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Phylogenetic Analysis of the Scarab Beetle Tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) Based on Adult Morphological Characters

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Phylogenetic Analysis of the Scarab Beetle Tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) Based on Adult Morphological Characters PHYLOGENETIC ANALYSIS OF THE SCARAB BEETLE TRIBE CYCLOCEPHALINI (COLEOPTERA: SCARABAEIDAE: DYNASTINAE) BASED ON ADULT MORPHOLOGICAL CHARACTERS A Thesis by Daniel Robinson Clark Bachelor of Science, Brigham Young University-Idaho, 2006 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 July 2011 © 2011 by Daniel Robinson Clark All Rights Reserved PHYLOGENETIC ANALYSIS OF THE SCARAB BEETLE TRIBE CYCLOCEPHALINI (COLEOPTERA: SCARABAEIDAE: DYNASTINAE) BASED ON ADULT MORPHOLOGICAL CHARACTERS The following faculty 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 ________________________________________ Karen Brown, Committee Member ________________________________________ William Parcell, Committee Member iii DEDICATION To my wife, my son, my parents, and all my loving family iv ACKNOWLEDGMENTS I would first like to thank my wife, Marnee Clark, whose love and support made this possible, as well as my father (Brad Clark) and my uncle (Shawn Clark) for sharing their love of insects with me. I would also like to thank my adviser, Mary Liz Jameson, for all of her thoughtful and patient help and guidance. I am grateful to the other members of my thesis committee, Karen Brown and William Parcell, for their suggestions throughout my thesis project. Thanks also go to my colleague, Matt Moore, as well as other students and faculty in the Department of Biological Sciences at Wichita State University, especially Jennifer Ellie, Maria Martino, and Debby Werth for all their assistance. Thanks to Brett Ratcliffe and Matt Paulsen (University of Nebraska Lincoln) for their assistance and input. This research was funded by Wichita State University; The Biofinity Project NSF DBI-0743783 to Scott, Henniger, Jameson, Moriyama, Soh, and Ocampo; and NSF-DEB 0716899 to Ratcliffe and Cave. Finally, thanks go to the University of Nebraska, University of Kansas, Brigham Young University, INBio, Carnegie Museum of Natural History, La Esquela Agricola Panamerica, and the Mary Liz Jameson personal collection for loaning specimens used in this research. v ABSTRACT The scarab beetle tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) includes 15 genera and approximately 500 species that are distributed primarily in the New World. Species in the group are important pollinators of aroids, palms, and lilies, some are agricultural pests, and a few are invasive in areas where they have been introduced. Evolutionary relationships of the group have not been examined, and monophyly of the tribe and relationships of genera have not been addressed. As a result, the classification remains unstable, and there is no foundation with which to interpret evolutionary patterns. To provide an evolutionary framework, a phylogenetic analysis using 77 adult morphological characters was performed. Representatives from all recognized cyclocephaline genera were examined as well as taxa that have historically been included in the tribe. Outgroups included exemplars from the dynastine tribes Oryctini, Oryctoderini, and Pentodontini and the ruteline tribes Anomalini and Rutelini. Based on the analysis, Cyclocephalini, as currently circumscribed, does not constitute a monophyletic lineage. Thus, the current classification does not reflect evolution, and revision is required. Based on strong character support, the genera Eriocelis and Stenocrates are not a part of the new cyclocephaline clade while genera Parapucaya and Neohyphus are members of the new Cyclocephalini. vi TABLE OF CONTENTS Page INTRODUCTION….……………………………………………………………………………..1 Classification and Phylogeny……………………………………………………………...3 Ecological Significance…………………………………………………………………....6 MATERIALS AND METHODS………………………………………………………………...10 RESULTS………………………………………………………………………………………..15 Character Analysis………….……………………………………………………………15 Phylogenetic Analysis……………………………………………………………………30 Discussion of Major Clades and Genera………………………………………................31 DISCUSSION……………………………………………………………………………………42 REFERENCES CITED…………………………………………………………………………..47 APPENDICES…………………………………………………………………………………...51 A. Preliminary, unpublished cladistical analysis of the tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) conducted by Brett C. Ratcliffe (1980)……...................................................................................................................52 B. Part of a preliminary, unpublished study of the superfamily Scarabaeoidea using the 28S rDNA D2-D3 and 18S gene regions conducted by “Team Scarab” at the University of Nebraska State Museum (2006)……………………………………………….……………………...54 C. Figures depicting the diversity of Cyclocephalini sensu Smith (2006), the newly defined Cyclocephalini, and the genus Cyclocephala…………………….55 vii LIST OF TABLES Table Page 1. Cyclocephalini genera, types species of the genus, number of species, and distribution…………….…………………….……..………………………...2 2. Cyclocephalini genera with described larvae………………………………………...…...7 3. Taxa used in the phylogenetic analysis indicating the subfamily and tribe. An asterisk (*) denotes the type species for the genus among Cyclocephalini…….……11 4. Data maxrix including 61 taxa and 77 morphological characters………………………..16 A.1 Cyclocephalini character states and consistency ratio…………………………………...52 A.2 Cyclocephalini data matrix using 17 characters…………………………………………52 viii LIST OF FIGURES Figure Page 1. Modified point card with (upper, left to right) maxilla, mentum, mandibles; (center, to left of pin) spiculum gastrale; (lower) male genitalia.………………...……...13 2. Showing body noticeably tapered at both ends (a), ends more rounded (b), and entire body somewhat circular (c)..................................…………………………….15 3. Clypeus exemplars described as apex/lateral middle/lateral base: emarginate/parallel/parallel (a), truncate/concave/convergent (b), broadly parabolic/convex/divergent (c), parabolic/convergent/convergent (d), tri-emarginate/convex/divergent (e), and acute/convergent/divergent (f)……………….18 4. Enlarged male foreclaw gently curved with conical unguitractor plate (a), claw bent at 90° angle with cylindrical unguitractor plate (b), and claw drastically bent with dorsoventrally flattened unguitractor plate (c)…………………….24 5. Protibial spur (dorsal view) strongly decurved at apex (a), protiba strongly curved with protibial spur fused and tooth near base (b) and protibial spur straight (c)…………………………………………………………………………...…...25 6. Spiculum gastrale with apodeme absent (a), only a minor nub (b), short (c), and long (d)…………………………………………………………………….29 7. One of three trees generated after successive reweight based on rescaled consistency index (RC). Trees only differ in hypothesized relationships in the clade that includes Cyclocephala aequatoria, C. amazona, Mimeoma acuta, and M. signatoides………………..………………………………………………33 8. Bootstrap 50% majority-rule consensus tree indicating the newly defined Cyclocephalini clade……………………………………………………………………..34 9. Tree from Figure 7 modified with nodes showing bootstrap support above nodes and character trace of major clades below and indicating major clades. Shaded boxes show polyphyly within the genus Cyclocephala…………….…..35 A.1 Cladogram of Cyclocephalini……………………………………………………………53 B.1 Section of Scarabaeoidea phylogeny that includes genera from the tribe Cyclocephalini…………………………………………………………………………...54 C.1 Generic exemplars for Cyclocephalini sensu Smith (2006)…………………………………..55 ix LIST OF FIGURES (continued) Figure Page C.1 Generic exemplars of the newly defined Cyclocephalini.…………………………………….56 C.2 Cyclocephala sub-clade 1 of the newly defined Cyclocephalini in order from basal to apical……………………………………………………………………………………..57 C.2 Cyclocephala sub-clade 2 of the newly defined Cyclocephalini in order from basal to apical……………………………………………………………………………………..58 x INTRODUCTION The family Scarabaeidae (Coleoptera) is one of the ten most species rich beetle families with more than 30,000 species (Ratcliffe and Cave 2006). Some highly impressive scarab beetles, in both size and form, are in the subfamily Dynastinae, commonly called the rhinoceros beetles. This common name is derived from many species having one or more horns. These horns make dynastine beetles very popular among collectors, behavioral biologists who study sexual selection, and evolutionary biologists who study highly variable characters. Characters typically used to circumscribe Dynastinae as a subtribe are as follows: 1) the labrum is hidden beneath the clypeus; 2) the clypeal apex is bidentate, rounded, acuminate, or truncate; 3) the antennae are nine to ten segmented with a club that is usually small and has three segments; 4) the scutellum is not enlarged; 5) the procoxae are transverse; 6) the mesotibial apex has two spurs that are mesad and adjacent (not separated by basal metatarsal segment); and 7) the tarsal claws are all subequal in size except for males of most Cyclocephalini and some Pentodontini where the protarsal claws are enlarged (Ratcliffe 2001). Male genitalia are diagnostic for nearly all species (Ratcliffe 2001). As currently recognized (Table 1) the dynastine tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) contains 15 genera and approximately 500 species (Jameson et
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