In some ways this may seem the wrong time to write on the systematics of the bees of the world, the core topic of this book. Morphological information on adults and larvae of various groups has not been fully developed or exploited, and molecular data have been sought for only a few groups. The future will therefore see new phylogenetic hypotheses and improvement of old ones; work in these areas continues, and it has been tempting to defer completion of the book, in order that some of the new information might be included. But no time is optimal for a systematic treatment of a group as large as the bees; there is always significant research under way. Some genera or tribes will be well studied, while others lag behind, but when fresh results are in hand, the latter may well overtake the former. I conclude, then, that in spite of dynamic current activity in the field, now is as good a time as any to go to press.
This book constitutes a summary of what I have been able to learn about bee systematics, from the bees themselves and from the vast body of literature, over the many years since I started to study bees, publishing my first paper in 1935. Bee ecology and behavior, which I find fully as fascinating as systematics, are touched upon in this book, but have been treated in greater depth and detail in other works cited herein.
After periods when at least half of my research time was devoted to other matters (the systematics of Lepidoptera, especially saturniid moths; the biology of chigger mites; the nesting and especially social behavior of bees), I have returned, for this book, to my old preoccupation with bee systematics. There are those who say I am finally finishing my Ph.D. thesis!
My productive activity in biology (as distinguished from merely looking and being fascinated) began as a young kid, when I painted all the native plants that I could find in flower in the large flora of Southern California. When, after a few years, finding additional species became difficult, I expanded my activities to drawings of insects. With help from my mother, who was a trained zoologist, I was usually able to identify them to family. How I ultimately settled on Hymenoptera and more specifically on bees is not very clear to me, but I believe it had in part to do with Perdita rhois Cockerell, a beautiful, minute, yellow-and-black insect that appeared in small numbers on Shasta daisies in our yard each summer. The male in particular is so unbeelike that I did not identify it as a bee for several years; it was a puzzle and a frustration and through it I
became more proficient in running small Hymenoptera, including bees, through the keys in Comstock's Introduction to Entomology.
Southern California has a rich bee fauna, and as I collected more species from the different flowers, of course I wanted to identify them to the genus or species level. Somehow I learned that T. D. A.Cockerell at the University of Colorado was the principal bee specialist active at the time. Probably at about age 14 I wrote to him, asking about how to identify bees. He responded with interest, saying that Viereck's Hymenoptera ofConnecticut (1916) (which I obtained for $2.00) was not very useful in the West. Cresson's Synopsis (1887) was ancient even in the 1930s, but was available for $10.00. With these inadequate works I identified to genus a cigar box full of bees, pinned and labeled, and sent them to Cockerell for checking. He returned them, with identifications corrected as needed, and some specimens even identified to species.
Moreover, Cockerell wrote supporting comments about work on bees and invited me to meet him and P. H. Timberlake at Riverside, California, where the Cockerells would be visiting. Timberlake was interested in my catches because, although I lived only 60 miles from Riverside, I had collected several species of bees that he had never seen. Later, he invited me to accompany him on collecting trips to the Mojave and Colorado deserts and elsewhere.
Professor and Mrs. Cockerell later invited me to spend the next summer (before my last year in high school) in Boulder with them, where I could work with him and learn about bees. Cockerell was an especially charming man who, lacking a university degree, was in some ways a second-class citizen among the university faculty members. He had never had many students who became seriously interested in bees, in spite of his long career (his publications on bees span the years from 1895 to 1949) as the principal bee taxonomist in North America if not the world. Probably for this reason he was especially enthusiastic about my interest and encouraged the preparation and publication of my first taxonomic papers. Thus I was clearly hooked on bees well before beginning my undergraduate work at the University of California at Berkeley.
As a prospective entomologist I was welcomed in Berkeley and given space to work among graduate students. During my undergraduate and graduate career, interacting with faculty and other students, I became a comparative morphologist and systematist of bees, and prepared a dissertation (1942) on these topics, published with some additions in 1944. The published version included a key to the North American bee genera, the lack of which had sent me to Professor Cockerell for help a few years before. Especially important to me during my student years at Berkeley were E. Gorton Linsley and the late Robert L. Usinger.
There followed several years when, because of a job as lepidopterist at the American Museum of Natural History, in New York, and a commission in the Army, my research efforts were taken up largely with Lepi-doptera and with mosquitos and chigger mites, but I continued to do limited systematic work on bees. It was while in the Army, studying the biology of chigger mites, that I had my first tropical experience, in Panama, and encountered, for the first time, living tropical stingless honey bees like Trigona and Melipona and orchid bees like Euglossa at orchid flowers. In 1948 I moved to the University of Kansas, and since about 1950 almost all of my research has been on bees.
Until 1950, I had gained little knowledge of bee behavior and nesting biology, having devoted myself to systematics, comparative morphology, and floral relationships, the last mostly because the flowers help you find the bees. In 1950, however, I began a study of leafcutter bee biology, and a few years later I began a long series of studies of nesting biology and social organization of bees, with emphasis on primitively social forms and on the origin and evolution of social behavior. With many talented graduate students to assist, this went on until 1990, and involved the publication in 1974 of The Social Behavior ofthe Bees. Concurrently, of course, my systematic studies continued; behavior contributes to systematics and vice versa, and the two go very well together.
Across the years, I have had the good fortune to be able to study both behavior and systematics of bees in many parts of the world. In addition to shorter trips ofweeks or months, I spent a year in Brazil, a year in Australia, and a year in Africa. The specimens collected and ideas developed on these trips have been invaluable building blocks for this book.
Without the help of many others, preparing this book in its present form would have been impossible. A series of grants from the National Science Foundation was essential. The University of Kansas accorded me freedom to build up a major collection of bees as part of the Snow Entomological Division of the Natural History Museum, and provided excellent space and facilities for years after my official retirement. Students and other faculty members of the Department of Entomology also contributed in many ways. The editorial and bibliographic expertise of Jinny Ashlock, and her manuscript preparation along with that of Joetta Weaver, made the job possible. Without Jinny's generous help, the book manuscript would not have been completed. And her work as well as Joetta's continued into the long editorial process.
It is a pleasure to acknowledge, as well, the helpful arrangements made by the Johns Hopkins University Press and particularly the energy and enthusiasm of its science editor, Ginger Berman. For marvelously detailed and careful editing, I thank William W. Carver of Mountain View, California.
The help of numerous bee specialists is acknowledged at appropriate places in the text. I mention them and certain others here with an indication in some cases of areas in which they helped: the late Byron A. Alexander, Lawrence, Kansas, USA (phylogeny, Nomada); Ricardo Ayala, Chamela, Jalisco, Mexico (Centridini); Donald B. Baker, Ewell, Surrey, England, UK; Robert W. Brooks, Lawrence, Kansas, USA (Anthophor-ini, Augochlorini); J. M. F. de Camargo, Ribeirao Preto, Sao Paulo, Brazil (Meliponini); James W. Cane, Logan, Utah, USA (Secs. 1-32 of the text); Bryan N. Danforth, Ithaca, New York, USA (Perditini, Halic-tini); H. H. Dathe, Eberswalde, Germany (palearctic Hylaeinae); Con-nal D. Eardley, Pretoria, Transvaal, South Africa (Ammobatini); the late George C. Eickwort, Ithaca, New York, USA (Halictinae); Michael S. Engel, Ithaca, New York, USA (Augochlorini, fossil bees); Elizabeth M. Exley, Brisbane, Queensland, Australia (Euryglossinae); Terry L. Gris-wold, Logan, Utah, USA (Osmiini, Anthidiini); Terry F. Houston, Perth,Western Australia (Hylaeinae, Leioproctus); Wallace E. LaBerge, Champaign, Illinois, USA (Andrena, Eucerini); G. V. Maynard, Canberra, ACT, Australia (Leioproctus); Ronald J. McGinley, Washington
D.C., USA (Halictini); Gabriel A. R. Melo, Ribeiräo Preto, Säo Paulo, Brazil (who read much of the manuscript); Robert L. Minckley, Auburn, Alabama, USA (Xylocopini); Jesus S. Moure, Curitiba, Paraná, Brazil; Christopher O'Toole, Oxford, England, UK; Laurence Packer, North York, Ontario, Canada (Halictini); Alain Pauly, Gembloux, Belgium (Malagasy bees, African Halictidae); Yuri A. Pesenko, Leningrad, Russia; Stephen G. Reyes, Los Baños, Philippines (Allodapini); Arturo Roig-Alsina, Buenos Aires, Argentina (phylogeny, Emphorini, Tapinotaspi-dini, Nomadinae); David W. Roubik, Balboa, Panama (Meliponini); Jerome G. Rozen, Jr., New York, N.Y., USA (Rophitini, nests and larvae of bees, and ultimately the whole manuscript); Luisa Ruz, Valparaíso, Chile (Panurginae); the late S. F. Sakagami, Sapporo, Japan (Halictinae, Allodapini, Meliponini); Maximilian Schwarz, Ansfelden, Austria (Coe-lioxys); Roy R. Snelling, Los Angeles, California, USA (Hylaeinae); Os-amu Tadauchi, Fukuoka, Japan (Andrena); Harold Toro, Valparaíso, Chile (Chilicolini, Colletini); Danuncia Urban, Curitiba, Paraná, Brazil (Anthidiini, Eucerini); Kenneth L. Walker, Melbourne, Victoria, Australia (Halictini); V. B. Whitehead, Cape Town, South Africa (Rediviva); Paul H.Williams, London, England, UK (Bombus); Wu Yan-ru, Beijing, China; Douglas Yanega, Belo Horizonte, Minas Gerais, Brazil, and Riverside, California.
The persons listed above contributed toward preparation or completion of the book manuscript, or the papers that preceded it, and also in some cases gave or lent specimens for study; the following additional persons or institutions lent types or other specimens at my request: Josephine E. Cardale, Canberra, ACT, Australia; Mario Comba, Cecchina, Italy (Tetralonia); George Else and Laraine Ficken, London, England, UK; Yoshihiro Hirashima, Miyazaki City, Japan; Frank Koch, Berlin, Germany; Yasuo Maeta, Matsue, Japan; the Mavromoustakis Collection, Department of Agriculture, Nicosia, Cyprus (Megachilinae).
The illlustrations in this book are designed to show the diversity (or, in certain cases, similarity or lack of diversity) among bees. It was entirely impractical to illustrate each couplet in the keys—there are thousands of them—and I made no effort to do so, although references to relevant text illustrations are inserted frequently into the keys. Drs. R. J. McGinley and B. N. Danforth, who made or supervised the making of the many illustrations in Michener, McGinley, and Danforth (1994), have permitted reuse here of many of those illustrations. The other line drawings are partly original, but many of them are from works of others, reproduced here with permission. I am greatly indebted to the many authors whose works I have used as sources of illustrations; specific acknowledgments accompany the legends. In particular I am indebted to J. M. F. de Ca-margo for the use of two of his wonderful drawings of meliponine nests, and to Elaine R. S. Hodges for several previously published habitus drawings of bees. Modifications of some drawings, additional lettering as needed, and a few original drawings, as acknowledged in the legends, are the work of Sara L. Taliaferro; I much appreciate her careful work.
The colored plates reproduce photographs from the two sources indicated in the legends: Dr. E. S. Ross, California Academy of Sciences, San Francisco, California, USA, and Dr. Paul Westrich, Maienfeldstr. 9, Tübingen, Germany. I am particularly indebted to Drs. Ross and Westrich for making available their excellent photographs. It is worth noting here that many other superb photographs by Westrich were published in his two-volume work on the bees of Baden-Württemberg (Westrich, 1989).
Svetlana Novikova and Dr. Bu Wenjun provided English translations of certain materials from Russian and Chinese, respectively. Their help is much appreciated.
The text has been prepared with the help of the bees themselves, publications about them, and unpublished help from the persons listed above. I have not included here the names of all the persons responsible for publications that I have used and from which I have, in many cases, derived ideas, illustrations, bases for keys, and other items. They are acknowledged in the text. Several persons, however, have contributed previously unpublished keys that appear under their authorship in this book. Such contributions are listed below, with the authors' affiliations.
"Key to the Palearctic Subgenera of Hylaeus" by H.H. Dathe, Deutsches Entomologisches Institut, Postfach 10 02 38, D-16202 Eberswalde, Germany.
"Key to the New World Subgenera of Hylaeus" by Roy R. Snelling, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007, USA. "Key to the Genera of Osmiini of the Eastern Hemisphere," "Key to the Subgenera of Othinosmia," and "Key to the Subgenera of Protosmia by Terry L. Griswold, Bee Biology and Systematics Laboratory, UMC 53, Utah State University, Logan, Utah 84322-5310, USA. "Key to the Genera of the Tapinotaspidini" by Arturo Roig-Alsina, Museo Argentino de Ciencias Naturales, Av. A. Gallardo 470, 1405 Buenos Aires, Argentina.
I have modified the terminology employed in these keys, as necessary, to correspond with that in use in other parts of this book (see Sec. 10). Several contributions became so modified by me that the original authors would scarcely recognize them. I have identified them by expressions such as "modified from manuscript key by . . ."
Names of authors of species are not integral parts of the names of the organisms. In behavioral or other nontaxonomic works I omit them except when required by editors. But in this book, which is largely a systematic account, I have decided to include them throughout for the sake of consistency.
A measure of the success of this book will be the need for revision as new work is completed and published. Not only does this book contain a great deal of information about bees, but, by inference or explicitly, it indicates myriad topics about which more information is needed. I hope that it points the way for the numerous researchers who will take our knowledge beyond what is here included, and beyond what is to be found in the nearly 2,500 items in the Literature Cited.
Lawrence, Kansas 1999
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