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Proquest Dissertation Femur 2-4 Tibia 2 Fibula 1 Patella Pretibial bone Tarsals Metatarsal I 8 Phalanx 1 9 Phalanx 2 9 Metatarsal II 4 Phalanx 1 8 Phalanx 2 9 Phalanx 3 9 Metatarsal III 2 Phalanx 1 8 Phalanx 2 9 Phalanx 3 9 Phalanx 4 9 Metatarsal IV 2 Phalanx 1 8 Phalanx 2 9 Phalanx 3 12 Phalanx 4 11 Phalanx 5 9 If an element is unnumbered, it was unossified in all specimens examined. If two numbers are given, these represent the range of ranks over which a variable element can ossify. 282 Evolution of Avian Ossification Sequences Erin E. Maxwell Department of Biology McGill University, Montreal April 2008 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Doctor of Philosophy © Erin Maxwell 2008 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de ('edition 395 Wellington Street 395, rue Wellington OttawaONK1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-53317-8 Our file Notre reference ISBN: 978-0-494-53317-8 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantias de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1*1 Canada ACKNOWLEDGMENTS Discussion and Support I would like to thank my supervisor Hans Larsson, my supervisory committee (Ehab Abouheif and Bob Carroll) and the members of the Larsson- Carroll lab for discussion. Special thanks to Audrey Heppleston, Maria de Boef and Sylvie Tissandier for assisting with the lab work, and to Luke Harrison and Nadia Frobisch for commenting on early drafts of Chapter 1. Luke Harrison also provided essential assistance with programming issues. Matt Vavrek and Maria de Boef helped with proofreading, and Julie Rousseau translated the abstract. The comments of anonymous reviewers improved Chapters 3, 4 and 5. Collections Access I appreciate the efforts of Virginie Millien and Anthony Howell for accessioning the Redpath Museum specimens for me. I would also like to thank Kristof Zyskowski, Richard Prum and the other Peabody Museum vertebrate zoology staff for their hospitality and allowing me access to the collections to collect the data used in Chapter 2. Thanks to Parker Cane for access to the cleared and stained Common Tern specimens, as well as Paul Sweet, Joel Cracraft and other members of the AMNH Department of Ornithology for allowing me access to the collections to assemble the data presented in Chapter 5. Thanks to David Evans for allowing me access to the ROM collections for comparative purposes. Specimen Collection I would like to thank Dr. Katherine Mehl and the members of the Ducks Unlimited NL/Labrador field team (2004/2005) for collecting Common Eider embryos for me. Thanks also to the Presqu'ile ProvincialPark staff (2005) for collecting Double-crested Cormorant eggs on my behalf. Thanks to the Biodome de Montreal for providing me with access to altricial embryos, used for comparative purposes. i Funding Funding for this project was provided by a Tomlinson Fellowship, a Wylie Memorial Fellowship, FQRNT, and NSERC CGS-M and CGS-D fellowships to myself, as well as an NSERC Discovery Grant and Canada Research Chair to H. Larsson which covered logistical expenses. A collections study grant helped cover the cost of the collections visit to the AMNH. 11 CONTRIBUTIONS OF AUTHORS Chapter 5, "Ossification Sequence of the Common Tern (Sterna hirundo) and Its Implications for the Interrelationships of the Lari (Aves, Charadriiformes)" (In Press, Journal of Morphology) is a manuscript I co-authored with Luke Harrison. I examined the embryos and reconstructed the ossification sequence. I wrote all portions of the manuscript not directly pertaining to PGi analysis, and made figures 2-5. 111 TABLE OF CONTENTS Acknowledgments i Contributions of authors Hi ABSTRACT/RESUME 1 INTRODUCTION 4 CHAPTER 1: Analysis of developmental sequence data: a review Introduction 8 Phenetic Methods 10 Phylogenetic Methods 16 Mapping sequences on an existing tree topology 16 The problem of simultaneity 17 The problem of polymorphism 18 The use of developmental sequence data for phylogenetic reconstruction 23 Conclusions 26 CHAPTER 2: Skeletal development in palaeognathous birds Introduction 28 Materials and Methods 30 Results Skull Dromaius novaehollandiae 31 Rhea americana 35 Struthio camelus 36 Eudromia elegans 39 Postcranial axial skeleton Dromaius novaehollandiae 42 Rhea americana 45 Struthio camelus 46 Eudromia elegans 47 Forelimb Dromaius novaehollandiae 48 Rhea americana 50 Struthio camelus 51 Eudromia elegans 52 Hind limb Dromaius novaehollandiae 55 Rhea americana 57 Struthio camelus 58 IV Eudromia elegans 60 Discussion Skull 63 Paedomorphosis and ratite cranial evolution 67 Postcranial axial skeleton 69 Forelimb 71 Hind limb 75 Incubation period, adult size and ossification 77 Mechanisms of skeletal reduction 78 Morphology and ossification sequence 79 CHAPTER 3: Comparative embryonic development of the skeleton of the Domestic Turkey (Meleagris gallopavo) and other galliform birds Introduction 87 Materials and Methods 88 Results Sequence variability 89 Skull 89 Postcranial axial skeleton 94 Forelimb 95 Hind limb 98 Discussion Skull 100 Postcranial axial skeleton 101 Forelimb 102 Hind limb 103 Factors affecting ossification sequence 105 CHAPTER 4: Ossification Sequence of the Avian Order Anseriformes, with Comparison to other Precocial Birds Introduction 115 Materials and Methods 117 Results Sequence variability 118 Skull 119 Postcranial axial skeleton 125 Forelimb 128 Hind limb 132 Discussion Sequence variability 137 Comparative ossification sequence Skull 137 Postcranial axial skeleton 140 Forelimb 141 Hind limb 142 v Life history and ossifcation sequence 143 CHAPTER 5: Ossification Sequence of the Common Tern {Sterna hirundo) and Its Implications for the Interrelationships of the Lari (Aves, Charadriiformes) Introduction 156 Materials and Methods 160 Results Skeletal development of Sterna hirundo Skull 163 Post-crartial axial skeleton 169 Forelimb 171 Hind limb 175 Intraspecific variability 178 Phylogenetic analysis Parsimony analysis of event-pairs 178 PGi analysis - 179 Discussion Skull 180 Post-cranial axial skeleton 181 Forelimb 182 Hind limb 183 Potential factors affecting ossification sequence 184 Intraspecific variability 186 Phylogenetic analysis 186 CHAPTER 6: Phylogenetic analysis of avian ossification sequence data Introduction 193 Materials and Methods 195 Results Topology reconstruction 197 Character evolution 198 Discussion 202 CONCLUSIONS AND FUTURE RESEARCH 250 BIBLIOGRAPHY 253 APPENDICES Appendix 1: List of specimens 274 Appendix 2: Ossification sequence of the Double-crested Cormorant 281 Appendix 3: Supporting documents 283 vi ABSTRACT The relative timing and sequence of events during embryonic development plays an important role in adult shape and thus in evolution. The sequence in which bones form in the developing embryo should therefore contain a component capable of revealing evolutionary history, however processes relating to ossification sequence and the sequences themselves are poorly known and rarely discussed. In this thesis, I describe the embryonic skeletal development of Meleagris gallopavo, Sterna hirundo, Somateria mollissima, Anas platyrhynchos, Cairina moschata, Dromaius novaehollandiae, Rhea americana and Struthio camelus for the first time in the scientific literature, focusing on ossification. All species exhibited intraspecific variation in ossification sequence, but the level of polymorphism present was generally quite low. Specimens collected from the wild did not show more variability in ossification sequence than those incubated under constant conditions in the lab. Dermal bones did not always ossify before endochondral bones, nor did neural-crest derived elements always form before elements derived from the paraxial mesoderm. All of this suggests that the factors controlling ossification sequence are complex, and more than one variable may play a role. In order to examine sequences in a more explicit phylogenetic context, I converted them
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