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MORPHOLOGICAL CHARACTERIZATION OF FOSSIL GWM10/P1, A PHALANX OF ARDIPITHECUS RAMIDUS by YVONNE P. McDERMOTT Submitted in partial fulfillment of the requirements for the degree of Master of Science Department of Biology CASE WESTERN RESERVE UNIVERSITY January 2019 CASE WESTERN RESERVE UNIVERSITY SCHOOL OF GRADUATE STUDIES We hereby approve the thesis of Yvonne McDermott candidate for the degree of Master of Science Committee Chair Karen Abbott, Ph.D Committee Member/Research Advisor Scott Simpson, Ph.D. Committee Member Michael Benard, Ph.D. Committee Member Bruce Latimer, Ph.D. Date of Defense August 31, 2018 We also certify that written approval has been obtained for any proprietary material contained therein. 2 Dedication This thesis is dedicated to Quinn, Shay, Tristane and Brian for all their love, support and encouragement. 3 Table of Contents List of Tables, 6 List of Figures, 7 Acknowledgements, 11 Abstract, 12 Introduction, 13 General features and functions of primate hands, 13 Modes of locomotion, 15 Bipedalism, 15 Knuckle walking, 17 Plantigrade, digitigrade, and suspensory locomotion, 20 Hands as predictors of locomotion, 24 Phalangeal curvature, 25 The hand and the emergence of tool use, 27 The fossil record with an emphasis on the hand, 28 Orrorin tugenensis, 28 Ardipithecus ramidus, 28 Australopithecus anamensis, 32 Australopithecus afarensis, 33 Australopithecus sediba, 34 OH 86, 36 Methods and materials, 38 Measurements, 39 Phalangeal curvature, 41 Articular surface area, 44 Torsion, 44 Graphical plots and statistics, 44 Results, 46 The morphology of proximal phalanges, 47 Proximal phalanx 1 of humans, gorillas, and chimpanzees, 47 Proximal phalanx 2 of humans, gorillas, and chimpanzees, 54 Proximal phalanx 3 of humans, gorillas, and chimpanzees, 59 Proximal phalanx 4 of humans, gorillas, and chimpanzees, 61 Proximal phalanx 5 of humans, gorillas, and chimpanzees, 62 GWM10/P1, 65 Metrical comparisons of proximal phalanges, 71 Lengths of proximal phalanges of humans, gorillas, and chimpanzees, 71 Proximal articular surface area (PAS) of phalanges as a proxy for body mass and its relation to phalangeal length, 75 4 Axial torsion, 82 Comparison of phalangeal shapes, 89 Phalangeal curvature, 97 Discussion, 101 The morphology of proximal phalanges, 101 Torsion as a tool for manual proximal phalanx ray assignment, 101 Morphology and metrical descriptions of manual proximal phalanges of humans, gorillas, and chimpanzees, 102 GWM10/P1 hand side and ray, 104 GWM10/P1 is long and curved, 105 Ar. ramidus had a hand similar to a Miocene ape, a pelvis for arboreality and bipedal terrestriality and a phalanx suited for arboreality, 107 The relation of phalangeal length and the transition between arboreality and bipedality, 109 Appendix, 112 References, 116 5 List of Tables Table R1.Distinguishing features of the proximal phalanges, 64 Table R2. Measurements of GWM10/P1 and other 4th proximal phalanges, 73 Table R3. Mean lengths of proximal phalanges of chimpanzees, gorillas and humans, 74 Table R4. Median torsion values, 87 Appendix Table 1, Gorilla specimen information, 112 Appendix Table 2, Chimpanzee specimen information, 113 Appendix Table 3, Human specimen information, 114 Appendix Table 4, Fossil specimen information, 115 6 List of Figures Figure 1. Bones of the hands of extant and extinct hominins and their predecessors, 15 Figure 2. Hand bone positions of knuckle walker, 19 Figure 3. Modes of locomotion by primates, 21 Figure 4. Orangutan engaging in quadrumanous clambering, above-branch quadrupedalism with a gripping hand, suspensory locomotion, and hand-assisted arboreal bipedalism, 23 Figure 5. Lateral view illustrating varying degrees of curvature of the manual proximal phalanges of extant anthropoids and early fossil hominins, 26 Figure 6. Dorsal and palmar view of the hand of Ar. ramidus, 31 Figure 7: Four views of proximal phalanx KNM-KP 30503, 33 Figure 8. Images of palmar and dorsal aspects of Au. sediba MH2 right hand, 35 Figure 9. Images of OH 86, a manual proximal phalanx, 36 Figure M1. Eleven linear measurements acquired from manual proximal phalanges, 40 Figure M2. Determination of included angle of a proximal phalanx, 42 Figure R1. Right hand proximal phalanges of rays 1 to 5 of a human, 48 Figure R2. View of the articular surfaces of the 1st proximal phalanges from a human, gorilla and chimpanzee from left to right, 48 Figure R3. Left hand proximal phalanges of rays 5 to 1 of a female gorilla and a female chimpanzee, 49 Figure R4. Human proximal phalanx 1 of the right hand, palmar surface, displays the distal ulnar condyle extending in the palmar direction and the distal direction, 51 7 Figure R5. Palmar surfaces of chimpanzee and gorilla proximal phalanges 1 of the right hand display the distal ulnar condyles extending in the palmar direction and distal direction, 52 Figure R6. Comparison of side views of proximal phalanges 1 of gorillas, chimpanzees, and humans, 53 Figure R7. Gorilla distal articular surface does not extend to the dorsal aspect for proximal phalanx 1, 53 Figure R8. Gorilla proximal phalanx 2 and 4 of right hand displaying proximal surfaces, 55 Figure R9 – Proximal articular surfaces of human proximal phalanges 2 and 4 of the right hand, 56 Figure R10. Chimpanzee proximal phalanges 2 (left) and 4 (right) of the right hand display proximal surfaces, 56 Figure R11. Human proximal phalanx 2 of the right hand shows that the ulnar side distal condyle extends further in the palmar direction than the radial condyle, 57 Figure R12. Gorilla proximal phalanx 2 of the right hand, 57 Figure R13. An oblique view of the palmar aspects of gorilla proximal phalanges, 2-5, left to right, 58 Figure R14. Palmar surface view of proximal phalanx 3 of humans, gorillas, and chimpanzees from left to right of the right hand, 60 Figure R15. Articular surfaces of proximal phalanx 5 of humans, chimpanzees and gorillas from left to right of the right hand, 62 8 Figure R16. Palmar sides of the 5th proximal phalanges of humans, gorillas and chimpanzees of the right hand, 63 Figure R17. GWM10/P1 in dorsal, palmar and side view, 66 Figure R18. Side view of GWM10/P1 and a human 4th proximal phalanx (bottom) comparing longitudinal curvature, 69 Figure R19. Proximal articular surface of GWM10/P1 cast, 69 Figure R20. Dorsal view of GWM10/P1 and A. L. 333-63 (Au. afarensis), 70 Figure R21. GWM10/P1 condyles at the distal surface, 70 Figure R22. Comparisons of PAS areas of proximal phalanges of ray 2 of chimpanzees, gorillas, and humans, 78 Figure R23. Evaluations of PAS areas of proximal phalanges of ray 3 of chimpanzees, gorillas, and humans, 78 Figure R24. Assessments of PAS areas of proximal phalanges of ray 4 of chimpanzees, gorillas, humans, Hadar, and GWM10/P1, 79 Figure R25. Plots of the square root of the PAS area of each proximal phalanx from ray 2, 3, and 4 against total length of each cognate bone from chimpanzees, gorillas, humans, Au. afarensis, and GWM10/P1, 80 Figure R26. Ratios of the means of the square root of the PAS of proximal phalanx/TL ± SEM for each species, 81 Figure R27. Image of axial torsion for proximal phalanx of Au. afarensis AL 333-62, a likely fourth proximal phalanx of the left hand, 83 Figure R28. Images of distal and proximal articular surfaces of proximal phalanges of gorillas, chimpanzees, and humans, 84 9 Figure R29. Torsion values in degrees, 86 Figure R30. Comparison of proximal phalanges 2 and 4 mean torsion values ± SEM of humans, gorillas, and chimpanzees, 88 Figure R31. Base shape comparisons, 90 Figure R32. Midshaft shape comparisons, 91 Figure R33. Head shape comparisons, 92 Figure R34. Proximal shaft shape comparisons, 94 Figure R35. Distal Shaft Shape ratio comparisons, 95 Figure R36. Linear regression performed on all distal shaft shape ratios demonstrate a correlation between distal shaft shapes and length, 96 Figure R37. Representative phalangeal curvature analyses performed on digital photos with ImageJ, 98 Figure R38. Curvature of the 4th proximal phalanges from fossil GWM10/P1 and those of extant and extinct hominoids, 100 10 Acknowledgments First and foremost, I would like to thank my advisor, Dr. Scott Simpson, for his constant support along this process. I thank him for giving so much of his time in guiding me through this fascinating and complicated field. For me, it was worth all the work. I am very grateful to Dr. Bruce Latimer for being so generous with his time, knowledge and guidance. I would like to thank Dr. Michael Benard for agreeing to be a committee member and for his advice and guidance. I would like to thank Dr. Karen Abbott for being my DGS representative. I would like to thank Dr. Yohannes Haile-Selassie for allowing me to have access to the Physical Anthropology Laboratory at the Cleveland Museum of Natural History and the research collection kept there. I am so grateful to Lyman Jellema for always being so accommodating and helpful at the Physical Anthropology lab. I would like to thank Dr. Linda Spurlock for her encouragement and friendship and all our wonderful conversations about this crazy field we love so well. I am very grateful to Dr. Mark Hans for making my dream come true and making me part of an incredible Archaeological Excavation. I would like to thank the wonderful scientists in Israel that I have had the extreme good fortune to get to know as part of the Manot Cave Project, such as Dr. Ofer Marder and Dr. Israel Hershkovitz. I thank them for their support and many thoughtful and inspiring conversations. I would like to especially thank my guardian angel, Julia Brown. I can’t imagine having been able to navigate this journey without her. 11 Morphological Characterization of Fossil GWM10/P1, a Proximal Manual Phalanx of Ardipthecus ramidus Abstract YVONNE P.
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    The Vertebral Remains of the Late Miocene Great Ape Hispanopithecus Laietanus from Can Llobateres 2 (Valles-Pened Es Basin, NE Iberian Peninsula)

    Journal of Human Evolution 73 (2014) 15e34 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol The vertebral remains of the late Miocene great ape Hispanopithecus laietanus from Can Llobateres 2 (Valles-Pened es Basin, NE Iberian Peninsula) * Ivette Susanna a, David M. Alba a, b, , Sergio Almecija c, a, d, Salvador Moya-Sol a e a Institut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICP, Campus de la UAB s/n, 08193 Cerdanyola del Valles, Barcelona, Spain b Dipartimento di Scienze della Terra, Universita degli Studi di Torino, Via Valperga Caluso 35, 10125 Torino, Italy c Department of Anatomical Sciences, Stony Brook University School of Medicine, Stony Brook, NY 11794-8081, USA d NYCEP Morphometrics Group, USA e ICREA at Institut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICP, Campus de la UAB s/n, 08193 Cerdanyola del Valles, Barcelona, Spain article info abstract Article history: Here we describe the vertebral fragments from the partial skeleton IPS18800 of the fossil great ape Received 15 November 2012 Hispanopithecus laietanus (Hominidae: Dryopithecinae) from the late Miocene (9.6 Ma) of Can Llobateres Accepted 7 May 2014 2 (Valles-Pened es Basin, Catalonia, Spain). The eight specimens (IPS18800.5eIPS18800.12) include a Available online 18 June 2014 fragment of thoracic vertebral body, three partial bodies and four neural arch fragments of lumbar vertebrae. Despite the retention of primitive