Fossil bees are rare, and with some exceptions only those in amber are well enough preserved to shed light on phy-logeny. The fossils so preserved are probably biased toward bees that used resin in nest construction, and therefore sometimes became mired and trapped as they collected the resin. Bees are prone to the development of convergent features in the wing venation, legs, and other parts often visible in fossils, whereas the mouthparts and male genitalia—so often important in phylogenetic studies—are almost never visible. Therefore the relationships, even the family, of a fossil are often not apparent. Nonetheless, some interesting results have emerged from studies of fossil bees, and more can be expected. The geological history ofbees was reviewed by Engel (2001b, 2004b) and Grimaldi and Engel (2005); see also Section 23. The only comprehensive account of fossil bees (Zeuner and Manning, 1976) was written by people not familiar with Recent bees, and accordingly it contributes nothing to our understanding of apiform history beyond bringing together and summarizing relevant literature. For photographs of bee fossils, see plates 17 to 20.
I have not tried to include fossil genera in the systematic account, except where their relationships are clear. I list in Table 22-1 the generic or subgeneric names based on fossil species, in the same format used for the synonymies of Recent bees. When the higher taxon to which a genus belongs is clear, further information may be given in the account of that taxon. In the table the higher taxa are indicated in bold face. It must be admitted that such taxa may sometimes be in doubt. For example, in the genera of "Halictinae" described by Engel (2002a), the basal vein is shown in his illustrations as curved, a halictine character, but it appears curved throughout its length or medially, not specifically toward the basal end as in Recent Halictinae. So while placement in Halictinae based primarily on the curvature of the basal vein is probably correct, some question remains.
The late Jurassic or early Cretaceous hymenopteran described as Palaeapis and placed in the Apidae by Hong (1984) is not a bee. The long stigma suggests a trigonalid, but it may be an aculeate wasp. The slender hind basitar-sus, the strong venation close to the wing apex, and the low position of the antennae, as well as the apparently almost globose scape, indicate that it is not one of the Api-formes.
See Addenda for additional material (including a new family) on fossil bees.
Table 22-1. Generic and Subgeneric Names of Bees Based on Fossils.
Anthophorites Heer, 1849: 97. Type species: Anthophorites mellona Heer, 1850, by designation of Cockerell, 1909d: 315. Anthophorini?
Apiaria Germar, 1839: 210. Type species: Apiaria dubia Germar, 1839, monobasic. [See also Germar, 1849: 66.] Osmiini?
Bombusoides Motshulsky, 1856: 28. Type species: Bombusoides mengei Motshulsky, 1856, monobasic.
Boreallodape Engel, 2001b: 78. Type species: Boreallodape baltica Engel, 2001, by original designation. Boreallodapini.
Calyptapis Cockerell, 1906a: 41. Type species: Calyptapis florissantensis Cockerell, 1906, monobasic. Bombini.
Apis (Cascapis) Engel, 1999b: 187. Type species; Apis armbrusteri Zeuner, 1931, by original designation. Apini.
Chalcobombus Cockerell, 1908d: 326. Type species: Chalco-bombus humilis Cockerell, 1908, by designation of Cockerell, 1909e: 11 = Protobombus Cockerell, 1908. Electrapini.
Megachile (Chalicodomopsis) Engel 1999a: 3. Type species: Megachile glaesaria Engel, 1999, by original designation. Megachilini.
Cretotrigona Engel, 2000a: 3. Type species: Trigona prisa Michener and Grimaldi, 1988, by original designation. Meliponini.
Ctenoplectrella Cockerell, 1909d: 314; also described as new by Cockerell, 1909e: 19. Type species: Ctenoplectrella viridiceps Cockerell, 1909, monobasic. Osmiini.
Cyrtapis Cockerell, 1908c: 339. Type species: Cyrtapis anomalus Cockerell, 1908, monobasic. Halictinae.
Eckfeldapis Lutz, 1993: 180. Type species: Eckfeldapis electrapoides Lutz, 1993, by original designation = Electrapis Cockerell, 1908. Electrapini.
Eickwortapis Michener and Poinar, 1997: 354. Type species: Eickwortapis dominicana Michener and Poinar, 1997, by original designation. Halictinae.
Electrapis Cockerell, 1908d: 326; also described as new by Cockerell, 1909e: 7. Type species: Apis meliponoides Buttel-Reepen, 1906, by designation of Cockerell, 1909e: 8. Electrapini.
Augochlora (Electraugochlora) Engel, 2000b: 32. Type species: Augochlora leptoloba Engel, 2000, by original designation. Augochlorini.
Electrobombus Engel, 2001b: 88. Type species: Electrobombus samlandensis Engel, 2001, by original designation. Electrobombini.
Electrolictus Engel, 2001b: 38. Type species: Electrolictus antiguus Engel, 2001, by original designation. Halictinae.
Eomacropis Engel, 2001b: 46. Type species: Eomacropis glae-saria Engel, 2001, by original designation. Eomacropidini.
Glaesosmia Engel, 2001b: 75. Type species: Glaesosmia genalis Engel, 2001, by original designation. Osmiini.
Glyptapis Cockerell, 1909d: 314; also described as new by Cockerell, 1909e: 13. Type species: Glyptapis mirabilis Cockerell, 1909, monobasic and by designation of Cockerell, 1909e: 14. Osmiini.
Table 22-1. Generic and Subgeneric Names of Bees Based on Fossils (continued)
Hauffapis Armbruster, 1938: 37. Type species: Hauffapis scheuthlei Armbruster, 1938 = Apis armbrusteri Zeuner, 1931, by designation of Zeuner and Manning, 1976: 243. [Hauffapis is not a valid name; see Section 121.] Apini.
Kelneriapis Sakagami, 1978 (June): 232. Type species: Trigona eocenica Kelner-Pillault, 1970, monobasic. Meliponini.
Kelnermelia Moure and Camargo, 1978 (Nov. 17): 565. Type species: Trigona eocenica Kelner-Pillault, 1970, by original designation = Kelneriapis Sakagami, 1978 (June). Meliponini.
Kronolictus Engel, 2002a: 256. Type species: Kronolictus volcanus Engel, 2002, by original designation. Halictinae.
Libellulapis Cockerell, 1906a: 42. Type species: Libellulapis antiquorum Cockerell, 1906, monobasic. Panurginae?
Liotrigonopsis Engel, 2001b: 135. Type species: Liotrigonopsis rozeni Engel, 2001, by original designation. Meliponini.
Lithandrena Cockerell, 1906a: 44. Type species: Lithandrena saxorum Cockerell, 1906, monobasic. Andreninae?
Lithanthidium Cockerell, 1911d: 225. Type species: Lithan-thidium pertriste Cockerell, 1911, monobasic. Anthidiini.
Electrapis (Melikertes) Engel, 1998: 95. Type species: Electrapis stilbonota Engel, 1998. Melikertini.
Meliponorytes Tosi, 1896: 352. Type species: Meliponorytes succini Tosi, 1896, by designation of Sandhouse, 1943: 570. Meliponini.
Melissites Engel, 2001b: 129. Type species: Melissites trigona Engel, 2001, by original designation. Melikertini.
Ocymoromelitta Engel, 2002a: 261. Type species:
Ocymoromelitta sorella Engel, 2002, by original designation. Halictinae.
Oligochlora Engel, 1997a: 336. Type species: Oligochlora eickworti Engel, 1997, by original designation. Augochlorini.
Paleoeuglossa Poinar, 1999: 29. Type species: Paleoeuglossa melissiflora Poinar, 1999, by original designation. Euglossini.
Paleomelitta Engel, 2001b: 42. Type species: Paleomelitta nigripennis Engel, 2001, by original designation. Paleomelittidae.
Pelandrena Cockerell, 1909f: 159. Type species: Pelandrena reducta Cockerell, 1909, monobasic. Andreninae?
Apis (Priorapsis) Engel, 1999b: 188. Type species: Apis vestuta Engel, 1998, by original designation. Apini.
Probombus Piton, 1940: 218. Type species: Probombus hirsutus Piton, 1940, monobasic. Bombini.
ProhalictusArmbruster, 1938: 48. Type species: Prohalictus schemppi Armbruster, 1938, monobasic. Halictinae.
Trigona (Proplebeia) Michener, 1982: 44. Type species: Trigona dominicana Wille and Chandler, 1964, by original designation. Meliponini.
Protobombus Cockerell, 1908d: 326; also described as new by Cockerell, 1909e: 9. Type species: Protobombus indecisus Cockerell, 1908, monobasic; also by designation of Cock-erell, 1909e: 10. Electrapini.
Protolithurgus Engel, 2001b: 51. Type species: Protolithurgus ditomeus Engel, 2001, by original designation. Protolithurgini.
Protomelecta Cockerell, 1908: Type species: Protomelecta brevipennisCockerell, 1908, monobasic. Melectini.
Pygomelissa Engel and Wappler, in Wappler and Engel, 2003: 909. Type species: Pygomelissa lutetia Engel and Wappler, 2003, by original designation.
Electrapis (Roussyana) Manning, 1961: 306. Type species: Apis palmnickenesis Roussy, 1937, by original designation. Melikertini.
Oligochlora (Soliapis) Engel, 2000b: 47. Type species: Oligochlora rozeni Engel, 2000, by original designation. Augochlorini. [Engel (2002a) considered this a synonym of Oligochlora s. str.]
Sophrobombus Cockerell, 1908d: 326; also described as new by Cockerell, 1909g: 21. Type species: Sophrobombus fatalis Cockerell, 1908, monobasic = Electrapis Cockerell, 1908. Electrapini.
Succinapis Engel, 2001b: 115. Type species: Succinapis proboscidia Engel, 2001, by original designation.
Apis (Synapis) Cockerell, 1907a: 229. Type species: Apis henshawi Cockerell, 1907, monobasic. Apini.
Thaumastombus Engel, 2001b: 109. Type species: Thaumasto-bombus andreniformis Engel, 2001, by original designation. Electrapini.
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