TAXONOMY, PHYLOGENY AND HOST RELATIONSHIPS OF THE TRICHODECTIDAE (PHTHIRAPTERA: ISCHITOCERA) by CHRISTOPHER HENRY GOUTTS LYAL, B.Sc. VOLUME 1 October 1983 A thesis submitted for the degree of Doctor of Philosophy of the University of London and for the Diploma of Imperial College, Department of Pure and Applied Biology, Imperial College, London SN7 and Department of Entomology, British Museum (Natural History), London SM7 2 ABSTRACT The external morphology of the Phthiraptera is discussed with particular reference to the Trichodectidae. Structures of the head, thorax and abdomen are examined and homologised, most attention being given to features of potential use in systematic analysis. The homologies of the component parts of the male and female genitalia, hitherto disputed, are established. The characters used by previous workers for systematic placement of the Trichodectidae, Ischnocera, Amblycera, Rhyncophthirina, Anoplura, Phthiraptera and Psocoptera are examined, and a cladistic analysis of these groups performed. The Psocodea and Phthiraptera are found to be holophyletic but the Psocoptera are paraphyletic. The Trichodectidae, Amblycera, Rhyncophthirina and Anoplura are all holophyletic, the Rhynco- phthirina is the sister-group of the Anoplura and the Amblycera the sister-group of all other lice. The Ischnocera is not demonstrably holophyletic, and the exact placement of the Trichodectidae is not determined. A cladistic analysis of the Trichodectidae is performed and the 351 species and subspecies reclassified on the basis of this. Five subfamilies are used to partition the twenty genera employed, ten of the latter being sub-divided into twenty-seven subgenera. One subfamily, three genera and four subgenera are described as new. Three genera are placed in synonymy, eight genera and subgenera are raised from synonymy, and four genera are reduced to subgenera,. The generic placements of 106 species and subspecies are changed. Keys to subfamilies and to genera and subgenera are produced. Relationships of the mammalian hosts of the Trichodectidae are compared to the phylogeny (cladogram) of the family in order to test Fahrenholz* Rule. This Rule, which holds that host and parasite phylogenies match, is found not to be fully applicable to the Trichodectidae, and inferences are drawn for the rest of the Phthiraptera. The ecological and evolutionary aspects of louse-hpst relationships are examined to indicate possible patterns of co-evolution. 3 ACKNOWLEDGEMENTS I must first thank the Director and Trustees of the British Museum (Natural History) for permitting me to carry out this research and submit it for examination under the Public Research Institute Scheme. I would also like to thank Br R. G. Davies of Imperial College and Br L. A. Found and Br G. 3. Vfhite of the British Museum (Natural History) for acting as my supervisors, and for their invaluable advice during the course of the stud;/. I am very grateful to all those who have given advice and read parts of the manuscript, particularly Dr T. Clay, Dr P. S. Cranston, Mr U. R. Dolling, Mr D. Hollis, Mr A. M. Hutson, Dr I. F. Hitching, Dr R. P.- Lane, Dr A. Vi. Gentry and Mrs J. M. Palmer of the British Museum (Natural History), Dr H. B. Boudreaux of Louisiana State University and Dr B. Feming of the University of Alberta. During my visit to the United States the following people were very helpful and hospitable: Dr K. C. Emerson, Dr and Mrs K. C. Kim of Pennsylvania State University, Dr and Mrs R. D. Price of the University of Minnesota, Dr 0. Flint of the United States National Museum, Washington, Dr R. Traub and Miss H. Starcke of the University of Maryland School of Medicine, Dr W. A. Drew and Dr D. Peters of Oklahoma State University. I would also like to thank Mrs J. Cranston for typing the thesis and my wife, Mrs J. Lyal, for typing the data matrix. Finally, I must thank my parents and my wife for supporting and encouraging me during the production of this thesis. 4 CONTENTS Page VOLUME 1 ABSTRACT - -- -- -- -- -- -- -- -- -- 2 ACKNOWLEDGEMENTS - -- -- -- -- -- -- -- - 3 CONTENTS - -- -- -- -- -- -- -- -- -- 4 I.IST OP FIGURES - -- -- -- -- -- -- -- - o LIST OP TABLES - -- -- -- -- -- -- -- -- 17 INTRODUCTION - -- -- -- -- -- -- -- -- 18 1. MATERIALS AND METHODS - -- -- -- -- -- -- 21 1.1. Literature - -- -- -- -- -- -- -- - 22 1.2. Specimens - -- -- -- -- -- -- -- -- 26 1.2.1. Introduction - -- -- -- -- -- -- -- 26 1.2.2. Nature of study material - -- -- -- -- -- 26 1.2.3. Collection and preparation of specimens ------ 27 1.2.3.1. Collecting - -- -- -- -- -- -- -- 27 1.2.3.2. Mounting - -- -- -- -- -- -- -- - 30 1.2.3.3. Observation -------------- - 32 1.3. Systematic analysis and classification ------- 33 1.3.1 • Introduction - -- -- -- -- -- -- -- 33 1.3.2. Homology - -- -- -- -- -- -- -- - 33 1.3.3. Methods of systematic analysis --------- 42 1.3*3*1* Introduction - -- -- -- -- -- -- - 42 1.3.3.2. Methods of phyletic analysis - -- -- -- -- 44 1.3.4. Classification - -- -- -- -- -- -- - 54 1.3*4*1* Relationship of classification and systematic analysis- 54 1.3.4*2. Structure of the classification -------- 55 1.4* Character survey - -- -- -- -- -- -- - 63 1.4.1. The taxonomic character - -- -- -- -- -- 63 1.4.2. Use of characters for grouping taxa - -- -- -- 63 1.4*2.1. Choice of characters and their relative values - - - 63 1.4*2.2. Determination of polarity for phyletic analysis - - - 63 1.4.2.3* Character weighting in phyletic analysis ----- 66 1.4*2.4* Problems of complex characters — - - ----- 68 1.4.2.5* Constitutive and diagnostic characters ------ 65 1.4*3* Lata recording in this study - -- -- -- -- - 69 1.4.3.1. Procedure - -- -- -- -- -- -- -- 65 1.4.3.2. Coding - -- -- -- -- -- -- -- - 70 5 Page 2. T'ORPHOLOGY AND CHARACTER ANALYSIS CF TRICEODECTIDAE - - - 75 2.1. Systematic position of Trichodectidae - -- -- -- 75 2.1.1. Introduction - -- -- -- -- -- -- -- 75 2.1.2. Relationship of Fhthiraptera to other insects - - - - 78 2.1.2.1. Introduction - -- -- -- -- -- -- - 73 2.1.2.2. Apomorphies proposed for relevant groups of P hall one opt era - -- -- -- -- -- -- -- -- 78 2.1.2.3. Significance of ahove characters -------- 89 2.1.2.4. Relationship of Phthiraptera to Liposcelidae - - - - co 2.1.3. Relationship of Trichodectidae to other Fhthiraptera - - 94 2.1.3.1. Introduction - -- -- -- -- -- -- - 04 2.1.3.2. Apomorphies proposed for families and supra-familial groups within the Phthiraptera - -- -- -- -- -- 05 2.1.3.3. Significance of ahove characters -------- 103 2.1.3.4. Ranking and classification - -- -- -- -- - 108 2.2. General morphology --------------110 2.2.1. Terminology - -- -- -- -- -- -- -- no 2.2.2. The head - -- -- - -- -- -- -- -- m 2.2.2.1. Introduction - -- -- -- -- -- -- - ni 2.2.2.2. Structure of the head capsule - -- -- -- --112 2.2.2.3. The tentorium - -- -- -- -- -- -- - 120 2.2.2.4. The antennae - -- -- -- -- -- -- - 121 2.2.2.5. The ocelli and compound eyes - -- -- -- -- 127 2.2.2.6. The mouthparts - -- -- -- -- -- -- -129 2.2.3. The thorax - -- -- -- -- -- -- -- -137 2.2.3.1. Introduction - -- -- -- -- -- -- -137 2.2.3.2. The neck region - -- -- -- -- -- -- 137 2.2.3.3. Segments of the thorax - -- -- -- -- --137 2.2.3.4. Description of the trichodectid thorax ------ 140 2.2.3.5. Goxal articulations - -- -- -- -- -- - 142 2.2.3.6. The legs - -- -- -- -- -- -- -- -142 2.2.4. The ahdomen - -- -- -- -- -- -- --148 2.2.4.1. Segmentation - -- -- -- -- -- -- - 148 2.2.4.2. Female genital and postgenital segments ----- 154 2.2.4.3. The male ahdomen - -- -- -- -- -- --166 2.2.4.4. The male genitalia - -- -- -- -- -- - 173 2.2.4.5- The tracheal system and spiracles ------- <jc-| 6 £age 2.3. Character survey of Trichodectidae - -- -- -- - qo6 2.3.1. Introduction - -- -- -- -- -- -- -- qc6 2.3.2. List of characters - -- -- -- -- -- -- -jcy 2.4. Character analysis - -- -- -- -- -- -- - 229 2.4.1. Introduction - -- -- -- -- -- -- -- 229 2.4.2. Identification of apomorphic states for phyletic analysis 229 2.4.3. Cladistic analysis - -- -- -- -- -- -- 248 VOLUME 2 3. CLASSIFICATION AND TAXONOMY OF TRICHODECTIDAE ----- 264 3.1. Taxonomic history of Trichodectidae - -- -- -- - 265 3.2. Proposed classification - -- -- -- -- -- - 282 3.3. Descriptions of genera and subgenera -------- 291 3.3.1. Introduction - -- -- -- -- -- -- -- 291 3.3.2. Bovicolinae - -- -- -- -- -- -- -- 292 3.3.2.1. Bovicola Ewing - -- -- -- -- -- -- - 292 3.3.2.2. Genus n. '2 - -- -- -- -- -- -- -- 303 3.3.2.3. Nerneckiella Eichler - -- -- -- -- -- - 305 3.3.2.4. Genus n. 3 - -- - - -- -- -- -- -- 307 3.3.2.5. Damalinia Fjttberg ------------ - 309 3.3.3. Eutrichophilinae - -- -- -- -- -- -- - 323 3.3.3.1. Eutrichophilus Mjbberg - -- -- -- -- -- 323 3.3.4. Dasyonyginae - -- -- -- -- -- -- -- 327 3.3.4.1. Cebidicola Bedford - -- -- -- -- -- - 327 3.3.4.2. Procavicola Bedford - -- -- -- -- -- - 331 3.3.4.3. Frocaviphilus Bedford - -- -- -- -- -- 338 3.3.4.4. Dasyonyx Bedford - -- -- -- -- -- -- 346 3.3.4.5. Eurytrichodectes Stobbe - -- -- -- -- -- 352 3.3.5. Trichodectinae - -- -- -- -- -- -- - 354 3.3.5.1. Frotelicola Bedford - -- -- -- -- -- - 354 3.3.5.2. Lutridia Kdler - - -- -- -- -- -- -- 358 3.3.5.3. Genus n. 4-------------- - 362 3.3.5.4. Nerneckodect es Conci - -- -- -- -- -- - 364 3.3.5.5. Trichodectes Nitzsch - -- -- -- -- -- - 366 3.3.5.6. Felicola Bring - -- -- -- -- -- -- - 383 3.3.5.7. Lorisicola Bedford - -- -- -- -- -- - 357 3.3.6. Subfamily n.--------------- - 409 3.3.6.1. Neotrichodect es Ewing - -- -- -- -- -- 409 3.3-6.2. Geomydcecus Ewing ------------