DEVELOPING SCARAB BEETLE IDENTIFICATION TOOLS FOR HAWAII AND THE PACIFIC A Thesis by Joshua Buck Dunlap Bachelor of Science, University of Western Washington, 2009 Submitted to the Department of Biological Sciences and the faculty of the Graduate School of Wichita State University in partial fulfillment of the requirements for the degree of Master of Science May 2016 © Copyright 2016 by Joshua Buck Dunlap All Rights Reserved DEVELOPING SCARAB BEETLE IDENTIFICATION TOOLS FOR HAWAII AND THE PACFIC The following faculty members have examined the final copy of this thesis for form and content, and recommend that it be accepted in partial fulfillment of the requirement for the degree of Master of Science with a major in Biological Sciences. ___________________________ Mary Liz Jameson, Committee Chair ___________________________ David Hughes, Committee Member ___________________________ Gregory Houseman, Committee Member iii DEDICATION To Hawaiian insects and the entomologists who love them iv “Sometimes you're the windshield. Sometimes your the bug.” ― Mark Knopfler v ACKNOWLEDGMENTS I would like to take the time to thank my graduate advisor, Mary Liz Jameson, whose guidance and forbearance made both the Hawaiian scarab project and my Masters of Science degree possible. I would also like to acknowledge those institutions and stakeholder organizations whose time, hard work and financial support enabled the project: The United States Department of Agriculture Plant Protection and Quarantine offices in Fort Collins, CO and Honolulu, HI, the Hawaiian Department of Agriculture, the University of Hawaii, the University of Nebraska Lincoln, the University of Kansas, the Florida State Arthropod Collection, the National Insect Collection and the University of Guam. I should like to extend a further thanks to Col. Dunlap for her spellchecking and rolly-polly wrangling efforts. Last but never least I must thank Wichita State University's Marcia Norton for her help with travel and budget issues, Dr. James Beck for his help with DNA protocols and the Entomology lab's trio of terror: Schmemmy, Stoneheart, and Jackie. vi ABSTRACT The last decade has seen the arrival of a number of pest scarabs (Coleoptera: Scarabaeidae) to the state of Hawaii and the US territory of Guam. Despite the arrival of such injurious species as the coconut rhinoceros beetle (Oryctes rhinoceros) and the masked chafer (Cyclocephala pasadenae) there remains no comprehensive resource pertaining to scarabs of the American Pacific or their identification. To allow for the ready identification of the 71 scarabs known from Hawaii and the 21 species reported from Guam an all-inclusive online "tool" was created in collaboration with the United States Department of Agriculture and Plant Protection and Quarantine (USDA PPQ) division. The tool was created from the assemblage of several distinct subcomponents: 1) A species checklist compiling all scarabaeoids (Coleoptera: Scarabaeoidea) recorded from Hawaii, with a second checklist for Guam. 2) "Fact sheets" or species pages that assemble relevant information for each species created by examination of preserved specimens and an exhaustive review of published literature. 3) Extensive imaging of relevant species supporting image-based identification for the fact sheets. 4) Creation of an image-based matrix key that facilitates the identification of specimens. The completed tool will fill the gap in taxonomic knowledge regarding Hawaiian and Guamanian scarabs, allowing port screeners, quarantine workers, and other end users to correctly and efficiently identify specimens. The complete, peer-reviewed identification tool was released in February 2016. The web-application and mobile- application identification tools may be accessed on the USDA-ITP (Identification Technology Program) ID Tools web site: http://idtools.org. vii TABLE OF CONTENTS Chapter Page 1. INTRODUCTION AND BACKGROUND 1 Scarabs as Pests 1 Scarabs in Hawaii and Guam 2 Purpose and Intent 4 2. MATERIALS AND METHODS 5 Included Taxa 5 Specimen Acquisition 6 Specimen Imaging 7 Fact Sheets 8 Key Characters 10 Introductory and Supplemental Content 10 Peer Review Website Development and Mobile App Development 12 3. RESULTS 13 USDA-ITP Identification Tools 13 Fact Sheets 13 Adoretus compressus Adoretus sinicus Anomala albopilosa Anomala cuprea Anomala orientalis Anomala sulcatula Anomala viridana Aphodiinae Apterocyclus honoluluensis Apterocyclus kawaii Apterocyclus munroi Apterocyclus palmatus Apterocyclus waterhousei Ateuchus lecontei Canthon humectus Canthon indigaceus Canthon pilularius Canthon viridis Catharsius molossus Copris incertus Copris remotus Cotinis mutabilis Cyclocephala pasadenae viii TABLE OF CONTENTS (continued) Chapter Page Dichotomius carolinus Digitonthophagus gazella Euoniticellus africanus Euoniticellus intermedius Figulus integricollis Holotrichia bipunctata Hybosorus roei Lepidiota carolinensis Ligyrus gibbosus Maladera castanea Maladera japonica Microserica guamensis Neosisyphus spinipes Omorgus procerus Omorgus suberosus Oniticellus cinctus Oniticellus militaris Onitis alexis Onitis phartopus Onitis vanderkelleni Onthophagus armatus Onthophagus binodis Onthophagus comperei Onthophagus cuniculus Onthophagus foliaceus Onthophagus granulatus Onthophagus incensus Onthophagus laminatus Onthophagus nigriventris Onthophagus nuchicornis Onthophagus oklahomensis Onthophagus sagittarius Onthophagus tuberculifrons Oryctes Oryctes rhinoceros Papuana Phanaeus daphnis Phyllophaga ephilida Popillia japonica Popillia lewisi Popillia quadriguttata Protaetia fusca Protaetia orientalis Temnorhynchus retusus ix TABLE OF CONTENTS (continued) Chapter Page Tomarus subtropicus Trox scaber Trypoxylus dichotomus Xylotrupes 4. CONCLUSIONS 271 REFERENCES 272 APPENDIX 294 Checklist to Scarabaeoidea of Hawaii 295 Checklist to Scarabaeoidea of Guam 297 x LIST OF TABLES Table Page 1. Checklist to Scarabaeoidea of Hawaii 295 2. Checklist to Scarabaeoidea of Guam 296 xi LIST OF ABBREVIATIONS C Celsius CRB coconut rhinoceros beetle cm centimeters FSM Fact Sheet Manager ITP Identification Technology Program mm millimeters PPQ Plant Protection and Quarantine sp. species (singular) spp. species (plural) USDA United States Department of Agriculture xii Chapter 1 Introduction Scarabs as Pests The family Scarabaeidae contains a number of well known and invasive pest species (Jackson and Klein, 2006). These pest scarabs include both generalist such as Popillia japonica, the infamous Japanese beetle, and more specialized pests such as the Oryctes rhinoceros, the coconut rhinoceros beetle. The economic importance of scarabs is highlighted by both of these species. The Japanese beetle is regarded as the single most destructive insect pest of lawns, gardens, and golf courses in the eastern U.S. (Klein, 2008). Indeed in 2000, the USDA estimated that $US 450 million were spent annually in the U.S. on control of P. japonica (Potter and Held, 2002). The coconut rhinoceros beetle, too, causes significant damage with palm damage costs exceeding $US 2.5 million between 2007-2009 just on the tiny island of Guam (Moore, 2009). Many of the most destructive pest scarabs are also invasive species that have now been widely spread by human activity. The previously mentioned Japanese beetle was spread to the eastern U.S. from Japan via commercial shipments of iris bulbs (Klein, 2008). Other widespread scarab species include Maladera castanea (the Asian garden beetle) (Ahrens, 2007; Skelley, 2012) and Anomala orientalis (the oriental beetle), both introduced to the U.S. from eastern Asia (Reding and Klein, 2006). In the Indo-pacific Protaetia fusca (the mango flower beetle) (Woodruff, 2006) and closely related Protaetia orientalis (the oriental flower beetle) (LeBlanc et al., 2013) have been spread via human activity, as has the destructive coconut rhinoceros Beetle (Molet, 2013; Bedford, 2015). The range of crops and plants damaged by scarab pests is considerable. The 1 Japanese beetle is known to damage over 300 different species of plants (Vittum et al., 1999), including both agricultural and horticultural species. Other broad generalists include Adoretus sinicus, the Chinese rose beetle, which has been recorded feeding on over 500 species of plants (Hession et al., 1994). The larvae of many scarabs (often referred to as grubs) are major turf grass pests, feeding on the roots of grasses and thereby killing the entire plant. Major turf grass pests include the Phyllophaga (Watschke et al., 1994) and Holotrichia species, sometimes called May beetles, as well as various members of the subfamilies Dynastinae (Jameson et al., 2009), Rutelinae (Suzuki et al., 1994; Krischik, 2011) and Aphodiinae (Shetlar and Niemczyk, 1999). In the tropics scarabs are also major pests of palms (coconut rhinoceros beetle)(Bedford, 2015) and sugarcane (Lepidiota species) (Kuniata and Young, 1992). Despite their clear importance and economic consequence there is no comprehensive tool useful for identifying scarabs on a worldwide scale. This is, in part, because of the sheer diversity of scarabs, with estimates placing the number of scarab species at 31,000 (Jameson and Ratcliffe, 2002). The impracticality of treating such a great number of species within a single key means identification on a global level is impeded by the limits of available keys, with most such materials being restricted to narrow taxonomic groups and/or specific geographic regions. Identification is further hampered by a dearth of taxonomic expertise, even when