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. . , . . (HYMENOPTERA, APOIDEA) ...

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III. : , 9. .. 9.1. (139). - (140). (141). (142). (142). (143). - (144). (145).

9.2. (145). (147). (148).

9.3. (150). - (150). (152). (152).

9.4. - (153). (153). (155). (157). (158).

9.5. (159). (160). (161). (163). . .

- (163).

9.6. Apis (164). (164). (166). (167). .

(169).

10. 10.1. ..... (171). (171). (172). (172). (173).

10.2. (178). (178). - (180).

10.3. : (182). (183). (184). (186).

10.4. (188). (189).

(191). (191).

10.5. (193). (194). (195).

11. ................................ 11.1 (196). (197). (198). , (198).

11.2. (199). (200).

11.3. (201). (201). (202). (203).

11.4. (203). (203). (204). (205). (206).

11.5. (207). (207).

(208).

11.6. (209). (210). (210).

11.7. (211). (212). (213).

11.8. (214). - (214).

12. 12.1. (217). (218). (218). (219). (220). (223). (225).

12.2. (228). (229). (232). (234). : (236).

12.3. : (237). (238). (239).

(241).

13. 13.1. (243). (244).

13.2. . 13.3. (247). (250).

13.4. ................. (251). (252). (253).

Summary CONTENTS Preface Part I. INTRODUCTION: DIVERSITY, DISTRIBUTION, LIFE CYCLES, A N D TROPHICAL LINKS OF BEES, METHODS OF STUDY OF THEIR BIOLOGY Chapter 1. General characteristics of bees 1.1. Origin and diversity 1.2. Biogeography and distribution Comparison the faunas of zoogeographical regions (8). Regional and local faunas (9). O l d and New W o r l d. Holarctic species (9). Synopsis of the fauna of Russia and neighbouring countries (10).

1.3. Life cycles and individual development Life history. Genetic control of sex (11). Form, size, and development of eggs (12). Emergence of larvae and their feeding (12). Defecation (15). Cocoon spining (16). Diapause, mating, and imago emergence (19). Phenology, voltinism (19).

1.4. Natural enemies and deseases Taxonomic and biological diversity (22). Parasites of immature phases and cleptoparasites (22).

Predators and nest-destroyers (25). Parasites of imago (27). Deseases (28).

Chapter 2. Trophical links and foraging behavior 2.1. Anthophily in bees M a i n sources of food (30). Collecting plant oils (30). Pseudocopulation and collecting sexual attractants (31).

2.2 Kinds of trophical links Terminology (32). Oligo- and polylectes (32).

2.3. Oligolectic bees in the fauna of Russia and neighbouring countries 2.4. Adaptations of the oligolectic bees........................................................... Territorial coincidence with food plants (35). Morphological and behavioral adaptations (35). On the heritability of the specialization (36).

2.5. On co-evolution of bees and angiosperm plants Conditions for reciprocal selection (37). On competition in the system plantpollinators* (38).

Coordination of phylogenies (41).

2.6. Foraging behavior Energetics (42). Learning (43). Flower constancy (44). Organization of foraging (44). On the theory of optimal foraging (45).

2.7. Pollination of entomophilous plants Importance of bees in agriculture (46). Breeding (48). Introduction (50).

Chapter 3. Cleptoparasitic bees 3.1. Taxonomic diversity 3.2. Origins of cleptoparasitism 3.3. Relation parasitehost: a taxonomic aspect 3.4. Relation parasitehost: a biological aspect Chapter 4. Methods for study of bee biology. Classifications of nests......... 4.1. Methods for study of nests and nesting behavior Search of nests (63). Study of nests in soil (63). Study of nests in plant (64). Timing (65).

Observation about behavior within a nest (65). Individual marking (66). Cameral investigations (67).

4.2. About classification of nests and kinds of the nesting of bees Initial classifications (68). Gutbier's classification (69). Malyshev's classification (70). Stephen's classification (71). Common defects of the classifications (72).

Contents Part II. N E S T I N G OF BEES A N D ITS EVOLUTION Chapter 5. Location and general structure of nests 5.1. Sites and ways of nests constructing Nests in soil (74). Nests within plants (74). Nests within natural cavities (75). Nests on exposed surfaces (76). Factors affecting the selection of nesting sites (76). Usurpation of else's nests (77).

Unusual sites for nests (77).

5.2. The main parts of the nest. Sequence of nest making up Entrance into the nest (77). M a i n burrow (80). Lateral burrows (80). B l i n d burrows (81). Nest chambers (81). Nest plug (82) 5.3. Principal types of nest patterns The approach to classification (83). Simple branched nests (83). Twice-branched nests (85). Linear unbranched nests (85). Linear-branched nests (85). Nests with sedentary cells (86). Chamber nests with the main burrow (88). Nests consisting of free cells without the main burrow (88). Nests without cells (88). Dependence of nest patterns on external factors (88) Chapter 6. Cell 6.1. General structure Form and orientation (90). Building materials (93). Vacant and other unhabited cells (96). C e l l re-use (96).

6.2. Methods for constructing and lining Methods for constructing (96). Constructing of nest chambers (98). Lining with excreted substances (99). L i n i n g with oils (100).

6.3. Defence functions of a cell Humidity control (100). Defence against pathogenous microorganisms (101). Initial desinfection (102). Protection against mechanical damages and enemies (102).

6.4. Formation provisions, laying of eggs and capping cells Composition, form, and consistence of provisions (103). Protection of provisions against soiling (104). E g g placement in a cell (106). Cell capping (107). Efficiency of females and sequence of forming cells (108).

Chapter 7. The proto-bee (ancestral bee) and its nest 7.1. Traditional and new hypotheses Wasp-like ancestor of bees (110). The proto-bee: traditional hypothesis (111). W h i c h bees are the most primitive (111). The proto-bee: a new hypothesis (112).

7.2. Pollen carrying on the body surface M a i n arguments (114). Flattened metabasitarsus (114). A scopa did not originate repeatedly (115).

7.3. Machining of cell walls The outline of argumentation (116). Functions of the pygidial and metabasitibial plates (116). Bees with the plates (117). Bees without the plates (118). The presence of the plates is a plesiomorphic character (119).

7.4. Nesting in soil, the role of mandibles 7.5. Cocoon spinning Developed spinning organs (120). Pattern of the character cocoon present vs. cocoon absent* (121).

7.6. About secretory lining of cells Structure of glossa (121). Negative correlation of lining and cocoon (122). Composition, sources, and methods for making of lining (123).

7.7. Provisions for larvae was dough-like M a i n arguments (124). Storage of liquid provisions or of relatively dry pollen grains (125). On transition to vegetable food (126).

7.8. Nest architecture............... General design (126). Orientation of cells, presence of cell caps (127).

Chapter 8. Evolution of bee nesting 8.1. Evolution of nesting in burrowing bees 8.2. Changes of the nest substrate Migration to rotten wood (129). Migration to firm plant materials (129). Nesting of Hylaeinae in plants (131).

Contents 8.3. Transition to nesting in natural cavities and on exposed surfaces 8.4. Evolution of nesting in Megachilidae Previous suppositions voiced before (133). A new hypothesis (135). Main directions of evolution (136).

8.5. The main trends in the evolution of bee biology Insurance of expansion (137). Care about offspring (138).

Part III. SOCIAL LIFE: T H E MAIN FORMS, ORIGIN, A N D EVOLUTION............... Chapter 9. The main forms of sociality in bees.... 9.1. The forms of sociality: terms and classification Eusociality (139). Mono- and polygyny (140). Subsociality (141). Eosociality (142). On p a r a social* colonies (142). On quasisociality (143). Semisociality - the polygynous foundation or terminal state of eusocial colonies (144). A synopsis of the classification (145).

9.2. Aggregations of individuals and nests Sleeping aggregations (145). Nest aggregations (147). Communal nests (148).

9.3. Subsocial colonies General characteristics (150). Primitive-subsocial colonies (150). Eosocial colonies (152). Cases of bisexual colony (152).

9.4. Primitive-eusocial colonies General characteristics (153). Eusocial colonies of the ceratinines (153). Eusocial colonies of the allodapines (155). Eusocial colonies of the xylocopines (157). Eusocial colonies of the euglossines (158).

9.5. Colonies of the meliponines " Nest structure (159). Caste differentiation and determination (160). Queen domination and division of labour (161). Egg-laying by workers (163). Mating. Swarming. Robber bees (163).

9.6. Colonies of apines Composition and distribution of the genus Apis (164). Nest structure (164). Caste differentiation and determination (166). Division of labour (167). Egg-laying by workers. Swarming (169).

Chapter 10. Eusocial colonies of the halictines.. 10.1. History of discovery and studying social life in the halictines Prehistory (171). Confirmation of eusociality (171). Peculiarities of studies of artificial nests (172).

M a i n discoveries (172). Distribution of eusociality in the halictines (173).

10.2. The foundation of colonies..... Foundation of nest (178). Polygynous foundation of a family (178). Structure of the foundress nest (180).

10.3. Eusocial life: castes and hierarchy Sex ratio in the first and subsequent broods (182). Caste differentiation (183). Maintenance of caste structure (184). Change of queens by workers and usurpation of nests by other females (186).

10.4. Family nest and nest behavior Within-nest behavior and general division of labour (188). Building labour (189). Foraging (191).

Nest defense and the recognition of the colony members (191).

10.5. Rearing of reproductive offspring and disintegration of the family Rearing of reproductive offspring (193). Mating (194). Family size and duration of family life (195).

Chapter 11. Bumblebee family.. 11.1. Founding of a family Preparation of a female for family founding and structure of its nest (196). The first brood (197).

Honey pot forming (198). Feeding of larvae of the first brood, formation of the second brood (198).

11.2. Rearing of the brood Types of feeding of larvae (199). Duration of immature development (200).

11.3. Microclimate of a nest and regulation of temperature Microclimate in a nest (201). Thermoregulation mechanisms and temperature conditions of incu bation (201). Ventilation of a nest (202). Body thermoregulation in foragers (203).

11.4. Caste differentiation and division of labour Caste structure of a family (203). Division of labour (203). Foraging (204). Within-nest labour (205). Nest guarding (206).

Contents 11.5. Maintenance of the caste structure Mechanisms of maintenance of caste structure (207). Egg-laying by workers (207). Elimination of gyne eggs by workers (208).

11.6. Rearing of reproductive offspring Beginning of rearing of reproductive forms (209). Rearing of males (210). Behavior of young females and males (210).

11.7. Disintegration of the family and gyne hibernating Family size (211). Seasonal rhythms of activity (212). G y n e hibernating (213).

11.8. Usurpation of nests Replacement of gynes and usurpation of nests by gynes of another bumblebee species (214). Cuckoo bumblebees (214).

Chapter 12. Origin of social life.. 12.1. Hypotheses for the mechanism of caste origin Natural selection and the problem of worker caste (217). Family selection (218). Group selection (218). Mutualism hypothesis (219). K i n selection (220). Polygynous family hypothesis (223).

Parental manipulation (225).

12.2. Haplodiploidy hypothesis Hamilton's discovery (228). K i n selection among individuals with different life strategies (229). M a i n predictions of the hypothesis (232). Conflict in rearing of males (234). Haplodiploidy and euso ciality: taxonomic aspect (236).

12.3. Polygynous founding of a family: a decision of the problem Essence and occurence of the phenomen (237). Statement of the problem (238). Approach to resolving the problem (239). Genetic gains of the strategies queen and sister (241).

Chapter 13. Prerequisites of origin and stages the evolution of eusociality in bees 13.1. Subsocial pathway Hypotheses on pathways to sociality (243). Subsocial hypothesis (244).

13.2. Evolutionary stages of sociality 13.3. Prerequisites for incipiency of eusociality Prerequisites for eusociality origin (247). Conservation of optimal reproduction (250).

13.4. M a i n tendencies and constraints in the evolution of eusociality Trends in the evolution of primitive eusociality (251). Inversion to solitary life (252). Evolution of eusociality in various groups of bees (253) References Summary Annotated index of Latin names of bees (HYMENOPTERA, APOIDEA) ( , ) ( , -) - 1991 .

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