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A Cladistic Analysis and Classification of the Subgenera and Genera of the Large Carpenter Bees, Tribe Xylocopini (Hymenoptera: Apidae)^

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A Cladistic Analysis and Classification of the Subgenera and Genera of the Large Carpenter Bees, Tribe Xylocopini (Hymenoptera: Apidae)^ 1 Scientific Papers Natural History Museum The University of Kansas 07 August 1998 Number 9:1-47 A Cladistic Analysis and Classification of the Subgenera and Genera of the Large Carpenter Bees, Tribe Xylocopini (Hymenoptera: Apidae)^ By Robert L. Minckley^ Division of Entomology. Natural History Museum, and Department of Entomology, The University of Kansas, Lawrence, Kansas 66045-2109, USA CONTENTS ABSTRACT 2 INTRODUCTION 3 Acknowledgments 3 MATERIALS AND METHODS 3 Selection ofTaxa 3 Selection of Characters 5 Characters List and Coding 7 Phylogenetic Analyses 13 RESULTS 14 Relationships among Larger Clades of the Xylocopini 15 Relationships among Members of the Major Clades andTaxonomic Recommendations 21 KEYS TO THE SUBGENERA OF XKLOCOPA 27 New World Females 27 New World Males 28 Old World Females 28 Old World Males 29 DIAGNOSES AND DESCRIPTIONS OF SUBGENERA OF XYLOCOPA 3 Subgenus Xylomelissa Hurd and Moure 31 Subgenus Nodula Maa 31 'Contribution No. 3198 from the Department of Entomology and Snow Entomological Division, Natural History Museum, The University of Kansas. -Present address: Department of Entomology, 301 Funchess Hall, Auburn University, Auburn, Alabama 36849-5413, USA, © Natural History Museum, The University of Kansas ISSN No. 1094-0782 Scientific Papers, Natural History Museum, The University of Kansas Subgenus Maaiana new subgenus 32 QL Subgenus Bihinn Maa 32 (X Subgenus Rhysoxylocopa Hurd and Moure 32 ^ I "-^ ^ Subgenus Pwxylocopa Hedicke, new status 33 A q- -2 Subgenus Nyctomelitta Cockerell 33 Subgenus Ctenoxylocopa Michener 33 34 V\ S'L:> Subgenus Xylocopa Latreille (sensu stricto) Subgenus Xylocopoides Michener 34 34 I I ( Subgenus Calloxylocopn Hurd and Moure Subgenus Copoxyla Maa 35 Subgenus Gnathoxylocopa Hurd and Moure 35 Subgenus testis Lepeletier and Serville, new status 35 Subgenus Notoxylocopa Hurd 36 Subgenus Schonnherria Lepeletier 36 Subgenus Dasyxylocopa Hurd and Moure 37 Subgenus Xylocopsis Hurd and Moure 37 Subgenus Nanoxyhcopm Hurd and Moure 37 Subgenus Cirroxylocopw Hurd and Moure 37 Subgenus Xylocopodn Hurd and Moure 38 Subgenus Stenoxylocopa Hurd and Moure 38 Subgenus Neoxylocopa Michener 38 Subgenus Prosopoxylocopa Hurd and Moure 39 Subgenus Zonohirsuta Maa 39 Subgenus Bomboixylocopa Maa 39 Subgenus Xenoxylocopn Hurd and Moure 40 Subgenus AUoxylocopa Maa 40 Subgenus Mesotricliia Westwood 40 Subgenus Koptortosoma Gribodo 41 LITERATURE CITED 42 APPENDICES 44 ABSTRACT Phylogenetic analyses of the genera and subgenera of the tribe Xylocopini (Large Carpenter Bees) were made using morphological characters coded for 51 species of the ingroup. These species represented all but three of the previously recognized subgenera. Twenty species of bees constituting the outgroup also were examined to evaluate primitive and derived character states within the Xylocopini. Four different analyses were made because of uncertainty in hypotheses of character transformations and among the relationships of the outgroup taxa. Cladistic parsimony analyses showed that previous classifications were based on combinations of characters and not unique synapomoiphies. Three major clades were supported in most analyses presented here; two are from the Old World, and one is from the New World. A biogeographically mixed group of subgenera are not cleariy placed in this analysis; among this group are likely to be found the basal members of the tribe. A new generic and subgeneric classification is proposed for the Xylocopini. I conclude that recognition of Pwxylocopa Maa and testis Lepeletier and Serville as genera creates a paraphyletic genus o'i Xylocopa: these should be recognized as subgenera of an expanded Xy/oco/ja. A new subgenus. Maaiana. is proposed here forX. hentoni and related species formerly recognized as members of the subgenus Nodiila. The following subgeneric synonymies also are proposed: Xylomelissa Hurd & Moure is expanded to include Epixylocopa Hurd & Moure, AUoxylocopa Hurd & Moure, Dinoxylocopa Hurd & Moure and Acroxylocopa Hurd & Moure; Ancylocopa Maa is synonymized with Proxylocopa Maa; Schonnherria Lepeletier is expanded to include loxylocopa Hurd & Moure and Xylocospila Hurd & Moure; Stenoxylocopa Hurd & Moure is expanded to include Xylocopina Hurd & Moure; Neoxylocopa Michener is expanded to include Megaxylocopa Hurd & Moure; Bomboixylocopa Maa is expanded to include Minioxylocopa Hurd & Moure; Mesotrichia Westwood is expanded to include Platynopoda Westwood, Hoplitocopa Lieftinck and Hoploxylocopa Hurd & Moure, and; Koptortosoma Gribodo is expanded Subgenera and Genera of Carpenter Bees. Xylocopini to include Ow.wlocopa Hurd & Moure, CyphoxYlocopa Hurd & Moure. Cycmcoderes Hurd & Mt)ure and Afroxxlocopa Hurd & Moure. A key to identify the subgenera i,s provided. Ke\i words: Insecta; Hvmenoptera; Apidae, Xylocopini; Cladistic analyses; Classification. INTRODUCTION it is worth The large carpenter bees, tribe Xylocopini, generally are catory changes were provided. Nevertheless, Cunha's robust, fast-flving insects, some of which are among the noting that the relationships generated by Da with caution, because similarity largest of all the bees. As their name implies, most mem- study should be used result in estimates of phylogeny (Hillis bers of this group place their nests in dead solid plant methods can poor especially the data used material, usually various kinds of wood, but some use the et al., 1994; Michener, 1970), when and not to infer phy- hollow culms of bamboo (subgenus Biluna; Maeta et al., were intended to diagnose groups 1985), or the pithv stems of Aloe (subgenus Giuitlioxylocopa; logeny. Watmough, 1974). Females oi Pwxylocopa construct their In this study, I use an exemplar approach in the cladis- nests in the ground. Some species of Xylocopini are im- tic analysis (see Phylogenetic Analysis). I also propose portant pollinators of particular plants (e.g., Corbet and some changes in the classification based on my phyloge- Wilmer, 1980; Gottsberger et al., 1988) but none seems to netic analyses. My approach for taxonomic decisions is to restrict its pollen collecting to one or a few plant taxa. Other combine terminal taxa that are monophyletic based on the well-studied features of the biology of the group include phylogenetic analyses and retain the names of the prob- its social behavior (reviewed in Gerling et al., 1988; Maeta lematic taxa as they were recognized before this study. and Sakagami, 1995; Michener, 1990), exocrine gland mor- Future analyses may result in the recognition of fewer phology (Mincklev, 1994; Vinson, 1994) and product chem- supraspecific groups than are accepted here. istrv (Gerling et al, 1988), and male reproductive behavior Acknowledgments (Alcock, 1992; GerUng et al, 1988; Minckley, 1994). This paper is part of a dissertation presented as partial Xylocopini is one of four tribes in the subfamily The requirement for the degree of Ph. D. to the Department of other three tribes Allodapini, Xylocopinae. The — Entomology, The University of Kansas. The curators of the Ceratinini, and Manueliini are mostly smaller than the — institutions listed in the section on Selection of Taxa (be- Xylocopini and build their nests in broken twigs or stems low) are gratefully acknowledged for the loan of the ma- (Daly et al., 1987; Michener, 1974, 1990). terial used in this study. Dr. Charles D. Michener provided The purpose of this study is a cladistic analysis of the generous support and guidance in this project from its in- phylogenetic relationships among the genera and subgen- ception. Others who were influential in the completion of world- era of the large carpenter bees, tribe Xylocopini. A this project include Drs. B. Alexander, J. S. Ashe, William J. wide revision of this group was last undertaken in 1963 Bell, E. O. Wiley, S. G. Reyes, R. A. B. Leschen, A. Roig- bv Hurd and Moure. They recognized 53 supraspecific taxa Alsina, D. Yanega and M. D. Greenfield. Bruce Cutler, who in the tribe: Three genera, testis, Pwxylocopa, and Xi/locopa; allowed me access to his laboratory and gave consider- 48 subgenera in Xylocopa and two subgenera in Proxylocopa. able guidance in the analysis of some of the internal mor- Although their work is the most comprehensive study of phological structures used in the study, deserves special the group to date, both with respect to the number of taxa commendation. I dedicate this paper to the late Byron examined and the number of characters considered, they Alexander; he substantially improved the quality of this did not provide an explicit hypothesis of phylogenetic re- study and piqued my appreciation for the importance of lationships for the members of the tribe. Da Cunha (1992) systematics in evolutionary biology. This work was facili- did a quantitative similarity analysis of the subgenera of tated by grants from the American Museum of Natural Xylocopa using the characters of Hurd and Moure (1963). History (Theodore Roosevelt Fund), Department of Ento- This study was largely concerned with phenetic method- mology at The University of Kansas (Hungerford Fund), ology and was not a taxonomic revision of the Xylocopini, and by the National Science Foundation grant DEB- so no discussion of phylogenetic relationships or classifi- 9302186 (C. D. Michener, principal investigator). MATERIALS AND METHODS Selection of Taxa in the analyses: Euxylocopa, In order to examine ingroup relationships, I used ex- Xylocopa were not included Diaxylocopa and emplar taxa of all supraspecific taxa of the Xylocopini rec- Lieftinckella, loxylocopa, Moiwxylocopa,
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