The Bees of the World 2nd Edition

931 Pages·2000·13.29 MB·English

by  Charles D. Michener

The Bees of the World
The Bees
of the World
Charles D. Michener
University of Kansas Natural History Museum
and Department of Entomology
The Johns Hopkins University Press
Baltimore and London
© 2000 The Johns Hopkins University Press
All rights reserved. Published 2000
Printed in the United States of America on acid-free paper
9 8 7 6 5 4 3 2 1
The Johns Hopkins University Press
2715 North Charles Street
Baltimore, Maryland 21218-4363
www.press.jhu.edu
Library of Congress Cataloging-in Publication Data
Michener, Charles Duncan, 1918–
The bees of the world / Charles D. Michener.
p. cm.
Includes bibliographical references.
ISBN 0-8018-6133-0 (alk. paper)
1. Bees Classification. I. Title
QL566.M53 2000
595.79(cid:1)9—dc21 99-30198 CIP
A catalog record for this book is available from the
British Library.
Title page illustration from H. Goulet and J. T. Huber
(1993). Used with permission.
Tomy many students, now scattered over the world,
from whom I have learned much
and to my family, who lovingly tolerate an obsession with bees

Contents
Preface ix 24. Diversity and Abundance 96
New Names xiv 25. Dispersal 99
Abbreviations xiv 26. Biogeography 100
27. Reduction or Loss of Structures 104
1. About Bees and This Book 1 28. New and Modified Structures 106
2. What Are Bees? 2 29. Family-Group Names 111
3. The Importance of Bees 3 30. Explanation of Taxonomic Accounts
4. Development and Reproduction 4 in Sections 36 to 119 112
5. Solitary versus Social Life 9 31. Some Problematic Taxa 114
6. Floral Relationships of Bees 13 32. The Identification of Bees 115
7. Nests and Food Storage 19 33. Key to the Families, Based on Adults 116
8. Parasitic and Robber Bees 26 34. Notes on Certain Couplets in the Key
9. Body Form, Tagmata, and Sex to Families (Section 33) 120
Differences 38 35. Practical Key to Family-Group Taxa,
10. Structures and Anatomical Terminology Based on Females 121
of Adults 40 36. Family Stenotritidae 123
11. Structures and Terminology 37. Family Colletidae 126
of Larvae 53 38. Subfamily Colletinae 130
12. Bees and Sphecoid Wasps as a Clade 54 39. Subfamily Diphaglossinae 164
13. Bees as a Holophyletic Group 55 40. Tribe Caupolicanini 165
14. The Origin of Bees from Wasps 58 41. Tribe Diphaglossini 168
15. Classification of the Bee-Sphecoid 42. Tribe Dissoglottini 170
Clade 60 43. Subfamily Xeromelissinae 171
16. Bee Taxa and Categories 61 44. Tribe Chilicolini 172
17. Methods of Classification 71 45. Tribe Xeromelissini 177
18. The History of Bee Classifications 72 46. Subfamily Hylaeinae 178
19. Short-Tongued versus Long-Tongued 47. Subfamily Euryglossinae 210
Bees 78 48. Family Andrenidae 225
20. Phylogeny and the Proto-Bee 83 49. Subfamily Alocandreninae 228
21. The Higher Classification of Bees 88 50. Subfamily Andreninae 229
22. Fossil Bees 93 51. Subfamily Panurginae 260
23. The Antiquity of Bee Taxa 94 52. Tribe Protandrenini 262
vii
53. Tribe Panurgini 273 89. Subfamily Nomadinae 614
54. Tribe Melitturgini 278 90. Tribe Hexepeolini 618
55. Tribe Protomeliturgini 281 91. Tribe Brachynomadini 620
56. Tribe Perditini 282 92. Tribe Nomadini 624
57. Tribe Calliopsini 292 93. Tribe Epeolini 627
58. Subfamily Oxaeinae 301 94. Tribe Ammobatoidini 633
59. Family Halictidae 304 95. Tribe Biastini 636
60. Subfamily Rophitinae 307 96. Tribe Townsendiellini 639
61. Subfamily Nomiinae 317 97. Tribe Neolarrini 640
62. Subfamily Nomioidinae 330 98. Tribe Ammobatini 641
63. Subfamily Halictinae 333 99. Tribe Caenoprosopidini 646
64. Tribe Halictini 339 100. Subfamily Apinae 647
65. Tribe Augochlorini 377 101. Tribe Isepeolini 652
66. Family Melittidae 396 102. Tribe Osirini 654
67. Subfamily Dasypodainae 399 103. Tribe Protepeolini 658
68. Tribe Dasypodaini 400 104. Tribe Exomalopsini 660
69. Tribe Promelittini 405 105. Tribe Ancylini 665
70. Tribe Sambini 406 106. Tribe Tapinotaspidini 667
71. Subfamily Meganomiinae 409 107. Tribe Tetrapediini 674
72. Subfamily Melittinae 412 108. Tribe Ctenoplectrini 676
73. Family Megachilidae 417 109. Tribe Emphorini 679
74. Subfamily Fideliinae 419 110. Tribe Eucerini 686
75. Tribe Pararhophitini 420 111. Tribe Anthophorini 720
76. Tribe Fideliini 421 112. Tribe Centridini 731
77. Subfamily Megachilinae 424 113. Tribe Rhathymini 739
78. Tribe Lithurgini 427 114. Tribe Ericrocidini 740
79. Tribe Osmiini 431 115. Tribe Melectini 747
80. Tribe Anthidiini 474 116. Tribe Euglossini 754
81. Tribe Dioxyini 521 117. Tribe Bombini 761
82. Tribe Megachilini 526 118. Tribe Meliponini 779
83. Family Apidae 570 119. Tribe Apini 806
84. Subfamily Xylocopinae 575
85. Tribe Manueliini 577 Literature Cited 809
86. Tribe Xylocopini 578 Addenda 871
87. Tribe Ceratinini 593 Index of Terms 873
88. Tribe Allodapini 600 Index of Taxa 877
Color plates follow page 32.
viii
Preface
Insome 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 in-
formation on adults and larvae of various groups has not been fully de-
veloped 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 re-
search 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 over-
take the former. I conclude, then, that in spite of dynamic current activ-
ity 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!
Myproductive activity in biology (as distinguished from merely look-
ing 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 rhoisCockerell, a beautiful, minute, yellow-and-black in-
sect 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
ix