# Beekeeping # Phillips, Everett Franklin, 1878-1951. # # # [1915?] # # Text of original volume reproduced my Albert R. Mann Library, Cornell University. # Scanning my Trigonix # OCR by Abbyy Finereader Pro 5.0 # No manual correction has been made to the OCR'd text. ######################################## tlbe tRurai Science Series EDITED BY L. H. BAILEY BEEKEEPING Efje i&ural Science Series EDITED BY L. H. BAILEY THE SOIL. King. THE SPRAYING OF PLANTS. Lodeman. MILK AND ITS PRODUCTS. Wing. Enlarged and Revised. THE FERTILITY OF THE LAND. Roberts. THE PRINCIPLES OF FRUIT-GROWING. Bailey. 20th Edition, Revised. BUSH-FRUITS. Card. FERTILIZERS. Voorhees. THE PRINCIPLES OF AGRICULTURE. Bailey. 15th Edition^ Revised. IRRIGATION AND DRAINAGE. King. THE FARMSTEAD. Roberts. RURAL WEALTH AND WELFARE. Fairchild. THE PRINCIPLES OF VEGETABLE-GARDENING. Bailey. FARM POULTRY. Watson. Enlarged and Revised. THE FEEDING OF ANIMALS. Jordan. THE FARMER'S BUSINESS HANDBOOK. Roberts. THE DISEASES OF ANIMALS. Mayo. THE HORSE. Roberts. How TO CHOOSE A FARM. Hunt. FORAGE CROPS. Voorhees. BACTERIA IN RELATION TO COUNTRY LIFE. Lipman. THE NURSERY-BOOK. Bailey. PLANT-BREEDING. Bailey and Gilbert. Revised. THE FORCING-BOOK. Bailey. THE PRUNING-BOOK. Bailey. FRUIT-GROWING IN ARID REGIONS. Paddock and Whipple. RURAL HYGIENE. Ogden. DRY-FARMING. Widtsoe. LAW FOR THE AMERICAN FARMER. Green. FARM BOYS AND GIRLS. McKeever. THE TRAINING AND BREAKING OF HORSES. Harper. SHEEP-FARMING IN NORTH AMERICA. Craig. CodpERATiON IN AGRICULTURE. Powell. THE FARM WOODLOT. Cheyney and Wentling. HOUSEHOLD INSECTS. Herrick. CITRUS FRUITS. Coit. PRINCIPLES OF RURAL CREDITS. Morman. BEEKEEPING. Phillips. Frontispiece. — The apiary of the Bureau of Entomology, Drummond, Maryland. BEEKEEPING A DISCUSSION OF THE LIFE OF THE HONEYBEE AND OF THE PRODUCTION OF HONEY BY EVERETT FRANKLIN PHILLIPS, PH.D. IN CHARGE OP BEE CULTURE INVESTIGATIONS, BUREAU OF ENTOMOLOGY, UNITED STATES DEPART- „ MENT OF AGRICULTURE THE MACMILLAN COMPANY LONDON: MACMILLAN & CO., LTD. 1917 Ml rights reserved COPYEIGHT, 1915, BY THE MACMILLAN (JOMPANY. Set up and electrotyped. Published August, 1915. Reprinted July, 1916; February, 1917. E 547 J. 8. Gushing Co. — Berwick & Smith Co. Norwood, Mass., U.S.A. Ho M. H. G. P. PREFACE THE present book is the result of an effort to present a logical discussion of the various phases of the complex subject of beekeeping. It was not planned as a book of rules to which one may go for directions for each day's work, for beekeeping cannot be treated correctly in such a way. The activities of bees vary during the seasons and no two localities present to the bees and their owners exactly the same environmental conditions, so that the successful beekeeper is one who has a knowledge of the activities of bees, whereby he can interpret what he sees in the hives from day to day, and who can mold the instincts of the bees to his convenience and profit. It has seemed desirable in the early chapters to discuss bees as they exist without man's interference, thus giving the foundation on which the practice of beekeeping rests. The beekeeper is not especially interested in the anatomy of the bee and, while it is necessary to use illustrations of various organs and to describe them briefly, an effort has been made to treat the bee as a living animal and to have the discussion deal with physiology and especially with activities, in so far as investigations have thrown light on these processes. In the preparation of the chapters devoted to the management of the apiary, an effort has been made to present the various systems of manipulations in such a way that the underlying principles shall be evident, rather than to attempt to describe each system as if it were separate. The author has been helped by the facilities of the office of the Bureau of Entomology with which he is connected and is viii Preface under obligations to Dr. Jas. A. Kelson and George S. Demuth for friendly advice and assistance. To F. V. Coville, of the Bureau of Plant Industry, thanks are due for assistance on the chapter on the sources of honey and to Dr. C. C. Miller for counsel on spring management and comb honey, on which subjects he is the highest authority. Especially to his wife, the author would express his gratitude for most valuable help. The illustrations with a few exceptions were either drawn for this book from material gathered from many sources or have been borrowed from publications prepared in the office of bee culture investigations of the Bureau of Entomology. The new drawings are by J. F. Strauss. A few illustrations copied directly from other sources are credited individually. In presenting a book to American beekeepers, the author would express the hope that it may be as helpful to them as the cordial assistance and cooperation of many of them have been to him in his work. E. F. PHILLIPS. WASHINGTON, D.C., March, 1915. CONTENTS CHAPTEK I BEEKEEPING AS AN OCCUPATION Two classes of beekeepers — Extent of beekeeping in the United States and Canada — The relation of apparatus to the development of beekeeping—Who should be a beekeeper ? — Beekeeping for women — Advantages in extensive beekeeping — Where bees may be kept—Results to be expected .......... 1-21 CHAPTER II APPARATUS Relative importance of equipment and skill — Apiary house — Hive stands — Hives and hive parts — Equipment for handling bees — Other equipment . 22-33 CHAPTER III THE COLONY AND ITS ORGANIZATION The point of view —Danger from poor work—Advantage of experience in behavior investigation — Zoological position of the honeybee — Bees not domestic animals — Necessity of colonial life — Size of the colony — Types of individuals in a colony — Queen — Workers — Drones — Brood — Natural nest — Contents of the cells — Arrangement of the nest — Color of the combs — Protection of the nest—Comparison with stingless bees .... 34-53 ix Contents CHAPTER IV THE CYCLE OF THE YEAR PAGES Brood-rearing — Brood-rearing during the season — Temperature of the hive — Swarming — Preparation for swarming— Issuing of the swarm—Stimulus to leave the hive — Behavior of issuing swarin — Clustering — Supposed aids to clustering — Scouts — Entering the new home — Parent colony — Mating flight — After-swarms — Activity of swarms — Swarming conditions induced artificially — Peculiarities of bees in swarming — Causes of swarming — Swarming-out — Gathering of nectar and storing of honey—Collection of other materials — Killing of the drones — End of brood-rearing — Winter cluster — Movements in winter — Responses to outside temperature — Conservation of heat — Source of heat—Effect of accumulation of feces . , 54-92 CHAPTER V THE LIFE OF THE INDIVIDUAL IN RELATION TO THE COLONY Developmental stages — Cellular structure of tissues — Egg — Early embryonic development — Later embryonic development — Segmentation — Fate of parts of the embryo —Larval development—Metamorphosis — Length of developmental stages — Cycle of duties of the adult worker bee — Division of labor — The labor within the hive — Comb building — Feeding of larvae — Composition of larval food — Feeding of queen and drones — Other inside work — The guarding of the colony—The labor outside the hive — Division of labor in gathering—Pollen gathering — Propolis collection — The collection of water — Duration of life — Work determines length of life — Practical applications—Possible determining factors ........ 98-131 CHAPTER VI THE LIFE PROCESSES OF THE INDIVIDUAL General plan of the body of the bee — Head — Thorax — Abdomen — Digestion — Circulation — Metabolism — Respiration — Excretion—Locomotion — Protective apparatus 132-161 Contents xi CHAPTER VII THE NERVOUS SYSTEM AND THE SENSES PAGES Nervous system — Sense organs — Sight — Smell — An-tennal sense organs — Taste — Touch — Hearing — Temperature sense — Finding of the flowers — Finding of the hive — Memory — Nature of bee activities ..... 162-180 CHAPTER VIII THE REPRODUCTIVE PROCESSES AND PARTHENOGENESIS Origin of the eggs — Origin of the male sex cells — Parthenogenesis— Sex determination — Practical applications — Hermaphrodite bees — Eggs which fail to hatch . . 181-191 CHAPTER IX RACES OF BEES Types of social bees — Species of the genus Apis—Varieties of the species mellifica — Egyptian — Syrian — Cyprian — Grecian — Caucasian — Italian — German—Carniolan — African races — Asiatic races — Chinese-Japanese — Best race of bees .......... 192-204 CHAPTER X REGIONAL DIFFERENCES WITHIN THE UNITED STATES Variation in intensity of honey-flows — Variation in the value of plants — Beekeeping regions—General regions — The white clover region — The alfalfa region — The southeastern region — The semi-arid region of the southwest — The sage region — Restricted regions — JJuckwheat — Sumac — Spanish needle — Willowherb — Sweet clover — Blue thistle — Raspberry — Beans — Heartsease — Variation within a region — Distribution of bees in the United States — Migratory beekeeping — Overstocking — Dadant out-apiaries .......... 206-221 xii Contents CHAPTER XI THE FIRST STEPS IN BEEKEEPING PAGES Purchase of colonies — Purchase of bees to be shipped from a distance — Requirements in purchased colonies — How to learn beekeeping—Value of reading — Merits of beekeeping courses — Beginner's outfit .... 222-227 CHAPTER XII THE APIARY SITE Apiary grounds — Exposure to the sun—Care of the apiary grounds — Arrangement of hives — Number of colonies in one apiary—Out-apiaries—Conveniences less essential in out-apiaries ......... 228-233 CHAPTER XIII THE MANIPULATION OF BEES Disturbance to be reduced to a minimum — Equipment for manipulation — When to handle bees — Opening a hive — Remedies for stings — Removing frames — Handling frames — Desirability of straight combs in manipulations — Closing the hive — Occasional manipulations — Feeding — What to feed — Feeders — Uniting—Influence of hive odor — Learning the new location — Transferring — The best time to transfer colonies — Methods — Transferring from walls of houses — Transferring from hollow trees — Preventing robbing in the apiary — Moving bees — Elimination of non-essential manipulations — Two essentials — Increase in efficiency through system ....... 234-254 CHAPTER XIV SPRING MANAGEMENT Object of spring, manipulations — Prevention of drifting — Spring protection—First examinations—Cleaning the hives — Equalizing the colonies—Clipping queens — Summary of favorable spring conditions— Questionable manipulations — Stimulative feeding — Spreading the brood — Substitutes for pollen ........ 256-264 Contents Xlll CHAPTER XV SWARM CONTROL AND INCREASE Loss from division of the working force — Variation in swarming — Variation in colonies in respect to swarming preparations — Direction of the beekeeper's efforts — Preventive measures — Breeding — Mechanical devices—Preventive manipulations — Miller's methods—Remedial measures — Control of natural swarms — Automatic hivers — Location for the swarm — Disposition of the brood after swarming — What to use in the brood chamber in hiving swarms — Remedial manipulations — Unbalanced condition of swarming colonies — Break in the emergence of brood — Requeening combined with dequeening — Removal of brood — Mechanical appliances — Increase ..... CHAPTER XVI THE PRODUCTION OF EXTRACTED-HONEY Increase in the production of extracted-honey — Advantages of extracted-honey—Disadvantages of extracted-honey — Extracted-honey hives — Choice of storage combs—Use of extracting combs smaller than brood combs — Number of supers — Manipulation of supers — Need of abundance of drawn combs — System in producing extracted-honey— Removing honey from the bees — House for extracting — Portable extracting outfits — Uncapping — Cans for cappings — Capping melters — Types of extractors — Extracting — Straining the honey — Storage tanks — Reduction of the lifting of honey — Returning combs to the hives . CHAPTER XVII THE PRODUCTION OF COMB-HONEY Purity of comb-honey—The " Wiley lie" — Decrease in comb-honey production — Demand for fancy comb-honey — Advantages of comb-honey—Disadvantages of comb-honey — Restrictions in comb-honey production — Honey-house— The best Jiive for comb-honey — The early development of the section —Types of sections — Types of supers — Other equipment — Preparation ot the sections — Manip- 266-286 286-306 XIV Contents ulation of the bees — Keeping bees in proper condition — Manipulation of supers — Removal of supers — Caring for the crop — Preparation of bait sections — Storage in supers — Bulk comb-honey — Bulk comb-honey for home use — Cut comb-honey for market . . . , . 301-319 CHAPTER XVIII MARKETING THE HONEY CROP Preparation of extracted-honey for market — Wholesale packages — Retail packages for local markets — High-class retail packages — Blending — Argument for blending — Heating honey — Preparation of comb-honey for market — Cleaning the sections of propolis — Grading—Shipping cases—Glazed sections — Use of cartons — Shipping comb-honey — Preparation of bulk comb-honey for market — Preparation of granulated honey for market — Wording of labels — Development of the home market — Co-operative selling 320-333 CHAPTER XIX THE PRODUCTION AND CARE OF BEESWAX Rendering the .wax — Wax presses — The residue — Removing wax by dissolving — Cleaning wax — Granulation of wax — Bleaching wax —- Adulteration of wax — Preparation of wax for market—Special production of wax—Uses of beeswax .......... 334-342 CHAPTER XX THE CARE OF BEES IN WINTER Losses in winter — Object of winter protection — Requirements of successful wintering—Winter stores — Cause and effects of humidity in the hive — Effects of ventilation — Source of heat and effects of changes of temperature — Disturbance—Methods of wintering bees — Outdoor wintering — Cellar wintering—Effects of confinement — Spring dwindling ........... 843-368 Contents xv CHAPTER XXI THE SOURCES OF NECTAR AND POLLEN Reasons for knowledge of nectar sources — Difficulties of identification — Study of neighboring locations — Function of nectar—Variations in nectar — Variations in secretion —Effects of climatic conditions on secretion—Advantages of swamp sources — Cultivation of plants for nectar — Value of the minor sources — Gathering of pollen — Value of bees in cross-pollination — Damaging effects of incorrect spraying— Bees do not puncture ripe fruit — Supposedly poisonous honeys — Plant honey-dews — Insect honey-dew — Annotated list of honey-plants ...... 359-396 CHAPTER XXII BEE DISEASES AND ENEMIES Brood diseases — American foul brood — European foul brood — Sacbrood or pickled brood — Methods of spread — Treatment — Shaking treatment — Fall treatment — Additional treatment for European foul brood — Diseases of adult bees — Dysentery — Nosema disease — Paralysis — Spring dwindling — Enemies of bees — The wax-moth (Gal-leria mellonella) — The lesser wax-moth (Achroia grissella) — Remedies—Other enemies ...... 397-416 CHAPTER XXIII THE REARING OF QUEENS Commercial queen-rearing — Systematic requeening — Conditions under which queens are reared — Saving natural queen cells — Having natural cells built—Miller method — Alley method — Hopkins method — Queen cells on artificial bases — Transferring larvae — Swarm box — Having cells built out — Nursery cages — Mating hives — Classification of queens — Mailing cages — Introducing queens — Improvement of stock — Study of breeding needed — Selection of drones — Desirability of pure races — Danger from inbreeding ....... 417-429 xvi Contents CHAPTER XXIV MISCELLANEOUS INFORMATION PAGES Literature on bees and beekeeping — Organizations of beekeepers—Laws — Supplies for beekeepers — The uses of honey—Honey crop reports—Educational work in beekeeping—The Bureau of Entomology . .... 430-487 APPENDIX Explanation of Symbols used in Anatomical Illustrations 439-448 ILLUSTRATIONS NOTE. —Illustrations marked by asterisk are from Farmers1 Bulletins Nos. 442 and 447 of the U. S. Department of Agriculture. Other illustrations are credited by authors individually. Apiary of the Bureau of Entomology, Drummond, Md. . Frontispiece PIG. PAGE 1. A primitive box-hive ......... 1 2. A bee and apple blossoms ........ 5 3. Mud hives in Palestine. (From photograph by Metcalf) . . 5 4. Group of Caucasian hives . . . . . . . 6 5. An old Greek hive. (From Wheler, 1682) .... 6 6. Gravenhorst hive, a combination of skep and frame-hive . . 7 7. German hive, opening at the rear ..... 7 8. Bee-house in Carniola, Austria ....... 8 9. Carniolan hive ......... 9 10. Bee-house mentioned in uThe Hoosier School Boy'7 . . 9 11. W. B. C. hive of England ........ 10 12. C. B. B. hive of Ireland ........ 11 13. Frame of C. D. B. hive ........ 11 14. A woman beekeeper ......... 16 15. Roof apiary in lower New York City. (Drawn from photograph by Root) .......... 18 16. Apiary on shed roof, to'economize space . . ^ . . . 19 17. Tropical apiary, San Sabastian, Porto Rico . . . .19 18. Honey-house door . . . . . . . t . .24 19. Porter bee escape *......... 24 20. Ten-frame Langstroth hive with queen excluder, comb-honey super and telescope cover * .' . . . . . .26 21. Diagram showing spacing of frame and rabbet in Langstroth hive 27 22. Spacing of Hoffman frames ....... 28 23. Spur wire-imbedder* . ........ 29 24. Smoker* ........... 29 25. German beekeeper's pipe . ...... 30 26. Cotton netting veil with silk tulle front * .... .31 27. Hive tools* .......... 31 28. German bee brush* ......... 31 xviii Illustrations FIG. PAGB 29. Tool-box seat .......... 32 30. Alley queen and drone trap * . . ... . . .32 31. Bee escape board *....*.... 32 32. Comb-foundation cutter ........ 33 33. Van Deusen hive clamp . . . . . . . . 33 34. The honeybee: worker, queen and drone * .... 40 35. The honeybee : egg, young larva, old larva and pupa * . . 40 36. Structure of comb *......... 47 37. Piece of new comb showing transition cells * .... 48 38. Queen cell ........... 49 39. Cappings of brood . . . . . . . .50 40. Capping of honey ....... Facing page 51 41. Colony in the open air . . . . . Facing page 61 42. Eggs in cells of the comb ...... Facing page 55 43. Larvae in cells of the comb, almost full grown . . Facing page 55 44. Concentric arrangement of the brood...... 57 45. Group of queen cells * ........ 62 46. A swarm cluster ........ Facing page 65 47. Swarm catcher ........ Facing page 67 48. Capturing a swarm ....... Facing page 65 49. Swarm entering a hive ...... Facing page 68 50. Group of tissue cells from skin of young salamander. (From Nelson) .......... 95 51. Three stages in the development of the embryo. (From Nelson) 97 62. Diagram of a longitudinal median section of a bee larva. (From Nelson) .......... 100 63. Ventral,plates of the abdomen of a worker bee. (From Casteel) 108' 64. Inner surface of the left hind leg of a worker bee showing a wax- • scale. (From Casteel) ....... 108 56. Ventral view of worker removing wax-scale. (From Casteel) . 109 66. Side view of worker removing wax-scale. (From Casteel ) . 109 57. Ventral view of worker passing wax-scale forward. (From Casteel) .......... 110 58. Side view of worker passing wax-scale forward. (From Casteel). 110 69. Median longitudinal section of head of worker, showing the glands. (From Snodgrass). . . . . . .111 60. Alimentary canal of worker, showing glands. (From Snodgrass) ........... 112 61. Longitudinal median section of base of oesophagus. (From Snodgrass) .......... 113 Illustrations xix 62. Lizard encased in propolis . . . . . . . .117 63. Outer surface of the left hind leg of a worker. (From Casteel) 124 64. Flying bee, showing movements of legs in pollen collecting. (From Casteel) ......'... 124 65. Flying bee patting pollen on the pollen basket. (From Casteel) ......... .124 66. Inner surface of the left hind leg of worker. (From Casteel) . 125 67. Flying bee loading the pollen baskets. (From Casteel) . . 125 68. Front and back views of head of worker bee. (From Snod- grass) ........... 134 69. Anterior views of head of worker, queen and drone. (From Snodgrass) .......... 135 70. Right mandibles of worker and drone. (From Snodgrass) , 136 71. Internal mandibular gland of worker. (From Snodgrass) . 136 72. Mouth parts of the worker. (From Snodgrass) . . . 137 73. Dorsal view of ventral wall and internal skeleton of worker. (From Snodgrass) ........ 139 74. Thorax of worker. (From Snodgrass) ..... 140 75. Lateral view of abdomen of worker. (From Snodgrass) . . 140 76. Tip of abdomen of worker with left side removed. (From Snodgrass) .......... 141 77. Histological details of alimentary canal. (From Snodgrass) . 145 78. Longitudinal median vertical section of body of worker. (From Snodgrass) .......... 149 79. Tracheal system of worker. (From Snodgrass) . . . 152 80. Fore and hind wings. (From Snodgrass) ..... 155 81. Legs of worker, queen and drone. (From Snodgrass) . . 156 82. Claws. (From Snodgrass) ....... 167 83. Ventral view of sting of worker and accessory parts. (From Snodgrass) .......... 159 84. Nervous system of worker. (From Snodgrass) . . . .163 85. Brain and suboesophageal ganglion of worker. (From Snod- grass) ........... 164 86. Section of compound eye and optic lobe ..... 166 87. Section of entire ommatidium ....... 167 88. Location of groups of olfactory pores — dorsal view. (From Mclndoo) .......... 170 89. Location of groups of olfactory pores — ventral view. (From Mclndoo) ...... ... 171 90. Cross-section of typical olfactory pore. (From Mclndoo) . . 172 xx Illustrations 91. Antennal organs. (Copied from Mclndoo, after Schenk) . 173 92. Reproductive organs, sting and poison glands of queen. (From Snodgrass) .......... 182 93. Reproductive organs of drone. (From Snodgrass) . . . 185 94. Propolis at entrance, built by Caucasian bees . . . . 197 95. Map of Florida showing distribution of honey plants. (From Baldwin) .......... 210 96. Map of United States, showing distribution of colonies of bees (Redrawn from map furnished by Bureau of Crop Estimates, U. S. D. A.) . . . . . . . . 214 97. Map showing distribution of Dadant apiaries. (Redrawn from Dadant) .......... 220 98. Apiary in the West, shaded by thatched shed. (Redrawn from Root) .......... 230 99. Former apiary of the Bureau of Entomology, College Park, Md.* 231 100. Hive-body resting on cover . . . . . . 236 101. Handling a frame, first position* ...... 238 102. Handling a frame, second position* . ..... 238 103. Handling a frame, third position* ...... 238 104. Hive leveling device . . . ..... 239 105. Division board (Doolittle) feeder *...... 242 106. Alexander feeder in collar under hive-body * ... . 242 107. " Pepper-box" feeder* ........ 242 108. Pan in super arranged for feeding * ..... 242 109. Cutting combs from a box-hive . / . . . . . 246 110. Hive ready for moving ........ 250 111. Manipulation to reduce population of parent colony — first position. (From* Demuth) ....... 275 112. Manipulation to reduce population of parent colony — second position. (From Demuth) . . . . . . . 275 113. Manipulation to reduce population of parent colony — third position. (From Demuth) ....... 276 114. Manipulation to reduce population of parent colony — fourth position. (From Demuth) ....... 276 115. Queen excluder (uhoney board") ...... 277 116. Uncapping knives* . . . . . . . .293 117. Steam-heated uncapping knife ...... 294 118. Comb for uncapping — used in Europe ..... 294 119. Capping melter ......... 295 120. Tank to receive cappings ....... 296 Illustrations xxi Extractor with stationary can * ..... 296 Power extracting outfit. (Photo by Root) . Facing page 297 Honey strainer. (Redrawn from Root) ..... 298 Honey storage tanks. (Drawn from photograph by Root) . 299 Diagram to show method of spacing bee-way sections. (From . Demuth) .......... 308 126. Diagram to show method of spacing plain sections. (From Demuth) .......... 308 127. Comparison of plain and bee-way sections. (From Demuth) . 308 128. Comparison of tall and square sections of equal capacity. (From Demuth) .......... 309 129. T-super. (From Demuth) ....... 309 130. Super for square bee-way sections with section holders. (From Demuth) .......... 309 131. Super for square plain sections with section holders. (From Demuth) ... .^ ...... 310 182. Super for tall plain sections. (From Demuth) . . . 310 133. Super for tall plain sections in wide frames. (From Demuth) 310 134. Old type of wide frame for holding sections. (Drawn from Miller) ........... 311 135. Section folder .......... 312 136. Diagram showing the arrangement of the supers. (From Demuth) .......... 315 137. Crate holding two 5-gallon honey cans ..... 321 138. Shipping cases for comb-honey *...... 328 139. Double boiler for melting combs ...... 335 140. Hot water (Hershiser) wax press ...... 336 141. Diagram showing the response of a colony of bees to changes in outer temperature. (From Phillips and Demuth) . . 348 * 142. An apiary in winter ......... 350 143. Roof of a bee-cellar away from a house . . . . . 353 144. Arrangement of hives in a cellar. (Drawn from Alexander) 354 145. Diagram showing the effects of an accumulation of feces. (From Phillips and Demuth) ........ 357 146. Alfalfa ........... 373 147. Basswood ........... 375 148. Buckwheat .......... 377 149. Spider flower (Cleome) . . . ... . . .378 150. Cotton ........... 379 151. Dandelion .......... 380 xxii Illustrations F16. PA expected," as beekeepers so often express it. For this reason, it is necessary for the beekeeper to know the behavior of bees in all its phases and in so far as they have been determined, which is not far, the causes of their various activities. Obviously, the successful beekeeper is a naturalist and such persons are born, not made successfully. Patience, 14 Beekeeping power of concentration and sympathetic understanding of the bees are essentials and, as a result, the bees become pets rather than beasts of burden to the true bee-crank. Persons who fail to appreciate bees from this point of view will probably find it more pleasant and profitable to let them alone. Like all general statements about bees, there are exceptions to this one. Some who are financially successful beekeepers are totally devoid of sympathetic interest in bees and have learned to handle bees as it were by force. Such men are out of place as amateur beekeepers and indeed fail to reach the highest success as professionals. The ardent bee-man finds pleasure in comparing experiences and observations with his co-workers, in conventions and out, and some of the best "conventions" are those in which two or three experienced beekeepers spend half or more of the night in talking over their latest ideas. They discuss new and supposedly improved apparatus and all the latest systems of manipulation, for there seem to be styles and fads in beekeeping as in clothes. The man who fails to find pleasure in such an interchange of views will find himself out of place among bee-enthusiasts. Not only is a knowledge of what to do necessary to success with bees, but it is equally necessary that the right thing be done at the right time. To put on comb-honey supers too late, to delay the necessary steps in swarm control or to neglect the preparation of bees for winter, all mean loss in bees, honey and money. In the make-up of the beekeeper must be promptness to do the things which his experience teaches. In the hands of the wise, the bees need remarkably little attention. They should not be manipulated *daily and the hive is better unopened unless some change is called for. The beginner errs almost universally in over-manipulation. It must not be forgotten, however, that the reduction in handling which comes with experience is not neglect, and the beekeeper must know daily whether the condition of the nectar-secreting plants or of his colonies calls for any manipulation. This requires experience and Beekeeping as an Occupation 15 observation and Finally promptness in doing what is necessary. Instead then of being an occupation fitted for everyone, beekeeping is well fitted only to the minority. The array of human excellences here enumerated are not all necessarily present in perfection, but the nearer the approach of these qualities to that happy state, the more satisfactory will beekeeping be found as a vocation or avocation. It is to be hoped that these formidable requirements will not deter the potential bee-crank from making a beginning. * Beekeeping for women. A question much discussed in books and journals on bees is that of beekeeping for women. Many women can and do handle bees (Fig. 14) with marked success. In those parts of the business which require delicacy of touch and minute attention, such as queen-rearing, women often surpass men in proficiency. As amateur beekeepers they are at home. The question which usually presents itself, however, is whether beekeeping is suitable for women as a means of earning a livelihood and repeatedly has the writer been asked for advice on this subject. Professional beekeeping on a scale sufficiently large to supply an adequate income requires long hours of work in the hot sun, heavy lifting and unremitting physical endurance, On a small scale these FIG. 14. — A woman beekeeper. 16 Beekeeping obstacles may be overcome, but in a commercial apiary the work must be done promptly, for delay means loss. While some women have found pleasure and profit in commercial beekeeping, it emphatically cannot be recommended for the majority of women, and this should be made clear to avoid disappointment for those who may be attracted to it. Of course, this applies only to those women who have no man in the company to do the heavy work. Many a professional beekeeper has received assistance of incalculable value from the women of the family. It should be made clear that the obstacles to the commercial success of women beekeepers are physical ones only. Advantages in extensive beekeeping. Several references have been made to the desirability of encouraging professional beekeeping, and this should be explained to avoid misunderstanding. Everyone who desires to keep bees, of course, has that privilege, so long as by so doing he does not interfere with the rights of others. • By common consent, a man's bees are not considered as trespassing when they go outside his land for forage and consequently a beekeeper cannot legally or morally claim the exclusive right to keep bees in a locality. The beginner, therefore, is not considered as overstepping his rights in getting bees. Taking a broader view of the subject, however, the professional beekeeper by his knowledge of the subject is able to produce larger crops, thereby utilizing the available nectar more economically. By this same knowledge and his better equipment, he is able to produce a better quality of honey. It is therefore evident that from the standpoint of conserving a resource to the best advaiftage there is reason to encourage the extensive beekeeper. In case a brood disease breaks out in a community, then there is every reason for taking sides with the professional beekeeper. The man with a few colonies is not financially interested to an extent which will compel him to care for the disease and in disease control it is usually necessary Beekeeping as an Occupation 17 that there be some incentive to compel action, the financial incentive being most efficient. The small beekeeper usually becomes a menace to the industry in such an outbreak and not until most of these men lose all they have is much progress made against disease. The most economical development of the larger honey markets for the beekeepers of any region can come only through co-operation in buying necessary supplies and in selling their products. So long as there are so many thousands of beekeepers with small financial interest in the industry, such co-operation is rendered virtually impossible and the industry is thereby retarded. In some of the western^ states, beekeeping is carried on chiefly by extensive beekeepers and they have found co-operation practical and profitable, while the beekeepers of the east still fight their battles individually, co-operation being made practically impossible because of the thousands of beekeepers who could not be reached by such a co-operative movement. Similarly, it is difficult to bring about concerted effort in having desirable laws passed for the protection of the industry or in instituting any agency for the advancement of the industry unless there are a number of men whose financial interest is sufficient to induce them to spend time and money in working for the things they need as beekeepers. Beekeepers are very human people, and "money talks" in this business as well as in other lines of human endeavor. There is therefore adequate reason in the view that the development of beekeeping to its true place in American agriculture depends on the making of a large number of professional beekeepers and this in turn implies the elimination of the beekeeper with a few colonies, little interest and still less of willingness to work for the industry. While the number of professional beekeepers is increasing in a way to give satisfaction to those interested in the best development of the industry, a word of caution may not be amiss. Some beekeepers feel that as professionals they must engage in no other business, whereas for certain months 18 Beekeeping they are not occupied for more than a small fraction of time. Without entering into a moral discussion on the virtues of industry or the various things that Satan is said to find for idle hands to do, it is obvious that the professional beekeeper may use other occupations to add to his income just as the amateur beekeeper uses his bees. As the beekeeper becomes more proficient he eliminates all unnecessary manipulation so that the care of a goodly number of colonies may take a relatively short time. When the crop is off and sold he has little to engage his attention until the next season, especially if his bees are wintered out of doors. Where bees may be kept. It has been the pleasure of the writer to visit apiaries on the roofs of city buildings (Fig. 15) and in the almost desert valleys of California (Fig. 166), in city backyards (Fig. 16) and in the mountain wilds, in small towns, on farms, in Canada and in the tropics (Fig. 17). In diversity of location these apiaries are as varied FIQ. 15. —Roof apiary in lower New York City. as their Owners. While recruits to the ranks of beekeepers may be found in all ages and conditions of men, so bees may be kept in places which would at first appear utterly unproductive, as well as in places which are obviously abundant in their nectar supply. The uninformed observer may fail utterly in his estimate of the value of a location from the standpoint of the bee. Most of the valuable nectar-secreting plants do not have Beekeeping as an Occupation 19 large highly colored flowers, and the cultivated varieties of the flower garden are of insignificant value. Bees fly for two or three miles for forage and may go even farther in emergency. In choosing a location, it is therefore necessary that FIG. 16. — Apiary on shed roof, to economize space. in the range of flight there be an adequate supply of nectar-producing plants. The ideal location is obviously one in _____________. which the nectar supply is near so that it may be obtained without the loss of energy inci-dent to long flights. Results to be expected. The stori]es sometimes told of the crops that have been obtained from single colonies or of the rapidity with which the number of colonies may be increased are apt to mislead the beginner. While several hundred pounds of honey may at times be obtained from a single colony in a season, this is by no. means usual. In apiaries managed for comb-honey production, it is perhaps fair to estimate the average FIG. 17. — Tropical apiary, San Sabastian, Porto Rico. 20 Beekeeping annual crop at 25 to 30 sections. For extracted-honey, larger averages may be expected, perhaps of 40 to 60 pounds. The financial returns depend entirely on the market and the method of selling the honey. If sold by the beekeeper direct to the consumer, a pound of extracted-honey brings from 10 to 20 cents, while a section of comb-honey sells at 15 to 25 cents. If sold to dealers, the return is less but there is less liability of financial loss and less time consumed in selling. Naturally these estimates must be dependent on the quality of the product and on the neatness of the final package. In addition to the labor there will be other expenses for supplies such as comb-foundation, sections and occasional new hives and fixtures, not counting the apparatus used in increasing the apiary. These may cost from 50 cents to $1.00 for each colony in a season. Estimates such as these arte really of little value since the returns differ so greatly according to the kind of honey obtained and the facilities for marketing. For example, the white clover honey of the North brings a higher wholesale price than the amber honeys which come from most regions of the South but, on the other hand, the southern beekeeper enjoys a longer nectar-secreting season and usually obtains larger crops from each colony. Another factor which must not be overlooked is the beekeeper. Anyone may reap a heavy harvest in the season when nectar is abundant but in the lean years, which come more often thaix desired, only the good beekeeper makes the most of the nectar at hand. And then come years of practically total dearth of nectar, when feeding is necessary to keep the colonies alive. Taking all these factors into consideration, it may be justly concluded that a successful beekeeper is usually well repaid for the time he spends in his work, if he considers the return in the sense of wage. He may also consider that he has received the interest on his original relatively small investment. He usually averages little morp than this, however, so that beekeeping is in no sense a "get-rich- Beekeeping as an Occupation 21 quick" business. Its advantage as a recreation over most other occupations of a similar character is that it is a means of occupying time not otherwise engaged to a financial profit and the returns therefore often add that part to the income which brings comforts and pleasures. Beekeeping yields a quick return on the investment, for frequently in a good year a colony will pay for itself. In fact there are few branches of agriculture which on so "small an investment will yield as great a return. It may at least be said for the person who decides to try out beekeeping that he does not stand to lose much. This chance calls to mind a conversation with a western friend. In recounting the present advantages and past glory of his beautiful city, he recalled the former gambling days when everything was "open." After a vivid description of those halcyon days and of some of the men of that time, he said, "I knew some of those men well. They were personal friends of mine and they saw nothing wrong in gambling. And I can appreciate their point of view — for I'm a beekeeper myself." In discussing the financial results, it is far from wise to overlook the other benefits. Beekeeping, to an enthusiast, means out of doors and intimacy with these interesting insects which have been studied for centuries and still remain an unsolved riddle in many of their activities. It may mean health to the person confined to an office. It means to a congenial spirit association with bee-enthusiasts, than whom no more optimistic and warm-hearted people exist. If these things make an appeal, then may apiculture be classed as yielding the greatest profits that can be conceived. If now we attempt to decide for the questioning prospective beekeeper whether he should take up bees, from the previous discussion the whole question is solved: if he will like beekeeping, he should take it up; if not, he would better never have considered it. And this is about as reliable and lucid a prophecy as is usually possible. CHAPTER II IP APPARATUS BEFORE discussing the phenomena observed in the activities of bees, on which the practical manipulations rest, it is desirable that some description be given of the^ hives and equipment used in beekeeping, since frequent* references are made to these things in the chapters dealing with behavior as well as in those concerning the practical work of the apiary. Since this subject is to be introduced early, it seems best to complete the discussion here, except for certain pieces of apparatus used in special manipulations. Relative importance of equipment and skill. It is important that the relation of the equipment of the apiary to the needs of the bees be understood. A hive is not only a home for the bees but it is, especially, a tool for the beekeeper and, being only a tool, it is of far less importance in apiary management than the skill and experience of the beekeeper. By many beekeepers, especially among beginners, the apparatus of beekeeping is given undue importance and the interest aroused by the work of putting together the carefully manufactured supplies is really quite excusable. In the American literature on beekeeping the description of apparatus plays too prominent a part. Tools alone do not make the mechanic. It is therefore proposed here to give only a brief description of the general equipment of beekeeping, leaving for the chapters on special phases of beekeeping, the description of the apparatus used in these manipulations. For greater detail, the reader is referred 22 Apparatus 23 to catalogues of supplies which manufacturers are quite willing to furnish. It would be interesting to trace the evolution of the various implements used in beekeeping, but this is beyond the scope of this book. For certain appliances, discussed in later chapters, such a method of treatment has seemed desirable and, in fact, to discuss all of the present apparatus in that manner would make the reasons for their construction clearer. There should some day be prepared a book on the evolution of hives and the beekeeper's equipment, if for no other purpose than to show the ardent inventor, who is usually a beginner, the steps that have already been taken and passed by and to prevent the repeated re-discovery of abandoned apparatus. In recent times, the industry is relatively free from the exploitation of worthless apparatus but, at about the time of the invention of the Langstroth hive, the beekeeping industry was well-nigh buried in bizarre hives. The industry has not ceased to advance, but beekeepers have outgrown, the belief "that success depends on tools. The recognized essentials of beekeeping are knowledge of the bees, skill in manipulation and simplicity in apparatus. The supplies of the beekeeper have few prerequisites. They must be simple in construction, strongly built and, above all, interchangeable throughout. The manufacturers of beekeeping supplies in the United States have done much to simplify the equipment. The best materials are usually employed. * Apiary house. In the main or home apiary, it is desirable to have a workshop, usually known by beekeepers as the " honey-house," where supplies may be prepared and the crop cared for and perhaps stored for a time. This house should be below the bees if the ground slopes (p. 292). It is perhaps needless to give plans for an apiary house since the experienced beekeeper will easily construct one that fits his individual needs 24 Beekeeping FIG. 18. — Honey-house door. The and the beginner will use what he has at hand. One suggestion is perhaps not amiss, if one may judge from the honey-houses usually seen. The house should be large enough to permit the storage of the surplus fixtures out of season and of the crop until it is shipped. Beekeepers frequently fail to provide adequate space for these uses. Windows and doorsl should be thoroughly screened to prevent the entrance of bees. The door should swing freely both ways (Fig. 18) so that the beekeeper may pass through with his arms full. The window screens are best made by tacking wire-cloth to the outside of the window casings, allowing it to extend about six inches above the opening. The upper border should be held out one-quarter of an inch by narrow wooden strips to provide abundant exits for bees which accidentally get into the house. Bees rarely enter such openings and those which fly to the screens from the inside immediately crawl upward and go out, promptly clearing the room of bees. Bee-escapes (Fig. 19) may be used at the corners of ordinary framed window screens but 1 A. C. Miller has recently called attention to the desirability of a solid door to the apiary house, so that bees will not be attracted to this opening by the odor of honey. The suggestion is good and the desirability of having such a door swing both ways still exists. * FIG. 19. — Porter bee-escape. Apparatus 25 these are less effective. The best arrangement of windows is to have the sash slide horizontally on runners so that the openings may be entirely free from glass. By this arrangement, bees are not imprisoned on single window panes and in hot weather the beekeeper appreciates all the breeze that may be allowed to enter the house. Benches, cupboards and racks for small supplies and tools can be arranged to suit individual needs, but these too should be large and roomy. It is a good plan to provide racks for surplus combs, the frames being hung in strips of wood properly spaced. The kind of honey produced determines the other features of the house. For comb-honey production, a well-supported second story is recommended for the storage of honey. In extracting, it is desirable that the extractor, uncapping boxes and tanks be so arranged that it is not necessary to lift heavy supers and cans and so that at no time the honey must be lifted by hand. Honey is best stored in a warm place and a second story or attic is ideal also for extracted-honey. By the use of a honey-pump, the honey can be raised to a high level and it can then be moved by gravity in future bottling or packing. While general advice on the construction and arrangement of honey-houses is difficult to give, it will profit the beekeeper carefully to study his needs in drawing his plans, so that labor will be reduced. For the out-apiary, a smaller house will serve and many beekeepers do not have any house in such yards. The portable extracting outfit is one solution, and for comb-honey production it is as easy to haul home in the supers as in shipping cases. For extracted-honey production, a small extracting house is usually preferable. If bees are wintered in a cellar (p. 353), this may be built under the apiary house. It is desirable to provide a cook stove, which is a comfort in chilly weather and is serviceable in wax-extraction. Running water in the honey-house will be found a great convenience. 26 Beekeeping Hive stands. The arrangement of the hives will determine the character of the stand. A wooden frame, bricks, tile (Fig. 20), concrete blocks or flat stones are equally good to raise the bottom board of the hive above the ground so that it will not rot. It is sufficient to raise it only a few inches to allow air to circulate freely under the bottom. In a permanent apiary, it is convenient to arrange the hive stands in the desired order and to number them by the system used in numbering the colonies for purposes of record. Hives and hive parts. The hive which opens at the top and in which the combs are built in freely movable frames is the one generally used in America/ 'It was invented by Rev. L. L. Langstroth, the Father of American beekeeping, in 1851. From this date, the development of modern beekeeping begins. The original Langstroth hive has been somewhat modified as the result of the experience of later years, but as now used (Fig. 20) it consists of a plain wooden box holding frames hung from a rabbet at the top (Fig. 21) and which do not touch the sides, top or bottom. The box is usually dovetailed and is commonly made of white pine dressed to f inch. The greatest advance of the Langstroth hive is not so much in the movable frames as in the free space (Fig. 21) all about them. The size of this space is of the greatest importance, it being such that bees pass through it freely but do not build wax nor deposit propolis in it» The manufacturers of beekeepers' supplies make this space a quarter of an inch. FIG. 20. — Ten-frame Langstroth hive with queen-excluder, comb-honey super and telescope cover. Apparatus 27 The plain box rests on a bottom board, so made that there is an entrance space (Fig. 20), and over the hive is a cover which can be entirely removed to permit the removal of frames. There are various types of bottoms and covers, with no marked advantages in one over the others. The telescope cover over a thin inner cover is a good type (Fig. 20). The size of frame standard in America is that of the Lang-stroth (or L) hive, 9| high by 17f inches long. Frames of other sizes, but having the same method of hanging, have been devised and a larger size has much to commend it, but the desirability of uniformity outweighs the advantages of the odd sizes. The number of frames in the hive is determined by the character of the locality and the kind of honey produced. Many comb-honey producers in the white clover region prefer the eight-frame hive FIG. 21. —Diagram while the majority of extracted-honey off^e anTrlb^ producers use the ten-frame size. Some bet in Langstroth prefer a twelve-frame hive. The sales of hive-supply dealers indicate a growing preference for the ten-frame size among all classes of beekeepers. In deciding which size of hive is preferable, the usual method is to determine the amount of brood that can be reared by a strong colony and to calculate the requisite number of combs from their area. This is not an entirely reliable criterion for the following reasons: (1) the outside combs are frequently unavailable for brood-rearing, because of inaccurate spacing, (2) the top rows of cells in combs built on comb-foundation usually sag, reducing the area available for brood by a depth of one to two inches, (3) there is frequently considerable drone comb or irregular comb. The comb area needed for brood depends on the character and time of the honey-flow and on the system followed. For example, if the main honey-flow conies early in the season (e.g. white clover in the North), it is desirable to build up the colony with great rapidity. This 28 Beekeeping may be done by stimulating breeding, and since more space is then needed it can be supplied by giving two hive-bodies for the brood. Later, when br6od is less to be desired, the breeding space may be reduced. Another type of frame is sometimes used and should perhaps be mentioned, although its use is decreasing. These frames have end-bars wide enough so that they touch each other and the bees cannot pass around the ends of frames. The chief advantage stated is greater warmth in winter. Some frames of this type are suspended from the top, others from the middle of the end-bar and some are supported from below. Frames of any description must be spaced so as to give room between the combs to allow brood to be reared in the cells and also to provide space enough for the bees between the combs. The spacing usually adopted is If inches from center to center but some beekeepers prefer 1| FlG 22 _Spacing inches.1 The closed-end frames when of Hoffman frames, brought together are properly spaced. While the larger number of beekeepers do not use the closed-end type, various devices are in use for the spacing of open-end frames. The frame most conynonly used has the end-bars wide enough for a short distance so that they touch at the top (Hoffman frames, Fig. 22). The metal-spaced frame is possibly an improvement. Some honey-producers object to spacing devices because they interfere in uncapping, and this objection is largely overcome by the use of staples in the side of the end-bar. To obtain regular cells in the comb, comb-foundation, a thin sheet of pure beeswax embossed to correspond with the bases of cells, is placed in the frames. On this as a guide, the bees build the side walls of the cells, utilizing to some extent the extra wax in the foundation. Foundation is made in various thicknesses, the thinnest being used for comb-honey, and in both worker and drone cell size. 1 The English frames are !•£$ inches from center to center. Apparatus *29 FIG. 23. — Spur wire-imbedder. To strengthen the combs, it is customary to wire the frames with fine (No. 30 gauge, tinned) wire. The wires are generally stretched horizontally, and most frames as they come from the manufacturer are pierced for wiring. After the wires are stretched tight, the foundation is fastened to the top of the frame and the wire is imbedded in the foundation, usually by pressure. The spur imbedder (Fig. 23) is generally used but is not especially good. Heat generated by a weak electric current is sometimes used, but perhaps the best method is to run along the wire a small warm soldering iron with a notch in the point. Whatever style of hive is adopted, the parts must be accurately cut so that the bee-spaces are of the right size and so that the apiary equipment may be interchangeable throughout. Hives or frames of different sizes or of improper dimensions are perhaps the worst inconveniences that can be found in an apiary. The materials used should be the best, for the equipment is often used for many years. As a rule, it is better to buy hives and frames and, in fact, practically all the necessary supplies from the regular manufacturers of such articles. This advice is not given as an advertisement for the manufacturer but is based on the recollection of ill-spaced, inaccurately cut, home-made outfits which have been encountered in traveling among beekeepers. Obviously, an expert wood-worker can do as well as the regular manufacturer, but even then the cost of home-made supplies usually exceeds the price charged by the dealers, when one considers the time consumed. The outside of hives should be painted to protect them FIG. 24. — Smoker. Beekeeping from the weather. It is most important that the joint or dovetail be painted as decay starts there in unpainted hives. White paint (white lead and raw linseed oil) is to be preferred as it makes a cooler hive than dark colors. For the sake of the appearance of the apiary, all hives should be of the same color. This is also important if one wishes to interchange hives in the apiary. • The hive, as, it has been discussed so far, is essentially the home of the bees and is occupied by them throughout the year. This portion is usually known as the* brood-chamber. For surplus honey, on which the beekeeper depends for his profit, additional parts are needed and these are discussed in connection with the production of the various kinds of honey. Equipment for handling bees. A few special tools are necessary in handling bees. A good smoker (Fig. 24), consisting of a tin or copper receptacle in which to burn rotten wood or other materials, with a bellows attached to force a draft, is indispensable. The medium-sized smokers are best for the beginner and the professional beekeeper may learn by experience what size is best suited to his needs. The German beekeeper often uses a specially constructed pipe (Fig. 25), which is naturally a dual-purpose tool. A veil of black material, preferably of cotton netting with a silk tulle front (Fig. 26), is needed to protect the face from stings. Even a seasoned beekeeper, who sometimes likes to brag that he never uses a veil, may find it convenient to have a veil thrown back on his hat, which can FIG. 25. — German beekeeper's pipe. Apparatus 31 FIG. 26. — Cotton netting veil with silk tulle front. be brought down when the bees become annoying. Black wire-cloth veils are often used and, while they are a better protection than the cloth veils, they are less convenient as they cannot so easily be thrown back. A steel tool of some kind is needed to pry up covers and to loosen and separate frames. A screwdriver will answer but some specially devised tools (Fig. 27) may be found preferable. Gloves of cloth or leather are sometimes used to protect the hands. The handling of frames is less impeded if the finger ends are cut out. Gloves are hot, usually sticky or stiff, and are as a rule abandoned after the early stages of bee- k^^^^rn keeping are passed. 'IT|T'ZII1 -—-_J^"ml II A. brush to sweep bees from -^ the combs is a convenience, especially in removing bees while taking frames from the hives at extracting time. The German brush with white bristles (Fig. 28) is perhaps the best of those manufactured, but a turkey feather, a long whisk broom or a bunch of weeds pulled as needed are as good. A tool box or portable FlG- 2a ~ German bee brush-seat (Fig. 29) and a wheelbarrow or cart for carrying supplies honey are among the. other conveniences used in handling FIG. 27. — Hive tools. 32 Beekeeping FIG. 29. — Tool-box seat. bees. A hive cover on edge makes a good temporary seat and has the advantage of being where it is needed, and when needed is not otherwise occupied. Other equipment. There are some additional appliances which may be useful in any apiary and which may be mentioned briefly. For making changes in supplies and in devising parts for special uses, the apiary equipment should include some carpenter's tools, among which may be mentioned hammers, saws (including a keyhole saw), brace and bits, square, planes and a good supply of nails of assorted sizes. Cement-coated nails are the best for most purposes FIG. 30. — Alley queen and drone trap. FIG. 31. — Bee-escape board. in the apiary. Queen and drone traps, usually known as Alley traps (Fig. 30), ar^ useful in catching undesirable drones or in preventing the escape of a queen at swarming time (p. 273). Bee-escapes (Figs. 19 and 31) are used in removing bees from supers of h$ney? especially comb- Apparatus 33 honey, before it is taken from the hive. An observatory hive with glass sides will be found instructive and entertaining to the beginner and even to the more experienced beekeeper, if placed where the bees may be watched frequently. A COmb-fOUn- FlG< 32. __ Comb-foundation cutter. dation cutter (Fig. 32) is convenient and better than an ordinary knife. If the beekeeper desires to make his own comb-foundation, there are various machines that may be obtained for that purpose. It is usually cheaper to buy foundation. In case it is necessary to feed colonies in order to stimulate brood-rearing or to provide stores for winter or dur- FIG. 33. — Van Deusen hive clamp. ing a period when no nectar is available, various types of feeders may be used. The construction of these is indicated in the illustrations (Figs. 105, 106, 107 and 108), given in connection with the discussion of feeding (p. 240). Clamps for holding the parts of the hive together (Fig. 33) are convenient in moving, but the wide (1J inch) staples sold by dealers in beekeeping supplies are as good. CHAPTER III THE COLONY AND ITS ORGANIZATION IN the proper management of bees, all manipulations must be based on their'normal activities, Bees are creatures of instinct and are limited in their ability to adapt themselves to changes in their environment. While in certain activities they show evidences of memory, learning, association and adaptive responses, in general they may be considered as responding to their environment in a "machine-like" manner. Because of the nature of most of their activities, it becomes necessary to know their normal behavior even more than would be the case were they more adaptive. In giving directions for handling bees, the systems of manipulations and apparatus are usually emphasized, but in the present book the normal activities will be made more prominent so that the reader may better understand the reasons for the usual rules and systems. Again, most of the American literature applies especially to the white clover region and the rules fail to apply elsewhere, so that there seems to be additional justification for a discussion of the more fundamental factors in beekeeping. It frequently happens that a supposedly new plan or system is published which is old, except that it is a new adaptation of well-known principles to slightly changed conditions. The success or failure of these plans when tried by others is often attributed to peculiarities of the various localities where they are tested. The word "locality" is called upon to cover a multitude of defects in our knowledge of bee activities. Bees respond to stimuli in but one way wherever a given stimulus is applied, the result is the m The Colony and its Organization 35 same. If one's knowledge of the circumstances surrounding his bees is not adequate there seems to be comfort in attributing to "locality" one's failure in the application of rules. Point of view. It may be worth while to extend these introductory remarks to explain the point of view held in the present discussion of bee activities. There are several distinct angles from which one may view the actions of a colony of bees and, since they lead to unreconcilable conclusions, they cannot all be correct. First to be mentioned among those who write concerning bees is the so-called student of nature who seemingly tries to find in bees a type of intelligence even higher than that possessed by man and who attributes to these insects thoughts and passions to which only the poetic may hope to attain. The complex colony life of bees offers to such a type of mind unlimited opportunity for speculation, which leads nowhere and is in fact a detriment to legitimate investigation. Allied to the just mentioned enthusiasts over nature are the amateur philosophers who hold up the bee as a brilliant example of industry. To all such speculative fancy, we may with profit turn our backs. In studying the behavior of any lower animal, there is but one source to which one should go for information. This is found in the actions of the animal in response to stimuli of its environment. If the bee makes a visible movement in response to a stimulus arising in its environment,1 that visible movement and nothing else is of value in forming a conclusion. If there is a movement or other response inside the animal or otherwise invisible, or if the bee perceives the stimulus but does not move in response, then the observer has a negative result. It is frequent in bee literature to find 1 The environmental factor may be inside or outside the hive, or even inside or outside the individual bee. For example, pathogenic microorganisms or irritating foods are inside but not part of the animal and are therefore environmental factors. 36 Beekeeping the words "think," "know," "suppose" and the like applied to bees. As a figure of speech such a form of expression may perhaps be admissible, but if used in its absolute sense then it is not warranted. It would result in a marked diminution of the literature on bees, and a great improvement therein, if such material could be wiped out of existence. Danger from poor work. There is but one source of erroneous theory more dangerous than those mentioned and that is the observer who makes false observations and unwarranted deductions. Here too the bee has not escaped. Because of the wide interest in bees there has been a demand for scientific information concerning them and this has induced several untrained or poorly trained men to undertake observations on the structure or behavior of bees, for which they were not equipped. Such work, being frequently presented in a more popular and attractive form than genuine scientific work, has had much influence among beekeepers so that, in attempting to present the results of thorough work, it is first often necessary to show the inaccuracies of work done by unqualified writers. Advantage of experience in behavior investigations. It must not be supposed that our present knowledge of the behavior of the bee is complete. It is, in fact, woefully meager. It is probably true, however, that a well-informed beekeeper has a wider and more accurate knowledge concerning bees than have many students of animal behavior concerning the species with which they work. The intimate acquaintance of the beekeeper with these insects results in a knowledge of their activities which, while faulty at times due to a lack of training in observation, is as a whole quite accurate. While this information is often fragmentary and is usually acquired without any special realization of the general principles of behavior, at the same time the data acquired through years of contact with the bees are perhaps The Colony and its Organization 37 as reliable as those obtained by the experimenter on other species in the course of a relatively brief investigation. A new worker in bee behavior should hesitate before denying the belief of the beekeeper until he is sure of his ground. Zoological position of the honeybee. The honeybee belongs to the order of insects known as Hymenoptera, to which belong also many parasites of other insects, the solitary and social wasps, ants and the entire group of bees, from the solitary species through various stages in the development of the bee colony to the honeybee. The honeybee is the highest of these colonial forms, highest because most specialized in its behavior and least able to exist alone. Yet, while it is highly specialized in its behavior, it is not so strikingly modified in its structure as are some of the other Hymenoptera, such as the Ichneumonidse. Among the Hymenoptera there are three groups of social insects, wasps, ants and bees, and the type of colony found in these three groups is fundamentally the same. The only other true colonial insects are the termites, "white ants," of a distinct order and with a quite different type of colony. The genus Apis to which the honeybee belongs also includes the species indica^ florea, dor&ata and zonata, all of which are natives of the far East and none of which is as useful to man as the species mellifica.1 These are briefly discussed in Chapter IX. 1 One of the cases of confusion originating from the application of the law of priority in scientific nomenclature is the attempted change of the name of the honeybee from mellifica, by which it has been known for so many years, to mellifera. In the 10th edition of Linnaeus' "Systema Naturae" (1758), the boundary of the prehistoric for the taxonomist, the name mellifera was used, while Linnaeus himself used mellifica in later years. The name mellifica is found in a vast literature, it is the scientific name by which the bee is known to most zoologists and beekeepers, the name which Linnaeus preferred and, last but not least, it is a correctly descriptive name. It should be recognized in taxonomy, as well as in civic legislation, that a law to be effective must be backed by public sentiment. It might therefore with propriety be suggested to the taxonomic purists that they cultivate public sentiment by allowing the zoologist, dealing in things not names of things, to live in peace among his old friends. 38 Beekeeping Bees not domestic animals. Bees have been kept by man from an early stage in the development of human civilization, yet it cannot be said that they are domesticated. In a^ll of their activities, bees under the care of man do not differ from bees in a wild state. The bee has been modified by breeding in various ways but, in so far as the natural instincts are concerned, it is doubtful whether any appreciable change has been brought about and in the greater number of phases of bee life no change has even been attempted. An escaping swarm takes up its abode in a hollow tree and the bees are often then spoken of as "wild/' but this adjective is just as applicable to the bees in the apiary. Certain animal trainers become proficient in handling savage animals through their knowledge of the ways of these beasts. Similarly the beekeeper, by studying the behavior of his bees, comes to know their habits and is governed by this knowledge. This comparison of bees and wild animals must be construed not as intended to inspire fear in the uninitiated but to point out that the beekeeper actually is dealing with animals unmodified in their instincts by their long association with man. By the proper use of smoke and especially by the way the colony is handled, the beekeeper can seemingly do with his bees as he pleases. The fact is, however, that he cannot overstep the bounds set by the instincts of these animals. It is therefore an incorrect conception of the ability of the beekeeper to state, as did Langstroth, that bees are capable of being tamed. In view of these facts, the necessity of a thorough knowledge of bee activities is most evident. Necessity of colonial life. Bees cannot live alone. Their structure and instincts fit them for life in a colony or community, where the various duties are divided among the individuals according to struc- Many zoologists refuse to take taxonomy seriously and there seems every reason for disregarding its laws in the present case. The Colony and its Organization 39 tural fitness and age. While an individual worker bee may live if forcibly isolated from its mates, it cannot reproduce itself, fails to care for itself adequately and soon dies. Most insects have the ability to hibernate in winter but the honeybee seems to have lost that ability. Sin after even a short confinement, will venture out for this purpose as soon as the outside temperature permits.1 Movements in winter. The cluster is first normally formed where the last brood emerged; here the bees find empty cells into which to crawl, so that they form themselves into a compact mass, separated only by thin walls of wax. They do not form the winter cluster where the combs are filled with honey and it would probably be impossible for them to conserve the heat of the cluster if sheets of honey separated the bees in adjacent spaces. As the contiguous stores are consumed and additional cells are emptied, the cluster is shifted so that the bees are always near stores. This shifting ig, however, apparently impossible in extreme cold weather, when colonies sometimes die in a way that can be explained only as due to starvation through inability to reach stores just a few inches distant. The early fall cluster is usually low on the combs, near the entrance, if there is considerable honey stored and the movement of the cluster is usually upward and toward the rear of the hive as winter progresses, and as stores are consumed. If a colony is in a two-story hive, the cluster is often in the upper story in the spring. Responses to outside temperature. The cluster varies in size with the outside temperature. After the emergence of the last brood in the fall, if the 1 Bees often remain in the hive at a temperature of 70° F. if a flight is not necessary, but will often fly when the temperature is about 50° F. if they have been confined for a time. 90 Beekeeping temperature is about 60° F., the bees do not cluster compactly and do not fly from the hive, even on bright days, but remain inactive on the combs, In this condition they are less active than at any other time in the cycle of the colony and approach most nearly to a condition of hibernation. During the warmer days there is no need of a tight cluster, for the function of the cluster is the conservation of the heat generated within. When the temperature is sufficiently high the bees generate no heat but, whenever the temperature of the air immediately surrounding the bees drops below 57° F. (the lowest temperature which normal bees ever experience in the hive), they form a definite cluster. As the outside temperature continues to fall, the cluster becomes more and more compact and the temperature of the inside of the cluster increases rapidly. After the generation of heat is begun, the temperature within the cluster soon reaches a point higher than that reached before heat generation was necessary. Within certain limits, the temperature of the cluster increases as the outside temperature drops and, as the outer temperature again rises, heat generation is reduced or discontinued while the temperature of the cluster drifts to meet the rising outside temperature. Heat generation is renewed if the outer temperature again drops, even though the temperature of the cluster and that of the outer air have not yet been equalized. This produces a peculiar inverse relationship between the outer and cluster temperatures. It is of particular practical importance that, within certain definite limits, the bees are not compelled to produce heat. Conservation of heat. The cluster consists of $ hollow sphere of bees several layers thick, those between the combs with their thoraces in contact and abdomens extending outward. The cells within the cluster are also filled with bees. The hair on the thorax assists in making this living shell an excellent non-conductor of heat, so effective in fact that a point in- The Cycle of the Year 91 side the cluster may sometimes be 100° F. warmer than a point a few inches away but outside the cluster. The number of inactive bees varies with the outer temperature, being larger at warmer temperatures when less heat production is required and smaller when more bees are engaged in activities involved in heat production. Source of heat. Within the hollow sphere are bees which move about freely, these being the ones most concerned in heat generation. They produce heat by muscular activity, such as movements of the legs and abdomen, but perhaps most effectually by vigorous fanning. The bees which form the shell constantly shift their positions and exchange places with bees from within. A bee from the center forces its way head first through the shell, then turns around and remains for a time on the outside layer. The shifting seems to be more rapid in cold weather than in mild. Effect of accumulation of feces. During the winter, the bees consume the honey stored during the summer. The undigested portion, which forms excreta, is retained in the rectal Ampulla (hind-intestine) until the bees have opportunity for flight, for normally no feces are deposited by the workers within the hive. During the cold winters of the North there are times when bees cannot fly for several weeks and the generation of heat during such a period 6f cold weather requires increased consumptioii of food arid causes an increase in the amount of feces. The presence of feces, on the other hand, causes the bees to become restless, to generate still more heat (see Fig. 145) arid to accumulate still more feces. Apparently a colony iri winter confinement is in the confines of a vicious circle and the successful wintering depends preeminently on good food. If, however, the colony is so placed that little or no heat must be produced, the situation is relieved and this the beekeeper accomplishes by placing colonies 92 Beekeeping in the cellar, provided conditions within the cellar are correct. Bees wintering in the open fly out whenever the outside temperature will permit, and after a considerable period of confinement many will fly out when it is so cold that they are unable to return. On these winter flights the feces are voided, consequently they are of the highest value to a colony wintered out of doors. While numerous other points concerning the activities of bees in winter are left to be discussed in the chapter on wintering, it is evident from what has been said that bees are highly sensitive to changes in temperature, and that they have a wonderful ability to overcome the adverse conditions of winter by the generation of heat. It is to be noticed especially that they usually do not warm the whole hive or cavity but confine heat production to the cluster. It might therefore be concluded that a hive is actually little protection for them in winter but it should be remembered that this protects them from wind, rain and snow. They further seal the hive with propolis to make the top watered even air-tight and some races contract the entrance with propolis. The practical bearing of these facts forms one of the most vital problems of the northern beekeeper and the discussion of this subject from the standpoint of practice forms a later chapter. CHAPTER V THE LIFE OF THE INDIVIDUAL IN RELATION TO THE COLONY IN the preceding chapter the activities of the colony are discussed, much as if the colony were an individual or unit. While this is a true picture of one side of the life of the species, it is not complete, for the individuals not only carry on their own life processes but pass through individual cycles. A knowledge of the interrelationships of the individuals within the colony is important for an understanding of the colony organization, for this complex society is based on a division of the labors of the hive, which is of the highest interest and of the greatest practical value to the beekeeper. While in this book there is no attempt at a complete discussion of the anatomy or development of the bee, it is necessary that these subjects receive some attention to outline correctly the little that is known concerning the physiology of the species. The discussion of physiology is reserved for another chapter. DEVELOPMENTAL STAGES This subject is one of mystery to the beekeeper. While the development of the bee in the egg has been investigated by several observers,1 the papers on the subject are not 1 Butschli, O., 1870. Zur Entwicklungsgeschichte der Biene. Zeit. f. wiss. Zool., XX. Kowalevski, A., 1871. Embryologische Studien an Wiirmern und Arthro-poden. M&m. acad. imp6r. sci. St. PStersbourg, (7) XVI, 12, pp. 1-70. Grassi, Battista, 1882-84. Studi sugli artropodi. Intorno allo sviluppo 93 94 Beekeeping readily accessible to beekeepers, and writers of books on beekeeping have not given to this subject as careful consideration as to the anatomy of the adult bee. The changes taking place during metamorphosis (pupa stage) are so wonderfully complex that an account of the transformation of the larva into an adult bee is almost unbelievable.1 Because of the lack of attention given to the development of the bee in the literature on beekeeping, relatively more attention is here given it than to the anatomy of the adult bee. Cellular structure of tissues. To form a correct understanding of the development of the bee or of the structure of the adult, one must know something of the units of which the tissues are formed, called cells. This word, as used by the biologist, has a special significance, being applied to a type of structure which makes up the tissues of all plants and animals. This unit of structure is usually microscopic and a single .organ of the bee may contain many thousands of them. The cell consists of a minute mass of protoplasm (living substance) containing a nucleus2 (Fig. 50). Protoplasm is a complex organic substance characterized by life; the nucleus is a differenti- dell Api nell' uovo. Atti dell' Acad. Gioenia di scienze nat. in Catania, Ser. 3, XVIII, pp. 145-222. Dickel, O., 1904. Entwicklungsgeschichtliche Studien ana Bienenei. Leipzig: Engelmann. The work of Carriere u. Biirger (1898, Entwicklungsgeschichte der Mauerbiene. Abhdl. der kaiserl. Leop. Carol. Deutsch. Akad. der Naturf., LXIX, 2) on the mason bee, Calicodoma, is of value in a study of this subject. The recent work of Dr. Jas. A. Nelson of the Bureau of Entomology (1915, The embryology of the honeybee. Princeton University Press) is the most complete on this subject and is the most thorough work on the development of any insect. It is the only discussion of the embryology of the bee in English and should be consulted by any one interested in this phase of the life of the bee. The author is indebted to Doctor Nelson for help in the preparation of this section. 1 The metamorphosis of the bee is described in detail by Anglas, J., 1900. Observations sur les metamorphoses internes de la guepe et de 1'abeille. Ill pp. Lille: Danel. 2 This word, like "cell," is one of various meanings. It is used by the beekeeper to designate a small colony. The Life of the Individual 95 ated portion of the protoplasm which is especially active during the division of cells and carries the special organs (chromosomes), instrumental as the bearers of hereditary characters. The nucleus and surrounding protoplasm are closely united in their functions and are incapable of separate existence. The nucleus is, in its resting condition, usually rounded in form, while the remaining protoplasm is of various shapes according to the special functions of the cell. Protoplasm is characterized by ability to takp in nourishment, to grow, to give off waste, to divide and to move in response to stimuli, but in each organ the cells become specialized to do some one thing especially well and they often lose some of the functions of primitive protoplasm. For example, a nerve cell loses its power of contractility but becomes specialized for transmitting nervous impulses, while a muscle cell has a marked power of contractility. A detailed discussion of the structure and function of the various,parts of the cell in different tissues is, of course, impossible here,1 but these few suggestions are sufficient to indicate the extreme complexity of the organization of each tissue that goes to make up any organism, such as the bee. The egg. The egg, as it leaves the ovaries of the queen where it is formed, is essentially a single cell. The eggs of most ani- 1 The interested reader is referred to Wilson, E. B., The cell in development and inheritance. New York: Macmillan, and to other works on cytology. FIG. 50. - - Group of tissue cells from skin of young salamander. 96 Beekeeping mals known to the layman require fertilization'(a union with one of the reproductive cells of the male) before they can develop, but there are many cases in which this is not necessary and the development of the drone bee is of this character. The eggs which develop into females are, however, fertilized. This difference has so important a bearing on practical beekeeping that a discussion of it is reserved for a future chapter. The egg of the bee is a small white cylindrical object about yg- of an inch long, somewhat larger at one end (future head end) and slightly curved. It is deposited on the base of the cell of the comb by the queen and is fastened in place by a secretion. The head end of the future larva is always formed away from the point of attachment. The egg is covered by chorion, a thin, tough membrane, the surface of which is ridged. These ridges are, however, quite minute and are not so conspicuous as most illustrations of bee eggs would indicate. In addition to the nucleus and surrounding protoplasm, the bee egg contains a relatively large amount of non-living stored food, yolk. The embryo is formed on the convex side of the curve of the egg, which becomes the ventral side of the larva. The fate of the various parts of the egg is therefore in a sense determined. Because of the presence of so much yolk, the early cells are not clearly marked off from one another.. Early embryonic development. Development consists of the repeated division of the egg cell into numbers of united cells and of the rearrangement and differentiation of the resulting cells to form definite organs. As development proceeds, the cells become more and more specialized until the final adult condition is reached, and even in the adult, certain changes in some cells continue through the life of the individual. As cell division (or rather, in this case, nuclear division, for the protoplasm is continuous in the early stages) progresses, the nuclei move from the interior to the surface. During the second half of the The Life of the Individual 97 second day, a thickening appears on the convex side, and, on the anterior end (larger end) of the egg, the first indications of the future appendages are soon visible (Fig. 51, a and b). These consist of the rudiments of the antennae (Ant) and mouth parts (mandible, Md and maxillae, IMx, 2MX) on the head and of the three pairs of legs (1L, 2L, 8L) on the thorax. These rudiments are simply slightly rounded swellings which are at first smaller toward the posterior end of the egg, since development progresses from the anterior end. The embryo shows at first no division into FIG. 51. — Three stages in the development of the embryo. head, thorax and abdomen, but the fate of the various swellings must be determined by following them through. The rudiments of the stigmata (Spt openings of the tracheal system) appear early and the first evidence of the silk glands (SlkGl) becomes visible about the same time just behind the second maxillae. The first external indication of the nervous system is in two pairs of swellings (Br) on the upper side of the head. Even in this early stage, a number of important organs are already outlined, 98 Beekeeping Later embryonic development. In a later stage (Fig. 51, c), the embryonic band on the ventral side of the egg has widened and in the next stage here illustrated (Fig. 51, d) the band completely envelops the egg. In the stage shown in Fig. 51, c the mouth (Mth) and anus (An) have appeared as pits. These continue to grow into the egg and ultimately join with certain cells on the interior to complete the alimentary canal. The portions formed by the two invaginations from the outside are the fore- and hind-intestine, while the part arising from the interior is the mid-intestine. The Malpighian tubes (MT), the excretory organs, arise as outgrowths from the anterior en4 of the hind-intestine. The pits (Sp) which are the rudiments of the spiracles, deepen and send branches forward, backward and downward to meet corresponding outgrowths from other pits, finally forming the tracheal trunks with their commissures and branches. The silk glands (SlkGl), which function only in the larva, project backward as long tubes. Segmentation. The most striking feature of the late embryo is the fact that it is constricted into a series of segments (metameres or somites) which are plainly recognized in the larva. These segments are characteristic of all insects and part of the metameres of the abdomen are still plainly marked off in the adult. From the fact that segmentation is recognizable in various parts of adult insects and is present in insect embryos, it is assumed that this form is characteristic of the primitive organism from which all insects have arisen. The typical appendages are arranged in pairs on the segments but in their later development these appendages are modified according to their fate. The stigmata and the ganglia of the nervous system are also arranged segmentally at first, but this primitive arrangement is later partially lost. The segmentation of various species studied does not wholly agree, but it is usually assumed that the first six or seven The Life of the Individual 99 segments coalesce to form the adult head, the next three the typical insect thorax, and the remaining ones, usually twelve 1 in number, form the abdomen. The thorax of the adult bee is not typical, as will be explained later. Fate of parts of the embryo. Some of the head appendages of the embryo disappear early, being rudimentary organs. For example, the appendages of the second segment become the antennse while those of the third disappear in insects, but in Crustacea (e.g. shrimps and lobsters) form the second antennse. Several of the segments of the primitive insect head are not recognizable in the bee. In the adult insect, these segments fuse completely and by growth of various parts are so distorted that an examination of the adult head does not suggest segmentation and, without a study of the developmental stages, this segmental origin would be unsuspected. The three thoracic segments are fused in the adult bee but, since the three pairs of legs arise from them, the segmental origin is suggested. The wings arise as secondary outgrowths or appendages, dorsal to the legs, from the two posterior thoracic segments and do not correspond with other appendages. In the adult bee, the first abdominal segment is also fused with the true thoracic segments to form the part known as the thorax, which therefore does not correspond exactly with the thorax of lower orders of insects. This fusion also occurs in most of the other Hy-menoptera. The remaining posterior segments form the abdomen of the adult but not all of the segments remain visible to the outside. In the adult worker and queen bee, the five posterior segments are turned in to form a pocket around the sting and anus. In the drone, only four segments are so turned in. The embryo, just before leaving the egg, shows no rudiments of antennse or legs, these temporarily disappearing. 1 Two of these segments are obscure and in later stages there appear to be present, only ten. 100 Beekeeping The nervous system is now well organized, consisting of the brain and a chain of ganglia arranged segmentally. The second maxillae fuse to form the lower lip (Lb). Larval development. At the end of about three days of embryonic development, the embryo breaks the chorion and becomes a young larva. During the larval period the most striking feature is the enormous growth of the animal. The illustration on page 40 (Fig. 35) shows an egg, a relatively young larva, a fully grown larva and a pupa drawn to the same scale and, when it is realized that the growth from the youngest larva to the fully grown larva takes place in a few days, the rapidity of growth is astonishing. It should be pointed out that the development of all insects is not similar. In the grasshopper, for example, a young insect hatches from the egg which resembles the adult in most respects. Such a development is known as incomplete metamorphosis. In the higher orders of insects, there hatches from the egg a larva unlike the adult and usually more or less worm-like, which when fully fed undergoes a complete aoid relatively sudden change into the adult. This type of development is known as complete metamorphosis. The bee larva is an extremely simple organism, lacking legs, wings, antennae a;nd eyes, and is unprotected by hairs or thick chitin. A longitudinal section through the larva (Fig. 52) shows that the largest organ is the stomach, as is necessary for excessive growth. Being protected from enemies and from adverse environmental condition^ in the FIG. 52. — Diagram of a longitudinal median section of a bee larva. The Life of the Individual 101 cells of the comb, the bee larva needs no protective covering and, being fed by the worker bees,1 it does not need organs which will enable it to seek or even to detect food or to masticate solid food. It is ideally adapted to the protected condition in which it is placed in the colony scheme and quickly perishes if removed and exposed to adverse conditions. Metamorphosis. After the excessive growth, the larva is sealed in the cell with a capping of wax (Fig. 39) and it then spins a delicate silken cocoon with the secretion of the silk glands (SlkGl) within the cell. Soon after this, all external motion ceases and the animal begins to undergo that wonderful series of changes known as metamorphosis. During the larval growth the mid-intestine and hind-intestine are not connected (Fig. 52) but this connection is made after sealing and the feces of the larva are then cast out. The organs which served the larva are of course not suitable for the adult insect and the changes necessary to obtain suitable adult organs take place in the pupal stage. Anglas has described many of these changes but the metamorphosis of insects is so complex and so much disputed by various workers that it is to be hoped that the changes in the bee may be again investigated. The simple alimentary canal of the larva is discarded and a new one is formed in its place. The segmentally arranged muscles of the larva either disappear or are changed into those of the adult. The nervous system apparently loses some of the segmental ganglia by the fusion of various ganglion pairs. The antennae, eyes, legs and wings develop from rudiments which have remained undeveloped in the larva. Not only do the internal organs change and new structures appear but the animal changes 1 The larva of the honeybee is fed frequently during the period of rapid growth. In bumblebees (Bombus) and stingless bees (Melipona and Trigona), a cell is filled with a mixture of pollen and nectar, after which the queen lays an egg on the mass. The cell is then sealed and the larva is not led further during the developmental stages. 102 Beekeeping its outward appearance. The small head of the larva grows to adult size, the thoracic segments and the first abdominal segment unite and undergo marked external changes to form the thorax of the adult. The abdomen changes least in external form but marked internal changes occur. This brief category of the vital modifications can give but a suggestion of the changes which the pupa undergoes. All of this occurs in an animal which externally seems lifeless, but the internal changes require such large expenditures of energy that the animal loses weight by the consumption of the food which the greedy larva stores up as fat in the fat body. The external changes of the pupa are interesting, even though of minor importance. The compound eyes first change from white to pink by the deposition of a pigment around the rhabdomes of the eye (p. 167) and later this pink pigment is gradually covered by a darker external pigment so that the eyes appear brown and then black. The thorax shows coloration earlier than the abdomen. Toward the close of the pupal period, the outside of the animal becomes covered over with a layer of hard chitin for the protection of the adult and to serve as a skeleton for the insertion of the muscles. The legs and wings originate as hollow bud-like outgrowths on the thorax and after the last moult of the larva these imaginations are suddenly extended by blood pressure. The wings are at first small thin sacs which grow and finally take on the adult form, after which the two sides of the sac fuse and the blood in the sac returns to the body cavity, leaving the wings as dry membranes. Length of developmental stages. The length of the various stages of development varies among the different types in the hive. The preceding account applies especially to worker bees, which have been most frequently investigated, probably because of the ease of obtaining material. The stages are essentially similar in queens and drones. While the rapidity of development The Life of the Individual 103 is slightly modified by changes in temperature of the hive, it is, in the main, quite uniform and it is therefore possible to* give the time from egg-laying to emergence of the adult. It must be understood that these vary somewhat and it is rather remarkable that the variation is not more pronounced. On account of the variation the various tables given for the length of stages are not uniform. The following table (II) is a fair average: TABLE II. DEVELOPMENTAL STAGES STAGE QUEEN WORKER DRONE Egg ..... 3 3 3 Larva .... 5| 6 6i Pupa ..... 7i 12 14| Total .... 16 21 24 The figures given in this table for the pupal stage include all the time that the developing bee is sealed up in the cell. During part of this time, the larval stage is continued but no additional food is taken. This is followed by a semi-pupa stage, when the insect resembles a larva but has undergone a moult and the hind- and mid-intestine are connected. The true pupa stage follows this and the transition to the adult is gradual, the separation between the two stages being marked by the emergence of the insect from the cell. The number of moults in the larval stage are sometimes given as probably six (Cheshire). This should be more carefully studied. The structure of the adult bee will be briefly discussed in conjunction with the functions of the various organs. When the young bee emerges from the cell it is structurally in the adult condition. It does not grow in size nor do any marked changes in most of the organs occur during adult life. This is true of all insects. While certain internal 104 Beekeeping organs undergo change, these are not of a character to change the outside appearance. The food taken by the adult is not stored up within the body, as in the larva, but is taken for immediate use. THE CYCLE OF DUTIES OF THE ADULT WORKER BEE When the worker emerges from the cell, it is covered with a soft skin, the last pupal moult, which is quickly removed. For a day or two the young bee remains on the combs, frequently on the one from which it emerged, and moves about but little. Numbers of young bees are often seen in the upper part of the hive and especially in the supers. In a few days they begin the inside work l of the hive which 1 An interesting opportunity for speculation is offered in attempting to determine the basis for the division of labor in worker bees according to age. In studying the structure of the compound eye, the author (Proc. Acad. Nat. Science, Philadelphia, Vol. LVII, pp. 123-157) was struck by the presence of enormous numbers of curved unbranched hairs which cover the eye of the young adult bee so completely that the facets are not visible. These hairs are broken off readily and in field bees most of the hairs have disappeared. It is probably impossible for the compound eyes to function while these hairs remain. These facts suggested the possibility that the young bees remain in the hive because they cannot see clearly enough to fly to the field and that when the hairs are lost the field work is begun. That the young bees are capable of flight is clearly shown by their ability to leave with a swarm. In this case, sight is probably not essential. In attempting to determine whether there is any ground for such a belief, numerous experiments were tried, by removing the hairs of young bees to see whether they were then more inclined to leave the hive. The hairs were scraped from some young workers and in other cases soft paraffin or beeswax and paraffin was applied to the eyes and then removed, the hairs breaking off with its removal. In every case the handling made the action of the bees abnormal, so that no conclusions of any value were obtained. That this is probably the correct interpretation .of the function of these hairs still lingers in the mind of the author, in spite of inability to obtain proof through experiments. It may be said in favor of this theory that it offers a structural basis for an instinct which is otherwise unexplained. The attribution of an action to "instinct" is a lazy way of explaining phenomena. Merely to classify an action and group it with others, to which a class name is given, does not throw any light on the behavior. When an action is attributed to "instinct" the study of the behavior often suffers a loss rather than gain, for the giving of a name, to some minds, constitutes an explanation. There is reason for the belief that instincts all have a physical basis, some The Life of the Individual 105 consists of feeding and caring for the larvae, feeding the queen and the drones, cleaning, ventilating, comb building when necessary, guarding the hive from intruders and other work inside the hive.1 When about a week old,2 on bright days, the young bees take "play flights" in front of the hive. Suddenly, as if in response to a signal, the young bees fly out, circle about the hive, usually with their heads toward the entrance, and as a rule they do not at first venture more than a few feet away. In a short time this flight is over and the young bees return to the hive. This flight of young bees is often mistaken by beginners in beekeeping for the attack of robber bees but the action in the two cases is so different that close observation soon makes the dissimilarity clear. When robbers are numerous, they dart toward the hive and alight about every crack, while young bees circle about, rarely alighting on the hive. The flight of the young bees is also sometimes mistaken for the beginning of swarming. Later flights are more extended, and when workers are from 14 to 21 days old (if during a honey-flow), they begin their field duties of gathering nectar, pollen, propolis and peculiar physical structure which determines the action. This physical basis may be a specialization of some nervous element or of some other organ, but it probably always exists. In the case under discussion, it is not enough to state that the division of labor inside and outside the hive is instinctive and such a statement is largely an evasion of the problem which the facts observed present to us. 1 In addition to the inside duties named, the young bees must sometimes serve as honey reservoirs during a heavy honey-flow. Especially in comb-honey production where the bees must be crowded to produce fancy honey, the comb built is often not sufficient to hold the nectar brought in and it is given to the young bees. They may be seen in the evening on the combs with abdomens distended, but usually before morning more comb is completed and the honey is deposited in cells. Possibly this may be part of the ripening process, which is poorly understood as yet. This function of young workers suggests the behavior of the honey ants, in which certain individuals serve as honey pots for the storage of honey until used. In this case the abdomen is abnormally distended. 2 In giving age in days or weeks it must be understood that this is variable, depending on season and honey-flows. The determining factor in the aging of bees is work, not days (p. 126). 106 Beekeeping water. Normally, they now abandon the work inside the hive. It sometimes happens that a colony will contain relatively too many young bees or too many old ones, these conditions often arising in practical manipulations. If there is a lack of young bees, the old ones act in their stead, but they secrete wax slowly (p. 108) and do not produce larval food adequately. If a colony is made up artificially of young bees, some of them begin field work earlier than normally. DIVISION OF LABOR From the preceding chapter, it is evident that there is a definite division among the different members of a colony. In a colony composed of perhaps 60,000 individuals, the very existence of the bees depends on an orderly performance of the various duties, and the development of colonial life, therefore, rests on the evolution of some system for the division of labor. The organization of the colony, already described, shows one of the most marked cases of apportionment of work, for the egg-laying is normally performed by but one individual, the queen, while all the other females (workers) are so constituted that egg-laying is not normal and mating is impossible. The drones or males are so specialized in function that they are probably useful to the colony only in the mating of young queens. While the duty of egg-laying devolves on the queen, the care of the brood falls entirely to the workers. Since they must do work both inside and outside the hive, there arises the further necessity of a division 1 of these functions and this, as has been stated, is based on the age of the individuals. 1 The division of labor is as highly developed among bees as in any insect community. Among certain species of ants, a greater diversity of structure accompanies the performance of certain duties. For example, there may be soldiers which serve only as protectors of the community and there may be two types of workers, differing structurally and in their duties. While structural differences do not occur in so marked a way, the members of the bee colony are fully as greatly specialized in their labor but the performance of specific duties is determined in some manner other than by structure. The Life of the Individual 107 Since it may not at first glance be clear how the age at which bees perform certain functions is determined, it may be well to explain the simple method by which this is accomplished. If the queen is removed from a colony of black (German) bees and a yellow (Italian) queen is at once introduced, for a period of twenty-one days after the removal of the old queen the young worker bees which emerge from the cells are black, since they are the progeny of the old queen. At the end of that time, however, the worker brood from the black queen has all emerged and yellow bees begin to appear. The time at which the yellow bees first perform certain functions may now be determined. This experiment may be variously modified, as by the removal of all the brood of the black queen at once or by the placing of a frame of brood from an Italian colony in a colony of black bees. The introduction of Italian bees into Germany and later into America has been an important factor in enabling investigators to learn many of the phenomena of the hive, for the use of bees of two colors 1 is often of the highest importance. The labor within the hive. When the workers first emerge from the cells they take no part in the work of the hive for a day or two, nor do they leave the hive. The first flight in front of the hive is usually when they are about a week old, if the weather is favorable, and these flights are continued on warm bright days until they are nearly three weeks old. Although they do not go far at first they may remain on the wing for a considerable period. That these early flights are necessary in enabling the young bees to void their feces is indicated by the fact that if confined they become restless.2 The abdomens of young bees are frequently distended with feces. 1 Another method of marking bees for observation is mentioned by Cas-teel, Cir. No. 161, Bureau of Entomology, p. 5. The method employed was to paint bees with different colors and also to number them. 2 This was observed when colonies containing young bees were placed 108 Beekeeping Donhoff1 states that he offered a stick dipped in honey to young bees daily. Until they were fifteen days old they did not lick the honey eagerly. The younger bees never attempted to lick it, but as they grew older they paid more attention to it. He concludes that the "impulse for gathering honey" is not developed in young bees. Not until his experimental bees were seventeen days old did he find any on his outdoor feeders and not until they were nineteen days old did any fly to the field. Comb building. If there is need for more combs, the workers form curtains by hanging on one another from the top of the hive or cavity. The temperature is raised and in a few hours wax-scales may be seen on the ventral sides of the abdomens of the hanging bees. Finally, some of these scales are removed and manipulated and the bees begin building new comb. The small pieces of wax are put approximately in the right place and are then sculptured and molded into FIG. 53. —Ventral plates of the abdomen of a worker bee. FIG. 54. — Inner surface of the left hind leg of a worker bee, showing a wax-scale. in a cellar for winter, in connection with work of Demuth and the author on winter activities. The entire colony became active and a high temperature was maintained. The condition was removed by taking the colonies from the cellar for a flight. Bees that emerged from brood combs were also kept in a warm room, away from older workers. These had distended abdomens and if one escaped from the hive it usually flew at once to the window, leaving a spot of feces on the pane. 1 Ddnhoff, 1855. EichstMt Bienenzeitung, p. 163. The Life of the Individual 109 FIG. 55. — Ventral view of worker removing wax-scale. Enlarged. their proper position and shape. In spite of the number of bees at work in building, the wax is quickly smoothed into its final form, becoming a part of the comb. Dreylingl has shown that in just emerged worker bees the cells of the wax glands are not fully§ developed and that as the worker grows older the cells elongate. As the bee ages, however, these cells decrease and degenerate. These results fully support the observations of beekeepers that bees secrete wax best before they become field bees. If, however, a colony of old bees is required to build comb, the bees can still secrete some wax, but for some reason not understood they usually build irregularly. Beeswax is secreted in pockets on the ventral side of the abdomen on the wax plates (Fig. 53) situated on the sternal plates of the last four visible segments of the abdomen. Each segment bears two of these plates, making eight in all. FIG. 66. — Side view of worker removing wax-scale. Enlarged. 1 Dreyling, L., 1903. Ueber die wachsbereitenden Organe der Honig-biene. Zool. Anz., XXVI. ——, 1905. Die wachsbereitenden Organe bei den gesellig lebenden Bienen. Zool. Jahrbiicher, Abth. Anat. u. Ont. d. Theire, XXII. 110 Beekeeping As the secreted wax comes in contact with the air, it hardens, forming the scales of wax. The manipulation by the bees of the wax-scales has been carefully described by Casteel.1 The scales are removed from the pockets by spines of the pollen comb (Fig. 54) on the first tarsal segment (planta) of the third pair of legs. The surface of the planta is passed over the ventral side of the abdomen (Figs. 55 and 56) and after the scale is loosened the third leg is bent forward (Figs. 57 and 58), thus passing the scale to the front pair of legs. It is then masticated by the mandibles, after which it is ready to put in place in the new comb. The various movements in manipulation are so well shown in CasteePs figures that further description is unnecessary. It is clearly shown that the so-called wax-shears, which are described by SO many authors as be- FIG. 58. — Side view of worker passing ing used to remove wax- wax-scale forward- Enlarged-scales, have in fact nothing to do with the wax manipulation. It is shown later that these are concerned in pollen gathering. FIG. 57. — Ventral view of worker passing wax-scale forward. Enlarged. 1 Casteel, D. B., 1912. The manipulation of the wax scales of the honey bee. Cir. No. 161, Bureau of Entomology, 13 pp. The Life of the Individual 111 Feeding of larvce. The feeding of the larvae is one of the most ardently disputed questions in bee activity. The chief controversy arises over the source of the food, some authors claiming that it is a secretion of glands, while others maintain that it is regurgitated from the ventriculus. The heat of controversy seems to have hidden from view the fact that this can be determined only by investigation. An explanation of the two current views involves some study of the glands emptying into the alimentary canal and of the ventriculus. There are in the head of the worker bee, two systems of glands (Fig. 59), the lateral pharyn-geal (supracerebral of Bordas, System No. 1 of Cheshire) (1GI) and the salivary glands of the head (postcerebral of Bordas, System No. 2 of Cheshire) (2GI), and in the thorax are found the salivary glands of the thorax (thoracic salivary of Bordas, System No. 3 of Cheshire) (Fig. 60, SGl). The ducts of the two systems of salivary glands unite into one median tube which enters the base of the labium and opens upon the upper surface of the ligula. These glands are homologous with the salivary glands of other insects and presumably their secretions assist in digestion although their exact function is unknown. They are found in queens, drones and workers. The lateral pharyngeal glands (1GI) are absent in the drone and never more than rudimentary in the queen, and this leads to the conclusion that they function in some way which is especially useful to the worker. They are claimed by FIG. 59. — Median longitudinal section of head of worker, showing the glands (1GI and 2Gt). 112 Beekeeping Schiemenz,1 and after him by Cheshire,2 to be the source of food given by the workers to the larvae of queens, drones and workers. It is claimed that the development of these glands is in proportion to the specialization of the species in the feeding of the larvae; in bumblebees (Bombus) they are as well developed as in the honeybee. They are decreas-ingly smaller in Psi-thyrus, Andrena and Anthophora. Since the feeding of some of these species is entirely unlike that of the honeybee, this evolution perhaps proves too much for this theory. Schonfeld,3 on the contrary, holds that the larval food arises in the ventriculus and not in these FIG. 60. — Alimentary canal of worker, showing glands, pharynx (Phi/), oesophagus (CE), honey-stomach (HS), proventriculus (Pvent), ventriculus (Vent), intestine (SInt), rectal ampulla (Reef) and Malpighian tubules (Mat). 1 Schiemenz, Paulus, 1883. Ueber des Herkommen des Futtersaftes und die Speicheldriisen der Bienen, nebst einem Anhange iiber das Reichor-gan. Zeit. f. wiss. Zool., XXXVIII, pp. 71-135. 2 Cheshire, 1886. Bees and beekeeping. 2 vols., London: L. Upcott Gill. 3 Schonfeld, 1886. Die physiologische Bedeutung des Magenmundes der Honigbiene. Arch. f. Anat. und Physiol. Abth., pp. 451-458. The Life of the Individual 113 glands. Cook 1 and Cowan2 both adhere to this view. The alimentary canal of the worker (Fig. 60), posterior to the pharynx, narrows to a slender oesophagus (OE) extending through the thorax. In the abdomen, this is enlarged into a thin-walled sac known in the honeybee as the honey-stomach (HS, crop of other insects), since it is used to carry nectar to the hive. At the posterior end this merges with the proventriculus, with heavy muscular walls, which contains a valvular apparatus (Fig. 61). Behind this is the stomach or ventriculus (Vent). Schonfeld claims that the brood food, especially that of the queen (royal jelly), is regurgitated from the ventriculus. The experiments of Schonfeld seem to show that the valve in the proventriculus opens and moves anteriorly even to the oesophagus when this is done, but Snodgrass 3 claims that this cannot be done without tearing the muscles of the proventriculus. Cowan and other authors figure this action in a diagram, but with no evidence from observation. Schonfeld and Cook fed charcoal in honey and found this in the brood food which would, in their estimation, be impossible if the food is of glandular origin, but they overlooked the fact that the charcoal might get into the brood food from the mouth of the worker. The charcoal could not pass through the walls of the ventriculus in FIG. 61. — Longitudinal median section of base of oesophagus. 1 Cook, A. J., 1904. The beekeeper's guide or manual of the apiary. 18th ed., Chicago. 2 Cowan, T. W., 1904. The honey bee, 2d. ed., London. 1 Snodgrass, R. L., 1910. The anatomy of the honey bee. Tech. Series, 18, Bureau of Entomology, pp. 162. I 114 Beekeeping digestion. According to Petersen, the peritrophic membrane in the ventriculus is so formed as to make regurgitation from the ventriculus impossible. While the work of Schiemenz and Schonfeld must be given due consideration, we must wait until some competent investigator takes up this problem. The various arguments are thus summarized by Snodgrass (p. 100): "1. The brood food itself is a milky-white, finely granular, and gummy paste having a strong acid reaction said to be due to the presence of tartaric acid. "2. The pharyngeal glands of the head are developed in proportion to the social specialization of the various species of bees; they are always largest in those individuals that feed the brood, and they reach their highest development in the workers of the honey bee. From this it would seem that they are accessory to some special function of the worker. "3. The contents of the stomach in the workers consist of a dark brown, slimy, or mucilaginous substance in no way resembling the brood food, even when acidulated with tartaric acid. Pollen is present in varying quantity, mostly in the posterior end of the stomach, and shows little or no evidence of digestion. Since the brown food is highly nutritious, it must contain an abundance of nitrogenous food material, which is derived only from pollen in the bee's diet. Therefore it is not clear how the stomach contents can alone form brood food. "4. The constituents of the food given to the different larvae, at different stages in their growth, and to the adult queens and drones show a constant variation apparently regulated by the workers producing it. A variation of .this sort cannot be explained if it is assumed that the brood food is produced by the glands alone. "5. Powdered charcoal fed to a hive of bees appears after a short time in the brood food in the cells, and this has been urged as proof that the latter is regurgitated ' chyle/ But it is certainly entirely possible that the charcoal found The Life of the Individual 115 in the food might have come only from the honey stomach or even from the oesophagus or mouth. "6. We have Schonfeld's word for the statement that a regurgitation of the stomach contents may be artificially induced by irritation of the honey stomach and ventriculus in a freshly dissected bee, but all explanations offered to show how this is mechanically possible in spite of the pro-ventricular valve are unsatisfactory when the actual anatomical structure is taken into consideration." TABLE III. COMPOSITION OF LARVAL FOOD$. — v. PLANTA QUEEN DRONES WORKERS Under 4 Days Over 4 Days Under 4 Days Over 4 Days Proteid . . . *Fat .... Sugar .... 45.15 13.55 20.39 55.91 11.90 9.57 31.67 4.74 38.49 53.38 8.38 18.09 27.87 3.69 44.93 Composition of larval food. The chemical composition of the larval food has been investigated by von Planta.1 This larval food is obviously not merely a mixture of honey and pollen nor is the food given the various kinds of bees at different ages uniformly the same. The following is a brief summary of von Planta's conclusions: The three kinds of bees require different food and, in the drone and worker larvae, the food changes after the third day, being mixed with half-digested pollen grains and honey in the case of the drone and honey only in the case of the workers.2 On the other hand the queen larva receives the rich food supplied the young larvsa of other 1 von Planta, Adolf, 1888. Ueber den Futtersaft der Bienen. Zeit. f. Phys. Chemie von Hoppe-Seyler, XII, pp. 327-354. 1889; idem, XIII, pp. 552-561. 2 Pollen grains are found plentifully in the mid-intestine of the older worker larvae, so that in this respect at least the results of v. Planta's work must be questioned. 116 Beekeeping castes throughout her entire larval period (called royal jelly) which is free from undigested pollen and completely predigested. The table (p. 115) gives the percentages of the various food constituents as determined by v. Plant a. Snodgrass (p. 93) reports finding undigested pollen grains in royal jelly, contrary to the statements of v. Planta. The larval food differs essentially in appearance from the contents of the ventriculus so that it is difficult to conceive of it being a regurgitated product to which is added merely an acid secretion of the glands. The beekeeping industry is under lasting obligation to v. Planta for his research in this and other subjects, but it is no disrespect to his work to express the belief that this subject should be thoroughly investigated by modern methods. The methods of analysis have been greatly improved since his work was done; they have, in fact, been so completely changed that v. Planta's results cannot be considered as conclusive in any respect. Feeding of queens and drones. In addition to the feeding of the various types and ages of larvae, the workers feed the queen and seemingly the drones also during their presence in the colony. The excessive egg-laying of the queen (p. 57) obviously calls for nourishment in large quantities and during the season of heavy laying the queen usually stops for a few minutes about every half-hour and during this resting period she is almost constantly fed. While the feeding of the drones is less easily observed, there is reason to believe that the feeding is discontinued at the close of the honey-flow, at which time the drones are first driven to the lower parts of the hive and finally are easily carried out, because of their weakened condition. Both queens and drones are capable of taking honey from cells, but apparently do not take pollen themselves. Other inside work. Little remains to be said in detail of the inside work of the hive which is performed by the workers. They clean The Life of the Individual 117 the hive, and in case they are unable to remove the debris, they may cover it with propolis. Lizards (Fig. 62), small snakes and other intruders to the hive, which are too large for the workers to remove, are sometimes found as " mummies" on the hive bottom, sealed in propolis. The ventilation of the hive is accomplished by fanning of the wings. The colony exhibits an astonishing degree of efficiency in its ability to protect itself and the brood from excessively high inside temperatures by rapid ventilation through a relatively small opening at the entrance. FIG. 62. — Lizard incased in propolis. The guarding of the colony from intruders is interesting and of great importance to the colony. This is done by bees which stand about the entrance and on the lower edges of the combs of the brood chamber. These bees usually do not remain long at this work for the guards are constantly changing. The hand may be placed right among them if the movement is slow, while a swift movement will cause them to dart out and will bring others to the entrance. The honeybee is capable of preventing the entrance of insects larger and more powerful than itself, such as wasps and bumblebees. During the summer of 1909, small yellow-jackets were especially abundant in the apiary of the Department of Agriculture, then at College Park, Maryland, and many dead ones were found daily in front of the hives. Numerous large wasps with hard chitinous covering are also killed by the bees. The bee-moth in some way often succeeds in entering the hive but usually the eggs or larvse are removed before any harm is done. Their success probably depends upon their habit of flying by night. 118 Beekeeping Of all these labors which the workers perform within the hive, none of them monopolizes the time of certain individuals as completely as does comb building, in which the bees hang in curtains from the comb support. Casteel has shown that even in this the bees change their duties frequently. Bees are constantly changing from guards to feeders of the brood or from ventilators to cleaners, and yet the work of the hive is done well and, one is almost tempted to say, systematically. The labor outside the hive. While the division of the inside duties may be explained to a certain degree, the division of the outside work presents problems of far greater perplexity, chiefly because of difficulty of observation. That there is an order to this work is an inevitable conclusion, but how this order is brought about among the thousands of field workers is not easily determined. Bees go to the field to obtain nectar, pollen, water and propolis. If there were no " system," we should expect to find colonies lacking one or more of these substances in sufficient quantity or, perchance, a colony with the brood nest choked with pollen or a hive over-propolized. There are, in fact, variations in all these things, but there are no cases which can be considered abnormal. Furthermore, on the grounds of an apiary of 200 colonies may be found heads of white clover or other nectar-secreting flowers right at hand. The bees in any case are not falling over each other to reacfi a certain flower and leaving other flowers untouched, as would be the case sometimes if bees were guided to nectar merely by the chance sight of a flower. Or, assuming only that there is a system whereby the individual colony divides up the surrounding territory, there would be cases of conflict between bees from the various colonies in their attempts to reach the same flowers. If then we dare to assume a pre-arranged plan, it must include the entire apiary and even more, all the apiaries within the range of flight. While bees get nectar from the flowers right beside the hive, they are no more numerous on The Life of the Individual 119 such flowers than on other flowers a quarter or half mile away. On one occasion, the author watched a head of white clover within two feet of a hive entrance. This flower was without a visitor for so long that it was almost concluded that there must be no nectar in it. All this time hundreds of bees were flying to and from the hive, many of them passing within six inches of the flower. Finally, a bee flew from the entrance directly to this flower and worked for a considerable time, sucking nectar, and, evidently getting a sufficient quantity after a time, it returned to the hive. That there was considerable nectar present in this flower is shown by the fact that other visits were made to this flower within the next half-hour from the same hive. At no time, in an hour's observation, were two bees on the head at once. Furthermore, when a bee flies from the hive, the flight is usually not uncertain but is directed toward a source of supply. It is usually stated that bees carry either nectar or pollen back to the hive but not both, but this is not correct. It may perhaps be stated that they usually gather from one species only on any given trip.1 Some additional 1 This feature is of the highest importance in a consideration of the value of the bee in the cross-pollination of plants. Since the trips are usually confined to one species, the beneficial results are increased many fold, for if they wandered promiscuously from one to the other species they would thereby scatter pollen where it would be ineffectual. That they fail to discriminate among various varieties may be considered as not a misfortune since certain varieties are pollinated better with pollen from another variety. Bulman (1902, The constancy of the bee, Zoologist, Ser. 4, VI, pp. 220-222) quotes from various authors to the effect that bees keep to one species on a single trip from the hive, and even "as long as they can, before going to another species" (Darwin, Fertilization of Plants, p. 415). This constancy is considered most highly developed in the honeybee but is claimed for certain Diptera (Bennett, Proc. Linn. Soc. Zool. XVII, p. 184). Ord (1897, The constancy of the bee, Trans. nat. hist. soc. Glasgow, n. s., V, Pt. 1, pp. 85-88) undertook to examine this as "one of the great pillars of the Law of Natural Selection" and finds that "only about 30% have proved inconstant while they were under my eye. ... In most cases when I was able to follow the bee for any considerable time, I found that, sooner or later, a change was made." He then records numerous observations which show inconstancy in a marked degree as from Leguminosse to 120 Beekeeping facts concerning the gathering of bees are of interest. If honey is exposed where it is accessible to bees, they go to it by the hundreds, if there is no nectar in the field, and under these circumstances they are on the lookout for openings in other hives so that they can rob. On the other hand, during a nectar-flow honey may sometimes be exposed in the apiary without a bee coining near it.1 This leads sometimes to the conclusion that bees prefer nectar to honey. Even if honey is placed in a feeder inside the hive, it is often not touched during a heavy nectar-flow. Division of labor in gathering. There has been little done on the division of labor outside the hive but Bonnier2 has written a paper of great interest on this subject. Whether his conclusions may be accepted must depend upon future experiments, but a resum6 of his paper is of interest. The field bees are divided by him into two classes, searchers and collectors. Searchers fly to various plants, gathering some nectar and some pollen and lighting on many neighboring objects, and behave much as do wasps, which are generally searchers. A bee is transformed Primulaceae or Composites or from yellow flowers to pink, white or purple. He concludes that the majority of bees are constant, but if watched long enough they are by no means so, that "few bees appear to be able to withstand the temptation of a Garden," where a variety of plants present themselves, and that "the Hive-bee appeared to be fully as inconstant as the wild Humble-bees." Bulman gives records of 48 observations on honeybees in a garden which were inconstant. That bees go from white clover (Trifolium repens) to alsike clover (T. hybridum) or to two species of another genus which are perhaps less readily distinguishable to an untrained human eye should not excite wonder. All that can be claimed from the known facts concerning the so-called constancy of the honeybee is that if enough flowers of one kind are easily accessible, they seem to prefer those of one kind. They usually do not fly from dandelion to apple blossom, although Ord records one such case. No more than this is needed to make bees more beneficial to the fruit-grower than they would be if their visits were entirely promiscuous. 1 Zander, Enoch, 1913. Das Geruchsvermogen der Bienen. Biol. Centralbl., XXXIII, pp. 711-716. 2 Bonnier, Gaston, 1906. Sur la division du travail chez les abeilles. Comptes rendus hebdomadaires des seances de I'academie des sciences, CXLIII, pp. 941-946. The Life of the Individual 121 from a searcher to a collector when a suitable source of nectar or pollen is discovered, and other bees come to the same source. During a good honey-flow, searchers are sent out only in the early morning and soon all become collectors (which may account for the lack of robbing and the indifference to honey about the apiary at such times) but during a dearth of nectar, searchers are out all day. Bonnier further claims that bees " commanded" to collect either nectar, water, pollen or propolis do not leave their work and will not stop even to collect honey placed in front of them. This claim is supported by experiments. The following translation of a portion of the paper cannot well be summarized: — ". . . I shall cite the following which shows . . , how the division of labor among bees of the same hive is organized and so a sort of tacit understanding, which is manifested among bees of different hives. I detached six branches of flowers of Lycium, each having about the same number of open flowers. I put each branch in a bottle filled with water. On placing these bottles in the same place from which I had taken the branches, I saw that the workers continued to visit the flowers of the branches put in water just the same as those on branches not detached from the plant. This verified, I carried the six bottles containing the branches to the fruit garden, September first, away from all nectar-bearing plants, consequently to a new place for the bees. I remained constantly watching the six bottles containing Lycium branches. No bees came to visit the flowers on these branches. The next day I saw the first bee as a searcher, which discovered them. She inspected all the branches and took some nectar and pollen; I marked her on the back with talc colored red. In about three minutes she returned to the hive. "Five minutes afterward the same first bee (which I call' A'), as shown by the red mark, came back with another and the two bees as collectors undertook a methodic visit to the branches, one to collect nectar and the other pollen. I call the second bee *B' and marked her white. 122 Beekeeping " Ten minutes after, there were three visiting bees. A new one 'C/ which I marked green, came to join the other two from the same hive, as I verified. "Later the same three workers, A, B, C, A and C always collecting nectar, and B only pollen, came back regularly to the flowering branches and visited them in the same order. All the next day these same three bees, A, B, and C, visited the six branches. "I then asked myself why other bees of the same hive or of other hives did not come to collect from these branches, as well as the three bees. Remaining under the branches, I observed attentively what took place on the second day. Early in the morning and several times in the forenoon, once in the afternoon, other searchers came to the branches of the flowers and each of these searching bees did the same thing as A. She observed the collectors with great care, their number, their manner of work, and, after two to four minutes of inspection, she flew away and did not come back. It seems that these bees, finding the place occupied, and the number of collectors sufficient for the small amount to be collected went elsewhere to search. "The fact is that the day after, I saw more and the same bees, A, B, and C, continued to visit the six branches in the same manner, A and C always for nectar, and B for pollen. "Then I replaced the six flowering branches of Lycium with twelve, branches which appeared to me about the same ; I saw two new recruits arrive, *D' and *E/ which I marked differently with colored powder; ten minutes after, two others, VJT fVDT FIG. 73. — Dorsal view of ventral walls and internal skeleton of worker. Much enlarged. iSnodgrass, R. E., 1909. Proc. U. S. Nat. Mus., XXXVI, pp. 511-595. 140 Beekeeping Oc, each side (pleural plates, PIS and pig). The tergum (dorsal plate) of the first abdominal segment (IT) is fixed to the metathorax. The posterior portion of this segment (propodium or median segment) forms the peduncle (Pd) to which the functional abdomen is attached. The legs and wings are discussed under organs of locomotion (p. 154). FIG. 74. — Thorax of worker, showing propodium or first abdominal segment (IT). Abdomen. The abdomen of the female bees (queens and workers) appears to consist of six segments (Fig. 75) but to this number must be added the modified abdominal segment on the thorax (IT). In the drones, there are additional segments partially visible externally. Snodgrass has figured (Fig. 76) the tip of the abdomen of the worker, showing that the eighth abdominal segment is in-vaginated and the eighth abdominal spiracle opens within the invagination. Zander1 further claims that the quadrate plate (Qd) is a part of the ninth tergum. The anal opening (An) is in a FIG. 75.- Lateral view of abdomen of worker. 1 Zander, Enoch, 1899. Beitrage zur Morphologie des Stachelapparates der Hymenopteren. Zeit. fur wiss. Zoologie, LXVII, pp. 288-333. The Life Processes of the Individual 141 tube (without chitin) which represents the tenth abdominal segment (X) so that the bee, like most other insects, has ten abdominal segments. In the drone, nine of these segments are partially visible. The plates of the abdomen are easily movable, being connected by membranes so that the abdomen may be distended by food, or in the queen by the growth of the ovaries. In the typical segments (II-VII), there is a tergum (T) covering the dorsal and lateral surfaces, overlapping a sternum (S) or ventral piece. The spiracles (see p. 151) (Sp) are on the terga (see Figs. 73 and 75). The eighth, ninth and tenth segments of the drone are not typical. The tergum of the eighth segment (Fig. 93, Z>, VIIIT) is partly covered by that of the seventh and carries the most posterior of the spiracles (Sp). The sternum of this segment (VII IS) is likewise concealed on the ventral side. The dorsal portion of the ninth segment is chitinized to but a small extent but the ventral portion (IXS) is chitinized and carries two pairs of clasping organs, used during copulation (iClsp and 2Clsp). The penis is extruded during mating between the clasping organs. AC*. FIG. 76. — Tip of abdomen of worker with left side removed, showing normal position of sting and anus. DIGESTION The workers take in food not only for their own nourishment but also that they may be able to provide food for the larvae. The queen eats frequently, especially during the period of active egg-laying, and a rapid metabolism 142 Beekeeping must take place to permit her to produce the large number of eggs which she lays in the height of her activities. The larvse, as has been explained in the previous chapter, take enormous quantities of food, given them by the worker bees, permitting the rapid growth during the short period of larval development. The food of the various members of the colony all comes from nectar and pollen. The workers eat honey and pollen for their own nourishment but modify the raw materials before feeding the larvee. They also normally feed the queen and the drones, but the composition of the material furnished is not determined. That the raw materials may serve their purpose, they must be so modified that they may pass through the walls of the alimentary canal and then remain in a soluble condition in the blood until taken up by the tissues. To accomplish this, various digestive enzymes are needed. The source of these will be discussed later. The digestive processes of the bee are not thoroughly understood. The usual discussions, which are abundantly numerous in spite of our lack of knowledge, are too often confined to the drawing of analogies with human digestion. No such analogies are permissible and it is, for example, entirely unwarranted to apply the name "chyle stomach" to the ventriculus, because of a supposed homology with human intestinal digestion. The whole structure of thie insect alimentary canal is different from that of man and it is, in fact, better not to apply names to any of the parts which are drawn from human anatomy. It is perhaps permissible to use the terms mouth, oesophagus and anus for both insects and man, but to call the ventriculus the chyle stomach or the rectal ampulla the large intestine is misleading. These parts do not seem to have homologous functions in man and bees. The structure of the alimentary canal has been well described by Snodgrass and by other workers and in so far as a knowledge of anatomy is helpful there is little room The Life Processes of the Individual 143 for criticism. There have been, however, very few investigations of the digestive processes. The digestion of insects is discussed by Biedermann,1 and more recently Petersen2 has published a discussion of the processes of digestion in the honeybee. This is the first good paper on this subject and the author is to be commended for taking a stand against the making of comparisons with human physiology. The mouth parts have already been described (p. 135), and those glands (1GI, Fig. 59) which are supposed to be concerned in the production of larval food have also been discussed (p. 111). Behind the mouth is an enlargement of the alimentary canal called the pharynx (Phy, Fig. 60), leading to a long narrow tube extending through the thorax, the oesophagus (CE). Behind the constriction between the thorax and abdomen, the alimentary canal widens to form the honey-stomach (HS), homologous to the crop of other insects. This is a thin-walled, muscular organ used by the worker in carrying nectar to the hive. Behind the honey-stomach is a valvular structure, the proventriculus (Pvent, the anterior part being often called the stomach-mouth) which separates the honey-stomach from the ventriculus (often called the chyle stomach). The proventriculus is of special interest in the bee, since when closed it prevents the nectar from mixing with the contents of the ventriculus and makes it possible for the honey-stomach to function as a carrying vessel. It is claimed by Schonfeld that the anterior end of the proventricular valve (nn, Fig. 61) may be moved forward to touch the posterior end of the oesophagus, so that the contents of the ventriculus may be forced out as larval food. Snodgrass has shown that this cannot happen without tearing the muscles of the 1 In Winterstein's Handbuch der vergleichenden Physiologic, vol. 2, Heft I. 2 Petersen, Hans, 1912. Beitrage zur vergleichenden Physiologic der Verdauung. V. Die Verdauung der Honigbiene. Pfliigers' Arch, fur die gesammte Phys. d. Menschen u. d. Tiere, XLV, pp. 121-151. 144 Beekeeping honey-stomach, and furthermore, as is shown later, the Contents of the ventriculus could not escape were this contortion possible. Cheshire claims that the hooks (at nny Fig. 61) of the pro ventriculus serve to separate the honey and pollen in the honey-stomach, but no proof is presented. The only known function of the proventriculus is that of opening to allow food to pass to the ventriculus. There is no evidence that it assists in the mastication of pollen. Behind the proventriculus is the ventriculus (Vent), a thick-walled organ, ringed by numerous constrictions. It consists (Fig. 77) of longitudinal and transverse muscles surrounding a much folded epithelium, the cells of which are supposed to produce some of the digestive enzymes. The inner depressions of this epithelium are filled with a gelatinous mass (pp) which extends into the lumen. The food contents of the ventriculus are surrounded by layers of membrane (peritrophic layers, Fig. 77, Pm&), formed from the gelatinous mass in the enveloping epithelium. The peritrophic layers are often described as chitinous but this, according to Petersen, is an error. These membranes do not seem capable of allowing the passage of the food contained in them to the ventriculus wall and probably little or no absorption of food occurs here. Furthermore, pollen is usually found at the posterior end undigested and, according to Snodgrass, it is not prepared for absorption until it reaches the rectal ampulla. Snodgrass observed in certain parts of the ventriculus wall a sloughing off of the ends of the epithelial cells (Enz), presumably enzymes, which are seen in the gelatinous peritrophic mass. Petersen, in confirmation of this view, found that the peritrophic layers (which come from the peritrophic mass on the epithelium) are not chitinous but contain proteolytic ferments. The peritrophic layers and their attachment to the pro-ventricular valve effectually prevent the regurgitation of the contents of the ventriculus into the honey-stomach. The contents of the ventriculus does not at all resemble larval food. It therefore appears clear that the theory that The Life Processes of the Individual 145 LMd, TMcl Epth TMcl iMcI FIG. 77. — Histological details of alimentary canal of worker: A, cross-section of ventriculus; B> section of ventriculus wall; C, section of Malpighian tubule; D, cross-section of small intestine; E, section of ventriculus wall, showing formation of enzyme cells; F, section of anterior end of rectum, showing rectal glands (RGl); G, slightly oblique section of posterior end of ventriculus, showing openings of Malpighian tubules. larval food is regurgitated can no longer be considered as at all tenable. Behind the ventriculus, the alimentary canal narrows to form the small intestine (Fig. 60, SInt) and at the point 146 Beekeeping where the ventriculus and small intestine join, the Mal-pighian tubes (Mal) empty into the alimentary canal. The small intestine is coiled and finally empties into the rectal ampulla (Rect, rectum or large intestine). The inner epithelium of the rectum is thrown into six longitudinal folds, the so-called rectal glands (Fig. 77, RGl) of unknown function. It is usually believed that they increase the absorbing surface but, since they are covered on the inner surface with chitin, this explanation seems improbable. The rectal ampulla is capable of considerable expansion and normally retains the feces when bees are confined to the hive, as in winter. It is supposed that most of the absorption of food takes place in the hind-intestine. Peter-sen advances the fantastic theory that the rectal glands are the source of the hive odor. The food of bees which must be acted upon by the various digestive juices to be prepared for absorption have their origin in the nectar and pollen collected from flowers. The chief food is honey, which consists largely of invert sugar. This name is given to a mixture of two sugars, levulose and dextrose, which by various means can be made from sucrose (cane sugar). In the higher animals, these sugars are capable of absorption without further change, and this is presumably true of bees also. The sugar in nectar is probably sucrose with some invert sugar. The preparation of this material for absorption therefore begins with the ripening of nectar into honey. As explained earlier (p. 85), this is by the action of an enzyme and, according to Petersen, such an enzyme was extracted from the head of the bee by Erlenmeyer and by v. Planta, presumably from the salivary glands. The absorption of sugars probably occurs in the ventriculus and any water in the honey which is not needed is ejected. The process of such an ejection is not clear. It is also stated by Petersen that tjie bee produces a diastatie ferment by which the digestion of starch is possible, but he was never able to prove from experiments that starch is changed into dextrin, maltose or dextrose in the honey- The Life Processes of the Individual 147 stomach. However; pollen contains no starch so that the breaking down of starch plays a small part in the digestion of the bee. The next important constituent of bee food is proteid, derived from pollen. Petersen shows the presence of pro-teolytic ferments in the salivary gland secretions and especially claims that the layers of peritrophic membrane consist largely of such ferments. He also makes the interesting observation that the bee is incapable of digesting the proteid from pollen unless the grains are broken before they enter the ventrieulus, any ones remaining unbroken simply passing out in the feces. It would therefore seem probable that the pollen is surrounded in the ventrieulus with the peritrophic layers containing the ferments and is then passed on to the small intestine without being broken up or absorbed. The use of pollen is more in evidence during brood-rearing and it is usually assumed that the workers predigest this, or perhaps more correctly, secrete a mixture rich in protein for the use of the larvae. However, since the mid-intestine of the older larvae contain considerable pollen, a large part of their proteid digestion is by their own proteolytic ferments. Pollen contains considerable oil but Petersen failed to find that any of it is digested by the bee, at any rate most of it passes through without being broken up or absorbed. The retention of feces by bees, so long as they stay in the hive, except when dysentery develops, is of importance in their management. The relation of this retention to the activities of bees in winter is discussed in a previous chapter (p. 91). Usually during the active season, when feces accumulate most rapidly, there are frequent opportunities for flight and the ejection of feces. For bees living only on honey or perhaps on a syrup of cane sugar, digestion is reduced to a minimum. It remains to be proved whether under such conditions the bees are fully nourished. That bees can live over winter without pollen is of course not proof that they do not need it then. 148 Beekeeping While bees can convert sucrose (cane sugar) into levulose and dextrose and can digest maltose, they cannot digest certain other sugars. There is also considerable evidence that dextrine cannot be digested and that the presence in the food of unusual amounts of dextrine may produce the condition known as dysentery. It has also been found that certain proteids which have been used as substitutes for pollen cannot be digested. The alimentary canal of the bee, therefore, appears to be a highly specialized system, incapable of any considerable flexibility. Bees would evidently fail to be nourished by the mixed diets of many other species, which is additional argument against attempted homologies with human digestion. CIRCULATION When the products of digestion are absorbed and traverse the alimentary canal wall, they must be carried to the various tissues for assimilation. This is done by means of the Wood. In the higher animals blood is normally confined in blood vessels which carry it throughout the body, but in the bee, as in other insects, the blood bathes the various organs, filling up the interstices between them. These spaces may, however, be so arranged that the blood flows in definite channels or sinuses. The blood is further confined to definite paths by membranes stretched across the dorsal and ventral walls of the abdomen (DDph and VDph, Fig. 78) which bound the chief sinuses. These diaphragms have a rhythmical motion and assist in the circulation of the blood. The heart (Hi) is located dorsal to the dorsal diaphragm, this sinus being therefore known as the pericardial chamber. The heart is a long muscular tube consisting of four chambers lying in the third, fourth, fifth and sixth segments of the abdomen. In each of these segments is a valvular opening (ostium, Ost) on each side for the admission of blood from the pericardial chamber, and there are also segmental valves to prevent a backward The Life Processes of the Individual 149 flow of blood. The posterior end of the heart is closed but on the anterior end it is continued in a long tube (aorta, Ao) extending in various convolutions and arches through 150 Beekeeping the thorax and opening by simple branches into the head cavity. The blood of the bee is a colorless liquid containing certain corpuscles, but no red ones such as are found in mammals. The blood is forced through the heart and aorta to the head cavity. It then flows backward through the sinuses of the thorax into the ventral sinus of the abdomen. Pumped backward by the pulsation of the ventral diaphragm, it flows through various definite cavities between visceral organs in the abdomen and into the pericardial cavity, from which it again enters the heart through the ostia. In its passage through the sinuses about the viscera, the blood takes up the food which has passed through the walls of the alimentary canal. This nourishment is promptly carried to all parts of the body by the circulation. METABOLISM It is not proposed at this time to enter into a long discussion of the ways by which each organ is rebuilt as needed. In the general discussion of the cells which make up the various organs (p. 94), it was stated that each cell is capable of taking up nourishment and of building this into protoplasm. It also utilizes oxygen furnished by the respiratory processes. To this process the name anabolism is given. Not all cells require the same constituents of the food presented or the same amount of oxygen, but by some mysterious process each cell is enabled to choose those parts which it needs. Similarly, as the activities of the cells progress, protoplasm is broken down and waste products are formed: this we know as katabolism. The final products of katabolism are carbon dioxid and water, together with various more complex chemical compounds usually containing nitrogen,, such as uric acid and urea. The elimination of the more complex waste products is discussed under excretion. The Life Processes of the Individual 151 KESPIRATION That an animal may live, it must have oxygen. The oxygen taken into the body in respiration does not go to form protoplasm in the various cells but it is used to combine with the products of katabolism to make simpler compounds which can be eliminated from the body. These products of the breaking down of the living substance are of such a character that they poison the cells unless they are promptly removed. The process is like ordinary combustion in that these products combine with oxygen to form carbon dioxid and water and to generate heat. In man, the oxygen is taken into the lungs 'and the blood is pumped there to meet the oxygen. But the bee does not have a closed circulation which will effectually carry the blood to the oxygen. Furthermore, the higher, animals have in their red blood corpuscles a substance, haemoglobin, which is capable of absorbing abundant oxygen, but this is lacking in the colorless blood of insects. In the bee, instead of the blood being carried to the oxygen, the oxygen is carried to the blood by means of tracheal sacs and a multitude of tracheal branches which go to every organ and to every part of the bee's body. These tracheae receive their air supply through openings in the outer wall, the spiracles, two pairs on the sides of the thorax and eight pairs on the abdomen. The tracheae are composed of a delicate epithelium lined with a thin layer of chitin. To prevent the collapse of the tracheal trunks, some of them are further strengthened with spirally placed rings of chitin, which are thickenings of the chitin lining. The finer branches lack these chitin rings and there are few heavy trunks in the bee, the walls usually being delicate. The oxygen is therefore carried to all parts of the bee's body, passes through the walls of the tracheal system, is absorbed by the blood and is carried to every cell. The products of katabolism are in turn carried by the blood, and the water vapor (at least most of it) and the carbon 152 Beekeeping dioxid enter the tracheal branches and are expelled through the spiracles. The more complex compounds are eliminated in the process of excretion. The tracheal system of the bee is shown sufficiently in the accompanying illustration (Fig. 79) so that a detailed description is unnecessary. The abdomen contains the unusually large tracheal sacs (TraSc) connected with each other by ventral commissures (TraCom). They are also connected with tracheal sacs of the thorax. From the most anterior spiracles of the thorax are heavy trunks to the air sacs of the head, above the brain. The tracheal system of the bee is more elaborate than that of most other insects and prob-ably in no other species is there more free access of oxygen to all parts of the body. The pumping of the •TraCcro Fio. 79. — Tracheal system of worker with dorsal sacs and trunks removed, from above. air through the body is accomplished by the respiratory movements of the abdomen, consisting of a lengthening and shortening of the abdomen and a slight dorso-ventraL movement. The muscles of the abdomen which function The Life Processes of the Individual 153 in respiration have been described by Car let,1 who distinguishes seven sets. It is stated by Djathchenko2 that in expiration the spiracles are momentarily closed, the contraction of the muscles thus forcing air to the minute branches. The spiracles are then opened and the air is expelled. EXCRETION The products of the breaking down of protoplasm consist, as previously stated, of carbon dioxid, water and various compounds containing nitrogen. Since the adult honeybee can live for long periods (especially in winter) on pure sugar, excretion must at times be reduced to a minimum. Sugar breaks down into carbon dioxid and water, both of which may pass off as gases through the tracheal system. It is only the other components of honey and the pollen which ultimately go to form nitrogenous compounds. The carbon dioxid is all expelled through the tracheae and probably most of the water escapes as vapor by this course, although some may be ejected with the nitrogenous compounds. The excretory organs are the Malpighian tubules (Fig. 60, MaJ), about 100 in number in the bee, which open into the alimentary canal at the junction of the ventriculus and the intestine. They are long delicate tubes which coil about the other viscera. These tubules are only one cell in thickness and the ends of these cells (Epih, Fig. 77, C) often bulge into the cavity of the tubule. The junction of the Malpighian tubules with the intestine is shown in Fig. 77, (?. Minute crystals of urates have been found in the Malpighian tubules. The excreted products empty into the intestine and are expelled in the feces. In the fat body (located in the body cavity) are found certain large cells of rather mysterious function, called 1 Carlet, G., 1884. Sur les muscles de I'abdomen de 1'abeille. Comptes rendues de 1'Acad. des Sci. de Paris, XCVIII, pp. 758-759. 2 Djathchenko, Sophie, 1906. Zur Frage der Athumsorgane der Biene, Ann. de 1'Inst. agron. de Moscou, XII, pp. 1-14. 154 Beekeeping oenocytes. Kosehevnikov1 states that the cenocytes of the young adult bee have a uniform, slightly pigmented protoplasm, while in old bees yellow granules begin to appear in these cells. After the winter confinement, these granules are numerous and in old queens they are especially abundant. According to the view of this author, oenocytes are excreting cells which take up waste products of katabo-lism and, after modifying them, deliver them again to the blood to be carried to the Malpighian tubules. The changes of age may be interpreted as due to an accumulation of these products in cells which are no longer able to discharge them. This failure of the oenocytes should be investigated from the point of view of the term of life of the bee. In some of the primitive insects the fat body is supposed to function as a permanent storage for urates. LOCOMOTION Bees are able to go from place to place by means of two systems of locomotor organs, the wings and the legs. Both of these are attached to the thorax and the muscles of flight are so well developed that they occupy almost the entire space in the thorax. The wings (Fig. 80) are membranous structures with a definite framework of veins attached to the sides of the thorax. As previously explained (p. 99) they are not primary embryonic appendages, but are secondary outgrowths from the second and third thoracic segments. The details of the venation of the wings need not be considered at length. This has been investigated in a careful manner by Comstock and Needham2 and the designations used in Fig. 80 are those decided upon by these authors after a study of the comparative venation of the various orders of insects. The symbols are explained in the appendix. The 1 Kosehevnikov, G. A., 1900. Ueber den FettkSrper und die (Enocyten der Honigbiene (Apis mellifera L.). Zool. Anz., XXIII, pp. 337-353. 2 Comstock, J. H., and Needham, J. G., 1898-99. The wings of insects. Am. Nat., XXXII and XXXIII: Reprinted, Ithaca, N. Y. The Life Processes of the Individual 155 attachment of the bee's wings to the thorax has been investigated by Snodgrass (I.e. pp. 61-63). The motion of the wings in flight is in four directions, up, down, forward and backward, and the combination of these movements causes the wing tips to describe the course of a figure 8, if the insect is held stationary. In flight, the 8 is of course modified. The hind wings are small and are attached by hooks on their anterior margins to thickenings on the margin of the front wings. They are not provided with large flight muscles of their own but are carried along by the action of the powerful muscles in the mesothorax which propel the fore * wings. The muscles of flight are in four sets, corresponding to the four directions of wing movement. The chief muscles are not attached directly to the bases of the wings, as in dragonflies, but the FIG. 80. — Fore and hind wings. wings are moved into the right position by muscles situated inside the pleura of the two thoracic segments. After the wings are in position for flight, the compression of the thorax by the vertical muscles lowers the dorsum and raises the wing while the contraction of the longitudinal muscles raises the dorsum and lowers the wing. The vertical muscles are therefore the elevators and the longitudinal muscles the [depressors. The movements of the wings during flight is therefore produced mainly by changes in the shape of the thorax. The forward and backward movements are accomplished by the action of the muscles on the pleurum, acting directly on the bases of the wings. Because of the enormous development of the two main sets of flight muscles, bees are capable of strong and rapid flight. They are also capable of arresting progress suddenly FIG. 81. —A, left front leg of worker, anterior view; B, spine of antenna cleaner; C, details of antenna cleaner; D, left middle leg of worker, anterior view; E, left hind leg of queen, anterior view; F, left hind leg of worker, anterior view; G, inner view of left hind leg of worker, showing pollen-combs; H, left hind leg of drone, anterior view. 156 The Life Processes of the Individual 157 or of getting under way rapidly. There is no reason to believe that flight is in any way dependent upon the amount of air in the tracheae, as has been claimed, for filling the air sacs obviously does not reduce the weight of the bee. The maximum rate of flight is not clearly established, for the currents of air must be eliminated in making such determinations; the rapidity of movement depends largely upon the load being carried. Bees are able to fly at a considerable angle for some distance as is seen in apiaries in mountainous districts. The power of the wing muscles is shown by the ability of a worker to fly from the hive carrying a drone, which weighs more than the worker itself. In walking, bees use all six legs (Fig. 81). In addition to their function in locomotion, the legs constitute a rather complex Set Of tools for nu- FIG. 82. —Dorsal (A), ventral (£) and meroUS Other purposes, lateral (C) views of last tarsal joint . ,, i • JT of first foot of worker. especially complex in the worker. On the front legs at the articulation of the tibia and first tarsal joint are the antennae cleaners. The middle leg has a spur to which has been attributed the function of prying pollen from the hind legs in storing it. The hind legs of the worker bees are highly specialized, carrying pollen baskets or corbicula on the outer side of the flattened tibiae and rows of spines on the inner side of the first tarsal joint. Between these two joints are the so-called wax-shears, which in fact have nothing to do with the wax, but function in pollen gathering (p. 123). Each leg is provided with a pollen brush for collecting pollen. 158 ' Beekeeping The muscles for moving the legs are located inside the joints and are inserted on the chitinous walls. The last tarsal joint on each leg carries a pair of bilobed claws (Fig. 82, CJa), which differ among the three types of bees. Those of the drone are bent more nearly at right angles than those in the workers and queens and those of the queen are larger than the claws of the workers. Between the claws is a lobe (empodium, Emp) used when the bee walks on a smooth surface. On such a surface the claws are useless and the sticky empodium is lowered and flattened, providing a good foothold. The motion of the legs in walking is typical of all insects. The legs move in two sets; the fore and hind legs on one side move in the same direction as the middle leg on the opposite side, thus giving a triangle for support at all times. In flight the legs hang freely and are forced somewhat backward, except when they are being used as in the manipulation of pollen (p. 123). PROTECTIVE APPARATUS Worker bees defend the colony by means of the sting, situated usually in a cavity at the tip of the abdomen (Fig. 76) but capable of marvelously rapid action when it is protruded. As was stated earlier (p. 140), this sting cavity is formed by the infolding of the eighth, ninth and tenth segments of the abdomen. The sting is homologous with the ovipositor of other insects (see Snodgrass, I.e. pp. 76- 77) and is made up of parts considered by some embry-ologists as comparable with the legs and mouth parts of the more anterior segments of the bee. The sting of the worker bee is straight while that of the queen is longer, curved and less strongly barbed. The sting (Fig. 83) and its accessory apparatus form a rather complex structure. The shaft consists of three parts, a dorsal sheath (ShS) along which move two barbed lancets (Let). The sheath is enlarged at the anterior end into a The Life Processes of the Individual 159 StnPIp. bulb (ShB) and is further continued in two arms (ShA) which curve outward. The lancets slide on a grooved track the full length of the sheath, past the bulb and diverge along the two basal arms. The sheath and lancets combine to form a hollow tube (PsnC) through which the poison flows. The arms of the sheath are attached at their anterior ends to oblong plates (Ob) which overlap the sides of the sting. To these plates are attached palpi (StnPlp), soft white projections provided with sense organs, by means of which the bee can tell when she is in contact with the object which is to be stung. The lancets are attached to triangular plates (Tri) which in turn articulate with the quadrate plates (Qd). By the movements of these plates on each other the lancets are slid along the sheath when the sting is used. It has been shown by Zander1 that the triangular plate (Tri) is part of the eighth sternum, the quadrate plate (Qd) is part of the ninth tergum and the oblong plate (06) is the ninth sternum. "In the accessory plates of the bee's sting we have a most excellent illustration of how parts of a segment may become modified to meet the requirements of a special function, and also an example of bow nature is ever reluctant 1 Zander, Enoch, 1899. Beitrage zur Morphologic des Stachelapparates der Hymenopteren. Zeit. f. wiss. Zool., LXVI, pp. 288-333. ShA- FIG. 83. — Ventral view of sting of worker and accessory parts, flattened out. 160 Beekeeping to create any new organ, preferring rather to make over some already existing structure into something that will serve a new purpose." — SNODGRASS, I.e. p. 78. The poison of the sting arises from two sets of glands. The conspicuous poison sac (PsnSc) which opens into the bulb of the sting is usually seen attached to the sting when the sting is pulled from a bee. The contents of this gland have an acid reaction and it was formerly believed to be formic acid. This acid comes from two long coiled tubes (AGID) on which are two small enlargements, supposed to be the secreting glands (AGl). The tubes (AGID) also probably have gland cells in the walls. The other poison glands (BGl), known as the alkaline glands, also empty into the bulb of the sting. Their secretion is supposed to have an alkaline reaction. According to Carlet,1 the secretions of these two sets of glands must be mixed to be fully effective. The secretions enter the bulb where they are mixed and are then forced down the canal (PsnC) formed by the sheath and lancets. In most books on bees, certain lateral openings in the lancets are described as paths of the poison in the process of stinging. Snodgrass showed, however, that these do not connect with the poison canal and supposed them to be ducts of some kind of subcuticular glands. Mclndoo has shown them to be olfactory pores (p. 170). The sting, as every beekeeper knows, is an effective weapon of defense. When used, it usually cannot be withdrawn because of the barbs (Brb) on the lancets. The sting with the accessory plates and poison sac are therefore usually torn from the body of the bee, causing so severe an injury to the abdomen that the worker dies within a short time. The defender is thus sacrificed for the good of the colony. The parts torn away include the muscles which operate the accessory plates and indirectly slide the lancets on the sheath. The sting may therefore be driven 1 Carlet, G., 1890. M6moir sur le venin et 1'aiguillon de 1'abeille, Ann. des sci. nat., Zool., 7 ser., IX, pp. 1-17, The Life Processes of the Individual 161 farther and farther into the wound if not promptly removed and the same reflex actions of the muscles serve to force more poison from the poison sac. In removing the sting, care should be exercised not to squeeze the poison sac thus emptying its contents into the wound. CHAPTER VII THE NERVOUS SYSTEM AND THE SENSES IN order that bees may respond to factors in the environment, obviously these influences must be perceived. The organs which receive the stimuli from without are the special organs of sense. The resulting nervous impulses are then transmitted through the nervous system, by means of which also the actions of the animal are coordinated and molded in response to the stimuli received. The nervous system and its various organs of special sense are therefore of the highest importance to the animal and the influence of the stimuli of the environment are so important in the behavior of these insects as to justify a separate chapter. Nowhere in the entire discussion of bee activities is it more necessary to avoid comparisons with our own actions than here. Man is capable of conscious and volitional acts while evidence of such acts in bees is lacking. Furthermore, the structure of the nervous system and of the sense organs is so unlike analogous structures in man that attempts at homologies are entirely unwarranted. NERVOUS SYSTEM This system of organs consists of a series of nerve masses called ganglia (Fig. 84, Gng) situated on the mid-ventral line of the body, the ganglia being connected by a pair of longitudinal cords, called connectives. The nerve cells are located in the ganglia while the delicate processes from these nerve cells, the nerve fibers, form the connectives and also go to all parts of the body, some serving to trans- 162 The Nervous System and the Senses 163 mit stimuli from the sense organs and some to carry stimuli from the nervous system to the various organs of the body. The nerve fibers therefore are often compared with wires used in conducting electric energy from place to place. In an hypothetical generalized insect embryo we should doubtless find a ganglion for each segment of the body, probably twenty in all, but the ganglia of the bee larva are modified from the primitive condition and in the adult still further specialization is observed, by the fusion of various ganglia. The brain (Fig. 85), situated above the oesophagus, consists of three consecutive ganglia, recognizable in the embryo, but completely fused and not readily recognizable in the adult. From the brain, two short connec- FlG' ^"Ne °f worker' tives (circum-oesopha-geal) pass one on either side of the oesophagus to the suboesophageal ganglion (SceGng) also located in the head. Continuous with the brain are the optic lobes (Opl) forming the nervous connection with the large compound eyes (E), and from the brain are nerves to the antennae (AntNv) and also to the frontal ganglion (FtGng), from 164 Beekeeping which the stomatogastric system (sympathetic system) has its origin* (StgNv). The suboesophageal ganglion gives off nerve branches to the mandibles (MdNv), maxillae (MxNv) and labium (LbNv). For a study of the minute structure of the brain and the paths of the various nervous elements, the reader is referred to the works of Kenyon l and Jonescu.2 OpLv FIQ. 85. — Brain and suboasophageal ganglion of worker, anterior view. In the thorax the number of ganglia is reduced to two (Figs. 78 and 84, IGng and 2Gng). The first innervates the first pair of legs while the second is a combination of four ganglia, as shown by the fact that it innervates the meso- 1 Kenyon, F. C., 1896. The brain of the bee. Jr. comp. neurol., VI, pp. 133-210. ——, 1897. The optic lobes of the bee's brain in the light of recent neurological methods. Am. nat., XXXI, pp. 369-376. 2 Jonescu, C. N., 1909. Vergleichende Untersuchungen iiber das Gehirn der Honigbiene. Jenaisqhen Zeit. f. Naturwiss., XLV, N. F., XXXVIII. The Nervous System and the Senses 165 and metathoracic segments (with the corresponding two pairs of legs and wings) and the first abdominal segment, which is fused with the thorax in the bee, as well as the first segment of the abdomen behind the constriction. It should be noted that nerves (W2Nv and WSNv) run to the bases of the wings to innervate sense organs (p. 170). In the abdomen are five ganglia (8-7Gng) which send nerve branches to the remaining abdominal segments. The third and fourth ganglia lie one segment in front of the segments which they innervate while the remaining ones are in their own segments, the last (7Gng) supplying the remaining posterior segments of the abdomen, it therefore being actually a fusion of four ganglia. The action of the nervous elements remains a matter chiefly of conjecture. These cells have lost their contractility and probably never regenerate nor divide in the adult bee. Their function is obviously important, for if this system is injured the coordination of the body is destroyed. However, the cutting of the nerve cord does not cause death and even if the thorax and abdomen are entirely separated the parts may function independently. If the head is removed, the animal can still walk and if the abdomen is removed it can still take in food. These facts indicate that the nervous control of the body is not centralized in the brain as completely as in man and in many other animals. Proper correlation of movement cannot, however, take place unless the nervous connections are intact. SENSE ORGANS So little is known of the structure and function of the sense organs of bees tfrat this subject must be discussed with caution. We know that the simple and compound eyes are the organs of sight and recently it has been found where the organs of smell are located. Beyond this is a vast field for investigation and a fertile field for speculation. 166 Beekeeping Sight. The organs which receive light stimuli are the three simple eyes or ocelli (0) and the two large compound eyes (£), all situated on the head. The compound eyes are located on inner ch. FIG. 86. — Section of compound eye and optic lobe of worker; Om, oramatidia. the sides of the head, each eye consisting of many units. In drones, the number of these units is larger than in the two types of females and the compound eyes are so enlarged as to meet on the vertex of the head. The structure of the units of the compound eye was described some years ago.1 On * Phillips, E. F., 1905. Structure and development of the compound eye of the honey bee. Proc. acad. nat. scl Phila., LVII, pp. 123-157. The Nervous System and the Senses 167 the outer surface, these units (ommatidia) are indicated by hexagonal facets in the chitinous covering of the eye. If a section is cut through the entire eye of a worker bee (including the optic lobes), the structure is that shown in Fig. 86. Numerous ommatidia are shown in full length and beneath these are the optic lobes, which need not be described here. An examination of a single ommatidium (Fig. 87) shows the following details of structure: (1) an outer corneal lens of chitin (CL) continuous with the chitin of the head, (2) the crystalline cone (CC) and (3) the rhabdome (rhb) surrounded by eight or nine sense cells or retinulse (ret). Surrounding the ommatidia are two types of pigment cells, (1) the corneal pigment cells (c.-p.c.), which in the pupal stage secreted the chitin of the corneal lens, and (2) the outer pigment cells (o.-p.c.). So far as can be determined, the functions of these parts are as follows: rays of light pass through the lens and crystalline cone cells and enter the transparent rhabdome where the stimulus is received. Any rays of light which enter obliquely or which strike the edge of the crystalline cone are absorbed by the surrounding pigment cells so that it seems probable that only those rays which strike the surface of the eye at a right angle ever reach the sense cells. J1-BM There is no apparatus for changing the focus of the lens. The type of image formed by the com- FIG. 87.-~ Section , , i . » of entire omma- pound eye has been the subject ot con- tidium. siderable speculation. The two theorites on this subject are (1) that each facet forms a separate 168 Beekeeping image1 and (2) that the impressions of the individual facets form a mosaic image.2 The latter theory has most to support it and is generally accepted. Forel3 gives an admirable discussion of these theories and adds considerable evidence to support the latter theory. As was stated earlier, it is probable that only the rays of light which strike the lens perpendicularly can reach the sensory cells. The image is probably not a distinct one. If an object in motion is within the range of vision of the bee, the image is transferred rapidly from one set of ommatidia to another, which probably accounts for the fact that bees perceive objects in motion more readily than they do still ones. In addition to the compound eyes, there are three simple eyes or ocelli (0), which Grenadier4 states are derived from the same primitive organ a& the individual ommatidia of the compound eyes; in fact, as shown by him and by Forel, compound eyes in some species are replaced by ocelli. The parts played by the ocelli and by the compound eyes in the vision of the bee are not clear. From a study of the angles of refraction, it has been inferred that the ocelli are for perceiving near-by objects, while the compound eyes are far-sighted. However, just the reverse has been claimed, and we have no reliable data on this subject. It has been shown by numerous experiments and by the experience of beekeepers that bees perceive differences in 1 Gottsche, C. M., 1852. Beitrag zur Anatomie u. Physiologie des Auges der Fliegen u. s. w. Miiller's Archiv. f. Anat. 2 Exner, Sigmund, 1875. Ueber das Sehen von Bewegungen und die Theorie des zusammengesetzten Auges. Sitzb. des K. Akad. der Wissensch., LXXII, Abth. III. This theory goes back to the work of J. Miiller, 1826. Zur vergleichenden Physiologie des Gesichtsinnes. Leipzig. 8 Forel, Auguste, 1908. The senses of insects. Eng. trans. by Yearsley. London: Methuen and Co. 4Grenaeher, H., 1874. Zur Morphologic und Physiologie des facet-tirten Arthropodenauges. Nachrichten v. d. K. Gesellsch. d. Wissensch. a. d. G. A. Univ. zu Gdttihgen, pp. 645-656. ——, 1877. Untersuchungen liber das Arthropoden-Auge. Beilageheft zu d. klinischen Monatsblattern f. Augenheilkunde, XV. Rostock. The Nervous System and the Senses 169 color. It is asserted that ants do not perceive red light and the same statement is made concerning bees, but this is incorrect for bees. It is also sometimes said that insects perceive some of the ultraviolet rays, beyond the range of human vision. The color preferences of bees have also been observed, it often being stated that they prefer blue. It seems certain that bees do not see objects distinctly and their vision is clearly far less acute than that of wasps and some other insects. Perhaps they do not perceive the form of objects at all. The relative intensity of light is probably an important part of their vision. When it is recalled that the hairs (p. 104) cover many facets of the compound eye, especially in younger bees, and that the structure of the eyes does not suggest a high degree of efficiency in vision, it becomes a matter of wonder that bees are helped by vision as much as appears to be the case. Smell. It is commonly believed that bees possess an acute sense of smell, and this belief is borne out by experiments on this subject. With the exception of qualifying statements by Lubbock * and Forel, this is usually conceded. The location of the olfactory organs is a matter of much less unanimity of opinion. Mclndoo 2 has recently performed a valuable service in gathering together the literature on the olfactory organs in insects and it is necessary only to gi^e a list of the organs which are supposed to carry the olfactory organs to show the confusion which has existed. These sense organs have been located by various authors on the following structures : (1) the spiracles, (2) organs close to the spiracles, (3) glands of head and thorax, (4) oesophagus, (5) "internal superior surface," (6) folded skin beneath antennae, (7) rhinarium, (8) plate between eyes and beneath antennae, 1 Lubbock, Sir John, 1899. The senses, instincts and intelligence of animals. Internat. Sc. Ser. London., vol. 65. * 2 Mclndoo, N. E., 1914. The olfactory sense of insects. Smithsonian misc. col. LXIII, no. 9, 63 pp. 170 Beekeeping (9) mouth cavity, (10) epipharynx, (11) palpi, (12) antennae, (13) various structures on the antennae, (14) caudal styles, (15) organs on base of wings and on legs, and (16) on different organs for different orders of insects. Notwithstanding this assortment of theories, it is probably correct to state that until recently it was the consensus of opinion that the olfactory organs are located on the antennae. However, Mclndoo1 shows that if the antennae of the honeybee are removed, the insect still reacts to odor stimuli. It is impossible to go into the details of this work here, but, hi brief, this author concludes that certain sense organs located at the bases of the wings, on the legs and on the stings of females are olfactory organs, named by him olfactory pores. His work covers not only a study of the structure and distribution of these organs but is supported by experimental evidence, which is usually omitted in other papers on this subject. The location of these organs is indicated on the diagrams from Fro. 88. — Diagram of dorsal Mclndoo's paper (Figs. 88 and view of worker, showing loca- 89), the organs being indicated Jkurf groups of olfactory by ^^ ^^ ^ ^ different groups being numbered (21 groups in all, Nos. 19, 20, and 21 being on the sting and not shown in the diagrams). The structure of a typical olfactory pore is shown in Fig. 90. From the sense cell OSC), a nerve fiber (SF) extends to the surface of the body through the pore aperture (PorAp), this aperture being 1 Mclndoo, N. E., 1914. The olfactory sense of the honey bee. Jr. exp. lool., XVI, pp. 266-346. The Nervous System and the Senses 171 within a flask (PorW) which lies in the chitinous body wall. These sense organs have protoplasm exposed to the outer air, not covered with chitin, while most of the other organs which have been supposed to have the olfactory function are covered with a chitinous layer. This is especially to be noted in the sense organs of the antennae and it is difficult to see how odors may be supposed to penetrate such layers. The structure of the olfactory pores therefore fits them for their olfactory function and Mclndoo has shown by experimental evidence that this is their office. He1 has also found these olfactory organs in spiders and in other Hymenoptera. Admitting that these olfactory pores are the true organs of smell, we are still confronted with some difficulty in deciding what part responses to odor stimuli play in the behavior of bees. That bees are attracted by odor to honey during a dearth of nectar cannot be doubted. Similarly it is believed that the recognition of hive-mates, the discovery of enemies and the reactions toward the queen are due to responses to odors. A difficulty encountered in this field of investigation is that the human sense of smell is so inefficient that it is difficult to comprehend the responses observed, 1 Mclndoo, N. E., 1911. The lyriform organs and tactile hairs of araneads. Proc. acad. nat. sc. Phila., LXIII, pp. 375-418. ——, 1914, The olfactory sense of Hymenoptera, Ibid., LXVI, pp. 294-341. FIG. 89. — Diagram of ventral view of worker, showing location of groups of olfactory pores. 172 Beekeeping which is perhaps but another way of saying that we are too prone to put human interpretations on all such observations, v. Buttel-Reepen,1 from his wide experience with bees, concludes that there are seven normal odors in a colony of bees which influence behavior. These are (1) an individual odor, (2) an odor common to the offspring of one queen, (3) brood and larval-food odor, (4) drone odor, (5) wax odor, (6) honey odor and (7) the hive odor, which is a combination of all or part of the other odors. Whether there are other normal odors is a matter of conjecture but, in cases of dysentery or a brood disease, abnormal odors occur which influence the behavior of the bees. On the dorsal side of the abdomen of the workers and queen on the articular membrane between the sixth and seventh terga (counting the propodium) is a transverse area which is the external portion of a scent-producing organ. This organ was described by Nassenoff,2 later by Sladen3 and more recently Mclndoo 4 has described the structure of the glands on the interior as well as the external structure. This organ may perhaps be considered as the source of the individual odor of the females. 1 v. Buttel-Reepen, H'., 1900. Sind die Bienen Reflex-maschinen ? Biol. Centralbl., XX; reprinted Leipzig: Georgi; Eng. trans. by Mary H. Geisler, Medina, O.: A. I. Root Co., 48 pp. * Nassenoff, see Zoubareff, A., 1883. A propos d'un organe de 1'abeille non encore decrit. Bul. d'apic. suisse rom., V, pp. 215-216. Trans. Brit, bee jr., No. 136. Nassenoff's paper is in Russian. 8 Sladen, F. W. L., 1901. A scent-producing organ in the abdomen of the bee. Gleanings in bee culture, XXIX, pp. 639-640; also in Ent. month, mag., XXXVIII, pp. 208-211. 4 Mclndoo, N. E., 1914. The scent-producing organ of the honey bee. Prod. acad. nat. sc. Phila., LXVI, pp. 542-555. FIQ. 90. — Cross-section of typical olfactory pore: *S<7, sense cell; 8F, sense fiber; PorAp pore aperture. The Nervous System and the Senses 173 If the queen is removed from a colony and a strange queen is placed among the bees in a cage, after a day or so she has, according to the current belief, acquired the hive odor and she will be accepted if released. If a strange bee attempts to enter a hive, it is usually recognized at once and repelled, this being considered as due to the possession of a different hive odor, but if a field bee returns to its own hive, it is admitted, because it has the hive odor. These responses may vary according to the honey-flow and other environmental factors. In these cases and many others, there is evidence of the importance of responses to odors in the behavior of bees, so that there is justification for FIG. 91. — Antennal organs: A, antennal believing that the sense (FFl); B, pore plates and Forel's flasks from drone's antenna; C, pore plates (PorPt), pegs (Pg) and tactile hairs (THr) from worker's antenna ; D, structure of pore plate and tactile hair; E, structure of peg; F, structure of tactile hair; G, structure of Forel's flask ; H, structure of pit peg. of smell is of primary importance. It must be admitted that the belief in this importance is based chiefly on the accumulated experiences of beekeepers rather than on careful experiments, which are sorely needed in an examination of these data in order to eliminate complicating environmental factors. Additional evidences of odor influences are given in the discussion of swarming. 174 Beekeeping Antennal sense organs. Before the work on the olfactory pores, just described, it was supposed that some of the sense organs on the antennae are olfactory organs. Just which of the organs serve in this way was not easy to decide. That these are sense organs can scarcely be doubted, but in view of the elimination of organs of smell from the antennae of bees, the only course at present is to describe these organs and leave their function to be decided by later experimental work. The accompanying illustration (Fig. 91) shows the distribution and structure of these organs. These organs are known as (1) pore plates, (2) pegs, (3) ForePs flasks, (4) pit pegs and (5) tactile hairs. In all of them the sensory cells are covered with chitin. Taste. To what extent bees have this sense has not been made clear. In human experience, the senses of taste and smell are so closely related that to determine these separately in the bee will prove a somewhat difficult task. There are sensory cells on the epipharynx, in the mouth cavity, on the palpi and perhaps on other mouth parts, some of which may prove to be organs of taste. The evidence that bees distinguish tastes is meager. It is well known that bees show preferences in the material collected. They will for example abandon honey-dew if nectar becomes plentiful, but this action may not be due to a sense of taste. Touch. This sense is probably well developed, and it is safe to assume that some of the antennal sense organs function in this way. The use of the antennae by the bees suggests this. Bees are remarkably sensitive to jars and respond promptly when touched on various parts of the body. Most of the hairs which cover the body are not sensory, however. .-.**» The Nervous System and the Senses 175 Hearing. No organ has so far been described for bees which is surely an organ of hearing nor is it definitely established that bees can hear. In experimenting on this subject, it is of course necessary that vibrations through solids be eliminated and that the stimulus come to the bee only through vibrations of the air. It is commonly believed by beekeepers that bees hear, the belief being based chiefly on the fact that bees make noises which are interpreted as purposeful. Various investigators share this belief, among whom may be mentioned v. Buttel-Reepenl (I.e.). Since this author has (pp. 12-18 Eng. trans.) gathered together the evidence on this subject, it is necessary here only to mention the various phenomena which he details. (1) Queenlessness of a strong colony is noticed in from one hour to'several hours. The bees no longer hum " contentedly," but this gives way to a "lamenting buzz." This change is said not to be due to the lack of the queen's odor, although the author admits that if a dead queen is placed in the colony the agitation ceases. (2) If a colony is made queenless and the caged queen is later placed in the upper part of the hive, the agitation ceases and v. Buttel-Reepen cannot believe that this is due to odor. (3) Bees disregard a queen in the open air a foot from the hive. From these observations, he believes that odor is not the only factor in "communication" of bees and he believes that bees communicate by sound. He further details some other evidence. (1) "It can hardly be doubted that sounds of some kind perhaps serve here [in swarming] for communication." (2) The "swarm tone" serves to draw out colonies scarcely ready to swarm. (3) The humming of bees is interpreted as leading the bees during the hiving of a swarm. 1 On p. 2 (Eng. trans.), v. Buttel-Reepen says: "No zoologist who has done any experimental beekeeping can have the least doubt that bees have an excellent sense of hearing, since observations yield him hundreds of proofs. The man who is not familiar with biological facts might recognize nothing of the kind with certainty, for up to the present the organ of hearing has not been discovered." 176 Beekeeping (4) The queen makes at least two sounds, "teeting" and "quahking." (5) When a queen is "flightened" she emits a peculiar sound. This author concludes by claiming that the fact that bees do not respond to artificial sounds is no proof of a lack of hearing. It need scarcely be pointed out that these statements are not conclusive evidence of a sense of hearing in bees; in fact most of the phenomena observed are as readily interpreted as evidence of a sense of smell. In §arlier chapters it is mentioned that the phenomena in swarming and in the hiving of a swarm are most plausibly explained as brought about by reactions to odors, v. Buttel-Reepen's statement that "only the dead bee is quiet" may be answered by the statement that a totally deaf man often makes more noise in walking and frequently by articulate sounds than does a man with acute hearing. To sum up, we are justified in concluding (1) that no organ or organs of hearing are recognized, (2) that the existence of a sense of hearing is doubtful, and (3) that the investigations so far carried out are inconclusive. Temperature sense. In the discussion of the activities of bees in winter (p. 90), it is stated that at about 57° F. the bees form a cluster and, if the outer temperature drops below that point, they begin to generate heat. When no cluster is formed the bees are more active at temperatures above 69° F. than at temperatures below this. In a discussion of the temperature of the hive at other seasons (p. 60), it is shown that the temperature of the hive while occupied by the bees rarely exceeds 97° F. and that during brood-rearing the temperature of the brood chamber is quite constant. This brief summary of the facts of hive temperature indicates that in some manner bees perceive changes in temperature and it may almost be believed that they have a temperature sense superior to our own. The nerve endings or sense organs which function in this response to temperature stimuli are not determined The Nervous System and the Senses 177 and perhaps this is a function of some of the problematical organs on the antennae. Finding of the flowers. In a previous chapter (p. 118), a discussion is given of the division of labor whereby bees are seemingly able to apportion the available forage to prevent duplication. In this connection the interesting question arises as to how the bees find the flowers. Considerable detailed and painstaking work has been done on this subject. Plateaul and his followers on the one hand believe that bees are guided to the nectar by odor, this being supported by experimental evidence as well as by an array of facts concerning the gathering of nectar from inconspicuous flowers. Forel2 and other writers assert, on the contrary, that color is the important stimulus and that flowers are found through the sense of sight. Plateau's work is open to- one important criticism, since he overlooks the possibility of the return of bees to his mutilated flowers through memory. Burton N. Gates, several years ago, showed that bees visit artificial flowers and also fly to natural flowers which have been sealed in glass tubes. The reaction to these unusual objects was entirely normal. These results point strongly to the belief that odor is of minor importance in the location of nectar- 1 Plateau, Felix, 1895-97. Comment les fleurs attirent les insects. Bul. acad. roy. d. Belgique, 3 ser., XXX, n. 11, XXXII, n. 11, XXXIII, n. 1, XXXIV, n. 9, 10, 11. See also Plateau, 1888. Recherches experimentales sur la vision chez les arthropodes, ibid., part. 3-5 and other papers. 2 Forel, Auguste, 1886-88. Recueil zoologiques Suisse, 1 ser., IV. ——, 1908. The senses' of insects. Eng. trans. Yearsley. London: Methuen and Co., 324 pp. See also: Andrese, Eug., 1903. Inwiefern werden Insekten durch Fabre und Duft der Blumen angezogen. Beihefte z. Bot. Centralbl., XV. Giltay, E., 1901. Ueber die Bedeutung der Krone bei den Bliiten und fiber das Farbenunterscheidungs vermogen der Insekten, I. Pringh. Jahrb. f. wiss. Bot., XL. Detto, Carl, 1905. Blutenbiologische Untersuchungen I u. II. Flora odor Allg. bot. Zeit., XCIV. Kienitz-Gerloff, 1898 u. 1903. Proffessor Plateau und die Blumentheorie I u. II. Biol. Centralbl., XVIII u. XXIII. 178 Beekeeping secreting plants. That bees differentiate between flowers which are encountered in their flights is shown by the fact that they usually visit but one species on a trip (p. 119). Finding of the hive. It is well known that bees normally return to the right hive. The fact that strange bees are not usually admitted may be explained on the basis of difference in colony odors but this does not explain the method by which they find the right hive in the majority of cases. Bethe 1 asserts that the bees are led back to the hive by an "unknown force" but, as v. Buttel-Reepen points out in his discussion of memory of place in bees, this explanation is not satisfactory, and cannot be accepted until the known forces are eliminated. It will be recalled (p. 105) that young bees take "play flights" on warm days. If bees which have not taken such flights are taken out a few feet from the hive, they fail to return. Bees that have had some experience on the wing are able to return from short distances, and, finally, old bees are often able to return if taken away two miles or more. They evidently increase in efficiency with experience. It is also known that if the hive is moved a foot or more in any direction the returning bees seek the entrance to the hive in the old place. If the. hive has been moved only a short distance they may soon find it by searching, but if it is moved several feet they may fail to find it. If bees were attracted to the hive by odor, the field bees would probably have no difficulty in finding it if it were moved perhaps a mile. Under such circumstances a short distance would make no appreciable difference and yet the moving of the hive a foot often delays their entering it. Odor is therefore evidently not the guiding sense. Bees in the field cannot always see their hive, and in all probability, they can see neither far nor distinctly. If 1 Bethe, A., 1898. Diirfen wir Ameisen und Bienen Psychische Quali-taten zusehreiben? Arch. f. d. ges. Phys., LXX, also as separate, 1898. Bonn: Etnil Strauss, with different paging. The Nervous System and the Senses 179 sight is their guide, they must remember various objects over or about which they fly as they go out and must return by known paths. This is actually the case. If bees are accustomed to fly in only one direction to the forage and are carried off a short distance into unknown environment, they fail to return. It is evident that bees are guided back to their hives by a memory of the objects encountered, as perceived by sight. If a hive is moved, they then follow over the accustomed paths to the old location of the entrance, but having no experience over the road from the old location to the new one, they fail to make the trip unless they accidentally encounter the hive. No "unknown force" need be called in here to explain the phenomena. Evidently the play flights and the early trips to the field are* the times during which bees acquire knowledge of their surroundings. If a colony is moved several miles, the bees must orient themselves anew, and in order that they may perceive the change and "recognize" the necessity for re-orientation, the beekeeper often places brush or grass about the entrance so that the change may be perceived when they first fly out. That sight is the important sense in the location of the hive is appreciated by beekeepers who have learned that irregularities in the rows of hives, landmarks of trees or shrubs in the apiary or differences in color of the hives are beneficial in enabling the bees to find their hives quickly. These customs are well founded on the behavior of the bees. Memory. It would appear from the preceding discussion that bees are not entirely bundles of reflexes but that they actually have memory. The finding of the hive is good evidence of this fact and it is also asserted (v. Buttel-Reepen) that they remember the location of the feeder in the hive and that scouting bees remember the paths to the locations chosen by them. The best evidence of memory is found in the fact that memory is sometimes lost. If bees are stupefied by tobacco 180 Beekeeping smoke, by the smoke of the puff ball (an old practice) or by some anesthetic, they are unable to return to their old location and must re-orient themselves after they revive. When bees swarm they usually do not again return to the location of the old hive (except when hived on the old stand by the beekeeper) and may safely be placed in a new location perhaps only a few feet from the old hive. The memory of the old location is not lost immediately, however, for if within a day or two the bees desert the new quarters they often return to the old hive. Here the old memories are, as it were, reserved, but they are lost in a short time. Similarly in artificial swarms, after drumming or after certain manipulations in which the colony becomes "demoralized/' the memory of the location is lost, either permanently or temporarily. If bees are confined for a few days they may be placed in any location and bees wintered in a cellar no longer remember their former locations. The loss of memory in these cases is not due to the formation of new associations. Bees obviously cannot lose what they do not possess and, if it is granted that memory is sometimes lost, the only conclusion is that they possess memory. Nature of bee activities. In the introduction to Chapter III, it was stated that bees are essentially creatures of instinct. While in the intervening discussions there are given evidences of the possession of memory, of limited powers of learning and association and of certain adaptations of the reactions of bees to circumstances, it should be clear that in the bee we have to do, not with human intellects and poetic passions, but with animals whose behavior is chiefly guided by mental capacities imprisoned in the chains of instinct, with animals most of whose activities are justly described as machine-like. If this discussion of the nervous responses of bees has destroyed some of the poetry of the hive, this can scarcely be considered as a serious loss, for it is not by such fancies that we can come to know the truth concerning the things about us. CHAPTER VIII THE REPRODUCTIVE PROCESSES AND PARTHENOGENESIS THE organs of reproduction are those which produce the cells from which individuals of the next generation develop and they also include the accompanying organs which serve to permit the proper disposition of the sex cells. The continuance of the species is the function of these organs. In the larger number of species, new individuals arise from eggs which have been fertilized by sex cells of the male of the same species. In the honeybee, we are not only interested in the methods by which new individuals arise but certain peculiar phenomena play an important part in practical apiary manipulations. The development of the drones or males from unf ertilized eggs must be considered, especially by the queen breeder. * Origin of the eggs. The eggs from which all the members of the colony develop are normally laid by the queen. In this individual, the only female in the colony whose reproductive organs are fully developed, the ovaries are large and, in fact, she is to a considerable extent simply an egg-producing machine. The ovaries of the queen (Fig. 92) consist of two groups (Ov) of egg tubes or ovarioles (ov). These tubes are small at the anterior end where the eggs are beginning their growth and toward the posterior end the individual tubes, as well as the total mass, increase in diameter. At the posterior end, the tubules in each mass open into the anterior end of an oviduct (OvD). The oviducts from the two ovaries unite farther 181 182 Beekeeping Ov back into a common tube or duct, the vagina (Vag), which opens to the outside below the base of the sting. The posterior portion of the vagina is enlarged, forming the bursa copula-trix (BCpx). Opening from the vagina is the spermatheca (Spm), a sac-like organ which serves to receive the male sex cells, the spermatozoa, from the drone at the time of mating and to retain them until they are needed. Attached to this are t wo accessory glands (SpmGl) the duct of which opens into the BGI BCpc- FIQ. 92. — Reproductive organs, sting and poison glands of queen, dorsal view. duct from spermatheca the vagina, duct from the to The the spermatheca is S-shaped and is surrounded by muscles forming the " sperm-pump" of Breslau.1 By the contraction of some of these muscles, the lumen of the upper end of the loop is enlarged and a small bundle of spermatozoa is taken from the spermatheca. By the contraction of other muscles, the sperma- 1 Breslau, Ernst, 1905-06. Die Samenblasengang der Bienenkonigin. Zool. Anz., XXIX, pp. 229-323. The Reproductive Processes and Parthenogenesis 183 tozoa .are forced on to the vagina. Cheshire 1 described this apparatus incorrectly by assuming that the muscles around the duct are sphincter muscles to hold back motile spermatozoa. The spermatozoa, according to Breslau, are not motile and no retaining muscle is needed. In copulation the spermatozoa are deposited by the drone in the vagina and must find their way to the spermatheca by this same duct. There is no special receiving duct as described by Cheshire. The spermatheca is not composed of muscle layers, as formerly supposed. Cheshire estimates that a normal vigorous queen may during her lifetime lay 1,500,000 eggs. Since mating occurs usually but once, those eggs which are fertilized must receive spermatozoa from the supply stored up in the spermatheca at the time of malftig. Since at each expulsion of spermatozoa a considerable number pass out and all but one are wasted, it is necessary that an enormous number be stored originally. Cheshire estimates the number at 4,000,-000 but it is enough to know that millions are then stored. The marvelous feature of the phenomenon is that these minute cells are able to live for perhaps five years away from the animal in which they were formed (the drone) and at the same time are so highly specialized that they can take no nourishment. There is no multiplication of spermatozoa in the queen as has been hypothecated by various beekeepers. The formation of the eggs has been studied by Paulcke.2 In the early stages of the formation of the egg at the anterior end of the ovarian tubes, the future egg nucleus is surrounded by other nuclei which later form nurse cells. There is at first no visible differentiation, no cell boundaries being seen, but farther down the tube the nuclei are surrounded by cell walls. Gradually the future egg cells begin to enlarge and 1 Cheshire, F. R., 1885. The apparatus for differentiating the sexes in bees and wasps. Jr. roy. micr. soc., ser. 2, V, pp. 1-15. 2 Paulcke, W., 1900. Ueber die Differenzirung der Zellelemente im Ovarium der Bienenkonigin (Apis mellifica). Zool. Jahrb. Anat. u. Ontog., XIV, pp. 177-202. 184 Beekeeping the individual egg cells are separated by a number of nurse cells, 48 to each egg, according to Paulcke. The egg cell increases in size chiefly by an accumulation of yolk which serves as food for the future embryo, this yolk being supplied by the nurse cells, which finally are exhausted and absorbed into the yolk of the egg. The egg and nurse cells are surrounded by an epithelium which grows thinner as the egg enlarges and which finally breaks when the egg passes into the oviduct. The egg is covered by a thin layer of chorion secreted around it by the epithelial cells and the boundaries of the cells may be seen in the lines which persist on the choriori, forming a delicate network on the surface. At the anterior end of the egg (where the head of the larva is formed and also toward the head of the queen) thtre is a peculiar arrangement of these lines, forming the micropyle. Here the spermatozoon which fertilizes the egg enters, but the mechanism has not been adequately described. In most insects there is a definite opening for the entrance of the spermatozoon and often a complex mechanism for the closing of the opening after fertilization. There is nothing so described for the bee egg. Origin of the male sex cells. The organs of the male (Fig. 93) in which the male sex cells originate are equally interesting. The spermatozoa develop in the testes (Tes), two organs homologous with the ovaries of the queen. The development of the spermatozoa probably occurs almost entirely during the pupal development of the drone and possibly not at all in the adult drone. From the testes, the spermatozoa pass through the vas deferens (VDef) into the vesicula semenalis (Fes) where they collect. The seminal vesicles open into the base of the accessory mucous gland (AcGl). These in turn open into a single duct, the ejaculatory duct (EjD), unusually large in the drone and curiously indented to conform to the structure of the vagina. The Reproductive Processes and Parthenogenesis 185 FIG. 93. — A, reproductive organs of drone, dorsal view, natural position; J3, inner surface of dorsal wall of bulb of penis; C, group of spermatozoa and intermixed granules; D, terminal segments of drone abdomen with penis partly protruded; E, lateral view of penis as invaginated within abdomen. 186 Beekeeping At the time of copulation, the penis, which is previously folded within the abdomen of the drone, is everted and projects into the vagina of the queen. The spermatozoa then pass through the ejaculatory duct as does presumably also the contents of the accessory mucous glands. The formation of the spermatozoa has been studied by Meves,1 by Mark and Copeland 2 and by Doncaster.3 The sudden expulsion of the penis causes the immediate death of the drone. The structure of the penis may be readily seen by gently squeezing the abdomen of a drone, by which means it is everted. In this case also the drone dies immediately so that his death at the time of mating should not be attributed to any action of the queen. As has been previously stated (p. 69), mating occurs in the air outside the hive. Parthenogenesis. The chief reason why the reproductive processes require extended discussion in a book on practical beekeeping is because of the development of the drones or males from unfertilized eggs. In most species, the sex cells disintegrate unless they unite with the products of the opposite sex of the same species, but there are numerous instances in the animal kingdom in which egg cells are produced, which, without fertilization, are able to develop into normal adults. To this phenomenon the name parthenogenesis4 is given. 1 Meves, Fr., 1903. Ueber Richtungskorperbildung im Hoden von Hymenopteren. Anat. Anz., XXIV, pp. 29-32. ——, 1907. Die Spermatocytenteilungen bei der Honigbiene (Apis mellifica L.) nebst Bemerkungen iiber Chromatinreduction. Arch, f. Microsk. Anat. u. Entwick., LXX, pp. 414-491. 2 Mark, E. L. and Copeland, Manton, 1907. Some stages in the sper-matogenesis of the honey bee. Proc. Am. acad. arts and sciences, XLII, pp. 103-111. 8 Doncaster, L., 1906. Spermatogenesis of the hive bee, Apis mellifica. Anat. Anz., XXIX. ——, 1907. Spermatogenesis of the honey bee. Ibid., XXXI, 4 For a more extended discussion, see Phillips, E. F., 1903. A review of 'parthenogenesis. Proc, Am. philos. soc,, XLII, No. 174, pp. 275-345. The Reproductive Processes and Parthenogenesis 187 In 1745, Bonnet described the parthenogenetic development of plant lice and just one hundred years later Dzierzon announced his theory that the drone is likewise a product of an unfertilized egg. This later paper, published in the Eichstadt Bienenzeitung, was the beginning of a long and heated discussion in which the leading zoologists of the day took part. Briefly his theory was as follows : (1) the queen is able "at pleasure" to lay either worker or drone eggs, the drone eggs being deposited just as they leave the ovary 1; (2) all eggs in the ovary are eggs which would normally develop into males and if fertilization occurs the sex is changed to female. It is well to divide Dzierzon's theory into these two parts for they are not equally capable of proof. The facts observed in the apiary on which this belief is based are as follows: (1) If a queen is unable to fly out to mate or is prevented from mating in some other way she usually dies (p. 70) but if she does lay eggs, as she may, after three or four weeks, the eggs which develop are all males; (2) if when a queen becomes old her supply of spermatozoa is exhausted, her offspring are all males; (3) if a colony becomes queenless and remains so for a time, some of the workers may begin egg-laying and in this case too only males develop. The author has found that many eggs laid by drone-laying queens fail to hatch and, in fact, are often removed in a short time by the workers. This makes it impossible for us to accept Dzierzon7s statement that all eggs laid by such a queen become males and the statement must be modified as follows : all of those eggs laid by a drone-laying queen which develop become males. The potentialities of the eggs which never hatch are not known. In addition to the facts here stated, the theory of the parthenogenetic development of the drone is supported by investigations of the phenomena of development in the egg. 1 Onions (1912, South African fertile-worker bees. Agricultural Journal of the Union of S. Af., May) claims that in South African bees females are also produced parthenogenetically. The claim is supported by considerable evidence. See also Van Warmelo, D, S., ibid^ 1913, who denies this statement. 188 Beekeeping Sex determination. The determination of sex is one of the most earnestly debated questions in zoology. Numerous theories have been proposed, most of which are not now seriously considered. From the observations and conclusions of Dzierzon and other observers it was long held that sex in bees and similar forms (ants and wasps) is determined by the presence or absence of fertilization. These species were seemingly an exception to the phenomenon observed in most species. Of recent years, sex determination has been the object of numerous investigations and it is now quite generally accepted that sex is inherited in accordance with the same laws which govern other phenomena of inheritance. It is, of course, impossible to attempt to record here or even to outline the observations which lead to this theory or to elaborate the theory, as has been done by various authors. It is now held that one of the chromosomes (the bearers of hereditary characters) of the sex cells bears the sex-determining character. If we take into consideration the important fact that not all the eggs of an unfertilized (drone-laying) queen hatch, then the bee does not appear as an exception in Nature. It seems clear, however, that the statement of Dzierzon that all the eggs in the ovary are male eggs cannot be accepted and it is, in fact, not improbable that the eggs destined to be females die for want of fertilization, while the eggs destined to be males, not requiring fertilization, are capable of development. It should be understood that the casting of doubt on Dzierzon's theory of sex determination does not invalidate his theory in so far as it pertains to the development of males from unfertilized eggs. In view of the fact that drone eggs are usually deposited in the larger cells, the theory has been advanced that the pressure on the abdomen of the queen when she is about to lay an egg in a worker cell, by some reflex, causes the sper-matheca to open, thereby enabling the egg to be fertilized. This is known among American beekeepers as the Wagner theory. Since fertilized eggs may be laid in comb foundation The Reproductive Processes and Parthenogenesis 189 when the side walls are only started and since drone eggs are often laid in worker cells, this simple explanation cannot be accepted. From the various phenomena observed in connection with parthenogenetic development, it appears that fertilization of the egg serves two purposes; it brings to the egg the hereditary characters of the male parent and also stimulates the egg cell to develop by cell division. If development can occur without this stimulation, the resulting individual contains the hereditary characters from one parent only. It should perhaps be mentioned that in plant lice both males and females sometimes develop from unfertilized eggs while in certain Lepidoptera only females develop from unfertilized eggs. The male sex is not a necessary result of parthenogenetic development. The theory that drones develop from unfertilized eggs has not been accepted without protest. From the beginning, it has been assailed by the publication of evidence and arguments which were supposed to contradict the theory. In the author's paper, to which reference has been made, the various contrary views are outlined and the interested reader is referred to this paper for references to the literature on the subject up to the date of publication (1903). Of recent critics, none is so insistent as Dickel, a German beekeeper, who claims that fertile queens cannot lay unfertilized eggs and that sex is determined by secretions of the nurse bees. These fantastic theories with others of a similar character have been adequately overthrown by DickePs critics and need not be discussed at length here. Practical applications. The development of males from unfertilized eggs is a fact of importance in various phases of apiary work. If, for example, an Italian queen mates with a black drone, the workers and queen offspring are hybrids,1 while the drone 1 Exception is sometimes taken to the use of the word hybrid as applied to a cross of two races, in which sense it is used by beekeepers. This 190 Beekeeping or male offspring is pure Italian. This fact is important to the breeder, for drones from mismated queens are just as good for breeding purposes as those from purely mated queens. It is true that this has been denied by various writers but the denial is based chiefly on variation in the color of the drones, it being overlooked that color is not a safe criterion for the purity of race of either queens or drones. Color is a much more stable characteristic in workers. The parthenogenetic development of drones must be considered in planning any breeding work with bees. In the selection of 'breeding material it does not necessarily follow because the workers of a colony have the quality desired that the drones of that colony will be best for breeding purposes, since the hereditary characters of the workers come from two parents while those of the drones come from only one of the two. Hermaphrodite bees. Many cases are recorded l of bees which show both male and female characters. These hermaphrodites or androgynous bees may have male characters on the head and female characters in the abdomen or they may be divided longitudinally in various combinations of characters. There is a mixture of male and female characters, varying in different individuals, in both external and internal organs. It is a peculiar fact, not easy of explanation, that when such cases occur there are often many in the same colony. Boveri2 suggests that in such cases fertilization is delayed until after cell division has begun and that only part of the cells receive criticism is probably based on the belief that sterility is characteristic of hybrids, as in the case of the mule, or it may be based on the belief that the word should be applied only to crosses of true species. There seems to be no objection to the word as beekeepers use it. It is most commonly applied to crosses of Italian and German bees. 1 v. Delia Torre, K. W. u. Friese, H., 1899. Die hermaphroditen und gynandromorphen Hymenopteren. Berichte d. naturw.-med. Ver. Innsbruck, XXIV. 2 Boveri, Th., 1901. Ueber die Polaritat des Seeigel-eies. Verb. Ges. Wurzburg (N. 8.), XXXIV, pp. 145-176. The Reproductive Processes and Parthenogenesis 191 the male chromosomes. While this theory would readily explain the great variation in such hermaphroditic bees it is based> on the assumption that sex is determined by fertilization, which may be questioned. Eggs which fail to hatch. In some cases, one of which came under the author's observation, queens are normally mated and lay eggs, but all the eggs fail to hatch. This is perhaps due to some abnormality of the queen, and in the case examined it appeared that the failure to hatch might have been due to the evaporation of the water in the protoplasm through the unusually thin and soft chorion of the eggs. Similar cases were described by Claus and v. Siebold l and also by Leuckart.2 1 Claus u. v. Siebold, 1873. Ueber taube Bienen-eier. Zeit. f. wiss. Zool., XXIII, pp. 198-210. 2 Leuckart, R., 1875. Ueber taube u. abortive Bieneneier. Arch. Naturgesch., XL. CHAPTER IX RACES OF BEES THE honeybee, so well known to beekeepers, has certain near relatives which are of interest, and it is quite probable that a careful study of the various phases in the behavior of these bees would throw considerable light on similar phenomena in the honeybee. The honeybee is usually considered as representing the apex of the evolution of the bees (Apidse), in that the social organization is the most complex found in this family of insects. The ants (Formicidse) and wasps (Vespidse) represent lines of parallel evolution in social life which has resulted in insect communities, comparable, but by no means identical, with that of the honeybee. TYPES OF SOCIAL BEES Among the Apidse are three great types of social "bees, the bumblebee (Bombus),1 the stingless bees (Melipona and Trigona) and the honeybees (Apis). The simplest forms, the bumblebees, have smaller colonies which die out during the winter, leaving the species to be continued from fertilized queens which hibernate. The stingless bees are tropical insects which store their honey and pollen in spherical vessels and rear their brood in "combs," one cell in thickness. In the honeybee colony, the architecture is the most perfect and the honey and pollen are stored and the brood is reared in hexagonal cells, which combine to form a comb two cells 1 For an excellent discussion of the biology of English bumblebees, consult Sladen, F. W. L., 1912. The humble-bee, the life history and how to domesticate it. London: Macmillan and Co. 192 Races of $ees 193 in thickness. The bumblebees and stingless bees fill a cell with pollen and honey, the queen then deposits an egg on this mass and the larva is not further fed or cared for. On the other hand, the queen honeybee lays -her eggs in empty cells and the larvae are fed a specially prepared larval food as they require it. For a further discussion of the more primitive bees as well as of the probable evolution of the Apidae, the reader is referred to the interesting paper of v. Buttel-Reepen.1 SPECIES OF THE GENUS APIS In the genus Apis there are other interesting honeybees but which have no special practical value. It is of interest to note first that specimens of Apis have been found in fossil form, preserved in amber, v. Buttel-Reepen mentions A. adamitica and A. meliponoides, the latter transitional between Melipona and Apis. Among recent species of this genus are A. dorsata, the giant bee of India, with its varieties zonata and testacea of the Philippines and the Malay peninsula, A. florea, a dwarf bee of India with several varieties and finally A. mellifica? the honeybee with the numerous varieties to be discussed later. Unsuccessful efforts have been made to introduce the giant bees into Europe and America, among which may be mentioned the trips of Benton, 1880 and 1905, and Dathe, 1883. Dathe succeeded in getting living dorsata bees to Germany but the effort was fruitless. The last mentioned trip of Benton was at the expense of the U. S.s Department of Agriculture. Dorsata builds a single comb in the open air, usually suspended on the limb of a 1 von Buttel-Reepen, H., 1903. Die stammesgeschichtliche Entstehung des Bienenstaates sowie Beitrage zur Lebensweise der solitaren und sozialen Bienen (Hummeln, Meliponinen, etc.). Leipzig: Thieme. 2 For a discussion of the propriety of mellifica as the specific name of the honeybee, see v. Buttel-Reepen, H., 1906. Apistica. Beitrage zur systematik Biologic u, s. w. Mitth. aus dem Zool. Mus. Berlin, and also the English translation of his paper "Are Bees Reflex Machines?" (Medina, O.: Root, 1907). See also p. 37 of this book, o 194 Beekeeping tree; there is no distinction between drone and worker cells and these bees do not take kindly to confinement in a hive.1 VARIETIES OF THE SPECIES MELLIFICA In the classification of insects, differences in structure and color are the characters on which classification is usually made, but in the differentiation of the varieties of honeybees there are no constant differences in these characters to guide the student. The varieties are established by beekeepers because of recognized and well-marked differences in the behavior of the bees from various regions. They are, however, valid biological varieties. While there are color differences, these are of little value in attempting a classification. Since beekeepers usually refer to these divisions of the species as races, this term is here adopted. Roughly the races are divided into three groups, (1) the eastern races, (2) the European races and (3) the African races. Certain characteristics of these groups are valid but the grouping is somewhat artificial. The principal races are here discussed in the order suggested by this grouping, the names given the races being indicative of their origin.2 Egyptian. These bees are somewhat smaller than the races best known to American beekeepers, the abdomen is slender and 1 For further data concerning the various species of the genus Apis, consult the above mentioned papers by v. Buttel-Reepen as well as the following: — , Gerstacker, 1862. Ueber die geographische Verbreitung und die Abande-rungen der Honigbiene nebst Bemerkungen iiber die auslandischen Honig-bienen der alten Welt. Reprinted in v. Buttel-Reepen's Apistica. Partial English translation by Dallas, Ann. and mag. of nat. history, 1863, III series, vol. 11. Koshewnikov, G. A., 1900-1905 [Material for the study of the genus Apis] Russian. Additional references are given in the v. Buttel-Reepen papers. 2 None of the races of the honeybee is native to America. The German bees were introduced early in the history of the country and are often designated native bees, but this is an error. After their introduction they Races of Bees 195 pointed and the cells of the comb are also said to be somewhat smaller. The first three segments of the abdomen are light yellow to reddish yellow with black border, being brighter than Italians. The abdomen is covered with grayish white hairs. The abdomen of the queen is marked with reddish brown on the first segment and the color areas are variable. Queens and drones are small and the queens are prolific. These bees sting furiously and are not subdued by smoke. They do not, according to v. Buttel-Reepen, form a winter cluster and therefore cannot withstand cold weather. Drones are reared in large numbers; the cappings are "watery";l the queen cells are small, very numerous, clustered and smooth. Fertile workers are abundant and are said to be present even when there is a laying queen. These bees were introduced into Germany in 1864 and to England and America in 1867. Here they attracted considerable attention but were promptly abandoned as worthless. Syrian. There are two races of bees in Palestine, one of which is, according to v. Buttel-Reepen, identical with the Egyptian. The other is known among American beekeepers as the Holy Land bees. The Syrians are larger than the Egyptians and in color they resemble Italians. These bees swarm excessively, build many queen cells and winter poorly. Many virgin queens go with after swarms and do not kill each other until one is mated. Young queens lay drone eggs in the first month. These bees were introduced into America in 1880 by Jones and Benton but were soon abandoned as valueless. They were introduced by Hopkins into New Zealand in 1883. multiplied rapidly and were soon found in the woods. It was formerly a common saying that a swarm always flies westward (to new territory). 1 Some races of bees fill their honey cells more completely than others and when the honey is in contact with the capping it gives the honey an appearance that is described as watery. * When the capping is separated from the honey by an air space the capping appears white (or yellow, depending on the color of the wax). In general the black races seem to produce whiter comb cappings than more yellow bees. 196 Beekeeping Cyprian. This bee has been given a thorough test by American beekeepers. It is somewhat smaller than the Italian and the abdomen is pointed, with three yellow bands, similar to that of Italians but somewhat lighter in color. The queens are small and very prolific. These bees winter well unless the colony wears itself out by breeding in winter. The workers are exceptionally cross, are not subdued by smoke and do not run on the combs. They build many queen cells (less than Syrians). Sent (unsuccessfully) to America by Gravenhorst in 1877 and first imported by Stahala in 1879: additional shipments by Jones and Benton in 1880. They have been widely advertised and tested but were abandoned because of their unmanageable qualities. Grecian. These bees resemble a hybrid between Italians and Germans. So far as known they have not been shipped to America. They were sent to Germany in 1860 by v. Roser. Caucasian. These bees vary in color from three bands of yellow on the abdomen to black or gray according to the region from which they come. The ones introduced into America have shown virtually no yellow color, having come from the more northern parts of the Caucasus. The yellow examples are said to resemble Italians markedly. This is the most gentle race known, although they defend their hives well against robbers. They seldom enter the wrong hive, winter well, cap their honey cells white and are, in the main, desirable bees. The hybrids are not gentle. They were first taken from their native country by Butlerov in 1877 and were shipped to Germany in 1879 to Vogel, who described them carefully. The first exportations were chiefly the yellow strains. In 1880 Julius Hoffman, Ft. Plains, New York, received two colonies of these bees but condemned Races of Bees 197 them because they did not work on buckwheat! Later Rauchfuss Brothers, Denver, Colorado, imported queens of this race and recommended them. Following this, additional queens were imported and American bred queens were distributed by the United States Department of Agriculture several years ago. The gentleness of this race is universally admitted, but Caucasians have some faults which have caused them to be abandoned by most beekeepers who have tried them. They use propolis most lavishly and in the autumn often build a wall at the entrance, leaving holes only large enough * ' 1 K *?IG< 94' — Propolis at entrance, built by Caucasian I or Single Dees beeg The entrance block on one side made a to pass (Fig. 94). propolis wall unnecessary there. They also build many burr and brace combs. The dark color makes it difficult to tell when the queens are purely mated and the dark queens are difficult to find on the combs. An additional factor which has led to the lack of interest in this race is the rapid spread of European foul brood within recent years. This has virtually necessitated the use of Italian bees in many localities and has discouraged experimentation with other races. There are still several prominent beekeepers who are enthusiastic in their praise of the Caucasians. Italian. This is the most popular race of bees among the best American beekeepers. The bees of Italy vary considerably in color, those in the north of the country being virtually identical with the German bees in color. The typical "three-banded" Italians are found farther south and in Sicily there is a still lighter strain. Some investigators believe Italians to be a cross between the German and 198 Beekeeping Egyptian bees. Typically, the yellow color covers three segments of the abdomen, the head and thorax and posterior segments of the abdomen being black with some traces of yellow on the mandibles, and the hairs have a yellow cast. The legs are brown. Queens and drones are variable in color from solid black to the yellow found on workers. Italians are gentle (but not equal to Caucasians in this respect), less prolific than the eastern races but usually better than black bees, build few queen cells, rarely develop fertile workers, keep the hive clean, drive out wax-moths, winter well, do not run on the combs, swarm less than Carnio-lans and some eastern races and cap their honey less white than Germans, Carniolans and Caucasians. The rearing of brood is quickly curtailed in a dearth of nectar and they cease rearing brood in the autumn sooner than most races. An important characteristic of Italians is the resistance to European foul brood. In this respect, they have been compared chiefly with German bees, to which race they are vastly superior. Italian bees were sent to Switzerland (by v. Baldenstein) in 1843, to Germany in 1853, to England (by Neighbor) in 1859 and to France aborut the same time (by Hamet), to Australia in 1862 and again in 1880, to German Guinea in 1887, from California to New Zealand in 1880, from Germany to Ceylon in 1882 and from Italy to New Zealand (to Hopkins) in 1883, to Guam in 1907 (from Hawaii by Van Dine). The first importation of these bees to America has been a matter of some dispute and was the basis of a sharp controversy. Their introduction marks an important milestone in American apiculture, almost equal in importance to the invention of the movable-frame hive. About 1855, Samuel Wagner and Edward Jessop of York, Pennsylvania, made an unsuccessful importation of an Italian colony, which died en route. In the winter of 1858-59, Wagner, Langstroth and Colvin (Baltimore) sent an order to Dzierzon (Germany), which was not delivered. Later in 1859, they received Races of Bees 199 seven living queens 1 from Dzierzon and reared two or three queens that fall, but the imported queens all died the following winter. On the same steamer that brought these queens, Mahan (Philadelphia), who had made a trip to Europe for these bees, brought over "one or more" queens (of doubtful purity). In June 1860, Wagner and Colvin received another consignment. In the meantime, S. B. Parsons (Flushing, L. I., New York) was commissioned by the Agricultural Division of the Patent Office to procure ten colonies, which he purchased from Herman of Tamins (reported by him January 3, 1860) and shipped from Havre, reaching the United States in May. In the annual report of the Division of Agriculture submitted January 29, 1861, the Superintendent reports that the effort was unsuccessful " owing to inattention to the instructions given by the agent of the Office." C. J. Robinson later asserted that he and Mahan had solicited an order from the Commissioner of the Patent Office in 1859 authorizing Mahan to proceed to Italy and procure bees. This request was refused but it was claimed that this instigated the movement to have Parsons (an agent of the Division then in Italy) get the bees. Robinson states that Parsons bought ten for the Government and ten for himself (this second purchase is denied by the friends of Parsons) and that he reported that all the bees consigned to the Government died. At any rate, Parsons in 1860 placed at least some of his stock in the hands of Gary, Langstroth, Quinby and others and the sale of Italian queens began in 1861. Riley, then Chief of the Division of Entomology, in 1892 claimed for the Government the credit of the first importation direct from Italy. Rose (New York) received colonies in 1861 and Colvin continued shipments from the Dzierzon apiary in 1863-64. Various other early shipments were made, and now many queens are received annually. In the early days of the enthusiasm 1 A cage dated May 1859 in Dzierzon's handwriting was found by C. H. Lake after he purchased the Colvin apiary (Beekeeper's Instructor III, No. 12, 1881). 200 Beekeeping over Italian queens they often sold for twenty dollars each. Italian bees are now found everywhere that beekeeping is conducted and are usually considered preferable to all others. In the United States, special attention has been given to the breeding of Italian bees and it is probably true that better Italian stock can now be obtained in America than in Italy. To distinguish differences in strains of Italian bees and in part to provide trade names, various names have been given by American beekeepers to certain types. The five-banded or "Golden-all-over" bees have been bred specially for an increase in the yellow color on the abdomen. The red clover Italians sold several years ago were supposed to have a tongue-length above the average, sufficient to allow* them to get nectar from the red clover blossom. Various strains are distinguished by the name of the breeder. The specially yellow bees are not usually considered as desirable for commercial beekeeping as the typical Italians, which are commonly designated as three-banded or leather-colored. German. These bees are black in color and are generally known among American beekeepers as "Black bees/7 It is of interest to note that according to Dzierzon there were yellow bees in Germany before Italians were introduced and this helps to explain the variation in the German bees. v. Buttel-Reepen, following distinctions made by Dzierzon and other early writers, divides the German bees into two varieties, the typical variety and the heath bees. The typical variety is native to Germany, Russia, Scandinavia, Denmark, Holland, England, Switzerland, Austro-Hungary and parts of other European countries. The heath bee is darker than the typical variety, swarms excessively and is especially adapted to honey-flows coming in late summer (buckwheat, heather): young queens in after swarms lay drone eggs abundantly the first season. This variety is found in Hanover, Holstein, Oldenburg and Holland. There is a possi- Races of Bees 201 bility that there are two varieties of German bees found in the United States, as is so often claimed, and that these have arisen from these two natural varieties. The German or black bees found in the United States seem to combine many of the undesirable qualities of all other bees. They are less prolific than Italians, they (and especially crosses with Italians) are cross but respond to smoke, they build more queen cells than Italians and develop fertile workers more readily (less in these last respects than the eastern races), they do not clean the hive well or resist moths completely, they run badly on the combs and fall off from the corners of the combs during manipulation and they swarm more than Italians. Their greatest fault is that they succumb so rapidly to European foul brood that it is most difficult to rid a colony of black bees of this disease. They cap comb-honey white and winter fairly well, but their nervousness is against them in this respect. While these bees are condemned by the best American beekeepers, some of the leading beekeepers of Europe (especially in Switzerland) claim them to be superior to Italians. Since no effort has been made to improve these bees in America, this may account for this difference of opinion. The German or black bees were introduced into New England (probably from England) in 1638. In 1644, John Bales was brought to Newburg from a neighboring town to instruct the people in beekeeping, indicating an early interest in the industry, but he later became a town charge. Black bees reached West Florida not later than 1763, Kentucky in 1780, New York in 1793, west of the Mississippi River in 1797, Cuba in 1764; San Domingo in 1781, New South Wales in 1822, Tasmania in 1831, New Zealand in 1839, Brazil in 1845 (or earlier), Chile about 1848, California in 1853, Columbia about 1855 and Argentine in 1857. Har-bison took 116 colonies (with a loss of only six) from Pennsylvania to California via Panama in 1857. The same year (August 20th) the first bees were shipped from San Jos6, California, to Hawaii. 202 Beekeeping Carniolan. These bees are grayish-black in color and the claim that yellow bees were native to parts of Carniola is often questioned. Professor Francis Jager is authority for the statement that the bees of the Wippach valley (Vipavska dolina) are yellow. Carniolan bees are large, gentle (second only to Caucasians), prolific, swarm excessively, are good honey-gatherers, build numerous queen cells, collect little propolis, winter admirably, cap their honey white and do not run on the combs during manipulation. It is claimed by some beekeepers that they resist European foul brood as well as Italians: this should be thoroughly investigated by disinterested persons. They are native to Carniola, Austria. These bees have been shipped repeatedly to Germany and other European countries and to America. While they have some ardent advocates in the United States, they are losing ground, especially on account of their swarming proclivities and the black color, which American beekeepers do not fancy. Queen breeders have distinguished other races, which are not distinct from the Carniolan, among which are the Banat (Banater) race, of recent importation into America, and the Dalmatian which appeared in American literature in the eighties. The names of other provinces have been used as trade names for different breeders. African races. While several races of bees have been distinguished from Africa, in addition to the Egyptian previously discussed, very little information is at hand concerning these bees in the hands of beekeepers. A yellow race, described as Apis adonsoni, is found in parts of Africa, having the abdomen a darker reddish-yellow than the Egyptian. On the north coast of Africa is found a black bee, known among beekeepers as Tunisian, Punic or North African. This race extends well into the continent. The bees of this origin that have been tried in the United States are extremely cross, propo-lize excessively and winter badly. They are not now known Races of Bees 203 to be present in the United States. A separate race of black bees is described from Madagascar and other islands and still another from Togoland. It is well known that honeybees are abundant in parts of Africa and careful explorations would doubtless reveal many interesting facts concerning these bees. Onions l claims that in the South African race the unmated workers lay eggs which develop into female bees. As the continent of Africa becomes more settled by white men and as apiculture advances, we may expect some interesting additions to our knowledge of the African races of bees. Asiatic races. v. Buttel-Reepen (Apistica) places A. indica as a variety of A-, mellifica. It is a smaller bee, which is said to bite rather than sting. It crosses with previously described races. Several sub-varieties are indicated. Chinese-Japanese. These bees are placed by v. Buttel-Reepen as sub-varieties of indica. The Chinese bee has a heavy coat of long dirty gray hair; the Japanese bee lacks this. BEST RACE OF BEES To answer the question as to which race of bees is best is difficult. For comb-honey production, the German, Carniolan and Caucasian races have the advantage of capping the honey white but the German bees are especially subject to European foul brood, Carniolans swarm excessively (especially in comb-honey production) and Caucasians propolize badly. Without going further into the merits and demerits of the various races, it may be as well to give the almost unanimous verdict of American beekeepers, 1 Onions, G. W.f 1912. South African "fertile worker-bees." Agricultural Journal of the Cape of Good Hope, May. See also Van Warmelo in the same journal, 1913. 204 Beekeeping which is in favor of the Italian race. It is probably true that the tests made cannot be considered as free from prejudice but the decision was made years ago and no special reason has been presented for changing it. Since this race became popular it has been carefully bred and it is easier to get good stock of this race than of any other in the United States. It was the first race brought to this country in the effort to improve on the early introduced black bees and proved so vastly superior that it soon took a firm hold on American beekeepers. It is doubtful whether any other race will be accepted as better or even as good by the majority of beekeepers and certainly no markedly better race has been tried in this country. CHAPTER X REGIONAL DIFFERENCES WITHIN THE UNITED STATES BEFORE discussing the different methods of manipulating bees in the successful production of honey, it may be helpful to point out some of the fundamental differences found between various parts of the United States in regard to the sources of honey and in climatic conditions, which influence the choice of the proper system of manipulation. In the American literature on beekeeping, these differences are frequently mentioned and the word "locality" in the beekeeper's vocabulary has come to be used as an all-inclusive argument or excuse for his particular practice and often partially to cover his ignorance of the actual reasons for differences observed. This term "locality" is the subject of the present chapter. As will be shown later, two apiaries but a few miles apart may give quite different results, not only in the amount or source of crops but in the effect of certain manipulations, and the facts here presented help to explain these differences. Since migratory beekeeping is practiced only between two unlik6 regions, this subject is also discussed in the present chapter. The system of manipulation to be followed and the manner in which honey may best be prepared for market depend on the color and quality of the honey and perhaps especially on the length and intensity of the nectar-flow. While the chief sources of honey are discussed in another chapter, it may be helpful here to present in outline the combination of floral and climatic conditions which so strongly influences 205 206 Beekeeping the business of honey-production and guides the beekeeper in choosing suitable locations for his apiaries. Variation in intensity of honey-flows. In general, the nectar-flows increase in rapidity or intensity as one goes northward and with this rapidity in the honey-flow usually comes a shortening of the period during which nectar is secreted. As a rule, the northern honeys are lighter in color, although there are many exceptions which will be pointed out later. In the more northern localities, the beginning and end of the honey-flows are usually sharply marked, while in the South there is a gradual increase in the honey-flow to the maximum and a correspondingly gradual cessation of the honey-flow. Variation in the value of plants. There is a striking difference in the value, from a beekeeper's standpoint, of plants according to locality, and the causes of these phenomena are in most cases not understood. A few of the more striking examples will serve to illustrate this variation. White clover yields nectar most abundantly in the northern range of this plant, while farther south the flow of nectar from this plant is less intense and the honey is often somewhat darker. Alfalfa yields nectar freely in the irrigated districts of the West, but is usually of no value to the beekeeper east of the Mississippi River. Buckwheat is the source of large honey crops in parts of southern New York and Pennsylvania, while in Indiana and Illinois it secretes much less nectar, again increasing in value to the beekeeper in Michigan. Exceptions to these general statements sometimes arise because of abnormal climatic conditions. For example, a heavy honey-flow from alfalfa was recently obtained in the vicinity of Syracuse, New York. This was probably not due to the plants becoming acclimated but occurred in a dry season. In some seasons white clover yields well much farther south than the limits previously given. Other examples are given later. Regional Differences Within the United States 207 BEEKEEPING EEGIONS It helps to an understanding of the differences in the practices of beekeepers in various parts of the United States if we divide the country into honey regions. This, as any experienced beekeeper will at once recognize, is a more or less arbitrary division and many exceptions might be cited to the following classification. In the main, however, the nature of the honey-flows justifies such an arrangement, and this plan is still more permissible if we consider the systems of manipulation found most advantageous by beekeepers. These regions may first be divided into general and restricted, depending chiefly on their area. The general regions are those not only of considerable extent, but of greater influence on the choice of manipulations. General regions. These are five in number and the division is based on differences in climatic conditions found in the United States, The placing of the sage region in the rank of a general region is justified mainly by the size of the crops obtained there in favorable seasons. (1) The white clover region includes eastern Canada, the New England States, except along the coast, and a belt along the northern United States as far west as the Dakotas. It is limited to the west by the arid region and again reappears on the Pacific coast, both in the United States and Canada. The southern boundary is approximately Mason and Dixon's line and the Ohio River. In this region, in addition to white clover, alsike clover, sweet clover, basswood, tulip poplar and locust contribute to the honey crops and, with the exception of that from tulip poplar, the honeys from these sources are light in color. Alsike clover is steadily increasing in importance with its wider planting for forage, while basswood is rapidly disappearing because of the extensive cutting of this tree for lumber. In this region the honey-flows are rapid and relatively short and the main 208 Beekeeping honey-flow is usually preceded by a honey-flow from spring flowers (fruit bloom, dandelion) followed by a dearth. In the more northern localities this interval is brief or entirely absent. After the main honey-flow there is usually a period when no nectar is available, followed again by a late summer or fall honey-flow (buckwheat, asters, goldenrod or Spanish needle, according to locality). This region is suitable for comb-honey production better than any other part of the United States, on account of the intensity of the honey-flows and the light color of most of the honeys. The necessity for a rapid building up of colonies in the spring and, the difficulty of swarm control make necessary special systems of manipulation in this region. The wintering problem is naturally most acute here also. Most of the American literature on beekeeping in both books and journals is based on systems applicable to the white clover region. The honeys of this region are in great demand, probably because the honey-consuming public is better educated to the flavors obtained here. In this region are thousands of beekeepers with only a few colonies, although the number of specialists is increasing satisfactorily. (2) The alfalfa region is located in the West, where this plant is chiefly grown for forage. Alfalfa is at its best as a nectar-producing plant under irrigation and usually at high altitudes. Colorado, Utah and Idaho are now the largest producing States in this region. Sweet clover is also an important contributor to the nectar supply in some sections. The honey of this region is usually of fine flavor and light in color, but alfalfa honey quickly begins to granulate and in consequence would seem best adapted to extracting. The honey-flow is not so rapid as in the white clover region, which also makes this region less favorable for comb-honey production. However, many beekeepers of this region produce comb-honey extensively. The system of manipulation is different from that in the white clover region because the honey-flows are usually longer and swarming is less difficult to control. The number of honey-flows de- Regional Differences within the United States 209 •« pends on the number of crops of alfalfa that are harvested in a season. This region is steadily increasing in importance, and a market is rapidly being built up for alfalfa honey. In this region, honey-production is conducted chiefly by extensive beekeepers. (3) The south-eastern region^ which varies greatly in the sources of honey throughout its extent, and which is an area of abundant rainfall, lies south of the white clover region and extends west to eastern Texas. The various rather distinct subdivisions of this territory might well be placed among the restricted honey regions, except that certain things in common in the honey-flows make the same type of manipulations necessary. Among the important plants of this district are sourwood, cotton, tulip poplar, tupelo, manchineel, mangrove, titi, palmettos, citrus trees, gallberry and partridge pea, with nectar from clovers in some sections in favorable seasons. Sweet clover is valuable in some localities. Most of the honeys are amber, and the chief characteristic of this region is a succession of honey-flows, often intergrading. The honey-flows are usually not rapid. This region is therefore best adapted to extracted-honey production. Swarming is much less troublesome than in the North. Beekeeping is developing in this region, but there is opportunity for many more beekeepers, there being now relatively few who rank as professionals. Colonies of bees may usually be bought at low prices in box-hives. This region is perhaps the best in the United States for commercial queen-rearing, except in parts of Florida where dragon flies are troublesome. The diversity of conditions in this region is well shown in the accompanying map (Fig. 95), which was prepared by E. G. Baldwin, De Land, Florida, who has studied the beekeeping possibilities of his State quite thoroughly. It will be seen that the sources of nectar are quite distinct geographically. The geographical position of this State and the diversity of soil and climatic conditions strongly influence the growth of the honey plants. In this State 210 Beekeeping two localities only a few miles apart may be quite unlike, when viewed from the standpoint of the beekeeper. This is also true in many other parts of the United States. (4) The semi-arid region of the south-west is located chiefly in the arid and semi-arid parts of Texas and Arizona and here too the honey plants are of somewhat restricted distribution. Among the plants which are important to the beekeeper are mesquite, horse-mint, catclaw, huisache and guajilla. Most of the honey pro-duced in this region is extracted, although a large number of beekeepers produce bulk comb-honey (chunk honey), cut from large combs (without sections) and placed in cans in which extracted-honey has been poured to fill the spaces. Such honey is usually sold locally. Beekeeping in this region is largely in the hands of professional beekeepers. (5) The sage region is confined to the cafions of southern California and should be considered a restricted region except for the fact that sage honey wields so important an influence on the honey market that the region may justly be placed among those of major importance. The various sages are all heavy yielders under favorable conditions and there is usually no other honey source of importance where this honey is produced. The influence of rainfall on the honey crop of this region is discussed in the chapter on honey Won of White Tupelo Money. • -Partridge Pea Hone? •Orange Blossom Money. Fio. 95. — Map of Florida, showing distribution of honey plants. Regional Differences within the United States 211 sources. If the crops of sage honey were uniformly heavy, this would be an ideal region for comb-honey production, for sage honey is mild in flavor, water-white and does not easily granulate. However, successful comb-honey production necessitates a rapid flow, which often fails to appear in this region and most of the honey is extracted. Here too there are many professional beekeepers, although a considerable number are not modern in their methods of manipulation and equipment. In addition to the geographical limits ascribed to these main regions, other localities situated outside the prescribed boundaries might well be included with certain of the general regions in discussing the type of flow. For example, California, north of the sage region, is comparable with the South, and along the Pacific coast to the north there are localities which belong to the white clover region. Restricted regions. In addition to the more general divisions named, there are other localities with special advantages for the beekeeper, but more limited in extent, which lie within the boundaries of the main divisions. As previously mentioned, the southeastern region is virtually composed of a number of such restricted regions. The list here given will be recognized as incomplete and is intended merely as a suggestion. Many more restricted regions will be recognized from the discussion of the sources of nectar. Among the more important limited regions may be mentioned those in which the following plants secrete nectar. (1) Buckwheat. — The honey of this plant is dark and of strong flavor, suitable chiefly for manufacturing purposes. "The variation in the secretion of nectar from buckwheat has been mentioned. (2) Sumac. — Valuable locally in New England. Another species of the same genus yields a surplus in limited areas in Georgia. (3) Spanish needle (numerous species).—Heavy yielders 212 Beekeeping of amber honey in the autumn in swamps. Among the best-known regions in which these plants are of value may be mentioned the lower Delaware River and Illinois River valleys and the Kankakee swamp. (4) Willowherb. — Important in northern Michigan in burned over forest areas. (5) Sweet clover. — In some sections, especially in limestone regions, this plant is exceptionally abundant and is the source of large crops of honey of a slightly greenish color. It is especially valuable in northern Kentucky and southern Indiana. _ (6) Blue thistle. — Important in the Shenandoah valley. (7) Raspberry. — Northern Michigan where the forests have been burned over and in parts of New York. (8) Beans. — In southern California, where beans of various kinds are grown in great quantity, beekeepers find it profitable to move their apiaries to the bean fields after the sage honey-flow. The honey is white, of excellent flavor and granulates quickly. (9) Heartsease. — Mississippi valley. A heavy yielder of nectar in late summer. Variation within a region. It must not be understood that the territory within either a general or a restricted region as here defined is equally good throughout. The cutting of forests, the extensive cultivation of some plant which restricts the growth of honey plants, local differences in soil or drainage, the presence of large towns and a multitude of other factors may so reduce the number of individual honey plants where they would normally grow as to make extensive beekeeping unprofitable. On the other hand, the cutting of forests may make a region better by allowing a honey plant to spread (e.g. willowherb) or the planting of some nectar-yielding species, either under cultivation (e.g. alsike clover, alfalfa) or in waste places (e.g. sweet clover), may greatly increase the value of a region to the beekeeper. In fact, the entire Regional Differences within the United States 213 alfalfa region is a man-made honey region. These factors, many of which are due to human interference with the natural environment, must be considered in choosing locations of apiaries and in manipulating colonies. Climatic influences may change an area from year to year. A lack of sufficient rainfall, for example, may kill white clover in certain areas and not in others. This occurred during 1914, when a severe drought killed clover over much of Illinois, while an abundance of rain fell in northern portions of the State, there being marked differences in localities only a few miles apart. DISTBIBUTION OF BEES IN THE UNITED STATES The relative importance of the various honey regions is indicated by the number of colonies of bees found in each one, although care must be exercised in examining these data to avoid misinterpretation. The only source of information on this subject is the United States Census, and the data from this source are not complete. However, while the number of colonies reported is far too low, it may perhaps be assumed that approximately the same percentage is omitted throughout the United States. The accompanying map (Fig. 96) was prepared in the Bureau of Crop Estimates of the Department of Agriculture from data furnished by the Census of 1910 and the author is indebted to this Bureau for permission to use it here. In this map will be found a dot for each county where bees are kept, the size of the dot being proportionate to the number of colonies reported. In the white clover region, it is evident that the more northern localities are most thoroughly stocked with bees. In the alfalfa region bees are less abundant, and this is true also in the sage region. The amount of honey produced in these regions is far below that of the moist regions of the country, but the honey goes to market in large shipments, because of the larger number of specialist beekeepers, and FIG. 96. — Map of United States, showing distribution of colonies of bees according to the Census of 1910. Regional Differences within the United States 215 as a result these crops are important in determining the wholesale price of honey. The enormous number of colonies in the southern States is a surprise. In the fifteen States usually included in the division of southern States are found forty-five per cent of all the colonies in the United States. In this region the box-hive and the farmer-beekeeper are still found in large numbers, there being few specialists except in Texas. Because the industry has not developed on modern lines, most of the honey from the South does not reach the larger centers of distribution, and it therefore has little influence on the wholesale honey markets. The number of colonies of bees found in the South is proof of the wonderful opportunities for the development of the industry, for many of these colonies are given no attention. As one beekeeper expresses it, these bees would die, if they could, to escape the ill-treatment to which they are subjected, but the environment is so favorable that they increase in spite of mismanagement. Attention should also be called to the larger number of colonies in southern New York and northern Pennsylvania, where buckwheat is plentiful. The other restricted honey regions seem to have less influence on the number of colonies. This map will repay considerable study in connection with other phases of beekeeping. To one familiar with the distribution of the diseases of the brood of bees in the United Statesl it is clear that there has been a severe loss from this cause, as indicated by the smaller dots in regions where diseases are most prevalent. New England was formerly well stocked with bees, but many colonies have been destroyed by disease. Certain areas in Pennsylvania, Ohio and Indiana, where disease is abundant, are inadequately provided with bees. One important reason for the larger number of colonies in the South is probably the scarcity of disease. 1 Phillips, E. F., 1911. The occurrence of bee diseases in the United States (Preliminary report). Circular No. 138, Bureau of Entomology, 25 pp. 216 Beekeeping MIGRATORY BEEKEEPING By this expression beekeepers designate the moving of apiaries from place to place during a single summer to take advantage of two or more honey-flows which do not occur in a single locality. This has been practiced since ancient times, and most extensive beekeepers cherish the hope that some day the subject may be sufficiently understood so that they may move their bees several times a season and thereby keep them working almost all the year. Some elaborate plans have been made for moving bees from south to north as the seasons advance, but most of the trials have been failures. Since success in beekeeping depends on an intimate knowledge of the honey sources of the locality and of the best manipulations to obtain maximum crops, such migratory beekeeping would necessitate detailed knowledge of many sections, so that the beekeeper may know when and where to move his colonies to advantage. The Mississippi River has long been considered an ideal avenue for transporting colonies in migratory beekeeping, especially since there is no better way to ship colonies than by boat. It has been proposed that the beekeeper place his apiary on a flatboat in the South in early spring and move northward by night, allowing his bees to gather nectar by day, and following the season as it extends northward. This plan so well illustrates the limitations of migratory beekeeping that it may be critically examined. One of the chief difficulties is the fact that the beekeeper must know just where to anchor after each move so that his bees will be in range of the best forage and this would involve too careful a study of the valley to make the plan practical. This objection might be overcome but there is a more fundamental difficulty which has not been sufficiently considered by those who have cherished this dream. If one species of plant furnished the main nectar-flow throughout the Mississippi valley, the beekeeper could move northward to prolong the gathering period, but this is not the case. Regional Differences within the United States 217 There are many plants which furnish nectar in the various portions in the valley and if the apiary were moved northward the bees often would leave behind them a honey-flow from another source. If migratory beekeeping from south to north and then back south with the closing season proves successful it will probably be within the nectar-secreting area of a single species of plant or perhaps of two species, one for each direction of the journey. The Mississippi River plan was tried several years ago on a rather extensive scale without success. The experiment of moving an apiary south for the winter for the purpose of making increase has recently been tried. If one is raising bees for sale and has a heavy demand for colonies that may pay but the chances of success in following this plan for honey-production are small. The limitations of migratory beekeeping, in so far as present successes indicate them, have not been previously pointed out. They are approximately as follows: The movement of bees must not be from one general region to another, as from the white clover region to the alfalfa region, but from a location where the flora is that of the general region'to a restricted region where the honey-flow comes at a different time, usually later. For example, it has been found profitable to practice the following plans in migratory beekeeping : (1) from white clover to sweet clover, buckwheat, Spanish needle or heartsease, (2) from sage to bean or (3) from one of the restricted regions in the South to another. As previously mentioned, the honey-flow at the temporary out-apiary usually comes after the main honey-flow at the permanent apiary. In most such cases, the conditions demand the production of extracted-honey, as comb-honey production and migratory beekeeping are not well suited to each other. In considering the possibility of migratory beekeeping it must be decided whether it is desirable to move the bees or simply the beekeeper. In other words, if the trip is a long one involving considerable expense and danger of loss 218 Beekeeping in moving a large number of colonies, it may be cheaper or easier for the beekeeper to own two or more lots of bees and supplies. The expense of transportation and the danger involved are probably the factors which determine the feasibility of moving from south to north or from sage in California to alfalfa in Utah or Colorado. In the South, especially where bees can be purchased at a low price, it would not seem profitable to move apiaries over long distances. The shipping of bees in wire-cloth cages may in the future remove the present limitations. It would certainly seem that a northern beekeeper is not embracing all his opportunities if he quits work when his bees can no longer get nectar, while there are still hundreds of places in the South or even in the tropics where he might maintain apiaries with profit in the winter. When it is recalled that the professional beekeeper is a relatively new factor in beekeeping, it may still be expected that the future development of the industry will show an increase in migratory beekeeping, or at least in migratory beekeepers. OVEBSTOCKING i The bugbear of the specialist beekeeper is the fear that he will overstock his localities, that is, place in each apiary so many colonies that there will not be enough nectar available to permit the colonies to store approximately the maximum profitable surplus. Since there are few places in the United States that are now overstocked, this subject worries the beekeeper more than the facts warrant. Some beekeepers have found it practical to keep several hundred colonies in one apiary. E. W. Alexander, Delanson, New York, found it more profitable, in an exceptionally good buckwheat region, to keep over 700 colonies in one yard than to establish out-apiaries. In the South and West large apiaries are not infrequent. While it is desirable to keep bees in as few places as possible to avoid duplication of apparatus and time lost in trans- Regional Differences within the United States 219 portation, there is another factor to be considered. The size of an apiary should be determined chiefly by the number of colonies that the beekeeper can manipulate in a single day during the honey-flow. If he finds that he can usually care for seventy-five colonies in a day under his system of management, then that number is ideal for his apiaries. He can then arrange his out-apiaries so that each will receive a visit as frequently as the conditions demand. The amount of work that can be done in a day will increase with experience and the out-apiaries correspondingly may be increased in size, for they should be large enough to furnish a full day's work, unless there is some means of rapid transportation available. With modern transportation facilities the distance to out-yards is of less importance than formerly and many beekeepers now have motor trucks to carry an extracting outfit and other apparatus and supplies from one apiary to another. Considering the day;s work as the determining factor in the size of the apiary, the out-apiaries may be more numerous and closer together than would be the case if each yard were increased to the maximum. In the present undeveloped condition of the beekeeping industry and with so many localities almost untouched by bees, it is not wise to run any risk of overstocking. The location of out-apiaries should be determined by the available forage, the minimum distance between them usually being determined by the distance that bees can fly. DADANT OUT-APIARIES To illustrate the problem which confronts the beekeeper in the establishment of out-apiaries there is here reproduced a map (Fig. 97), made from one by C. P. Dadant, Hamilton, Illinois, of the apiaries near his home in 1891. He then owned the Home, Sherwood, Villemain and Sack apiaries, the other four shown being apiaries of other beekeepers. All of these are located on land sloping toward the Mississippi River. The Sherwood apiary was the best, giving crops in 220 Beekeeping the spring and fall. The Villemain apiary appears to have been in the poorest location, the range of the bees being restricted by the river, but it was near the only basswood grove in the country and the bees gathered honey in the fall from the islands. The Sack apiary seems to have been too near other apiaries but was actually second only to the Sherwood yard. The bees did not work more than a mile along the bluff but went three miles to the river, having the bottom lands covered with fall flowers within their range of flight. They were separated from the adjacent apiaries by hills and timber. The two small circles show sites of former apiaries, used before the Sherwood apiary was established. The bees in the home apiary were only a mile and a half from abundant pasturage on an island but did not reach it, although they sometimes went two miles or more in another direction. This description of conditions in 1891 * may not represent the condition of the Dadant apiaries to-day. The map, however, shows the locations decided upon in that region by an experienced beekeeper who had kept bees in that district for twenty years. It shows that distance from one apiary to the next is not the sole consideration but that contour of the land, timber tracts and other barriers must be taken FIG. 97. — Map showing distribution of Dadant apiaries, Hamilton, 111. ' * Dadant, C. P., 1891. Arrangement of out-apiaries. Bee Culture, XIX, pp. 60-61, Gleanings in Regional Differences within the United States 221 into account. Similar maps and descriptions of out-apiaries appeared in the same journal at about the same time, one of the apiaries of E. France, Platteville, Wisconsin, and another from A. E. Manum, Bristol, Vermont. The effect of contour of the land is especially well illustrated in the Manum map. It would probably profit any extensive beekeeper who has several out-apiaries to make a similar map of his region, especially if he includes the honey sources. CHAPTER XI THE FIRST STEPS IN BEEKEEPING MANY persons begin beekeeping accidentally. The interest of many of the present beekeepers in the honeybee has first been aroused by a swarm passing over the premises or perhaps lighting on a tree near by. The desire not to allow anything to go to waste or not to allow a valuable article to elude him, coupled perhaps with a dare-devil impulse to risk a combat, has induced many a man or woman to attempt to hive the stray swarm. Or perhaps there is a temptation to exhibit one's prowess before the other members of the family. Having hived the swarm in a box or barrel with no loss of life or limb — and bees are never easier to handle than when swarming — it is by easy steps that one goes on until an attack of what is commonly known as " bee-fever" has developed, from which recovery seems hopeless. If a swarm is caught and put into a box, the owner should obtain a modern hive as soon as practical and lodge his new possession in a home where they may be manipulated. The necessary equipment is discussed in Chapter II. Purchase of colonies. There are those, however, who decide to begin beekeeping without this accidental impulse. The desirability of beginning on a small scale may be again emphasized here. Colonies should if possible always be* purchased near at hand to prevent the loss which may follow, especially if colonies must be transported by inexperienced persons. The further advantage in this procedure is that the former owner may usually be induced to assist in the moving and he will also 222 The First Steps in Beekeeping 223 prove helpful in the early days with the bees. It is best to have the bees already housed in the hive which is to be adopted but if this is not possible then colonies in any hive or in boxes or barrels may be purchased and transferred (p. 245). Transferring is a difficult operation for a novice, in fact it is not relished by an experienced beekeeper, and is to be avoided. Early in the summer is usually the best time for making the start, although the price charged for colonies is usually higher at that season. There is less opportunity for making such mistakes in management as will result in loss of bees during the first few months of ownership if these come during the summer. It matters little what race or strain of bees is obtained at the beginning except that it is desirable to avoid bees with too great a percentage of black blood in them, such bees being difficult to manipulate. After the apiary is established any desired race may be obtained by removing the queens and replacing them with mated queens purchased from commercial queen breeders. If these are introduced to replace the old queens of the colonies, the progeny of the new queens will rapidly replace the offspring of the discarded ones as they die from natural causes. '^Purchase of bees to be shipped from a distance. Another method of buying bees which is growing in favor is to buy them by weight, without frames. Bees are now easily shipped in cages specially constructed for the purpose, even though the journey require several days. By this method the possibility of carrying some brood disease is obviated and there is less likelihood of damage to the bees by the breaking or melting of combs or by suffocation during the journey. Nuclei or small colonies with frames may also be shipped a considerable distance. These will usually build up to full colonies during the season if bought early, but of course surplus honey can scarcely be expected the first year from such a small colony. Still another plan is to buy an empty hive and leave it with some beekeeper so 224 Beekeeping that a swarm may be hived in it, after which it is removed to the desired location. Requirements in purchased colonies. If there is opportunity to examine the colonies before purchasing them, there are several important things which should be insisted upon: (1) get as little drone comb or crooked or defective comb as possible; (2) see that the colony is free from disease (p. 397); (3) the colony should be provided with plenty of honey and (4) the amount of brood should be adequate for the time of year. It is perhaps asking too much of the beginner to expect him to determine whether disease is present in colonies purchased. In many states and counties there are official apiary inspectors whose duty it is to give advice on the subject of disease and these men may be asked to assist in this work. At any rate, even the novice can tell whether there is any dead or discolored brood and it is at least safe not to accept colonies in which any discolored brood is found, normal brood being pearly white. Formerly the sale of bees was believed to bring ill-luck and the customary way to acquire colonies was to go at night to the apiary and after the removal of the colony, to leave coins to the value of the bees on an adjacent hive. The possibility of an insufficient pile of coins or perhaps none at all is probably a factor in causing modern beekeepers to prefer to *sell bees according to present-day methods. The beginner can scarcely be advised to adopt the ancient manner, for the custom might be found faulty when explained to a magistrate. How to learn beekeeping. To acquire skill in manipulating bees and to learn the proper management of the apiary so as to obtain maximum results, the best method is to spend some time in the apiary of an experienced beekeeper. If one contemplates making beekeeping an important part of the occupation, this is especially to be desired. It is usually possible to arrange The First Steps in Beekeeping 225 for employment at a small wage in the apiary of a specialist for a season. Not all extensive beekeepers, however, manipulate their bees well and many of them fail to get the maximum returns through faulty systems, especially in comb-honey production, but after some experience in such an apiary the prospective beekeeper is better able to read the details of manipulations understandingly, and he can correct in his own practice the mistakes which may have been taught him by his teacher. Value of reading. The many books on bees all have points of merit and reading the various journals devoted to beekeeping is to be commended. Reading alone does not make a beekeeper. The " book-beekeeper" may be well informed concerning the behavior of bees and may know the different systems of management so that he can discuss them in detail, but only by practice do these things become an actual part of his beekeeping equipment. Merits of beekeeping courses. A good way to learn beekeeping is to attend some school where a thorough course in this subject is given. Until recently beekeeping was not included in the work of the agricultural colleges in the United States, but interest is now being aroused in this work and it is spreading in a manner to give deep satisfaction to those interested in the development of the industry. In the apiary of an experienced beekeeper, the beginner perhaps gets more personal attention than he does in a class, but usually in the rush of honey-production, the theoretical side of the work is neglected and frequently the beekeeper is not able to offer much help to his student on such subjects. Consequently when he begins to keep bees for himself, he may find local conditions quite unlike those in the apiary in which he worked and, not knowing the fundamental facts about bees, he may be at a loss to know what to do. In, a regular course of study, 226 Beekeeping the proper emphasis may be placed on the various subjects, although naturally there is less opportunity for practice with the bees. The ideal plan is to take the prescribed course and then spend the following summer in the apiary of the best beekeeper available. The student is then able to understand more clearly what he sees and hears and is better able to recognize and perhaps mentally to condemn the little peculiarities in practice which one encounters occasionally in the manipulations of practical beekeepers. If the teaching of beekeeping is conducted wisely, it should result not only in increased knowledge of bees, but in the training of more professional beekeepers. Beginner's outfit. It is only with experience that one is able to judge of the comparative merits of different hives and other equipment, but the beginner usually desires definite information concerning the equipment which should be purchased. The giving of such advice is attended with some risk, for one hesitates to advise an equipment which may be discarded when the beekeeper becomes more familiar with the business. In the following lists, the choice is made on the basis of the equipment which is preferred by the majority of "good beekeepers and not alone on the author's personal preferences. General equipment: — Bee veil. Smoker — medium size. Gloves (for the beginner only). Some kind of hive tool — a screwdriver will answer. For each colony: — Bottom board of 1 inch material. itf-frame Langstroth hive — preferably single-walled. Self-spacing frames, punched for wiring. Medium brood foundation, li Ib. for each hive body. Telescope cover. The First Steps in Beekeeping 227 For comb-honey production (minimum): — 3 supers for 10-frame hive (if possible one made up for sample). The 4J inches square section, If inches wide, is usually preferred. The purchase of only one super to the colony is to be condemned. Thin foundation, 2 oz. to super. For the beginner the use of small starters of foundation is sometimes preferable. If full sheets are used, 8 oz. to the super should be provided. For extracted-honey production: — 2 extra hive bodies identical with those used in brood chamber, with full sheets of comb-foundation (see p. 28 concerning spacing devices in surplus chambers). 1 2-frame non-reversible extractor. 1 Bingham uncapping knife. For bulk comb-honey production: — 3 10-frame supers with shallow extracting frames. | Ib. thin-super foundation for each super. CHAPTER XII1 THE APIARY SITE IN the establishment of a commercial apiary, the chief requisite is proximity to the sources of nectar. To fulfill this, it is essential that the honey resources of the region be studied carefully. While it is possible to keep bees in almost all of the habitable parts of the United States, it is not everywhere profitable to practice extensive beekeeping, so that if one contemplates making honey-production a major portion of his business, it is best to go to the best available location rather than to struggle along in a mediocre locality. A second requisite is ease of transportation to the apiary and to market. Apiary grounds. In the North, a plot of ground sloping to the south or east is usually preferable and in any region it is advisable to face the apiary so as to protect the entrances of the hives from the prevailing winds. If the contour of the land or a n6ar-by forest does not afford protection from the wind, a windbreak may be planted. A row of evergreens is efficient in the North and is serviceable in winter when it is most needed. A solid fence or building is less desirable, since such a windbreak, instead of breaking the force of the wind, often simply deflects the currents into the midst of the apiary with disastrous results. In moist regions, a slope is desirable for drainage. The hives should be so placed that they receive sunlight in the early morning. This is helpful in winter and perhaps equally so in summer as it starts the bees to the field earlier, so that they get the nectar from 228 The Apiary Site 229 plants which furnish it only in the mornings. If possible, the apiary should not be near the public road and should be situated where the bees will not prove a nuisance to passers-by or sting live-stock. If the only convenient location is near the road, the line of flight of the bees may be deflected upwards by a high hedge or a solid fence, for after they fly over such an obstruction they will keep above the line of travel on the road and will not molest teams or pedestrians. This is an important consideration as bees sometimes sting horses fatally. The apiary should preferably be located away from the clothes-drying yard so that they will not spot the clothes with their feces. This applies especially in the North, and this objection may largely be overcome by removing the cellar-wintered bees when no clothes are to be hung out. Perhaps it would be more in keeping with the usual practice to advise that no clothes be hung out just after the bees have been placed on their summer stands. The hives should, if practicable, occupy the higher ground of the plot chosen for the apiary, so that in carrying heavy supers to the apiary house the load will be carried downhill and the empty supers uphill. Such an arrangement will materially reduce the labor in a commercial apiary. Exposure to the sun. While exposure to the sun is to be advised in the early morning, it is often well to protect the hives from the sun in the middle of the day, so that the bees will not hang out in frpnt of the hive and to prevent the melting down of combs. On the other hand, too dense a shade is not advantageous and usually it is not best to locate an apiary in woods. To provide shade, the hives may be placed in two rows under a shed or arbor with the hive entrances to the outside (Fig. 98). Such sheds usually run north and south, but in hot, dry countries an east and west direction is sometimes better. In temperate climates, sheds are not in favor, but many beekeepers use shade-boards, so constructed that they extend about a foot in all directions from the hive except to 230 Beekeeping the north. These must be held in place by a heavy weight and are rather objectionable because they have to be removed each time the colony is manipulated. As the extreme of protection from the sun's rays may be mentioned the house apiary, in which the entrances to the hives are through holes in the wall of a specially constructed house. Such arrangements meet with little favor among American beekeepers because of the difficulty in manipulating the colonies inside the house. In Europe, however, the beekeepers often construct elaborately designed and decorated house apiaries (Fig. 8) in which an American com-mercial beekeeper would find himself seriously hindered. Care of the apiary grounds. The ground on which the hives are located should be smooth so that a wheelbarrow or cart may be utilized in carrying supplies or honey. This is also desirable if a lawnmower is used to keep grass and weeds from obstructing the entrances. High grass about the entrances is a hindrance to the bees on returning to the hive and should be avoided. Few commercial beekeepers find time to use a lawnmower during the rush season, but prefer to lay boards in front of the hive or to kill the grass with salt. It is sometimes convenient to pasture one or more sheep in the apiary inclosure. Raising the hives above the grass on high stands is another solution of the difficulty, but is not desirable in a heavy flow of honey, since bees often FIG. 98. — Apiary in the West, shaded by thatched shed. The Apiary Site 231 fall to the ground with their loads and since the stands may break down under the weight of honey. Arrangement of hives. The hives may be variously arranged according to the preference of the beekeeper. Each hive should be inclined FIG. 99. — Former apiary of the Bureau of Entomology, College Park, Md. The use of this apiary for experimental work accounts for the divergence in the height of the hives. The hives, were here arranged singly about four feet apart. . so that the entrance is about an inch lower than the back of the hive to prevent water from collecting on the bottom board. It is usually desirable that the hives be so placed that the beekeeper will not need to pass in front of the hive entrances as he goes about his work. The usual practice is to place hives in rows either close together on a slightly raised platform or singly on individual stands. The placing of hives in pairs on a single stand is also common. Where economy of space is a consideration, it is found advantageous 232 Beekeeping to place hives close together in groups of four, back to back, two facing east and two west. This allows a space beside each hive for the beekeeper while manipulating and is greatly to be preferred to hives in long rows close together. Where space will permit, the placing of hives singly (Fig. 99) is preferable. Number of colonies in one apiary. The number of colonies which may profitably be kept in one apiary depends entirely on the nectar supply. In the white clover region, it is considered best to have not more than one hundred colonies in an apiary, with apiaries located at least two miles apart. This number can be increased in many localities. In the other general honey regions of the United States, it is usually profitable under favorable local conditions to keep from 200 to possibly 500 colonies in one apiary. There are so many factors to consider in determining this point that no general rules may be laid down. Since this is not a question which the beginner is called on to answer, it may be advised that the beekeeper decide each case individually from a study of the honey flora, the experience of other beekeepers and his own experience. Out-apiaries. In locating an apiary away from the central apiary, usually called an out-apiary or out-yard, easy transportation, especially to the main apiary, is most desirable. It is also an advantage to have the out-yard, especially one in which comb-honey is produced, near to the home of some person who can hive swarms which may issue in spite of precautions taken, and to protect the colonies from depredation. If these things are not practical, it is better to have the bees where they are not easily seen from the highway. Conveniences less essential in out-apiaries. In establishing an out-apiary, the points previously mentioned should be* considered as desirable but not essential. The Apiary Site 233 Since bees may be kept on roofs, in woods and in other places lacking many desirable features, it will be seen that it is not profitable to consider the desirable features too seriously. Another distinction should be made. If the out-apiary is permanent, it will pay better to plan the location thoroughly. However, many commercial beekeepers, especially those outside the white clover belt, find it well to change the locations of their out-apiaries to meet changing conditions in this region and they therefore do not find it profitable to consider the conveniences in equipment and in apiary planning to any great extent. To the commercial beekeeper the only essentials are the things which bring the greatest return. The amateur can better afford to spend time cutting grass and arranging hive stands since his living does not depend on the crop and he has fewer colonies for which to plan. CHAPTER XIII THE MANIPULATION OF BEES THE work which the beekeeper does with his bees has for its object an increase in their productiveness. Bees gather nectar and pollen when they are available in response to their own instincts to gather; they build wax when it is needed if space and food are available. The duties of the beekeeper are not concerned with creating these impulses. However, bees do not always work so as to accomplish the most efficient results, when measured by the commercial standards of the beekeeper, and the care which he bestows on his bees serves to provide conditions suitable for the turning of their natural instincts into those channels which will yield the greatest profit. Disturbance to be reduced to a minimum. Bees should be handled so that their work will be disturbed as little as possible, for the manipulations of the beekeeper are only accessory to their labors. Stings should be avoided. This is not so much because they are painful, but chiefly because the odor of the poison irritates bees and makes them difficult to manage. A veil (Fig. 26) and a good smoker (Fig. 24) are practically indispensable. By the use of smoke, the bees may be quieted so that they may be handled readily, the guards are disorganized and the bees gorge themselves with honey, after which they are not easily provoked to an attack. Too much smoke must be avoided as it disorganizes the entire colony and considerable time elapses before the bees fully return to their normal activities. 234 The Manipulation of Bees 235 Hasty movements and the jarring of the hive are to be avoided. The organization of the bee's eyes enables it to see movement more readily than still objects. On seeing bees flying about the face, the beginner often strikes at them or moves quickly to escape the sting, thus provoking an attack. It requires quiet nerves not to jerk a frame or even to drop it when the hand in which it is being held is stung. Equipment for manipulation. Aside from a smoker, veil and hive-tool, the beekeeper needs no other equipment in opening a hive, but the beginner may find gloves (better those with the fingers removed) desirable. If special clothing is worn in the apiary, and it is desirable for both comfort and economy not to wear'one's best, white suits are most satisfactory. They are the most comfortable in the heat of summer and the beekeeper has a good excuse for this comfort because they are best for apiary use. When to handle bees. The best time to open hives is in the middle of warm days, especially when the bees are busily engaged in collecting nectar. Bees should never be handled at night nor on w^t, cold days. It is not always possible for the extensive beekeeper to choose the ideal time but it is well to plan to open hives in favorable times, not only for the comfort of the operator but principally because it interferes least with the work of the colony. Opening a hive. Before opening the hive, a little smoke should be blown in the entrance. When the cover is slightly raised, a little more smoke should be directed over the frames before the bees have an opportunity to escape. If the frames are covered by a mat or oilcloth, which is not desirable but often used, the outer cover may be entirely removed and one 236 Beekeeping corner of the mat lifted to admit smoke. The covering then may be removed and the manipulation begun. In case the bees become troublesome at any time during the work, as they probably will if it is continued for a time, more smoke may be blown over or directed down between the frames to disorganize new guards. No directions need be given as to the way to recognize trouble and it need only be stated that the most common fault is to use smoke too freely. Inuring all manipulations the operator should stand at the side or back of the hive and not in front of it, to prevent interference of bees leaving and returning to the hive. If one wishes to examine the brood chamber when the colony is in two or more hive bodies or has FIG. 100. -Hive-body resting on cc^er COmb-honey Supers, the during manipulation. hive £OVCr Serves as a good support for the removed bodies. They are placed diagonally on the cover (Fig. 100) with only four points of support, ttius avoiding the crushing of bees. If a second body is removed, it may be placed out of line on the first in the same manner. If the bees show signs of robbing, combs in removed bodies should be more carefully protected. Remedies for stings. Various remedies for stings have been advocated but they are all valueless. The puncture made by the sting is so small that no liquid can enter it after the sting is removed and the opening has closed. As soon as practical, immediately if possible, the sting should be removed, care being taken not to squeeze the attached poison sac. This can be done by scraping the sting out with a knife blade or the finger nail. After this is done the injured spot should not be The Manipulation of Bees 237 rubbed and the* usual advice of the beekeeper is to "forget it/' Bathing with liniment or any other irritation serves only to spread the poison through the tissues. The intense itching soon disappears. As a comfort to the novice, it may be stated that repeated stings usually cause an immunity to the poison to develop, after which no after-swelling occurs. In case of severe stinging, the injured parts may be covered with an ice bag or with cloth wet with ice water. Removing frames. After the frames are exposed, the propolis which often fastens them may be loosened by prying gently with a hive-tool and the frames may be crowded somewhat closer together to permit the removal of one of them. It is immaterial which frame is removed first, unless the special object for opening the hive determines it. In cool weather.the propolis may be brittle and care should be taken not to jar the hive as this is broken. During manipulation, a side frame is often removed and leaned against the hive to allow more room for moving the other frames. In leaning a frame against the hive, it should be in a nearly upright position to prevent breakage and leaking of honey. The frame on which the queen is located should not be left outside the hive unless necessary, for she may crawl away and be lost. The frame should be leaned against the hive on the side away from the operator so that he will not be annoyed by bees crawling up his legs. In all the handling of the colony, bees should not be crushed, for this excites the others, and if frames are crowded too closely together the queen may be killed. Handling frames. In examining a comb, it should be held so that if any bees fall they will drop into the hive, except when it is necessary to carry away a frame for some purpose. Freshly gathered nectar sometimes drops out if the comb is improperly handled. If this falls into the hive the bees clean it up, whereas outside 238 Beekeeping the hive it may cause robbing and is at least untidy. The •beginner should early form the habit of keeping combs in a vertical position. While sometimes it does no harm to tip a frame, it is rarely necessary and may cause honey to leak or the comb to break, especially if the frame is not wired. As a comb is taken from the hive, it should be lifted by the ends of the top-FIG. 101. — Handling a frame, first position, bar, two hands being used. This brings the comb up vertically with one side toward the operator (Fig. 101). To examine the reverse side without tilting the comb, raise one end of the top-bar until it is perpendicular (Fig. 102), turn the frame on the top-bar as an axis until the reverse side is brought into view, and then lower to a horizontal position with the top- FIG. 102. — Handling a frame, second position. FIG. 103. — Handling a frame, third position. bar below (Fig. 103). In actual practice these steps are not taken successively but the turning on the top-bar is simultaneous with the raising and lowering of the end of the frame. This is a good operation for the beginner to practice a few times. The Manipulation of Bees 239 Desirability of straight combs in manipulation. The use of comb-foundation in the frames is desirable to insure uniform comb, all of worker cells, except in places where the foundation may sag or become torn. Drone comb is undesirable except in raising drones for queen-rearing. The use of comb-foundation in wired frames insures straight combs and reduces the danger of bees being crushed in removing or in returning frames. A frame-hive with combs built crosswise is more difficult to handle than a box-hive and this should never be permitted. The entrance of the hive should be exactly horizontal so that the combs will hang parallel with the sides of the hive and so that the outer ones are not fastened to the hive-body, if they are properly spaced at the top. The back of the hive should be about one inch higher than the front to allow condensed moisture to escape. A hive leveling device made by Burton N. Gates is shown in the accompanying illustration (Fig. 104) which needs no description. This has been found useful, especially with the tile hive-stands used in the Bureau of Entomology apiary. FIG. 104. — Hive leveling device. In the upper figure the wedge-shaped piece is on edge so that when the top is level the hive entrance is one inch lower than the back. In the lower figure the wedge is on its side. 240 / Beekeeping Closing the hive. In closing a hive, after the frames are replaced and spaced properly, the cover should be put on in such a way as to crush no bees. If necessary the bees may be driven down by the use of smoke, but if bees are on the top edges of the hive, the cover may be slid on from the end or side so that none will be crushed. OCCASIONAL MANIPULATIONS In the discussions which follow, manipulations will be described which may be useful at almost any time or at different times in the season but which are not part of the regular work of most apiaries. The plan followed in this book is to give the various manipulations in the order in which they are used during the season. The manipulations discussed under this heading are most frequently useful in the spring. Feeding. To stimulate brood-rearing or to provide stores in the spring, in preparing colonies for winter and at other times during a shortage of stores, it may be necessary to feed the bees. Obviously, it is desirable to allow the bees to keep sufficient honey and if this can be done it is always preferable to feeding. No better stimulation to heavy breeding in the spring can be found than adequate protection and an abundance of stores, but a large amount of food is needed at this season and the beekeeper should feed if he finds that he has failed to leave enough. In small hives, the giving of additional stores in the spring is usually desirable, either in the form of combs of honey or as a syrup. The feeding of sugar syrup to produce comb-honey has of course been tried and some beekeepers have believed that the product is honey. This is not the case and the fraud may readily be detected. Fortunately, even at the lowest prices of granulated sugar, the sections actually cost the The Manipulation of Bees 241 beekeeper more than he gets for pure comb-honey and this fact effectually keeps adulterated comb-honey off the market. What to feed. Honey from an unknown source should never be fed, because of the danger of introducing disease. Beekeepers usually feel that it is cheaper to feed sugar syrup because of the higher market value of honey, but no food for bees better than honey has yet been found. If extracted-honey is fed, it should be somewhat diluted. The best plan is to give combs of honey. As a substitute for honey, a syrup made of granulated sugar is best. For spring feeding, a thin syrup may be used, even as dilute as two parts of water to one of sugar (by volume). Ordinarily equal parts of each are used. For supplying winter stores, a thick syrup, 2} to 2| parts of sugar to one of water, is preferable. To prevent granulation of the sugar in the thick syrup, it is inverted (changed chemically to levulose and dextrose) by the addition of a teaspoonful of tartaric acid to 20 pounds of sugar while the syrup is being heated to dissolve the sugar crystals. In early spring and late fall, syrup may be fed while warm and fall feeding should be done as rapidly as the bees will take the syrup. In making syrup, the greatest care must be taken not to allow it to be discolored by scorching the sugar; it should be as clear as if made with cold water. Glucose, other cheap syrups or molasses and the cheaper grades of sugar should not be fed to bees, especially for winter stores, since they contain substances indigestible to bees, causing dysentery. Candy and cube sugar are sometimes used for supplying bees in winter after their stores are exhausted. These should be used only in emergency and nothing but granulated sugar should be used in making candy for this purpose. A soft white sugar, known in the trade as "coffee A," placed in a division board feeder is sometimes used as a stimulant to brood-rearing. 242 Beekeeping Slow feeding to prevent robbing is sometimes desirable during extracting or other manipulations or while rearing queens. A thin syrup of one part sugar to nine of water is used, being fed in large feeders in the open. FIG. 105. — Division board (Doolittle) feeder. FIG. 106. — Alexander feeder in collar under hive-body. Feeders. There are numerous types of feeders, used for different purposes. The division board feeder (Fig. 105) is hung in the hive like a frame. It may be filled without being re- FIG. 107. — " Pepper-box" feeder. FIG. 108. — Pan in super arranged for feeding. The Manipulation of Bees 243 moved and a float must be used to prevent bees from drowning. The Alexander feeder (Fig. 106) is useful mainly for stimulation. It may be placed under the rear of the brood-chamber if the bottom board is moved forward, but this often causes robbing and a better plan is to place it in a collar under the brood-chamber as shown in the illustration. For feeding small quantities, a "pepper-box feeder" (Fig. 107) may be inverted over the brood frames in an empty hive-body. Mason jars may be used in the same way, special pierced covers being sold by dealers in beekeeping supplies. For rapid feeding in the fall, a large pan in an empty super (Fig. 108) is perhaps best. Green grass should be thrown in the syrup to give support to the bees while feeding, this being better than excelsior or chips as it does not absorb the syrup. The empty super and pan may be placed under the brood-chamber for late feeding, in which case the bees take the syrup more rapidly. Uniting. If a colony becomes queenless in late fall, it is usually not profitable to give it a queen, but it may be united with a normal colony to save the bees. It is not wise to try to winter weak colonies, but if two or more are united to make a strong colony, keeping the best queen, the risk in wintering is reduced and better results are obtained in the spring. It is also more profitable to unite weak colonies in the spring than to build them up, Influence of hive odor. Every colony of bees has a distinctive odor by which bees recognize individuals from their own colony, normally resenting the entrance to their hive of those from other colonies. In uniting colonies, the different odors may be hidden by smoking both vigorously. Tobacco smoke may be used, but if too much is employed the bees become stupefied. If bees are stupefied by tobacco smoke, chloroform or other anaesthetics, they lose their memory of former locations 244 Beekeeping and may be united and placed wherever desired, but American beekeepers rarely have occasion to use such methods. During a honey-flow, when the field bees are coming in heavily laden with nectar, the field bees of two colonies that are close together may be allowed to enter one hive and they do not molest each other. The queen to be saved should be caged for a day or two to overcome the danger of the strange bees killing her. When brood-rearing is reduced, as in the fall, the colony odor is apparently less influential, for less precaution is necessary in uniting. Learning the new location. . Field bees return to the location of their hive and they remember the old location and return to it if the hive is moved. If two colonies to be united are not close together, they should be moved gradually nearer, perhaps a foot every day that the bees can fly, until they are side by side. The bees learn each location in succession anli after uniting they will not return to the original position and be lost. If it is necessary to move the colonies faster, they may be put into the new place and a pile of brush or weeds or a slanting board placed in front of the entrance so that when the bees fly out they will perceive a change and learn the new location. If it is desired to unite two weak swarms, this may be done simply by placing them together, either in the hive or on the hive entrance. Swarming bees abandon the memory of the old location (p. 180); they are full of honey and may be placed anywhere. The better queen should be saved and the other removed or the bees may separate into clusters. Swarms may be added to newly established colonies if desired. If queenless colonies are found in early spring which are to be united with normal colonies, the usual practice is to place them on top of the normal colony. Few bees return and there is usually no trouble as such bees seem ready to go to any colony. The Manipulation of Bees 245 Transferring. In increasing the apiary, it is sometimes profitable to buy colonies in box-hives because of their small .cost. They should, of course, be transferred to movable-frame hives as speedily as possible, for bees in box-hives are of small value as producers, because of the impossibility of manipulating the combs. The advice is often given to beginners to buy colonies in box-hives and transfer them, but this advice is questionable. There is no more trying work connected with beekeeping, unless possibly it is the moving of a large apiary, and if a beginner can successfully transfer a colony from a box-hive he has proved his right to become a beekeeper. The best time to transfer colonies, if there is opportunity for choice, is the spring (during fruit bloom in the North) when the amount of honey and the population of the colony are at a minimum. However, the work can be done at any time during the active season, but there should be nectar coming to the hives so that while combs are exposed robbing will not be induced. If necessary, transferring may be done in a tent or cage of netting or wire cloth to keep robbers away, but the odor of honey may cause excitement in the apiary. If the field bees are out of the hive, the work is lessened. Methods. There are several methods of transferring and one may be chosen according to the plans and wishes of the beekeeper. Plan 1. — The box-hive is set a few feet to one side and in its place is put a hive with movable frames, containing full sheets of foundation or drawn combs. As the field bees return, they go at once to the new hive. The box-hive is turned upside down and a small box is inverted over it. The box-hive is now pounded continuously (the operation being known as drumming or driving) in such a manner as to transmit the jar to the combs and the bees desert their combs for the upper box. They cluster in this box like a 246 Beekeeping natural swarm and they may then be thrown in the new hive. If possible, the queen should be seen so that the operator may be sure that she is off the old combs. It is necessary that she be obtained, unless one desires to requeen at this time, in which event the old queen should be captured and the new one may be run in with the bees and will be promptly accepted. The box-hive containing the brood is now placed right side up in a new location. In 21 days all of the worker brood will have emerged and possibly some new queens will be reared. These bees may be driven out and united with their former hive-mates by allowing them to run in the entrance. They should, however, be compelled to go through perforated zinc or a queen and drone trap (Fig. 30) to keep out the young queens. The old combs may now be melted after removing the honey. By this method straight combs are obtained. If nectar is not being collected, the newly established colony should be fed. Plan 2. — Wait until a swarm issues from the box-hive and then move the old hive to a new location. The swarm is then placed in a new hive on the old stand and it is further increased by returning field bees. After 21 days the bees which have emerged are united with the bees in the new hive, as described under Plan 1. Plan 3. — If the beekeeper desires to save the combs in the box-hive, the bees may be drummed into a box, after FIG. 109. — Cutting combs from a box-hive. The Manipulation of Bees 247 which the brood combs and any other good combs are cut out (Fig. 109) and fitted into frames, being fastened with string, rubber bands or strips of wood until the bees have an opportunity to repair them. These frames are hung in a hive on the old stand and the bees are then allowed to run in. The cutting of combs, especially those containing brood and honey, is a disagreeable job and, since combs from a box-hive are usually of little value, this method is not recommended. Plan 4. — Another method which is in some respects better than those just given is to place the box-hive with its largest surface uppermost. If the bottom is now open, it is closed except for an entrance and a piece is removed from the upper side of the box-hive. The hive in which the colony is to be located is now put over the large opening and all cracks and openings around it are closed. The upper hive is filled with drawn combs or, if these are not available, with sheets of foundation. When the queen needs more room for egg-laying, she will go to the upper hive and, after she is located there, a queen excluder is put between the box-hive and the new hive to prevent her return. As the brood emerges below, the colony becomes established above. If there is difficulty in getting the queen'to go to the new hive, the box-hive may be drummed. After the brood in the old combs has all emerged, the bees may be drummed from the box-hive and it may be treated as desired. Transferring from walls of houses. Swarms often locate in the walls of houses and it is sometimes necessary to remove them to prevent damage from melting combs. If the cavity in which the combs are built is accessible, the method is the same as in transferring under the third plan, except that drumming is impractical and the combs must be cut out with the bees still adhering to them. A liberal use of smoke will subdue them. If it is impossible to open the cavity without doing considerable damage to the building, a bee-escape (Fig. 19) may be put 248 Beekeeping over the entrance so that the bees can leave but cannot return, any other openings to the combs being carefully closed. Even better than a bee-escape is a cone of wire-cloth eight inches high with a hole at the apex, just large enough for a single bee to pass. This is tacked on the house and the bees issue through the hole in the apex but do not find it again to return. A hive (with drawn combs in it if possible) is then placed so that its entrance is as near as practical to the entrance which the bees have been using. A queen should soon be introduced to the bees in the hive. The old queen does not desert her combs and continues laying eggs, but, as her colony is depopulated, the amount of brood rapidly diminishes. As brood emerges, the young bees also leave through the wire-cloth cone and join the bees in the hive until in four or five weeks the queen is left practically alone. When nearly all of the bees are out of the cavity and there is little or no brood, the bee-escape is removed, the entrance to the cavity is made larger if possible and if there is no honey-flow, the bees rob their old home and carry the honey to their new hive, leaving only the empty combs. These will usually do no damage as wax-moths soon destroy them. The entrance to the cavity should now be carefully closed to prevent another swarm from taking up quarters there and the-hive is removed. This method takes considerable time, but is often best where the cavity is inaccessible. It is often difficult to close the cavity to prevent the bees from establishing a new entrance when a bee-escape is placed over the one to which they are accustomed. Transferring from hollow trees. The method to be used will depend on the accessibility of the cavity and the value of the tree. Usually the bees cannot be drummed out and the combs must be cut out after subduing the bees with smoke. If the colony is high in the tree and the tree is felled, the bees are disorganized by the jarring so that they can be handled easily. The hunting The Manipulation of Bees 249 of colonies in the woods is interesting and the cutting of a bee tree is an experience which everyone interested in bees should have, but the time consumed is considerably more than the value of the bees and honey justify. It does not pay to build up the apiary in this way if the beekeeper's time is valuable. Preventing robbing in the apiary. At any time during warm weather, bees are inclined to rob other colonies, if there is no honey-flow. Every precaution should be taken to prevent this. Feeding often attracts other bees and, if there are indications of robbing, it should be deferred until late in the day. Honey left where bees can get it or combs left out of the hives during manipulations may at times lead to serious robbing. As soon as robbers are noticed, manipulation should be discontinued and the hives should be closed. If serious robbing occurs, the entrances should be contracted and the hive fronts wiped with a cloth moistened with kerosene or carbolic acid. If brush is thrown at the entrances, robbers are less likely to enter. Outdoor feeding to prevent robbing is described in a previous section (p. 242). A wire-cloth cage, five feet square and six feet high, may be used if manipulations are necessary during a time when robbing is probable. This cage should not be closed at the top and bees which fly from the colony under manipulation escape, while robbers will rarely enter. A folding tent or cage made of mosquito netting may also be used. A smaller wire cage closed at the top may be set over a colony that is being robbed. If the cause of robbing is suddenly removed, this may produce more excitement than if the robber bees were allowed to complete their work. For example, if a colony is being robbed and is suddenly removed to save it, the robbers turn their attention to other colonies, or if a piece of exposed comb has attracted robbers, its removal may divert them to more serious devastation. 250 Beekeeping The beginner in beekeeping may mistake the play flights of young bees for robbing, but after the latter has once been observed this error will not be repeated. Bees appear old soon after they begin robbing; they are dark and thin, their actions are nervous, and the hairs on the body are lost, probably by being torn off by defending bees and by squeezing through narrow openings. When a colony is abnormal, as in queenlessness or disease, it may be robbed of its stores slowly, without any excitement, usually by the bees of a single colony. Moving bees. In migratory beekeeping, in bringing purchased bees to the apiary and under various other circumstances the beekeeper will find it necessary to move bees. The frames must be fastened firmly in place. If self-spacing frames are used, especially if there is considerable propolis on them, no precautions need be taken to prevent the swinging of the frames, but it is often desirable to nail a f-inch strip over the tops of the frames so that they cannot fall out of place if the hive is tipped. Abundant ventilation should be given, the amount depending on the temperature. In cold weather, the entrances may simply be closed with a block and the cover fastened securely, but in extremely hot weather it is desirable to remove the cover or both the cover and the bottom board and nail on wire-cloth (Fig. 110). A 2-inch collar may be fastened to the top of the hive-body and wire-cloth put above this, to give clustering space for the bees. Colonies rarely suffer from exposure during moving so that the FIG. 110. — Hive ready for moving. In this case the bottom board is left on. The Manipulation of Bees 251 beekeeper should not err by giving inadequate ventilation. In transporting colonies on a wagon, the length of the frames should be across the wagon bed, while on a train they should be parallel with the length of the car. It is customary to ship colonies in hives with combs, but recently the shipping of bees in wire-cloth cages without combs has been practiced. This has much to commend it, especially in reducing the danger of introducing bee diseases into new locations. The cages contain numerous slats on which the bees hang, and they are provided with food for the bees en route. The bees are put into the cage by means of a funnel, either with or without a queen. Bees are now regularly sold by the pound in such packages and shipped to all parts of the United States. It is probable that as this method is perfected it will be used in migratory beekeeping, thus avoiding danger of carrying disease to the home apiary and reducing the transportation charges. ELIMINATION OF NON-ESSENTIAL MANIPULATIONS While it is necessary in any discussion of beekeeping which aims at completeness to describe the various manipulations which may be needed during the course of the year, the beekeeper should early in his experience establish a system for the care of his bees so that unessential movements and manipulations may be avoided. If bees are kept solely for pleasure, it matters little whether they are disturbed in their work, and the time of the beekeeper need not be considered an important consideration, but when bees are kept for profit, these factors become vital. Every manipulation which does not benefit the beekeeper by increasing his profit should be ruthlessly eliminated, and every time a colony is opened it should be for some definite purpose. Two essentials. There are two factors necessary to the production of the maximum honey-crop over which the beekeeper has no 252 Beekeeping control. He cannot govern the weather or produce honey-plants with profit. There are on the other hand two other factors with which his work must deal. His efforts should be for the purpose (1) of getting plenty of bees of the right age in time for the harvest and (2) of keeping these bees in proper condition for gathering the maximum crop. The first essential is far-reaching and obviously includes the entire care of the colonies to prevent starvation or loss from other causes. It applies especially to the work in the spring. The second essential applies chiefly to the control of swarming. It is well for the beekeeper to keep these two essentials always before him and to ask himself, when he plans any work with the bees, whether it comes under one of th$Se heads. /^Fne beekeeper may profitably go one step further in the analysis. For example, stimulative feeding in the spring is mentioned earlier in this chapter. He should first of all determine whether stimulative feeding is more profitable than the giving of abundant stores. If he finds that he gets more bees by stimulative feeding, he should then determine whether he gets enough more to justify the expenditure of time and money, or whether he can get a larger total crop by keeping a few more colonies, combined with the giving of abundant stores. An example taken from life may not be amiss, the names being here omitted. Two beekeepers are located in exceptional situations which maybe assumed to be equally good. One of these men is skilled in the improvement of his stock and has made significant progress, but the work occupies considerable time. The other beekeeper feels that he has not the time for this (and he may not have the skill), but he keeps 100 colonies more than his co-worker. In the case just given the beekeeper with the larger number of colonies makes more money, but this illustration is by no means given to discourage breeding work. It shows, however, that for that particular region the greater profits come with extensive beekeeping, while in other regions more intensive work might yield better financial returns. The Manipulation of Bees 253 Both men have the same object in view — to produce workers on time for the harvest. Both are successful while all about them are beekeepers with indifferent or poor success, attributed probably to bad luck. Increase in efficiency through system. Not only must these essentials be emphasized, but the necessary manipulations must be systematized. After some effort in this respect, the beekeeper is usually astonished at what may be accomplished. This may be illustrated by another case. This beekeeper began work with bees on a business basis after keeping a few colonies for pleasure for several years. At the beginning of his experience he taught school, thus having his Saturdays for the bees as well as the summer vacation. At first the vacation was six months, but later the school year was increased, giving him only three months. By systematizing his work, he was able to do as much as formerly and gradually increased his colonies to 250 in three apiaries, all run for comb-honey. He then gave up teaching and accepted a position which kept him away from his bees except during thirty days' vacation in the summer. At first he was frequently near his bees so that if any work was necessary he could arrange to have it done by others. Finally he accepted another position which took him entirely away from his old home and he now goes back just before the honey-flows and leaves as soon as they are over. He still produces comb-honey and is still successful. He has probably almost reached the maximum number of colonies that he can run for comb-honey in so short a time. It is obvious that many manipulations usually considered necessary must be eliminated in these apiaries. By leaving plenty of stores and by giving the bees abundant protection many of these are rendered unnecessary. This case is not by any means recorded as ideal, but it illustrates what the elimination of superfluous manipulations may accomplish. Anyone can produce honey in a time of plenty, but only 254 Beekeeping the good beekeeper gets an adequate return in less abundant seasons. The ideals toward which the beekeeper should work are: (1) to handle the bees as little as possible; (2) to manipulate them only when he has some definite object in view; (3) to follow a definite system, not based on rules but on a knowledge of bees, capable of modification as occasion may arise, but working for one end — maximum honey-production. CHAPTER XIV SPRING MANAGEMENT IN attempting to give the work of the apiary in chronological order, it is difficult to decide where to begin. To a large degree, success depends on the results obtained in wintering, so that preparation for winter might be considered the first step in the annual cycle, and practical beekeepers usually so consider it. However, winter is a period during which the beekeeper has little work with his bees, and it is perhaps better to begin the cycle with the first evidences of activity outside the hive. As has been shown, bees do not< hibernate, and consequently their early flights are not evidences of an awakening after a period of inactivity. With their first return to the open air in the spring, the beekeeper knows that the active season with his bees has arrived. As will be shown in the chapter on wintering (see also p. 91), bees are often compelled to retain their feces for long periods in winter. This, together with the excessive generation of heat, may deplete the colony, causing conditions Imown as spring dwindling and dysentery, one or both of whipti may be present. l/li will also be shown later that it is not desirable to manipulate bees in winter. Brood-rearing may begin during the severe weather of January or February in the North in colonies wintered out of doors, but this can scarcely be considered as an activity of spring. With the opening of the earliest spring flowers and the accompanying rise in temperature, the bees venture forth to get the small amounts of nectar and pollen thus provided. 255 256 Beekeeping As the weather becomes warmer the supplies rapidly increase and the bees are greatly stimulated to build up the colony. The old bees that emerged the previous autumn have been called upon, under ordinary winter conditions, to expend considerable energy, and their ability to do the collecting and the inside work in the spring is in general in inverse ratio to the expenditure of energy in the winter. Brood-rearing, however, begins before or as soon as new supplies come to the hive, provided, of course, that the colony is normal, and as the first bees emerge they in turn increase the capacity of the colony for brood-rearing, so that with a good queen and other favorable conditions the brood is rapidly increased. Object of spring manipulations. The main object of the work in the spring is to insure an abundance of bees in time for the harvest. In the more northern localities, summer comes on with a rush and often the principal nectar-secreting plants are in bloom so soon after cold weather that the colony is frequently not in condition to obtain the maximum crop, or there may be a period in the spring when, from lack of nectar, the bees are not stimulated to the maximum breeding. If left to themselves and if honey is already present in the hive, bees will naturally rear brood and thereby rapidly increase the size of the colony, and the work of the beekeeper is to provide the most favorable conditions for the manifestation of this instinct. Prevention of drifting. Colonies which are wintered in the cellar need not be put in the same locations that they occupied the previous year, when they are removed. In setting them out, some care is necessary to prevent mixing. If they can fly as soon as they are set out, they may rush forth and then be unable again to, locate the proper hive, in which case they often drift^that is, bees enter the wrong colonies with the result some colonies will be increased in size at the expense others, If bees can be set out at night or on a cloudy or Spring Management 257 chilly day, this is generally avoided. The entrances may also be reduced or, if necessary, may be closed with wet cloths. Spring protection. If the colonies have been wintered in the cellar, breeding will normally not begin so soon as in colonies that were left outside. When the hives are carried to their summer stands, the bees are subjected to sudden changes in temperature and to low temperatures and, unless the wintering has been exceptionally good, they may be able to withstand adverse conditions less well than colonies that were wintered in the open. It is therefore preferable to provide packing for these colonies, even if it is only a wrapping of waterproof paper over the hive. After colonies are removed, the bees need a cleansing flight to rid themselves of the accumulated feces and they should be put out at a time when this will probably soon be possible. First examinations. During early breeding, the beekeeper ought to have no occasion to open a hive, but, if he finds that certain colonies are not up to standard, he may choose a warm day to open them to do whatever conditions ' may demand. After a winter away from the bees, the beekeeper is usually anxious to look at them. On a fine warm day when the bees are flying freely, he should make his first general examination of the apiary but, if he has previously supplied the colonies with abundant stores and has them protected from changes of temperature, he may well put off a general examination of the apiary. If he desires to learn whether the bees have sufficient stores he can determine this by lifting the hive, or the size of the cluster may be determined, without breaking the propolis which seals the cover, by looking at the combs from below. On the first examination, the beekeeper should look especially for queenless colonies. If any are found, it is 258 Beekeeping best to unite these with normal colonies, although queens may now be obtained early from southern breeders. He should also examine the stores, for bees require large amounts of food during the spring and, while they usually get considerable nectar, it rarely is enough to provide stores for excessive breeding. If food is needed, it may be given rapidly in the form of a thick sugar syrup, or it is even better to give combs of honey. If hives are soiled with the spottings of dysentery or if there are dead bees present, the hives may be cleaned out somewhat, but the first examination should be brief, unless the weather is exceptionally warm. Spring dwindling. The old bees die rapidly and are replaced by young bees, which, in a good colony, emerge more rapidly than the old ones disappear. If, on the contrary, mortality among the old bees exceeds the rate of emergence, the condition arises which is known as spring dwindling. Obviously, prevention is better than treatment, but by giving extra protection and by making the collection of stores unnecessary by feeding, the energy of the old bees may be conserved so that it is utilized chiefly in rearing brood and the colony may often be saved. The brood-chamber may also be reduced to conserve the heat of the cluster. Need of water. Bees need water for brood-rearing and it sometimes happens in the spring that bees are lost in trying to obtain it. If there is no water close at hand, it is often advantageous to provide a watering place in a warm sheltered spot in or near^he apiary. ^njniiinq. If exceptionally weak colonies are found, it is economy not to attempt to build them up, but to unite them. In uniting colonies in the spring, two weak colonies should not be placed together, but a weak colony should be placed Spring Management 259 with a strong one. If desired, the number can be restored by subsequent division. This is one of the most important points in spring management. Cleaning the hives. When the weather becomes settled, it is desirable, especially where comb-honey is produced, to subject the hive to a spring house-cleaning. If the bottom board is cleaned of debris and the propolis is scraped from the frames and rabbets, it will not only facilitate future manipulations but, when the sections are put on, there will be less propolis available to discolor them. Beekeepers, however, are not so devoted to a spring house-cleaning as are housewives. While Caucasian bees were kept in the apiary of the Bureau of Entomology the removal of propolis in the spring was practically a necessity. This may be done quickly in the spring, while the propolis is brittle. Dr. Miller uses a hoe to remove propolis and burr combs from the top-bars of the brood frames. Equalizing the colonies. Not all colonies increase in population equally fast, even with the best of management. The differences may be due to a variety of causes. If some colonies have more stores than they need, thereby reducing the space available for brood-rearing, combs of honey may be removed and given to colonies that need more stores, returning to the rich colonies empty combs removed from those to which honey is given. Similarly, if some hives contain more brood than the average, colonies may be equalized by taking combs of emerging brood with the adhering workers away from those abundantly supplied, giving them to weaker colonies, care being exercised not to transfer the queen. The weakest colonies in the apiary should be assisted in this way only after all the others are equalized; -then they are given any frames of brood still available, and are thus built up as rapidly as possible. Another method of equalizing is to shake bees 260 Beekeeping from the frames of a strong colony in front of the entrances of those to be helped. The young bees go in and are accepted, while the field bees return to their original hive. The queen course, not be shaken in this way. The advantages colonies develop at about the same rate and of reaching the honey-flow equally strong in bees, are as follows : (1) the colonies are ready for a given manipulation at the same time, thus allowing the work to be well systematized ; (2) less hive-bodies are needed than if strong colonies are given supers in accordance with their individual needs; (3) when properly done, equalizing probably results in an actual increase in the total number of bees in the apiary, since every queen is more nearly capable of egg-laying to her full capacity and no queen is restricted by having only a small number of workers to feed her brood; (4) less manipulation is necessary when the honey-flow begins (especially in comb-honey production) in sorting combs of brood and in reducing the brood to one hive-body, if this is practiced; (5) the brood is compact and this is especially desirable in comb-honey production. The work of equalizing colonies is considerable and the beekeeper must determine for himself whether it is profitable. In the management of out-apiaries, this work necessitates extra trips which come at a time when the average beekeeper has all the work that he can do. Making colonies all in one mold often fails to bring to light the deficiencies of some queens and the superior qualities of others for breeding stock. If a brood disease is present or is even known to be present in the neighborhood, there is danger in moving combs about so freely. Clipping queens. The clipping of the wings of queens is advantageous in the control of swarms, as will be shown later, and to find queens in the spring is easier than later. In clipping the queen, she is lifted from the comb, held securely but gently between the thumb and index finger of the left hand and a Spring Management 261 wing is cut off with fine scissors. The queen may be held with her wing against wood, when it may be cut with a sharp penknife, but scissors are safer, at least for the beginner. Some beekeepers clip the queen's wing when she is introduced, in case queens are mated from nuclei, but some colonies may rear queens without the knowledge of the owner and an examination for undipped queens in the spring will greatly reduce swarms which issue with queens capable of flight. Some beekeepers, so that they can tell a queen's age, clip opposite wings on alternate years or make a distinctive cut each year. It may perhaps be well to warn the beginner against clipping the wings of an unmated queen. If egg-laying is progressing regularly the queen is of course mated. Summary of favorable spring conditions. The conditions favorable to the rapid increase in the size of the colony in the spring may be restated as follows: (1) a large number of vigorous workers, due to successful wintering, (2) a prolific queen, (3) abundant stores properly located in the hive so as to be easily accessible to the bees, (4) favorable weather conditions, (5) fresh pollen and nectar and water for the bees, (6) a prolific race or strain of bees, (7) good brood combs of worker cells in quantity sufficient for the needs of the colony. QUESTIONABLE MANIPULATIONS The manipulations previously discussed in this chapter are not all practiced by all beekeepers but they are not the subject of controversy. On the other hand, there are two spring manipulations that have been the subject of much discussion by beekeepers and they are still in dispute. These are spring stimulative feeding and the spreading of the brood. Stimulative feeding. So long as nectar is coming to the hive in abundance, the colony is stimulated to the maximum brood-rearing possible 262 Beekeeping in proportion to the population. Frequently, however, there are periods when from a lack of nectar-secretion or inclement weather, no fresh supply of food is obtained. It is asserted by some that the feeding of a thin sugar syrup at this time in small quantities acts just as a natural honey-flow, stimulating the bees to greater activity in brood-rearing and resulting in the maximum strength of colony at the time of the honey-flow. Since feeding requires some manipulation of the colony which is not beneficial in inclement weather, many beekeepers believe that by providing an abundance of food in the fall or by giving stores rapidly in the spring the colony receives all the stimulus to brood-rearing that it should have and that stimulative feeding is not desirable. This is obviously a question especially for the northern beekeeper. If a colony is short of stores, combs of honey may be given it. If a colony has wintered well, has a good queen, is in a large hive abundantly supplied with stores and is well protected from changes in temperature, it is doubtful whether it can be stimulated to much greater brood-rearing than these conditions will bring about. Even if stimulative feeding results in increased brood-rearing, as it may under some circumstances, the beekeeper may still find it to be an unprofitable practice. If he is managing several apiaries, the work of going to all of them daily, or even less frequently, is considerable, and he may find it more profitable simply to operate more colonies to make up for the difference. If stimulative feeding is practiced, it is usually best to feed warm syrup in the evenings so that the bees will not fly as a result of the feeding and so that robbing will not be started. Spreading the brood. Spreading the brood is an even more debated question. If the brood-cluster is divided and an empty comb is inserted, the bees will attempt to cover all the brood and, in so doing, that part of the empty comb which intervenes will be kept warm enough so that the queen will lay eggs therein. When Spring Management 263 this new brood is well started, the manipulation may be repeated and still more eggs will be laid. This is attractive in theory but in practice is attended with danger. The bees may not attempt to cover both portions of the divided brood, resulting in loss, or, because of exceptionally cold weather, they may contract the cluster and leave the outside brood to die. The beginner should by all means leave the amount of brood to be determined by the bees, confining his work to the supplying of protection, stores and room for the expansion of the brood. If the giving of abundant protection, stores and room for the maximum advantageous expansion of the brood will cause the colony, from its own instinct, to reach its maximum strength in time for the storage of the crop, then additional manipulations in stimulative feeding and in spreading of the brood, even though they may do no harm, are non-essential. They are, therefore, to be condemned for the commercial apiary. If the favorable conditions enumerated do not bring the necessary strength of colony and if stimulative feeding will, then these manipulations are justifiable. If the period for brood-rearing previous to the beginning of the honey-flow is short, rapid brood-rearing becomes more important. This is usually the case in northern localities. It is safe to say, however, that stimulative feeding and spreading of the brood should not be practiced early in the spring but should be confined to a period of six or eight weeks just previous to the particular honey-flow for which the beekeeper is building up his colonies. If the main crop is in late summer, the beekeeper need not force his bees in the spring. In some localities, the season is made up of a series of honey-flows of about equal importance. If there are long intervals between honey-flows, the beekeeper must see that brood-rearing is at its best during a period of six or eight weeks before each flow. Substitutes for pollen. Beekeepers have repeatedly noticed that during a shortage in the supply of pollen, bees will pack meal or sawdust 264 Beekeeping on their legs, just as they do pollen, and will carry it to the hive. There are also reports of coal dust being carried in this way. After observing bees carry in rye flour from a neighboring mill, Dzierzon put some in the apiary where the bees could get it readily and they carried it in eagerly. It is still held by many beekeepers that bees should be given rye flour or pea, oat or corn meal in the spring, these being considered more suitable foods than wheat flour. These substitutes are chemically not very similar to pollen, and observations as to the effects of them on the development of the brood are badly needed. In fact, it can scarcely be said that we know that the giving of substitutes for pollen is serviceable in brood-rearing, and one cannot but wonder what Dzierzon's advice would have been if his apiary had been near a coal mine. Because of the unusual things that bees do, we are not justified in concluding that the giving of substitutes for pollen is useless, however, and no harmful results are recorded from the practice. It is a promising field for study, for there is sometimes a scarcity of pollen just when it is most needed. CHAPTER XV SWARM CONTROL AND INCREASE AT the close of the chapter on the manipulation of bees, it is pointed out that success in honey-production depends (1) on getting bees in time for the harvest and (2) on keeping them in the proper condition for storing. The first requirement is discussed in the chapter immediately preceding. However, if a colony of bees builds up rapidly to full strength, the beekeeper is confronted by the problem of preventing it from dividing its forces, thereby causing him to fail in getting the maximum crop, or even to get no surplus from it. This second problem is mainly involved in swarm control, but also includes the providing of other conditions favorable for storing. Loss from division of the working force. In the days of the box-hive, success in beekeeping was measured by the number of swarms that issued, but the beekeeper now knows that he cannot increase the number of his colonies during a honey-flow without curtailing his crop, unless the increase is made from brood that would emerge too late for the resulting bees to assist in gathering nectar. Success in manipulation is now measured by the results the beekeeper attains in preventing swarming. If swarms issue, as they will at times in spite of every known precaution, the beekeeper then aims to use the bees so as still to keep them together and thus to overcome the danger of a reduced crop. Because the experienced beekeeper so well knows that swarming endangers his crop and also that swarms may be lost, the usual statements concerning the 265 266 Beekeeping beauty of a swarm fail to meet a ready response from him. To him, swarming is the one great handicap in beekeeping. The necessity of keeping the bees together cannot be overestimated. If a colony is divided just before or during the honey-flow, the two parts fail to produce as much surplus honey in that honey-flow as the same bees would if they had remained in one colony and in normal condition. Furthermore, when bees are preparing to swarm, their condition is not so favorable for gathering. Whether there is some physiological difference or whether the lack of concentrated effort in gathering is due to an unbalanced condition of the colony population is not known, but the results of the swarming preparations are shown in a decrease in the crop. In successful honey-production, it therefore becomes essential that every effort be made to reduce and to overcome the tendency to swarm. Variation in swarming. It is interesting to note that, in any region, swarming usually occurs at a certain season or seasons and rarely occurs throughout the entire active season. It is most common in those sections of the North in which the main honey-flow occurs in early summer. If there are two heavy honey-flows, swarming may occur in connection with each one, although it is usually less troublesome in the later one. The crowded condition of the hive in the production of comb-honey is favorable for the development of the swarming tendency and, since the early summer flows of the North are the best for comb-honey, the control of swarming is most difficult in northern comb-honey apiaries. That many comb-honey producers crowd their colonies more than is necessary or desirable will be shown in a later chapter, but, even with the most skillful manipulation of the supers and with the proper manipulations throughout, there is always more crowding than is necessary in extracted-honey production. Swarm control is therefore chiefly a Swarm Control and Increase 267 problem for the northern comb-honey producer, and from these men we have obtained the best systems for controlling swarming and also the most light on its cause. In the South, where the honey-flows are longer and less intense, swarming is less frequent, and this is also true in the irrigated regions of the West. In those regions of the tropics where the honey-flows are practically continuous, there may be a kind of swarming season, but swarms are so much less frequent that the northern beekeeper would not consider the control of swarming a serious problem in such a locality. Variation in colonies in respect to swarming preparations. In any apiary and in almost any season, colonies differ greatly in their propensity to swarm. (1) There are some which show no indication of swarming. These are the very best for honey-production and the beekeeper should aim to increase their number. (2) Other colonies show a tendency to swarm by starting queen cells, but may be deterred either by cutting out the newly started queen cells or by taking away some combs of brood. (3) Still other colonies are more persistent and will swarm if the honey-flow continues unless they are subjected to some radical manipulations. (4) Some colonies whose queens fail swarm in connection with the supersedure of the old queen. "Of 160 colonies run for comb honey that were fair subjects for comparison, 13| per cent went through the season without ever offering to start queen-cells; 12f per cent started cells one or more times, but gave it up when their cells were destroyed; and 73f per cent seemed so bent on swarming that they were treated by being kept queenless 10 or 15 days. The colonies that were left with their queens all the time averaged 36| per cent more sections than those that were treated. But that's better than they would have done if left queenless for 21 days, which would be the case practically if swarms were shaken." — C. C. Miller, 1905, "Gleanings in Bee Culture," XXXIII, p. 1174. 268 Beekeeping Direction of the beekeeper's efforts. The work of the beekeeper in swarm control may be divided into two phases, for his manipulation of a colony depends on his recognition of the degree of persistence in swarming which a colony exhibits. He may try to increase the number of colonies which make no preparations to swarm and may prevent swarming in the colonies which respond to simple measures. To these manipulations may be given the name preventive measures. However, the beekeeper finds some colonies which he knows from experience cannot be kept from swarming by cutting out queen cells, by the removal of a frame or two of brood or by other simple expedients. To describe the difference which the beekeeper recognizes is somewhat difficult, but, in general, if the larvae in the queen cells are still small, preventive measures may be used. In cases of the building of queen cells obviously due to supersedure or when the working force is relatively not so strong as the brood, an artificial swarm should not be made. To the more drastic measures, used on colonies with advanced larvae in the queen cells which will persist in their preparations to swarm, the name remedial measures l is proposed. PREVENTIVE MEASURES These may be grouped under the three heads given below. Whatever the system of management, the earliest manipulations in swarm control will usually be preventive measures, for the beekeeper cannot know very far in advance which colonies will fail to respond to the less drastic manipulations and in any event these will deter swarming in the larger number of colonies. 1 In Demuth's bulletin on "Comb Honey," he uses the term "control measures," but the words "preventive" and "control" are not mutually exclusive. Swarm Control and Increase 269 Breeding. Some beekeepers make a practice of requeening colonies which swarm with young queens which are the progeny of queens whose colonies have not swarmed, in an effort to eliminate swarming by selection of non-swarming stock. In a sense, this work has failed, for after years of such selection the bees still swarm under favorable conditions, but the testimony of many practical beekeepers indicates that the percentage of colonies that swarm under proper management is reduced by selection of non-swarming stock. Since the men who are making this selection are, at the same time, improving the manipulations in swarm control and are becoming more skilled in this work, it is somewhat difficult to measure the value of this effort. Since requeening from good stock is a highly commendable practice for other reasons, it seems advisable to choose breeders from those which have not swarmed, wherever possible. If breeding queens are chosen from the colonies which show the best results in honey crops, these queens will usually be from colonies that have not swarmed during the season. Mechanical devices. Efforts have been made to devise a hive which will give to the bees an environment in which the swarming tendency will usually not be developed, a well-known example of which is the Aspinwall hive, with slatted frames between the combs. Similar slats between the frames of ordinary hives have been used. Since a non-swarming hive is needed especially in the production of comb-honey, it would appear that there should be provision for more crowding of the bees than is given in the Aspinwall hive, but it is perhaps too early to pass judgment on the efforts in this line. A deep (two inch) bottom board with a large entrance (Miller, " Fifty Years among the Bees ") leaves space under the frames in which may be placed a slatted rack during the active season. This provides abundant ventilation and room for bees and may 270 Beekeeping act as a preventive of swarming, although it is not so claimed by Doctor Miller. The use of large hives in the production of extraeted-honey, which so successfully reduces swarming, may be considered as the giving of an environment unsuitable for the development of the swarming tendency rather than the control of swarming by manipulation. Preventive manipulations. The most common methods of preventing swarming are by manipulation, probably because success, if attained, is immediately recognizable. Greater progress has been made in the devising of manipulations for this purpose than is shown in breeding or in the invention of mechanical devices. The manipulations used by the beekeeper in swarm prevei}-tion fall into the following classes: (1) the introduction of young queens (preferably from superior stock, possibly the progeny of queens whose colonies have not swarmed); (2) the prevention of crowding in the brood-chamber previous to the honey-flow, the crowding incident to comb-honey production being brought about only after supers are put on. This is often accomplished by giving an extra hive-body for early brood-rearing, so that there is abundant room for brood and stores; (3) the use of bait sections or extracting combs (Fig. 133) in the first super in comb-honey production, thus inducing the bees to begin work in the supers promptly to avoid excessive and unnecessary crowding in the brood-chamber; (4) the proper manipulation of supers in comb-honey production (p., 314) to reduce crowding as far as possible (possibly also to remove young bees from the brood-chamber) ; (5) the use of only good worker comb in the brood-chamber, to reduce the number of cells unavailable for worker eggs; (6) ventilation (by raising the hive on blocks, or by large entrances); (7) shade, to prevent overheating; (8) the removal of combs of brood which are replaced by empty combs or sheets of foundation to relieve the congestion (see also this manipulation under remedial measures); (9) the removal of queen cells soon after they Swarm Control and Increase 271 are started, since if queen cells are well advanced, their removal is not so effective in preventing swarming. This usually requires an examination of the brood-chamber once in seven to ten days. Miller's methods. To make these manipulations clear, it may be well to recapitulate by describing the system used by C. C. Miller. To provide abundant bees in time for the harvest, as well as to eliminate any tendency to early swarming, strong colonies are given an extra hive-body, during the rapid spring breeding, all the combs being built to the bottom bar of the frame so far as practical. Colonies are requeened whenever a queen shows signs of inability to keep up the full strength of colony, these queens being from mothers whose colonies have not swarmed. When the honey-flow begins, a single hive-body for each colony is filled with full combs of brood (any additional combs of brood being used in other less populous colonies, for increase or for other purposes) and each colony is given a super containing one or more bait sections, into which the bees go at once, if the honey-flow permits.1 Doctor Miller is a master in the manipulation of supers and the system used by him is described in a later chapter (p. 314). His hives have wide entrances (2 inches deep) and are protected by trees from the heat of the sun. Frequent examinations are made to remove newly started queen cells. The crops which Doctor Miller obtains are so much greater than those of other beekeepers similarly situated, or even than those in better locations, that his methods should be carefully studied. He uses the 8-frame Langstroth hive, but does not especially recommend it. It should also be added that Doctor Miller is a firm advocate 1 Doctor Miller once asked the author, in all seriousness, what beekeepers mean by their reported difficulty in getting bees to work in the supers promptly. Probably his bees are so much better prepared to gather a surplus than are those of many beekeepers that in his own apiary he has not seen for years conditions which occur yearly in the apiaries of many beekeepers. 272 Beekeeping of the improvement of stock by selection and he attributes much of his success to his efforts in this line. REMEDIAL MEASURES The preventive measures previously mentioned are usually sufficient to control swarming in a colony used in extraeted-honey production but, in the crowded conditions of the comb-honey hive, in a good season, there will probably be some colonies that will persist in their preparations to swarm. In a poor season, when the colony lacks the stimulus of nectar coming to the hive, it has not the conditions nor the number of bees necessary for swarming, but when conditions during early brood-rearing are favorable and when there is abundance of nectar during the main honey-flow, there is also usually a larger population, and preparation for swarming may be begun and often completed in most of the colonies in the apiary. It then rests with the beekeeper so to manage these colonies that, by keeping the bees together and by keeping them in working condition (p. 85), he may obtain practically as large a crop as if swarming had not occurred. He now aims not so much to prevent swarming as to satisfy the instinct and to overcome the evils incident to division of the working force. The method to be adopted depends largely on the size and location of the apiary. If the bees are all in one apiary, where they are under the immediate care of the beekeeper every day, the bees may be permitted to swarm naturally but, in comb-honey production especially, colonies in out-apiaries can be expected to produce more, without the loss of swarms, if by some remedial manipulation the swarming tendency is controlled to suit the convenience of the beekeeper. If an assistant is kept at each apiary, it becomes essentially like the home apiary, but it rarely pays to keep so much help. Control of natural swarms. Swarms which issue may be managed in several ways. (1) They may be allowed to fly into the air and cluster on Swarm Control and Increase 273 some support, after which they may be hived and placed in the desired location. When the bees have clustered they may be shaken into a box or basket and then placed in front of the hive that they are to occupy. They should be placed so that some of the bees find the entrance promptly, otherwise the bees may begin their characteristic march in the wrong direction (p. 68). If the bees cluster on a limb which can be cut, this may be removed with the adhering bees and carried to the hive and the bees shaken off. If the cluster forms on the trunk of a tree or post or in some other place from which they cannot be readily removed, a box containing a piece of comb (Fig. 47) may be placed above and preferably in contact with the cluster and the swarm will soon move into the box, where it may be handled. Care should be taken to get the queen, as otherwise the bees may again take wing and return to her. (2) If the queen's wings are clipped (!p. 260), she is unable to fly with the swarm and, after leaving the hive, she falls to the ground. The swarm generally does not cluster if the queen is not with it or, if it does cluster, it soon takes wing and returns to the old hive, provided it does not have a virgin queen with it as is sometimes the case if swarming has been unduly delayed. In the meantime, the beekeeper should find the queen on the ground and place her with the returning bees, after adjusting the hives as described later. (3) If a queen and drone trap (Fig. 30) is placed over the entrance, workers can leave, but when the queen attempts to leave she is retarded by the trap. She then, in her attempts to escape, usually goes into the upper part of the trap and is unable to return. The swarming bees then behave as they do when the queen's wings are clipped, and soon return. To allow the queen to go below with the bees it is necessary only to pull the tin slide which is shown partly drawn in the illustration. Here too the hives are adjusted as when the queen is clipped and of course the swarm is not left in the old brood-chamber. The inexperienced beekeeper should perhaps be warned not to leave a queen trap on the 274 Beekeeping entrance at all times for it will prevent virgin queens from leaving the hive to mate. Drones of course are also prevented from leaving and if they are abundant they may crowd the entrance, with disastrous results. (4) As the swarm issues, a wire-cloth cage may be placed over the hive or fitted over the entrance. The bees then cluster in the top of the cage, without causing confusion in the apiary, and may be hived when convenient. Automatic hivers. Several years ago the desirability of some automatic hiving device was much discussed and numerous efforts were made to devise apparatus which would deposit or lead the issuing swarm to a new hive. These arrangements were devised to place the swarm in a new location and beekeepers now prefer that it be returned to the old location. Location for the swarm. After a swarm has issued, the old practice was to hive it in a new location, thus dividing the working force. The beekeeper should manipulate the two parts of the original colony so as to prevent such a division. A method sometimes used is to return the swarm without the queen to the old hive and about a week later (before the developing queens emerge) the queen cells are cut and the colony is requeened later. The usual method is to remove the hive-body containing the brood while the swarm is out and to return the swarm to a new hive on the old stand. By either of these methods, the swarm is augmented by the returning field bees and, if there were supers on the colony before swarming, they are kept with the swarm and the bees promptly return to their work. The most common error of the inexperienced beekeeper in swarm management is to put the supers on the "parent colony" (the one which retains the brood). The population of the parent colony is reduced by the loss of the field bees and after-swarming is thereby made less probable. These manipulations make it Swarm Control and Increase 275 necessary that the beekeeper be present when the swarm issues, or soon after, and they are therefore not suitable for out-apiary management. Disposition of the brood after swarming. The so-called parent colony may be sufficiently populous to cast an after-swarm and should therefore be managed so as to prevent this and also so that the emerging bees shall be useful, especially if the honey-flow will probably be of long duration. The parent colony may be broken up at once by the distribution of the brood to other colonies, while the adhering bees are added to the swarm. Another method is to destroy all queen cells except one and to allow the parent colony to remain intact. If the parent colony is left to requeen itself by the emergence of the developing queens, it often casts an after-swarm, so it is safer either to remove all queen cells except one or to remove them all and give a laying queen or virgin queen. Still another method is to reduce the population of the parent colony just before the young queens emerge and to add the emerging bees to the swarm. If the parent colony is put back beside the swarm after the swarm is hived, is left there for a week and is then removed to a new location, it is so reduced when the virgin queens emerge that an after-swarm is not cast. A modification of this method to be preferred when the clipping of queens is practiced or when the queen trap is used is to set the parent colony to one side with its entrance about 90° from its former FIG. 112. —Manipulation to reduce population of parent colony — second position. Parent colony is now in hive without supers. 11. — Manipulation to reduce population of parent colony — first position. Previous A swarming. to 276 Beekeeping FIG. 113. — Manipulation to reduce population of parent colony — third position. location (Figs. Ill and 112), so that all returning field bees join the swarm. As the brood emerges, the young'bees become accustomed to the location of their hive. In a couple of days the parent colony is turned about half way around toward its former position (Fig. 113), and, after the bees again become accustomed to the change, it is moved to a position parallel to that of the new colony (Fig. 114). If within seven or eight days of the issuing of the swarm, the parent colony is removed to a new location, the young bees in flying out join the swarm, thereby considerably reducing the parent colony. When the parent colony is moved, part of the bees may be brushed in front of the entrance of the swarm, leaving some to care for the brood but not enough to induce an after-swarm. The parent colony may be used for increase or the bees as they emerge may still be added to the swarm or to some other colony. Other methods of using some young bees or sealed brood to advantage will be found by the beekeeper. They may be used to build up weak colonies or, if the honey-flow will probably be long enough to warrant it, two parent colonies may be placed side by side. By giving one a queen and removing the queen cells from the other, they may be united about two weeks after the swarm issues, when most of the brood has emerged from the queenless colony, and they are then ready for supers. What to use in the brood-chamber in hiving swarms. The use of full sheets of foundation in the brood-frames has the marked advantage of resulting in straight combs of worker FIG. 114. — Manipulation to reduce population of parent colony—fourth position. Swarm Control and Increase 277 cells. The comb is built up rapidly, in fact so rapidly as to be considered a disadvantage at times, in that brood is so quickly reared that the increase in population may again induce swarming. The use of full sheets of foundation may increase the work done in the brood-chamber, at the expense of the surplus. Narrow strips of foundation, perhaps an inch wide or less, may be used, and this usually results in slow progress in the construction of combs in the brood-chamber. The bees then do more work in the supers, if they have been started, and it is so long before the colony can rear much brood that they rarely attempt to swarm again in the season. However, combs built on strips of foundation often contain many drone cells, especially if the comb building in the brood-chamber progresses faster than the FIG. 115.—Queen excluder ("honey board"). cells are filled with eggs by the queen or when comb is built outside the space needed for brood. If the parent colony has a brood disease, the use of strips of foundation is preferable, thus combining swarm management and disease treatment. When either strips of foundation or full sheets are used and partly drawn combs are present in comb-honey supers, the queen may go above to lay eggs and this should be prevented by the use of the queen excluder (Fig, 115) for a feW days or until there is room for egg-laying below, when the excluder may be removed. If the supers are left off for a time, work will progress in the brood-chamber so that space for egg-laying will be available there and the queen will not go to the supers. One or two empty combs may be