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The Genus Category and Cranial Morphometrics of the Catarrhini with Implications for Fossil Hominins

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The Genus Category and Cranial Morphometrics of the Catarrhini with Implications for Fossil Hominins The genus category and cranial morphometrics of the Catarrhini with implications for fossil hominins Jack Andrew Coate A thesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy Postgraduate Board University of New South Wales September 2007 PLEASE TYPE THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet Surname or Family name: Coate First name: Jack Other name/s: Andrew Abbreviation for degree as given in the University calendar: PhD School: SOMS Faculty: Anatomy Title: The genus category and cranial morphometrics of the Catarrhini with implications for fossil hominins Abstract 350 words maximum: (PLEASE TYPE) Recently, the number of hominin genera has increased dramatically. Prior to the announcement of Ardipithecus, only two genera were used by paleoanthropologists: Australopithecus and Homo. Presently, up to eight hominin genera are used: Sahelanthropus, Orrorin, Ardipithecus, Australopithecus, Praeanthropus, Kenyanthropus, Paranthropus and Homo. Unlike species concepts, the genus category has not received wide critical examination. To investigate the use of the genus category in paleoanthropology, a comparative framework drawing on morphometric data from a large number of catarrhines is developed. Cranial variables include 36 standard linear measurements from representatives of catarrhine genera across the major tribes/families. This study seeks to assess whether too few or too many hominin genera have been recognized compared with extant catarrhines. Moreover, two published hypotheses about the use of Homo are examined: 1) Wood & Collard’s (1999) proposal to transfer Homo habilis/rudolfensis to Australopithecus; and 2) Goodman et al’s (1998) classification of both humans and chimpanzees in Homo. To analyze these cranial variables and a number of shape indices calculated from them, as well as to assess competing hypotheses, univariate, bivariate and multivariate statistical approaches are used. The results allow the identification of a set of variables and shape indices which distinguish genera across the catarrhines. Importantly, body size seems to be the major separator of catarrhine genera, reinforcing the idea that they occupy discrete adaptive zones. Moreover, differences between these genera mostly represent contrasts in the size of the neuro- versus the viscerocranium. When applied to hominins, a picture emerges which distinguishes them from extant catarrhines: cranial shape rather than size is the major component distinguishing them; this suggests that extinct hominins occupied similar habitats and adaptive zones; variability in size and shape within hominin genera is much lower than extant catarrhines; and the major differences seen in shape among hominins are the result of encephalization in Homo. It is concluded here that both Wood & Collard’s (1999) and Goodman et al.’s (1998) proposals appear to be premature. Moreover, while the earliest hominins may be too finely split at the genus level, the evidence for distinction of Australopithecus and Paranthropus is solid. Declaration relating to disposition of project thesis/dissertation I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only). …………………………………………………………… ……………………………………..……………… ……….………4/14/07...…….… Signature Witness Date The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing. Requests for a longer period of restriction may be considered in exceptional circumstances and require the approval of the Dean of Graduate Research. FOR OFFICE USE ONLY Date of completion of requirements for Award: THIS SHEET IS TO BE GLUED TO THE INSIDE FRONT COVER OF THE THESIS Abstract: Recently, the number of hominin genera has increased dramatically. Prior to the announcement of Ardipithecus, only two genera were used by paleoanthropologists: Australopithecus and Homo. Presently, up to eight hominin genera are used: Sahelanthropus, Orrorin, Ardipithecus, Australopithecus, Praeanthropus, Kenyanthropus, Paranthropus and Homo. Unlike species concepts, the genus category has not received wide critical examination. To investigate the use of the genus category in paleoanthropology, a comparative framework drawing on morphometric data from a large number of catarrhines is developed. Cranial variables include 36 standard linear measurements from representatives of catarrhine genera across the major tribes/families. This study seeks to assess whether too few or too many hominin genera have been recognized compared with extant catarrhines. Moreover, two published hypotheses about the use of Homo are examined: 1) Wood & Collard’s (1999) proposal to transfer Homo habilis/rudolfensis to Australopithecus; and 2) Goodman et al.’s (1998) classification of both humans and chimpanzees in Homo. To analyze these cranial variables and a number of shape indices calculated from them, as well as to assess competing hypotheses, univariate, bivariate and multivariate statistical approaches are used. The results allow the identification of a set of variables and shape indices which distinguish genera across the catarrhines. Importantly, body size seems to be the major separator of catarrhine genera, reinforcing the idea that they occupy discrete adaptive zones. Moreover, differences between these genera mostly represent contrasts in the size of the neuro- versus the viscerocranium. When applied to hominins, a picture emerges which distinguishes them from extant catarrhines: cranial shape rather than size is the major component distinguishing them; this suggests that extinct hominins occupied similar habitats and adaptive zones; variability in size and shape within hominin genera is much lower than extant catarrhines; and the major differences seen in shape among hominins are the result of encephalization in Homo. It is concluded here that both Wood & Collard’s (1999) and Goodman et al.’s (1998) proposals appear to be premature. Moreover, while the earliest hominins may be too finely split at the genus level, the evidence for distinction of Australopithecus and Paranthropus is solid. II Acknowledgements: I would personally like to thank my supervisor and co-supervisor, Dr. D. Curnoe and Prof. K. Ashwell, as well as, many other close friends and colleagues, including; H. Cohen, E. Danielson and family, T. Furlong, H. Green, D. Harris, Dr. A. Herries, S. Kiker, J. Louys, K. McQualter, D. Neuweger, Dr. S. van Holst, and L. van der Weyde. In addition, I would like to especially thank my parents and family for emotional and financial support throughout the years and during this thesis project. Furthermore, I would also like to personally thank the following institutions and their Physical Anthropology, Zoology and/or Mammalogy staff for allowing me to gather craniometric data from primate skulls in their care – Dr. S. Ingleby and T. Ennis at the Australian Museum, Sydney, Australia; Prof. F. Thackeray, Dr. T. Kearney, S. Potze and T. Perregil at the Transvaal Museum, Pretoria, South Africa; Prof. M. Raath, M. Dayal, M. Spocter, B. Billings and J. Hemingway at the University of Witwatersrand, School of Anatomical Sciences, Jo-burg, South Africa; J. Spence and E. Sarmiento at the American Museum of Natural History, NYC, NY, USA; Dr. Y. Haile-Selassie, L. Jellema and Prof. B. Latimer at the Cleveland Museum of Natural History, Cleveland, OH, USA, which houses the Hamann-Todd Collection; M. Schulenburg at the Field Museum, Chicago, IL, USA; L. Gordon at the Smithsonian Institute, National Museum of Natural History, Washington, D.C., USA; and P. Jenkins and D. Hills at the Natural History Museum, London, UK. In addition, Dr. R. Raaum and Dr. A. Tosi provided genetic information and distance matrices for some of the primates studied herein and both are greatly appreciated by the author for their data and assistance. III Table of contents: Abstract: II Acknowledgements: III Table of contents: IV Abbreviations: V-VII Chapter 1: Introduction, pp.1-18. Chapter 2: Materials and Methods, pp.15-50. Chapter 3: Results for genera of Cercopithecini, pp.51-113. Chapter 4: Results for genera of Papionini, pp.114-179. Chapter 5: Results for genera of Colobinae, pp.180-277 Chapter 6: Results for genera of Hominoidea, pp.278-337 Chapter 7: Summary and Discussion of results for the extant Catarrhini genera, pp.338- 356. Chapter 8: Results for genera of the Hominina, pp.357-412. Chapter 9: Conclusions, pp.413-420. References: pp.421-489. Appendix: pp.490-500. IV Anatomical, Genus and Journal Abbreviations (in alphabetical order): AJP – American Journal of Primatology AJPA – American Journal of Physical Anthropology Al. – Allenopithecus An. – Ankarapithecus Ar. – Ardipithecus Au. – Australopithecus Bas – Basion Bien – Interentoglenoid Breadth bp – basepairs Br - Bregma Cc. – Cercocebus Cp. – Cercopithecus Ch. – Chlorocebus Co. – Colobus Cum. – Cumulative CV – Coefficient of Variation
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