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THE MEGACHILINE BEES of CALIFORNIA (Hymenoptera: Megachilidae)

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THE MEGACHILINE BEES of CALIFORNIA (Hymenoptera: Megachilidae) BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 3 THE MEGACHILINE BEES OF CALIFORNIA (Hymenoptera: Megachilidae) BY PAUL D. HURD, JR. (Department of Entomology and Parasitology, University of California, Berkeley) AND CHARLES D. MICHENER (Department of Entomology, University of Kansas, Lawrence) UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES 1955 BULLETIN OF THE CALIFORNIA INSECT SURVEY Editors: E. G. Linsley, E. 0. Essie, S. B. Freeborn, R. L. Usinger Volume 3, pp. 1-248 Submitted by Editors, May 14, 1954 Issued May 31, 1955 Price $3.50 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES CALIFORNIA CAMBRIDGE UNIVERSITY PRESS LONDON, ENGLAND PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA Contents Introduction ....................... 1 Bionomics ...................... 1 Floral relationships ................... 2 Distribution ..................... 2 Acknowledgments .................... 3 Systematics ....................... 6 Family Megachilidae ............ ....... 6 Phyletic relationships within the Megachilini ............. 8 Key to the Genera of Megachilini found in the Western Hemisphere .......10 Heriades .............. .......11 Chelos toma .................... 24 Chelostomopsis ................... 37 Hoplitis .............. .......44 Anthocopa ............. .......94 Proteriades ............. Ashmeadiella ..................... 153 Diceratosmia ............. .215 Osmia ...................... 217 Megachile .............. .......217 Coelioxys .............. .......222 Literature Cited ...................... 223 Index to the California Megachiline bees ...............239 Index to the floral visitational records ................243 Introduction This study is the result of an effort to provide sorts carried to the nest. These materials are usable keys, pertinent descriptive comments, and quite constant for each species but vary consid- detailed dam on distribution and floral visits of erably among the species and genera. Thus most certain of the megachiline bees found in the state Megachile use pieces of leaves, but some species of California. The genera Anthocopa, Ashmeadiella, use pebbles, mud, resin, and other materials in Chelostoma, Chelostomopsis, Diceratosmia, combination with leaves or without leaves. Most Hetiades, Hoplitis, Proteriades are treated in Hoplitis chew up the leaf fragments to make a detail, but the genera Coelioxys, Megachile, and pulp to form partitions between cells. Osmia are considered only to the subgeneric The cells may be placed in natural cavities level. The above mentioned genera are our rep- or in burrows constructed by the female bee in resentatives of the tribe Megachilini of the family advance of cell construction. They may be lined Megachilidae . on all sides by the foreign material brought for The genera which are treated in detail have this purpose, as is usual in Megachile, or only been the subject of special studies by one of partitions between cells may be constructed of us (Michener), who is largely responsible for the such material (as in Hoplitis), in which case the keys, discussions, and descriptive comments. The sides of the cell consist of the walls of the other author (Hurd) has assembled the distribu- burrow. tional and floral data, prepared the maps, and Situations in which nests of these bees are supervised much of the preparation of the manu- likely to occur include holes (often made by script for the press. beetles) in logs, pine cones, galls, and sticks; The terminology is essentially that utilized by burrows in pithy stems such as those of Rubus Michener (1944~)except that, in the light of and various weeds; and burrows or cavities in the newer studies of head structures in insects, the soil. Sometimes they make use of abandoned nests word frons is used instead of the complicated of other bees or wasps and, especially in Europe, term, "supraantennal area," and the segments of of abandoned snail shells. the metasoma (=second and following true abdomi- Each cell is provisioned with a mass of pollen nal segments) are numbered starting with one. and honey occupying the lower end of the cell, Thus the segment behind the first morphological and on top of this a single large, elongate, gently abdominal segment or propodeum is the first curved egg is laid. After hatching, the larva eats metasomal segment. As a result, the numbering downward through the pollen mass, generally of the segments corresponds to that most gener- growing rapidly and reaching maturity in less than ally used in taxonomic works on the Hymenoptera. two weeks. It then spins a cocoon consisting of two or more layers of silk, at least one of which Bionomics is impregnated with a solid brown material applied by the larva as a liquid. This substance fills the The bees of the genera discussed in this bulletin interstices in the silken sheet and probably makes are all solitary, that is, each female constructs the cocoon relatively airtight except for a small and provisions her own nest or nests; there is area of loose threads often projecting as a nipple no worker caste. Each nest consists of one to at one end of the cocoon. several cylindrical cells. As usual in the mega- After the cocoon is complete the larva may go chilids, but unlike most other bees, the cell into a resting stage, often called the prepupa, in walls are built of foreign materials of various which it remains for many months. Among the 2 BULLETIN OF THE CALIFORNIA INSECT SURVEY genera considered in this study, most of the native bees have enabled us to evaluate more species that have but one generation per year properly the economy of these bees. As mentioned are in flight in the spring and reach the prepupal earlier, they are associated with the native stage before midsummer and remain in this stage floras; more particularly the majority are associ- through the rest of summer, fall, and winter, ated with the floras of the broad desert and emerging after a brief pupal stage in the spring. chaparral or scrub areas of the state. Other species (e+, Megachile brevis Cresson) The pollination of floras of such areas is have as many as four generations per year so that significant in connection with problems of range the prepupal resting stage is very short except management, watershed, and soil conservation. for the overwintering generation. As may be seen by reference to the floral listings, In many species males appear in flight before these native bees visit many of these econom- the females and disappear long before the females. ically important plants and therefore are likely Thus collections often consist entirely of one to be responsible for much pollination of them. sex or another, and proper association of sexes It is therefore advisable that in the planning and is often a problem. A number of species are still practice of the management programs, considera- known from only one sex. Mating usually occurs tion be given to perpetuation of these bee faunas. very soon after emergence of females. Considerable public interest has developed The above generalizations on the biology of in the appreciation and preservation of wild these bees are based on meager data concerning flowers within California. It is our obligation to few species. We urgently need exhaustive etho- society to contribute tangibly to a knowledge of logical and ecological studies of several species the ecology of these plants. It is apparent that in order to interpret better the comparative be- among the flowers visited by this group of bees havioral characteristics and in order to judge are many of the more aesthetically appreciated more intelligently the importance of these bees wild flower forms. As cross-pollination is often to man. important in the reproduction of such plants, it is likely that these bees play a significant role Floral Relationships in ensuring survival of these natural assets of the state. Many of the megachiline bees are probably im- portant pollinators of native plants. Only in a few known instances, for example Dicerutosmia Distribution subfasciata, are they known to assume any role in the pollination of agriculturally importan Among the eight genera discussed in detail in crops. Even here, their role may be regarded as this paper, 124 species (144 species and sub- but incidental in relation to the principal economy species) are known or anticipated in California. of these bees. This is not especially surprising A few additional species represented by one or if we realize that the majority of megachiline a very few specimens occur within the state but bees are restricted in their pollen-collecting have not yet been described. A total of 116 habits to certain types of flowers, and sometimes named species is recognized at present from to the flowers of a particular family or even California. This is a surprisingly large propor- genus. It is probable that this floral selection tion of the 154 species of these genera known by the bees is operative in restricting them to from the entire Western Hemisphere. This is the native floras and, further, is responsible to partly owing to the fact that these genera are a large degree for accentuating the differences of northern distribution, several of them Holarc- among geographically differentiating faunas. It tic; no species in the eight genera concerned seems probable, because of the floral preferences, occurs in South America and only one or two in that this group of native bees is not likely to the tropical regions of Central America. It is contain many representatives which wiil assume possibly
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