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Download Download Morphological phylogeny of Megachilini and the evolution of leaf-cuter behavior in bees (Hymenoptera: Megachilidae) Victor H. Gonzalez, Grey T. Gustafson, & Michael S. Engel Journal of Melitology No. 85 ISSN 2325-4467 3 July 2019 On the cover: A female of Megachile sp. preparing to take a freshly cut coinvine [Dal- bergia ecastaphyllum (L.) Taub. (Fabaceae: Faboideae: Dalbergieae)] leaf section back to her nest (Frenchman’s Forest Natural Area, Palm Beach County, Florida; photograph by Bob Peterson; used with permission). Journal of Melitology Bee Biology, Ecology, Evolution, & Systematics The latest buzz in bee biology No. 85, pp. 1–123 3 July 2019 Morphological phylogeny of Megachilini and the evolution of leaf-cuter behavior in bees (Hymenoptera: Megachilidae) Victor H. Gonzalez1,2, Grey T. Gustafson2,3, & Michael S. Engel2,3,4 Abstract. A unique feature among bees is the ability of some species of Megachile Latreille s.l. to cut and process fresh leaves for nest construction. The presence of a razor between the female mandibular teeth (interdental laminae) to facilitate leaf-cuting (LC) is a morphological novelty that might have triggered a subsequent diversifcation in this group. However, we have a lim- ited understanding of the phylogeny of this group despite the large number of described species and the origins and paterns of variations of this mandibular structure are unknown. Herein, using a cladistic analysis of adult external morphological characters, we explored the relation- ships of all genera of Megachilini and the more than 50 subgenera of Megachile s.l. We coded 272 characters for 8 outgroups and 114 ingroup species. Depending on the weighting scheme (equal or implied weighting), our parsimony analyses suggested the monophyly of Megachile s.l. and that either Noteriades Cockerell or the clade Coelioxys Latreille + Radoszkowskiana Popov is the extant sister group of all other Megachilini. In addition, we conducted Bayesian total- evidence tip-dating analyses to examine other possible hypotheses of relationships and paterns of variation of the interdental lamina. Our analyses suggest that interdental laminae developed asynchronicaly from two diferent structures in the mandible, and difer in their phenotypic plasticity. Character correlation tests using phylogenetic pairwise comparisons indicated that the presence of interdental lamina is not associated with head size, mandible size and shape, and pubescence on the adductor interspace. We discuss the implications of our fndings for the classifcation of Megachilini and the development of novel evolutionary, ecological, and func- tional hypotheses on this behavior. New taxa established are Pseudoheriadini Gonzalez & En- gel, new tribe, Ochreriadini Gonzalez & Engel, new tribe, Cremnomegachile Gonzalez & Engel, new genus, Rozenapis Gonzalez & Engel, new genus, and Saucrochile Gonzalez & Engel, new genus, along with the following new combinations: Cremnomegachile dolichosoma (Benoist), new combination, Rozenapis ignita (Smith), new combination, and Saucrochile heriadiformis (Smith), new combination. 1 Undergraduate Biology Program, Haworth Hall, 1200 Sunnyside Avenue, University of Kan- sas, Lawrence, Kansas 66045, USA ([email protected]). 2 Department of Ecology & Evolutionary Biology, Haworth Hall, 1200 Sunnyside Avenue, Uni- versity of Kansas, Lawrence, Kansas 66045, USA ([email protected]). 3 Division of Entomology, Natural History Museum, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66045-4415, USA ([email protected]). 4 Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, USA. doi: htp://dx.doi.org/10.17161/jom.v0i85.11541 Copyright © V.H. Gonzalez, G.T. Gustafson, & M.S. Engel. Creative Commons Atribution-NonCommercial-NoDerivatives 4.0 International (CC BY-NC-ND 4.0). ISSN 2325-4467 2 Journal of Melittology No. 85 CONTENTS Abstract ..................................................................................................................................... 1 Introduction ............................................................................................................................. 3 Diversity of Megachilidae .................................................................................................. 6 Diversity and Phylogeny of Megachilini ......................................................................... 7 What is the Genus Megachile? ............................................................................................ 7 Fossil Record ....................................................................................................................... 9 Material and Methods ......................................................................................................... 10 Morphological Phylogeny of Megachilini ..................................................................... 10 Taxon sampling .............................................................................................................. 10 Morphological data ....................................................................................................... 11 Data compilation ........................................................................................................... 11 Data characterization .................................................................................................... 26 Phylogenetic analyses .................................................................................................. 26 Phylogenetic signal of morphological characters .................................................. 28 Origin of the Interdental Lamina in the Female Mandible ......................................... 28 Evolutionary Origins of the Interdental Lamina .......................................................... 29 Molecular data ............................................................................................................... 29 Combined data .............................................................................................................. 30 Phylogenetic analyses .................................................................................................. 30 Paterns of variation of the interndental lamina ....................................................... 33 Results .................................................................................................................................... 33 Morphological-based Phylogeny of Megachilini ......................................................... 33 Phylogenetic signal ....................................................................................................... 34 Origin and Evolution of the Interdental Lamina ...................................................... 35 Combined phylogenies ................................................................................................ 35 Types of interdental lamina ....................................................................................... 36 Evolution of the interdental lamina ......................................................................... 39 Discussion .............................................................................................................................. 40 Phylogenetic Relationships within Megachilini .......................................................... 40 Monophyly of subgenera of Megachile s.l. ................................................................ 43 Phylogenetic signal of morphological characters .................................................... 44 Origins and Paterns of Variation of the Interdental Lamina ................................... 45 Origin of Leaf-cuting Behavior ...................................................................................... 45 Classifcatory Implications .............................................................................................. 48 Family Megachilidae .................................................................................................... 48 Tribe Megachilini .......................................................................................................... 50 Conclusions and Future Directions ............................................................................... 54 Limitations of the Study .................................................................................................. 55 Descriptions of New Taxa ............................................................................................... 56 Key to extant tribes of Megachilinae ........................................................................ 56 Pseudoheriadini Gonzalez & Engel, new tribe ..................................................... 57 Ochreriadini Gonzalez & Engel, new tribe ........................................................... 58 Cremnomegachile Gonzalez & Engel, new genus .................................................... 58 Rozenapis Gonzalez & Engel, new genus ................................................................ 60 Saucrochile Gonzalez & Engel, new genus ................................................................ 62 Acknowledgements .............................................................................................................. 63 References .............................................................................................................................
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