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A Monograph of the Baltic Amber Bees and Evolution of the Apoidea (Hymenoptera)

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A Monograph of the Baltic Amber Bees and Evolution of the Apoidea (Hymenoptera) A MONOGRAPH OF THE BALTIC AMBER BEES AND EVOLUTION OF THE APOIDEA (HYMENOPTERA) MICHAEL S. ENGEL Research Scientist, Division of Invertebrate Zoology, Presently: Research Associate, Division of Invertebrate Zoology, American Museum of Natural History; Fellow, the Linnean Society of London; Assistant Curator, Division of Entomology, Natural History Museum and Biodiversity Research Center; Assistant Professor, Department of Ecology and Evolutionary Biology, 1460 Jayhawk Boulevard, Snow Hall, University of Kansas, Lawrence, Kansas 66045-7523 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 259, 192 pp., 123 ®gures, 7 color plates, 15 tables Issued April 16, 2001 Price: $17.00 a copy Copyright q American Museum of Natural History 2001 ISSN 0003-0090 Photomicrograph of the holotype (IMGP-K72) of Glyptapis mirabilis Cockerell. CONTENTS Abstract ....................................................................... 5 Zusammenfassung .............................................................. 6 Introduction .................................................................... 7 Historical Review ............................................................ 13 Age and Origin of Amber .................................................... 15 Materials and Methods ......................................................... 17 Specimen Preparation and Study ............................................... 18 Format ..................................................................... 18 General Morphology ........................................................... 19 Prosoma .................................................................... 20 Mesosoma .................................................................. 23 Metasoma .................................................................. 32 Systematic Paleontology ........................................................ 33 Superfamily Apoidea Latreille ................................................. 34 Division Apiformes .......................................................... 35 Short-Tongued Versus Long-Tongued Bees ..................................... 35 Short-Tongued Bees .......................................................... 36 Family Halictidae Thomson ................................................... 36 Subfamily Halictinae Thomson .............................................. 36 Tribe Halictini Thomson ................................................. 37 Genus Electrolictus, new genus ......................................... 38 Family Paleomelittidae, new family ............................................ 41 Genus Paleomelitta, new genus ......................................... 42 Family Melittidae Schenck .................................................... 44 Subfamily Macropidinae Robertson .......................................... 46 Tribe Eomacropidini, new tribe ........................................... 46 Genus Eomacropis, new genus ......................................... 46 Long-Tongued Bees .......................................................... 49 Family Megachilidae Latreille ................................................. 49 Subfamily Lithurginae Newman ............................................. 50 Tribe Protolithurgini, new tribe ........................................... 51 Genus Protolithurgus, new genus ....................................... 51 Subfamily Megachilinae Latreille ............................................ 53 Tribe Osmiini Newman .................................................. 54 Genus Glyptapis Cockerell ............................................. 56 Genus Ctenoplectrella Cockerell ........................................ 68 Genus Glaesosmia, new genus .......................................... 75 Family Apidae Latreille ...................................................... 76 Subfamily Xylocopinae Latreille ............................................ 77 Tribe Boreallodapini, new tribe ........................................... 77 Genus Boreallodape, new genus ........................................ 78 Subfamily Apinae Latreille ................................................. 86 Tribe Electrobombini, new tribe ........................................... 88 Genus Electrobombus, new genus ....................................... 88 Tribe Electrapini Engel ................................................... 90 Genus Electrapis Cockerell ............................................. 92 Genus Protobombus Cockerell .......................................... 99 Genus Thaumastobombus, new genus ................................... 109 Tribe Melikertini, new tribe .............................................. 112 Genus Succinapis, new genus .......................................... 115 Genus Melikertes Engel ............................................... 123 3 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 259 Genus Melissites, new genus .......................................... 129 Genus Roussyana Manning ............................................ 131 Tribe Meliponini Lepeletier de Saint Fargeau .............................. 133 Genus Kelneriapis Sakagami .......................................... 134 Genus Liotrigonopsis, new genus ...................................... 135 Apoidea Incertae Sedis ...................................................... 137 Genus and Species Indeterminate ......................................... 137 Missing Taxa of Uncertain Identity ....................................... 138 Cladistic Analyses ............................................................ 139 Lithurginae ................................................................ 139 Xylocopinae ............................................................... 140 Corbiculate Apinae ......................................................... 142 Discussion ................................................................... 150 Origin of Bees ............................................................. 150 The Eocene Bee Fauna ...................................................... 159 Epilogue ................................................................... 161 Acknowledgments ............................................................ 161 References ................................................................... 162 Appendix 1: Bees in Amber and Copal .......................................... 175 Appendix 2: Tribal Classi®cation and Geological Distribution of Bees .............. 177 Appendix 3: Catalog of Family- and Genus-Group Names for Fossil Bees ........... 180 Appendix 4: Full Titles of Journals Abbreviated in References ..................... 182 Index of Taxonomic Names .................................................... 185 Color Plates .............................................................. 186±192 2001 ENGEL: BALTIC AMBER BEES 5 ABSTRACT The bees found as Baltic amber inclusions are revised and the history of studies on these fossils is brie¯y reviewed. In total this subtropical Eocene fauna contains 36 species and 18 genera, all extinct. These are classi®ed here into nine tribes, six subfamilies, and ®ve families of which six tribes and one family are unknown in the modern fauna. The following taxa are described as new to science: PALEOMELITTIDAE, new family; Boreallodapini, Electro- bombini, Eomacropidini, Melikertini, Protolithurgini, new tribes; Ctenoplectrellina, new subtribe; Boreallodape, Electrobombus, Electrolictus, Eomacropis, Glaesosmia, Liotrigon- opsis, Melissites, Paleomelitta, Protolithurgus, Succinapis, Thaumastobombus, new genera; Boreallodape baltica, B. mollyae, B. striebichi, Ctenoplectrella cockerelli, C. grimaldii, Elec- trapis krishnorum, Electrobombus samlandensis, Electrolictus antiquus, Eomacropis glae- saria, Glaesosmia genalis, Glyptapis densopunctata, G. disareolata, Liotrigonopsis rozeni, Melikertes clypeatus, Melissites trigona, Paleomelitta nigripennis, Protobombus basilaris, Protolithurgus ditomeus, Succinapis goeleti, S. micheneri, S. proboscidea, Thaumastobom- bus andreniformis, new species (seven new family-, 11 new genus-, and 22 new species- group taxa). The genus Electrapis is found to be paraphyletic and the subgenera Melikertes, Roussyana, and Protobombus are given generic status outside of Electrapis. The subtribe Electrapina is elevated to tribal rank among the corbiculate Apinae and the subfamily Glyp- tapinae of Cockerell is reduced to subtribal rank within Osmiini. The genera Chalcobombus and Sophrobombus are newly synonymized with Protobombus. Glyptapis reducta Cockerell is synonymized with G. fuscula Cockerell, Electrapis minuta Kelner-Pillault with Apis palm- nickenensis Roussy, Ctenoplectrella splendens Kelner-Pillault and C. dentata Salt both with C. viridiceps Cockerell, Electrapis apoides Manning and Chalcobombus humilis Cockerell both with Protobombus indecisus Cockerell, and Electrapis bombusoides Kelner-Pillault with E. tornquisti Cockerell (new synonymies). The following new combinations are proposed: Electrapis martialis (Cockerell), Melikertes proavus (Menge), M. stilbonotus (Engel), Kelner- iapis eocenica (Kelner-Pillault), Protobombus fatalis (Cockerell), P. hirsutus (Cockerell), and Electrapis martialis (Cockerell) (new combinations). A lectotype is designated for Electrapis minuta Kelner-Pillault and neotypes designated for Apis meliponoides Buttel-Reepen, A. palm- nickenensis Roussy, Chalcobombus humilis Cockerell, C. hirsutus Cockerell, C. martialis Cockerell, Ctenoplectrella dentata Salt, C. viridiceps Cockerell,
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