Coleoptera: Nitidulidae) Andrew R
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Louisiana State University LSU Digital Commons LSU Doctoral Dissertations Graduate School 2005 Revision of Pocadius Erichson (Coleoptera: Nitidulidae) Andrew R. Cline Louisiana State University and Agricultural and Mechanical College, [email protected] Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_dissertations Part of the Entomology Commons Recommended Citation Cline, Andrew R., "Revision of Pocadius Erichson (Coleoptera: Nitidulidae)" (2005). LSU Doctoral Dissertations. 561. https://digitalcommons.lsu.edu/gradschool_dissertations/561 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Doctoral Dissertations by an authorized graduate school editor of LSU Digital Commons. For more information, please [email protected]. REVISION OF POCADIUS ERICHSON (COLEOPTERA: NITIDULIDAE). A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Entomology by Andrew R. Cline B.S. University of Alabama - Huntsville, 1996 M.S. University of Missouri, 2000 December 2005 © Copyright 2005 Andrew R. Cline All rights reserved ii DEDICATION This dissertation is dedicated to my wife JoAnna. Through the many years we have spent together you have often been my balance and point of reason. Your patience and understanding were always a comfort and inspiration. The work produced herein would not have been possible without you. iii ACKNOWLEDGEMENTS I thank my graduate advisor Dr. Christopher Carlton for his advice throughout the completion of this research. I thank former and current entomology department head’s Drs. Frank Guillot and Timothy Schowalter respectively for their support of my graduate studies. Drs. Dorothy Prowell, Meredith Blackwell, Mark Hafner and Daniel Burba graciously served as graduate committee members. Alexey Tishechkin, a lab associate during my tenure at LSU, provided an immeasurable amount of inspiration and encouragement through our daily discussions and across-office chats, and not only broadened my scientific horizons but also developed my global point of view on cultural, socioeconomic, and historical matters. Dr. Shaun Winterton aided in the analysis of data. I graciously thank the numerous curators and directors that generously loaned specimens used in this study, and without whom this research would have been impossible. In particular, I thank Al Newton and Margaret Thayer at the Field Museum; Robert Anderson, Francois Genier, and Henry and Anne Howden at the Canadian Museum of Nature; Alexander Kirejtshuk at the Zoological Institute in St. Petersburg, Russia; Max Barclay and Malcolm Kerley at the National Museum of Natural History - London; Phillip Perkins at the Museum of Comparative Zoology at Harvard University; and Mike Thomas and Paul Skelley at the Florida State Collection of Arthropods for their kindness during visits made by the author to their respective institutions. Paul Skelley aided in the production of the SEM photographs. Richard Leschen of the New Zealand Landcare; Jose Luis Navarette-Heredia of the University of Guadalajara; Steve Ashe of the Snow Entomological Collection at the University of Kansas; Roberta Brett and Dave Kavanaugh at the California Academy of Sciences; Nancy Adams, and Gary Hevel and David Furth of the United States National Museum - Smithsonian Institute provided iv numerous specimens for this work. Robert Turnbow, Roy Morris, Jim Wappes, Bill Warner, and Alan Gillogly all graciously donated material. Funding for this research was provided by the Louisiana State University Agricultural Center, an Ernst Mayr grant from the Museum of Comparative Zoology at Harvard University, visiting systematist grants from the Field Museum and the Canadian Museum of Nature, travel grants from the Florida Entomological Society and Louisiana State University Graduate School, an A.W. Mellon grant from the Organization for Tropical Studies, a Louisiana State University Sigma Xi Grant-in-Aid of Research, and a National Science Foundation Doctoral Dissertation Improvement Grant DEB 0308764. v TABLE OF CONTENTS ACKNOWLEDGEMENTS………………………………………………… iv ABSTRACT……………...…………………………………………………. vii CHAPTER 1. INTRODUCTION………………………………………….. 1 General Background………………………………………… 1 Taxonomic Background. I. Cucujoidea Latrielle 1802……... 2 Taxonomic Background. II. Nitidulid-Lineage ……………. 7 Taxonomic Background. III. Nitidulidae Latrielle 1802…… 8 Systematics and Classification of Nitidulidae……………… 13 Biological / Ecological Considerations…………………….. 19 Morphological / Molecular Considerations………………… 25 Pocadiini / Pocadius Background………………………….. 27 2. MATERIALS AND METHODS…………………………… 34 Taxonomic Materials / Methods……………………………. 34 Systematic Methods. I. Ingroup / Outgroup Discussion..….. 39 Systematic Methods. II. Character Discussion……………... 44 Systematic Methods. III. Phylogenetic Reconstruction……... 55 3. SYSTEMATIC TREATMENT...………………………….. 58 Generic Redescription……………………………………… 58 Key to Species……………………………………………… 62 Species Accounts............…………………………………… 72 Pocadius Phylogeny….……………………………………... 299 Checklist of Pocadius Erichson.............................................. 329 4. DISCUSSION……………………………………………….. 332 Monophyly of Pocadius and Its Placement in Nitidulinae…. 332 Origin and Distribution of Pocadius..…………….…………. 335 Host Fungal Evolution……………………………………… 340 Phenological Considerations……………………………….. 342 Future/Continuing Research...……………………………… 343 REFERENCES CITED……………………………………………………... 346 APPENDIX: DATA MATRIX…………………………………………..….. 369 VITA………………………………………………………………………… 375 vi ABSTRACT A revision of Pocadius Erichson (Coleoptera: Nitidulidae) was completed. A total of 46 species were (re)described, including 25 new species, a key to species constructed, and phylogenetic analysis performed. Taxonomic changes, including nomenclatural emendations, reinstatement of species, and description of new taxa, within this dissertation do not constitute formal changes as defined by the International Code of Zoological Nomenclature. Illustrations of the genitalia, key characters, and dorsal habitus of species are provided. The phylogeny demonstrates a monophyletic Pocadius. The phylogeny suggests a Palearctic origin of the genus with subsequent speciation into the Old World tropics and New World. Host specialization on gasteromycetes fungi by Pocadius species was shown not to be obligate but rather facultative. Some sympatric species were shown to have temporally disjunct occurrences that may provide reproductive isolation. vii CHAPTER 1. INTRODUCTION “In all things of nature there is something of the marvelous.” Aristotle, On the Parts of Animals, circa 350B.C. GENERAL BACKGROUND Nitidulidae, or sap beetles, are small to minute Clavicornia taxa (e.g. beetles having clubbed antennae) with a variety of body forms, ecologies, life history strategies, and evolutionary peculiarities. Characterization of the family is difficult, and the literature is replete with misplacements of members of other, sometimes unrelated families, into Nitidulidae. Until the 20th century, Nitidulidae contained several other families or parts of families (e.g. Cyclaxyridae, Smicripidae, Kateretidae, Protocucujidae, and Phloeostichidae). These small convex beetles with drab coloration have had historically unclear systematic affinities thereby making this family a quintessential “dumping ground” for small beetles with clubbed antenna. Characterization of what a nitidulid is remains questionable as remarked upon by Murray (1864) prior to his monograph, “this I knew to be no easy task, no journey of the Sabbath day.” Through comparative morphology, Lawrence et al. (1999a, 1999b) and Habeck (2002a, based on Parsons 1943 and Audisio 1993) provided the following that differentially define Nitidulidae: antennae 11 segmented with at least the three terminal antennomeres forming a well-defined club, labrum free and visible, maxilla with a single lobe, procoxae transverse, tarsal formula 5-5- 5, larvae with a complex mandibular prostheca and pretarsal setae present. Of these, I find two most useful when identifying material, i.e. transverse procoxae and compact 3-segmented club. However, only the former character as well as a single-lobed maxilla and complex larval mandibular prostheca are synapomorphic, as evidenced by their absence in other beetle lineages. 1 To understand and appreciate the complexity of the taxonomic and subsequent systematic condition of Nitidulidae fully, it is best first to assess the superfamily in which it is contained and continue down the classification hierarchy (Table 1). The following sections deal with our current understanding of nitidulids and their closest allies. Taxonomic efforts focusing on nitidulids and their relatives, how nitidulids are subdivided, the lifestyles and evolutionary histories some nitidulids exhibit, and studies underlying comparative morphology and molecular biology in the family also are treated. Finally, an introduction to the focal taxa of this dissertation, i.e. the genus Pocadius Erichson and tribe Pocadiini Seidlitz, are given. TABLE 1. Hierarchy demonstrating the placement of Pocadius, Pocadiini, and other taxa. Taxa in bold indicate some of the taxa considered in this study. KINGDOM Animalia PHYLUM Arthropoda CLASS Insecta ORDER Coleoptera SUBORDER Archostemata, Myxophaga, Adephaga, Polyphaga