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Durham E-Theses Durham E-Theses Studies on the morphology and taxonomy of the immature stages of calliphoridae, with analysis of phylogenetic relationships within the family, and between it and other groups in the cyclorrhapha (diptera) Erzinclioglu, Y. Z. How to cite: Erzinclioglu, Y. Z. (1984) Studies on the morphology and taxonomy of the immature stages of calliphoridae, with analysis of phylogenetic relationships within the family, and between it and other groups in the cyclorrhapha (diptera), Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/7812/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 studies on the Morphology and Taxonomy of the Immature Stages of Calliphoridae, with Analysis of Phylogenetic Relationships within the Family, and between it and other Groups in the Cyclorrhapha (Diptera) Y.Z. ERZINCLIOGLU, B.Sc. The copyright of this thesis rests with the author. No quotation from it should be published without his prior written consent and information derived from it should be acknowledged. ..being a thesis presented in candidature for the degree of Doctor of Philosophy in the University of Durham 1984 11. FEB.19G^ ABSTRACT The present study deals with the taxonomy and morphology of the immature stages of the Diptera family Calliphoridae (blowflies). The eggs were studied, using both the light and the scanning electron microscopes, and structural features were found that enable specific identification. The larvae and puparia of some sixty species were studied, and a large number of new characters were found that enable the separation of species reliably. In addition, various anatomical details, especially of the cephalopharyngeal skeleton, have been elucidated. The cephalopharyngeal skeletons of various related families were studied and compared with the Calliphorid structure. Descriptions of the larvae of sixty species are presented, and keys to various groups, especially those of medical or veterinary importance, are given. A discussion of the evolution and phylogeny of the Calliphoridae and related families is presented, based on the above-mentioned morphological studies and other evidence. A N N E M E (TO MY MOTHER) though there are animals which have no attractiveness for the senses, yet for the eye of science, for the student who is naturally of a philosophic spirit and can discern the causes of things. Nature which fashioned them provides joys which cannot be measured. Aristotle De Partibus Animalium - (v) - CONTENTS Page Acknowledgements (x) 1. INTRODUCTION 1 1.1 The Value of Anatomical and Taxonomic Studies on Calliphorid larvae 1 1.2 The Aims and Scope of the Present Work 4 1.3 Previous Work 6 1.4 Species Studied 8 2. MATERIALS AND METHODS 11 2.1 Sources of Material 11 2.2 Culture Methods 11 2.3 General Laboratory Methods 12 2.4 Sectioning Methods 13 2.5 Scanning Electron Microscopy 13 2.6 Illustration 14 3. THE MORPHOLOGY OF THE EGGS 15 3.1 The Literature on Calliphorid Eggs 15 3.2 General Morphology of Calliphorid Eggs 16 3.3 Taxonomic Value of Egg Characters 17 3.3.1 Colour 18 3.3.2 Size and Shape 18 3.3.3 Chorionic Structure 20 3.3.4 Micropyle and Micropylar Plate 21 3.3.5 Median Area 22 3.3.6 Hatching Pleats 20 3.3.7 Plastron Craters 26 3.3.8 Examination and Identification of Eggs 27 3.3.8.1 Tentative Key to the Eggs of British Genera of Carrion-breeding Calliphoridae and all Species of British Calliphora 28 4. MORPHOLOGY OF THE LARVAE AND PUPARIA 30 4.1 Introduction: Literature Review 30 4.2 Morphology of the Larvae 33 4.2.1 Colour 33 4.2.2 Size and Shape 34 4.2.3 Segmentation 35 4.2.4 Processes, Papillae and Surface Structures 37 4.2.4.1 The Cephalic Segment 37 4.2.4.2 The Twelfth Segment 39 4.2.4.3 The Thoracic and Remaining Abdominal Segments 42 - (vi) - Page 4.2.5 Spinulation 44 4.2.6 The Tracheal System 48 ,3 Morphology of the Puparia 56 4.3.1 Pupariation 56 4.3.2 Colour 57 4.3.3 Surface Sculpture 58 4.3.4 Size and Shape 58 4.3.5 Pupal Respiratory Apparatus 59 4.3.6 Adult Eclosion 61 THE CEPHALOPHARYNGEAL SKELETON 63 5.1 Introduction 63 5.2 Homology of the Sclerites 64 5.3 Musculature 66 5.4 The First Instar Skeleton 67 5.5 The Second Instar Skeleton 69 5.6 The Third Instar Skeleton 71 5.7 Ecdysis 85 5.8 The Variation of Sclerotisation 86 5.9 Biometrics 89 THE CEPHALOPHARYNGEAL SKELETON IN NON-CALLIPHORID CALYPTERATA 92 6.1 Introduction 92 6.2 Sarcophagidae 93 6.3 Tachinidae 97 6.4 Oestridae 100 6.5 Cuterebridae 102 6.6 Gasterophilidae 103 6.7 Hypodermatidae 107 SYSTEMATIC ACCOUNT: 1. CALLIPHORINAE 108 7.1 Introduction 108 7.2 Genus Calliphora Robineau-Desvoidy 109 7.3 Genus Cynomya Robineau-Desvoidy 137 7.4 Genus Cynomyopsis Townsend 140 7.5 Genus Triceratopyga Rohdendorf 141 7.6 Genus Aldrichina Townsend 143 7.7 Genus Eucalliphora Townsend 144 7.8 Genus Lucilia Robineau-Desvoidy 145 7.9 Genus Hemipyrellia Townsend 157 7.10 Biology of Calliphorinae 160 7.11 Mew Host Records for Calliphorinae 161 SYSTEMATIC ACCOUNT: 2, CHRYSOMYIINAE 162 8.1 Introduction 162 8.2 Genus Chrysomya Robineau-Desvoidy 162 8.3 Genus Cochliomyia Townsend 179 8.4 Biology of Chrysomyiinae 183 8.5 New Host Record for Chrysomyiinae 183 - (vii) - Page 9. SYSTEMATIC ACCOUNT: 3. PHORMIINAE AND OTHER GROUPS 184 9.1 Introduction 184 9.2 Phormiinae 184 9.2.1 Genus Phormia Robineau-Desvoidy 184 9.2.2 Genus Boreellus Aldrich and Shannon 188 9.2.3 Genus Protocalliphora Hough 189 9.3 Other Genera 194 9.3.1 Genus Amenia Robineau-Desvoidy 194 9.3.2 Genus Stomorhina Rondani 198 9.3.3 Genus Tricyclea Wulp 199 9.3.4 Genus Auchmeromyia Brauer and Bergenstamm 200 9.3.5 Genus Elephantoloemus Austen 202 9.3.6 Genus Booponus Aldrich 204 9.3.7 Genus Cordylobia Grunberg 205 9.3.8 Genus Pollenia Robineau-Desvoidy 208 9.3.9 Biology of Phormiinae and Other Groups 209 9.3.10 New Host Records 211 10. IDENTIFICATION OF SPECIMENS (] 212 10.1 Introduction 212 .10.2 Examination of Specimens 213 10.3 Key to British Carrion-breeding Calliphorid Genera 216 10.4 Key to World Genera of Third Instar Larvae Causing Myiasis in Mammals, including Man 220 10.5 Key to the British Species of Calliphora 224 10.6 Key to African Calliphora 230 10.7 Key to Carrion-breeding Phormiinae 232 10.8 Genus Chrysomya 234 10.8.1 Ch. albiceps and Ch. rufifacies 234 10.8.2 Ch. marginalis and Ch. megacephala 225 10.8.3 Ch. putoria and Ch. chloropyga 236 10.9 Genus Lucilia 236 10.10 Key to Third Instars of Cochliomyia 238 10.11 Key to Third Instars of Cordylobia 239 10.12 Nomenclature 240 11. EVOLUTION AND PHYLOGENY 241 11.1 Introduction 241 11.2 Materials and Methods 242 11.3 The Taxonomic Value of Characters of Immature Stages 248 11.4 Lines of Evidence 252 11.4.1 Evidence from Palaeontology 252 11.4.2 Evidence from Parasitology 257 11.4.3 Evidence from Comparative Morphology 263 11.4.4 Evidence from Ontogeny 264 11.4.5 Evidence from Genetics 264 11.4.6 Evidence from Physiology 265 - (viii) - Page 11 .4.7 Evidence from Zoogeography 266 11 .4.8 Evidence from Climatology 268 11 .4,9 Evidence from Previous Taxonomic Work 269 11 5 Character States 269 11 .5,1 Characters of the Third Instar Larva 270 11 5,1.1 Spine Structure and Pigmentation 270 11 .5.1.2 Spine Banding 270 11 5.1.3 Number of Lobes of Anterior Spiracle 271 11 5.1.4 Posterior Spiracle Size and Distance Factor 271 11 5.1.5 Ecdysial Scar Pigmentation 272 11 5.1.6 Posterior Spiracle Anastomoses 272 11 5.1.7 The Sun-Ray Structure 273 11 5.1.8 Body Processes 273 11 5.1.9 Posterior Papillae Size and Position 274 11 5.1.10 Shape Factor 274 11 5.1.11 Mouth-Hook Length 274 11 5.1,12 The Oral Sclerite 275 11 5.1.13 The Liguloid Arch 270 11. 5.1.14 The Dental Sclerite 276 11. 5.1.15 The Hypostomal Plates 277 11. 5.1.16 The Hypostomal Sclerite 277 11. 5.1.17 The Parastomal Bars 278 11. 5.1.18 The Dorsal Cornu 278 11. 5.1.19 The Ventral Cornu 279 11. 5.1.20 The Lateral Plate 279 11. 5.1.21 The Pharyngeal Ridges 280 11. 5.2 Characters of the Second Instar Larva 280 11. 5.2.1 The Mouth-Hook 280 11. 5,2.2 The Hypostomal Process 281 11. 5.3 Characters of the First Instar Larva 281 11.
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