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Taxonomy of Proconsul: an Issue of Species

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Taxonomy of Proconsul: an Issue of Species TAXONOMY OF PROCONSUL: AN ISSUE OF SPECIES NUMBERS OR SEXUAL DIMORPHISM? HUGH WEAVER Submitted in total fulfilment of the requirements of the degree of Master of Philosophy October 2015 Department of Anatomy and Neuroscience University of Melbourne ABSTRACT I have investigated the alpha taxonomy of Miocene primate genus Proconsul by performing measurements on photographs of fossil dental material obtained from museum sources. My aim was to assess levels of variation, and thereby evidence for sympatric species, by comparing results with those obtained from sex-matched samples of extant hominoids. The holotype of the initial species described, Proconsul africanus, came from a mainland site in western Kenya. Additional examples of the genus were obtained subsequently from adjacent sites and from islands within Lake Victoria. By 1951, three species of progressively-increasing size were recognised, seemingly present at both island and mainland localities. However, subsequent investigations questioned whether, instead of two sympatric species at a particular site, particularly Rusinga Island, size disparities noted reflected the presence of a single sexually dimorphic species. I addressed this debate by comparing results of measurements of 144 Proconsul specimens with those from sex-matched samples comprising 50 specimens each of four extant primate genera: Pan, Gorilla, Pongo and Hylobates/Symphalangus. The protocol consisted of measuring occlusal views of adult molars to obtain linear measurements and cusp areas. By selecting samples on a 1:1 sex-matched basis for extant groups, and matching these against those for Proconsul, I minimised the potential that variations within the Proconsul material reflected sexual dimorphism within one species. I have investigated the situation relevant to each of two clusters of primate fossil sites, respectively on Rusinga and Mfangano Islands (Area 1) and at mainland sites around Koru (Area 2). The study has addressed a null hypothesis: that at each Area, only one Proconsul species was present from the relevant time-horizon. Analyses confirmed that levels of variation within Proconsul, at each of Areas 1 and 2, outweighed those within extant hominoids. More than one species of Proconsul existed contemporaneously at each cluster of Miocene fossil sites. i DECLARATION This is to certify that (i) the thesis comprises only my original work towards the MPhil except where indicated in the preface. (ii) due acknowledgement has been made in the text to all other material used. (iii) the thesis is less than 50,000 words in length, exclusive of tables, maps and bibliographies. ……………………………………………….. 26 October 2015 ii PREFACE The primary material utilised in this study has consisted exclusively of two-dimensional photographic images of primate teeth; this material has been subjected to appropriate measurement using a dedicated software package and the results obtained in turn submitted to statistical analysis. The material was obtained by my supervisor, Dr. Varsha Pilbrow, from museum sources in 1999. Suitable acknowledgement of this primary source has been provided at appropriate points throughout the study which follows. However, I can confirm that I performed all measurements undertaken on this material, employing an appropriate software package in order to do so. I checked the data set and then performed a sequence of statistical analyses, from which I was able to draw the conclusions identified later in this paper. To this extent, the work undertaken has been entirely my own. No part of the work has been submitted for purposes of consideration for the award of any qualification other than the current application for the degree of Master of Philosophy. No work of any kind, either utilising the primary source material, or of any other kind, had been undertaken prior to my enrolment for candidature. iii ACKNOWLEDGEMENTS The genesis of this project was provided for me by Dr. Varsha Pilbrow, Lecturer in the Department of Anatomy and Neuroscience, University of Melbourne. I approached Varsha in 2012, asking if she would consider providing me with some suggestions whereby I might undertake a project within the field of physical anthropology. Although I had for many years possessed an amateur interest in this area of academic study, I had never previously had the opportunity to pursue such an interest at any professional level. Varsha kindly agreed to take me on as a postgraduate student and was entirely responsible for the concept of the project which I have undertaken, namely an attempt to address the continuing debate regarding taxonomy within the primate genus Proconsul. It was her suggestion that I might attempt to address levels of variation within Proconsul dental material by comparing them with levels obtained from similar examples from extant taxa. As has been indicated in the Preface, Varsha was able to furnish the source material which provided the basis for my project. She has been responsible for suggesting the programs of statistical analysis which were best suited to assess the results obtained from the data set; and she has supplied a critical eye throughout, particularly with regard to the conclusions which might be drawn from the entire process. It goes without saying that I am grateful to her for her support throughout. Any deficiencies which afflict the final product are my responsibility entirely. My gratitude goes also to Associate Professor Christopher Briggs, Department of Anatomy & Neuroscience, University of Melbourne. Chris has provided continuing moral support to me throughout the process. The first draft of the thesis was assessed by two external examiners and I am grateful for the criticisms which they have provided, the result of which I trust is an improved study overall. I am grateful to my brother, Phillip Weaver, for the assistance which he provided in helping to prepare my bivariate plots. His help was invaluable in making up for some of my deficiencies in word processing. My thanks go to my wife, Pam, who has, as in so many other areas throughout the course of our partnership, provided her ongoing love and support. Pam encouraged me constantly iv to persevere on many occasions when I felt as if the project was becoming too much for me. I can only trust that the finished project will go some way towards justifying the faith she showed in me. v TABLE OF CONTENTS Abstract ...................................................................................................................... i Declaration................................................................................................................. ii Preface ...................................................................................................................... iii Acknowledgements ................................................................................................... iv Table of Contents ...................................................................................................... vi List of Tables .............................................................................................................. x List of Figures ........................................................................................................... xii INTRODUCTION 1 Proconsul: Issues of Taxonomy and Phylogeny ............................................................ 1 This Study................................................................................................................... 2 CHAPTER ONE: HISTORICAL SURVEY AND LITERATURE REVIEW 4 Background ................................................................................................................ 4 Geochronology ........................................................................................................... 8 The island and mainland fossil sites........................................................................... 10 Songhor/Meswa Bridge ............................................................................................ 12 Rusinga Island; The British-Kenya Miocene Expeditions 1948-50; further work at Songhor/Koru ........................................................................................................... 13 Collections on Rusinga Island: Site R106; Whitworth’s ‘pothole’ at R114; R5, the Kaswanga Primate Site ............................................................................................. 14 Review of Earlier Findings ........................................................................................ 16 Subsequent Issues: species numbers or sexual dimorphism? ..................................... 17 Updated Positions .................................................................................................... 21 vi CHAPTER ONE: HISTORICAL SURVEY AND LITERATURE REVIEW (cont.) The Alternative View: two species, rather than extreme sexual dimorphism .............. 21 Distinctiveness of Proconsul heseloni at Rusinga from Proconsul africanus at Koru ..... 22 Support from postcranial material ............................................................................ 24 Proconsul meswae .............................................................................................. 25 Rangwapithecus .................................................................................................. 25 Ugandapithecus .................................................................................................. 26 The Initial Concept….………………………………………………………………………………………….
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