Studies on the Taxonomy of Strongyloides (Nematoda; Strongyloididae)
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ResearchOnline@JCU This file is part of the following work: Speare, Richard (1986) Studies on the taxonomy of Strongyloides (Nematoda; Strongyloididae). PhD thesis, James Cook University. Access to this file is available from: https://doi.org/10.25903/5e1bf2068703b Copyright © 1986 Richard Speare. If you believe that this work constitutes a copyright infringement, please email [email protected] STUDIES ON THE TAXONOMY OF STRONGYLOIDES (NEMATODA; STRONGYLOIDIDAE) Thesis submitted by Richard Speare, B.V.Sc., M.B., B.S. University of Queensland, St Lucia, Queensland for the Degree of Doctor of Philosophy in the Graduate School of Tropical Veterinary Science James Cook University of North Queensland March 1986 DECLARATION I declare that this thesis is ray own work and has not been submitted in any form for another degree or diploma at any university or other institute of tertiary education. Information derived from the published or unpublished work of others has been acknowledged in the text and a list of references is given. Richard Speare March, 1986 I, the undersigned, the author of this thesis, understand that the James Cook University of North Queensland will make it available for use within the University Library, and, by microfilm or other photographic means, allow access to users in other libraries. All users consulting this thesis will have to sign the following statement; "In consulting this thesis I agree to make proper written acknowledgement for any assistance which I have obtained from it". Beyond this, I do not wish to place any restriction on access to this thesis. In particular I wish to emphasise that I have no objection to this thesis being copied in whole or in part without my permission, and I deny any institution or organisation the right to impose this restriction on those wishing to do so. £ - 3- ££ (Signature) (Date) Richard speare TABLE OF CONTENTS. ABSTRACT.............. i KEY TO ABBREVIATIONS iii GENERAL INTRODUCTION 1 CHAPTER 1 CLASSIFICATION OF STRONGYLOIDES 1.1 Higher Taxon Level.................... .. 5 1.2 Generic Level...................................... 7 1.3 Name of the Genus............................. 8 1.4 Summary............. ..................................... 14 CHAPTER 2 DEFINITION OF THE GENUS 2.1 Introduction................................................................ 15 2.2 Definition..................................................................... 22 2.2.1 Parasitic Female.................................................. 22 2.2.1.1 Description................................................................. 22 2.2.1.2 Additions to Little's Definition............. .. 25 2.2.1.3 Points of Disagreement.................... ............. 30 2.2.2 Free-Living Female................................. ............. 32 2.2.2.1 Description........... ................................ ............. 32 2.2.3 Free-Living Male.................... 36 2.2.3.1 Description......................................................... 36 2.2.3.2 Additions to Little * s Definition........... 38 2.2.3.3 Disagreements with Little's Definition 40 2.2.4 Eggs 40 2.2.4.1 Description............... ..................... 40 2.2.4.2 Comment......................................... 42 2.2.5 First stage Larva............................ 43 2.2.6 Second Stage Larva........................... 43 2.2.6.1 Second Stage Rhabditoid Larva 46 2.2.6.2 Second Stage Filariform Larva 46 2.2.6.3 Comment............................................... 46 2.2.7 Third Stage Filariform Larva... 46 2.2.8 Third Stage Rhabditoid Larva... 51 2.2.9 Fourth Stage Rhabditoid Larva.. 51 2.2.9.1 Description............................... 51 2.2.9.2 Comment............................................... 51 2.2.10 Fourth Stage Parasitic Female.. 54 2.2.10.1 Description............................... 54 2.2.10.2 Comment...................................... 54 2.3 Generic Boundaries........................... 56 2.3.1 Buccal Capsule............................... 56 2.3.2 Buccal Teeth............................... .... 57 2.3.3 Seminal Receptacle...................... 57 2.3.4 Caudal Papillae.............. 60 2.3.5 Clarity of Generic Boundaries 60 2.4 Summary. .................... ............................ 61 CHAPTER 3 STATUS OF NAMES USED FOR SPECIES OF STRONGYLO IDES 3.1 List of Published Names.................................................. .. 62 3.2 Unavailable Names............................... .......................................... 63 3.2.1 Comment on Unavailable names.................................. .. 70 3.3 Invalid Names..................................................................................... 77 3.3.1 Comment on Invalid names...................................................... 79 3.4 Descriptions not Sighted. ........................................................ 89 3.5 Summary.................................................................................................... 89 3.6 Further Aims of This Thesis...................... ............................ 91 CHAPTER 4 TECHNIQUES AND EXPERIMENTS 4.1 Introduction..................................................... ................ 92 4.2 Techniques.................................................................................. 92 4.2.1 Collection of Strongyloides.............................. 92 4.2.1.1 Parasitic Female.................................................. 92 4.2.1.2 Other Parasitic Stages............................... 93 4.2.1.3 Free-living Stages............................. 94 4.2.2 Fixation........................... ....................................................... 96 4.2.3 Examination............................................................................ 96 4.2.3.1 General............................................................... ................ 96 4.2.3.2 Transfering into Glycerol..................................... 97 4.2.3.3 En face Preparations............ .. ................................ 98 4.2.4 Depigmentation technique.................................. 99 4.2.5 Measurement................. ......................................................... 100 4.2.6 Drawing..................................................................................... 100 4.2.7 Photography...................... ..................................................... 100 4.2.8 Electron Microscopy, ................................ ..................... 101 4.2.9 Histology........................ ...................................................... 101 4.2.10 Culture. .................................................................................. 101 4.2.11 Experimental Infections.................................. ................ 102 4.2.11.1 Percutaneous Penetration...................................... 102 4.2.11.2 Subcutaneous Infection............................. .............. .102 4.2.12 Statistical Analysis............ ............................ .. 102 4.3 Specific Experiments................................................. 103 4.3.1 Post-Mortem Degeration................................................ 103 4.3.2 Host Immunity. .................................................................. 104 4.3.2.1 Experimental Infections......................................... 104 4.3.2.2 Natural Infection....................................................... 104 4.3.3 Fixation Experiments...................................................... 105 4.3.3.1 Type of Fixative................................................... .. 105 4.3.3.2 Temperature of Fixative......................................... 106 4.3.4 Effect of Host........................................................ 107 4.3.5 Change of Morphology with Different Hosts 107 4.3.6 Effect of Temperature on Free-living Stages 108 CHAPTER 5 ARTIFACTS 5.1 Introduction............. ............ .. 109 5.2 Parasitic Female................................... 111 5.2.1 Cuticle.......................................... .. 111 5.2.1.1 Wrinkling.................................... 111 5.2.1.2 Splitting........... .. ........................... 114 5.2.1.3 Anterior Prolapse................ 117 5.2.1.4 Loss of Transverse Striations 117 5.2.1.5 "Cuticular Moulting" ............................... .. 120 5.2.2 Reproductive Tract.......................................... 121 5.2.2.1 Degeneration.................................................. 121 5.2.2.2 Misinterpretations of the Ovary Type 122 5.2.2.3 Failure of Ovaries to Spiral................ 123 5.3 Changes in Dimensions........................................ 124 5.3.1 Too Large.................. ........................................... 124 5.3.1.1 Squashing.......................................................... 124 5.3.1 Too Small................................... .......................... 126 5.3.2.1 Shrinkage. ....................................................... 127 5.4 Shape............................. 134 5.4.1 Conformation..... 134 5.5 Extraneous Material 136 5.5.1 Bacteria........... ................. 136 5.5.2 Immune Precipitates... 136 5.5.3 Pigment. .................... .. 138 5.5.4 Sheaths............. ................. 138 5.6 Free-living Adults........... 143 5.6.1 Death.................................... 143 5.6.2 Bacterial Attack........... 143 5.6.3 Temperature of Culture 144 5.6.5 Age of Culture........ 148 5.6.6 Pigmentation.......... 150 5.7 Summary................................. .. 150 CHAPTER 6 CRITERIA FOR THE DIFFERENTIATION OF SPECIES 6.1 Introduction.................................................................