Trichodagmia Enderlein (Diptera: Simuliidae: Simulium) in the New World
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SYSTEMATICS OF THE BLACKFLY SUBGENUS TRICHODAGMIA ENDERLEIN (DIPTERA: SIMULIIDAE: SIMULIUM) IN THE NEW WORLD Luis M. Hernández Triana Thesis committee Thesis supervisors Prof. dr. M.S.M. Sosef Professor of Biosystematics Wageningen University Prof. dr. R.J. Post Research Entomologist The Natural History Museum, London, United Kingdom Other members Prof. dr. M. Dicke, Wageningen University Prof. dr. M. Schilthuizen, University of Groningen & NCB Naturalis Dr. H. de Jong, University of Amsterdam Prof. dr. R.A. Cheke, University of Greenwich, UK ii SYSTEMATICS OF THE BLACKFLY SUBGENUS TRICHODAGMIA ENDERLEIN (DIPTERA: SIMULIIDAE: SIMULIUM) IN THE NEW WORLD Luis M. Hernández Triana Thesis submitted in fulfilment of the requirements for the degree of doctor at Wageningen University by the authority of the Rector Magnificus Prof. dr. M. J. Kropff, in the presence of the Thesis Committee appointed by the Academic Board to be defended in public on Tuesday 29 March 2011 at 4 p.m. in the Aula. iii Luis M. Hernández Triana Systematics of the blackfly subgenus Trichodagmia ENDERLEIN (Diptera: Simuliidae: Simulium) in the New World, 546 pages Thesis, Wageningen University, Wageningen, NL (2011) With references, with summaries in Dutch and English ISBN 978-90-8585-865-2 iv The research described in this thesis was financially supported by The Natural History Museum, London. Financial support from Wageningen University is gratefully acknowledged. v To my wife Nathalie David-Hernández, to my children Mina, Lani, Kiran, Ariadna and Ian to my mother Francisca Triana, my late father Alberto Martin, to my sister Arais Martin, to all my family in Cuba vi CONTENTS 1. GENERAL INTRODUCTION ..................................................................................................................... 1 1.1. Background.............................................................................................................................................. 1 1.2. History of the Higher Classification of Simuliidae............................................................................ 2 1.3. A Phylogenetic Approach to Blackfly Systematics............................................................................ 5 1.3.1. Morphological Characters ..................................................................................................................... 5 1.3.2. Molecular (DNA) Characters................................................................................................................ 7 1.3.2.1. DNA Barcoding...................................................................................................................................... 8 1.3.3. Impact of other Biochemical Methods (Enzyme Electrophoresis and Cuticular Hydrocarbons) and Cytogenetics on Simuliidae Systematics....................................................................................... 9 1.4. Development of a Classification System for the Simuliidae in the Neotropical Region.............. 9 1.4.1 Classical approaches to Taxonomy, including Supraspecific Division in Neotropical Simuliidae. .................................................................................................................................................................. 9 1.4.2. Phylogenetic approaches to Neotropical Simuliidae taxonomy .................................................... 12 1.5. Current Classification of the Family Simuliidae in the Neotropical Region................................ 13 1.6 Perspectives and Future Developments............................................................................................ 13 1.7. Systematics of the Subgenus Trichodagmia ENDERLEIN in the New World.................................. 14 1.8. APPENDIX 1. TABLES. ........................................................................................................................... 15 2. MORPHOTAXONOMY OF THE SUBGENUS TRICHODAGMIA ENDERLEIN ........................................ 19 2.1. Summary ................................................................................................................................................ 19 2.2. Introduction .......................................................................................................................................... 19 2.3. Material and Methods .......................................................................................................................... 20 2.3.1. Collecting, Rearing and Preparation .................................................................................................. 20 2.3.2. Digital Imaging and Measurements ................................................................................................... 21 2.3.3. Preservation........................................................................................................................................... 22 2.3.4. Structures Used by other Authors in Taxonomic Studies on Neotropical Simuliidae............... 22 2.3.5. Methods Used in Taxonomic Research on Simuliidae in the Current Work .............................. 23 2.3.6. General Figures and Terminology Used in this Work .................................................................... 24 2.3.7. Acronyms Used for Depositaries of Simuliidae............................................................................... 28 2.4. Review of the Subgenus Trichodagmia ENDERLEIN, 1934 ............................................................... 28 2.4.1. Previous work ....................................................................................................................................... 29 2.4.2. Remarks.................................................................................................................................................. 36 2.4.3. Checklist of Trichodagmia species......................................................................................................... 38 2.4.4. Taxonomy of the subgenus Trichodagmia........................................................................................... 42 2.4.5. Identification keys to species groups of the subgenus Trichodagmia .............................................. 44 2.4.6. Identification keys to species of the New World CANADENSE species group....................... 47 2.4.7. Identification keys to species of the New World ORBITALE species group ............................ 51 2.4.8. Identification keys to species of the New World PICTIPES species group ............................... 55 2.4.9. Identification keys to species of the New World TARSATUM species group........................... 56 2.5. Morphological Description, Taxonomic Discussion, Distribution, and Biology and Medical Importance of species of the subgenus Trichodagmia in the New World..................................... 61 2.5.1. The CANADENSE species group.................................................................................................... 61 2.5.1.1. Simulium ayrozai...................................................................................................................................... 62 2.5.1.2. Simulium burchi........................................................................................................................................ 65 2.5.1.3. Simulium canadense .................................................................................................................................. 68 2.5.1.4. Simulium capricorne.................................................................................................................................. 72 2.5.1.5. Simulium carolinae.................................................................................................................................... 76 2.5.1.6. Simulium contrerense................................................................................................................................. 78 2.5.1.7. Simulium dalmati ..................................................................................................................................... 82 2.5.1.8. Simulium delatorrei................................................................................................................................... 84 2.5.1.9. Simulium estevezi...................................................................................................................................... 88 vii 2.5.1.10. Simulium ethelae....................................................................................................................................... 90 2.5.1.11. Simulium falculatum................................................................................................................................. 94 2.5.1.12. Simulium gorirossiae.................................................................................................................................. 96 2.5.1.13. Simulium johnsoni..................................................................................................................................... 99 2.5.1.14. Simulium juarezi..................................................................................................................................... 102 2.5.1.15. Simulium larvispinosum........................................................................................................................... 105 2.5.1.16. Simulium menchacai...............................................................................................................................