and an auditory bulla, the composition of which is uncertain, but best interpreted as petrosal. P. cookei and P. tricuspidens appear derived in having external auditory meati that are tubular and in having a maxilla that lacks extensive dorsal exposure of the molar tooth roots within the orbit. P. tricuspidens is apparently autapomorphic among plesiadapids in having an increased relative size and posterior projection of the premaxilla, consequent narrowing of the frontal contact with the nasal, an increased relative size of the glenoid fossae of the squamosal and decreased prominence and posterior projection of the nuchal crests. N. gidleyi is autapomorphic in having an increased relative size of the glenoids, and in the apparent lack of an intratympanic route for the internal carotid plexus. P. cookei and P. tricuspidens have been traditionally thought to be close relatives. P. tricuspidens is also thought to be closely related to Platychoerops. The skull of P. cookei differs from that of P. tricuspidens in ways that make it similar to earlieroccurring North American plesiadapids. Unlike P. tricuspidens or other North American plesiadapids, P. cookei has a dentition indicating it had a diet focused on leaves. These features and a cladistic analysis of other dental characters suggest that P. cookei is a closer relative of Platychoerops than is P. tricuspidens. The postcranial skeleton of Plesiadapis cookei, while generally similar to that of other plesiadapids, has more elongate limbs, possibly reflecting a more specialized arboreal habitus. Finally, new information from plesiadapid skulls and postcrania increases cladistic support for previously proposed higher-level clades. Specifically, the new data support the hypotheses that plesiadapids, carpolestids, saxonellids and Chronolestes comprise the Plesiadapoidea, and that Plesiadapoidea is the sister taxon to Euprimates. Therefore the results of this dissertation corroborate the hypothesis that euprimates evolved their grasping features from a fine-branch foraging, common ancestor with plesiadapoids. iv
Page 1 and 2: NEW CRANIAL AND POSTCRANIAL REMAINS
Page 3: Stony Brook University The Graduate
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 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
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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 2.18. MNHN CR 125 Plesiadapi
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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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