A Molecular Phylogenetic Assessment of North America Lichen Moths (Lepidoptera; Erebidae: Arctiinae; Lithosiini) with Life Histo
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A Molecular Phylogenetic Assessment of the North American Lichen Tiger Moths (Lepidoptera; Erebidae; Arctiinae; Lithosiini) with Life History Observations and a Description of a New Species from Central Arizona Item Type text; Electronic Dissertation Authors Palting, John Douglas Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction, presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 24/09/2021 12:38:11 Link to Item http://hdl.handle.net/10150/642057 A MOLECULAR PHYLOGENETIC ASSESSMENT OF NORTH AMERICA LICHEN MOTHS (LEPIDOPTERA; EREBIDAE: ARCTIINAE; LITHOSIINI) WITH LIFE HISTORY OBSERVATIONS AND A DESCRIPTION OF A NEW SPECIES FROM ARIZONA by John D. Palting _________________________ Copyright © John D. Palting 2020 A Dissertation Submitted to the Faculty of the GRADUATE INTERDISCIPLINARY PROGRAM IN ENTOMOLOGY AND INSECT SCIENCE In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 2020 1 2 TABLE OF CONTENTS ABSTRACT................................................................................................................... 4 CHAPTER 1 .................................................................................................................. 5 CHAPTER 2 .................................................................................................................. 9 REFERENCES ............................................................................................................ 11 APPENDIX A .............................................................................................................. 16 APPENDIX B .............................................................................................................. 46 APPENDIX C .............................................................................................................. 47 3 ABSTRACT The tribe Lithosiini (Lepidoptera: Erebidae: Arctiinae) includes some 3600 described species worldwide, with probably another thousand or more awaiting description. As such, they represent one of the larger radiations among the ditrysian Lepidoptera. Yet the taxonomy of the group remains uncertain, their life histories largely unknown and at least 25% of their biodiversity undescribed. We inferred the phylogeny of North American lithosiines using molecular sequence data from three genes that have been show to phylogenetically informative in Lepidoptera: cytochrome oxidase subunit I (COI), ribosomal protein S5 (RPS5) and the large subunit 28S ribosomal DNA (28S). Results provide new insights as to how representatives of the 16 genera and 63 species that comprise the lithosiine fauna of the US are related. We also investigated a unique relationship that has evolved between ants and the caterpillars of lithosiines, the latter of which feed on lichens growing on rocks and tree trunks, habitats where ants are primary predators. Finally, we describe a new species of Hypoprepia, H. lampyroides, from the mountains of central Arizona. This is the fifth member of the genus Hypoprepia, all of which occur in the US. 4 CHAPTER 1. INTRODUCTION 1.1 Lichen Tiger Moths (tribe Lithosiini) Tiger Moths (Erebidae, Arctiinae--with about 11000 species worldwide) are recognizable to most people because of their relatively large size and gaudy coloration. Their larval stages, popularly referred to as woolly bears, are also familiar to most. While nearly two-thirds of the subfamily is comprised of these familiar forms (tribe Arctiini contains about 7000 species worldwide), the much more diminutive and secretive Lichen Tiger Moths, tribe Lithosiini, make up about one third of the group (about 3600 species). The Lichen Tiger Moths are so-named because those few with known life histories specialize in feeding on lichens (Comstock and Henne 1967, McCabe 1981, Wagner 2005, Wagner et al. 2008), symbiotic organisms comprised of algae, fungi and a cyanobacterium that are found throughout most of the world, as are the moths. As with other types of Tiger Moths, the diversity of the Lichen Tiger Moths is greatest in tropical regions, with diversity gradually dropping as one moves north and south from the equator (Holloway 2002, Wagner et al. 2008). The lithosiine fauna of the US consists of at least 63 species in 16 genera (Covell 2005, Schmidt and Opler 2008, Powell and Opler 2009), with nearly half of these occurring in the Southwest. This is likely due, in part, to the high diversity of lichens found here, representing some 1000 species, about 20% of the lichen fauna of North America (CNALH website 2020). The Southwest is also known as a biological blend zone, where typically northern or Rocky Mountain taxa meet more southern neotropical elements (Warshall 1995). Lichen moths may be small, but they are often brilliantly colored, a situation in nature which usually indicates them to be poisonous or harmful in some way. The fascinating chemical ecology of these aposematically-colored moths is just beginning to be studied, and preliminary data demonstrate that the larvae and adults of some sequester toxic polyphenolic compounds derived from the lichen on which they feed (Hesbacher et al. 1995, Scott et al. 2014, Scott-Chialvo et al. 2018). This and a few other tantalizing details known of their life histories suggests the group possesses a wealth of biological secrets, including potential myrmecophily (Ayre 1958, Komatsu and Itino 2014), mimicry rings and acoustic communication (Connor 2009, Nakano et al. 2003, Corcoran et al. 2013). 5 1.1.1 Taxonomy and Systematics The taxonomic status of the Lichen Tiger Moths has been reconsidered many times, from family to subfamily and finally tribe, this latest taxonomic placement being determined using molecular phylogenetics (Zahiri et al. 2011, 2012, 2013). While the monophyly of the group as a whole is well supported, the major lineages within this large group are still being investigated, and it was only twenty years ago that systematists tried to place them into subtribal groups based on morphology (Bendib and Minet 1998, 2000, Jacobson and Weller 2001). More recently, molecular systematics was used to evaluate if these proposed subtribes represent natural groups (Scott et al. 2014, Zenker at al. 2017, Scott-Chialvo et al. 2018). Morphology has been the cornerstone of taxonomy for more than two centuries, organisms bearing characteristics inferred as being both shared and derived being grouped together. The hierarchical ordering of kingdom, phylum, class, order, family, genus and species being based on the work of Linnaeus and others. Under consideration here is the order Lepidoptera, family Erebidae. Between family and genus are the subcategories of subfamily, tribe and subtribe. This is the area of our focus on Lichen Tiger Moth taxonomy. The field of molecular phylogenetics has been evolving rapidly in the last few decades, and involves using DNA sequence data to infer organismal relationships. By using phylogenetic inference methods that incorporate models of nucleotide evolution, evolutionary relationships can be inferred independent of the morphological features of the organisms (Bybee et al. 2010). The order Lepidoptera as a whole contains well over 100,000 species. By far the largest group of lepidopterans are the Noctuoidea, a huge radiation of moths containing nearly 40,000 species. About half of these are placed in the family Erebidae, and more than half of the Erebidae are the subfamily Arctiinae, with about 11,000 species. For many decades the Noctuiodea has been reorganized by taxonomists focusing on different suites of morphological characters. It was not until very recently that the use of molecular phylogenetics has allowed us to see through the dizzying array of morphological character loss and gain amongst this group, and allowed for a more stable taxonomic framework (Zahiri et al. 2011, 2012, 2013). Using this new framework, we are now able to refine our assessment of how genera and species are related, with an ultimate goal of more fully understanding how their magnificent diversity evolved. 6 1.1.2 Myrmecophily and other ant associations Ants evolved rather early on during the Cretaceous period and have successfully radiated as a group across the planet (Moreau et al. 2006). The resource-rich nests of hard-working ants offer an attractive target for other organisms to exploit. Those which have evolved obligate dependency on ants are called myrmecophiles. Insect examples of myrmecophiles include ground beetles of the tribe Paussini (Maurizi et al. 2012; Moore and Robertson 2014) and certain butterflies belonging to the family Lycaenidae. In the latter family, there exists a spectrum of myrmecophily, the larvae of most lycaenids having special glands that the produce secretions that compel the ants to protect them like cattle (Devries 1992, Pierce et al. 2002). Others, such as the European genus Maculinia, chemically compel the ants to collect the larvae and bring them into their nests as if they are their own, at which point the caterpillars feed upon the ant larvae to complete their development into a butterfly (Thomas and Settele 2004, Thomas et al 2010). Published observations suggest the caterpillars of lithosiines have some sort of relationship with ants, with some even calling them myrmecophiles