The Biology and External Morphology of Bees
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3?00( The Biology and External Morphology of Bees With a Synopsis of the Genera of Northwestern America Agricultural Experiment Station v" Oregon State University V Corvallis Northwestern America as interpreted for laxonomic synopses. AUTHORS: W. P. Stephen is a professor of entomology at Oregon State University, Corval- lis; and G. E. Bohart and P. F. Torchio are United States Department of Agriculture entomolo- gists stationed at Utah State University, Logan. ACKNOWLEDGMENTS: The research on which this bulletin is based was supported in part by National Science Foundation Grants Nos. 3835 and 3657. Since this publication is largely a review and synthesis of published information, the authors are indebted primarily to a host of sci- entists who have recorded their observations of bees. In most cases, they are credited with specific observations and interpretations. However, information deemed to be common knowledge is pre- sented without reference as to source. For a number of items of unpublished information, the generosity of several co-workers is ac- knowledged. They include Jerome G. Rozen, Jr., Charles Osgood, Glenn Hackwell, Elbert Jay- cox, Siavosh Tirgari, and Gordon Hobbs. The authors are also grateful to Dr. Leland Chandler and Dr. Jerome G. Rozen, Jr., for reviewing the manuscript and for many helpful suggestions. Most of the drawings were prepared by Mrs. Thelwyn Koontz. The sources of many of the fig- ures are given at the end of the Literature Cited section on page 130. The cover drawing is by Virginia Taylor. The Biology and External Morphology of Bees ^ Published by the Agricultural Experiment Station and printed by the Department of Printing, Ore- gon State University, Corvallis, Oregon, 1969. Contents Introduction 3 External Morphology of Bees 3 Adult Bees 3 Bee Larvae 21 Bee Pupae 29 Systematic List of Included Taxa 32 Key to the Genera of Northwestern Bees 33 Characteristics of the Bee Taxa 43 Distribution and Estimated Number of Species of Northwestern Bee Genera 65 The Biology and Behavior of Bees 66 Geographical and Ecological Distribution 66 Solitary and Social Behavior 67 Bionomics 68 Nesting Activities 73 General Nest Architecture 76 Classification of Nest Patterns 85 General Nest Features 90 Cell Characteristics 96 Provisions 101 Oviposition and Development 107 Foraging . .116 Flight Periods 121 Parasitic Bees 122 Natural Enemies of Bees 124 Defense Against Natural Enemies 128 Literature Cited 130 Sources of Figures 134 Index 135 The Biology and External Morphology of Bees With a Synopsis of the Genera of Northwestern America W. P. STEPHEN, G. E. BOHART, and P. F. TORCHIO Introduction ENTOMOLOGISTS, BOTANISTS, and AGRICULTURISTS all also presented an opportunity to speculate on several have occasion to concern themselves with bees. The en- facets of systematics and ethology. tomologist is fascinated by their complex behavior pat- This work reviews present knowledge of the mor- terns, the botanist needs to evaluate their significance in phology and biology of bees in general, but emphasizes floral biology, and the agriculturist must take them into northwestern forms and, in many instances, uses them account as vital factors in crop production. Finally, the as standards of reference. No attempt is made to syste- apiculturist often wishes to extend his knowledge of matically cover morphological and biological informa- bees beyond the confines of the honey bee. Recent years tion for each taxon, although morphological information have seen a revival of interest in bees by all of these of particular value for the separation of taxonomic groups and, with it, a profusion of taxonomic and bio- groups is included. The taxonomic treatment does not logical literature. This interest among scientists has been extend below the generic level and is confined to north- stimulated by exciting biological and behavioral discov- western forms except for a more comprehensive treat- eries, attempts to determine the value of biological in- ment of families. The Northwest, as here interpreted, formation, and a concern in pollinator management by includes the area west of the Rocky Mountains, bounded farmers engaged in the production of seed, fruit, and on the south by the latitude 41° N, and extending north vegetable crops. to include British Columbia and Alaska. (See the map It is our hope that the followir j general synthesis inside the front cover.) Several genera included in the of knowledge about bees, with special emphasis on key have not yet been taken in the above-defined area, northwestern genera, will be of general interest to many but because they are known to occur immediately to the entomologists, botanists, and agriculturists and may south or east there is a possibility that they may have have special value as a handbook for workers in the escaped detection or else that they may soon become Northwest. The study was prepared so that it would members of our bee fauna. be useful to students for class and field use, and apprise No species authors names are cited in the text; workers of the present state of knowledge in bees so that rather, they are included in the index at the end of this existing gaps may be more systematically filled. It has publication. External Morphology of Bees Adult Bees not to imply that absolute structural homology may not exist throughout Insecta and that our ultimate goal The sections dealing with the morphological termi- should not be directed towards its realization, but rather nology used in this series of studies are presented not as that we should adopt a standardized terminology so that definitive treatises on bee morphology, but as guides to one apoid taxonomist may know immediately to which those structures having taxonomic significance. Much of structure the other is referring. the terminology used herein is that of M'ichener (1944a) Classically, three body regions are designated among which has been widely accepted in apoid systematics. insects: the head, thorax, and abdomen. Michener Our knowledge of comparative morphology in insects in (1944a) states that the use of the term mesosoma for general, and Hymenoptera in particular, is still woe- the fused thoracic region in the Clistogastra is morpho- fully inadequate, and any attempt to establish a fixed logically preferable to thorax. This region is composed general terminology for the morphologically recogniz- of four actual segments: the prothorax, mesothorax, able structures in the bees would be premature. This is metathorax, and the first abdominal segment (the pro- podeum), which is sharply constricted posteriorly to gum of the second true abdominal segment is referred form the petiole common to bees, wasps, and ants. Mit- to as the first metasomal tergum, and the tergum of the chell (1960) reverts to the use of the term thorax for third true morphological segment of the abdomen is re- this region, indicating that he prefers the older and more ferred to as the second metasomal tergum, and so on generally used term for ease of reference. We follow (see Fig. 1). Mitchell's use of the term thorax even though it includes The following section covers the principal morpho- the first abdominal segment as an integral portion of logical structures employed in species discrimination that structure. among bees. The discussion of these structures includes The third body region, generally referred to as the as much comparative information as is deemed neces- abdomen, is separated from the thorax by a very marked sary for persons unfamiliar with the group. The illus- constriction. This has been referred to by Michener as trations, however, are based principally on the halictid the metasoma, for it consists of the second and subse- bee, Nomia melanderi, as it is generally one of the quent morphological segments, the first being immovably most readily available species in northwestern America. attached to the thorax. In this bulletin the term abdomen Although this bee is not as "primitive" as members of is applied to this general body region. The term meta- the Colletidae, it is more so than Apis and Anthophora, somal is used only in its adjectival sense when referring which have been used as standards of reference in pre- to the various abdominal terga or sterna. Thus the ter- vious studies (Snodgrass, 1956; Michener, 1944a). Scuto-Scutellar Pronotum \ Scutellum \Lateral Lobe vMetanotum Metasoma-- \\ Scutum v\Propodeum Metasomal Tergum I /Gradulus Compound Eye\ \ "Metasomal Sterna Basitibial Plate -r Ba si tars us Tarsal Segment ^-Claw Nomia melanderi FIGURE 1. Lateral view of Nomia melanderi female. Vertex Ocelli Genal Area Supra-antennal Area (Frons) Inner Orbital Margin Paraocular Area Antenna! Socket-.. Antennal Sclerite-^ -Antennal Suture— Subantennal Suture -Supraclypeal Area- Epistomal Suture— -Ant. Tentorial Pit Clypeus- Malar Space- --Labrum --Labral Fimbria- Mandible "Inner Tooth of Mandible b Preoccipital Ridge Foramen Magnum Proboscidial Fossa 2 e Nomia melanderi FIGURE 2. Frontal, lateral, and ventral views of the head of Nomia melanderi. Head and Apis (Fig. 4) the compound eyes extend over the The head is hypognathous with the face perpendic- top of the head and meet at the mid line. The inner mar- ular to the longitudinal axis of the body (Fig. 1). The gins of the compound eyes, inner orbital margins, may large convex compound eyes occupy much of the lateral converge below as in most colletids and halictids (Figs. surface of the head (Fig 2a). In some genera, 5, 6); may be parallel as in andrenids and megachilids such as Ceratina (Fig. 3), the compound eyes do (Figs. 7, 8) ; or more rarely diverge as in some dufour- not nearly reach the upper margin of the sides of the eines and andrenids. In most genera of bees the eyes are head, whereas in the males of certain species of Boinbus bare. However, in the American Coelioxys, Apis, and in the single species of Holcopasites that occurs in The antennal bases in Nomia are located about mid- the Northwest, the eyes are densely covered with short, way between the vertex and the apical margin of the erect hairs.