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Brooks 1988 Anthophora Comp sl t:l:l:!:l:i:l:l:3:l:l':l:li:l:l i :3:3f:3:!:3:3:!:3:3'i'.3..!.3, 11111:;;3:;:;1ç;:;1 THE UNIVEI3SITY OF'I(ANSAS )SCIENCE BULLETIT{ Systematics and Phylogeny of the Anthophorine Bees (Hy*enoptera: Anthophoridae; Anthophorini) By Robert Iry. Brooks Vol.53, No.9, pp.436-575 September 30, 19BB i v I t F r I E L^ t i ; t. ? É. To I I ProfessorCharles Duncan Michener this study is dedicated in anticipation of the aa I fl occasionof his seventiethbirthdav 22 September19BB : his retirement ; and F 1., from the departmentsof Entomology and Ecology r;. and Systematics 18 Mav 1989 !': l+ THE UNIVERSITY OF I(ANSAS SCIENCE BULLETIN Vol. 53, No. 9, pp. 436-575 September30, 19BB Systematics and Phylogeny of the Anthophorine Bees (Hymenoptera: Anthophoridae; Anthophorini) Ronnnr W. Bnoors CONTENTS Introduction +39 Acknowledgem;;,; :::: +39 Historical Review . ++0 Methods. ++0 Characters Employed. ++2 Tribe Anthophorini . ++7 Unplaced Species . 450 Anthophorini incertae sedis . 450 K.y to Genera of the AnthoPhorini 450 Genus Anthophora Latreille 450 Kty to Subgenera of AnthoPhora. 4s3 Subgenus Anthophora Latreille s. str. +57 Pyganthophora new subgenus 460 Lophanthophora new subgenus +6+ Mystacanthophora new subgenus +66 Caranthophora new subgenus +70 Subgenus Paramegilla Friese +70 Subgenus Melea Sandhouse. +76 Subgen us AnthophoroidesCockerell and Cockerell 478 Subgen us Anthomegilla Marikovskaya . 481 Rhinomegilla new subgenus +82 Petalosternonnew subgenus +B+ Daslmegilla new subgenus . 486 Subgenus C/asodon Patton . 486 Subgenus HelioPhila Klug . ..489 Anthophora incertae sedis . +96 Genus Amegilla Friese ..+96 K"y to Subgenera of Amegilla 498 Zebramegilla new subgenus 502 Subgenus Amegilla Friese s. str. 504 Dizonamegilla new subgenus 505 Megamegilla new subgenus 505 Subgen us Aframegilla PoPov 506 Ackmonopsis new subgenus 508 Micramegilla new subgenus 508 Subgenus Zonamegilla PoPov . 510 Notomegilla' new subgenus . 511 Glossamegilla new subgenus . 5t2 'opo{otqog snue8 leluar.ro pue rrpr"Ioq ^ ou âql repun pezrurduou(s 'seureu st sts(otot1(u.gsnue8 eql uortrppg uI cgrcads ot xepû u" pu" .rurpodorqell pue rurroqdoçuy 3o sarcedsqnspue sercedso1 paqdde ueeq e^"q reqr seurcu I[e jo rsq " r*r,ro -uds ivrou l(urduouroq pazru8ocer d1ir,leuro3 g xrpueddy pue ,suorleuEtsep cràd1oeu pue crddlopal 'pepnlcur ro3 7 xrpueddy eog sr sdnor8 crraue8qnsJo uorssnrsrp 'sle^âl '[€qul lecrqde.r8oeE-orq V cr.reuotqns pue crreue8 eql 1z uezrrEare sdrqsuorl€ler Jr]srp€IJ elqeqord 'Nuotmor! 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' ' 'I Zgç rurroqdoqluy;o sarcadg,roN xrpusddy LZ9"' PellCerruerâll'I '.dqder8oa8org €2E... "sdrqsuorlela1rrrsrpelC l1g"' "'telq-J IZç"' "'gâlquJ 0zç"' "'z3twJ 0zç"' "'lelqsr 9Iç"' .....srsdpuvrpsrp"lC SIg... ""'srp?saoyatutrl1zîtwy çIg"' .' "'1ârâIf,o3rpo{otosy €Iç"' snua8qng Nrrsl-rnfl scNsrDS svsNvx do ^rrsus^rN.[I sHJ Bffi SysrpvrATrcs AND PHyrocENy oF THE ANtHopHoRrNB BsBs +39 INTRODUCTION Natuurlijke Historie, Leiden, who allowed me to stay in his home and study his fine collection of anthophorine and This study provides a classification bees. I will remember him for his painstaking and proposes a phylogeny for the genera and beautifully illustrated works on bees and for the many subgenera of anthophorine bees. The tribe warm interesting discussions in his garden. I wish to Anthophorini, worldwide in distribution, thank Yu. A. Pesenko of the Zoological Institute of Leningrad for his help in establishing the status of considered here to include only two and Anthophorabactriana Popov and A. pubescenslabronigro genera, Anthophora and Amegilla, is distin- Lebedev. guished from other Anthophorinae by three Mick C. Day of the British Museum (Natural His- submarginal cells, the first recurrent vein tory) and Astrid Loken of the Zoological Institute of Blindern, Norway, brought to my attention many over- meeting the posterior margin of submarginal looked details concerning the identity of Apis aceraorum with cell 2 near its midpoint, the forewing Linneaus. simple aJar papillae, the second abcissa of I am grateful to Lorraine Hammer of the Electron vein M + Cu of the hind wing subequal in Microscope Facility of the University of Kansas for the microscope. length to the cu-v crossvein, and the elon- opportunity to use the scanning electron I am indebted to the following collectors and mu- gate, narrow gonostylus (if present) never seum curators who have generously lent material: C.J. Habro- flattened and paddle shaped. The van Achterberg*, Rijksmuseum van Natuurlijke Histo- podini consisti.g of Deltoptila, Elaphropoda, rie, Leiden, The Netherlands; A. V. Antropov, Zoo- Habropoda (: EmQhoropsis rr. syn.), Habro- logical Museum of Moscow, U.S.S.R.; P. Arnaud and Academy of Sciences, San phorula and Pachymelus, and formerly in- W. Pulawski, California Francisco; R. Ayala,Instituto de Biologîa, México; C' in Anthophorini, are as distinct cluded the Baroni-urbani*, Naturhistorisches Museum, Basel, from the Anthophorini as the Emphorini are Switzerland; G. E. Bohart and F. D. Parker, Bee from the Eucerini. Biology and Systematics Laboratory Utah State Uni- and Chemical Anthophorine bees are mainly xerophilic .rersity,Logan; G. R. Brown, Biological ResearchInstitute, Rydalmere, Australia; C. Besuchet, and abundant in Mediterranean climates. Muséum d'Histoire Naturelle, Geneva, Switzerland; J. over Among the North American species C. Cardale, Australian National Insect Collection, 90% occur in the dty southwest. In the Old cslRo, canberra; H. V. Daly, IJniversity of califor- World, where most anthophorine bees occur, nia, Berkeley; C. Darling (presently at the Royal Toronto, Canada) and W' P' the most speciose area is the Mediterranean Ontario Museum, Stephen, Oregon State LJniversity, Corvallis; E. Diller, L+ subgenera of Anthophora Region. Of the ZoàlogischeStaatssammlurrg, Munich, West Germany; eight are found in the New World; only one M. Dylewska, ZoologischesInstitut, Polska Akadernia is restricted to that area. They are scarce in Nauk, Krakow; G. C. Eickwort, cornell university, Else*, British Museum tropical forests but are often found in trop- Ithaca, New York; G. R. (Natural History), London; M. Fischer*, Naturhis- ical montane environs. Amegilla is strictly iorisches Museum, Vienna, Austria; S. Frommer and Old World and flies in the summer, while Riverside; Y' J.-Hirashima, C. Hall, University of California, Anthophora is worldwide in distribution and Kyushu University, Fukuoka, Japan; the commonly flies in the sPritg. late P. D. Hurd, J.. and R. J. McGinley* (McGinley was formerly of the Museum of Comparative Zoology, Cambridge, Massachusetts),National Museum of Nat- ACKNOWLEDGMENTS ural History Washington, D.C.; A. Jansson, Zoo' I am sincerelygrateful to CharlesD. Michenerof the logiska Museum, tlniversity of Helsinki, Finland; J. University of Kansas,who providedencouragement, Càsevitz Weulersse and the late S. Kelner-Pillault*,
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