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A Dissertation By WEB-INTEGRATED TAXONOMY AND SYSTEMATICS OF THE PARASITIC WASP FAMILY SIGNIPHORIDAE (HYMENOPTERA, CHALCIDOIDEA) A Dissertation by ANAMARIA DAL MOLIN Submitted to the Office of Graduate and Professional Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Chair of Committee, James B. Woolley Committee Members, Mariana Mateos Raul F. Medina Robert A. Wharton John M. Heraty Head of Department, David Ragsdale December 2014 Major Subject: Entomology Copyright 2014 Anamaria Dal Molin ABSTRACT This work focuses on the taxonomy and systematics of parasitic wasps of the family Signiphoridae (Hymenoptera: Chalcidoidea), a relatively small family of chalcidoid wasps, with 79 described valid species in 4 genera: Signiphora Ashmead, Clytina Erdös, Chartocerus Motschulsky and Thysanus Walker. A phylogenetic analysis of the internal relationships in Signiphoridae, a discussion of its supra-specific classification based on DNA sequences of the 18S rDNA, 28S rDNA and COI genes, and taxonomic studies on the genera Clytina, Thysanus and Chartocerus are presented. In the phylogenetic analyses, all genera except Clytina were recovered as monophyletic. The classification into subfamilies was not supported. Out of the four currently recognized species groups in Signiphora, only the Signiphora flavopalliata species group was supported. The taxonomic work was conducted using advanced digital imaging, content management systems, having in sight the online delivery of taxonomic information. The evolution of changes in the taxonomic workflow and dissemination of results are reviewed and discussed in light of current bioinformatics. The species of Thysanus and Clytina are revised and redescribed, including documentation of type material. Four new species of Thysanus and one of Clytina are described. The taxonomy of Chartocerus is reviewed, including redescriptions of 25 out of 33 currently valid species, most based on type or topotypical material. ii DEDICATION To my grandparents. iii ACKNOWLEDGEMENTS I would like to thank everyone who helped me complete this work by providing academic, personal or financial support. Special thanks to my major advisor, Jim Woolley, and my advising committee: John Heraty, Mariana Mateos, Raul Medina and Bob Wharton, for making their time, knowledge and lab space available to me. I also want to extend my thanks to the students at the Woolley Lab and colleagues in Entomology program at Texas A&M, the guys at Heraty Lab, especially the ones that put up with my general pestering—Jason Mottern, James Munro and Roger Burks—, to Matt Yoder, Katja Seltman, Debbie Paul and all who put up with my computer-related pestering (it’s not over BTW), the Brazos Cluster and CIPRES Portal teams, Ann Ellis and Mike Pendleton at the Microscopy Center, the many curators and researchers who sent us material and/or welcomed us at their institution, to John Noyes and the BMNH for the extraordinary amount of specimens contributed to this study, to the many people who have welcomed me during museum and meeting trips, to the Texas A&M Libraries, the student services at Texas A&M, the staff of the Department of Entomology, to all the authors who choose to share their publications and software freely on the Internet, and everyone else who helped in the development of this project. Thanks to the National Science Foundation, the Texas Ecolab program, the Department of Entomology, the EEB program, the College of Agriculture and Life Sciences and sponsors of the Bessoudo Scholarship, the sponsors of the Charles H.’17 iv and Frances Fleming Scholarship, ISS, Entomological Society of America, and all those who provided financial support. I also want to thank the dear friends I met here and those who helped me deal with this brave new world of Texas: Carla and Jim, Aubrey, Katrina, Kira and Brad, Jussana, Kamis, Cacá and the folks at the BRSA, Susanne, Ruth, Lauren, Djalleh and Oscar, Maic, Mikel, Bia, Bruna, Carrie Carolyn, the Ashleys, and many others. Thanks to Rodrigo for his support and not letting me become “one-dimensional”. Thanks to Gil, Maureen, Flavia, Karin, Guilherme, Elaine, Pamela, Bruno, Eduardo and all the other folks from Brazil for the exchange of experiences, ideas, methods, the continued dialogue (which may or may not have involved semi-conscious discussions of species concepts very late hours) and for not forgetting me. I am indebted to my B.S. and M.Sc. major advisor, Gabriel Melo, first for making me write the web site that set this all in motion and later for pushing me into trusting the offer from gringos to attend a workshop that just seemed too good to be true. Finally, I would like to thank my family, especially my parents, for their love, strength, and everything—I have no words to express how grateful I am and how much I look up to you. v TABLE OF CONTENTS Page ABSTRACT ........................................................................................................................... ii DEDICATION ..................................................................................................................... iii ACKNOWLEDGEMENTS ................................................................................................... iv TABLE OF CONTENTS ...................................................................................................... vi CHAPTER I INTRODUCTION ............................................................................................ 1 Summary of Chapter Contents ........................................................................................ 3 Dissertation Objectives .................................................................................................... 4 CHAPTER II BACKGROUND ............................................................................................. 5 Overview ........................................................................................................................... 5 Introduction ..................................................................................................................... 6 “Taxonomic Impediment” ............................................................................................... 6 Issue 1: Publication vs. Dissemination ............................................................................ 8 Publication Format ...................................................................................................... 8 Taxonomy as Multi-Layered Discipline ..................................................................... 11 Issue 2: Hypertext Is Not Just Text ................................................................................ 14 Shifts in Media Format ............................................................................................... 14 Unique Identifiers and Semantics .............................................................................. 17 Stability ...................................................................................................................... 20 Issue 3: Tools and Data Repositories ............................................................................ 22 Conclusions .................................................................................................................... 25 CHAPTER III MOLECULAR PHYLOGENY OF THE PARASITIC WASPS OF THE FAMILY SIGNIPHORIDAE (HYMENOPTERA: CHALCIDOIDEA), WITH NOTES ON CURRENT SUPRASPECIFIC CLASSIFICATION ...................................................... 28 Overview ......................................................................................................................... 28 Introduction ................................................................................................................... 29 Diversity, Distribution and Biology ........................................................................... 30 Classification and Relationships within Signiphoridae ............................................ 32 Relationships of Signiphoridae within Chalcidoidea ............................................... 37 Age of the Group ........................................................................................................ 40 Objectives of This Study ............................................................................................ 40 Materials and Methods ................................................................................................... 41 Taxonomic Sampling .................................................................................................. 41 Collection and Treatment of Molecular Data ............................................................ 41 vi DNA Extraction .......................................................................................................... 42 DNA Sequencing ........................................................................................................ 43 Molecular Markers and Partitioning ......................................................................... 43 Sequence Alignment .................................................................................................. 44 Alternative Alignments .............................................................................................. 45 Phylogenetic Analyses ............................................................................................... 46 Results ...........................................................................................................................
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