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4. the Phylogeny and Disparity of the Odontopleurida (Trilobita)

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4. the Phylogeny and Disparity of the Odontopleurida (Trilobita) University of Bath PHD Trilobita: phylogeny and evolutionary patterns Pollitt, Jessica R. Award date: 2006 Awarding institution: University of Bath Link to publication Alternative formats If you require this document in an alternative format, please contact: [email protected] General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 10. Oct. 2021 TRILOBITA: PHYLOGENY AND EVOLUTIONARY PATTERNS Jessica R. Pollitt A thesis submitted for the degree of Doctor of Philosophy University of Bath Department of Biology & Biochemistry September 2006 COPYRIGHT Attention is drawn to the fact that copyright of this thesis rests with its author. This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with its author and that no quotation from the thesis and no information derived from it may be published without the prior written consent of the author. This thesis may be made available for consultation within the University Library and may be photocopied or lent to other libraries for the purposes of consultation. JjUJUbb UMI Number: U601710 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. Dissertation Publishing UMI U601710 Published by ProQuest LLC 2013. Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 ..'Nf/ERSITY OF BATH r LIBRARY I 6 JUL 2007 P h .T > . ABSTRACT TRILOBITES have been cited in the scientific literature for nearly 200 years but their evolutionary relationships are contentious. Studies resolving this issue are necessary in order to assemble a stable trilobite classification and also to facilitate further macroevolutionary studies on the group. However, less than fifty papers have been published using modern cladistic methods. This work investigated the phylogenies of four large trilobite groups: Lichoidea, Calymenina, Odontopleurinae and lllaenoidea. These four taxa have distinctive gross morphologies and present different challenges. Some taxa are well-known, others poorly; some are spiny and some exceptionally effaced. Fine resolution was attained in all resulting phylogenies. The systematic palaeontology of each group was reassessed accordingly and phylogenetically valuable character states were listed. A Bayesian phylogenetic method was employed in Chapter 2 to analyse morphological data for the first time: lichoids were used as a case study. The trees obtained were similar to those inferred using parsimony, with the exception of relationships between the deeper branches. Chapters 4 and 5 investigated the phylogeny of two large groups (odontopleurids and illaenoids respectively) and used the same character data to explore the disparity (morphological variety) of both groups. The disparity of odontopleurids decreased significantly through time, but that of illaenoids did not. The relative disparity of major clades within each group was also investigated. In summary, the results of this study provide a platform for future trilobite workers: important morphological distinctions have been identified between clades, phylogenetically-important character states recognised and temporal patterns of disparity for two higher taxa have been investigated. 2 ACKNOWLEDGEMENTS FIRST, many thanks to my supervisors, Matthew Wills and Richard Fortey, whose writings inspired me for many years before we even met. (What a wonderful opportunity you gave me: to be able to study my very favourite thing for so long!) Thanks to Matthew for everything: especially his help with all things numerical. I am very grateful to Richard for trilobite (and fungi) inspiration and lots of thought-provoking, happy Oxford afternoons. I am enormously indebted to Euan and Cecilia for agreeing to supervise me all those years ago in Edinburgh - for opening my eyes to the wonderful world of fossils - and for their continued support and valued friendship. Professor Hugh Drummond (UNAM) appeared unexpectedly in my life at a crucial time, armed with coffee. I thank him for helping me redefine and reflect on my ideals for the future and for his gentle guidance and friendship. Many thanks to Dr. Alan T. Thomas (University of Birmingham) and Dr. Tam&s Szekely (University of Bath) for being the examiners of my viva voce and for making it such a positive and constructive experience. The Bath University Biodiversity Lab has shaped this work, and all involved are thanked: especially Peter and Al for being carrier-pigeons in the last year. The Geological Society Publishing House is much appreciated: it appointed me in 2005 and has offered support and tea whenever the night-writing took its toll. To 10 Belgrave Crescent: for being a stimulating haven to write-up in. Helen Haste kindly provided a comfortable and happy atmosphere. And to the late Bev Halstead, who provided me with a wonderful, well-cherished library and, if he still disagrees with my cladistic methods, let it be... To Mary, Kate and Rachel: for your continued friendship, support and soothing of the trilo- traumas over the last few years. I couldn’t have done it without you. To John: my best friend. For always seeing what I mean (regardless of what I say!). For your unreserved support and for encouraging me to look at everything with fresh eyes. Lastly, to my parents and Simon: for your continued love and support and for appreciating the geek in your lives. I thank my parents especially for all their personal sacrifices in order that I can make life- choices selfishly and without restriction. This work is dedicated, in its entirety, to them. Thank you so much. 3 TABLE OF CONTENTS TITLE PAGE 01 ABSTRACT 02 ACKNOWLEDGEMENTS 03 TABLE OF CONTENTS 04 LIST OF ILLUSTRATIONS 09 LIST OF TABLES 14 LIST OF PLATES 15 CHAPTER 1: INTRODUCTION 1.1 WHAT ARE TRILOBITES: WHY ARE THEY USEFUL SUBJECTS? 17 1.1.1 General morphology 19 Dorsal 23 Ventral 24 Internal 26 Behavioural and sensory 26 1.1.2 Trilobite ontogeny and body patterning 27 1.2 PHYLOGENETICS: A (VERY) BRIEF HISTORY 28 1.3 THE PHYLOGENY OF TRILOBITES: AS CURRENTLY KNOWN 30 1.3.1 The Trilobita sensu stricto 30 1.3.2 Who makes up the group and what other cladistic studies have there been? 32 1.3.2.1 Order Redlichiida 1.3.2.2 Order ‘Ptychopariida' Swinnerton, 1915 1.3.2.3 Order Asaph ida Salter, 1864 1.3.2.4 Order Harpetida Ebach & McNamara, 2002 1.3.2.5 Order Proetida Fortey & Owens, 1975 1.3.2.6 Order Phacopida Salter, 1864 1.3.2.7 Order Lichida (Fortey in Kaesler, 1997) 1.3.2.8 Order Corynexochida Kobayashi, 1935 1.4 CHARACTERS AND THEIR SELECTION 42 1.5 REFERENCES 43 CHAPTER 2: SYSTEMATICS OF THE TRILOBITE FAMILIES LICHIDAE HAWLE & CORDA. 1847 AND LICHAKEPHALIDAE TRIPP. 1957: THE APPLICATION OF BAYESIAN INFERENCE TO MORPHOLOGICAL DATA 2.1 INTRODUCTION 57 2.2 METHODS 59 2.2.1 Maximum-parsimony analysis 59 2.2.2 Bayesian inference analysis 60 2.2.3 Outgroup 60 2.2.4 Characters 60 Cranidium 60 4 Hypostome 63 Thorax 63 Pygidium 64 2.3 RESULTS 64 2.3.1 Maximum-parsimony analysis 64 2.3.2 Bayesian inference analysis 66 2.4 DISCUSSION 66 2.5 SYSTEMATIC PALAEONTOLOGY 68 Order Lichida Moore, 1959 Family Lichidae Hawle & Corda, 1847 Genus Leiolichas Schmidt, 1885 Subfamily Lichinae Hawle & Corda, 1847 Tribe Platylichini Phleger, 1936 Tribe Tetralichini Phleger, 1936 Tribe Echinolichini Phleger, 1936 Tribe Lichini Phleger, 1936 Tribe Dicranopeltini Phleger, 1936 Subfamily Trochurinae Phleger, 1936 Family Lichakephalidae Tripp, 1957 2.6. ACKNOWLEDGEMENTS 70 2.7 REFERENCES 71 2.8 APPENDIX: Character dataset for the Lichoidea 73 CHAPTER 3: THE PHYLOGENY OF THE SUBORDER CALYMENINA (TRILOBITA) 3.1 TAXONOMIC HISTORY OF THE CALYMENOIDEA 75 3.2 DISTRIBUTION OF THE CALYMENOIDEA 76 3.3 PHYLOGENETIC ANALYSIS 77 3.3.1 Taxonomic sampling 77 3.3.2 Methods 80 3.3.2.1 Outgroup 82 3.3.2.2 Characters 82 Whole exoskeletal characters 82 Cranidium 83 Librigenae 89 Rostral plate and hypostome 90 Thorax 91 Pygidium 91 Ontogeny 92 3.3.3 RESULTS 93 3.4 DISCUSSION 96 3.5 SYSTEMATIC PALAEONTOLOGY 97 Order Phacopida Salter, 1864 Suborder Calymenina Swinnerton, 1915 Family Bavarillidae Sdzuy, 1957 Family Homalonotidae Chapman, 1890 Subfamily Eohomalonotinae Hupe, 1953 Subfamily Homalonotinae Thomas, 1977 Subfamily Trimerinae Hupe, 1953 5 Family Bathycheilidae Pribyl, 1953 Genus Protocalymene Ross, 1967 Superfamily Calymenoidea Fortey in Kaesler, 1997 Genus Calymenella Bergeron, 1890 Genus
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