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PDF Compressor Pro Diving Beetles of the World PDF Compressor Pro Diving Beetles Systematics and PDF Compressor Pro of the World Biology of the Dytiscidae Kelly B. Miller Department of Biology and Museum of Southwestern Biology University of New Mexico Albuquerque, New Mexico, USA and Johannes Bergsten Department of Zoology Swedish Museum of Natural History Stockholm, Sweden Johns Hopkins University Press Baltimore PDF Compressor Pro © 2016 Johns Hopkins University Press All rights reserved, Published 2016 Printed in the United State of America on acid-free paper 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218-4363 www.press.jhu.edu ISBN 13: 978-1-4214-2054-7 (hardcover: alk. paper) ISBN 10: 1- 4214-2054-6 (hardcover: alk. paper) ISBN 13: 978-1-4214-2055-4 (electronic) ISBN 10: 1-4214-2055-4 (electronic) Library of Congress Control Number: 2015958608 A catalog record for this book is available from the British Library. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410-516-6936 or [email protected]. Johns Hopkins University press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 percent post-consumer waste, whenever possible. PDF Compressor Pro To Mom and Dad, who made me curious, and to all my students, who keep the adven- ture alive. — K. B. Miller To Anders N. Nilsson for being an endless source of inspiration and for contagiously sharing his passion for water beetles, entomology, and systematics. — J. Bergsten PDF Compressor Pro This page intentionally left blank PDF Compressor Pro Contents Preface ix 19. Tribe Eretini 123 20. Tribe Aciliini 125 1. Introduction 1 21. Subfamily Coptotominae 133 2. Taxonomy and Morphology 21 22. Subfamily Hydrodytinae 135 3. Keys to Major Groups of Dytiscidae 23. Subfamily Hydroporinae 138 Subfamilies, Adults 39 24. Tribe Laccornini 145 Subfamilies, Larvae 43 25. Tribe Laccornellini 147 Subterranean & Terrestrial Genera 45 26. Tribe Hydroporini 150 4. Subfamily Matinae 50 27. Subtribe Hydroporina 154 5. Subfamily Lancetinae 53 28. Subtribe Deronectina 162 6. Subfamily Agabinae 55 29. Subtribe Siettitiina 172 7. Tribe Hydrotrupini 57 30. Subtribe Sternopriscina 180 8. Tribe Agabini 62 31. Tribe Vatellini 190 9. Subfamily Colymbetinae 69 32. Tribe Methlini 194 10. Subfamily Copelatinae 78 33. Tribe Hydrovatini 196 11. Subfamily Laccophilinae 87 34. Tribe Pachydrini 199 12. Tribe Agabetini 89 35. Tribe Hygrotini 201 13. Tribe Laccophilini 91 36. Tribe Hyphydrini 207 14. Subfamily Cybistrinae 103 37. Tribe Bidessini 219 15. Subfamily Dytiscinae 111 16. Tribe Dytiscini 114 Literature Cited 259 17. Tribe Hydaticini 118 Index 307 18. Tribe Aubehydrini 121 PDF Compressor Pro This page intentionally left blank PDF Compressor Pro ix Preface Discovering and organizing the diversity of life on We fi rst and foremost thank our graduate academic Earth (the “natural system”) are some of the great- mentors, Boris Kondratieff, Anders N. Nilsson, and est scientifi c undertakings of mankind. Students of Quentin Wheeler, who inspired us and provided the diving beetles have benefi ted from some of the best liberty and resources to explore our taxon. Also, to historical systematists who built a strong foundation the following colleagues, we humbly offer our sin- for our current generation. Study of diving beetle cere thanks: M. Samuel Adams, Yves Alarie, Robert systematics has progressed dramatically in the past Angus, Stephen Baca, Michael Balke, Luca Bar- several years with great numbers of new species de- tolozzi, David Bilton, Olof Biström, Rafael Braga, scribed, many from habitats only newly discovered Gracen Brilmyer, Rasa Bukontaite, Stephen Cam- to have diving beetles. Fortunately, modern phylo- eron, Gilbert Challet, Emma Cleary, Lauren Clea- genetics using DNA sequence data and sophisticated vall, Jason Cryan, Aurélie Désamoré, William Edel- analytical techniques has made the evolutionary his- man, Georgia Evans, Erin Fenton, Hans Fery, Garth tory of diving beetles more accessible, and a natu- Foster, Marco Gaiani, Joja Geijer, Hemant Ghate, J. ral classifi cation based on their phylogeny is being Randy Gibson, R. Antonio Gomez, Traci Grzymala, constantly improved. Organization and improve- Grey Gustafson, Jiri Hajek, Lars Hendrich, Anna ments to the historical nomenclature have been Hjalmarsson, Alicia Hodson, Emily Hodson, San- largely completed. Finally, much has been advanced dra Holmgren, Juri and Nicholas Homziak, Heidi about the potential utility of diving beetles for stud- Hopkins, Roger Härdling, Toshio Inoda, Benjamin ies of biogeography, evolution, community ecology, Isambert, Manfred Jäch, April Jean, Sarah Jogi, Luis macroecology, chemical ecology, and sexual strat- Joly, Kristina Karlsson Green, Martita Lara, David egy evolution. Given all this, it seemed to us a good Larson, Matthew Leister, Richard Leschen, Nathan time to summarize the known world diversity of the Lord, Shelley MacNeil, Rachael Mallis, Timothy Mc- group in a book form. Cabe, Michael Medrano, Mariano Michat, Elizabeth Montano, Jérôme Morinière, Gino Nearns, Shuhei Systematics knowledge is acquired over Nomura, Fernando Pederzani, Philip Perkins, Pyotr long periods of time, and changes are to be expected Petrov, Felix Picazo, Roberto Poggi, Aaron Prairie, as new taxa and characters are discovered. We ex- pect that this book is not (and should not be) a fi nal Tolotra Ranarilalatiana, Jacquelin Randriamihaja, Caroline Rempe, Ignacio Ribera, Robert Roughley, word on the study of dytiscid biodiversity. Rather, Desi Sanchez, Kayla Sayre, Emily Schmeltzer, Am- we hope it will inspire further research and testing ber Schwettmann, Helena Shaverdo, Andrew Short, of the systematics conclusions presented here. Also, Robert Sites, Paul Skelley, Jessica Smith, Paul Span- this is a large assemblage of information to gather, gler, Warren Steiner, Gavin Svenson, Nicole Telles, and it is likely that there are errors or omissions that Geoff Thomson, Mario Toledo, Patricia Torres, Em- we hope can be forgiven. A number of great diving manuel Toussaint, Julie Urban, Ernie Valdez, Bo beetle systematists has inspired us, including but not Wang, Chris Watts, Hans Weeks, Gunther Wewalka, limited to Frank and John Balfour-Browne, Henry Michael Whiting, G. William Wolfe, Karen Wright, Fall, Felix Guignot, John LeConte, Robert Rough- Donald Yee, and Isabelle Zürcher-Pfander. We also ley, Maurice Régimbart, David Sharp, Paul Span- wish to express considerable thanks to our families, gler, Frank Young, and Alois Zimmermann. To them who, over the years, provided much needed support and other pioneers we are ever grateful. and encouragement in so many ways. This book would also not have been possi- Portions of this project were funded by ble without the generous help of many mentors, col- several sources, including US National Science laborators, and students over the years who trained us, did the lab work, suffered hardships in the fi eld, Foundation grants #DEB-0515924, #DEB-0738179, #DEB-0816904, #DEB-0845984, and #DEB- laughed with us, talked us out of bad ideas, encour- 1353426 to K. B. Miller, Swedish Research Council aged us, challenged us, and otherwise formed the grants #2009-3744, #2013-5170, and fi nancial sup- scaffolding on which we were able to build this proj- port from the Swedish Museum of Natural History ect. Where the book is excellent, they deserve con- to J. Bergsten. siderable credit. The errors, however, belong to us.