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Novitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3046, 56 pp., 89 figures, 6 tables July 27, 1992 Systematics and Host Relationships of the Cuckoo Bee Genus Oreopasites (Hymenoptera: Anthophoridae: Nomadinae) JEROME G. ROZEN, JR. CONTENTS Abstract ............................................. 2 Introduction ............................................. 2 Acknowledgments ............................................. 3 Relationships of Oreopasites to Other Nomadinae ............... .................. 3 Genus Oreopasites Cockerell .............................................. 6 Phylogenetic Relationships of Species .......................................... 7 Host Associations, Host Specificity, and Effects of Host Evolution ....... ......... 12 Key to Species ............................................. 18 Subgenus Oreopasites Cockerell ............................................. 20 Oreopasites vanduzeei Cockerell ............................................. 20 Oreopasites albinota Linsley ............................................. 33 Oreopasites scituli Cockerell ............................................. 35 Oreopasites arizonica Linsley ............................................. 37 Oreopasites hurdi, new species ............................................. 39 Oreopasites euphorbiae Cockerell .................................... ........ 40 Oreopasites collegarum, new species ......................................... 43 Oreopasitesfavreauae, new species ........................................... 44 Oreopasites powelli, new species ............................................. 47 Perditopasites, new subgenus ............................................. 49 Oreopasites linsleyi, new species ............................................. 49 Oreopasites barbarae, new species ........................................... 53 References ............................................. 55 ' Curator, Department of Entomology, American Museum of Natural History. Copyright © American Museum of Natural History 1992 ISSN 0003-0082 / Price $5.30 2 AMERICAN MUSEUM NOVITATES NO. 3046 ABSTRACT The western North American cuckoo bee species have been discovered and are listed. genus Oreopasites is revised. Evidence indi- All hosts belong to the highly specialized Cal- cates that it is most closely related to the Old liopsini and Perditini, exclusively New World World ammobatine genera Ammobates, Par- tribes of the andrenid bee subfamily Panur- ammobatodes, and Melanempis. Eleven spe- ginae. Oreopasites s.s. is restricted to certain cies are recognized, six of which are new: 0. North American subgenera ofCalliopsis: No- hurdi, 0. collegarum, 0. favreauae, 0. pow- madopsis, Macronomadopsis, Micronoma- elli, 0. linsleyi, and 0. barbarae. Oreopasites dopsis, and Hypomacrotera. Perditopasites linsleyi is designated the type species ofa new has been associated with three subgenera of subgenus Perditopasites, which also includes Perdita. An analysis of the relationships of 0. barbarae. A key to species is presented, the monophyletic clade to which these pan- all species are described (or redescribed), and urgines belong with all genera (Holcopasites, their important taxonomic features are illus- Caenoprosopis, Caenoprosopina, Neolarra, trated. Species distributions are mapped, and and Oreopasites) of cleptoparasitic bees phylogenetic relationships are evaluated. Or- known to attack it failed to reveal a single eopasites vanduzeei is the most wide-ranging, case where the same host species supports polytypic species. Although it had been di- more than one cleptoparasitic genus. How- vided into subspecies, these subdivisions are ever, different species in Perdita are parasit- no longer justified in light of new knowledge ized by Oreopasites and Neolarra, and dif- of anatomical variation, hosts, and distri- ferent species of Calliopsis (Hypomacrotera) butions. Consequently the following subspe- are attacked by Oreopasites and Holcopasites. cific names applied to it are placed in syn- Although some host species are parasitized onymy: diabloensis Linsley, melanantha by more than one species ofOreopasites, there Linsley, vernalis Linsley, and xerophila Lin- are no known cases where host sharing occurs sley. sympatrically. A number ofOreopasites have The hosts or suspected hosts ofall but three more than one host. INTRODUCTION This is a revision of the genus Oreopasites 1944) which is otherwise restricted to the Pa- (Ammobatini) consisting of small to minute learctic and African regions. cleptoparasitic bees restricted to the western Most individuals are uniformly colored part ofNorth America. The genus ranges from with a dark brown to black head and meso- the state of Washington to Baja California soma, a red metasoma, and reddish to black Sur and eastward to western Nebraska and legs. They have been rarely collected, prob- the Big Bend region of Texas (fig. 9). All of ably because of their small size and unifor- its species attack ground-nesting bees be- mity of color pattern and because, like most longing to the specialized New World tribes parasitic bees, they do not commonly visit Calliopsini and Perditini of the Panurginae flowers, where most bee collecting is carried (Andrenidae). Oreopasites adults enter open out. They can, however, be abundant at brood cells ofthe host while the host females times-flying over the surface of the ground are absent and hide their eggs by inserting at the nesting sites of their hosts. them into the cell walls. Upon hatching the When Linsley (1941) revised the genus, he hospicidal first instar searches out and kills found that the interpretation of the species the host egg or larva and then feeds for the in Oreopasites was difficult. With only 125 rest of its larval life on the provisions that specimens, he recognized that large Oreo- had been supplied by the host for its young. pasites had larger hosts and small ones tended The 11 recognized species of this parasitic to have smaller hosts and that the structural genus are the only known New World mem- differences between parasitic species were of- bers of the Ammobatini (sensu Michener, ten limited. He concluded that the host spe- 1 992 ROZEN: OREOPASITES 3 cies of Oreopasites might influence the ana- The following institutions through persons tomical features of their cleptoparasites in charge oftheir collections kindly lent spec- through the amount and perhaps quality of imens used in this investigation: Academy of food (pollen and nectar) they used in provi- Natural Sciences of Philadelphia, Pennsyl- sioning their cells. Although recognizing this vania (D. Otte); Bee Biology and Systematics to be a potential problem, he had no way to Laboratory, USDA-ARS, Utah State Uni- determine the "nature or extent ofthis influ- versity, Logan (G. E. Bohart, P. F. Torchio); ence." I have dealt with this problem by fo- R. M. Bohart Museum of Entomology, Uni- cusing my investigations on extensive collec- versity of California, Davis (L. S. Kimsey, tions ofOreopasites that have been associated the late R. 0. Schuster); British Museum of with their hosts. Natural History, London; California Acad- Fortunately Luisa Ruz undertook a mas- emy of Sciences, San Francisco (W. J. Pu- sive, generic-level investigation on the phy- lawski); California Department of Food and logenetic relationships of the Panurginae on Agriculture, Sacramento (M. S. Wasbauer); a worldwide basis (Ruz, 1987). Part of this Essig Museum ofEntomology (formerly Cal- investigation (Ruz, 1991) treats that group of ifornia Insect Survey), University of Califor- specialized, New World panurgines to which nia, Berkeley (the late P. D. Hurd, J. A. Pow- all known hosts of Oreopasites belong. Her ell); Cornell University (G. C. Eickwort, J. work makes possible the analysis of the host K. Liebherr); National Museum of Natural relationships of Oreopasites presented in this History, Smithsonian Institution, Washing- paper. ton, D.C. (R. J. McGinley); Natural History Museum of Los Angeles County, California (R. R. Snelling); Oregon State University, ACKNOWLEDGMENTS Corvallis (J. D. Lattin); University of Ari- I especially wish to extend my appreciation zona, Tucson (F. Werner); Snow Entomolog- to Marjorie Favreau for her years ofresearch ical Museum, University of Kansas, Law- assistance on this project in the American rence (R. Brooks, C. D. Michener); collection Museum ofNatural History and at the South- of R. C. Bechtel, Reno, Nevada; and Uni- western Research Station. I acknowledge with versity of California, Riverside (S. I. From- gratitude the efforts in laboratory and/or field mer). of Byron Alexander, Jaun Barberis, Lillian Early phases of this research were sup- Floge, Ronald J. McGinley, Barbara L. Roz- ported in part by National Science Grant G- en, and Kenneth C. Rozen. Rose Ismay, 13315, and National Science Grants GB-5407 Phyllis Browne, Beatrice Brewster, and Deb- and GB-321193 provided subsequent fund- orah J. Hickman have helped in typing and ing for fieldwork. organizing versions of the manuscript. This manuscript has been reviewed by Bry- The resident directors ofthe Southwestern an N. Danforth and Roy R. Snelling, and E. Research Station, Mont A. Cazier, Vincent Gorton Linsley, James S. Miller, and David D. Roth, and Wade C. Sherbrooke, offered K. Yeates have read sections of it. I wish to their hospitality and assistance during my thank all of these people for the time they stays there. Without the Station
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