Phylogenetics of Bees Editors Rustem Abuzarovich Ilyasov Doctor in Division of Life Sciences Major of Biological Sciences, and Convergence Research Center for Insect Vectors Incheon National University Songdo-dong, Incheon South Korea and Ufa Scientific Center Institute of Biochemistry and Genetics Russian Academy of Sciences Ufa Russia Hyung Wook Kwon Professor in Division of Life Sciences Major of Biological Sciences, and Convergence Research Center for Insect Vectors Incheon National University Songdo-dong, Incheon South Korea p, p, A SCIENCE PUBLISHERS BOOK A SCIENCE PUBLISHERS BOOK Cover credit: Cover illustration reproduced by kind courtesy of the first editor, Rustem Abuzarovich Ilyasov. 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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identifi cation and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Names: Ilyasov, Rustem Abuzarovich, 1980- editor. | Kwon, Hyung Wook, 1968- editor. Title: Phylogenetics of bees / editors: Rustem Abuzarovich Ilyasov, Hyung Wook Kwon. Description: Boca Raton, FL : CRC Press, [2020] | Includes bibliographical references and index. Identifiers: LCCN 2019037641 | ISBN 9781138504233 (hardcover) Subjects: LCSH: Bees--Phylogeny. | Bees--Variation. Classification: LCC QL563 .P495 2020 | DDC 595.79/9--dc23 LC record available at https://lccn.loc.gov/2019037641 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Preface The first idea of writing this book came about after reading the books of Ruttner “Biogeography and Taxonomy of Honeybees” (1988), Michener “The Bees of the World” (2000), Kipyatkov “Life Cycles in Social Insects Behaviour, Ecology And Evolution” (2006), and Hepburn “Honeybees of Asia” (2011). Meanwhile, I met with Professor Hyung Wook Kwon who invited me to a postdoctoral position at Incheon National University, South Korea. In Korea, we are working very hard and met many times with local beekeepers and scientists at conferences of The Korean Beekeeping Society. Compared with western honeybees, eastern honeybees have received very thin coverage on honey bee taxonomy, phylogenetics, physiology, biochemistry, genetics, and pathology. Especially, the eastern honey, bee Apis cerana, has no clearly defined subspecies like the western honey, bee Apis mellifera. In order to address this disparity, we decided to write this book, which combines the efforts of western and eastern teams of scientists. The book contains works not only about phylogenetics but also about current aspects of adaptivity and expansion to the New World. We fervently hope that this collation will provide a stimuli to broaden the base of the development of taxonomy and phylogenetics of the Asian and European honeybees. We also hope that it will stimulate further research, both empirical and theoretical, in this important and promising field. We wish to thank all contributing authors as well as the referees for their collaboration and help. Also, I appreciate my scientific advisors Professor Alexey Nikolenko of the Institute of Biochemistry and Genetics of Ufa Federal Research Center of Russian Academy of Sciences and Professor Hyung Wook Kwon of the Incheon National University for their help in editing and for the constructive discussions on basic ideas in this book. Rustem Abuzarovich Ilyasov Incheon National University Songdo-dong, Incheon, Yeonsu-gu September 24, 2019 Acknowledgments We are grateful to the CRC Press publisher, Russian Government Assignments (RGA) (the Grant No. No. AAAA-A16-116020350026-0), Russian Found of Basic Research (RFBR) (the Grant No. 17-44-020648 Povolzhye), the National Research Foundation of Korea (NRF), the Korea government (MSIP) (the Grant No. 2016R1A2B3011742), the Cooperative Research Program for Agriculture Science and Technology Development (RDA) (the Grant No. PJ012526), and the Postdoctoral Fellowships in the Incheon National University (2017–2019) for kindly supporting both editors– Sc.D. R.A. Ilyasov and Prof. H.W. Kwon. Contents Preface iii Acknowledgments v Introduction ix 1. The Origin and Evolution of the Colony in Apidae 1 Eskov, E.K. 2. Phylogenies of Asian Honey Bees 28 Raffiudin, R. and Shullia, N.I. 3. The Origin of the European Bees and their Intraspecific Biodiversity 58 Brandorf, A.Z. and Rodrigues, M. 4. The Classic Taxonomy of Asian and European Honey Bees 72 Dar, S.A., Dukku, U.H., Ilyasov, R.A., Kandemir, I., Kwon H.W., Lee, M.L. and Özkan Koca, A. 5. Genetic Diversity of Honey Bee Apis mellifera in Siberia 97 Ostroverkhova, N.V., Kucher, A.N., Konusova, O.L., Kireeva, T.N., Rosseykina, S.A., Yartsev, V.V. and Pogorelov, Y.L. 6. Current Drivers of Taxonomic Biodiversity Loss in Asian and 127 European Bees Hatjina, F., Gajda, A. and Dar, S.A. 7. The Loss of Taxonomic Biodiversity of Honey Bees Apis mellifera 144 and Main Breeds in Russia Berezin, A.S., Borodachev, A.V., Borodachev, V.A., Mitrofanov, D.V. and Savushkina, L.N. 8. Breeding Better and Healthy Honey Bees is the Only Way to 178 Save A Native Biodiversity Gregorc, A. 9. Honey Bees in Latin America 206 Requier, F. 10. The History of Honey Bees in North America 222 Seeley, T.D. Conclusion 233 References 237 Index 281 Color Plate Section 283 Introduction Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living and over 2,000 extinct species of Hymenoptera have been described (Mayhew 2007, Janke et al. 2013, Aguiar et al. 2013). Honey bees are flying eusocial insects closely related to wasps and ants, and are known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea and are presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families (Michener 2000, Danforth et al. 2006). They are found on every continent except Antarctica, in every habitat on the planet that contains insect- pollinated flowering plants. Some species, including honey bees, bumble bees, and stingless bees live socially in colonies. Bees are adapted for nectar and pollen nutrition, the former primarily as an energy source, and the latter for protein and other nutrients. Most pollen is used as food for larvae. Bee pollination is important both ecologically and commercially. The decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The size of the bees ranges from tiny stingless bee species whose worker bees are less than two millimeters long to Megachile pluto, the largest species of the leafcutter bee, whose females can attain a length of forty millimeters. The most common bees in the Northern Hemisphere are the Halictidae or sweat bees, but they are small and often mistaken for wasps or hoverfly species. Vertebrate predators of bees include birds, such as bee-eaters; insect predators include beewolves and dragonflies. Human beekeeping or apiculture has been practiced for a thousand years, since at least the times of Ancient Egypt and Ancient Greece. Apart from honey and pollination, honey bees produce beeswax, royal jelly, and propolis. Bees have appeared in mythology and folklore, through all phases of art and literature, from ancient times to the present day, though primarily focused in the Northern Hemisphere, where beekeeping is far more common. Some big questions in the study of honey bees are their evolution and phylogenetics. The resolution hinges on understanding the phylogenetic relationships of the four bee subgroups: orchid bees, bumble bees, stingless bees, and honey bees. These problems are studied by three methods, based on morphological data, nuclear DNA (nDNA) characters, and mitochondrial DNA (mtDNA) features. The fact that no consensus is reached reflects the complexity of the challenges, and shows the need for additional research.