Resolving the Pygmy Borers (Curculionidae: Scolytinae: Cryphalini)
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RESOLVING THE PYGMY BORERS (CURCULIONIDAE: SCOLYTINAE: CRYPHALINI) By ANDREW J. JOHNSON A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2017 © 2017 Andrew J. Johnson To my parents ACKNOWLEDGMENTS This work would not be possible without a network of collaborators, including Bjarte Jordal, Miloš Knížek, Thomas Atkinson, Anthony Cognato, Sarah Smith, Emily Moriaty Lemmon, Alan Lemmon, Randy Ploetz and Roger Beaver who provided specimens, assistance and advice. Max Barclay (BMNH), Harald Schillhammer (NHMW) and Natalia Vandenberg (USNM), Paul Skelly and Kate Fairbanks (FSCA) provided assistance and access to museum specimens including type material. Makail Mandalstam, Matt Kasson, Ching- Shan Lin, Sangwook Park, Zvi Mendel, Paul Kendra and Francisco Infante provided many specimens critical for this study. Hagus Tarno and Karinda (Universitas Brawyjaya, Malang, Indonesia) assisted with fieldwork. Lab members, particularly Craig Bateman, Caroline Storer, Martin Kostovcik, Li You, Yin-Tse Huang, Sedonia Steininger, Demian Gomez, Paige Carlson, Zachary Nolen, Conrad Gillett and James Skelton, for providing specimens, advice, support and a productive working environment. Committee members, Matt Gitzendanner, Marc Branham, Jason Smith, Fernando Vega and Jiri Hulcr provided valuable advice and comments for this dissertation. Funding was provided by National Science Foundation, the Florida Forest Service, Florida Department of Agriculture – Division of Plant Industry, USDA forest Service, USDA Farm Bill section 100007, and by the School of Forest Resources and Conservation. Additional support was provided by Dr. Duane McKenna, University of Memphis. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES .......................................................................................................... 10 LIST OF FIGURES ........................................................................................................ 11 ABSTRACT ................................................................................................................... 14 CHAPTER 1 INTRODUCTION .................................................................................................... 16 Background ............................................................................................................. 16 Aims and Goals ...................................................................................................... 17 A Review of Cryphalini in America North of Mexico ......................................... 17 Resolving the Pestiferous Hypocryphalus, Global Pests of Mango and Fig ..... 17 Using Phylogenomics to Explore the Evolutionary Innovations of Bark and Ambrosia Beetles .......................................................................................... 18 2 A REVIEW AND IDENTIFICATION GUIDE OF THE PYGMY BORERS (SCOLYTINAE: CRYPHALINI) OF AMERICA NORTH OF MEXICO. .................... 19 Introduction to Chapter 2 ........................................................................................ 19 Introduction to Ecology and Biology of Cryphalini ............................................ 19 Publication and Species Description History .................................................... 20 Materials and Methods for Chapter 2 ...................................................................... 21 Morphology of North American Cryphalini Genera ................................................. 22 Frons ................................................................................................................ 22 Eye ................................................................................................................... 22 Antennae .......................................................................................................... 23 Pronotum .......................................................................................................... 24 Elytra ................................................................................................................ 24 Legs ................................................................................................................. 25 Phylogenetics and Systematics of Cryphalini ......................................................... 26 Key to Cryphalini Genera of North America ............................................................ 27 Cryphalus Erichson, 1836 ....................................................................................... 28 Diagnosis .......................................................................................................... 28 Biology and Ecology ......................................................................................... 29 Notes ................................................................................................................ 29 Key to Species ................................................................................................. 30 Cryphalus pubescens Hopkins, 1915 ............................................................... 30 Cryphalus rubentis Hopkins, 1915 .................................................................... 31 Cryphalus ruficollis Hopkins, 1915 ................................................................... 31 Cryptocarenus Eggers, 1937 .................................................................................. 32 5 Diagnosis .......................................................................................................... 32 Remarks ........................................................................................................... 33 Distribution ....................................................................................................... 33 Biology and Ecology ......................................................................................... 33 Key to Species ................................................................................................. 33 Cryptocarenus diadematus Eggers, 1937 ........................................................ 34 Cryptocarenus heveae (Hagedorn, 1912) ........................................................ 34 Cryptocarenus seriatus Eggers, 1933 .............................................................. 35 Ernoporicus Berger, 1917 ....................................................................................... 35 Remarks ........................................................................................................... 35 Ernoporicus kanawhae (Hopkins, 1915) ........................................................... 35 Hypocryphalus Hopkins, 1915 ................................................................................ 36 Diagnosis .......................................................................................................... 36 Remarks ........................................................................................................... 36 Distribution ....................................................................................................... 36 Biology and Ecology ......................................................................................... 37 Key to Species ................................................................................................. 37 Hypocryphalus mangiferae (Stebbing, 1914) ................................................... 37 Hypocryphalus sp. “1422”................................................................................. 38 Hypothenemus Westwood, 1834 ............................................................................ 38 Diagnosis .......................................................................................................... 38 Distribution ....................................................................................................... 39 Biology and Ecology ......................................................................................... 39 Identification to Species.................................................................................... 40 Key to Species ................................................................................................. 41 Hypothenemus areccae (Hornung, 1842) ........................................................ 45 Hypothenemus birmanus (Eichhoff, 1878) ....................................................... 46 Hypothenemus brunneus (Hopkins, 1915) ....................................................... 48 Hypothenemus californicus Hopkins, 1915....................................................... 49 Hypothenemus columbi Hopkins, 1915 ............................................................ 50 Hypothenemus crudiae (Panzer, 1791) ............................................................ 50 Hypothenemus dissimilis (Zimmermann, 1868) ................................................ 51 Hypothenemus distinctus Wood 1954 .............................................................. 52 Hypothenemus erectus LeConte, 1876 ............................................................ 52 Hypothenemus eruditus Westwood, 1836 ........................................................ 53 Hypothenemus gossypii (Hopkins, 1915) ......................................................... 56 Hypothenemus hirsutus (Wood 1954) .............................................................