IN THE UNITED STATES

Agriculture Handbook No. 335

Agricultural Research Service

UNITED STATES DEPARTMENT OF AGRICULTURE ySDA. National Agriourtural Library

10301 Baltimore Blvd Beitsville. MD 20705-2351 BEEKEEPING IN THE UNITED STATES

Agriculture Handbook No. 335

Agricultural Research Service

UNITED STATES DEPARTMENT OF AGRICULTURE

Washington, D.C. ^^^^^^ ^^^^^^ ^^^^

For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $1 Trade names are used in this publication solely to provide specific in- formation. Mention of a trade name does not constitute a guarantee or warranty by the U.S. Department of Agriculture and does not signify that the product is approved to the exclusion of other comparable products. Caution: If pesticides are handled or apphed improperly, or if unused portions are disposed of improperly, they may be injurious to humans, domestic animals, desirable plants, bees and other pollinating msects, fish, and wildlife, and may contaminate water supphes. Use pesticides only when needed and handle them with care. Follow the directions and heed all precautions on the container label. CONTENTS Page Introduction ^ History of beekeeping in the United States _ 2 Seasonal colony activity and individual development 5 Development of American ^ Nectar and pollen plants. ^^ Beekeeping regions in the United States _ 17 Managing colonies for high honey yields 23 Bee behavior ^^ Beehive and honey handling equipment 37 Honey processing, packing, and distribution 42 nutrition ^^ Honey, its composition and properties ^ö Queen and package bee production ^^ Breeding and genetics of bees— _ ^^ Pollination '' Brood diseases of honey bees ^" Diseases and pests of adult honey bees ^^ Pesticides ' Hymenopterous insect stings ^^^ Observation beehive ^^' Management of wild bees ^^^ Federal and State bee laws and regulations H^ Federal and State apiculture activities 1^^ Collections of bee hterature ^"^^ Nongovernment beekeeping organizations 1^1 Honey marketing aids Honey price support program Statistics on bees and honey World production and trade in honey ^^^ Glossary III

BEEKEEPING IN THE UNITED STATES INTRODUCTION

By S. E. MCGREGOR, Chief, Apiculture Research Branch, Entomology Research Division, Agricultural Research Service

Requests are frequently received by the U.S. Extreme variations in climate and flora affect Department of Agriculture for ^^all available in- bees. For example, a beekeeper in the extreme formation on honey bees/^ Such requests usually Northern States places his colonies where they indicate the writer is unaware that several thou- will obtain maximum sunlight, whereas a bee- sand books are written on bees, over a hundred keeper in the arid Southwest will supply them with bee magazines are issued regularly, and hundreds shade. Bees may be storing nectar in the fall in of technical articles on bees are published annu- the Deep South when colonies in the North are ally in every major language of the world. buried by snow. After a mild winter in the South, What is really wanted and is supphed is a bees may again be storing a crop of honey from pamphlet, ARS 33-10-4, Information About Bee orange blossoms, while those in the Far North are Culture. This pamphlet gives basic information still in their winter cluster. In general, however, about a hive of bees and where other bee informa- basic information appHes to the bee colony wher- tion and suppUes can be obtained. ever it is located, cold or hot, wet or dry. This This handbook on beekeeping is for the estab- information is included in this handbook. lished beekeeper, the extension specialist, teachers, This publication has been prepared by U.S. and those who desire to know more about bees. Department of Agriculture specialists in bee- It includes information about the colony and its keeping or bee products. Like other groups, these activity as affected by external environment or speciahsts do not agree on all details in every beekeeper manipulations, the makeup of the bee- phase of apiculture. An attempt has been made hive, bee behavior, honey production, honey plants, use of bees in pollinating crops, and bee here to give basic information without personal diseases. bias. HISTORY OF BEEKEEPING IN THE UNITED STATES

By ERIC V. NELSON, apicuUurist, Entomology Research Division, Agricultural Research Service ^

Beekeeping was unknown in the Western Hem- and inspection of bees impossible. This situation isphere until after the first European settlers continued for another century and a half. Not arrived. The natives of the West Indies used until the middle of the 19th century did a signifi- honey and waxes, which probably came from the cant in beekeeping methods occur. native stingless bees (Meliponinae). Wax may also have been obtained as a plant derivative. In 1828, Moses Quinby took up beekeeping in The Indians of North America referred to honey New York; in 1837, the Reverend Lorenzo bees as the '^he white man's fly/' and regarded Langstroth acquired two hives in Andover, Mass. their presence as indicating the coming of white These two men revolutionized the bee industry in settlers. the United States. Publication of their books in about 1852 signaled the advance to begin. The actual date of importation of the first colonies of honey bees {Apis mellijera L.) to From then to the beginning of World War I, North America is unknown. It was during early virtually every modern beekeeping method and colonization, since bees were important in the much of the modern equipment were devised It rural European economy. In 1622, bees were was truly ^The Golden Age of Beekeeping.^' m Virginia; by 1648, and honey were abundant. In 1640, the town of Newbury, Mass., Movable-Frame Hive and Bee Space established a municipal . (The expert put in charge became the town's first pauper.) In Prior to the time of Langstroth, bees were kept 1641, bee colonies in New England were sold for m a section of hollow log, a wooden box, or a 5 pounds apiece, the equivalent of 15 days of labor straw skep, in which they were allowed to build by a skilled craftsman. comb in any way that instinct indicated. Inspec- In 1763, the English introduced colonies of tion of the combs and removal of honey could bees into Florida, although they may have been only be accomplished by seriously disrupting or brought earlier to St. Augustine by the Spanish. destroying the colony. In a few instances a honey In 1773, a single colony was taken from Florida ^'super'' (a second box placed over the first and to Mobile, Ala. By the end of the 18th century, the two connçcted by a bee passageway) was bees were fairly common throughout the eastern employed. The beekeeper could remove honey half of the continent. from the super without destroying the colony, but Bees reached the West from two very diverse could not inspect or manipulate the makeup of sources. In 1809, bees were imported to Alaska the colony. from Russia by a Russian monk. In 1830, they For centuries the Greeks had used , in were carried to California from Alaska. There is which the combs were suspended from bars of no record to indicate that these colonies survived. wood placed across the top that permitted removal However, in 1856, gray-banded bees were found in of individual combs. This method apparently the wild in California. These may have been made little impact on the rest of Europe until descendants of the original Alaskan imports. In Dzierzon, a German clergyman, improved and 1853, the first importations to California from the used It in Germany during the early 19 th century. Eastern United States occurred, followed by ex- Huber, a blind Swiss naturalist, then designed a tensive importations beginning in 1856. California hive, in which frames supported the comb on all soon became one of the most important beekeeping four sides and were hinged to spread apart like areas in the country. pages of a book. Langs troth's discovery of ''bee space'' was the one feature needed to unite the The Golden Age of Beekeeping hive designs of Dzierzon and Huber into a suc- cessful hive. Previous to Langstroth's discovery, Beekeeping in the American colonies declined practical use of a four-sided frame failed because rapidlj after 1670, probably because of the disease It was designed to fit closely to the sides and top now known as . The primitive of the hive. Langstroth found that bees preferred beekeeping practices of the day made proper care a passageway of about three-eighths inch. A narrower space was sealed with propolis, which 1 In cooperation with Louisiana Agricultural Experiment made removal of combs difficult. The bees were .Station. inclined to build comb in wider spaces. BEEKEEPING IN THE UNITED STATES

the 19th century were descended from the British Development of Modern Equipment and German stocks brought over by the early Langstroth's discovery of bee space led directly colonists. They were highly susceptible to Euro- to the development of the modern hive. With the pean foulbrood, swarmed excessively, and were publication of his results, the U.S. Government difficult to handle. The renewal of beekeeping Patent Office was deluged with designs for new stimulated a worldwide search for new and more hives. Every imaginable variation of Langstroth's satisfactory strains or races of bees. Langstroth, design was attempted, many to avoid infringe- Dadant, Frank Benton, and others led the way ment on his patent. The fact that the patented in this search. feature was bee space, and not the frame itself, After many disappointments, three significantly was ignored. Although Langstroth's patent was superior races were established and accepted in clearly vaUd, he had neither financial nor physical the United States. These were the Itahan, the resources for the long court battles that followed, Caucasian, and the Carniolan races. and he soon gave up the struggle to protect his It is unlikely that any of the strains or races patent. The deluge of hive designs ended; only a brought from Europe were pure when they arrived, few sizes of hives are still in use today. and less likely that they remained pure after During this period the industrial revolution arrival in the United States. Only since the devel- affected beekeeping, resulting in machine produc- opment of techniques for controlled mating has tion of standard-sized interchangeable frame and maintenance of pure strains or races of bees become hive. Two inventions, the wax comb foundation possible. and the centrifugal honey extractor, completed The new stock of bees arrived at a time when the evolution of beekeeping from a tiny farming great strides were being made in forcing colonies sideline to its present commercial status. Wax to rear queens. This development, combined with comb foundation is simply a sheet of wax im- the discovery that queens could be shipped pressed with the shape of worker cell bases and through the mail, made the purchase of queens placed in the frame. The first foundation was by every beekeeper an easy matter. The new produced by a German, Johannes Mehring, m stock spread rapidly across the country; the 1857. A roller press to produce foundation com- ItaUan race soon became the most popular. mercially was developed in 1876 by A. I. Root. Finally the use of wire bracing in the foundation The Modern Era was developed by J. E. Hetherington. Further development of the has been a At the beginning of World War I, the ^^Golden process of refinement rather than innovation. Age of Beekeeping'' drew to a close. Interest m The idea of a centrifugal honey extractor came bees became increasingly commercialized and small from an Austrian, Major Hrushka. Prior to his farm declined. Beekeeping as a hobby was invention, honey could only be removed by cutting increasing. the comb from the frames, crushing it, and strain- The 20th century has been one of rapid tech- ing the honey from the wax. This was both time nological achievement in beekeeping as in other consuming and costly. Until the development of industries. Apiculture research faciUties have been the extractor, the biggest seasonal honey crop for established. The honey bee has become important one beekeeper was obtained by Quinby and not only as a producer of honey and wax and as a amounted to only 10 tons. Hrushka's extractor pollinator but also as a research animal in biolog- was modified by Langstroth, Dadant, Quinby, ical studies. . and many others. The first commercial extractor The package bee industry, by which bees raised was marketed in 1870 by H. O. Peabody. The ex- in the Southern States and California are sent all tractor made Kquid honey available on a commer- over the United States and to other countries, cial scale previously unknown. However, many was started just prior to World War I. It has been people refused to believe that it was pure honey, of great value to beekeepers in the Northern and several years passed before it was accepted States and Canada, where cold weather makes by the public. Once public acceptance was won, successful wintering of colonies difficult. Many extracted honey became the principal product of beekeepers in these areas kill their bees in the the American beekeeping industry. fall, then restock with a young queen and bees Another equally indispensable development of in the spring. the period included the . The Porter With controlled mating of bees now a refined bee escape and metal queen excluders were also and dependable method, at least two strains of developed. All are used by beekeepers today and hybrid bees have been made available to the bee- have changed only shghtly since their original keeper; more are certain to follow. In the future development. most of the queens used by beekeepers will prob- ably be artificially inseminated, and the modern New Strains or Races of Bees beekeeper will use different strains of bees for The bees in North America until the middle of different areas and purposes. 4 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULfTURE

References (2) OERTEL E. 1945. HISTORICAL NOTES ON HONEY BEES AND BEEKEEPING IN LOUISIANA AND OTHER (1) LANGSTROTH, L. L. SOUTHERN STATES. La. Acad. Sei. Proc. 1865. A PRACTICAL TREATISE ON THE HIVE AND /ON p^^ ^ ^^^ J?^W^~^^' HONEYBEE. 409 DD. J. B. LippinCOtt W; rELLETT, 1^. O. Co Philadelnhin i-ippincott. 1938. HISTORY OF AMERICAN BEEKEEPING. 213 ^0., l-miadelphia. Collegiate Press, Ames, Iowa. SEASONAL COLONY ACTIVITY AND INDIVIDUAL BEE DEVELOPMENT

By NORBERT M. KAUFFELD, apiculturist. Entomology Research Division, Agricultural Research Service ^

Seasonal Colony Activity characteristics respond in various ways that affect the activity of the bee colony. Basically a colony of honey bees comprises a Reduced incoming food causes reduced brood cluster of several thousand workers (sexually im- rearing and diminishing population. The pro- mature females), a queen (a sexually developed portion of old bees in the colony will begin to female), and, depending on the colony population, increase, since young bees emerging in the fall an undeterminate number of drones (sexually de- live longer. Many of the fall bees survive the veloped males) (fig. 1). A colony normally has winter. Propolis collected from the buds of trees only one queen, whose sole function is egg laying is used to seal all cracks and reduce the entrance and not colony government. The bees cluster to keep out cold air. loosely over several wax combs, the cells of which When nectar supplies in the field become scarce, are used for the storage of honey (carbohydrate the workers drag the drones out of the nest and food) and pollen (protein food) and for the rearing do not let them return, causing death by star- of young bees to replace old adults. vation. This elimination of drones reduces the The activities of a colony vary with the seasons. consumption of winter honey stores. If the queen The period from September to December might is old, she may be killed by the workers during be considered the beginning of a new year for a late summer and replaced by a young mated one colony of honey bees. The condition and activity before the drones disappear. of the colony at this time of year to a great degree When the temperature drops into the 50's, the affect its prosperity for the next year. In the fall bees begin to form a tight cluster. Within this the night temperatures decrease and the days cluster the brood (consisting of eggs, larvae, and shorten. Plant growth, nectar secretion, and flower pupae) is kept warm, with heat generated by the 1 In cooperation with Wisconsin Agricultural Experi- activity of the bees feeding on the stored honey. ment Station. The egg laying of the tapers off

FIGURE 1.^—Worker, queen, and bees. 6 AGRICULTURE HANDBOOK 335^ U.S. DEPT. OF AGRICUI/TURE

and may stop completely during October or drones, the bees also prepare to rear a new queen. November, even if pollen is stored in the combs. A few larvae that would normally develop into During the winter the colony is put to its severest worker bees are fed a special food called royal test of endurance. jelly, their cells are altered to handle the larger As temperatures drop, the bees draw closer queen, and her rate of development is speeded up. together to conserve heat. The outer layer of The number of queen cells produced will vary with bees is tightly compressed, insulating the bees races and strains of bees as well as individual within the cluster. As temperatures rise and fall, colonies. the cluster expands and contracts. The bees Regardless of its crowded condition, the colony within the cluster have access to the food stores. will try to expand by building new combs if food During warm periods, the cluster shifts its position and room are available. These new combs are to cover new areas of comb containing food. An generally used for the storage of honey, whereas extremely prolonged cold spell can prohibit the older combs are used for pollen storage and cluster movement, and the bees may starve to brood rearing. death only inches away from food. When the first virgin queen is about ready to The queen stays within the cluster and moves emerge and prior to the main nectar flow, the with it as it shifts position. About mid-January colony will swarm during the warmer hours of to early February if the colony is well supplied the day. The old queen and about half of the bees with honey and pollen, it will begin to stimula- will rush en masse out the entrance hole. After tively feed the queen and she begins egg laying. flying around in the air for several minutes, they This new brood aids in replenishing the number of will cluster on a limb of a tree or similar object. bees that have died during the winter. The extent This cluster usually remains an hour or so, de- of this early brood rearing is determined by pollen pending on the time taken to find a new home by stores gathered during the previous fall. A lack scouting bees. When it is found, the cluster of pollen delays brood rearing until more is col- breaks up and flies to it. On reaching the new lected from spring flowers, and the colony emerges location, combs are quickly constructed, brood from winter in a weakened condition. rearing starts, and nectar and pollen are gathered. The colony population during the winter generally occurs in the Central States decreases because few young bees emerge to during May or June, although it can occur at replace those dying. Colonies with plenty of almost any time from April to October. young bees during the fall and an ample supply of After the swarm departs, the remaining bees in pollen and honey for winter usually have a the parent colony continue their field work of strong population in the spring. collecting nectar, pollen, propolis, and water. During early spring the lengthening days and They also care for the eggs, larvae, and food, new sources of pollen and nectar stimulate brood guard the entrance, and build combs. Emerging rearing. The bees also gather water to liquefy drones are nurtured so that there will be a male thick or granulated honey in the preparation population for mating the virgin queen. When of brood food. Karely will there be any drones she emerges from her cell, she eats honey, grooms within the colony at this time of the year. herself for a short time, and then proceeds to Later in the spring the population of the colony look for rival queens within the colony. Mortal expands rapidly and the proportion of young combat eliminates all queens except one. When bees increases. As the population increases, the the survivor is about a week old, she flies out field-worker force also increases. It may collect to mate with one or more drones in the air. The nectar and pollen in greater amounts than is drones die after mating, but the mated queen needed to maintain brood rearing, and surpluses returns to the nest as the new queen mother. of honey or pollen may accumulate. This, of Nurse bees care for her, whereas prior to mating course, will depend upon the availabihty of plants she was ignored. Within 3 or 4 days the mated that produce pollen and nectar. queen begins egg laying. As the days lengthen and the temperature During hot summer days the colony temper- continues to increase, the cluster expands further ature must be held down to 93° F. The bees do and drones are produced. With increase in brood this by gathering water and causing it to evapo- rearing and the accompanying increase in adult rate within the cluster by its exposure to air bees, the nest area of the colony becomes crowded. circulation. More bees will be evident at the entrance of the During the early summer the colony reaches nest. A telltale sign of overcrowding is to see the its peak population necessary for the collection bees crawl out and hang in a cluster around the of the greatest amount of nectar and the storage entrance on a warm afternoon. of honey for the coming winter. After reproduction, Combined with crowded conditions, the queen all colony activity is geared toward winter also increases drone egg laying in preparing for survival. the natural division of the colony population by Summer is the most favorable time for the swarming. In addition to rearing workers and storage of food supplies. More food is available BEEKEEPING IN THE UNITED STATES and warm weather permits free flight. The day- About the 12th day of its life the wax glands, hght period is the longest then, permitting located between the fourth and seventh segments maximum foraging, although rain or drought may on the undersurface of the bee's abdomen, de- reduce flight and the supply of nectar and pollen velop. Through some still unknown body process available in flowers. It is during the summer that honey is eaten and converted into beeswax, stores are accumulated for winter. At times enough which is secreted as small scales between these is stored that man can remove a portion and still segments. The fiakes of wax are kneaded by leave ample for colony survival. mandibular action into cell walls in the construc- tion of combs. When the bee is about a week old, it takes short orientation flights during the Individual Bee Development afternoon to memorize its home location. Various The queen lays small, elongated, white eggs, activities such as the conversion of nectar into one attached to the base of each cell. The egg re- honey, packing pollen pellets in the cells, hand- mains in an upright position for 3 days, then a ling water brought in for air-conditioning, fanning larva hatches. for ventilation, and guarding the entrance are The larva, a small white grub, is mass fed royal performed until about the 21st day, after which jelly by nurse bees for the first 2 days to such an it begins foraging for food. extent that it literally floats in this food. For the Adult honey bee workers live about 6 weeks next 4 days the worker larva is fed a less nutritious during the peak activity periods of late spring, food at regular intervals (progressive feeding) summer, and early faU. Most of them die in the by a different group of nurse bees. During these field. Those that die in the nest are carried out feeding periods it grows rapidly to several times some distance from the entrance and dropped to its size at hatching, and soon occupies the greater the ground. part of its cell. Its cell is capped on the ninth day The development period of drones from egg after hatching. No feeding occurs after the cell stage to adult is 24 days compared with 21 days is capped. for workers and 16 for queens. They are larger The next 12 days are spent in the capped cell than workers, do not have stings, and are gen- in prepupal and pupal stages. Following the erally fed by the workers. Their activity consists fifth or last larval molt, which occurs 2 or 3 merely in eating honey and making flights from days after the ceU is capped, the distinct adult the nest for possible mating with a virgin queen body parts appear, such as legs, antennae, wings, bee. They live about 8 weeks during the mouth parts, head, thorax, abdomen, and eyes. season but are killed off in the fall at any age. Initially the pupa is white and extremely soft The queen bee, which is longer than the worker textured, but as it grows, to emerge from the or drone (fig. 1), has the shortest period of develop- cell as an adult, it changes from white to a darker ment from egg stage to adult. Her diet from the gray and finally to its adult color. Hair develops larval stage throughout her entire life seemingly is on the various body areas, and on the day before , although she can eat honey herself. The emergence it completes its sixth and final molt. nurse bees groom and care for her bodily needs On the 21st day the young cuts its from feeding to removal of feces. Her sole func- way through the capping and crawls out of the tion within the colony is to lay eggs for the pro- cell as a physically adult honey bee. duction of the necessary bee population needed to The activity of a honey bee within the colony store sufficient food supplies for survival. Her life varies according to its age and the development expectancy is about a year, but some queens have of its internal glands. During the first 3 days been reported to live up to 6 years. She gives off a following emergence, it aids in cleaning the nest substance that has a stabilizing effect on the area. From the fourth to seventh day after emer- colony. gence, it feeds older larvae with mixtures of honey and pollen. About the seventh day after emer- The coloration of honey bees varies with the gence its pharyngeal glands become developed and races and strains. Usually the more distinctive it produces royal jelly. As such a nurse bee, it patterns separate one species from another. mass feeds the young larvae or queen until it is 10 For additional information about colony ac- to 13 days old. tivity and bee development, see pages 23-30. DEVELOPMENT OF AMERICAN BEEHIVE

By SPENCER M. RIEDEL, JB., ayiculturist, Entomology Research Division, Agricultural Research Service '

The modern American beehive (fig. 1), com- The shallow super is frequently used in produc- monly called the , permits devel- tion of liquid or extracted honey, when opment of a strong colony and production of a is to be cut from the frame, or when special comb large honey crop. With it the beekeeper has con- sections are to be filled with honey and then trol over the bees. It is simple in design, mobile, individually removed. light, durable, and economical. Its components Several variations of the Langstroth hive are are interchangeable with those of other hives. used to a limited extent. They include the 8-frame The dimensions for a Langstroth 10-frame hive Langstroth, the modified Dadant that holds 11 are given in figure 2. Basically the queen and her frames, the 12-frame Langstroth, and a square brood are confined in the brood chamber. The hive that holds 13 frames but is only 6K inches worker bees can pass through the deep. The most popular is the full-depth 10-frame and store honey in the super. Since one shallow Langstroth for both brood nest and is not enough space for a populous colony, storage. beekeepers often use several supers. The brood chambers and supers are used interchangeably. Early-American beekeepers kept colonies of bees If the combs contain brood, the section is referred in hollow tree trunks, which for convenience and safety were gathered together in an apiary, or a to as a brood chamber; if no brood is present, it is called a super. "bee yard." Since many of these logs came from gum trees, the log hive became a "bee gum." This term is rapidly disappearing, but the terms "hive" and "colony" remain synonymous. In 1921, Dadant & Sons perfected a method of inserting vertical wires into the foundation. This extra support of the comb was beneficial when beekeepers began using high-speed extractors. An aluminum comb was developed but was unaccept- able to the bees. Both aluminum and plastic foundations have also been developed. They are embossed with the cells and coated on each side with beeswax. Since aluminum is a good conductor of heat, it is not satisfactory in brood rearing areas of the hive. Neither of these permanent base mate- rials is readily accepted by the bees unless properly installed in the frames and supplied to the bees during a heavy honey flow. The best foundation is still beeswax held firmly in the frame with embedded wires. The best type of hive under most conditions is the Langstroth 10-frame hive.

References

(1) GROUT, R. A., ed. 1963. THE HIVE AND THE HONEY BEE. 556 pp. Dadant & Sons, Hamilton, 111. FIGURE 1.—Modern beehive cut away to show interior (2) PELLETT, F. C. and placement of movable frames: Bottom, full-depth 1938. HISTORY OP AMERICAN BEEKEEPING. 213 hive body; middle and top, shallow hive bodies. pp. Collegiate Press, Ames, Iowa. (3) ROOT, A. I. ■ In cooperation with Louisiana Agricultural Experiment 1962. ABC AND XYZ OF BEE CULTURE. 712 pp. Station. A. I. Root Co., Medina, Ohio. BEEKEEPING IN THE UNITED STATES

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ssssssssssssss:

^4 Space

A. Corner of lO-frome hive body, showing construction and position of frames Reversible bottom_board B. Port of end of hive body, shov»(ingi rabbet, which should be mode of tin or galvanized

CROSS SECTION OF HIVE BODY AND FRAME -— 1 VA Outside cover

ai" i Inside cover 1 !-=■ ■ =^ 1 ''» VA I Shallow super 1 CROSS SECTION OF SHALLOW SUPER

Queen excluder

Wire Brood chamber

Reversible bottom e board SIDE,END,AND TOP ELEVATION OF FRAME BN-30052

FIGURE 2.—Plans and dimensions for Langstroth lO-frame beehive. NECTAR AND POLLEN PLANTS

By E. OERTEL, apiculturist, Entomology Research Division, Agricultural Research Service ^

In order to manage bee colonies for successful acres in 1949 to about 32 million in 1964. In some honey production, a beekeeper must have avail- States certain soybean varieties are reported to able data on the nectar and pollen plants in the be valuable sources of nectar. However, because of vicinity of his apiary. Such information enables contradictory reports, the beekeeper should test him to calculate when he should install package an area for several years to determine whether bees, divide colonies, put on supers, use swarm- the soybean produces nectar in amounts adequate control measures, remove honey, requeen, prepare for his purpose. colonies for winter, and locate profitable apiary Introducing a new crop into a locality can bene- sites. fit beekeepers if the plant provides nectar or pollen. VanselP listed 150 species of nectar and pollen Land in the Soil Bank reserve, 28 million acres in plants in California, but only six are principal 1957, may benefit beekeepers, depending on what sources for commercial honey production. He ^ plant species are growing on the land. Beekeepers listed about 90 species of nectar and pollen plants should survey areas taken out of cultivation, be- in Utah, but noted that the main sources of com- cause they may be suitable for apiary locations. mercial honey are alfalfa and sweetclover. Wilson Other changes in agricultural practices include et al.* observed honey bees visiting the blossoms the use of herbicides and power mowing machines. or extrafloral nectaries of 110 species of plants in They reduce or eliminate plants that are impor- Colorado, of which the most important honey tant sources of nectar or pollen. Farmers are sources are alfalfa, yellow sweetclover, and depending less on legumes to add nitrogen to the dandelion. sou and are using more liquid ammonia fertilizer. Beekeepers are advised to record the blossoming Pests, such as insects or nematodes, may cause period for the nectar and pollen plants in their so much damage to some plant species that farmers vicinity. Most State agricultural extension ser- change to other crops. In Ohio in 1966 the alfalfa vices have bulletins available on beekeeping. These weevil (Hypera postica (Gyllenhal)) became so usually contain a list of the important nectar and destructive that there was serious concern farmers pollen plants. Unknown plants can be sent to the would stop growing alfalfa, an important nectar botany department of the State university or source. The sweetclover weevil (Sitona cylin- agricultural college for identification. dricollis Fahraeus) has destroyed much of the sweetclover that formerly was grown in the Mid- Beekeeping Locations west. The acreage planted for seed has decreased over 50 percent since 1950. Plant breeders have Beekeepers, especially commercial operators, introduced a nectarless cotton so that destructive have learned that the nectar- and pollen-producing insects will not be attracted to the plant. plants may change considerably over the years. There is no information concerning the effects Variations may be caused by droughts, changes in of air pollution caused by factories, motor vehicles, agricultural crops and practices, irrigation proj- radioactivity, and major metropolitan areas on ects, and subdivision development. Changes have nectar and pollen plants except in limited areas. been particularly rapid since World War II and Productive locations for the commercial bee- are likely to continue. keeper will become more difficult to find. However, Acreages planted to buckwheat, alsike clover, if a housing development or new subdivision forces and cotton have decreased, whereas those with a beekeeper to move his colonies from his home, alfalfa hay, mustard, saflîower, and soybeans have he may have difficulty finding a desirable location increased. Soybean crops increased from 12 million nearby.

1 In cooperation with Louisiana Agricultural Experiment Station, Poisonous Honey Plants 2 VANSELL, G. H. NECTAR AND POLLEN PLANTS OF CALIFORNIA. Câlif. Agr. Expt. Sta. Bui. 517, 55 pp. 1931. Fortunately the American beekeeper seldom ^ VANSELL, G. H. POLLEN AND NECTAR PLANTS OF needs to be concerned about plants that are poi- UTAH. Utah Agr. Expt. Sta. Cir. 124, 28 pp. 1949. 4 WILSON, W. T., MOFFETT, J. O., and HARRINGTON, sonous to honey bees. Locations with abundant H. D. NECTAR AND POLLEN PLANTS OF COLORADO. Colo. growth of California buckeye {Aesculus spp.), Agr. Expt. Sta. Bui. 503-S, 72 pp. 1958. deathcamas {Zigadenus venenosus Wats.), loco- 10 BEEKEEPING IN THE UNITED STATES 11 weed (Astragalus or Oxytropis spp.), laurel (Kalmia produce nectar from tiny specialized areas in the sp.), or rhododendron (Rhododendron spp.) should leaves or stems called extrafloral nectaries. be avoided, if possible, while these plants are in bloom. Damage to colonies from poisonous Honeydew nectar or pollen may be severe in some years, but of small consequence in others. Honeydew is the sweet liquid secreted by certain insects, such as aphids or plant lice, scale insects, gall insects, and leafhoppers, and Nectar Secretion also by the leaves of certain plants. Honeydew honey differs chiefly from floral honey in its Beginners in beekeeping frequently ask: ''Are higher dextrin and mineral content. The quality there any plants that I can grow that will increase of honeydew honey varies greatly. Some are my yield of honey?'' In general, it is not econom- fairly palatable, whereas others are undesirable ically practical to grow a crop for the bees alone. for human food or for wintering bees in northern Beekeepers are dependent on cultivated crops areas. grown for other purposes or on plants growing wild. Certain nectar and pollen plants, such as Pollen Plants alfalfa, the clovers, and sweetclover, are grown widely for agricultural purposes and they are PoUen is an essential food used in the rearing of wild to some extent. These plants, together with the honey bee larvae. It supplies the necessary less important ones, such as citrus (orange, grape- protein. The adult honey bee can Uve on carbo- fruit, lemon, limes, tangelos), cotton, sage, and hydrates alone, but the developing larvae must túpelo, furnish the greater part of the Nation's have some protein. A good, strong colony of bees commercial honey. may collect and use 50 to 100 pounds of pollen Sometimes friendly farmers will seed sniall during the season. Lack of pollen slows colony areas near an apiary with nectar-producing development in many localities in the spring and species if the beekeeper provides the seed and in some locations in the summer and fall. Pollen thus honey production increases. A few orna- may be available in the field, but cold or rainy mental flowers or trees on a city lot are of small weather may prevent the bees from gathering value to an apiary or a colony of bees. From one it. Some beekeepers feed pollen supplements, to several acres of abundant flowers are usually alone or mixed with bee-gathered pollen, to their necessary to provide sufficient nectar for one colonies. Pollen supplements are sold by bee- colony.^ supply dealers. Plants valuable only for the pollen that honey Nectar secretion or production is affected by bees obtain from them include corn, oak, and such environmental factors as soil type, soil ragweed. Examples of plants that produce nectar condition, altitude, latitude, length of day, light and pollen at a time when brood rearing is im- conditions, and weather. Such soil conditions as portant are dandelion, maple, elm, willow, wild fertility, moisture, and acidity may affect not cherry, wild plum, boneset, goldenrod, and aster. only the growth of the plant but also the secre- The clovers produce large amounts of both nectar tion of nectar. Luxuriant plant growth does not and pollen. necessarily imply that maximum nectar secretion For information on the honey bee as a pollinat- will take place. In some instances limited growth ing agent and the nutritional value of pollens, results in increased nectar production. Clear, see pages 77 and 52, respectively. warm, windless days are likely to favor nectar secretion. Most of our information on nectar production is based on casual observation rather Nectar and Pollen Plant Regions than on experimentation. Nectar is secreted by an area of special cells In table 1 the nectar and pollen plants are in the flowers called a nectary. Certain species, Hsted by region (see fig. 1, p. 17). Some of the such as vetch, cotton, partridgepea, and cowpeas. species are limited to a small area within a region; for example, thyme in New York, fireweed in the northern part of the north-central region and the ^OERTEL, E. NECTAR YIELDS OF VARIOUS PLANT SPECIES West, gallberry in the Southeast, and citrus in AT BATON ROUGE IN 1955. 10th Internatl. Cong. Ent. Proc. 4, pp. 1027-1029. 1958. the Southeast, Southwest, and West. 12 AGRICULTURE, HANDBOOK 335, U.íS. DEPT. OF AGRICULTURE

TABLE 1.—Nectar and pollen plants by regions

North- North- South- Plains Moun- South- Plant east central east 1 region tainous west West» Alaska ^ Hawaii 5 region region 2

Alder {Alnus spp.) _ X X X Alfalfa {Medicago sativa L.) X X X X X X X Algaroba {Prosopis chilensis (Mol.) Stuntz) X Alkaliweed (Hemizonia spp.) X Almond (Prunus amygdalus Batsch.) X Amsinckia (Amsinckia spp.) X Ash (Fraxinus spp.) X X Aster (Aster spp.) X X X X X X Baccharis (Baccharis spp.) X Balsam root (Balsamorrhiza spp.) X Basswood (Tilia americana L.) X X X X X Bermudagrass (Cynodon dactylon (L.) Pers.) X X Bindweed (Convolvulus spp.) X X Birdsfoot trefoil (Lotus corniculatus L.) X X Bitterweed (Helenium amarum (Raf.) Rock) X Blackberry (Ruhus spp.) X X X X Black wattle (Acacia spp.) X Bladderpod (Lesquerella gordonii (Gray) Wats.) X Blueberry (Vaccinium spp.) X X X Bluecurls (Trichostema spp.) X Blue thistle (Echium vulgäre L.) X X Bluevine (Gonolohus laevis Michx.) X Blueweed (Cichorium intyhus L.)_ X Boneset (Ewpatorium spp.) X X x X Boxelder (Acer spp.) X Broomweed (Gutierrezia texana (DC.) T.&G.) X X X Buckbrush (Symphoricarpos spp.)_ X X X Buckeye (Aesculus californica (Spach) Nutt) X Buckwheat (Fagopyrum esculentum Moench) X X Burroweed (Haplopappus tenuisectus (Greene) Blake ex Benson) X X Cacti (Cactaceae family) X X Camphorweed (Heterotheca suhaxillaris (Lam.) Britt. & Lusby) X X Cascara (Rhamnus vurshiana DC.) :_ X Catclaw (Acacia greggii Gray) X Ceanothus (Ceanothus spp.) X X Cedar elm (September elm) (Ulmus serótina Sarg.) __ X Citrus (Citrus spp.) X X X X Cleome (Cleome serrulnta Pursh) X X X Clethra (Clethra alnifolia L.) __ X Clover: Alsike (Trifolium hyhridum L.) __ _ X X X X X Crimson (Trifolium incarnatum L.) X Red (Trifolium pratense L.) X X X X X X X Sweetclover (Melilotus spp.) X X X X X X X X See footnotes at end of table. BEEKEEPING IN THE UNITED STATES 13

TABLE 1.—Nectar and 'pollen plants by regions—Continued

North- North- South- Plains Moun- South- Plant east central east 1 region tainous west Wests Alaska * Hawaii ^ region region 2

Clover—Continued White {TrifoUum repens L.) X X X X X X X X Coffee {Coffea arabica L.) X Corn (Zea mays L.) X X X X X X Cotton (Gossypium spp.) X X X X Cottonwood (Populus spp.) _ X x X X Cowpea (Vigna sinensis (Torner) Savi) Cranberry {Vaccinium macrocarpon Ait.) _ _ _ X X X Creosote bush (Larrea tridentata (DC.) CoviWe). __ _ X X Crownbeard ( Verbesina spp.) X X X X Cucurbits : Cantaloup (Cucumis melo L.) __ _ X X X X X X Cucumber (Cucumis spp.) X X X X x X X Gourds (Cucúrbita spp.). X X X Molon (Citrullus spp.) ^ X X X X X Pumpkin (Cucúrbita spp.) X X X X X Squash (Cucúrbita spp.)_ X X X X Dande ion (Taraxacum spp.)... X X X X x X Ear drop vine (Brunnichia cirrhosa Gaertn.) X Elm ( Ulmus spp.) X X X X X X Eucalyptus (Eucalyptus spp.)__ X X X Filaree (Erodium spp.) X X Fireweed (Epilobium angustifol- ium L.) _ _ _ X X Fruit bloom: Apple (MaZt^s spp.)_ X X X X X X X Apricot (Prunus spp.) X "x X Cherry (Prunus spp.)_ X X X X Citrus (Citrus spp.) X "x X X Peach (Prunus spp.) X X X X X X X Pear (Pvrus SDD.) _ _ X X X X X X Plum (Prunus spp.) X X X X X X Gallberry (Ilex glabra (L.) Gray) X Goldenrod (Solidago spp.) X X X X X X X Grape (Vitis spp.) _ X Greasewood (Sarcobatus vermicul- atus (Hook.) Torr.) X Guajillo (Acacia berlandieri Be nth.) X Guava (Psidium guajava L.)_ X G um weed (Grindelia spp.) X X X Hemp (Cannabis sativa L.) X Holly (Ilex opaca Ait.)__ X Horsemint (Monarda spp.) X X Huckleberry (Gaylussacia spp.) X Hue (Lagenaria siceraria (Mol.) Standley) X Ilima (Sida spp.) X Johnsongrass (Sorghum halepense (L.) Pers.)__ X X Kly (Acacia spp.) _ _ _ X Knopweed (Centaurea repens L.) X Koa haole (Acacia spp.)__ X Lantana (Lantana spp.) X Lima beans (Phaseotus limensis Macf.) X X Locoweed (Oxytropis or Astragalus spp.) X X Footnotes at end of table. 261-907 0--67—2 14 AGRICULTURE HANDBOOK 335, U./S. DEPT. OF AGRICULTURE

TABLE 1.—Nectar and pollen plants by regions—Continued

North- North- South- Plains Moun- South- Plant east central east 1 region tainous west Wests Alaska * Hawaii ^ region region 2

Locust: Black {Robinia pseudo-acacia L.) X X X X X X Thorny (Gleditsia triacanthos L.) X Water (Gleditsia aquatica Marsh) X Loosestrife (Lythrum spp.) X Lupine (Lupinus spp.) X X Macadamia {Macadamia spp.) X Mamane (Sophora spp.) X Mangrove: Black (Avicennia nitida Jacq.) X Red (Rhizophora mangle L.)_ X White (Laguncularia racemosa (L.) Gaertn. F.) X Manzanita (Arctostaphylos spp.)_ X X X Maple (Acer spp.) _ _ X X X X X X Matchweed (Gutierrezia saraihrae (Pursh) Britt. & Rusby) X X Mesquite (Prosopsis juliflora (SW.) X X Mexican clover (Richardia scabra L.) . X Milkvetch (Astragalus spp.) X Milkweed (Asclepias spp.) X X X X Mint (Mentha spp.) X Monkeypod (Samanea spp.) X Mountain apple (Eugenia malac- censis L.) V Mule ear (Wyethia spp.) X Mustard (Brassica spp.) X X X X X Nohu (Tribulus cistoides L.) X Oak (Quercus spp.) X X X X Ohia lenhua (Metrosideros spp.) _ _ X Oi (Fer6ena spp.) X Oregon grape (Berberis nervosa Pursh) X Oregon maple (Acer macrophyl- lum Pursh) '_ X Paintbrush (Castilleja spp.) X Palmetto (Sabal spp.) X Palmetto, saw (Serenoa repens (Bartr.) Small) X Palm trees (Palmaceae family) X Partridgepea (Chamaecrista spp.)_ X Peppervine (Ampélopsis arbórea (L.) Koehne) X X Persimmon (Diospyros virginiana L.) X X Pili (Heteropogon contortus (L.) Beauv. ex Roem. & Schutt.) __ X Pine (Pinus spp.) X Pluchea (Pluchea spp.) X Poplar (Populus spp.) X Rabbitbrush (Chrysothamnus spp.) X X Ragweed (Ambrosia spp.) _ X X X X Rape (Brassica napus L.) X Raspberry (Rubus spp.) X X X X Rattanvine (Berchemia scandens (Hill) K. Koch) X X Footnotes at end of table. BEEKEEPING IN THE UNITED STATES 15

TABLE 1.—Nectar and pollen plants by regions—Continued

North- North- South- Plains Moun- South- Plant east central east ^ region tainous west West 3 Alaska ^ Hawaii 5 region region ^

TÎpHhiirl (dprrist conadenfiifi L ) X X TÎPQinix/ppH (Crvi71 dplin ^r>r> ^ X X T?iissian-thistlp (SCLISOICL SDD.) X ~X X X Safflower (Carthamus tinctorius X X X Sfîffp (SnJvin, SDD ") X Saguaro (Carnegiea gigantea fi^nrrt^lrrt ^ Thrift Ár Tíosíp'^ X X X X Santa maria (Parthenium X X Silver oak (Grevillea robusta A. X Smartweed (Polygonum spp.) X X X X X X "x X Snow vine {Mikania scandens (T. ^ Wiiifi ^ X Sorsrhuni (Sorohutn SDD.^ X X X Sourwood {Oxydendrum X X Soybeans (Glycine max (L.) IVTprr ^ X X Spanish-needles (Bidens X X X X X X X Summer farewell (Petalostemun «¡■HT» ^ X SnnfloTtrpr (TTpJi/rnthiift sirn ^ X X X Tamarix (Tamarix aphylla (L.) Karst.) (Tamarix articúlala Vahl) X X TíirwppH ( T-TpTYii í'nn'ín SUD ) X Thistle (Sonchus arvensis L. and X Canadian (Cirsium arvense (L ) ScoD ) X Stíir (f^ptif/TjjTPn sirn ^ X Thvme ( Thi/mua SD ) X Tievine (Convolvulus or Ipomoea bpp.; X X Titi: Black (Cliftonia monophylla Í^Tiíim "i Tirittnn PY SA re* ^ X Spring (Cyrilla racemiflora X Snmmpr ( dirilln sirn ^ X Toyon (Photinia arhutifolia Lindl ) X Tulip poplar (Liriodendron X X X TiTnpln ( J^ii'i'in «iirn ^ X X Vetch (Vicia SDD.) X X X X X Vine maple (Acer circinatum Pursh) X WilH nlfnlffl (T.nfijfi snn ") X Wild buckwheat (Eriogonum snn ) X X X X Wild dandelion (Hymenopappus arenosus Heller) X Footnotes at end of table. 16 AGRICULTURE HANDBOOK 335, U.;S. DEPT. OF AGRICULTURE

TABLE 1.—Nectar and pollen plants by regions—Continued

North- North- South- Plains Moun- South- Plant east central east 1 region tainous west West 3 Alaska * Hawaii 5 region region 2

Wild snowberry (Symphoricarpos spp.) ___ X Willow (SaZzx spp.) ___ X X X X X Wingstem {Actinomeris alternifolia (L.) DC.) X Yellow ginger {Hedychium flavescens Carey) X Yellow-rocket (Barbarea vulgaris R. Br.) X X

1 Morton, J. F. Honeybee Plants of South Florida. ^ Washburn, R. H. Beekeeping in the Land of the Mid- Fla. State Hort. Soc. Proc. 77, pp. 415-436. 1964. night Sun. Gleanings Bee Cult. 89: 720-723, 756. 1961. 2 Wilson, W. T., Moffett, J. O., and Harrington, H. D. ^ Botanical names taken from Neal, M. C, Gardens of Nectar and Pollen Plants of Colorado. Colo. Agr. Expt. Hawaii, (Honolulu) Bishop Mus. Spec. Pub. 50, 924 pp., Sta. Bui. 503-S, 72 pp. 1958. Vansell, G. H. Pollen 1965. Nectar and pollen plant names taken from Dyce, and Nectar Plants of Utah. Utah Agr. Expt. Sta. Cir. E. J., Beekeeping in the 50th State, Gleanings Bee Cult. 124, 28 pp. 1949. 87: 647-651, 1959, and Eckert, J. E., and Bess, H. A., 3 Vansell, G. H. Nectar and Pollen Plants of California. Fundamentals of Beekeeping in Hawaii, Hawaii Univ. Bui. Calif. Agr. Expt. Sta. Bui. 517, 55 pp. 1931. 35, 32 pp., 1952. References (5) JEPSON, W. L. 1957. MANUAL OF THE FLOWERING PLANTS OF CALIFORNIA. 128 pp. California University (1) ABRAMS, L., and FERRES, R. S. Press, Berkeley. 1960. ILLUSTRATED FLORA OF THE PACIFIC STATES, (6) LovELL, H. B. WASHINGTON, OREGON, AND CALIFORNIA. 1956. HONEY PLANTS MANUAL. 64 pp. A. I. 4 V. Stanford University Press, Stanford, Root Co., Medina, Ohio. Calif. (7) NEAL, M. C. (2) BAILEY, L. H. 1965. IN GARDENS OF HAWAII. (Honolulu) 1949. MANUAL OF CULTIVATED PLANTS. REV., Bishop Mus. Spec. Pub. 50, 924 pp. 1116 pp. Macmillan Co., New York. (8) SMALL, J. K. (3) BLAKE, S. F. 1933. MANUAL OF THE SOUTHEASTERN FLORA. 1954. GUIDE TO POPULAR FLORAS OF THE UNITED 1554 pp. Science Press Printing Co., STATES AND ALASKA. U.S. Dept. Agr. Lancaster, Pa. Bibliog. Bui. 23, 56 pp. (9) VINES, R. A. (4) FERNALD, M. L. 1960. TREES, SHRUBS AND WOODY VINES OF THE 1950. GRAY'S MANUAL OF BOTANY. Ed. 8, 1632 SOUTHWEST. 1104 pp. Texas University pp. American Book Co., New York. Press, Austin. BEEKEEPING REGIONS IN THE UNITED STATES

By W. P. NYE, apiculturist, Entomology Research Division, Agricultural Research Service ^ Based on flora, beekeeping methods, and land in the North and mountainous areas must be topography the continental United States can be protected from the cold (fig. 3), in certain forested divided into seven geographical regions (fig. 1). areas from bears, and on the desert from drifting Each of these regions is discussed here from the sand. standpoint of honey production and methods of Some locations must be paid for by the bee- beekeeping operations. keeper, others are furnished free. Where the bees The flora, climate, and nature of the terrain are desired for pollination, the beekeeper is usually determine the system of management practiced paid for their services. by the beekeeper. For example, in the Apalach- Most beekeepers move their colonies at night icola swamps of the Southeast, hives are placed (if moving is necessary) when the bees are inside on scaffolding for protection from flood waters. the hive. But when daytime temperatures exceed In the Southwest, shade must be provided to 110° F. in the Southwest, bees are more easily protect the hives from the hot sun (fig. 2). Colonies moved at midday when they are inside the hive 1 In cooperation with Utah Agricultural Experiment rather than at night when they tend to cluster Station. on the entrance.

U. S. DEPARTMENT OF AGRICULTURE BUREAU OF AGRICULTURAL ECONOMICS

BN-30051 FIGURE 1.—Beekeeping regions of the United States. 17 18 AGRICULTUEE HANDBOOK 335, UJS. DEFT. OF AGRICULTURE

FIGURE 2.—Typical apiary under a ramada that partially shades colonies in hot Southwest.

FIGURE 3.—Typical winter pack in plains region. Colonies placed side by side in groups with 6 or more inches of dry straw on top and wrapped in single layer of tar paper. BEEKEEPING IN THE UNITED STATES 19 Northeast honey and pollen in locations protected from wind and exposed to warming sunlight. Still others The severe winters, short summers, and hilly have most of the honey removed, and the hive or mountainous nature of the Northeast produce is reduced to a single brood nest that is trucked a variety of plants but no major source of honey. to the Southeast, where it is allowed to build up Whiteclover, basswood, black locust, birdsfoot and be divided to form new colonies. It is returned trefoil, various berries, and wild flowers contribute to the North in the spring for fruit pollination to producing a mixture of honey, much of which before the main honey flow. Midsummer shade is is sold locally to residents acquainted with the beneficial (fig. 4). Little migratory beekeeping types produced, and some of the highest prices occurs other than movement of the colonies to for honey are obtained here. Few commercial the Southeast for increase. beekeepers are in the Northeast. Some colonies are rented for pollination of The average honey production per colony is fruits, legumes, and cucumbers. only 33 pounds, but occasionally locations produce much higher averages. An estimated 187,000 Southeast colonies are in this region. The colonies are seldom moved except the few The production of honey per colony in this belonging to commercial or semicommercial bee- region, 33 pounds, is the same as in the Northeast keepers, who rent their bees for pollination of but lower than elsewhere. An estimated 1,595,000 blueberries, cranberries, other fruits, or cucumbers. colonies are located permanently in the Southeast. Colonies are located where there is good air In addition, many thousands of colonies are drainage, protection from the cold winds, and trucked here from the northern areas during the exposure to as much winter sun as possible. winter, then returned to the North in the spring. For additional protection from cold winters, Most of the queen breeders and package bee many colonies are ^^packed,^' or wrapped with shippers of the country are located within the insulation and tar paper, leaving only the entrance Southeast. An estimated 660,000 pounds of live exposed. Winter loss is usually high and is re- bees and an equal number of queens are shipped placed with packages of bees and queens purchased from here annually. Some of these beekeepers pro- from southern beekeepers. Shade in summer is duce no surplus honey for sale to supplement their unnecessary. sale of live bees and queens. Some of the northern Most beekeepers overwinter their colonies in beekeepers pick up their package bees and queens two- or three-story, 10-frame-deep Langs troth in van-type air-conditioned trucks for safe trans- hives. When the honey flow starts, they add one portation to their northern locations. or two extra deep supers for surplus honey storage Except for sizable areas in Florida, little polli- or one or two shallow supers for section or comb nation is provided on a cost basis in this region. honey production. Bees are rented for occasional fruit orchards and legume seed and melon fields. In Florida, bees are North-Central Region rented for citrus, cucurbits, and other fruits and vegetables. The bulk of the honey from the north-central In the mountainous area sourwood is the pre- region comes from whiteclover, alsike clover, and vailing quality honey, along with tulip poplar and alfalfa, with minor surpluses from basswood, the clovers. Sourwood honey is almost water black locust, and raspberry. All of this is high white, does not granulate readily, and is so es- quality honey. Clover and alfalfa are the predomi- teemed that it usually passes directly from pro- nant American honeys. Less desirable grades come ducer to consumer at far above the price of other from aster, goldenrod, and smartweed. The honeys. variety of other plants, however, insures something In the lower elevations gallberry becomes the for the bees to work on from spring until frost. predominant source. In the Apalachicola swamp The bulk of produced by the bees area, túpelo, famous for its high lévulose content in 1-pound sections comes from this region. and nongranulating characteristics, is also an ex- There are approximately 1,205,000 colonies, many cellent honey. Farther south in Florida, citrus is of which belong to commercial beekeepers. The the major source, with clovers the major source average production of surplus honey per colony toward the Mississippi Delta, then cotton becomes is 58 pounds. important. Various other honeys from light to Some colonies are killed in the fall and the equip- dark and from mild to strong are produced in the ment is stored; then hives are restocked in the Southeast. spring with packages of bees and a queen is pur- Considerable migratory beekeeping occurs, for chased from southern beekeepers. Others are the long season permits harvest of a crop of honey wrapped with insulation and tar paper for winter in one area before another harvest commences protection. Some are left with ample stores of elsewhere. 20 AGRICULTURE HANDBOOK 3 35, U.S. DEPT. OF AGRICULTURE

FIGURE 4.—Apiary sheltered by hardwood forest in north-central region. Hives composed of 11-frame 65^-inch-deep hive bodies.

Cut-comb honey production is common, that is plains region. One reason is that the sweetclover a chunk of comb in a jar of liquid honey. Little and alfalfa fields are relatively large and can sup- section honey is produced. port many colonies, and many of the apiaries Little work is necessary to prepare bees for belong to commercial beekeepers. winter. They are usually wintered in two- or three- Some of the colonies are trucked to southern story hives. The problem is to have ample stores areas for the winter, some are packed (fig. 3), of honey and pollen in the colony in the fall. This some are killed and then restocked in the spring, is necessary for the strong colonies needed in the and others receive no special winter treatment. early spring for package bee production or the Colonies are used to a limited extent in the pol- early honey flows. lination of alfalfa, sweetclover, and cucumbers. Colonies benefit from shade during the summer From this region westward to the Pacific, where in the Southeast, and shade is essential in the migratory beekeeping is piacticed to a greater southern part for maximum colony production. extent than elsewhere, the California-style top and bottom rather than the telescoping top and Plains Region reversible bottom are used as they permit better stacking of colonies on a truck. The bulk of the honey from the plains region comes from sweetclover and alfalfa, much of it prodviced by commercial beekeepers. Mountainous Region In this region about 396,000 colonies produce 76 pounds of honey per colony. Colonies are win- The major source of honey is alfalfa (fig. 5). tered and operated similarly to colonies in the About 434,000 colonies produce on an average 52 north-central region. Shade is not generally neces- pounds of honey per colony. More than half the sary, although partial or thorough shading during colonies belong to commercial beekeepers, who extremely hot midsummer days is beneficial. The may manage 2,000 or more colonies with only highest production per colony is obtained in the part-time summer help. BEEKEEPING IN THE UNITED STATES 21

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-*"*'."■'"'■■■''■«. ■j• — jV~ '•j„,a■-,■ .ji_ " '-VJEI ■. MJ

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•\ii-,'\.

Xlkt BN-30056 FIGURE 5.—Unsheltered colonies located for alfalfa honey production and pollination.

Honey production is almost entirely dependent on irrigation, although in recent years alfalfa is Southwest being grown on dry land. Weed spraying is In this hot semiarid region there are 230,000 reducing the sweetclover acreage. colonies that produce 63 pounds of honey per In the migratory beekeeping from this area colony. The major sources of honey are alfaKa, west and south, the colonies are usually moved at cotton, and mesquite. Other sources include citrus, night. The hive entrances are not closed but the catclaw, tamarix, sañíower, wild bvickwheat, and truckload is usually covered with a plastic screen. other desert shrubs. Some colonies are packed during the winter, which is extremely cold and dry—factors that im- Sunimer shade is highly important (fig. 2). prove survival chances over damp cold areas. Many Artificial shade is often provided. Winter pro- of the colonies are placed three or four back to tection is unnecessary. Some colonies are wintered back, then packed as a group. Colonies not packed in a single brood nest with one or two shallow are located where they have wind protection and supers, but most are in two or three deep supers. good air drainage. Nearby water is essential, and if it disappears Spring buildup is slow and fall flows are rare. even for only a day, the colonies may perish. Shading is unnecessary. Ample comb space for Migration from one honey flow to another is ripening honey is important and seems to act as common. heat insulation. Colonies are used extensively in pollination of Migratory beekeeping is extensive. For ex- alfalfa and melons and to a less degree for citrus, ample, in the Delta area of central Utah in the onions, and cotton. A few package bees and queens summer of 1966, there were over 40,000 colonies, are produced, but largely bees are kept for pro- but almost none wintered there. Some colonies duction of honey by commercial operators. are moved hundreds of miles to desirable areas. Apiaries of 100 colonies or more are not unusual. Many are moved south or west for the badly needed spring buildup, then returned for the West summer flow. Some colonies are killed in the fall and restocked in the spring. About 717,000 colonies in this region produce Not much honey is retailed by the producer in 58 pounds of honey per colony. This production the mountainous region. is rather meaningless because of difi'erences due 22 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

to extreme variations in temperature, rainfall, This excessive migratory operation has increased elevation, and flora. Clover, alfalfa, citrus, sage, the bee disease problem, because the colonies wild buckwheat, cotton, star-thistle, and fire- have so many more chances to be located in weed are the major honey sources. areas where disease exists. Conversely, the migra- The region varies in rainfall from only an inch tory blends well with the use of colonies or two in the desert areas to more than 60 in the rain-forest area, in elevation from below sea for pollination. The placement of 2,000 colonies level to snowcapped mountains, and in temper- from several beekeepers in a solid square mile ature from dry and hot to humid and extremely of alfalfa grown for seed is not unusual. Use of cold. Colonies in the mountains must be protected bees for pollination is extensive. An estimated from the cold and with fences from bears, where- half of all colonies are used some time during as colonies in the lower ranges to the south must the year for pollination hire. be protected from the heat. In addition to honey production and polli- Migratory beekeeping is practiced by most of the commercial beekeepers, and four or more nation services, there are wax salvage plants. moves per year are not uncommon. An average Diseased equipment is taken to these plants and beekeeper will winter his bees on the coast, move the wax is steamed from it and salvaged. The to almonds, then manzanita and sage, then to equipment is used again. alfalfa and cotton, and back to the coast for the Some of the beekeepers operate many thou- fall flow. Productive accessible locations are diffi- sands of colonies. Under such operation the apiary cult to find. Many colonies are reduced to a single rather than the colony is considered a unit. Such story for moving, then given extra supers in which to expand. The California-style top and manipulations as requeening, supering, and re- bottom are almost exclusively used in this region. moving honey are performed on all colonies Most commercial beekeepers use mechanical hive regardless of their relative condition. More than hoists that lift one or more colonies at a time 300,000 pounds of bees and 250,000 queens are onto the truck. shipped annually from the West. MANAGING COLONIES FOR HIGH HONEY YIELDS

By F. E. MoELLER, apiculturist, Entomology Research Division, Agricultural Research Service

Colonies of bees existing in the wild away from weaker ones, (2) temperamental bees are more man's control will produce small surplus crops of reluctant to accept a new queen than gentle bees, honey above their requirements for survival. Such (3) young bees accept a queen more readily than surplus will vary depending on the region or older bees, (4) the colony to be requeened should locality, but will seldom exceed 30 to 35 pounds. first be made queenless, and (5) the queen to In the same area subject to the same nectar re- be introduced should be in egg-laying condition. sources, colonies properly managed will produce There is less risk in requeening a colony by surplus honey crops of 150 to 200 pounds. Inten- giving it a laying queen with some of her own brood sive two-queen colony management can often and bees than by giving it a queen in a shipping result in surplus crops of 500 pounds or more with cage. A new or valuable queen should first be the same resources available. The key to these introduced into a small colony or divisions of one differences is management. in a queen shipping cage. After she is laying, the Proper management employs practices that small colony can be united with a large one. harmonize with the normal behavior of bees and A drone-laying queen can be replaced if she is brings the colony to its maximum population discovered while the colony is still strong. If the strength at the start of the bloom of major nectar- colony is weak, the bees should be removed and producing plants. Management practices are simi- the equipment added to another colony. lar in basic principle wherever bees are kept and Assuming colony conditions and the condition of vary only as regards timing for the desired nectar the queen are favorable, the effect of environmen- source of the region or locality concerned. tal or working conditions and the time of year are Regardless of the type of hives or equipment factors that affect queen acceptance. Best accept- used, proper management aims at providing ance is usually obtained when some nectar is colonies with unrestricted room for brood rearing, available in the field. ripening of nectar, and storage of honey, plus One possible period for requeening is during the provision of adequate food requirements, both broodless period of late fall. Queens are easily in- pollen and honey, for the time of year concerned. troduced at this time, and the bees are passive to Swarming is minimized and the storing instinct their presence. However, the uncertainty of the encouraged when proper management is used. weather, the difficulty of finding old and shrunken queens, and the danger of inciting robbing make Preparing Colony for New Season this time of year less desirable for requeening than the summer. In the temperate regions of the northern hemi- Brood rearing declines in late summer and fall, sphere, August to October is the time when the and many normal colonies are completely brood- beekeeper prepares his colonies for the coming less during much of November and December, year. This is when the major honey flows are particularly if the colony has no pollen. Older usually past and the bees must be made ready queens stop brood rearing sooner than younger for the coming winter. queens. All queens of questionable performance with Brood rearing should be encouraged as late in only a small amount of brood of irregular pattern the season as possible. This can be assured by (fig. 1, A) should be replaced. Frequently providing vigorous young queens in late summer, the bees of the colony will replace or supersede by preventing undue overcrowding and restriction queens of subnormal performance even before the of the brood nest with honey, and by encouraging beekeeper senses a problem. Some queens may be pollen storage. satisfactory in their second year; queens less than a year old are usually best. In areas where fall honey flows occur, partially To requeen a colony, certain principles of queen filled supers should be kept on the colonies, acceptance must be borne in mind: (1) Strong especially if the brood nest is heavy. colonies more reluctantly accept a queen than If brood rearing is restricted by a crowded brood nest or because of poor queens, the colony may 1 In cooperation with Wisconsin Agricultural Experi- enter the winter with a high percentage of old bees ment Station. that will die early in the winter. Such colonies may 23 24 AGRICULTURE HANDBOOK 33 5, U.iS. DEFT. OF AGRICULTURE

FIGURE 1.—Queens with (A) irregular and (B) good brood pattern. BEEKEEPING IN THE UNITED STATES 25 later develop serious nosema infections and perish they used over winter, whereas other colonies that before spring. A colony should start the winter have had normal, unimpeded rearing of brood will with about 10 pounds of bees and plenty of honey soon be able to replace all the honey they con- to carry it to the next spring. sumed over winter plus a substantial surplus. Preparing Colony for Winter Organization To accommodate the best queens in standard Population Langstroth 10-frame hives, a minimum of two hive bodies and preferably three should be used for The strength of a colony of bees is relative and year-round management. In the fall most of the difficult to describe. A ''strong^' colony to one bee- honey should be located in the top hive body. With keeper might be ^ Veak'' to another. Colonies with experience the beekeeper can soon learn to estimate less than 10 pounds of bees should be united to the weight of hive bodies or frames by lifting them. stronger ones or several weaker ones combined. A frame full of honey should weigh approximately At between 40° and 50° F., 10 pounds of bees will 5 pounds. The top hive body should contain 40 to cover practically all the combs of a three-story 45 pounds of honey. This means that all frames in hive wall to wall and top to bottom. Naturally as the top hive body will be full of honey except for the temperature drops the cluster will contract. two or three frames in the center. The second body The beekeeper must see that at no time is the should contain 25 to 30 pounds of honey and some available space for brood rearing reduced because pollen. The bottom hive body should contain 20 to of overcrowding with honey from the fall flow. A 30 pounds of honey plus pollen. If in the fall the balance must be maintained between crowding the combs in the top hive body are not filled, the colony to get the brood chambers well filled with beekeeper should reorganize them and if necessary honey and adding space to relieve brood rearing feed additional sugar sirup so that this top hive restriction. Partially filled supers kept on colonies body is well filled with stores. in the fall may be necessary. Any subnormal colony As the winter progresses the cluster of bees will should not be overwintered, but should be united shift its position upward as the stores are con- with another colony. sumed. A colony of bees in a cold climate can A colony may appear to have an adequate fall starve with abundant honey in the hive if the population, but if the bees are old, it will weaken honey is below the cluster. rapidly as winter advances and may starve to With the advent of cold weather, the bees death, even with abundant honey in the hive, cluster tightly in the interspaces of the combs. because the cluster is too small to cover the honey Usually there are no bees in the bottom part of stores. the hive near the entrance. For this reason an entrance cleat or reducer should be used to ex- Food Reserves clude mice. One-inch auger holes drilled into the hive bodies of the brood nest just below the hand- The colony should have a minimum of 500 holds are helpful. In late summer these auger-hole square inches of comb filled with pollen in the fall. entrances are closed with corks so that the bees To insure uninterrupted brood rearing in late win- will fill the combs near them. During winter the ter and early spring, the beekeeper may need to top auger-hole entrance should be open. This supplement this. The average colony of bees under allows the escape of moisture-laden air and affords intensive management may consume about 60 a flight exit for the bees during warm spells. pounds of honey between the last fall flow and the (Fig. 2.) first available food from the field in the spring. A weak colony may consume 20 pounds or less, but the very best colony will consume 80 pounds Packing the Hive or more. To insure the survival of the top quality Many beekeepers in the coldest parts of the colony, 90 to 100 pounds of honey should be left country consider that some form of protection on it in the fall. A colony of bees not rearing brood around the hive is essential. Others believe that will consume an average of about one-eighth pound colonies with strong populations and ample stores of honey per day or 5 pounds per month. When need no further protection. Factors to consider in brood rearing begins, the consumption of honey deciding whether or not to pack are the cost of is greatly accelerated. Brood rearing should com- material and labor and any savings in honey or mence in midwinter and accelerate as temperatures bees. Packing will not replenish colonies deficient moderate in late winter and early spring. in honey, pollen, or bees, replace poor queens, or When brood rearing is discouraged or curtailed, cure bee diseases. Packed colonies will consume the colony will consume less winter stores but will slightly less honey. However, the difference is neg- emerge in the spring much weaker and with a ligible. The most important consideration in pre- population of primarily old bees. Such colonies will paring colonies for winter is a strong population have difficulty replacing the small amount of honey and adequate stores. 26 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE From this period until the bees can forage, such colonies can be fed either full combs of honey, or if these are not available, a gallon or two of heavy sugar sirup (two parts sugar by volume to one part :fc 'i water) can be poured directly into the open cells of empty combs.

# Spring Buildup Overwintered colonies will usually start brood rearing in midwinter and continue into the summer unless the stored pollen is all consumed before fresh pollen is available. If the supply is exhausted and not supplemented, brood rearing -will slow down or stop entirely when it should proceed without interruption. For best results in honey production, a bee- keeper should have strong populations of young bees for the honey flow. Colonies emerging in the spring with predominantly old bees must build a population of young bees for later flows by using the early sources of pollen. Some beekeepers trap pollen at the hive en- trance from incoming bees by means of a pollen trap such as that described in U.S. Department of Agriculture ARS 33-111, A Simplified Pollen Trap for Use on Colonies of Honey Bees. This pollen is dried or frozen until needed, then mixed \vith sugar, water, and soy flour, and fed to the FIGURE 2.- -Colony during winter allowing top auger-hole colony as a supplement to its natural supply entrance. (fig. 3). Various other types of pollen supplements and substitutes have been described and some are When outside temperatures are near freezing, available on the open market. the temperature at the surface of a cluster of bees Supplements containing pollen are eaten more ranges between 43° and 46° F. As the temperature readily by bees and generally give better results decreases, the cluster contracts and the bees in the than those containing soy flour or other material outer insulating shell concentrate to provide an without pollen. Pollen supplement is preferred insulating band of 1 to 3 inches in depth. Metab- by the bees in direct proportion to the amount olism and activity of the bees in the center of the of pollen it contains. The less pollen the supple- cluster maintain a desired temperature. This may ment contains, the less is eaten. Substitutes be around 92° if brood rearing is in progress. The made without pollen tend to be dry and gummy. temperature of the area of the hive not occupied A pound of pollen will make approximately 12 by bees will be similar to the external tempera- pounds of pollen supplement (see p. 55). ture. This is true whether the hive is packed or not. The difference is that the temperature in the Swarm Control in Single-Queen unpacked hive changes more rapidly and responds more quickly to that outside the hive. Heavy Management packing is worse than no packing, because during warm periods in midwinter when the bees should After pollen becomes abundantly available in fly, those heavily packed do not fly at all. the spring, the beekeeper should provide ample space for brood rearing and honey storage. The natural colony behavior is to expand its Late Winter Manipulation brood nest upAvard, and a simple manipulation utilizing this tendency is to shift the empty If colonies are inspected in late winter or early frames or emerging brood to the top of the hive spring, adjustments can be made to save colonies and the youngest brood and honey to the bottom that might be lost otherwise. Even weak or me- part. This permits the expansion of the brood dium-strength colonies can often be saved if honey rearing upward into this area (fig. 4). Subsequent is moved into contact with the cluster. A strong reversal of brood chambers can be made at about colony with insufficient honey can starve if addi- 10-day or 2-week intervals until the honey flow tional food is not provided at this time. starts. BEEKEEPING IN THE UNITED STATES 27

FIGURE 3.—Strong colony feeding on pollen supplement cake during March. As soon as the three brood chambers are filled with bees, the first super should be given whether Two-Queen System or not the honey flow is in progress. If this is done, most colonies with a vigorous queen will The establishment of a two-queen colony is not swarm. However, any queen cells the bee- based on the harmonious existence of two queens keeper sees as he reverses the brood chambers in a colony unit. Any system that insures egg should be removed. A simple method of reversing production of two queens in a single colony brood chambers is to lower the hive backward for about 2 months prior to the honey flow will to the ground, separate the brood chambers, inter- boost honey production. change the first and third hive bodies, and return The population in a two-queen colony may to position. be twice the population of a single-queen colony. After the honey flow starts, the danger of Such a colony will produce more honey and pro- swarming lessens and brood chamber reversal can duce it more efficiently than will two single-queen be discontinued. At the start of the honey flow, colonies. A two-queen colony usually enters winter "bottom supering" should be used. The empty with more pollen than a single-queen colony. As super should be placed above the top brood a result of this pollen reserve, the two-queen chamber but below the partially filled supers colony emerges in the spring with a larger popu- (fig. 4). lation of young bees and is thus a more ideal unit After the supers have been fiUed and the honey for starting another two-queen system. has been extracted, they should never be put To operate two-queen colonies, start with strong directly over the brood nest, but should be placed overwintered colonies. Build them to maximum on top of the partially fiUed supers to prevent strength in early spring. Obtain young queens the queen seeking them and laying eggs in them. about 2 months before the major honey flows Why such combs are attractive to the queen is start. When the queens arrive, temporarily divide not known. the colony. Replace the old queen and most of the 28 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

second queen excluder is used to keep the queen SUPER 1 out of them. The top brood nest may tend to be- 2/3 FULL come honey bound. If this occurs, reverse the upper and lower brood nests around the queen Pulls older bees excluder. This puts the top honey-bound brood up for honey SUPER 1 SUPER 2 nest on the bottom board and the lighter brood 2/3 FULL 1/3 FULL storage nest with the old queen above the excluder. There is no advantage in having a second queen I when about a month of honey flow remains, be- SUPER 2 SUPER 3 cause eggs laid from this time on will not develop 1/3 FULL EMPTY SPACE into foragers before the flow has ended. However, CENTER OF entering the brood nest during the middle of the HEAVIEST COLONY ACTIVITY BROOD EMERGING (Congestion relieved) flow to remove one of the queens is impractical. CHAMBER BROOD Uniting back to a single-queen status can be done AND OR YOUNGEST SPACE after the bulk of the honey is removed from the BROOD colony. By this time some of the colonies may SEALED SEALED have already disposed of one queen. When this AND AND Pulls young bees EMERGING EMERGING happens, all that needs to be done is simply to BROOD BROOD down for nursing remove the queen excluder and operate the colony HEAVIEST as a single-queen unit. EMERGING BROOD BROOD CHAMBER AND (OR) 1 AND SPACE YOUNGEST Improved Stock BROOD BEFORE AFTER Production of honey is one major criterion in selecting honey bee stock and breeding for im- MANIPULATION BN-30049 provement. Superior stock must also be reason- ably gentle, not prone to excessive swarming, FIGURE 4.—Basic colony manipulation for swarm control. maintain a large but compact brood nest, and winter well. It should ripen its honey rapidly, seal the cells with white wax, and use a minimum of younger brood plus about half of the population burr comb. To obtain all the desirable characters in the bottom section. Cover with an nner cover in a superior stock, specific inbred lines from many or a thin board and close the escape hole. The divi- sources must be selected and developed and then sion containing most of the sealed and emerging recombined into a genetically controlled hybrid. brood, the new queen, and the remainder of the When this is done, hybrid vigor or heterosis population is placed above. The upper unit is usually results. provided with an exit hole for flight. Queens of common stock reared under favorable At least two brood chambers must be used for conditions and heading well-managed colonies the bottom queen and two for the top qiueen. Two will probably be more productive than poorly weeks after her introduction, remove the division reared queens of superior stock. Queens of superior board and replace it with a queen excluder. The stock reared under favorable conditions will re- supering is double that required for a single-queen quire a higher standard of management than is operation. In other words, where three standard demanded of common stock. To realize the maxi- supers are necessary for a single colony, six will be mum benefits from improved stock, the beekeeper required for a two-queen colony. must provide unrestricted room for brood rearing, When supering is required, larger populations in ripening of nectar, and storage of honey. two-queen colonies require considerably more room To realize the maximum benefits from improved at one time than is required for single-queen colo- stock, the queen breeder should produce the best nies. If a single-queen colony receives one super, a queens possible, and the honey producer receiving two-queen system may require two or even three these queens should manage them in such a way empty supers at one time. that they can develop their maximum colony The brood chambers should be reversed to allow populations. normal upward expansion of the brood area about every 7 to 10 days until about 4 weeks before the expected end of the flow, after which the honey Disease Control as Affected by crop on the colony may be so heavy as to preclude Good Management any brood nest manipulations. Thereafter give supers as they are needed for storage of the crop. If colonies are operated for highest honey yields, As the honey is extracted, the supers are returned they must be kept in optimum condition (fig. 5). to the hive to be refilled. They should never be This includes rigid control of all bee diseases. For replaced directly over the top brood nest, unless a information about bee diseases, see pages 86-96. BEEKEEPING IN THE UNITED STATES

FiGUEE 5.—Brood combs showing (A) healthy brood necessary for high honey production and (B) diseased brood, which results in weakened colonies and low honey production.

(4) References 1944. PRODUCTIVE MANAGEMENT OF HONEYBEE (I) FARBAR, C. L. COLONIES IN THE NORTHERN STATES. U.S. 1930-31. INFLUENCE OF THE COLONY'S STRENGTH Dept. Agr. Cir. 702, 28 pp. ON BROOD REARING. Ontario Beekeep- (5) ers' Assoc. Ann. Rpt. 51-52, pp. 126- 1952. ECOLOGICAL STUDIES ON OVERWINTERED 130. HONEY BEE COLONIES. Jour. Econ. Ent. (2) 45: 445-449. 1937. INFLUENCE OF COLONY POPULATIONS ON (6) HONEY PRODUCTION. Jour. Agr. Res. 54: 1960. OLD AND NEW IDEAS ABOUT WINTERING. 945-954. Amer. Bee Jour. 100: 306-310. (3) (7) HARP, E. R. 1942. NOSEMA DISEASE CONTRIBUTES TO WINTER 1966. A SIMPLIFIED POLLEN TRAP FOR USE ON LOSSES AND QUEEN SUPERSEDUBE. Glean- COLONIES OF HONEY BEES. U.S. Agr. ReS. ings Bee Cult. 70: 660-661, 701. Serv. ARS 33-111, 4 pp. 261-907 0—67—3 30 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

(8) HooPiNGARNER, R., and FARRAR, C. L. FECTED BY HONEY BEE STOCK LINES. U.S. 1959. GENETIC CONTROL OF SIZE IN QUEEN HONEY Dept. Agr. Prod. Res. Rpt. 55, 20 pp. BEES. Jour. Econ. Ent. 52: 547-548. (11) and HARP, E. R. (9) MOELLER, F. E. 1965. TWO-QUEEN SYSTEM SIMPLIFIED. Glean- 1956. BEHAVIOR OF NOSEMA-INFECTED BEES AF- ings Bee Cult. 93: 679-682, 698. FECTING THEIR POSITION IN THE WINTER (12) ScHAEFER, C. W., and FARRAR, C. L. CLUSTER. Jour. Econ. Ent. 49: 743-745. 1946. USE OF POLLEN TRAPS AND POLLEN SUP- (10) PLEMENTS IN DEVELOPING HONEYBEE COL- 1961. THE RELATIONSHIP BETWEEN COLONY POP- ONIES. U.S. Bur. Ent. and Plant Quar ULATIONS AND HONEY PRODUCTION AS AF- E-531, rev., 13 pp. BEE BEHAVIOR By STEPHEN TABER HI, apiculturist, Entomology Research Division, Agricultural Research Service ^ Bee behavior refers to what bees do—as indi- There is no known governmental hierarchy viduals and as a colony. By studying their be- giving orders for work to be done, but a definite havior we may learn how to change it to our effect on the colony is observed when the queen benefit. disappears. This effect seems to be associated Two very practical discoveries of bee behavior with a complex material produced by the queen made our beekeeping of today possible. One was that we refer to as ^^ queen substance.'^ There is the previously mentioned discovery by Langstroth also evidence that the worker bees from 10 to 15 of bee space. The other was the discovery by G. days old, who have largely completed their M. Doolittle that large numbers of queens could nursing and household duties but have not begun be reared by transferring larvae to queen cells. to forage, control the ^^governmentaF^ structure. The discovery of the ^language" of bees and their Just what controls them has not been determined. use of polarized light for navigation have attracted These and many other factors make an considerable interest all over the world. organized colony out of the many thousands of Much has been learned about the behavior of individuals. insects, including bees in recent years. As an ex- The Domicile ample, the term ^'pheromone'^ had not been coined in 1953, when Ribbands summarized the subject When the swarm emerges from its old domicile of bee behavior in his book, The Behaviour and and settles in a cluster on a tree, certain ^'scout Social Life of Honeybees. A pheromone is a sub- bees'' communicate to it the availability of other stance secreted by an animal that causes a spe- domiciles. At least some of these domiciles may cific reaction by another individual of the same have been located by the scout bees before the species. Now many bee behavior activities can be swarm emerged. The various scouts perform explained as the effect of various pheromones. Un- their dances on the cluster to indicate the direc- fortunately for apiculture much of the study of tion, distance, and desirabiUty of the donaiciles. pheromones has been connected with insects other Eventually the cluster becomes united in its than the honey bee. approval of a particular site. Then the swarm moves in a swirling mass of flying bees to it. Recently we have learned how certain bee be- Agreement is always unanimous. havior activities are inherited, and this informa- tion gives us a vast new tool to tailor-make the Wax Combs honey bee of our choice. Further studies should reveal other ways to change bees to produce As soon as the swarm enters the new domicile, specific strains for specific uses. food is collected and wax comb is built. The wax they use is secreted in tiny flakes from glands on the underside of the worker bee abdomen—but The Honey Bee Colony only when fresh food is available. The wax is mold- The physical makeup of a colony has been de- ed by the bee mouth parts to form the intricate scribed (p. 5). An additional requirement of a comb. The first comb consists of about 25 cells per colony is a social pattern or organization, prob- square inch. This is the size worker bees are ably associated with a ^'social pheromone.'^ It reared in. After there is a considerable amount causes the bees to collect and store food for later of worker brood and increased population of bees, use by other individuals. It causes them to main- comb containing larger cells is built. This comb tain temperature control for community survival is used for rearing drones. when individually all would perish. Individuals Although most references indicate that the within the colony communicate with each other presence of drone cells, even in small quantities, but not with bees of another colony. Certain bees IS objectionable, it is natural and normal for these in the colony will sting to repel an intruder even cells to be present. They may have a morale- though the act causes their death. All of these, boosting or possibly some other beneficial effect. and perhaps many other organizational activities, The space between combs inside a colony are probably caused by the social pheromone. varies greatly. Worker brood comb is about an inch thick. The open space between brood combs 1 In cooperation with Arizona Agricultural Experiment in a natural cluster is about three-eighths inch Station. but varies from one-fourth to 1 inch. 31 32 AGRICULTURE HANDBOOK 3 35, U.S. DEPT. OF AGRICULTURE

The space between honey storage combs is flowering period, usually the more intense the much more uniform than between brood combs. swarming ''fever'' in an apiary. Some genetic The space left between capped honey cells is lines of bees are much more prone to swarm than usually one-fourth inch or even less—room others. enough for one layer of bees. One likely cause of swarming may be a change As the colony ages, the combs that were first in production of certain pheromones by the queen used for rearing worker bees may be converted bee that affects the workers. Swarming can often to honey storage comb ; areas damaged in any way be discouraged by giving the bees more room above are rebuilt. These changes usually affect the bee the brood nest area into which they can expand. space and result in combs being joined together with ^'brace'' comb. Strains of bees show genetic variation in building these brace combs. Housecleaning All these cells are horizontal or nearly so; Certain waste material accumulates in a normal vertical cells are used for rearing queens. Why colony. Adult bees and immature forms may die. horizontal cells are used for the rearing of brood Wax scales, cappings from the cells of emerging and for honey and pollen storage, whereas ver- bees, particles of pollen, and crystallized bits of tical cells are built only for queen production is honey drop to the floor of the hive. Intruders, such unknown. as wax moths, bees from other colonies, and pred- ators, are kiUed and faU to the floor. Worker bees Flight Behavior remove this debris from the hive. In some ants this behavior pattern is controlled by certain pher- When several thousand bees and a queen are omone-like chemicals. When these chemicals are placed in new surroundings—^which happens applied to live ants of the same colony, their sis- when the swarm enters its new domicile or a ters bodily haul them out to the graveyard. A package of bees is installed, or a colony is moved pheromone of a similar nature may occur in the to a new location^—^normal flight of some workers honey bee colony. from the entrance may occur within minutes. The cleaning behavior associated with removal If flo^yering plants are available, bees may be of larvae and pupae that have died of American returning to the hive with pollen within an hour. foulbrood in the cells is known to be genetically Bees transferred by air from Hawaii to Louisiana controlled by two genes. It is modified by honey- and released at 11:30 a.m. were returning to the flow conditions. What makes this discovery impor- new location with pollen loads within an hour. tant is not only that it should help in developing Package bee buyers in the Northern States have bees more resistant to this and related bee diseases, noticed similar patterns in bees shipped from the but also that we can now expect to find that many South. of the other behavior characteristics of bees can be What causes this virtually instant foraging by modified to suit special needs, with mutual benefit bees? What determines whether they collect to beekeepers and farmers. pollen, nectar, or water? If food and water in the hive are sufficient, why should they leave to forage? Does a pheromone or a hormone cause Brood Rearing this flight activity? Answers to these questions The seasonal changes in egg production by the can lead to directing bees to specific duties we queen and the subsequent rearing of brood by the desire accomplished. workers are thought to be mainly dependent on the supply and abundance of nitrogenous and car- Swarming bohydrate food. However, brood rearing slows down and stops in the fall, then starts again during The basic causes of swarming are not under- midwinter in both Louisiana and Wisconsin, States stood. Usually a consistent sequence of events geographically and cUmatically dissimilar. The occurs prior to swarming, but the actual swarm buildup of fat bodies in fall and winter bees coin- may not emerge even if afl the events occur. cides with similar physiological changes in other Workers are first reared in great numbers; species of insects, such as the boll weevil going then comes a period marked by the rearing of into a period of diapause or inactivity. Initiation both workers and drones. Large quantities of or control of diapause has been shown to be regu- pollen and nectar are brought into the hive. This lated by quantity and quality of light. crowds the brood nest and restricts the number of An important aspect of bee behavior is the pos- eggs the queen is able to lay. From 10 to 50 queen sible effect of the 24-hour or circadian rhythm on cells are produced, and shortly after the first daily brood rearing activities. one is sealed the swarm issues. Much of the brood rearing activity cycle of colo- Swarms usually emerge from the hive in the nies during the year can be related to the available late morning and often prior to the main flower- forage or plants in bloom. Colonies in different ing or honey-flow period. The more intense the parts of the country show fluctuations in brood BEEKEEPING IN THE UNITED STATES 33 production and in their subsequent colony popula- careful manipulations. The right amount of smoke tions that are coordinated with forage conditions. to use on a colony is learned only by experience. Colonies in the Northeastern and Midwestern United States increase their brood rearing and bee Temperature Control by Bees populations during maple-dandelion-fruit bloom periods and show an intense swarming tendency Bees are paradoxical in that they are individ- prior to bloom of clovers. The brood and adult ually cold-blooded insects but collectively behave populations generally and slowly dechne during like a warm-blooded animal. During periods of late summer, but increase in areas with abundant high temperature they bring in water, which on acreages of aster, goldenrod, and smartweed. A evaporation cools the cluster to the desired tem- severe decline in brood rearing follows the first perature. During periods of low temperature they killing frost. But in areas of New York, Pennsyl- cluster tightly and generate and conserve heat to vania, and Wisconsin, where the late flowering hold the temperature up so that the center of the buckwheat is grown, greatly increased brood bee cluster rarely gets colder than 80° F. Neither rearing, population increase, and even swarming the exact mechanism of cooling nor the building occur in the early fall. up of heat in the cluster is understood. None of Beekeepers in the Western United States de- the proposed theories have been proved. pend on flowering of irrigated agricultural crops and indigenous plants stimulated by infrequent Colony Morale rains for buildup of brood and adult bee popu- lations. In the Southwest the farmers on irrigated ^'Colony morale^' generally refers to the well- land need bees on specific, predictable dates for being of the colony. If the colony morale is good, pollination of their crops. The beekeeper who the bees are doing what is desired of them, in- must supply these bees is dependent on unpredict- cluding increasing the colony population, making able rainfall to provide natural food for his col- honey, and pollinating flowers. Many factors onies to develop brood and bee populations needed seem to affect colony morale. For example, if the for this service. queen is removed from a colony during a honey In the Southeastern States many different floral flow, the daily weight gains inmiediately decrease, sources bloom abundantly at different but specific although the bee population for the next 3 weeks times of the year. Brood rearing and colony ex- is unaltered. Also when a colony is preparing to pansion commence early, and high populations are swarm, the bees practically stop gathering pollen maintained with relatively little difficulty from and nectar. Improper manipulations or external swarming. Brood rearing usually declines some- environment also affects colony morale. what during high summer temperatures when few flowering plants are available for bee activity, Known Pheromone Activity then picks up again late in the summer and con- tinues until a killing frost. Some of the bee colony pheromones and their Variation in the quantity of brood reared by biological action are known to beekeepers. The colonies of bees nationwide is obviously affected Nasanov or scent gland was described over a by the immediate local environment. Yet with hundred years ago. The biological behavior activ- these apparent differences there are also similar- ity of its pheromone is best seen when a swarm ities associated with photoperiods or other external is hived. When the bees first enter the new domi- stimuli rather than the quantity of food available. cile, some bees stand near the entrance and fan. At the same time, they turn the abdominal tip downward to expose a small, wet, white material Temper and Its Control on top of the end of the abdomen. This seems The temper or gentleness of bees determines to affect the other bees, for within several minutes their inclination to sting. Many factors affect all will have settled and entered the new hive. their temper, including the genetics or inheritance When bees find a new source of food, they mark of the bee, environment of the hive, and manip- it with the same scent gland. A Canadian re- ulation of the colony by the beekeeper. The temper search team has recently reported isolation and of the colony can be temporarily controlled by identification of this pheromone. man with a certain amount of smoke. Exactly Colony odor refers to the odor of an individ- why and how smoke affects bees is unknown, even ual colony. Because each colony odor is different, though it has been used by beekeepers worldwide the colonies cannot be combined in the same for over a hundred years. The amount of smoke hive without fighting and killing one another. needed to control the temper of a colony varies Colony odor probably results from a combina- with time, temperature, and various other ex- tion of endogenous (pheromone or pheromone-like ternal factors, as well as with the inherent gentle- materials) and exogenous (food accumulation and ness of the colony. Ruthless manipulations that food interchange) materials in each hive and injure or kill bees create more need for smoke than seems to be recognizably different for every colony. 34 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

The usual procedure for the beekeeper when this sound could lead to protection of bees from colonies are to be combined is to place a news- harmful pesticides. paper between the two sets of bees. By the time Experienced beekeepers recognize a difference the bees have eaten through and disposed of the in sound between a colony with a queen and one newspaper, their odors have intermingled and without, between a ''mad" bee and an undisturbed become indistinguishable. During heavy honey one, and between normal and cold or starving bees. flows, differences between colonies seem to dis- Individual queens and even worker bees emit appear, and colonies can be united without squeaky sounds called ''piping" and "quacking." difficulty. When a bee returns from a foraging trip and One of the most interesting and complex performs a dance, she communicates the kind pheromones, originally termed '^queen substance," of "plant" or "flower" on which she was foraging is now believed to be a complex of different by releasing a taste or the perfume of the flower chemical pheromone compounds, which stimulates through nectar régurgitation or on body hairs. a large number of complex behavior responses. This has prompted other experiments designed Its presence in virgin queens in flight attracts the to train or force bees to collect or work a desired drone for mating from an unknown distance. Its crop. These experiments have so far been unsuc- presence in virgin and mated queens prevents the cessful. The reason for the failures may well be ovaries of the worker bee within the hive from that the bee language code has not been completely developing and the worker bees from building translated. We are still unable to "talk" effectively queen cells. It keeps swarming bees near the queen. to the bees and "tell" them what we want done. Its decrease is a cause of swarm preparation or Bees also recognize and are guided by different supersedure. Queen substance is produced in colors but are unable to communicate these glands in the queen's head. If these glands are colors. Their eyes are receptive to the polariza- removed, the queen presumably produces no more tion of the light in the sky and this aids in their queen substance, yet the queen and the colony go navigation. on much as they did before. The quantity and quality of queen substance vary in virgins and Age Levels of Bees Correlated With mated queens of different ages. Work Habits The alarm or sting pheromone, which also may be a complex of pheromones, has been tentatively The bee is adaptable to many environments. identified by a Canadian research team. When a Honey bees native only to Europe, Asia, and bee stings, other bees in the immediate vicinity Africa have adapted well to all but the polar also try to sting the same spot. In this case the regions of the world. Part of this adaptability sting pheromone provokes other bees to sting the lies in the capacity of the individual bee to "sense" same place, and they in turn provoke additional what must be done, probably through reception stings ad infinitum. Smoke blown onto the area of colony pheromones, then to perform the neces- seems to neutralize this effect. sary duty. Whether the genetic basis for a difference in Under normal conditions there are bees of all temper is in the quantity of the alarm pheromone ages in the hive. The age of the bee determines or pheromones produced or in the reception organs in general its daily activity. However, when con- of other bees is unknown. ditions become abnormal, age is no longer a criterion of duties. If a colony is made up of old Other Known Methods of Bee bees, for instance, some of them will become physiologically young, and conversely if it is made Communication up of all young bees, some will become physiolog- Besides the known and possible methods of bee ically old and take on duties normal to such bees. The reasons and causes for bees changing w^ork communication or language that have been men- tioned involving the chemical pheromones, there in this manner are unknown. are, of course, others. The best known is the so-called ^'dance'' of the returned forager bee. Behavior and Activities Outside This dance tells other bees precisely in which direction to search for food and how far they the Hive should fly, the type of food they will find, and the Genetically we have found that some bees pro- relative quantity. Minor genetical variations in duce more honey than others, but we do not kno\\ individual dance behavior exist. How this genetic why. The individual bee may collect more be- variance can be used in an economic way is cause of its own genetic inheritance. The colony unknown. The cluster itself and individual bees in may store more honey because of the queen's the field make subtle noises or sounds, all of which inherited ability to lay more eggs, resulting in are not at present understood. One such sound a greater total population of bees in the hive, seems to tell the bees to stay home. Utilization of or because the bees are inherently longer lived. BEEKEEPING IN THE UNITED STATES 35

These activities would have the same gross effect Control of Foraging of increasing the population of the colony. We can, as ''bee controllers/' affect the bee's A major goal is to control the foraging bee and environment in conjunction with its inheritance. get it to more effectively pollinate the plant. Bees There is no reason to believe that bees work should be attracted to areas where they are better when in want. Since evidence indicates desired, e.g., pollination areas, and repelled from the reverse, there should always be an ample areas where there is danger to them from insec- supply of reserve honey in the hives. ticides or where they endanger people. Work Another environmental factor is colony manip- with other insects—both social and nonsocial— ulation. A beekeeper's disturbance of the colony indicates that this could be accomplished by during the honey flow results in a marked decrease chemical and physical means. in the amount of honey stored for that day and Actually beekeepers have been using various even the following day. Colonies of bees should not chemicals to control bees in the hive for some be needlessly disturbed; however, manipulation to time. Smoke, a combination of chemicals in give them extra room to expand the cluster or gaseous form, and various chemical repellent store food is necessary. compounds have been used for years to repel bees. There is considerable evidence that different Pollination—Crop Problems plant species produce varying attractant com- A problem associated with bee behavior is in pounds associated with their nectar and pollen. getting bees to work on a particular field. Growers Bees are highly attracted to the scent of recently of seed and fruit crops rent bees in ever-increasing extracted honeycomb and to the scent of honey being extracted or heated. During periods ^ of quantities. They want the bees they rent to stay intense honey flow, bees much prefer collecting on their acreage. nectar of lower sugar concentration than exposed One method being explored at the present time takes advantage of inherited behavior differences. honey. Obviously certain chemical scents associ- Today a bee is being bred that is specifically suit- ated with certain flowers and to some extent incorporated in the collected honey are highly able for alfalfa pollination. Bees specifically de- attractive to bees. signed genetically to pollinate certain other crops Certain chemical extracts of pollen strongly will eventually be developed. indicate that some pollens contain compounds In conjunction with these is genetic variabiUty that stimulate collection response in bees. in the attractiveness of plants to bees. Nectar and Isolation and identification of these bee-attractive pollen availabiUty in plants can be accidentally compounds and the application of the attractant eliminated by breeding. When this occurs, there to areas requiring high bee populations for is a loss of a potential honey crop, but more impor- pollination should attract bees to the area. tant can be the loss of a seed or fruit crop because Synthetic chemical compounds, not necessarily the plant no longer attracts pollinators. A bee- attractive, can probably be used if the bees are keeper visiting Europe and seeing the large acre- properly trained. Bees associate certain smells ages of potatoes cannot help but wonder what with food sources and communicate the smell at would happen to the European bee industry if the food source to the colony. Isolated chemicals, a strain or variety of potatoes were developed that which may or may not have anything to do with would yield pollen and nectar for bees. pheromones produced by the bee or attractant compounds produced by plants, profoundly affect Contradictory Behavior of Foraging bee behavior. Some of these chemicals cause the Bees bees to fight or flee from the source, but others attract the bees. Probably the biggest problem in adequate pol- Of immediate concern is the need for a repel- lination that must be solved pertains to the for- lent that can be applied to a field to drive off aging activity of the bees. Individual bees usually all pollinators while a pesticide is applied. Then, confine their foraging area to a relatively small hopefully, when the toxic effect of the spray has block of a few square yards or to a single tree. On disappeared, the repellent will dissipate and the the other hand, the foraging area of a colony com- bees return to work. prises entire square miles—a circle with a radius Research should not be confined to chemicals approximating 2 miles—with the collective effect alone, but should be shared equally with various that foraging bees shift about, readily affected by physical factors that can possibly attract or the dance intensity of other returning foragers. repel bees. In other entomological fields the This contradiction could be resolved by breeding research on physical methods of controUing bees with greatly expanded individual foraging insects is receiving intensive investigation. Dif- areas but with a reduced radius of colony activity, ferent insects respond in differing ways, they possibly three-fourths mile. are attracted to certain Hght wavelengths and 36 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

repelled by others. Night-flying moths are A young, fully mated queen rarely lays drone repelled or go into defensive maneuvers because eggs before she is several months old. After that of bat sonar signals, whereas crickets and other time she seems capable of controlling the sex of members of their insect group can be collected the eggs by laying either fertihzed or nonfertiUzed by reproducing certain stridulations. eggs. The worker bees occasionally kill their queen. Particular Behavior Activities of Bees More frequently they will kill a newly introduced or virgin queen. To do this, 15 or 20 worker bees The Drone collect about her in a tight ball until she starves. The time of day that drones fly in search of a It has generally been thought that bees ''balled'' mate depends on many factors, such as the strange or introduced queens because they did geographic location, day length, and temperature. not have the proper ''colony odor.'' The reason Drones usually fly from the hive in large num- for balling is probably more complicated than bers between 11:00 a.m. and 4:30 p.m. Morning that, because bees will occasionally ball their or early afternoon flights last about 2 or 3 hours. own queen. Even if the ball is broken up, the Later flights are shorter. An individual drone queen seldom survives. If it is broken up or after flies for about 15 minutes before returning to the queen is dead, the stimulus is powerful the hive and may fly four or five times in one day. enough that the bees taking part in the queen When flying, drones seem to congregate in balling are sometimes subsequently balled ^'mating areas/' which may serve to attract themselves by other bees. virgin queens. These areas are usually less than 100 feet from the ground and seem to be associated with land terrain. References Some control of the time of drone flight can be obtained. The usual afternoon flight can be (1) BUTLER, C. G. prevented by placing the drone-containing col- 1955. THE WORLD OF THE HONEYBEE. 226 pp. ony in a cool, dark room. On the following day Macmillan Co., New York. the drones will fly several hours earlier than (2) VON FRISCH, K. 1955. THE DANCING BEES. 183 pp. Harcourt, normal. Brace & Co., New York. (3) HAYDAK, M. H. The Queen 1963. ACTIVITIES OF HONEY BEES. In Grout, The virgin queen becomes sexually mature R. A., The Hive and the Honey Bee. about 5 days after emergence. She is relatively Rev., 556 pp. Dadant & Sons, Hamilton, quiet in the morning and most active in the (4) LINDAUER, M. afternoon. She may begin her mating flights 5 1961. COMMUNICATION AMONG SOCIAL BEES. 143 or 6 days after emergence and may go on a num- pp. Harvard University Press, Cam- ber of them over several days. Mating with 8 to bridge, Mass. 12 drones will stock her spermatheca with 6 or (5) RIBBANDS, C. R. 7 million sperm. She will begin to lay in 2 to 5 1953. THE BEHAVIOUR AND SOCIAL LIFE OF HONEYBEES. 318 pp. Dover Publications, days and may continue for years. Inc., New York. BEEHIVE AND HONEY HANDLING EQUIPMENT

By CHARLES D. OWENS, agricultural engineer, Agricultural Engineering Research Division, Agricultural Research Service ^ Mechanization can save time and reduce lifting a similar colony exposed to the sun. Under partial of heavy hives and supers or other containers of shade the increase in production was proportional honey. To use labor-saving devices efficiently, cer- to the amount of shade. tain changes in methods of operation and different A portable shade (fig. 1) has been designed for use in types of equipment may be needed. areas needing shade and where permanent shades The hives should be easily accessible by truck cannot be used. The construction of this shade is giv- and arranged so that several can be manipulated en in detail in U. S. D ep artment of Agriculture Leaflet as a unit. Some type of hive hoist is needed so the 530, Shade and Water for the Honey Bee Colony. beekeeper can handle heavy colonies alone. After In cooler parts of the country a windbreak and the supers of honey are lifted from the hive onto good air drainage are more important than shade. the truck, they should be placed on easily movable skid boards for moving into the honey house. And Water finally, when the liquid honey is removed from the supers, other warehouse equipment is needed to Water is used by honey bees to dilute their food move or load it for shipment to the retail packing and to cool the hive. On a hot day a colony may plant. use as much as a gallon of water. Therefore, Today bees are moved frequently to take ad- hives should be located near a supply of water. vantage of nectar flow and to furnish pollination Under no condition should they be placed more services. In some areas they are moved several than a mile from water; and in warmer parts of times a year and over long distances. These moves the country they should be less than one-half require equipment for loading the hives. Shade and mile. The water should be as close as possible so water should always be considered when colonies the bees utilize a minimum of energy in trans- are to be relocated. If these are not readily avail- porting it to the hive. able, they can be furnished to bees by artificial Bees will usually collect water at the nearest means. satisfactory source; this may be a birdbath, swim- ming pool, pond, or stream. The source may be Artificial Shade on a neighbor's property, and therefore the bees become a nuisance to him. You can prevent this The importance of shade is determined by the by making water available nearby with some type prevailing local temperature. Bees benefit from of watering device. shade in all areas where the máximum summer The water can be furnished to the bees from a temperature normally exceeds 80° F. If the range barrel, tank, or directly from a pressure water is between 80° and 95°, partial shade all day or system into a w^atering device (fig. 2). A trough full shade in the afternoon is sufficient. If the filled with coarse gravel makes a good watering average maximum temperature exceeds 95°, bees device. A perforated tube in the bottom of the benefit from total shade all day. trough gives uniform water distribution. In southern California and southwestern Arizona, A coarse cellulose sponge floating in a pan or beekeepers build shades for the bees. These shades trough of water also makes a good w^aterer. The are usually 7 feet high, 12 feet wide, and 50 or pan should be 2 inches deeper than the thickness more feet long. A 50-foot shade will accommodate of the sponge. A float on the supply line, similar to 50 hives arranged in two rows back to back, with to the one in figure 2, will maintain the desired a 4-foot walkway between rows. Such shades are water level in the pan. built with the long dimension running east and Such watering units should be thoroughly cleaned west. The shade should extend down at least half- about every 2 weeks to reduce the possible spread way to the ground on each end to protect the end of bee diseases. In some areas the watering unit hives from morning and afternoon sun. will require protection from other animals. Tests made in Arizona show that a colony under solid shade produces 50 percent more honey than Moving Colonies

1 In cooperation with Arizona Agricultural Experiment Many mechanical devices can be used in load- Station. ing and unloading colonies. The choice of the 3Î 38 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

FIGURE 1.—Portable shade made of steel tubing and covered with shade cloth.

appropriate one will depend on the number of Some beekeepers have designed pallets the colonies to be moved and the frequency and width of the truck. These are pulled on or off distance of the moves. the truck by special drag chains. The least expensive lifting device is a crane Boom hoists are most satisfactory for loading or hand winch and boom attached to the rear and unloading hives (fig. 4). With them a hive corner of a truck. Many types are built for in- can be picked up and placed in position on the dustrial use. Some can even be converted to truck. Presently there are three companies manu- floor cranes for use in the shop or honey house. facturing these hoists for lifting hives. These cranes have about a 4-foot reach and their The hoist is mounted on the truck frame, length of reach is not changeable. Where they usually between the cab and the truck bed. Two are used, a hand cart or similar device is needed types of drives are used, electrical and hydraulic. to take the hive to them and to move it after An electrically powered hoist operates from the it is on the truck. With the lift, one man can truck batteries or from a gasoline-driven gener- place a hive on or off the truck. For the small ator. Today most hoists are powered by batteries. beekeeper with infrequent moves, this may be To keep the battery charged, the truck motor satisfactory. is kept running while the boom is being operated. A hydraulic tailgate will lift several hives at Battery-operated hoists are usually more depend- one time. Tailgates are made to fit all sizes of able than those operated by gasoline engines. trucks. The tailgate is useful for loading hives The hydraulic hoist operates like the tailgate, and supers of honey (fig. 3) and also as a platform but it has electrical valves to control the move- on which to stand when working tall hives. The ment of the boom. hydraulic system is powered by belt from the The hoists have boom lengths of 12 to 22 feet. truck engine or power takeoff from the trans- The simplest hoist only raises the load and the mission. operator moves it by hand. Others level auto- BEEKEEPING IN THE UNITED STATES 39 it must be hauled on a trailer behind the truck, and the combined length may exceed the State high- way permissible limit. Also in fields where bees are supplied for pollination purposes, there may be insufficient room to maneuver the tractor around the truck. Moving the Hive Many migratory beekeepers now use either forklifts or hoists. They use cleats on the ends of the hive, and they nail the bottom board under- neath. Hives are usually loaded at night, hauled to the new location, and unloaded before dawn. If the hives are loaded with open entrances, a plastic screen covering the entire load should be used. Inspecting Hives The hoist can also be used to lift ofl^ the supers for inspection of the colony brood nest. One com- mercial cart for hand moving hives is also designed to lift the supers and tip them back out of the way, so the brood nest can be inspected. There is also a stationary unit that clamps to the hive, then lifts and swings the supers aside so the brood nest can be inspected. Removing Honey Supers For the small beekeeper, the simplest way to remove bees from the honey super is to brush them off each comb. Another method of removing bees from supers is to use an inner cover with FiGUEB 2.—Four-inch eaves trough filled with coarse a bee escape. The escape board, as it is called, gravel and equipped with float water-level control. is placed between the supers and the brood nest the day before the supers are to be removed. All other openings into the supers must be closed so matically and can move the load along the boom. that the bees cannot return. This method works The choice depends largely on how much it will best in cool weather when the bees move at be used. A good hoist should support a 300-pound night from the to the brood nest. load. A recent innovation for removing bees from A fork on the end of the boom is used to pick up supers of honey is to blow them out. A large the hive. The following three methods are employed: vacuum cleaner with a clevis tool attachment Fork prongs are inserted under the bottom board, works satisfactorily. The larger household tank- fingers are dropped into the hand holes, or fingers type vacuum cleaner can be used if the dust bag are inserted under a iK-inch cleat nailed on each is removed. A velocity of about 18,000 feet per end of the hive. The cleats are usually the most minute is required. There is now an engine-driven satisfactory. blower built specially for removing bees. This On special order a hoist can be obtained that method does not irritate the bees and it is will lift a pallet bearing two or four hives with effective. a total maximum weight of 1,000 pounds. A short truck and trailer can be used if the hoist is placed on the rear of the truck where it can service Handling Supers both. Hives on pallets can also be loaded by a Supers of honey removed from the hive are usu- forklift mounted on a tractor. The tractor can ally placed on dripboards or on pallets on the also haul hives to places difficult to reach by truck. The dripboard is the size of the super and truck. If the tractor is equipped with a blade, it it has a metal sheet or pan to catch dripping honey. can level the area before the hives are unloaded. It is made to fit a handtruck for moving a stack of The disadvantage of a tractor forklift is that supers (fig. 5). The pallet holds two or more stacks 40 AGRICULTUREi HANDBOOK 3 3 5, U.S. DEPT. OP AGRICUUTURE

BN-22119 FiGUEE 3.—Loading supers on truck with hydraulic tailgate. and is moved with a hand- or power-operated pal- wholesale container, and small beekeepers still use let truck. The supers are moved into the honey it. Although single cans may be handled by hand, house in these stacks. Pallets with casters are some- they are more easily handled by pallet truck. times used in the honey house, but they are usually Larger quantities of honey are more easily handled not satisfactory. Generally handling the single in 55-gallon drums. The drums are more durable stack on a dripboard is most satisfactory. than cans and are reusable. Several industrial hand The handling of individual supers should be re- and power trucks are available for handling drums. duced as much as possible from the time they are If drums are to be stacked, a motor-driven lift removed from the hive until they are returned. truck is needed. All commercial bottlers of honey are equipped to handle both cans and drums. Handling Honey Some beekeepers have built large tanks for stor- age of 1,000 gallons or more of honey, which is then Honey is usually sold wholesale to packers in pumped into tanks on the trucks for transfer to 5-gallon cans, drums, or in bulk. For many years bottlers. One company uses air pressure to transfer the 5-gallon (60 pound) can was the principal honey into and out of the truck tanks. BEEKEEPING IN THE UNITED STATES 41

BN-22120 FIGURE 4.—Loading hives with electrically powered boom loader.

A-

FIGURE 5.—A two-wheeled handtruck, a versatile piece of equipment around honey house, is being used to move empty supers. HONEY PROCESSING, PACKING, AND DISTRIBUTION

By B. F. DETROY, agricultural engineer, Agricultural Engineering Research Division, Agricultural Research Service ^

Honey is at its peak quality when properly cured Two-story structures may have both floors and sealed in the comb by the honey bee. When it above ground or may have one floor above is converted from this state by man to suit his ground and a full basement. If both floors are particular need, deterioration begins. The extent above ground, it is necessary to provide a ramp of deterioration depends on the processing methods to the second floor so that it is accessible to trucks used between the time the honey is extracted from or to install elevators to move equipment and the comb and its use by the consumer. It is the material from one floor to the other. A hillside responsibility of the industry to provide a top building site will provide access to both floors quality product to the consumer if acceptance without ramps and both floors are at least par- is to be expected. tially above ground (fig. 1). The main advantage Since most honey harvested is extracted from of the two-story building is that the supers of the combs in an extracting plant, the beekeeper honey can be unloaded at the upper level where should equip this plant so that the operations can the extracting plant is located. Extracted honey be done in the most efficient possible manner to can then flow by gravity to storage or further provide a high quality product for market. processing on the lower floor.

Honey House Space Requirements The extracting plant is generally located in a Careful planning prior to building a honey honey house. The honey house is the center of ac- house may save costly additions later. The oper- tivities for the beekeeper, represents a goodly por- ations necessary in extracting honey and the se- tion of his investment, and may contribute greatly quence in which they are performed should be to the overall efficiency of the entire operation. considered in detail for filling the needs of the The honey house may contain various other beekeeper. Ample space should be provided for facilities in addition to the extracting plant, such all extracting and processing equipment. The as storage space for hive equipment and honey, equipment chosen should be expected to oper- workshops, office space, and possibly a packing or ate at near-rated capacity and should be compactly salesroom or both. The building should be de- arranged so that the material flows smoothly from signed for the work to be done in it and it should be operation to operation with a minimum of move- properly equipped. Efficient arrangement, clean- ment by the operators from area to area. liness, and ample space are of prime consideration Storage space for full honey supers should be in planning the honey house. figured on the basis of the maximum nuraber anticipated in the honey house at any one time. Types of Honey Houses Uniform stack height of full supers should be used Honey houses may be one- or two-story struc- throughout the operation for efficient handling. tures. The one-story structure is probably the Warm storage areas for full honey supers aid in most common and is used by both large and small the extracting operation. An area for storage of beekeepers. The building construction is more liquid extracted honey must also be provided. simple and the choice of a building site is not Required space should be figured on the basis of limited by terrain. Honey handling cannot be the type of bulk containers used and the height readily adapted to gravity flow, but proper use they are to be stacked. of honey pumps can overcome this disadvantage. Other areas to be included in the building will Small beekeepers may use this type of building to vary in type and size according to the individual particular advantage. Since all equipment can be beekeeper and the operations he desires to per- compactly arranged on one level, it is easier to form. They may include a room for rendering closely regulate all operations. cappings, space for shop facilities, equipment- ^ In cooperation with Wisconsin Agricultural Experiment assembling area, truck storage, small packing Station. plant, salesroom, and office space. 42 BEEKEEPING IN THE UNITED STATES 43

BN-30037 FIGURE 1.—Apiculture laboratory and honey house at University of Wisconsin. Noto that lower floor is partially below ground and that entrances are at both floor levels.

Special Features facilities in another building, and if so, the build- Kegardless of type or size, the building should ings should be separated by at least 20 feet. be bee tight and should provide a means of un- If handtrucks, motorized" lifts, and fork or barrel loading filled honey supers in a closed-in area. A trucks are to be used, doorways should be large sunken driveway or raised platform to permit enough to permit their free movement to all parts loading or unloading with the truck bed at floor of the building. level will prove to be a great labor and time saver. The floors, walls, and ceiling should be made of materials that can be easily cleaned and main- Honey supers should be removed from the hive tained. Walls and ceilings should be a light color as soon as the honey is sealed. Extraction soon and should be of a material or paint that can be after removal may prevent crystallization in the washed often. Floors should be of concrete where comb. It is often possible to reuse the super on the possible. If hardwood floors are necessary, they colony before the flow ceases. Equipment in the should be covered with ceramic tile. Any floor extracting area should be arranged so that the used must withstand heavy loads and be free of material flows smoothly through the various oper- vibration. A smooth surface with ample drain facilities will make cleaning easier. ations with a minimum of interruption and with as little physical effort as possible. Illumination and ventilation should be carefully Extracting equipment will differ in almost every considered in planning the building. In areas where close operator attention is required, it is honey house. Choice of equipment is dependent on the size of operation, physical properties of the highly important to provide adequate illumina- tion, 50 to 75 foot-candles. The entire building honey, availability of labor supply, and the per- should have adequate lighting. Windows to pro- sonal selection of the individual. In many cases vide both light and ventilation should be strate- the beekeeper has designed and built his own gically located. Fans may be installed to reduce equipment or remodeled commercial equipment odors and lower humidity. All windows must be to meet some particular need. screened and provided with bee escapes. Rooms in which fires might originate, such as Care and Storage of Supers the furnace room or wax-rendering room, should Supers are most easily handled if they are kept be buut of fire-resistant material or lined with in uniform stack heights from the time the full asbestos board. It may be desirable to have such supers are removed from the colony until the emp- 44 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE ties are returned. Stack boards or skid boards Extractors are commonly used. Skid boards placed on the truck bed receive the supers as they are removed Two types of extractors are in use in this from the colonies and are moved through the country today—the reversible basket and the entire extracting process and back onto the truck radial. Both use centrifugal force to remove the with an ordinary warehouse truck. honey from the comb. More expensive types of lift trucks are some- Reversible-basket extractors range in size from times used that can handle skids containing two 2 to 16 frames per load. Honey is extracted by or four stacks of supers. Castered dollies may also applying centrifugal force to first one side of the be used, but these are not easily hauled to and from comb then the other. The comb is reversed three the yards on trucks. or four times—turned 180°—^during the extracting Hot rooms are sometimes used for storage of cycle. On some of the small extractors the frames filled supers prior to extraction, especially in are reversed by hand with the machine stopped, regions where cool temperature or high humidity whereas on others the frames are reversed by is common. If the only requirement is to keep the use of a brake while the machine is running. honey warm to facilitate extraction, the room Extracting time ranges from 2 to 4 minutes at should be kept at 75° to 100° F. and should have constant speed. a circulating fan. Warm, dry air may also be used Radial extractors range in size from 12 to 50 to remove moisture from the honey, in which frames per load. Both sides of the comb are case slatted skid boards should be used or the extracted simultaneously as the combs are rotated, supers stacked crisscross. The air should be in- the centrifugal force acting radially across the troduced at floor level and drawn up through the face of the comb. The extracting cycle is started stacks. Moisture-laden air should be discharged at 150 r.p.m. and is gradually increased during from the room. Dehumidifiers may also be used the cycle to 300 r.p.m. The time required to extract to speed the lowering of the moisture in the honey. a load of combs will vary from 12 to 20 minutes depending on the temperature and density of Supers stored for a long time should be fumi- the honey. gated to prevent damage by the larvae of the wax Special extractors have been built and used moth. that are larger than those described and that extract the combs in the supers, special boxes, Uncapping Devices or baskets. None of these are available com- mercially. To extract honey it is first necessary to remove Automatic electric controls have been devel- the capping from the comb cells. A wide range of oped for both the radial and reversible-basket equipment is available for uncapping combs, from extractors. These controls change the r.p.m., unheated hand knives to elaborate mechanical reverse the baskets, and shut off the motor when machines. the cycle is completed. Mechanical controls are A cold knife can be used to uncap warm combs also available that automatically increase the or it can be heated by placing it in hot water. speed of the radial extractor during the extracting It is most commonly used by the hobbyist bee- cycle. keeper who has only a few hives. The steam and electrically heated hand knife and hand plane Care of Cappings are probably the most widely used uncapping devices in this country today. In a more refined Cappings and honey removed from the combs version, the knife is mounted in a frame on spring in the uncapping operation must be separated to steel mounts and vibrated by an electric motor. salvage the honey and wax. Caution must be taken This type, referred to as the jiggler knife, may be in recovering the honey to prevent impairing the mounted in a vertical, horizontal, or inclined flavor, color, and aroma. The following methods position. The knife vibrates in the direction of are used: its length. (1) Draining hy Gravity.—The cappings are accumulated in screened or perforated containers Machines that carry the combs of honey and allowed to drain, usually for at least 24 hours through an uncapping device after being fed in a warm room. Stirring and breaking up the into the machine by hand are available com- cappings facilitate draining. mercially and their use is becoming more common (2) Centrijuging.—The cappings are placed in (fig. 2). The uncapping devices on these machines a specially constructed centrifugal drier or in may be vibrating knives, rotating knives, flails wire baskets that fit into a radial extractor. Honey of various kinds, or perforating rollers. is removed from the cappings by centrifugal force A few machines have been built by individuals as the cappings rotate. that uncap the frames in the super; however, (3) Pressing.—A basket-type perforated con- none of these machines are available commercially. tainer is used to catch the cappings where some BEEKEEPING IN THE UNITED STATES 45

zi IHB WllliMlllii ¡Mfriiiii / ? •'^sL

FIGURE 2.—Semiautomatic, perforating-roller uncapping machine removes frames from super, uncaps combs, and deposits frames in extracting tubs. gravity draining takes place prior to pressing. maintained between the honey level and the heat Usually the container is placed directly vmder the source. Liquid wax accumulates at the top of the press ram and jjressure applied to squeeze the tank and is discharged into solidifying containers. honey from the cappings. Heat may be supplied by steam coils, electric Honey removed from the cappings by any of heaters, heat lamps, or radiant gas heaters. these methods is undamaged. Usually the remain- Various models of this type of separator are ing cappings will contain as much as 50 percent marketed. If they are properly operated, the honey honey by weight, which may be recovered when obtained can be added to the remainder of the the cappings are melted. This honey is generally crop without damage to grade, color, or flavor. damaged by overheating and should be handled (5) Centrifugal Separator.—Recent develop- separately. ment of a centrifugal separator (fig. 3) that auto- (4) Flotation and Melting.—The cappings mel- matically separates the honey and dries the cap- ter is widely used to separate honey and cappings. pings has greatly advanced the use of mechanical The cappings and honey enter the melter tank uncappers. Usually all honey and cappings from near the bottom and are separated by gravity. the uncapper and extractor are run through the Separation is facilitated by heat that softens the machine. Large pieces of cappings should be broken cappings and increases the fluidity of the honey. up to assure proper feeding into the separator. The cappings being less dense rise to the top where they are melted. The honey level is con- Processing trolled by an adjustable height overflow enclosed by a baffle to prevent the entry of wax. A layer Processing the honey crop beyond the extraction of cappings in various stages of liquefaction is stage may be done by the producer, the packer, or 261-90i7O—67 4 46 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

FIGURE 3.—Centrifugal separator that separates honey from cappings. both. Regardless of where these operations take The Sump and Pump place, they are necessary to provide the consumer with a high quality product. It is important, how- Honey from the uncapping and extracting opera- ever, that the heating be controlled, since the tion usually flows into a sump. The sump is a tank, flavor, color, and aroma of honey can be seriously usually water jacketed, that collects honey from impaired by excessive temperature over a given the extracting process so that it can be delivered period of time. for further processing at a uniform rate. The sump BEEKEEPING IN THE UNITED STATES 47

may contain a series of baffles or screens or both over long periods of time as well as high tempera- for removing coarse wax particles and other foreign tures for short periods of time. material. A honey pump is generally used in con- Several methods of heating are used successfully. junction with the sump; however, in some systems Shallow pans with inclined surfaces heated by gravity flow can be used and the pump eliminated. water jackets are commonly used. As the honey Gear pumps or vane pumps are commonly used. flows into the pan, it should be distributed over In cases where the centrifugal separator is used and the surface by suitable baffles. Jacketed tanks may it is necessary to pump large quantities of cap- be used for heating, in which case the honey should pings, some other type of pump may be required. be slowly but continuously agitated to insure Pumps used in a continuous now system should be uniform heating throughout the tank. Heat ex- supplied with honey in sufficient quantity to allow changers in which the honey is pumped quickly uninterrupted operation. To prevent introduction through a passage contained in hot water are used of air into the honey, the pump should run at low very successfully as flash heaters. speed and the level of honey in the sump kept well One design of heat exchanger is shown in figure above the pump intake. Automatic pump controls, 5. This exchanger consists of three concentric tubes either float type or electric liquid level control in which the honey is pumped through a ^{e-inch- type, can be used to eliminate continual operator thick annular space between two layers of flowing supervision. hot water. Honey enters the exchanger at the bottom and is pumped in a direction opposite to Strainers the flow of water. These units may be connected in series to provide the desired amount of heating. After the bulk of the wax has been removed from Precautions for cooling honey after heatirg are the honey by the sump tank, coarse straining, or seldom practiced to a suitable degree. Immediate centrifugal separators, it is necessary to remove cooHng following flash heating is essential to pre- very fine material. Settling of honey may prove vent honey damage. Equipment similiar to that satisfactory for some processors. The honey is first used for heating can be used effectively for cooling screened in a sump and then pumped into settling by using cold water instead of hot. Heat exchang- tanks at a temperature of at least 100° F. Suffi- ers are particularly effective, but may cause ex- cient time should be allowed to permit the required cessive line pressure as the honey becomes more separation. viscous upon cooling. To be certain that all honey packed will meet the desired grade requirements, it is necessary to Storage of Honey use some type of strainer. Many types and sizes are used and the straining media may be metal Honey in bulk containers, 60-pound cans, or screen, crushed granite, silica sand, or cloth. Re- 55-gallon drums should be stored in a dry place at gardless of the material used, the mesh must be as near 70° F. as possible. Long periods of storage fine enough to produce the desired result. Cloth above 70° wiU damage the honey the same as has the advantage of being easily cleaned ; further- excessive heating. Storage of unheated honey at more, since the initial cost is low, a cloth may be 50° to 70° is indu ci ve to granulation and fermen- used only once and discarded. tation. This also holds true for honey packed in Honey may be moved through the strainer by bottles and other small containers. These should be pressure (pumping) or by gravity flow. When cloth stored in shipping cases to protect them from light. strainers are used in a pressure system (fig. 4), a Most deterioration in honey during storage can pressure switch should be installed in the honey be prevented by maintaining storage temper- line to prevent excessive pressure that could rup- atures below 50°. Honey stored at freezer tem- ture the strainer cloth. peratures, 0° to -10°, for years cannot be dis- Heating the honey to 115° will greatly facihtate tinguished from fresh extracted honey in color, the straining process. This increases the fluidity flavor, or aroma. of the honey without softening the wax particles appreciably. Higher temperatures will soften the Producer Marketing wax so that it may be forced into or through the The producer has a choice of methods in disposing straining media. of his honey crop. He may sell his entire crop in bulk containers to a packer or dealer or he may Heating and Cooling pack a part or all of his crop and sell direct to Heat properly applied can be a great aid in retail stores or the consumer or both. The pro- handling honey. Heat also dissolves coarse crystals ducer may be a member of a cooperative through and destroys yeasts, and thus prevents fermenta- which his honey is processed and sold. tion and retards granulation. Heat may also seriously damage the color, flavor, and aroma of Wholesale Marketing honey unless particular precautions are taken. The producer who markets his honey in bulk Damage may result from a small amount of heat should keep in mind the market to be supplied 48 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

FIGURE 4.—Cloth strainer showing containing shell, supporting framework with strainer cloth, gasket, and cover plate. when he chooses the type of container to use. packer who knows exactly what he buys can Generally these will be either the 60-pound can readily process and blend to meet his particular or the 55-gallon drum. A limited quantity of standards without concern for discrepancies or honey is moved from the producer to the packing variation. plant in tank trailers. Careful sampling is neces- Approximately 50 percent of the honey produced sary when the honey is extracted. Representative in the United States is marketed by the producer samples should be taken from each tank, each in bulk. yard, or each day's run and should be carefully marked on both the sample and the containers Producer-Packers for accurate identification. Honey producers who bottle and sell part or all Honey is generally sold at wholesale prices on of their honey crop are referred to as producer- the basis of samples, and accurate sampling will packers. Almost half the honey produced in the result in building confidence, understanding, and United States is marketed in this manner. satisfaction for both producer and buyer. The The producer-packer receives a higher price per producer who knows exactly what he has for pound for his honey; however, he may have many sale can demand and get top market prices. The additional costs. Processing equipment that will BEEKEEPING IN THE UNITED STATES 49

FIGURE 5.—Concentric tube heat exchangers connected in series and used for flash heating honey. (Note strainers ahead of flash heaters.) 50 AGRICULTURE HANDBOOK 33 5, U.S. DEFT. OF AGRICULTURE yield a product meeting the desired grade stand- Much honey is sold in bulk for industrial con- ards must be utilized. His honey must compete sumption, such as for the baking industry, restau- with other brands of honey and other foods rant trade, honey candies, and honey butter. backed by aggressive sales and promotion pro- Many other industries use honey in varying quan- grams. He may employ a broker to move his tities. honey into retail channels. Many producer-packers confine their sales to Liquid Honey salesrooms in their homes or honey houses, road- side stands, door-to-door sales, or local stores. Liquid honey is packed in glass, tin, plastic, and Some have established regular sales routes to paper containers. Glass is the most popular and supply retailers over a wide area, and these is used in a wide variety of shapes and sizes. routes are serviced at regular intervals. Plastic containers in various shapes are becoming more ^ and more popular. The 12-ounce plastic Cooperative Marketing container makes a very satisfactory table dis- penser. Special glass and plastic containers are There are several cooperative marketing organi- used effectively in novelty and gift packs and are zations in the United States. These organizations popular on the retail market. may buy the meniber producer's crop and process, Bottled honey should be free of air bubbles or pack, and distribute the products under the any foreign particles and the containers must be cooperative label. Other organizations may only spotlessly clean. Honey bottled by floral source pool and market the member's production in should be clearly labeled as such to insure customer bulk containers. satisfaction. Generally the cooperative may operate as Honey selected for bottling should be from floral follows: Member producers are furnished with sources that granulate slowly. Proper heating in bulk containers. When the crop is harvested, the the processing and bottling operation will also honey is put in these containers and shipped or help retard granulation. Any bottled honey in a trucked to the cooperative by the producer. The sales display that shows signs of granulation honey is then graded and the producer is paid a should be replaced immediately. part of the total price. The cooperative then processes, packs, and sells the honey through its Granulated or Creamed Honey sales organization. At a later date the producer is The popularity of granulated or creamed honey paid the remainder of his selling price. is increasing in the United States. This honey is Cooperative marketing offers many advantages, presently available in many retail food stores. It but there are also some disadvantages just as in is packed in various paper, plastic, and glass other types of marketing. The producer must containers. The desired consistency of creamed decide which method of marketing is the most honey is soft and smooth to allow easy spreading advantageous to him and market his crop ac- at room temperature. cordingly. Honey used for this purpose should be from a floral source that granulates rapidly into a product Packing and Distribution of soft, smooth, fine, creamy consistency. Honeys that granulate slowly may be used by adding Honey packed for market must be of high about 10 percent of finely ground crystallized quality, neatly packaged in clean, attractive honey. To encourage granulation the honey containers, and attractively labeled. Every cau- should be refrigerated immediately after the fine tion should be taken in processing and packing to honey crystals have been added to prevent any insure a product of quality as near as possible to air bubbles rising to the surface. After rapid that sealed in the cell by the bee. All honey cooling, the honey should be stored at 55° to 57° F. packed under a given label should be as uniform and that temperature maintained until the honey as possible to assure consumer satisfaction. An is completely crystallized. Cool storage is desirable. attractive, eye-catching display in a prominent lo- Creamed honey will remain firm at room tem- cation is desirable. perature, but will break down if subjected to Most large honey packers have automatic high temperature or high humidity. Once it has labeling, filling, and capping equipment. Their softened or partially liquefied, recooling will not honey is distributed and sold under their ad- make it firm again. vertised brand, usually in a limited area. Few, if any, have nationwide distribution. Some have Comb Honey sales personnel, whereas others employ food Comb honey is marketed in the form of section brokers or other sales agencies to market their comb, cut-comb, and chunk. All forms require product. Many use warehousing facilities in special care and handling, and when properly pre- areas of concentrated retail outlets. pared they have excellent consumer appeal. BEEKEEPING IN THE UNITED STATES 51

Section comb honey is produced in a special usually glass jars. The containers are then filled super. When removed, the sections are carefully with liquid honey that has been heated to retard scraped with a suitable instrument to remove the granulation. The liquid honey should be run down propolis. The sections are then sorted, graded, and the inside of the container to prevent introduction placed in window cartons or wrapped in cellophane. of air bubbles and should be at a temperature of Some packers seal the sections in clear plastic 120° F. The containers should be capped and laid bags before placing them in window cartons. on their sides immediately after filling to prevent Sections that do not meet the required grade damage to the comb because of its buoyancy. standards should not be marketed. Special widemouthed jars are used for packing Cut-comb honey is produced in shallow supers chunk honey. The chunk should be as wide as on foundation similar to that used for sections, possible and still slip readily into the jar, and the but in frames instead of sections. The comb honey length should extend from the top to the bottom is cut from the frames into the desired size for of the container. marketing. Sizes of cut-comb honey vary from a Probably the greatest deterrent to packing 2-ounce individual serving to large pieces weighing chunk honey is the tendency of the liquid honey nearly a pound. The cut edges of the comb must to granulate. be drained or dried in a special centrifugal drier, so that no liquid honey remains. The pieces are References either wrapped in cellophane or heat sealed in (1) GROUT, R. A., ed. polyethylene bags and packaged in containers of 1963. THE HIVE AND THE HONEY BEE. ReV., 556 various styles. pp. Dadant & Sons, Hamilton, 111. One of the most attractive and appealing packs (2) OWENS, C. D., and DETROY, B. F. of honey is the chunk honey pack. It consists of 1965. SELECTING AND OPERATING BEEKEEPING EQUIPMENT. U.S. Dept. Agr. Farmers' a chunk of comb honey in a glass container Bui. 2204, 24 pp. surrounded with liquid honey. (3) TOWNSEND, G. F. When packing chunk honey, the pieces of well- 1961. PREPARATION OF HONEY FOR MARKET. drained cut-comb are placed in the container, Ontario Dept. Agr. Pub. 544, 24 pp. HONEY BEE NUTRITION

By L. N. STANDIFER, apiculturist, Entomology Research Division, Agricultural Research Service

A general knowledge of honey bee nutrition aids of worker bees. Workers probably use the secretion in understanding how the individual bee grows from their glands to prepare and manipulate and how the colony develops and maintains itself. wax for building comb. It may also be used to The anatomical and vital physiological systems soften the pupal cocoons of bees and is found in usually associated with living animals are present royal jelly. in the honey bee. Food enters the alimentary canal The primary function of the postcerebral (fig. 1) of the adult bee by way of the mouth and glands, located in the back of the head, is to the long tubelike esophagus, which extends provide the enzymes necessary for the digestion through the thorax and into the abdomen, where of foodstuff consumed by the bee. it enlarges to form the honey stomach. Nectar is The thoracic or salivary glands in the anterior transported in the honey stomach from the flower part of the thorax secrete a carbohydrate-splitting to the hive. Immediately behind the honey stom- enzyme, invertase, in large amounts. ach is the proventricular valve. It retains the The function of the postgenal glands in the nectar load in the honey stomach, lets food pass lower inner wall of the head and the sublingual into the midgut, but prevents food from returning. glands at the base of the bee tongue is unknown. The midgut or ventriculus is a relatively large Two pairs of rectal glands or pellets on the sides segment of the alimentary canal, lined on the of the rectum are associated with fat absorption. inside with the peritrophic membrane. Beyond the Beeswax is secreted by specialized cells called midgut are the short, small intestine, the large wax glands on the underside of the abdomen of intestine or rectum, and finally the anus. the worker bee. Generally wax glands become The adult honey bee has six sets of paired glands fully developed about the 15th day of adult life. located in the head and thorax. The labial glands Secreted beeswax appears as thin, delicate scales are generally believed to be associated with the or flakes. It is a byproduct of metabolism and alimentary canal. They dehver their secretions at directly follows the digestion of large quantities the base of the labrum. Their function is dependent of nectar or other sugars. The bees use wax to form on the age of the bee and on the work in which the cells of the comb, in which food is stored and it is engaged. brood is reared. The hypopharyngeal or brood food glands pro- When wax is needed for comb building, the 16- duce the food called royal jelly. Most of the larval to 24-day-old bees fill their honey stomachs, then food comes from these glands. They also supply hang together in vertical sheets or festoons. The the food for the adult queen and possibly adult wax scales are secreted, then removed with the drones. They are fully developed only in the hindlegs, and passed forward to the mouth, where worker bee. Royal jelly is milky in appearance, they are worked by the mandibles and subse- slightly acid, and rich in digestive enzymes, quently applied to the edge of the comb. The proteins, carbohydrates, fats, and vitamins. wax glands shrink and become nonfunctional be- The mandibular glands are saclike, single tween the 20th and 25th day of adult life, or about structures located immediately above the man- the time worker bees become field-foraging bees. dibles. They are extremely large in the queen, smaller in the worker, and vestigial in the drone. They do not vary in size with age or occupation. Digestion However, if the newly emerged bee does not The movement of pollen through the alimentary consume adequate protein during the first few canal of the adult bee reveals something of the di- days of adult life, these glands do not develop fully. gestive process. Ten minutes after the material is fed to the bee, the pollen grains are clustered at The secretions from these glands in queens the proventriculus. Thirty minutes after feeding, contain a compound called queen substance, they are within the peritrophic membrane in the which is essential for social unity of the colony. forepart of the ventriculus. Ninety minutes after This substance is not in the glandular secretions feeding, the peritrophic membrane-enwrapped pollen mass enters the anterior or small intestine. iln cooperation with Arizona Agricultural Experiment At the end of 2 hours the pollen is within the btation. small intestine or just entering the large intestine 52 BEEKEEPING IN THE UNITED STATES 53

BROOD FOOD GLANDS

SALIVARY GLANDS SALIVARY GLANDS ESOPHAGUS

MALPIGHIAN TUBULES

SMALL m HONEY STOMACH INTESTINE

STOMACH OR STOMACH OR VENTRICULUS VENTRICULUS

LARGE INTESTIN OR RECTUM

FIGURE 1.—Alimentary canal of adult worker honey bee. or rectum. The peritrophic membrane that encir- the midgut of the adult worker and drone. Its cles the pollen grains in the midgut persists in the value in digesting the lipids is unknown. In higher rectum for a considerable time before all is voided animals lipids are digested by lipase or esterase in the feces. into free fatty acids and glycerol. The fatty acids in pollen are niade water soluble by neutralization with alkalies in the alimentary Nutritional Requirements canal secretion. The proteins are broken down into peptides, and these are further hydrolyzed into Honey bees and other insects have no unusual amino acids. . nutritional requirements. They require carbohy- The lipids of bee food occur chiefly in pollen. drates, proteins, fats, minerals, vitamins, and The lipid-splitting enzyme Upase is abundant in water for growth, development, maintenance, and 54 AGRICULTURE HANDBOOK 33 5, U.S. DEFT. OF AGRICULTURE reproduction. Nectar and honeydew are the chief Plant Juices sources of supply for carbohydrates in the diet of bees, and pollen furnishes all the other indispen- Bees often collect juices from overripe fruit sable constituents. and various plant exudates that are rich in sucrose Adult bees can live on the carbohydrates glu- or related sugars. They usually do this only when cose, fructose, sucrose, trehalose, maltose, and me- nectar is not available. lezitose. They cannot utilize the carbohydrates galactose, mannose, lactose, raffinose, dextrin, in- Pollen sulin, rhamnose, xylose, or arabinose. Beekeepers Pollen furnishes all the other indispensable often feed sucrose if a shortage of nectar or honey constituents of the diet, except water, that are exists. Bees also utilize fruit juices and certain required for vital activity, including rearing occasional plant juices. A small amount of carbo- young bees. Not all pollens are alike nutritionally, hydrates is also obtained from pollen. The adult and bees grow and develop better on some than bee can survive on carbohydrates; however, pro- on others. teins, fats, minerals, and vitamins are necessary in In nature bees generally utilize a mixture of pol- rearing the immature stages. lens in their diet. This is eaten by adult bees and is Proteins of a precise quaUty and definite amino fed to worker and drone larvae after they are 3 acid composition are required for optimum growth days old. Consumption and digestion of pollen by and development of the brood food-producing adult bees are essential, since they can only pro- hypopharyngeal glands and no doubt others. duce brood food from pollen that they have eaten. When nursing duties are finished (between lOth This brood food, or royal jelly, is fed to all larvae and 14th day of adult life) and field duties are the first 3 days of life and to the queen bee through- undertaken, the requirements for protein decrease, out her larval and adult life. Royal jelly has the and the chief dietary constituent becomes the following approximate chemical composition (per- carbohydrates obtained from nectars and honey. cent) : Water 66, dry matter 34; of the latter, car- Fats, like carbohydrates, are also used as sources bohydrate 13, protein 12, fat 5, ash 1, and undeter- of energy. Bees probably require and utilize mined matter including vitamins, enzymes, and some of the fats in pollen they consume. However, coenzymes 3. chemical analyses of feces show that large amounts Bee-collected pollens are comparatively rich in of fats in the pollen consumed by bees pass the carbohydrates. Reducing sugars range from 15 through the digestive tract and are not utilized. to 43 percent, with an average of about 29 percent. Water is necessary in the diet for diluting con- The glucose, fructose, sucrose, raffinose, and stach- centrated honey. It is also used in air-conditioning yose content is not significant, although the bees the cluster. Normally bees do not store water apparently utilize those that are available. Corn as they do nectar and pollen. It is collected only pollen, for example, has a high starch content. The when needed. pollen shell is not utilized by bees, but is eliminated The nutritional value of the enzymes, coenzymes, with the feces after the internal matter has been and pigments found in pollen is largely unknown. removed by digestive processes. The protein value of pollen varies from 10 to 36 percent. The amino acid content of average pollen Sources and Chemical Composition and sweet corn pollen with a crude protein of 26.3 of Foods and 26.9 percent, respectively, is shown in table 1. Nectar TABLE 1.—Amino acid content of average pollen and sweet corn pollen, expressed as percent of crude When nectar is collected, it may contain from protein 5 to 75 percent soluble solids (sugars), most of which is in the 25 to 40 percent range; the remain- Component Average Sweet corn der is water. The primary sugars are sucrose, pollen pollen glucose, and fructose. As the nectar is ma- nipulated and finally stored as honey, much of Percent Percent the sucrose is inverted to glucose and fructose, Arginine 5.3 4. 7 usually in about equal amounts. Histidine 2.5 1.5 Isoleucine 5. 1 4.7 Leucine 7. 1 5.6 Honeydew Lysine 6. 4 5.7 Various species of insects secrete a material Methionine__. 1.9 1. 7 Phenylalanine 4. 1 3. 5 called honeydew, which bees collect and store in Threonine 4. 1 4.6 the comb. Honeydew has a high percentage of Tryptophane. 1.4 1.6 dextrins and melezitose. It is generally considered Valine 5.8 6. 0 a poor source of carbohydrates for bees. BEEKEEPING IN THE UNITED STATES 55 All the amino acids in table 1, except threonine, Pollen substitute (dry mix) : are essential for normal growth of the young Soybean flour 20 adult bee. With the exception of histidine and Casein 30 perhaps arginine, they cannot be synthesized by Brewer's yeast 20 Dry skim milk 20 bees and must be obtained from the pollens Dried egg yolk 10 consumed. Other constituents of pollen are as follows: Few problems facing the apiculture industry Amount today require immediate research attention as Constituent Percent much as the development of an artificial or chem- Fats 1. 3-19. 7 Minerals (ash) : ically defined diet for honey bees as a substitute Calcium 1. 0-15. 0 food for pollen. Work on nutrition and physiology Chlorine . 6- 9 of the honey bee may soon lead to an artificial diet Copper . 05-. 08 for bees. Iron . 01-12. 0 Magnesium 1. 0-12. 0 Phosphorus . 6-21. 6 Potassium 20. 0-45. 0 Silicon 2. 0-10. 4 References Sulfur . 8-1. 6 (1) BERTHOLF, L. H. MicTograms 1927. THE UTILIZATION OF CARBOHYDRATES AS per gram Vitamins : identified FOOD BY HONEYBEE LARVAE. Jour. Agr. Res. 35: 429-451. Ascorbic acid 131. 0-721. 0 Biotin . 19-. 73 (2) BUTLER, C. G. D .2-. 6 1949. THE HONEY BEE'. AN INTRODUCTION TO HER SENSE-PHYSIOLOGY AND BEHAVIOUR. 139 E 0-. 32 pp. Oxford University Press, London. Folie acid 3. 4-6. 8 Inositol . 3-31. 3 (3) LUNDEN, R. 1956. LITERATURE ON POLLEN CHEMISTRY. Grana Nicotinic acid 37. 4-107. 7 Pantothenic acid 3. 8-28. 7 Palynologica 1 (2), 17 pp. (4) NELSON, J. A., STURTEVANT, A. P., and LINEBURG, B. Pyridoxine 2. 8-9. 7 1924. GROWTH AND FEEDING OF HONEYBEE LAR- Riboflavin 4. 7-17. 1 VAE. U.S. Dept. Agr. Dept. Bui. 1222, Thiamine 1. 1-11. 6 38 pp. Bees also occasionally collect spores and store (5) PHILLIPS, E. F. them as pollen. Although spores can be utilized 1927. THE UTILIZATION OF CARBOHYDRATES BY as a proteinaceous food, they do not stimulate HONEY BEES. Jour. Agr. Res. 35: 385-428. brood rearing and are generally considered a (6) RIBBANDS, C. R. 1953. THE BEHAVIOUR AND SOCIAL LIFE OF HONEY poor substitute for pollen. BEES. 352 pp. Bee Research Association Limited, London. (7) ROEDER, K. D. Artificial Diets 1953. INSECT PHYSIOLOGY. 1100 pp. Johu Wiley & Sons, Inc., New York. The beekeeper can supplement the diet of nec- (8) SHUEL, R. W., and DIXON, S. E. tar or honey with sucrose. This is usually mixed 1960. THE EARLY ESTABLISHMENT OF DIMORPHISM with about an equal amount of water and fed IN THE FEMALE HONEY BEE, APIS MELLIFERA as sirup. L. Insectes Sociaux 7: 265-282. There is no substitute for pollen. Various mate- (9) SMITH, M. V. rials, including brewer's yeast, soybean flour, 1959. QUEEN DIFFERENTIATION AND THE BIOLOG- ICAL TESTING OF ROYAL JELLY. N.Y. dry skim milk, and egg albumin, mixed with (Cornell) Agr. Expt. Sta. Mem. 356, 56 pp. honey or sugar water, have been fed to bees, but (10) STANDIFER, L. N. the colony stimulation is minor compared to that 1966. FATTY ACIDS IN DANDELION POLLEN derived from fresh pollen. The addition of dried GATHERED BY HONEY BEES, APIS MELLIFERA (HYMENOPTERA : APIDAE). Amer. Ent. Soc. pollen trapped from colonies earlier increases the Ann. 59: 1005-1008. stimulation slightly. (11) The two most commonly used artificial diets 1966. SOME LiPiD CONSTITUENTS OF POLLENS are the pollen supplement diet and the pollen COLLECTED BY HONEYBEES. Jour. ApiC substitute diet. Their composition is as follows: Res. 5: 93-98. Materials Percent (12) WAHL, O., and BACK, E. Pollen supplement : 1955. EINFLUSS DER IM POLLEN ENTHALTENEN Sugar-water (2 parts sugar to 1 part water by VITAMINE AUF LEBENSDAUER, AUSBILDUNG weight) 67 DER PHARYNXDRUESEN UND BRUTFAEHIG- Pollen-soy mix (1 part fresh dry pollen to 3 KEIL DER HONIGBIENEN (APIS MELLIFICA). parts soybean flour by weight) 33 Naturwissenschaften 42 (4): 103-104. HONEY, ITS COMPOSITION AND PROPERTIES

By J. W. WHITE, JR., Chief, Plant Products Laboratory, Eastern Utilization Research and Development Division, Agricultural Research Service Honey is the sweet viscous fluid elaborated by- considerable difference from other sweeteners, bees from nectar obtained from plant nectaries, chemists have long been interested in its composi- chiefly floral. After transportation to the hive in tion and food technologists sometimes have been the honey stomach, this fluid is ripened and stored frustrated in attempts to include honey in prepared in the comb for food. Other definitions may be food formulas or products.Limitations of methods more restrictive. For example, the U.S. Food and available to earlier researchers made their results Drug advisory definition for honey states that only approximate in regard to the true sugar ^^Honey is the nectar and saccharine exudation of composition of honey. Although recent research plants, gathered, modified, and stored in the comb has greatly improved analytical procedures for by honeybees {Apis mellvfera and A. dorsata); is sugars, even now some compromises are required levorotatory; contains not more than 25 percent in order to make possible accurate analysis of water, not more than 0.25 percent ash, and not large numbers of honey samples for sugars. more than 8 percent sucrose.'^ Although this An analytical survey of U.S. honey is reported definition once served a useful purpose, it is con- in Composition of American Honeys, Technical sidered today to allow much too high a content of Bulletin 1261, published by the U.S. Department water and sucrose and is too low in ash. Both of Agriculture in 1962. In this survey, consider- definitions exclude honeydew. able effort was made to obtain honey samples The color of U.S. honey may vary greatly, from all over the United States and to include from a nearly colorless fireweed or sweetclover enough samples of the commercially significant type through yellow, yellow green, gold, ambers, floral types that the results, averaged by floral dark browns or red browns to nearly black. The type, would be useful to the beekeeper and packer variations are almost entirely due to the plant and also to the food technologist. In addition to source of the honey, though climate may modify providing tables of composition of U.S. honeys, the color somewhat through the darkening action some general conclusions were reached in the of heat. bulletin on various factors affected by honey The flavor of honey varies even more than the composition. color. A honey may appear to have only a simple Where comparisons were made of the com- sweetness or may be mild, spicy, fragrant, aro- position of the same types of honey from 2 crop matic, bitter, harsh, medicinal, or objectionable. years, relatively small or no differences were found. This is again almost entirel}^ governed by the The same was true for the same type of honey from floral source. In general, a light-colored honey is various locations. As previously known, dark hon- expected to be mild in flavor and a darker honey ey is higher than light honey in ash (mineral) and to be of pronounced flavor. The exceptions com- nitrogen content. Averaging results by regions mon to all rules sometimes endow a light honey showed that eastern and southern honeys were with very deflnite specific flavors. Since flavor darker than average, whereas north-central and judgment is personal, one's favorite may be an- intermountain honeys were lighter. The north- other's ^^unflavored sugar sirup'' or ^411-tasting central honey was higher than average in moisture, medicine." With the tremendous variety avail- and the intermountain honey was more heavy able, everyone should be able to flnd his own bodied. Honey from the South Atlantic States favorite honey. showed the least tendency to granulate, whereas the intermountain honey had the greatest tendency. Composition of Honey The technical bulletin includes complete anal- yses of 490 samples of U.S. floral honey and 14 The characteristic physical properties of honey— samples of honeydew honey gathered from 47 of high viscosity, '*stickiness," great sweetness, rel- the 50 States and representing 82 *'single" floral atively high density, tendency to absorb moisture types and 93 blends of ''known" composition. from air, and immimity from some types of For the more common honey types, many samples spoilage—all stem from the fact that honey is were available and averages were calculated by naturally a very concentrated solution of several computer for many floral types and plant families. sugars. Because of its unique character and its In this bulletin are also given the average honey 56 BEEKEEPING IN THE UNITED STATES 57 composition for each State and region and detailed Most of these constituents are expressed in discussions of the effects of crop year, storage, percent, that is, parts per hundred of honey. area of production, granulation, and color on The acidity is reported difierently. In earlier composition. Some of the tabular data are included times acidity was reported as percent formic acid. in this handbook. We now know that there are many acids in honey, Table 1 gives the average and the range of with formic acid being one of the least important. values found for each constituent. The range Since a sugar acid, gluconic acid, has been found shows the great variability. to be the" principal one in honey, these results Nearly all the constituents are familiar. The could be expressed as ''percent gluconic acid'' by lévulose and dextrose are the simple sugars making multiplying the numbers in the table by 0.0196. up most of the honey. Sucrose (table sugar) is Since actually there are many acids in honey, present in high concentration in nectar, from the term "miUiequivalents per kilogram" is used which honey is made. ''Maltose'' represents a to avoid implying that only one acid is found in group of several more complex sugars, which honey. This figure is such that it properly ex- collectively are analyzed and reported as maltose. presses the acidity of a honey sample independently Higher sugars is a more descriptive term for the of the kind or kinds of acids present. material formerly called honey dextrin. In table 1, the differences between fioral honey The undetermined value is found by adding and honeydew honey ^ can be seen. Floral honey all the sugar percentages to the moisture value is higher in simple sugars (lévulose and dextrose), and subtracting from 100. The active acidity of lower in disaccharides and higher sugars (dextrins), a material is expressed as pH; the larger the num- and contains much less acid. The higher amount of ber the lower is the active acidity. The lac tone mineral salts (ash) in honeydew gives it a less is a newly found component of honey. Lactones active acidity (higher pH). The nitrogen content, may be considered to be a reserve acidity, since refiecting the amino acids and protein content, is by chemically adding water to them (hydrolysis) also higher in honeydew. an acid is formed. The ash is, of course, the The carbohydrate composition or principal material remaining after the honey is burned sugars in the more common types of honey are and represents mineral matter. The nitrogen is a measure of the protein material, including the 1 Strictly speaking, honeydew is an excretory product of enzymes, and diastase is a specific starch-digesting several species of insects when sucking plant juices. If it is enzyme. gathered and stored by bees, it becomes honeydew honey.

TABLE 1.—Average composition ojfloral and honeydew honey and range oj values ^

Floral honey Honeydew honey Characteristic or constituent Average values Range of values Average values Range of values

Color 2 _ Dark half of white. Light half of water Light half of Dark half of extra white to dark. amber. hght amber to dark. Liquid to complete Granulating tendency 3_ Few clumps of Liquid to complete Me-to^^-inchlayer crystals, }i- to }\' hard granula- of crystals. soft granulation. inch layer. tion. 17.2 13.4-22.9 16.3 12.2-18.2. Moisture percent 23.91-38.12. Lévulose do. _ 38.19 27.25-44.26 31.80 31.28 22.03-40.75 26.08 19.23-31.86. Dextrose do_- .44-1.14. Sucrose do_ _ 1.31 .25-7.57 .80 7.31 2.74-15.98 8.80 5.11-12.48. Maltose do_ _ 1.28-11.50. Higher sugars do__ 1.50 .13-8.49 4.70 Undetermined do-_ 3.1 0-13.2 - 10.1 2.7-22.4. 3.91 3.42-6.10 4.45 3.90-4.88. pH* 30.29-66.02. Free acidity 22.03 6.75-47.19 49.07 7.11 0-18.76 5.80 .36-14.09. Lactone * 34.62-76.49. Total acidity 4___ 29.12 8.68-59.49 54.88 Lactone-r-free acid .335 0-.950 .127 .007-.385. .169 .020-1.028 .736 .212-1.185. Ash percent-_ .047-.223. Nitrogen do .041 0-.133 .100 Diastase ^ 20.8 2.1-61.2 31.9 6.7-48.4.

1 Based on 490 samples of floral honey and 14 samples of 3 Extent of granulation for heated sample after 6 months' honeydew honey. undisturbed storage. 2 Expressed in terms of U.S. Dept. Agr. color classes. 4 MiUiequivalents per kilogram. 5 270 samples for floral honey. 58 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

shown in table 2. In all cases lévulose predomi- lévulose, with small amounts of more complex nates. There are a few types, not represented in sugars. Many other substances also occur in honey, the table, that contain more dextrose than lévu- but the sugars make up by far the greater propor- lose, such as dandelion and bluecurls. This excess tion of honey. The principal physical character- of lévulose over dextrose is one way that honey istics and behavior of honey are due to its sugars, differs from commercial invert sugar. Lévulose is but the minor constituents, such as flavoring more soluble than dextrose. Even though honey materials, pigments, acids, and minerals, are has less dextrose than lévulose, the former is the largely responsible for the differences among indi- sugar that crystallizes when honey granulates or vidual honey types. ^'sugars.'' The sucrose level in honey never reaches zero, even though it may contain an active sucrose-splitting enzyme. Water Content As noted, honey varies tremendously in color Beekeepers as well as honey buyers know that and flavor, depending largely on its floral source. the water content of honey varies greatly. It may Although many hundreds of kinds of honey are range between 13 and 25 percent. According to the produced in this country, only about 25 to 30 are United States Standards for Grades of Extracted commercially important and are available in large Honey, honey may not contain more than 18.6 quantity. As also noted, even these may show percent moisture to qualify for U.S. grade A considerable variation in composition and proper- (U.S. Fancy) and U.S. grade B (U.S. Choice). ties. Until this survey of U.S. honey was reported, Grade C (U.S. Standard) honey may contain up to the degree of variation was not known and it re- 20 percent water; any higher amount places a tarded the use of honey by the food industry. honey in U.S. grade D (Substandard). Honey consists essentially of a highly concen- These values represent limits and do not indi- trated water solution of two sugars, dextrose and cate the preferred or proper moisture content for

TABLE 2.—Carbohydrate composition oj "honey types

Number of Floral type Dextrose Lévulose Sucrose Maltose Higher samples sugars

Percent Percent Percent Percent Percent 23 Alfalfa 33.40 39. 11 2.64 6.01 .89 25 Alf alf a-sweetclo ver. 33.57 39.29 2.00 6.30 .91 5- Aster 31.33 37.55 .81 8.45 1.04 3. Basswood 3L59 37.88 1. 20 6.86 L44 3_ Blackberry 25.94 37.64 L27 1L33 2.50 5. Buckwheat 29. 46 35.30 .78 7.63 2.27 4_ Buckwheat, wild__ 30.50 39.72 .79 7.21 .83 26 ''Clover" 32. 22 37.84 1. 44 6.60 1.39 3- Clover, alsike 30.72 39. 18 L40 7.46 L55 3_ Clover, crimson 30.87 38. 21 .91 8.59 L63 3- Clover, Hubam 33.42 38.69 .86 6.23 .74 10 Cotton 36.74 39. 28 L 14 4.87 .50 3. Fireweed 30.72 39.81 L28 7. 12 2.06 6- Gallberry 30. 15 39.85 .72 7.71 1.22 3- Goldenrod 33. 15 39.57 .51 6.57 .59 2. Heartsease 32.98 37.23 L95 5.71 .63 2_ Holly 25. 65 38.98 LOO 10.07 2. 16 3_ Honey dew, cedar. _ 25.92 25. 16 .68 6.20 9.61 5- Honey dew, oak 27.43 34.84 .84 10. 45 2. 16 2. Horsemint 33.63 37.37 LOI 5.53 .73 3_ Locust, black 28.00 40. 66 LOI 8. 42 L90 3. Loosestrife, purple. 29.90 37.75 .62 8. 13 2.35 3- Mesquite 36.90 40.41 .95 5.42 .35 4. Orange, California. 32.01 39.08 2.68 6.26 L23 13 Orange, Florida 31.96 38.91 2.60 7.29 1.40 4_ Raspberry 28.54 34.46 .51 8.68 3.58 3_ Sage 28. 19 40.39 1. 13 7.40 2.38 3-, Sourwood 24. 61 39.79 .92 1L79 2.55 4_. Star-thistle 31. 14 36.91 2.27 6. 92 2.74 8., Sweetclo ver 30.97 37.95 L41 7.75 1. 40 3_. Sweetclover, yellow 32.81 39.22 2. 93 6.63 .97 4_. Tulip tree 25.85 34. 65 .69 11. 57 2.96 5-. Tupelo 25. 95 43.27 1. 21 7.97 1. 11 7_. Vetch 3L67 38.33 L34 7.23 1.83 9_. Vetch, hairy 30. 64 38.20 2.03 7.81 2.08 12. Whiteclover 30.71 38. 36 L03 7.32 1.56 BEEKEEPING IN THE UNITED STATES 59 honey. If honey has more than 17 percent moisture Acids of Honey and contains a sufficient number of yeast spores, it The acids of honey, though nearly negligible will ferment. Such honey should be pasteurized, on the weight basis (less than one-half percent), i.e., heated sufficiently to kill such organisms. This have a pronounced effect on the flavor. They also is particularly important if the honey is to be may be responsible in part for the excellent sta- ''creamed" or granulated, since this process results bility of honey against micro-organisms. At least in a slightly higher moisture level in the liquid 18 organic acids have been reported in honey, part. On the other hand, it is possible for honey with varying degrees of certainty. Until recently to be too low in moisture from some points of view. it was thought that citric and malic acids were In the West, honey may have a moisture content the principal ones. Now it is realized that gluconic as low as 13 to 14 percent. Such honey is somewhat acid is the acid present in the greatest amount difficult to handle, though it is most useful in in honey. It arises from dextrose through the blending to reduce moisture content. It contains action of an enzyme recently found in honey over 6 percent more honey solids than a product of called glucose oxidase. Other acids reported in 18.6 percent moisture. honey are formic, acetic, butyric, lactic, oxalic, In the 490 samples of honey analyzed in the succinic, tartaric, maleic, pyroglutamic, pyruvic, Department's Technical Bulletin 1261, the average a-ketoglutaric, and glycollic. moisture content was 17.2 percent. Samples ranged between 13.4 and 22.9 percent, and the standard Proteins and Amino Acids deviation was 1.46. This means that 68 percent of the samples (or of all U.S. honey) will fall within It will be noted in table 1 that the amount of the hmits of 17.2 ±1.46 percent moisture (15.7- nitrogen in honey is low, on the average 0.04 18.7) ; 95.5 percent of all U.S. honey will fall within percent, though it may range to 0.1 percent. If the limits of 17.2±2.92 percent moisture (14.3- this were all from protein in honey, the corre- sponding protein values would be about 0.25 to 20.1). 0.8 percent. Since other nitrogenous substances In the same bulletin a breakdown of average are known to occur in honey, the true values for moisture contents by geographic regions is shown. protein content are somewhat lower. Little is These values (percent) are North Atlantic 17.3, known of the proteins of honey, except that the East North Central 18.0, West North Central 18.2, enzymes fall into this class. The peculiar physical South Atlantic 17.7, South Central 17.5, Inter- properties of heather honey, which thickens to a mountain West 16.0, and West 16.1. solid on standing and thins when stirred, are reported to be due to a protein, which if added Sugars of Honey to clover honey will confer these same properties Honey is first and foremost a carbohydrate. on it. Sugars make up 95 to 99.9 percent of the solids The presence of proteins causes honey to have of honey and their identity has been studied a lower surface tension than otherwise, which for may years. produces a marked tendency toward foanaing and Honey was long thought to be mainly lévulose scum formation and encourages formation and and dextrose, with some sucrose and dextrins. retention of fine air bubbles. Beekeepers familiar These were considered to be poorly defined com- with buckwheat honey know how readily it tends plex sugars of high molecular weight. With the to foam and produce surface scum, which is largely advent of new methods for analyzing and due to its relatively high protein content. separating sugars, workers in Europe, the United The amino acids are simple compounds obtained States, and Japan have found many sugars in when proteins are broken down by chemical or honey and in some cases isolated and identified digestive processes. They are the ''building blocks'' them by suitable physical and chemical methods. of the proteins. Several of them are essential to life and must be obtained in the diet. The quan- Dextrose and lévulose are still the principal tity of free amino acids in honey is small and of sugars, but at least 12 more have been found in no nutritional significance. Recent breakthroughs honey, namely maltose, isomaltose, turanose, in the separation and analysis of minute quan- maltulose, nigerose, kojibiose, leucrose, meiezitose, tities of material (chromatography) have revealed erlöse, kestose, raffinose, and dextrantriose. Most that various honeys contain from 11 to 21 différ- of these sugars probably do not occur in nectar, èrent free amino acids. Proline, glutamic acid, but arise because of either enzymic action during alanine, phenylalanine, tyrosine, leucine, and iso- the ripening of honey or chemical action during leucine are the most common. storage in the concentrated, somewhat acid sugar Amino acids are known to react slowly, or mixture we know as honey. Individual honey more rapidly by heating, with sugars to produce types vary in the relative amounts of the various yellow or brown materials. Part of the darkening sugars, but each seems to have the same kinds of honey with age or heating may be due to of minor sugars. this. 60 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

Minerals measure of honey quality by several European When honey is dried and burned, a small residue ^^^^ ^^^' of ash invariably remains. This is the mineral Another enzynie recently found m honey is glu- content. As shown in table 1, it varies from 0.02 ^^^^ oxidase. This converts dextrose to a related to slightly over 1 percent for a floral honey, material, a gluconolactone, which m turn forms averaging about 0.17 percent for the 490 samples glucomc acid the principal acid in honey. Since analyzed. ^^^^ enzyme had previously been shown to be in Honeydew honey is richer in minerals, so much ^^^F^u^^^^^I^ ^^^""^ ""^^^ ^''''^^ ^^^^ '^ f ^'^^^^ so that its mineral content is said to be a prime ^^^^ ^^'^ ^1^^® ^^^^^^•. -^f.^^ ^S^^^, ^^ "^^^^ other cause of its unsuitabiUty for winter stores. Schuette enzynies, the amount m different honeys is van- and his colleagues at the University of Wisconsin f^^^' T T ^^^^^ ^^ gluconolactone, this enzyme have examined the mineral content of light and t^^"^^ hydrogen peroxide during its action on dex- dark honey. They reported the following average ^^^l®' ^^}^.^^^ been shown to be the basis of the values: ^ ^ & & neat-sensitive antibacterial activity of honey. Light honey Dark honey Other cnzymcs havc been reported in honey, Potassium ^""íoo itu including inulase and phosphatase. Except for Chlorine___I-"'"''"'"]'"']]^] 52 ' 113 catalase, the results have not been sufficiently Sulfur 58 100 confirmed. Sodium^ ~ 18 76 ^^^ ^^^^^ enzymes can be destroyed or weak- PhosphorusI----^'"--'"""!"!!! 35 47 P^d by heat. For the use of enzyme levels to Magnesium 19 35 indicate heating history of honey, see page 63. Silica 22 36 irng"an"e¡¿::::::::::::::::::::: ^: 30 109 Properties of Honey ■g^^^^g^ ^£ j^^ unusual composition, honey ex- EnzymeS hibits some properties that may make its handling ^ ^ and use somewhat difficult. Means have been de- One of the characteristics that sets honey apart veloped to cope with these problems. Honey also from all other sweetening agents is the presence has some interesting attributes, of enzymes. These are complex protein materials A x-i. j. • i A ^- -^ o TX that under mild conditions bring about chemical Antibacterial Activity of Honey changes which niay be very difl^icult to ac- it was once thought that since milk can be a comphsh m the laboratory^^ Enzymatic reactions carrier of some diseases, honey might Hkewise be are the very basis of life. Enzymes in honey can such a carrier. Some years ago this idea was ex- conceivably arise from the bee, pollen, nectar, or amined by adding nine common pathogenic bac- even yeasts and niicro-orgamsms. Those most teria to honey. All of them died within a few hours prominent are added by the bee in the conversion or days. Honey is not a suitable medium for bac- 01 nectar to honey. teria for two reasons—it is fairly acid and it is too Invertase, also known as sucrase or saccharase, high in sugar content for growth to occur. This spHts sucrose into its constituent simple sugars, kilHng of bacteria by high sugar content is called dextrose and lévulose. Other more complex sugars the osmotic effect. It seems to function by Hter- have been found recently to form in small amount ally drying out the bacteria. However, in tlie during this action and in part explain the com- resting spore form, some bacteria can survive, plexity of the minor sugars of honey. Although though not grow in honey. the work of invertase is completed when honey Another type of antibacterial property of honey is ripened, the enzyme remains m the honey and is that due to inhibine. The presence in honey of retains its activity for some time. Even so, the an antibacterial activity was first reported about sucrose content of honey never reaches zero. 1940 and confirmed in several laboratories. Since bince the enzyme also synthesizes sucrose, perhaps then several papers have appeared on the subject, the ñnal low value for the sucrose content of it has been generally agreed that inhibine (name honey represents an equilibrium between sphttmg used by Dold, its discoverer, for antibacterial and formation of sucrose. activity) is sensitive to heat and fight. The effect Another enzyme known to be in honey is diastase of heating honey on its inhibine content has been (amylase). Since this enzyme digests starch to sim- studied by several investigators. It is apparent pier compounds and starch has not been found in that heating it sufficiently to reduce markedly or nectar, it is not clear what its function in honey destroy its inhibine activity would deny it a might be. Diastase appears to be present in vary- market as first-quality honey in several European ing amounts in nearly aU honey and it can be countries. In fact, the use of sucrase and inhibine rneasured. It has probably had the greatest atten- assays together to determine the heating history tion in the past, because it has been used as a of commercial honey has been proposed. Until BEEKEEPING IN THE UNITED STATES 61 recently no information on the actual nature or plied heat. Honey that has granulated can be re- constitution of inhibine has been published. turned to liquid by careful heating. Heat should be It is now known that this inhibine effect is due to applied indirectly by hot water or air, not by direct the accumulation of hydrogen peroxide in diluted flame or high-temperature electrical heat. Stirring honey. This material, well known for its antiseptic accelerates the dissolution of crystals. For small properties, is a byproduct of the formation of containers, temperatures of 140° F. for 30 minutes gluconic acid in diluted honey by an enzyme that will usually suffice. occurs in honey, glucose oxidase. The peroxide can If unheated honey is allowed to granulate nat- inhibit the growth of certain bacteria in the diluted urally, several difficulties may arise. The texture honey. Since it is destroyed by other honey con- may be fine and smooth or granular and objection- stituents, an equihbrium level of peroxides will able to the consumer. Furthermore, a granulated occur in a diluted honey, its magnitude depending honey becomes more susceptible to spoilage by fer- on many factors such as enzyme activity, oxygen mentation, caused by natural yeast found in all availabihty, and amounts of peroxide-destroying honeys and apiaries. Quahty damage from poor tex- materials in the honey. The amount of inhibine ture and fermented flavors is usually far greater than (peroxide accumulation) in honey depends on any caused by the heat needed to eliminate these floral type, age, and heating. problems. A chemical assay method has been developed Finely granulated honey may be prepared from that rapidly measures peroxide accumulation in a honey of proper moisture content (17.5 percent diluted honey. By this procedure different honeys in summer, 18 percent in winter) by several proc- have been found to vary widely in the sensitivity esses. All involve pasteurization to eliminate fer- of their inhibine to heat. In general, the sensitivity mentation, foUowed by addition at room temper- is about the same as or greater than that of invertase ature of 5 to 10 percent of a finely granulated and diastase in honey. "starter" of acceptable texture, thorough mixing, and storage at 55° to 60° F. in the retail containers Food Value of Honey for about a week. The texture remains acceptable if storage is below about 80° to 85°. As a carbohydrate food, honey is a most delec- table and enjoyable treat. Its distinctive flavors Deterioration of Honey Quality cannot be found elsewhere. The sugars are largely the easily digestible ''simple sugars,'' similar to Fermentation.—Fermentation of honey is those of many fruits. caused by the germination and growth of yeasts Honey because of its content of simple sugars is that are normally found in afl honey. These an excellent source of energy. It can be regarded yeasts, which may be in the soil of every apiary, as a good food for both infants and senior citizens. in the honey house, and in the hive, differ from The enzymes of honey, though used as indicators ordinary bread or wine yeasts. The obvious differ- of heating history and hence table quality of honey ence is that these yeasts can grow at much higher in some countries, have no nutritional value and sugar concentrations than other yeasts, and are are destroyed in the digestive process. Mineral con- therefore called "osmophilic." Even so, there are tent of honey is variable, but some darker honeys upper limits of sugar concentration beyond which may have significant quantities of trace minerals. these yeasts wiU not grow. Thus the water content Although some vitamins may be demonstrated in of a honey is one of the factors concerned in honey, the amounts are far too low to have any spoilage by fermentation. The others are extent meaning in human nutrition. of contamination by yeast spores (yeast count) and temperature of storage. Granulation of Honey Honey with less than 17.1 percent water will not A large part of the honey sold to consumers in ferment in a year, irrespective of the yeast count. the United States is in the Uquid form, much less in Between 17.1 and 18 percent moisture, honey a finely granulated form known as ''honey spread'' with 1,000 yeast spores or less per gram will be or finely granulated honey, and even less as comb safe for a year. When moisture is between 18.1 honey. The consumer appears to be conditioned and 19 percent, not more than 10 yeast spores to buying liquid honey. At least sales of the more per gram can be present for safe storage. Above 19 convenient spread form have never approached percent water, honey can be expected to fer- ment even with only one spore per gram of honey, those of liquid honey. Smce the granulated state is natural for most of a level so low as to be very rare. the honey produced in this country, processing is When honey granulates, the resulting increased required to keep it hquid. Careful appUcation of moisture content of the hquid part is favorable heat to dissolve "seed" crystals and avoidance of for fermentation. Honey with a high moisture subsequent "seeding" will usually suffice to keep content AviU not ferment below 50° F. or above^ a honey Hquid for 6 months. Damage to color and about 80°. Honey even of relatively low water flavor can result from excessive or improperly ap- content wifl ferment at 60°. Storage at temper- 261-907 0—67—^5 62 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

atures over 80°, to avoid fermentation, is not processing temperatures and storage on honey practical as it will damage honey. quality. V, ^' .^j-^^''i;^'^' °^ Michigan State University, To be of highest quality, a honey—whether has studied the mechanism and course of yeast liquid, crystallized, or comb—must be well ripened fermentation in honey in conjunction with his with proper moisture content; it must be free of work on the hygroscopicity of honey. He con- extraneous materials, such as excessive pollen firmed that when honey absorbs moisture, which dust, insect parts, wax, and crystals if liquid- it occurs when It is stored above 60-percent relative must not ferment; and above all it must be' of humidity the moisture content at first increases excellent flavor and aroma, characteristic of the mostly at the surface before the water diffuses particular honey type. It must, of course, be free into the bulk of the honey. When honey absorbs of off-flavors or odors of any origin In fact the moisture, yeasts grow aerobically (using oxygen) more closely it resembles the well-ripened honev at the surface and multiply rapidly, whereas as it exists in the cells of the comb, the better it is below the surface the growth IS slower. Several beekeeping practices can reduce the J^ermentmg honey IS usually at least partly quahty of the extracted product. These include granulated and is characterized by a foam or combining inferior floral types, either by mixing Iroth on the surface It will foam considerably at extracting time or removing the crop at incor- when heated An odor as of sweet wme or fer- rect times, extraction of unripe honey, extraction mentmg fruit may be detected Gas production of brood combs, and delay in settling and straining may be so vigorous as to cause honey to overflow However, we are concerned here with the handling or burst a contamer. The off-flavors and odors of honey from its extraction to its sale. During associated with fermentation probably arise from this time improper settling, straining, heating the acids produced by the yeasts and storage conditions can r^iake a superb honey Money that has been fermented can sometimes into just another commercial product, be reclaimed by heating it to 150° F. for a short The primary objective of all processing of honey i ■ f X '*Tfl*^^ fermentation and expels is simple-to stabilize it. This means to keep it some of the off-flavor Fermentation in honey free of fermentation and to keep the desired may be avoided by heating to kill yeasts. Minimal physical state, be it liquid or finely granulated, treatments to pasteunze honey are as follows: Methods for accomplishing these objectives have Heating ^^^^ fairly well worked out and have been used Temperature CF.) (JÍZes) ^^'^ I^^^l ^^^''^■- Probably improvements can be 128 kmmutes) made. The requirements for stability of honey are lSOmill'lil[lllllllllll[llll[lllllll[ll 170 ™^^® stringent now than in the past, with honey }35 1 60 a world commodity and available in supermarkets 145 ^7 ^^^ ^^^^ around. Government price support and l5o'.'-'/.iy.'///.'_'_'_ 2 8 ^^^^ operations require storage of honey, and I55...\llllllllll^llllllll[ll\l\l\imy 1.0 market conditions may also require storage at 160 'V.l '.4: any point in the handling chain, including the ru,„ju„. r „ Ï. rr a- JO. rry, producer, packer, wholesaler, and exporter. nfW iÜrffl /^ ^'fr^ """"^ .-Storni.-The The primary operation in the processing of W ?f,i^l P ^5^1''^- ^"""^y spoilage, damage honey is the application and control of heat. If Spd tn r^r^.r i^ ™proper storage are re- we consider storage to be the application of or ^tt nnltl f ■■ ^^,S^"?^^1' t^^^^^ ^^^^ t'^ke exposure to low amounts of heat over long periods, t r T.S f^^^ f"*'"^ ^^-fu ''fr' "^^"i ^' it «^° be seen that a study of the effects of heat on n3n/nnl« r"^ Storage With the rate de- honey quality can have a wide application. llf^^SZ if *f7P"^^t"^«- These include dark- Any assessment of honey quality must include Ti^.lrLrlJlt- r^' ^""^ formation of flavor considerations. The objective measurement off-flavor (caramehzation). of changes in flavor, particularly where they are 1 o keep honey in its original condition of high gradual, is most difficult. We have measured the quality and delectable flavor and fragrance is accumulation of a decomposition product of the possibly the greatest responsibility of the bee- sugars (hydroxymethylfurfural or HMF) as an keeper and honey packer. At the same time it index of heat-induced chemical change in the is an operation receiving perhaps less attention honey. Changes- in flavor, other than simple loss trom the producer than any other and one re- by evaporation, may also be considered heat- qmring careful consideration by packers and induced chemical changes. wholesalers To do an effective job, one must To study the effects of treatment on honey, know the factors that govern honey quality, as we must use some properties of honey as indices well as the effects of various beekeeping and of change. Such properties should relate to the storage practices on honey quality. The factors quality or commercial value of honey The are easily deterniined, but only recently are the occurrence of granulation of liquid honey, lique- facts becoming known regarding the effects of faction or softening of granulated honey, and BEEKEEPING IN THE UNITED STATES 63 fermentation as functions of storage conditions honey in which a good enzyme level is needed, as has been reported; also, color is easily measured. for export. As indicators of the acceptability of honey for The damage done to honey by heating and by table use, Europeans have for many years used storage is the same. For the lower storage temper- the amount of certain enzymes and HMF in honey. atures, simply a much longer time is required to They considered that heating honey sufficiently obtain the same result. It must be remembered to destroy or greatly lower its enzyme content or that the effects of processing and storage are addi- produce HMF reduced its desirability for most tive. It is for this reason that proper storage is so uses. A considerable difference has been noted in important. A few periods of hot weather can offset the reports by various workers on the sensitivity the benefits of months of cool storage—^10 days to -heat of enzymes, largely diastase and invertase, at 90° F. are equivalent to 100-120 days at 70°. in honey. Only recently has it been noted that An hour at 145° in processing will cause changes storage alone is sufficient to reduce enzyme content equivalent to 40 days' storage at 77°. and produce HMF in honey. Since some honey Perhaps the easiest way for an individual to types frequently exported to Europe are naturally decide whether he has storage or processing dete- low in diastase, the response of diastase and rioration in his operation is to take samples of the invertase to storage and processing is of great fresh honey, being careful that they are fairly rep- importance for exporters. resentative of the batch, place them in a freezer A study was recently made of the effects of for the entire period, then allow them to warm to heating and storage on honey quality and was room temperature, and compare them by color, based on the results with three types of honey flavor, and aroma with the honey that has been stored at six temperatures for 2 years. The results in common storage. In some parts of the United were used to obtain predictions of the quality States the value of the difference can reach 1}^ life of honey under any storage conditions. The cents per pound in a few months. Such figures cer- following information is typical of the calculations tainly would justify expenditures for temperature based on this work. control. At 68° F., diastase in honey has a half-life of The individual storing honey is on the horns of 1,500 days, nearly 4 years. Invertase is more a dilemma. He must select conditions that will heat sensitive, with a half-life at 68° of 800 days, minimize fermentation, undesirable granulation, or about 2M years. Thus there are no problems and heat damage. Fermentation is strongly re- here. By increasing the storage temperature to tarded below 50° F. and above 100°. Granulation 77°, half the diastase is gone in 540 days, or 1% is accelerated between 55° and 60° and initiated by years, and half the invertase disappears in 250 fluctuation at 50°-55°. The best condition for days, or about 8 months. These periods are still storing unpasteurized honey would seem to be rather long and there would seem to be nothing below 50°, or winter temperatures over much of to be concerned about. However, temperatures the United States. Warming above this range in in the 90^s for extended periods are not at all the spring can initiate active fermentation in such uncommon: 126 days (4 months) will destroy half honey, which is usually granulated and thus even the diastase and about 50 days (2 months) will more susceptible. eliminate half the invertase. As the temperature Some progressive producers and packers are increases, the periods involved become shorter now using controlled temperature storage for and shorter until the processing temperatures are honey, particularly in the warmer regions. Using reached. At 130°, 2K days would account for half the data we now have, we can definitely state that the diastase and in 13 hours half the invertase is reducing the storage temperature of honey by 10° gone. to 15° F. will reduce the rate of honey deteriora- A recommended temperature for pasteuriza- tion to about one-third to one-sixth of that at the tion of honey is 145° F. for 30 minutes. At this higher temperature. Such a temperature reduc- temperature diastase has a half-life of 16 hours tion, which can easily be attained by air-condi- and invertase only 3 hours. At first glance this tioning, would reduce HMF production to one- might seem to present no problems, but it must third, enzyme loss to one-fifth to one-sixth, and be remembered that unless flash heating and darkening rate to about one-sixth of the rate at immediate cooling are used, many hours will the higher temperature. Loss of fiavor and fresh- be required for a batch of honey to cool from ness would be expected to be reduced similarly. 145° to a safe temperature. Thus honey can at any time of the year be more If we proceed further to a temperature often nearly like honey at its very best—fresh from the recommended for preventing granulation, 160° F. comb at extraction time. for 30 minutes, the necessity of prompt cooling becomes highly important. At 160°, 2K hours will References destroy half of the diastase, but half of the more (1) TOWNSEND, G. F. sensitive invertase will be lost in 40 minutes. This 1961. PREPARATION OF HONEY FOR MARKET. treatnient then cannot be recommended for any Ontario Dept. Agr. Pub. 544, 24 pp. 64 AGRICULTURE, HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

(2) WHITE J. W, JR. . ^, ... (3) RIETHOF, M. L, SUBERS, M. H., and 1963. HONEY. In Grout, R. A., ed., The Hive KUSHNIR, I. » ^ ??^i \^? c. ^^^Í7^^?; ^?^'' ^^^ PP- 1962. COMPOSITION OF AMERICAN HONEYS. U.S. Dadant & Sons, Hamilton, 111. Dept. Agr. Tech. Bui. 1261, 124 pp. QUEEN AND PACKAGE BEE PRODUCTION

By WILLIAM C. ROBERTS and WARREN WHITCOMB, JR., apiculturists, Entomology Research Division^ Agricultural Research Service *

High quality queens are necessary for repeat sales, Rearing Queens and quantity production is necessary for profit- able operations. Customer demand determines For the amateur who desires to rear a few the kind or breed of queens produced. Each queen queens, many methods are available. Most of breeder uses methods that suit his own particular them are described in the references at the end conditions. of this section. For those desiring to rear less than 20 queen cells at one time and not more than Queen cells are started when worker larvae are 50 queens during a season, we suggest the fol- transferred or grafted into wax cells and placed in lowing procedure: Select the best colony as a special cell-starting colonies. Two queenless types breeder to produce the queen cells. Rear them are in common use. The ^'swarm box^' is composed after the main honey flow is over by two easy of approximately 4 pounds of bees confined in a manipulations. box with combs of honey and pollen. The ''queen- First, remove the queen with one frame of brood less colony'' is a hive with 6 pounds or more of and two frames of honey with adhering bees to a bees and an open entrance permitting the bees to hive body in another location. The queenless fly. In each of these types about 50 queen cells colony will start a dozen or more queen cells can be started. The cells are left in these starters within 24 hours. These will be produced through- for 24 to 36 hours and then transferred to the cell- out the brood nest. Cut out and save the largest finisher colonies. If the queen breeder uses the and most uniform cell. Be sure that the cell ''double graft'' method of producing queens, he selected is not damaged. Destroy all unused cells. removes the larvae from the started queen cell Return the queen and three frames to the center and replaces them with other young larvae that of the brood nest of the parent colony. If the bees are allowed to remain. are not robbing, there will be little chance of Some cell-finisher colonies are queenless, others losing the queen by this method. are queenright. They are populous colonies with Finally, make an additional colony or nucleus 10 pounds or more of bees and with both brood and by again removing from another colony three bees of all ages. They must have ample food—both frames of brood, honey, and adhering bees. Gouge honey and pollen. The queenless cell finisher a cavity near the top of the center comb about % requires the addition of brood and bees at regular inch wide and IK inches long. Gently press the intervals. The queenright cell finisher has a brood queen cell into it so the queen will have room to nest and queen below the excluder. The queen cells emerge from the cell tip. Replace the comb and are placed between frames of brood in the upper leave the new hive closed for 2 weeks. Repeat this part of the colony. Started queen cells are added process for each queen desired. every 3 or 4 days. The frame containing the started The commercial queen breeder does not use this queen cells is placed between the frames of un- inefficient method. He transfers female larvae sealed brood. from worker cells into artiñcial queen cell cups A few queen breeders start and finish queen cells and uses special queenright or queenless cell- in the same queenless colony. Others start and building colonies to rear them. By this method finish the cells in a queenright cell-builder colony. hundreds of cells can be produced continuously. Good cell builders properly handled produce good queen cells and poor cell builders produce Commercial Queen Production poor queen cells. The developing queen larva feeds for only 5 days. If it is properly and adequately In the United States there are many com- fed during this period, a good virgin queen results. mercial queen breeders who rear annually several If inadequately fed during this period, a poor queen thousand queens each. They are concerned with results. The best queen rearing methods are those quantity production of high quality queens. that provide for well-fed larvae during the 5-day feeding period. Malnutrition results in small 1 In cooperation with Louisiana Agricultural Experiment Station. queens with fewer ovarian tubules. 66 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

Queen cells are removed from the cell-builder honey, and about 1}^ pounds of bees. A three- colonies 9 or 10 days after grafting. Ten-day old frame standard nucleus box with screen top and cells go directly to the small mating hive or nuclei, bottom makes a satisfactory carrier package for but 9-day cells are kept for 1 or 2 days in an incu- shipment. bator maintained at brood rearing temperatures. A commercial queen breeder will obtain 200 to On the 11th day after grafting, the virgin queen 400 queen cells from each cell-building colony each emerges from the queen cell. Emergence may be month of operation. This is enough to supply queen delayed by keeping the queen cells at a slightly cells for 100 to 200 nuclei. Few queen breeders lower temperature during the late pupal stage of average more than 50 mated queens for each 100 the queen. cells put into nuclei. Many types and sizes of mating nuclei are in Improvements in the technique of artificial in- use. Most queen breeders use a three-frame nucleus semination of queen bees may soon eliminate the containing a feeder. The frames vary in size from need of nuclei and possibly cut the cost of queen about 3 or 4 inches to deep Langstroth frames. production. Ninety-five percent survival of queens Many are established with bees, brood, and honey. mated artificially is not uncommon with present Others are started without brood or honey, and techniques. the feeder is filled with sugar sirup. The amount of The greatest customer demand for package bees bees needed to stock a nucleus varies from one- and queens occurs in March, April, and May. Con- fourth to three-fourths pound. If brood is given, sequently, the large producers are located in areas only one frame of brood is placed in each nucleus. with a climate that permits economical production Many nuclei are started with only two combs, before and during these months. Locations in the allowing space for adding the bees to the nucleus South and in California, with mild winters and box. Nuclei are usually closed for 12 hours if early springs, are ideally suited for early produc- brood is given, but may be kept closed for 2 or 3 tion of populous colonies. The choice locations are days if no brood is used. When only two frames are those that have not only a favorable climate but used in installing the bees in the nucleus, the also good floral sources of nectar and pollen. third frame is added 4 days later. Many queen An area without a good nectar flow can be sat- breeders make up nuclei in a building ana confine isfactory, but a good pollen flow is essential. Many them there for a day or two before moving them producers feed sugar sirup to their colonies during to the mating yard. February and March. Others find it necessary to To insure proper mating, the queen breeder feed continuously during their most active season. should have an adequate supply of mature drones There is no substitute for an abundant pollen in- at all times during the mating periods. Special come during the spring and fall. Pollen storage in drone-producing colonies should be maintained the fall is necessary for early spring buildup. in the apiaries containing the nuclei. Producers Without adequate pollen sources after the middle who do not maintain plenty of drones usually of February the colonies will curtail brood produc- have many inadequately mated queens. tion. Fourteen days after a queen cell is given to a The queen and package bee production in the nucleus, it should have a mated queen with eggs United States today is a $5 million industry. More and young larvae. Such queens are ready for than 1 million queens and over 500 tons of bees shipment and can be removed. Another queen cell are produced and sold to beekeepers in the United can then be given. Many breeders prefer to wait States and Canada each year. until the next day after removal of a laying queen before giving another queen cell. If the queen is to be shipped by mail, the queen Package Bee Production cage should have candy and 7 to 10 attendant worker bees. If the queen is to be placed in a Commercial shipments of package bees for re- package of bees for shipment or if she is to be placing or starting new colonies in the honey-pro- placed in a queen bank for storage, she should be ducing regions were apparently first made in 1912 caged without attendant bees and without queen by H. C. Short from Fitzpatrick, Ala. The demand cage candy. for package bees increased greatly after World A queen bank is a colony without a laying queen War I and shipments of queens reached a peak and sonietimes without brood. Often 200 to 500 about 1947. The Railway Express Agency prob- queens in cages are stored in one such colony. ably handled 80 to 90 percent of all bee shipments They can be kept in a queen bank for a month or during the 1920's and 1930's. Since then truck, air, longer if sufficient populations are maintained. and mail shipments have become increasingly Some queen breeders prefer to ship large orders important. Approximately 80 percent of all package of queens in queen cage carriers rather than adding bees are now trucked to northern honey producers. queen cage candy and attendants to each queen The requirements for producing a maximum cage. A carrier consists of two Langstroth frames crop of bees and a maximum crop of honey are holding 36 queens in each frame, one frame of basically similar. Prolific queens, adequate equip- BEEKEEPING IN THE UNITED STATES 67 ment, abundant stores at the proper time, a more than 6 pounds of bees at a single shaking knowledge of local nectar and pollen sources, and results in decreased brood production. a system of management planned for the particular Three-story colonies are necessary during the environment are necessary for both crops. Funda- buildup period to supply the area for eggs and mental differences are (1) the length of time during brood that a prolific queen requires together which colonies must be stimulated for maximum with the comb space necessary for honey and pol- brood production, (2) the amount of equipment len stores. In order to decrease labor in handling used, and (3) the maintenance. The shipping hive bodies, most of the honey can be placed in season lasts for 2 to 3 months for the bee-producing the bottom body at the beginning of the shaking colony as opposed to a honey-flow period of 3 to period and only the top two bodies manipulated. 6 weeks for the honey-producing colony. Three This three-story colony can be manipulated with deep bodies are used for the bee-producing colony essentially the same labor requirements as a two- and a minimum of three deep bodies for the story colony, but because of the space and food honey-producing colony. After each shaking, a reserves in the bottom body the brood produc- population of about 18,000 bees—the level that tion will be higher. A maximum of 37 pounds of gives the maximum production of brood per bee— bees has been shaken from a single colony and is required for the bee-producing colony as opposed an average of 32 pounds of bees per colony from to a population of 60,000 or more bees for maxi- 10 colonies during a 60-day period without miun honey production. detriment to the colonies. Methods of producing package bees have been Methods of getting the bees into the shipping carefully studied. A system of management that package vary widely depending on local honey- is successful in Baton Rouge, La., will be described, flow conditions, the type of package used, and and it can be easily adapted to other areas. how it will be transported. Two widely used methods include (1) shaking the bees from combs Preparing colonies for package bee production in the brood nest, which requires more labor but should begin in August, with a check of colonies allows a constant check on the condition of the for good queens and the replacement of failing queen and colony, and (2) the smoke-up or drive- queens. August and September can be a danger up method, by which bees are smoked or driven period in many parts of the shipping area because by a repellent up onto combs in the top body, of lack of honey flow—almost a necessity for then shaken off. By this method the brood nest successful queen introduction—and especially be- combs are not removed. The recently developed cause of lack of pollen. Productive colonies will forced-air method of removing bees from honey have mostly young bees in mid-November, supers may be adapted for package bee removal. although the colony may be nearly broodless. Bees are sometimes shaken through a funnel di- There should be 50 to 70 pounds of honey and rectly from the comb into the package, or into a 200 to 600 square inches of pollen in three deep ^'shaker box,'^ from which they are later measured bodies of good combs. About one-half the honey into packages. Both of these methods are should be in the top body and the remainder fairly satisfactory and widely modified by individual equally divided between the other two bodies of shippers. the brood nest. Three to six good combs of pollen A recent survey of commercial package bee should be placed near the center of the lower brood shippers shows that most of them ship about chambers. In good colonies most of this pollen will 10 pounds of bees from each colony during the be used before the start of brood rearing in Janu- shipping season. ary. Lack of honey stores can be corrected by feeding sugar sirup. A deficiency in pollen can be partially corrected by giving pollen cakes after References brood rearing starts. (1) DOOLITTLE, G. M. Colonies should reach maximum populations 1915. SCIENTIFIC QUEEN REARING. 126 pp. Amer- at the start of the shipping season. This means ican Bee Journal, Hamilton, 111. that colonies should be stimulated to sustained (2) LAIDLAW, H. H., and ECKERT, J. E. brood production for 8 to 10 weeks before colonies 1950. QUEEN REARING. 147 pp. Dadant & Sons, Hamilton, 111. are to be shaken. The greatest production of (3) PELLETT, F. C. brood per bee is in colonies with approximately 1918. PRACTICAL QUEEN REARING. 103 pp. Amer- 18,000 bees. The greatest colony yield of bees ican Bee Journal, Hamilton, 111. (4) SMITH, J. is obtained by shaking about 4 pounds of bees 1923. QUEEN REARING SIMPLIFIED. 119 pp. A. at regular 10-day intervals so that a popula- I. Root Co., Medina, Ohio. tion level of approximately 18,000 bees is main- (5) tained. At the start of the shipping season the 1949. BETTER QUEENS. 100 pp. Published by strongest colonies may have 35,000 to 45,000 or author. (6) SNELGROVE, L. E. more bees. Such colonies may have 5 pounds of 1946. QUEEN REARING. 344 pp. Purnell & Sons, bees shaken at 10-day intervals, but removing London. BREEDING AND GENETICS OF BEES

By OTTO MACKENSEN, apicuUurist, Entomology Research Divisioriy Agricultural Research Service i

Until recently, progress in inheritance studies the gametes followed by the creation of new com- and breeding better bees has been slow. This binations by fertilization is called segregation and has been due not only to the inability to control recombination. This is the mechanism that brings mating but also to a general lack of understand- about genetic variation between individuals in a ing the breeding biology of the honey bee. After family. Since all the sperms produced by a drone the technique of artificial insemination was per- are identical, no variation between sisters (queens fected, progress accelerated. or workers) is introduced from the drone. With the tremendous gene pool provided by A gene may change so that it has a different the many geographic races of the world, the effect. Such a change is called a mutation. Both possibilities should be great for developing better the mutant and the parent gene are referred to bees. Since much of the beekeeper's basic oper- as alíeles. The position of these genes on the ating expenses are the same regardless of the chromosomes is called a locus. There can be sev- quality of bees used, a more productive bee can eral alíeles of a given locus, each having a different increase his profits rapidly. Greater uniformity effect on the body part or the behavior, which is of an improved stock would help to reduce operat- affected by genes of this locus. ing expenses. A gene may be dominant to another alíele, A single superior strain is not enough. We need completely masking its effect. The other gene is different strains adapted to different environments then called recessive. When the two genes at a and methods of management and also strains or locus are alike, the individual is said to be homo- hybrids bred for specific purposes such as pol- zygous for that gene, and if they are different, the lination of specific crops. individual is said to be heterozygous. The drone, having only one gene of a pair, is said to be hemi- Honey Bee as a Breeding and zygous. A recessive gene must be homozygous to Genetic Subject find expression in queens and workers but always finds expression in the hemizygous condition in The honey bee presents special problems and the haploid drone. also several distinct advantages to the breeder and The two members of a gene pair may have student of genetics. These arise from its partheno- different expressions when they are heterozygous genetic and social nature as well as its mating than when each is homozygous. Genes at different habits and peculiar method of sex determination. loci may also react with each other to cause var- Of course, the basic laws of genetics apply. It is a ious effects. When one considers that there may be matter of understanding how they operate in the thousands of loci with interaction of genes at honey bee. different loci as well as interaction of alíeles, it The honey bee queen and worker develop from becomes apparent that the possibilities for genetic fertilized eggs and therefore their bodies contain variation are tremendous. two sets of chromosomes and genes and are said The honey bee colony is a family consisting of the to be diploid. The drone develops from an un- queen mother, the father represented by the sperm fertilized egg and has only one set of chromosomes in the spermatheca of the queen, all the biparental and genes in all the cells of his body and is said to worker progeny developing from fertilized eggs, be haploid. Eggs and sperms are called gametes by and the maternal drones developing from unfer- the geneticist. tilized eggs. If the queen mated with one drone, In the formation of eggs a reduction division the workers are full sisters in the sense that they takes place so that each egg receives a random all have the same father and mother. Genetically assortment (only one member of each pair) of the they are more closely related than the full sisters genes of the queen. Since the drone has only one of diploid animals, because the sperms of the drone set of genes, no reduction takes place and his are identical. These have been called '^super- sperms are all identical to the composition of the sisters.'^ If the queen mated with several drones, drone himself. The separation of genes going into as is the case in nature, there are as many groups 1 In cooperation with Louisiana Agricultural Experiment of half sisters as there were drones involved in the Station. mating. The normal colony established by natural 68 BEEKEEPING IN THE UNITED STATES 69 mating may then be looked upon as a superfamily Italian queens, reared daughters, let them mate made up of several subfamilies. naturally to common drones, and then used the The fact that the drone develops from an un- resulting colonies to produce pure Italian drones. fertilized egg has several consequences of interest He then reared virgins from an Italian colony and to the bee breeder. Each egg of a queen contains a let them mate naturally with the mixture of Ital- random assortment of her genes, and since her ian and common drones. In the following gener- eggs develop into drones without fertilization, all ation he selected the yellowest virgins, which were genes find expression and these drones show the probably Italian. Thus by rigid selection he could gametic ratio. This is illustrated in figure 1, which change his stock to Italian in two generations and shows the inheritance of a single pair of genes, in one season. This emphasis on selection for the recessive mutation hairless (h), which causes yellow contributed to the development of the almost complete hairlessness over the whole body, American Golden . and its dominant wild-type alíele (H), the gene Parthenogenesis makes possible the equivalent determining development of normal body hairs. of self-fertilization of queens. A virgin queen is This diagram shows a heterozygous queen (h/H) first made to start laying by carbon dioxide treat- mated to a wild-type drone (H). The queen pro- ment, caged outside the hive for a few days to duces two types of eggs (h and ¿?), which without shrink her ovaries and stop egg production, and fertilization develop into drones, one-half of which then is inseminated with sperm taken from her are hairless (h) and one-haK wild type (H). This sons. This makes possible very fast inbreeding if is called the gametic ratio. The drones are, there- such is desired. fore, in effect ^^flying gametes.'^ Since they can be The cells of the drone body, and, therefore, produced in great numbers, this arrangement is the sperm he produces all have the same set of a great convenience to the geneticist. The w^orker chromosomes and genes. This means that one-half bees are all wild type, having developed from eggs of the genetic makeup of the worker bees of a fertilized by sperm carrying the dominant wild- colony is identical provided the mother mated type alíele (H) and are indistinguishable. This is with a single drone. This adds uniformity to the the simplest possible illustration. Of course, there workers of a colony. When a series of queens is are thousands of pairs of genes acting and the mated individually to drones of one queen, dif- queen's eggs represent random assortments of ferences between the resulting colonies are largely these genes, all of which can find expression in the due to the genetic differences between the drones, haploid drone. which represent the gametes of their mother. If we mate a black drone with a yellow queen, When we select from among these colonies, we the worker progeny will be hybrid containing are selecting the best gametes of the queen. When genes from both races and will be intermediate in we rear breeding individuals (queens) from these color, whereas the drones in this colony will be selected colonies, we are breeding descendants of yellow like the mother. This phenomenon is very the desired gametes. The colonies of such a series useful to the breeder. When he wants many drones are, therefore, in a way equivalent to individuals. of a given stock, he can let queens of this stock We can take advantage of these facts in making mate naturally to any type drone and still produce genetic analyses of colony behavior, which is dif- from them the type drones he wants. ficult to study because it is a product of the action When the Italian race was first introduced into of many individuals. First, inbred lines are estab- this country, it was yellower than the common lished by inbreeding and selection for extremes of dark bee. If the beekeeper wished to change com- the characteristic under study. For example, one mon to Italian stock, he first obtained a few might select for high and low preference for aKalf a pollen. The two lines are crossed and drones of a hybrid queen are mated individually to queens of the high and low lines. It is assumed that the maternal inheritance of the worker bees is identical in each of the resulting colonies, since it came from an inbred line, and that differences between colonies are mainly due to the genetic contributions made by the drones, each drone having contributed a random sample of the genes of his mother and thus a study of gene segregation is possible. PROGENY Since some characteristics such as body color can be recognized in the breeding individuals themselves, selection of breeding individuals is FIGURE 1.—Inheritance of recessive gene hairless (h) and its wild-type alíele (H) in cross of heterozygous queen simple. Many of the characteristics of economic with wild-type drone. importance, such as swarming tendency and pro- 70 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE polizing tendency, are determined by the worker tinguished from the white mucus. The average population of the colony as a whole, and for such drone delivers about 1 microliter of semen. characteristics the colony is the test unit. We Queens inseminated with semen from a single cannot breed this unit. We can in effect breed the drone will nearly all lay fertilized eggs initially, queen by using her sons, because, as we have and a few of them may continue to do so for a seen, they represent her germ cells. For breeding year or more. To approximate natural insemi- females we can only use virgin queens that are nation, either two injections of 4 to 6 microliters sisters to the workers of the colony. on separate days or a single injection of 12 micro- liters can be given. Larger injections of up to 18 Control of Mating microliters can be given without injury, but ef- ficiency (percentage of sperms reaching the sper- In recent years the bee breeder has learned much matheca) decreases as the dose increases. Ninety about mating. Queens fly some distance from percent or more of the queens inseminated can their hive to mate. How far is not known. Queens be expected to start laying. are almost invariably flying alone when they have Carbon dioxide initiates oviposition in addition completed their circling of the queen yard and to acting as an anesthetic, but the exposure during disappear into the distance. Using genetically two inseminations is sometimes not enough. There- marked stock, more queens have been shown to fore, an additional treatment for 10 minutes is mate with drones at a distant location than at given on the day following the second insemina- the home location. Just where mating takes tion. When one insemination is made, a treatment place under various circumstances has not been is given on each of the following 2 days. determined. What the bee and queen breeder would like to know is what arrangement he needs to make to Sex Determination get the maximum pure mating. Until better ex- perimental data are available, this goal can prob- Beekeepers have long noticed that after several ably best be reached by saturating the queen yard years of line-breeding, the brood of their strain and the area around it with as many drones of the began to deteriorate in viability, having a ^^spot- desired type as possible. To accomplish this, the ted'' or a '^pepper box'^ appearance. They found drone-producing colonies should be placed from 1 that introducing ^^new blood'^ by purchasing to 3 miles from the mating yard in all directions breeding queens from another bee breeder would as well as in the mating yard. At the same time quickly correct the situation. The deterioration the area should be free of colonies producing was thought to be due to a weakening through undesired drones for at least 5 miles. the general bad effects of inbreeding, which may Since queens mate with several drones, single have been true to some extent, but now it has drone mating in nature is probably extremely rare. been determined that much of this is due to the This fact together with the ever-present chance that peculiar sex determination mechanism of the honey the queen has met with an undesired drone makes bee. It is essential that the bee breeder understand natural mating of limited value to the scientific this mechanism. breeder and geneticist, who must use artificial Dzierzon in 1845 was the first to recognize insemination. This is impractical for the com- that males (drones) are parthenogenetic, that is, mercial bee breeder. develop from unfertilized eggs, and that females The equipraent needed for artificial insemination (queens and workers) develop from fertilized eggs. includes a stereoscopic microscope, a device for This still holds true, but in recent years it has been administering carbon dioxide as an anesthetic, and discovered that this is only a part of a very much the insemination apparatus itself (fig. 2), consist- more complicated system. It has now been well ing of the manipulating stand, holding hooks, established that sex is determined by a series of queen holder, and syringe. The queen is allowed allelic genes at the sex (X) locus, which are desig- to back into the queen holder tube until the end nated xa, xb, xcy etc. Females (queens and workers) of her abdomen projects. She is secured by means are always heterozygous, that is, they contain of a stopper, through which carbon dioxide is flow- two of these alíeles that are different, as, for ing, and placed in the manipulating stand. Then example, xa and xb. A queen of this composition the chitinous plates at the tip of the abdomen are produces eggs of which one-half are xa and one- separated with the holding hooks to expose the half are xb. Since the drones normally found in genital opening. Semen is taken into the syringe the hive develop from unfertilized eggs, one-half from the drone and injected into the oviducts of the sons are xa and produce only xa sperm and while a valvelike fold is held aside with a probe. the other half are xb and produce only xb sperm. To collect semen the drone is made to partially All haploid drones are viable. ejaculate by squeezing the head and thorax, and When an egg such as xa is fertilized by a sperm then the abdomen is squeezed to complete the carrying a different sex alíele such SiS xb, a queen process. The cream-colored semen is easily dis- or worker having the composition xa/xb will result. BEEKEEPING IN THE UNITED STATES 71

FIGURE 2.—Artificial insemination apparatus.

If it is fertilized by a sperm carrying a similar alíele of females among fertilized eggs. It is, therefore, such as xa, this homozygous combination xajxa is more important to avoid inbreeding in bees than in also a male, but it is a biparental male that will be other domesticated animals and plants. eaten by the nurse bees within a few hours after Multiple mating obscures the effect of sex alíeles. hatching. When a queen (xa/xb) is mated to a single If a queen mates only with drones having sex al- drone carrying a different sex alíele such as xc, then íeles similar to her own, the viability of her brood all the progeny resulting from fertilized eggs will is 50 percent, and its poor quality is very notice- have dissimilar sex alíeles {xa/xc and xb/xc) and able. Ordinarily this rarely occurs in nature be- will be viable females (fig. 3, A). Efficiency in the cause drone populations contain several sex alíeles. brood nest will be high and a populous colony will If all her mates carry dissimilar alíeles, the viabil- result. If, on the other hand, she is mated with a ity of her brood is theoretically 100 percent. If she single drone having a similar sex alíele such as xa, mates with a mixture of the two types of drones, then one-half of her progeny resulting from fertil- the viability ranges between 50 and 100 percent ized eggs will be xa/xb females and viable and one- depending on the proportion of the two types of half will be homozygoiis biparental (xa/xa) males sperm. The reduced viability of a multiple mated and be eaten by the bees (fig. 3, B). In addition queen could easily be overlooked if only one or two to wasting the queen's energy, the diploid males of the eight to 10 drones she mated with had simi- live for more than 3 days before removal and thus lar alíeles to her own. Multiple mating and the reduce efficiency in the brood nest and a weak colony results. tendency to mate far from the hive may be evolu- Inbreeding reduces the number of sex alíeles in tionary adaptations to the sex-determining mech- the population and increases the proportion of bi- anism, since both would be favored in natural parental males. Outbreeding brings new sex alíeles selection because they both tend to increase effi- into the population and increases the proportion ciency in the brood nest. 72 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

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PoUmann is native to the Caucasus Mountains The Gene Pool of Russia. In appearance this bee is much like The honey bee population of the world varies the Carniolan bee except that it is not so gray tremendously. Through the ages many rather and has variable amounts of yellow on the distinct geographic races have evolved through abdomen depending on the region from which natural selection. These races with their various it comes. It is a good brood producer, building characteristics, some desirable and some undesir- to strong colonies, but is slow to reach full strength. able, form the gene pool, or raw material, avail- It has the longest tongue of all races. It is gentle, able to the bee^breeder. The four races of honey is calm on the combs, caps honey flat, swarms bees (Apis mellijfera L.) that have been of most very little, propolizes very heavily, and is in- value to the bee industry are briefly described. clined to drifting and robbing. It stores honey The European brown bee or dark bee A. close to the brood nest and winters well. mellifera mellifera L. is native to Europe north The honey bees that were brought to America and west of the Alps. It is black with short legs by the early settlers and spread throughout the and broad body. Colonies of these bees develop land to become the common bee of this country slowly in the spring to populations of medium were very probably of the A. mellifera mellifera strength in the summer and winter. They winter race because their characteristics were similar. well in severe climates, being conservative of The worst characteristics of our early common bee their winter stores. The bees run from the combs were susceptibility to European foulbrood, a tend- very readily when manipulated, are often aggres- ency to swarm and sting, and a strong tendency sive, have a weak disposition to swarm, and are to run off the combs and out of the hive when susceptible to brood diseases and the wax moth. manipulated. The Italian bee imported in 1859 The Dutch or ''heather bee'' is a special strain became popular rapidly and soon was almost of this race that was selected for swarming tend- universally accepted by beekeepers in this country. ency by the beekeepers of the heather areas of It still remains the dominant race. A great deal northern Europe. Each year they could kill the of mixing must have occurred as this transition bees in the heaviest skeps, obtain the honey, took place. then the following spring restock each skep with Both Caucasian and Carniolan bees have been a new sw^arm. imported at various times and are still being prop- The Italian bee A. mellifera ligustica Spinola agated; particularly the former. In addition, is native to Italy north of Sicily. It is smaller several other races have been introduced and and has a more slender abdomen then A. m. propagated on a limited scale until they were mellijera, with yellow bands on the front segments discovered to be inferior. Among these were Cyp- of the abdomen. It is variable in color in Italy. rians, Syrians, and Egyptians, all of which sting It breeds from early spring to late fall, developing excessively. Vestiges of the genes of these races exceptionally strong colonies and using up reserve may still remain in this country. This, then, is stores. It winters with a high population and high the gene pool that has been in the hands of the food consumption. It is calm on the combs, is American beekeeper since importations were gentle, builds comb readily, caps honey white, stopped in 1922 by law to prevent introduction propolizes sparingly, and has a weak disposition of the mite Acarapis woodi (Rennie), the cause to swarm. It tends to rob and drift more than of acariñe disease. other races. The Carniolan bee A. mellifera cárnica PoUmann is native to the Austrian Alps, the northern Balkan Importation and Maintenance of region, and the Danube Valley. Early Carniolan Stock bees had a bad reputation for swarming, but since about 1930 a well-planned breeding program Special methods for importing stock without in Austria has resulted in productive strains danger of introducing acariñe disease have been with little tendency to swarm. This is a slender, developed. One of these is a method of shipping long-tongued bee, with black chitin and short semen. The semen of 30 to 40 drones is placed in gray hairs that give it a gray appearance. Brood a capillary glass tube, 1.8 to 2.0 mm. in diameter, rearing starts early and advances rapidly, build- with an artificial insemination syringe. Each time ing the colony to a strong population early. This the syringe is emptied, the semen is forced down bee reduces brood rearing later in the year, to the bottom of the tube with a centrifuge at winters with a small cluster, and conserves 10,000 r.p.m. When loaded, the tube is sealed on its food. It is disposed to swarming because of a flame as near the semen as possible. When its rapid spring development. It is quiet on the finished, the tube contains about 20 mm. of semen combs, is very gentle, drifts very little, does not and 10 to 15 mm. of air. Properly protected these rob, and is resistant to brood diseases. It is an tubes are placed in an envelope, sent airmail, and excellent bee for an early honey flow. used immediately on arrival at destination. Sev- The Caucasian bee A. mellijera caucásica eral successful overseas shipments have been 74 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

made. Some viable offspring have been obtained the hive and therefore very heterogeneous. Great by using semen handled in this way and stored for variation occurs not only between races but within 68 days at room temperature. them. Our task is to sort out the good qualities Another method consists in transporting eggs, from the bad and at the same time not be misled larvae, and pupae. Mites do not live on these by the effects of the environment. stages. Shipment by air led to only very limited survival because of rough handling, but transport Testing for Selection in an incubator carried as personal luggage was In testing colonies for the purpose of selection, very successful. Queen and drone pupae were the bee breeder must be certain that the test removed from the comb a few days before emer- colonies have an equal opportunity to develop and gence and placed in gelatin capsules, the eggs produce. They should be in one location if possible. were put on wax-paper slips in vials, and worker If single colonies are selected at different locations, larvae 24 hours old were deposited on royal jelly. they should be evaluated and chosen on the basis At destination the queens and drones were intro- of the average performance of the yard they are duced into queenless nuclei, and queens were in. A colony bringing in a medium honey crop in reared in the usual manner from the worker eggs a low producing location may be as good a selec- and larvae. Offspring were obtained after arti- tion as a much higher producing colony in a high ficial insemination. To date, six stocks have been producing location. When groups of colonies are imported rather successfully. to be compared and they cannot all be placed at No doubt these techniques will be used to great a single location, equal numbers should be placed advantage in the future. However, the importation at each of several locations. of stock should be purposeful. The stocks should be A test yard should be established in an area studied in their native land, and only those of where there are some trees and shrubs for bees to proven superiority or with a specific desired use as landmarks in locating their hives rather character should be considered for importation. than in an open field. The colonies should be placed A recent importation of the A, in irregular positions as far apart as possible and mellifera adansonii Latreille to Brazil has appar- equally exposed to the elements. If the colonies ently been regretted because of its viciousness, are placed in regular rows, returning field bees and this bee is spreading throughout the country. tend to drift into the end ones and thus these On account of the techniques involved, this colonies have a distinct advantage. Manipulation method can only be utilized by research should be such that all colonies have an equal laboratories. opportunity to perform. Once imported, the stocks must be maintained Selection is simple for characteristics expressed during their period of immediate use and also for in the queen and drone. However, we have noted future use. Since only a limited amount of breed- that for most characteristics of economic impor- ing material of any one stock can be imported and tance, the colony as a whole is the unit of perform- maintained and since artificial insemination must ance on which selection of breeding individuals is be used, inbreeding and reduced vigor are unavoid- based. A superior colony is chosen on the basis of able. This often leads to greater susceptibility to brood rearing qualities expressed through the disease and to wax moths. Several queens must queen and in the behavior of the workers. be maintained at all times to insure survival of In basing selection on performance of a single the stock. Mutations are much easier to carry, colony, one takes the risk that this high perform- because inbreeding can be avoided and several ance is accidental and will not be transmitted. It mutations can be carried in the same stock. is, therefore, -safer to hold colonies over to the Nevertheless as mutations useful to research next year while the progeny colonies are being accumulate, the maintenance will become a burden tested. Then those whose progeny colonies perform in research laboratories. Many valuable stocks best can be chosen as breeders. have been lost because of neglect due to the more Important qualities to look for in the queen are immediate urgency of other research. There is a longevity and egg-laying capacity. If at all pos- definite need for a center, the principal function sible, a queen should be tested for 2 years. Workers of which is the maintenance of valuable breeding should be observed for such characteristics as and genetic stocks, of both domestic and foreign fast buildup in the spring, wintering ability, origin, for distribution to research workers and willingness to enter supers and draw comb, swarm- bee breeders who can properly use them. ing tendency, good handling qualities such as quietness on the combs, temper, resistance to Breeding Methods disease, and honey production. Such character- istics as length of tongue, size of honey stomach, Selection is only effective when there is heritable wing area, and other physical characteristics are variation. As we have seen, this variation is very secondary. If they are advantageous, they will great in the honey-bee, because it is a naturally automatically be selected along with honey pro- outcrossed species, mating in the air away from duction. Long-tongued bees do seem to gather BEEKEEPING IN THE UNITED STATES 75 more honey from red clover, which has a long If after several years his stock begins to show narrow corolla tube that makes the nectar out of a spotted brood pattern, he may introduce ^^new reach for the shorter tongued bees. blood,'' but before doing so he should first test several promising stocks from other bee breeders to determine whether they have qualities he Methods for the Commercial Breeder wishes in his own stock. At least 10 colonies of The common method of breeding practiced each stock should be compared in a single yard by queen breeders is known as line-breeding. It outside the mating range of his queen mating can be defined as breeding and selection within yard. a relatively small closed population. The bee These are methods that the commercial queen breeder's colonies constitute such a population to breeder can use without undertaking complete the extent that mismatings with drones outside control of mating by isolated mating stations or his stock do not take place. In line-breeding some artificial insemination and extensive records. inbreeding is inevitable. Its main effects are (1) fixation of characteristics so rapidly that ef- fectiveness of selection for good qualities is reduced Hybrid Breeding and the stock loses vigor and (2) the detrimental When one crosses inbred lines, strains, or races results from homozygous sex alíeles. These effects or even some related individuals, one often gets can be largely avoided by using as many breeding hybrid progeny superior to either parent in growth individuals as possible for every generation. and vigor that does not carry through to the The bee breeder will naturally breed from the following generations. This is called hybrid vigor best colonies, and this is good practice. However, or heterosis. The cause is not thoroughly under- there are good and poor methods of breeding the stood, but it is thought to be due in part to selected colonies. He may select one or more combining the dominant genes of the two parents outstanding colonies and rear queens from them, in the hybrid and thus are covered up many det- which are mated to the general run of drones of rimental recessive genes that were homozygous his stock. These are used to requeen his pro- in one of the parents. Many of the genes that ducing colonies, which he observes the following become heterozygous will have their greatest year. From these he again selects several of the favorable effect in that state and thus contribute best colonies to use as breeders for his queen vigor to the hybrid. Since the honey bee is a production and repeats the procedure. highly outcrossed animal in nature, one would If he uses any one breeder for 2 years in suc- expect an accumulation of such genes in its germ cession, some virgin queens will mate with some drones plasm. of their sisters reared the previous year. If only Hybrid breeding seems to be a good solution to one breeder was used the previous year, then all the bee breeder's special problems; however, since virgins will mate with sons of their sisters. Such it requires good control of mating, it is not for the close inbreeding brings together similar sex alíeles average bee breeder. Hybrid breeding programs and reduces the effectiveness of selection by are now being conducted by qualified organi- increasing homozygosity too rapidly. Using a zations. The procedure is very much like that single but different breeder each year would also practiced in the production of corn hybrids. The be too much inbreeding. He should use as many first step is to develop many inbred lines. Then breeders as is practical, the more the better, and these are crossed in various combinations to deter- requeen all his production colonies each year to mine those that produce the best hybrids, which prevent backcrossing to the same source of drones. can then be produced in quantity as long as the The bee breeder can go a step further to prevent component lines are maintained. inbreeding if he is not averse to keeping some Usually four lines are combined to make four- records. He can breed from several outstanding way hybrids. For example, such a hybrid involving queens that are not highly related, for example, lines 1, 2, 3, and 4 might be produced as follows: A, B, C, and D, keep track of their queen progeny, An inbred queen of line 3 artificially mated to and thus establish queen lines A, B, C, and D, drones of line 4 is used as a breeder queen to pro- respectively. In requeening his stock he should duce hybrid (3 X 4) queens. These are allowed to use equal numbers of daughter queens from each mate naturally and are used to produce drones. line, and after testing for a season, select at least Queens of line 1 are then mated to drones of line 2 one from each line to be used as grafting mothers and hybrid virgins (1 X 2) reared for mating to the the following year. The larger the number of queen above-mentioned drones. The cross is then (1 X 2) lines the better. If the bee breeder selects for X (3 X 4). As foundation stock for his hybrid certain physical characteristics, but without sacri- queen production, the queen breeder receives ficing other qualities, to make his stock uniform queens of line 3 mated to line 4, from which he and distinctive, he will be better able to recognize rears many queens and lets them mate naturally mismated queens and can avoid using them for use in his mass drone production. He also re- as breeders. ceives inbred queens of line 1 mated to line 2, from 76 AGRICULTURÍE, HANDBOOK 335, U.S. DEPT. OF AGRICULTURE which he rears the queens (1 X 2) to be mated to for abnormalities and reporting them to a research (3 X 4) drones and sold. Thus with very few laboratory. queens of the foundation stock, he can rear thou- Since characteristics of economic importance sands of queens for the market. The resulting respond to selection, we can assume that they colonies will be headed by two-way hybrid queens, are heritable to at least some extent. Few thorough which will be uniform in appearance, whereas the investigations have been made to determine just worker bees will be four-way hybrids and variable how they are inherited. Some work has shown in appearance unless the color markings of the that tongue length, stinging behavior, and distri- bees of the parent lines were very similar. bution of body pigment are determined by many Comparative tests of hybrids have shown their genes. Preference for alfalfa pollen also appears superiority. Increased productivity of 34 to 50 to be an inherited character, because high and percent over the average of line-bred strains has low preference lines have been selected. Resist- been reported.. Segregation and random mating ance to American foulbrood seems to be a matter in the generations following hybridization are of removal of diseased larvae by the worker bees likely to result in only mediocre colonies. Hybrids and is governed entirely by two recessive genes. are an end product, and to make proper use of As our knowledge of the inheritance of specific them it is necessary to requeen every year. For characteristics grows, the bee breeder^s task these reasons they have been unpopular with should become much easier. some beekeepers. In the present hybrid breeding programs, lines are inbred rapidly with practically no selection. References The highly inbred lines are difficult to maintain while tests are being made, and most of them are (1) ADAM, B. discarded, resulting in a considerable waste of 1954. BEE BREEDING. Bee World 35: 4-13, 21-29, 44-49. effort. Lines bred on a broader basis with selection (2) GoETZE, G. K. L. for desirable characteristics and for different sex 1964. DIE HONIGBIENE IN NATÜRLICHER UND alíeles may in the future produce far better hy- KÜNSTLICHER ZUCHTAUSLESE. Teil I and brids. This type of hybrid breeding and race II. Entomologie 19, 120 pp., and 20, 92 pp. crossing needs to be explored more thoroughly. (3) KERR, W. E., and LAIDLAW, H. H. 1956. GENERAL GENETICS OF BEES. In H. Mutations and Inheritance of Demerec (ed.), Advances in Genetics. V. 8, pp. 109-153. Academic Press, Characteristics Inc., New York. (4) MACKENSEN, O. More than 20 useful mutations have been 1954. SOME IMPROVEMENTS IN METHOD AND SYRINGE DESIGN IN ARTIFICIAL INSEMINA- discovered and an ever larger number are being TION OF QUEEN BEES. Jour. Econ. Ent. maintained by research laboratories. In addition 47: 765-768. to being the working tools of the geneticist, they (5) and ROBERTS, W. C. are useful in bee behavior and other studies. For 1948. A MANUAL FOR THE ARTIFICIAL INSEMINA- TION OF QUEEN BEES. U.S. Bur. Ent. example, the recessive gene cordovan {cd) that and Plant Quar. ET-250, 31 pp. changes the black body pigment to brown with- (6) and ROBERTS, W. C. out reducing vigor has been used as a marker in 1952. BREEDING BEES. lu Insects. U.S. Dept. studies on mating flight range, multiple mating, Agr. Ybk. 1952: 122-130. (7) ROBERTS, W. C., and MACKENSEN, O. and pollination behavior. Eye color mutations 1951. BREEDING IMPROVED HONEY BEES. Amer. have been used to prove the sex-determining Bee Jour. 91: 292-294,328-330,382-384, mechanism. Mapping the loci of these genes on 418-421, 473-475. the 16 chromosomes will have to await discovery (8) ROTHENBUHLER, W. C. 1958. GENETICS AND BREEDING OF THE HONEY of many more mutants. Only three cases of linkage BEE. Ann. Rev. Ent. 3: 161-180. (two gene loci on the same chromosome) have (9) been reported. Since most mutations are recessive 1960. A TECHNIQUE FOR STUDYING GENETICS and since the drone is haploid, mutants are most OF COLONY BEHAVIOR IN HONEY BEES. Amer. Bee Jour. 100: 176-198. easily found by examining the drones. A mutation (10) SMITH, M. V. may appear in a single drone or in many drones 1962. ESTABLISHMENT OF HONEYBEE STOCKS of a single colony. Since worker bees tend not to BY TRANSPORTING IMMATURE STAGES AND tolerate abnormal individuals, many of the muta- SEMEN. Jour. Apic. Res. 1: 19-23. (11) TABER, S. III. tions that occur escape discovery. Beekeepers can 1961. SUCCESSFUL SHIPMENT OF HONEYBEE SE- be very helpful to the geneticist by watching MEN. Bee World 42: 173-176. POLLINATION

By M. D. LEVIN, apiculturist, Entomology Research Division, Agricultural Research Service '

A farmer can provide the best agronomic flowers or at different stages of the life cycle of practices—proper seedbed preparation, fertiliza- the flowers. For such plants, cross-pollination tion, soil moisture, cultivation, pest control, and usually results in better seed or fruit production, harvesting methods—and fail to obtain a bounti- but automatic self-pollination insures that some ful crop if he neglects to provide for pollination. seeds will form even if external agencies fail. Many of his fruit crops, legumes, vegetables, and Some plants produce pollen that is light and oilseed crops depend on bees for assistance in easily blown about by wind. They are referred to reproduction, and therein lies the basis for the as wind-pollinated plants. Others produce pollen most important contribution made by any insect that is heavy and sticky, not blown easily from to agriculture—pollination by bees. flower to flower, but they require other agents, The function of a flower is to produce seed. usually insects, to transfer it. They are usually In a typical flower (fig. 1), the essential fenaale referred to as insect-pollinated plants. parts are the ovary, style, and stigma. The typical male part is the anther. It produces pollen, which contains the male sex cell. These parts Agents of Pollen Transfer may vary greatly in shape or number. The ovary becomes, botanically speaking, the fruit; the style Wind, gravity, and insects are the naain external is a columnar extension of the ovary. At the tip agents of pollination. Wind or gravity disperses of the style, which may be expanded or otherwise pollen of corn and other grasses, coniferous trees, modified, is the receptive stigma. In many plants many deciduous fruit and nut trees, and such crops the stigma has a sticky surface to which pollen as castorbeans, spinach, and beets. grains adhere and on which they germinate. To produce seed, pollen must move from the Bee Pollination anther to a receptive stigma. This is pollination. Many of our fruits, vegetables, legumes, and When pollen adheres to the stigma, it germinates oilseed crops are insect pollinated. Although many and produces a pollen tube that grows through kinds of insects visit flowers and effect accidental the stigma and the style to the ovary. Fertili- pollination, the amount is small. Bees are the most zation takes place in the ovary when the nuclei efficient and only dependable pollinators, because of the male germ cell in the tip of the pollen tube they visit flowers methodically to collect nectar unite with the nuclei of the female germ cell. and pollen and they do not destroy the plant by Pollen transfer from an anther to the stigma feeding on it in the pollination process. Although of the same flower or to the stigma of another various species of bees contribute to the pollination flower on the same plant is referred to as selj- of our crops, an estimated 80 percent of this pollination. The transfer of pollen from an anther pollination is done by the honey bee. of one flower to the stigma of a flower on another Modern agriculture has come to depend greatly individual plant is called cross-pollination. There- on the honey bee to fulfill its pollination needs. fore, cross-pollination in the horticultural sense This species has several valuable qualifications refers to the transfer of pollen between flowers on plants with different genetic constitutions. for this role. In plants adapted for self-pollination, the Beekeepers maintain honey bees at a high pop- pollen grains are often transferred from the anther ulation level in many agricultural areas of the to the stigma by gravity or the movement of United States for the honey and wax they produce. parts within the flower. Some plants are self- Bee colonies can be easily concentrated when and sterile or have floral parts modified or so arranged where needed to satisfy pollination requirements, that the plant is clearly unable to pollinate and by using techniques developed for honey itself. In such cases, pollen has to be transferred production their number can be increased in a by external agents. Many plants combine adapta- relatively short time. The honey bee is adapted to tions for both cross- and self-pollination in different many climates and can successfully revert to its original wild condition in most parts of the world, iln cooperation with Arizona Agricultural Experiment quickly becoming part of the natural reservoir of Station. pollinators. 261-907 0—617^—^—6 77 78 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

■ANTHER •POLLEN GRAIN) STAMEN f I LAMENT

OVARY WALL OR PERICARP^ SPERM ) UNITE TO FORM INTE6UMENTS(DEVEL0F) EGG / EMBRYO iNrOSEEDCOATS) r~-- EMBRYO SAC FLORAL TUBE OVULE (DEVELOPS INTO SEED) NECTARY OVARY {DEVELOPS INTO FRUIT)

RECEPTACLE

PEDUNCLE (DEVELOPS INTO STEM OF FRUIT)

FIGURE 1.-—Longitudinal section of a typical flower.

As a species the honey bee has an extremely wide and nectar to carry the colony through the winter host range. It visits a larger variety of flowers and other dearth periods stimulates it to visit than any other insect. However, an individual bee flowers long after its current needs are met. on a single foraging trip usually confines itself to Honey bees are most active between 60° and one species of plant, avoiding nonproductive 105° F. At temperatures exceeding 90° many interspecific visits. foragers shift to collecting water to cool the hive. Its size and hairiness enable it to accumulate Strong winds slow down flight activity and stop on its body pollen grains that later come in con- it completely if they exceed 25 miles per hour. Hot tact with the floral stigma during subsequent bee winds reduce nectar secretion and indirectly visits. Its natural instinct to store enough pollen reduce flight. Cloudy days usually reduce flight BEEKEEPING IN THE UNITED STATES 79 and cause bees distantly located from a pollen will return to this area repeatedly if it has source to be more easily deterred from foraging adequate points of orientation. On large poorly than bees located nearby. landmarked fields of uniform bloom, they seem The population of the colony affects its response unable to find their way back repeatedly to the to unfavorable climate. A strong colony is less same spot. The effect this has on the foraging sensitive to unfavorable climatic conditions than efficiency of the bee or the pollination of the crop a weak colony and sends out many more foragers is unknown. It could favor cross-pollination. at marginal temperatures. Also genetic differences between bees affect their foraging. Directing Bees to Target Crops Therefore, the crop to be pollinated should be in Because bees do not always visit flowers of a good physiological condition to produce the maxi- particular crop where pollination is desired, many mum amount of pollen and concentration of nectar attempts have been made to direct bees to a crop. to attract bees and compete with other crops or Feeding the bees sugar sirup flavored or scented flowering weeds within the foraging area. Some with extracts of flowers from the field where crops compete well as sources of nectar (white- visitation is desired does not increase pollination clover, alfalfa, cucurbits), others as pollen sources in the field. Spraying the field with a sugar solution (safflower, red clover), and still others attract both is also valueless. Some tree fruit pollination has nectar and pollen collectors (sweetclover, crucifer- been achieved with pollen dispersal by bees from ous crops). hive-entrance inserts. However, bees will not In many instances the plants competing for bee continue to visit a crop that does not yield nectar visitation are weeds rather than agricultural or pollen, and no technique has ever resulted in plants. Most weeds can be eliminated from an area sustained visitation. by chemical or cultural treatments, although this may not be practical or even desirable, because Progress is being made in breeding plants more weeds often serve as a needed food source for the attractive to bees and in breeding bees that show bees. When other agricultural crops are involved, preference for a particular crop. Recent research the grower can sometimes reduce competition by has demonstrated conclusively that the preferences timing the bloom period of the target crop so the shown by bees for certain crops or even varieties pollination period will miss that of the competing can be intensified by selective breeding. The crop. If this is impractical, he may need enough bees ^^tailormade bee'' is not too far off. to saturate both the competing crop and the crop to be pollinated. This last is the method most often Pollination of Agricultural Crops attempted for overcoming plant competition. The following list includes seed and fruit crops grown in the United States that require insect Pollinator Competition pollination or that show definite increases in yield Occasionally where wild pollinators are abun- or quality as a result of the pollinating activity dant, competition may develop between them and of bees. Not included are ornamentals, medicinal honey bees. Beekeepers in such situations complain plants, and most spices, many of which are also that the wild bees pollinate the flowers before the insect pollinated. honey bees can collect a crop of honey. On the other hand, growers sometimes object to the Leguminosae : Leguminosae—Continued Alfalfa Lespedeza (bush) presence of honey bees in areas where wild bees are Clovers : Lima bean i effective pollinators. Neither complaint has been Sweetclovers: Trefoil verified experimentally. Annual Vetch (hairy) white Annual Liliaceae (Allium) : Forager Distribution yellow Garlic Hubam Leek The first orchard pollination recommendation Sour Onion stated that bees within 2 to 3 miles would supply True clovers: adequate pollination. We now know that bees Alsike Malvaceae: Cotton 1 pollinate most effectively within 200 yards of Arrowleaf Ball Kenaf their hives. Therefore, to achieve the most effi- Berseem Okra 1 cient distribution of foragers, the colonies should Crimson be distributed at intervals of 175 to 200 yards. Persian Cruciferae: Red Broccoli Also beginning foragers work close to the hive, Strawberry ^ Brussels sprouts then disperse more widely with age and orien- White, Cabbage tation. Therefore, it is essential that a pollinating Ladino Cauliflower colony should have plenty of these beginning Guar Chinese cabbage Horsebean Collard foragers. Kale Individual foragers tend to limit their work area (broadbean) i on any one trip to a few square yards. They 1 Pollination beneficial. 80 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

Cruciferae—Continued Fruits and nuts: lated during World War II by the need for in- Kohlrabi Almond Mustard Apple creased legume seed production. Since 1940, Radish Apricot research and agronomic practices have largely Rape Avocado centered around the use of honey bees as Rutabaga Blackberry- pollinators of legume seeds. Turnip Blueberry Cherry After scientists in the U.S. Department of Cucurbitaoeae: Chestnut Agriculture proved that high populations of honey Cantaloup Coffee 1 bees combined with insect control could greatly (muskmelon) Cranberry increase alfalfa seed production, the practice Citron Dewberry Cucumber Gooseberry of renting bees for alfalfa seed production rapidly Honeydew melon Grape expanded. The recognition of grower benefits from Persian melon Huckleberry renting bees carried over to other crops, and today Pumpkin Macadamia nut paying for pollination by honey bees is a common Squash Mandarin Watermelon Mango practice in many parts of the United States. Growers of tree fruits, legume and vegetable Umbellif erae : Nectarine Carrot Orange ^ seeds, all kinds of cucurbits, oilseeds, and many Celeriac Passion fruit other crops rent bee colonies to distribute pollen Celery Peach Pear on the large scale required by modern agriculture. Parsley Bees are usually rented only where pollination Parsnip Persimmon, native Plum problems are extreme; far more incidental polli- Miscellaneous : Prune nation is supplied free by colonies managed by Asparagus Raspberry beekeepers for honey production. Buckwheat Strawberry Flaxi Tángelo Lotus Tangerine Specific Crops Safflower ^ Tung Sunflower Legumes 1 Pollination beneficial. Most legumes are attractive to bees and adapted to benefit from their visits. Many of the clovers The 1963 farm value of the crops included in produce seed abundantly when bee pollinated. this list that required bee pollination amounted to Alfalfa.—Alfalfa is one of the legumes that approximatily $1 billion. The farm value of the presents several unusual pollinating problems. additional crops benefited by bee poUination was Before pollination can take place, the sexual approximately $6 billion. column, containing the stamens and pistil, must When hybrid seed and fruit-production methods be released from the keel petals confining it. This are developed for other crops, the list will doubtless irreversible ^'tripping'^ occurs when a bee inserts increase. its tongue into the throat of the flower. When the Pollination Practices sexual column is released, it strikes the underside of the bee's head and deposits pollen. At the same History time it contacts pollen left on the bee from a prior The pollination of plants by bees was first des- flower visit. Since most lines of alfalfa are relatively cribed by Sprengel in 1793. However, fruit pol- self-incompatible, the pollen must come from a lination requirements were not intensively studied genetically different plant to cause cross-polli- until M. B. Waiters publication on Pollination nation and seed production. of Pear Flowers in 1895. Most of the research on Many efforts have been made to contrive me- pollination in the early 1900's was conducted by chanical devices to trip alfalfa flowers. None have horticulturists, who became aware ol the need for been successful, mainly because mechanical trip- pollen vectors and cross-pollination to obtain com- ping without pollen distribution results in self-pol- mercial crops of fruit in the many varieties being lination and low seed set. Furthermore, self- developed. They recognized that interplanting pollinated seeds produce less vigorous plants. of compatible varieties should be based on knowl- Honey bees visit alfalfa flowers readily, but edge of bee foraging behavior. Renting of colonies many of them learn to collect nectar from the side for apple pollination started about 1910. This of the flower and avoid the tripping mechanism. practice grew rapidly in the next 20 years. Now Accidental tripping by high populations of *'nectar thousands of colonies are used each year in collectors'' can account for large seed yields. Also Washington, New York, and other apple- a certain percentage of the bees always will collect producing States. pollen from alfalfa. At certain times and under The need by horticulturists for this information certain conditions these ''pollen collectors" make encouraged apiculturists to study bee foraging in up a high percentage of the total population. This orchards during the early decades of the century. is most likely to occur in the hotter, drier regions Their research provided a good background for of the Southwest—probably from lack of compe- the expanded pollination research that was stimu- tition by other pollen-producing plants. BEEKEEPING IN THE UNITED STATES 81

About 38 million flowers must be tripped and Deciduous Tree Fruits.—In tree fruits the degree cross-pollinated to set a 500-pound-per-acre seed of self-fertility varies tremendously, even within crop. In California and Arizona, two to three colo- species. In apples and pears, differences in seH- nies per acre are used. They supply a ñeld popula- fertility occur between years or locations even tion of two to three bees per square yard, 5 to 20 within the same variety. percent of which are pollen collectors. In northern Most fruit trees are propagated by grafts, and Utah, where pollen collectors rarely exceed 1 to 3 an orchard of one variety therefore is genetically percent, eight to 10 bees per square yard are re- one plant. No matter how efficiently bees carry quired. Farther north and into Canada, high yields pollen between trees of the same self-sterile vari- of seed from honey bee pollination are rare. In the ety, a commercial crop cannot be set. In addition, Midwest, alfalfa poUination is largely left to not all varieties are compatible with each other. chance. Seed production per acre in this area is Therefore, orchards must be carefully planned to low. provide adequate distribution of good poUenizer Red Clover.—Red clover competes well with varieties, which must (1) bloom in synchrony, other crops as a pollen source, and pollen-collect- (2) produce pollen abundantly, (3) be equally ing honey bees visit the flowers freely. Because attractive to bees, (4) bloom heavily each year, of the corolla tube length, nectar collectors have and (5) produce fruit commercially useful or easily difliculty reaching the nectar unless it is especially distinguishable. abundant. Usually two or more colonies per acre Much of the recent research has shown the are adequate. limited foraging areas of bees on fruit trees. This Other Legumes.—Alsike, whke, and crimson behavior pattern must be considered when plac- clovers, sweetclover, and trefoils require bees for ing poUenizer trees for the best cross-polhnation. pollination, but are visited freely. Colonies store An optimum poUenizer arrangement is to have surplus honey crops from them even at a high pop- every main variety tree adjacent to a poUenizer ulation density. In contrast. Ladino clover re- tree. Planting poUenizers in every third space in quires two or more colonies per acre to supply the every third row will achieve this. Since such a two bees per square yard required for good pol- scattering of individual trees would interfere A^ith lination. But because it is a poor nectar yielder, efficient orchard management, growers prefer to colonies at this density will have difficulty main- plant all of every third row to the poUenizer taining themselves. variety or alternate two rows of each variety as a Hairy vetch is rather attractive to bees. Small satisfactory compromise to provide good cross- fields surrounded by uncultivated land are usually poUination. well pollinated, but honey bee colonies must be im- All varieties of sweet cherries, Japanese plums, ported for large acreages. and avocados require cross-pollination. Most varieties of apples, pears, and European plums and Nuts and fruits prunes also require cross-pollination, as do some varieties of grapes, citrus, apricots, and peaches. Almonds require large numbers of bee visitors Some kinds, for example peaches, may not re- for adequate pollination. The almond grower wants quire cross-pollination, but they still need bees to the heaviest set possible, because a heavy set transfer their pollen to the stigma. means nuts with smaller kernels, which are in In most cases, one colony per acre and the greatest demand. In California, where most of colonies placed in groups of 15 to 20 in the center the almonds in the United States are produced, of each 15- to 20-acre block wiU provide adequate the pollinating season frequently starts in Febru- bee visitation. About 300 to 350 yards or one- ary. Cool cloudy weather at that time often in> fifth mUe between groups wiU give this spacing, hibits bee fiight to only an hour or two each day. which is a compromise between the orchardist's Strong, vigorous colonies capable of supplying ideal of individual colonies weU scattered through suflicient foragers are sometimes difficult to ob- the orchard and the beekeeper's ideal of aU the tain. This emphasizes the need for at least two colonies in one location. populous colonies per acre, which many beekeep- With pears, which compete poorly for the serv- ers cannot easily supply so early in the season. ices of bees because of low nectar-sugar con- Also the colonies should be distributed in the centration, at least two colonies per acre are groves, which is difficult after heavy or continued necessary. Pears are unusual in one other respect. rains. Therefore, supplying bees for almond polli- One of the most popular varieties, the Bartlett, is nation presents a bee-management problem. In considered self-unfruitful in the Eastern United addition, many compatibility problems occur States and in most other areas where it is grown. among the many varieties grown, and alrnond In spite of the usual recommendations that groves must be carefully planned with this in poUenizers be interplanted with Bartlett trees, mind. most of the orchards in California are in solid Except for macadamia nuts in Hawaii, most plantings, and yet commercially profitable yields other nuts are self-fertile or wind pollinated. are produced. Horticulturists have found that 82 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

most of the fruit obtained under these conditions some varieties as much as 25 percent. Most varieties is due to vegetative parthenocarpy (seedless fruit are beneflted to some extent by bees. This benefit development without the immediate stimulation can be in higher yields, earlier set, or better grades. of pollination). Cross-pollination usually gives For many years plant breeders have been inter- higher j^ields of seeded fruit. ested in the possibilities of increasing production Distribution of colonies for the pollination of by utilizing hybrid vigor in cotton. When hybrid all these tree crops should be synchronized with cotton production is developed on a large scale, cultural and pest-control programs. This can be only honey bees will be able to supply the vis- difficult because of the unfavorable weather that itation required. often prevents placement or removal of colonies at The long blooming season of cotton, the re- the appropriate time. luctance of honey bees to collect and utilize cotton Berry Fruits.—Horticulturists have long known pollen, and the frequent lack of pollen sources in of the differences in self-fertility between varieties of many of the areas where cotton is grown combine cane fruits and other berry fruits. In the older to create problems in maintaining colony popula- literature they frequently mentioned the necessity tions at a level suitable for continued pollination. for poUenizer varieties. Rarely, however, was the This is a situation where a suitable pollen sub- need for insect pollinators, recognized. Perhaps stitute, the development of a cotton pollen- this was a reflection of the abundance of natural collecting strain of bees, and techniques for pollinators in the small acreages at the turn of the maintaining brood rearing are necessary. century. With specialized agriculture, increased Recent work indicates that bees can further acreages, and associated side effects came the benefit cotton by removing the extrafloral nectar recognition that many of the berry fruits were on which harmful insects must feed to lay their dependent on insect pollinators. normal number of viable eggs. Most cranberry and blueberry growers now Saffiower.—This close relative of the common use honey bees for pollination. Many varieties of thistle is attractive to bees and a good source of blackberries, raspberries, and strawberries benefit nectar and pollen, although saffiower honey is from bee pollination, but these crops are fre- a low quality table honey. Recently the acreage quently grown in areas where resident populations of this crop has been increased sharply. of bees are available to pollinate relatively small The value of bees to saffiower depends on the acreages. Where larger acreages are grown, bees degree of self-fertility of the particular variety. should be provided for maximum production of Honey bee activity in saffiower is being studied high quality berries. in anticipation of the need for bees as cross- Many of the common grape varieties grown in this pollinators of hybrid saffiower. Only a few thou- country are self-fertile and do not benefit from bee sand colonies are rented for saffiower pollination. visitation. However, some varieties require bees Flax.—The blossoms of flax are not strongly and some yield better as a result of bee visitation. attractive to bees, and pollination by bees is gen- Citrus.—^The nectar of most citrus flowers is erally not considered by growers. Actually bee polli- extremely attractive to bees, and a citrus tree in nation will increase seed production, and growers flower without bee visitors is rare. Because of the should encourage bee visitation to their flax crop. large numbers of flowers on the tree, the number of Sesame.—There is little information on pollina- bee visits per flower during the period of pollen tion of sesame. It is highly attractive to bees and receptivity may be small. benefited by bee visitation. Most varieties of citrus have been considered to Soybeans.—Many varieties of soybeans are be parthenocarpic or self-fruitful without the aid grown under various conditions throughout the of bees. But the role of bees in citrus pollination United States. In some places bees work soybeans has not been sufficiently studied to eliminate their heavily, and particularly in Arkansas good crops consideration completely. In the Algerian (Clem- of soybean honey are reported. Evidence is build- entine) tangerine and the tángelo (a cross between ing up that bees are of value at least to some grapefruits and tangerines), poUenizer varieties varieties under some conditions. More work on and honey bees to carry the pollen are as important the pollination of this highly important crop as in any other self-sterile fruit variety. The should be done. association of pollination with seed production Sunflower.—Insect pollination is essential for and of seeds with size of the popular Valencia seed production of sunflower. The acreage of this orange and several other citrus varieties indicates crop seems to be on the increase, and cultural that bees are highly beneflcial to them. programs that do not include insect pollination Oilseed crops may result in a crop failure. Cotton.—The principal varieties of cotton grown in this country are largely self-fertile. But the Plant breeders need to be aware of the possible structure of the flower is well adapted to cross- increases in yield that bee activity can stimulate pollination, and bees can increase production of in those varieties in which self-fertility is low. BEEKEEPING IN THE UNITED STATES 83

The value of bees when varietal selections are for each seed produced. Honey bees are the main made in small plots well visited by bees is easily pollinators. If pollination is not adequate, no overlooked. Yields may dwindle when these marketable fruit is harvested. varieties are grown in large acreages with High bee populations are associated with more inadequate pollination. seeds per fruit and earlier, sweeter, larger, and In many cases the relationship of honey bee more fruit (fig. 2). Dependable production of top foraging to production of many kinds of oilseeds quality fruit should not be left to chance. still remains an unexplored, potentially fruitful Commercial recommendations call for one field' of investigation. colony per acre. This should provide the equiva- lent of one bee for each 10 perfect fiowers, or at Vegetable seed crops least 12 bee visits per flower during the forenoon. Carrots are attractive to a wide variety of If not, more colonies may be needed. In cooler insects including honey bees. However, only areas bee visits are probably of value over a honey bees can be provided in sufficient numbers longer period; therefore, the number of colonies consistently, and carrot seed producers are necessary will vary with the location of the fleld. usually conscientious about supplying colonies. Some honey bees range far afield from their Nectar and pollen collectors are equally effective colony. Melon fields without a few bees present pollinators. Good pollination in carrots not only are rare. Visits by such bees can result in a high insures optimum yield but affects seed size and percentage of fruit set from the fewer flowers germination. Inadequately pollinated flowers pro- they visit. This may explain why some growers duce fewer and objectionably oversized seeds with obtain moderate yields without purposely placing lower rates of germination. honey bee colonies in their fields. Such inade- Onions and closely related crops are self-fertile, quate chance visitation usually results in a reduced but require visitation by bees or other insects yield, a later crop, and a high percentage of culls. for either cross- or self-pollination. Many of the Watermelons and cucumbers have pollination insects attracted to onions are small and not requirements similar to cantaloup, and successful hairy enough to satisfactorily transfer pollen. growers usually see that colonies are near the Nectar-collecting honey bees are less effective crop. The optimum pollinator populations or pollinators than those collecting pollen because colonies per acre needed are undetermined. of their stance while working the florets. When Where watermelons are grown on large acreages, locating onion seed fields, all attractive sources honey bee colonies are usually provided at the of pollen like sweetclover or safflower should be rate of one colony per acre. Recent work in Florida avoided. Production of hybrid onion seed has showed that this should supply the minimum of expanded greatly since 1940, and honey bees are eight visits per flower necessarv to set a commer- cial crop. regularly supplied for pollination. Doubtless many more colonies per acre are Most tomato varieties are attractive to pol- needed for cucumbers than watermelons. Many linators. Honey bees work the newer varieties acres of cucumbers are grown in greenhouses in and probably contribute to their pollination. Bee the Eastern States, and providing bees for them activity has proved beneficial to tomatoes grown has become an important beekeeping specialty, in greenhouses. The development of hybrid toma- with problems peculiar to keeping bees in toes will require bee pollination. confinement. Other vegetable seed crops that require visits Squash and pumpkins, too, require bee transfer by bees include most of the cruciferous plants such of pollen, but since these are native to America, as cabbage, cauliflower, and radish, many legumes several species of native bees as well as honey such as beans, and such other crops as peppers, bees visit them. However, wild bees are usually asparagus, and celery. Where acreages of these inadequate where acreages are large, and again crops are relatively small, wild pollinators or we must rely on the honey bee. resident populations of honey bees may be ade- Many of the examples mentioned point up the quate. However, many growers of these crops deficiencies of pollination information on our are insuring themselves against insufficient levels crops. In many cases the importance of bees is of pollinators by renting colonies of honey bees, recognized, but detailed information on the quan- especially where acreages are large or crops are titative and biological aspects is either localized competitive. or completely lacking. Cucurbits Practical Aspects of Current Practices Cantaloups produce hermaphrodite (female) During the last few decades, honey bees in and imperfect (male) flowers. Both produce nec- the United States have increasingly been used tar. Female flowers have both ovaries and pollen- as an additional ^^tooP' in the production of fruit bearing anthers. Male flowers bear anthers only. and seed crops. As is necessary with all coni- At least one pollen grain must reach the stigma plicated tools, skills must be developed in their 84 AGEICULTUHE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

15-18 NUMBER OF BEE VISITS

BN-30045 FIGURE 2.—Effect of bee visits on cantaloup fruit set, Yuma, Ariz., 1963. use. We must learn what bees can do, their limi- pollinate the earlier flowers. However, for others, tations, and the factors that affect their use. As increased numbers of colonies should be moved a biological organism, the honey bee is much into the field as flowering progresses. more susceptible to factors beyond our control than are most tools, and only recently have we Moving even developed techniques for studying some of The logistics of moving large numbers of col- these factors. onies quickly and frequently for pollination service stimulated the invention of loading equip- Colony Standards ment and other new techniques. Colonies now Information on the number of colonies per are merely smoked, picked up by hand or hoist, acre needed for many crops is lacking. Further- and stacked with open entrances on large flat-bed more, apiculturists are aware that the colony trucks. They are roped down, usually covered is not a standard unit. Greater percentages of with a wet burlap tarpaulin or plastic screen, the colony population fly from ^'strong'' colonies and moved to their new location. than from weak colonies and under more adverse conditions. Flight activity and pollen collection Water, Shade, and Windbreaks are correlated with the amount of brood rearing. Every effort should be made to provide a de- Therefore, evaluations of colony strength or pendable water source as close as possible to the quality are based on population and brood. bees. Time spent carrying water or cooling hives Some beekeepers supply a one-story colony is time lost to pollination. Air temperatures ex- containing bees and brood on only five to seven ceeding 90° to 100° F. greatly increase the need frames. However, pollination colonies need to be for water and can upset the normal work in the populous and vigorous at the beginning of the hive. Shade greatly reduces the heating effects of flowering season and should occupy two or more the sun. Water needs are reduced under shade, standard Langstroth hive bodies and have 600 to although not eliminated, and the bees can be more 800 square inches of brood in all stages. productive. Wherever possible, colonies should be placed under at least partial shade even in areas Timing where high temperatures are not often a problem. The best time to place the colonies in the field Colonies pollinating very early crops are vul- or orchard is not the same for all crops. For some nerable to chilling winds, which cut down flight crops, the colonies should be taken to the crop as activity. Windbreaks effectively reduce these soon as flowering starts so that the bees can detrimental effects. They also cut down on drifting BEEKEEPING IN THE UNITED STATES 85 of bees between colonies, which is frequently A strong, prosperous, beekeeping industry is troublesome in windswept locations. needed to supply the pollinators demanded by modern agriculture. Until beekeepers receive full Costs, Contracts, and Middlemen value for the pollination services of their bees, When beekeepers undertake pollination work, they must rely on income from honey and wax they encounter new business problems. When a production, and their problems in doing this are of grower pays for the use of bees, both he and the importance to all agriculture. beekeeper assume responsibilities. Difficulties arise The increasing use of honey bees as a tool in when responsibilities are not clearly stated in a agricultural crop production has greatly augmented written and binding contract. Paying for bees is the value of the beekeeping industry to the econ- a new concept to many growers, and some- omy of our country, and the dependence of times they hesitate to accept the associated agriculture on honey bees will continue to increase. responsibilities. Since some contracts include the possibility of a partial honey crop as part of the fee, the beekeeper manipulates the colonies for maximum honey References production, which should be of secondary concern. Ideally, the pollination fee should allow the bee- (1) BOHART, G. E. 1960. INSECT POLLINATION OF FORAGE LEGUMES. keeper to concentrate on pollination efficiency and Bee World 41: 57-64,85-97. ignore honey production. This should ultimately (2) and TODD, F. E. be most profitable to both grower and beekeeper. 1961. POLLINATION OF SEED CROPS BY INSECTS. In 1967, pollination fees range from $1 per In Seeds. U.S. Dept. Agr. Ybk. 1961: 240-246. colony, which scarcely pays for the cost of moving, (3) FREE, J. B. to $15, which should compensate for a potential 1960. POLLINATION OF FRUIT TREES. Bee World honey crop. Probably more colonies are rented for 41: 141-151, 169-186. $4 to $5, and this is rarely enough to permit a (4) GRIGGS, W. H. 1953. POLLINATION REQUIREMENTS OF FRUITS AND beekeeper to ignore honey production. NUTS. Calif. Agr. Expt. Sta. Cir. 424, Most successful pollination services are con- 35 pp. tracted through a third party. Business-oriented (5) LovELL, H. G. beekeepers, seed companies, commodity coopera- 1963. THE HONEY BEE AS A POLLINATING AGENT. In Grout, R. A., ed.. The Hive and the tives, or enterprising individuals not connected in Honey Bee. Rev., 556 pp. Dadant & any other way with crops or bees are more likely Sons, Hamilton, 111. to be successful ''third-party contractors.^' (6) MEEUSE, B. J. D. A pollination service, as such, can '^selF' 1961. THE STORY OF POLLINATION. 243 pp. Ronald Press Co., New York. growers on the benefits of efficiently utilized bees, (7) RIBBANDS, C. R. contract with beekeepers, coordinate the logistics 1957. THE BEHAVIOUR AND SOCIAL LIFE OF HONEY of moving colonies, conduct an impartial inspec- BEES. 352 pp. Dover Publications, Inc., tion service, and in other ways protect the interests New York. (8) TODD, F. E., and MCGREGOR, S. E. of both parties. In many ways the real future of 1960. THE USE OF HONEY BEES IN THE PRODUC- pollination lies in the development of efficient TION OF CROPS. Ann. Rev. Ent. 5: 265- ^'third-party contractors'' to run pollination- 278 (9) WAITE, M. B. ' service businesses. 1895. POLLINATION OF PEAR FLOWERS. U.b. Dept. Agr. Div. Veg. Path. Bui. 5, 86 pp. BROOD DISEASES OF HONEY BEES

By H. SHIMANUKI, microUologist, Entomology Research Division, Agricultural Research Service

Bee diseases result in losses amounting to mil- Workers, drones, and queens are equally suscep- lions of dollars annually. In 1963, various States tible to American foulbrood. Only spores are ca- spent an estimated $1 million in apiary inspection pable of inciting the disease. The spores are ex- for American foulbrood. In the same year the value tremely resistant to heat and chemical agents. of colonies destroyed under State laws to control A severely infected American foulbrood comb American foulbrood exceeded $470,000. European has a mottled appearance due to a mixture of foulbrood, sacbrood, nosema disease, septicemia, healthy capped brood and diseased or empty cells paralysis, and other diseases and enemies are formerly occupied by diseased brood. The cappings estimated to cause losses in excess of 10 times that of diseased cells appear moist and darkened. The of American foulbrood. In addition to these figures, convex cappings found on cells of diseased larvae bee diseases cause considerable losses, which are become concave as the disease progresses. Another difficult to estimate, through reduced honey and symptom commonly associated with the disease beeswax production and crop pollination. It is is the punctured capping. Larvae are susceptible apparent, therefore, that both beginning and ad- to American foulbrood only when they are less vanced beekeepers should learn to recognize and than 3 days of age. A healthy larva has a glisten- control bee diseases. ing, pearly white appearance. Normally it begins A guide for diagnosing brood diseases of honey development curled on the base of the cell. As it bees is given in table 1. grows, it elongates to the full length of the cell. It is in the elongated position that the larva or pupa American Foulbrood dies. As the infection progresses, the larva or pupa changes to creamy brown and eventually becomes American foulbrood disease occurs throughout dark brown. The remains become ropy and can be the world where honey bees are kept. About 5 per- drawn out as threads of an inch or more. Also a cent of all colonies inspected in the United States very characteristic odor develops at this stage de- are found to be infected. scribed by some as '^typical gluepot odor.'' Bacillus larvae White, the causative organism of The remains of diseased brood finally dry out to American foulbrood, is a spore-forming bacterium. form scales, which adhere strongly to the sides of

TABLE 1.—Differentiating characters oj brood diseases

Characters to observe American foulbrood European foulbrood Sacbrood

Appearance of brood Sealed brood; discolored, Unsealed brood; some sealed Sealed brood; scattered cells comb. sunken, or punctured brood in advanced cases. with punctured cappings, cappings. with discolored, sunken, or often with 2 holes. punctured cappings. Age of dead brood Usually older sealed larvae or Usually young unsealed lar- Usually older sealed larvae; young pupae; occasionally vae; occasionally older occasionally young unsealed younger unsealed larvae. sealed larvae. larvae or young pupae. Color of dead brood Dull white, becoming, yellow. Dull white, becoming yellow- Grayish or straw colored. light brown, coffee brown ish white to brown, dark becoming brown, grayish to dark brown or almost brown, or almost black; black, or black; head end black. often unevenly colored. darker. Consistency of dead Soft, nonsticky, becoming Watery to pasty; rarely Watery and granular; tough brood. sticky to ropy. sticky or ropy. skin forms sac. Odor of dead brood Slight to pronounced typical Slight to penetratingly sour. None to slightly sour. odor. Scale characteristics Brittle; rough texture; lies Rubbery; smooth texture; Brittle; rough texture; lies flat on lower side of cell; usually twisted in cell; flat on lower side of cell; adheres tightly to cell wall; does not adhere tightly to does not adhere tightly to head lying flat; fine, cell wall. cell wall; head prominently threadlike tongue of dead curled up. pupae adheres to opposite wall of cell. BEEKEEPING IN THE UNITED STATES 87 the cells. If death occurs in the pupal stage, the Larvae die of sacbrood in capped cells in the mouth parts may adhere as a fine thread to the op- elongated position. As the disease progresses the posite side of the cell. This is a positive symptom larval skin forms a sac, which separates itself of American foulbrood disease (figs. 1 and 2). from the pupal skin. Between these two layers of The infection can be transmitted to a larva from skin is an accumulation of ñuid. The outer skin nurse bees or from spores remaining in the bottom toughens and as a result the larva can be picked of the brood cell. Early detection of the disease is up in its entirety without the release of the fluid. helpful in preventing further spread. A colony The larva changes from pearly white to off- that is weakened by American foulbrood may be white, then brown, and finally almost black. The robbed, and the robber bees inadvertently carry head end is usually the darkest. It usually curls honey containing spores of B. larvae to healthy up from the cell fioor. A loosely adhering scale is colonies. formed from the larval remains. No odor is attributable to the sacbrood virus although heavily infected combs may have a yeastlike odor European Foulbrood (fi^.4). Like European foulbrood, sacbrood is most com- In some areas European foulbrood poses a more monly found in the spring. No chemotherapeutic serious threat to beekeepers than American foul- agent is effective against sacbrood. Fortunately brood. This disease is serious because it occurs colonies appear to recover spontaneously from the most frequently at the time that colonies are build- disease. ing their peak populations. The cause of this disease is Streptococcus pluton (White), a nonspore-forming bacterium. Other Other Brood Diseases bacteria commonly associated with the disease are Bacillus alvei Cheshire & Cheyne and Bacterium Chalk brood and stonebrood are both caused by- eurydice White. fungi. Chalk brood is caused by Ascosphaera apis Superficial examination of diseased combs Olive & Spiltoir. This disease is characterized by shows the same mottled effect and puncturing as the chalky white appearance of the larvae after seen in American foulbrood. In most cases death death. To date there has been no report of this usually occurs in the larva stage. Workers, drones, disease in the United States. and queens are equally susceptible to European Aspergillus flavus Link is usually isolated from foulbrood. bees that have stonebrood. This disease is unusual . Larvae that die from European foulbrood are in that it infects both brood and adults. Bees dying found in a variety of positions. Some are in a from this disease form mummies. The fruiting curled stage and others elongated. The normal bodies of the fungus make the infected bee appear pearly white appearance of a healthy larva changes yellowish green or brown. to a dull white, then yellow, and finally brown. Purple brood is a nutritional disease of larvae Ropiness and sour odor are caused by secondary and pupae. It is believed to be caused by nectar invaders. The elasticity of the ropy material is or pollen from Cyrilla racemifloral L., also called less than that associated with American foulbrood. southern leatherwood, black titi, red titi, summer The tracheae appear as fine silvery tubes im- titi, and he-huckleberry. This problem exists only mediately below the skin, especially as the larvae in the Southern States where southern leatherwood tvirn brown. This symptom is highly charac- is found. Diseased larvae and pupae are purple. teristic of European foulbrood. Loosely adhering Other conditions that mimic contagious diseases scales also differentiate this disease from American are chilled and starved brood. Chilled brood is foulbrood (fig. 3). usually attributed to insufficient bees to keep the European foulbrood can be transmitted by brood area warm. The same cause can also result contaminated food, stores, and equipment. The in starved brood. However, starved brood is gen- disease is usually most serious in the spring and erally caused by insufficient nectar. At times wax clears up during the summer when nectar and moth damage may cause brood to appear diseased. pollen are abundant. However, outbreaks of European foulbrood in the late summer are not Disease Prevention and Control unusual. A beekeeper can spread bee diseases by transfer- Sacbrood ring combs from colony to colony to equalize stores or numbers of bees. This practice can lead to Death of a colony by sacbrood is rare. However, serious consequences if disease is present. Records because of the similarity to other diseases, the kept on the source of combs are helpful so that a beekeeper should learn to distinguish sacbrood beekeeper can trace a possible source of infection. from the more serious diseases. The etiologic Honey from colonies with American foulbrood agent in sacbrood is a virus. should never be fed to healthy colonies. 88 AGRICULTUKE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

I J

FIGURE L—-Honey bee larvae killed by American foulbrood, as seen in cells: A, Healthy larva at age when most of brood dies of American foulbrood; B-F, dead larvae in progressive stages of decomposition (remains shown in F is scale) ; G, longitudinal view of scale. BEEKEEPING IN THE UNITED STATES 89

FIGURE 2.—Honey bee pupae killed by American foulbrood, as seen in cells: A-C, Heads of pupae in progressive stages of melting down and decay; D-E, scales formed from drying of dead pupae. In B, C, and E tongue is shown adhering to roof of cell. 90 AGRICULTURE HANDBOOK 33 5, U.S. DEFT. OF AGRICULTURE

DN-2218 FIGURE 3.—Honey bee larvae killed by European foulbrood, as seen in cells: A, Healthy larva at earliest age when brood dies of European foulbrood; B, scale formed by dried-down larva; C, one of several positions of sick larvae prior to death; D-E, longitudinal views of scales from larvae that were in lengthwise position prior to death. BEEKEEPING IN THE UNITED STATES 91

DN-2217 FIGURE 4.—Honey bee larvae killed by sacbrood, as seen in cells: A~B, Larvae in different stages of decomposition; C, erect head of dead larva showing through opening made by bees in capping; D~E, views of scale; note how head remains erect; F, remains of larva, head of which has been gnawed away by bees. 92 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

Purchasing used bee equipment may be eco- a definite diagnosis in the apiary is sometimes dif- nomical but can lead to serious disease problems. ficult. Examination by laboratory methods is then Infected honey, combs, hive bodies, or other equip- necessary. Sometimes the diagnoses made in the ment are important disseminators of bee diseases apiary and verified in the laboratory are also far beyond the flight range of the honey bees. desirable. Consequently, beekeepers should thoroughly in- Diagnosis of the disease in the laboratory is spect colonies or equipment before making the a service made available to beekeepers and State actual purchase. Even then, used hive equipment apiary inspectors by the U.S. Department of should be washed in hot soapy water or scorched Agriculture. before it is used. The wash water should be dis- A sample of brood comb for laboratory exam- posed of in such a way that it would be inacces- ination should be 4 or 5 inches square and contain sible to foraging bees. A common practice among as much of the dead brood as possible No honey beekeepers is the capturing of swarms. These should be present, and the comb should not be should be isolated and inspected for disease before crushed. A sample of adult bees should consist they are moved into a healthy apiary. of at least 200 sick or recently dead bees. A beekeeper should always keep a clean, tidy Mail the samples in a wooden or strong card- apiary. Scrapings of comb and propolis should be placed in a sealed container. Honey should not be board box. Do not use a tin, glass, or plastic exposed. If sugar sirup is fed, it should not be container, and do not wrap the comb or bees in allowed to run out the front entrance. These waxed paper or aluminum foil. Send all samples measures are important in the prevention of to the Bee Pathology Laboratory, Entomology robbing of a diseased colony. Building A, Agricultural Research Center. Belts- The U.S. Food and Drug Administration has ville, Md. 20705, Your name and address should approved labeling for Terramycin, sulfathiazole, be plainly written on the box. If the sample is and fumagillin as aids in the control of bee dis- forwarded by an inspector, his name and address eases. Terramycin has been used to control Amer- ican and European foulbrood, whereas sulfathiaz- should also appear on the box. ole is effective only on American foulbrood. Fumagillin has been used with limited success in controlling nosema disease. It is well to remember References that the label carries specific instructions for the (1) BAILEY, L. use of these materials for bee diseases. Therefore, 1963. INFECTIOUS DISEASES OF THE HONEY BEE. these drugs can be used subject to State laws and 175 pp. Lard Books, London. regulations in the manner specified, but should (2) DADE, H. A. never be used at a time or in such a way that 1949. THE LABORATORY DIAGNOSIS OF HONEY would result in contamination of the marketable BEE DISEASES. Rev., 19 pp. Williams and Norgate, London. [Monographs of honey. Terramycin and sulfathiazole do not kill Quekett Microscopical Club No. 4.] spores of Bacillus larvae. Consequently, some (3) GOCHNAUER, T. A. States require that the bees, contaminated combs, 1963. DISEASES AND ENEMIES OF THE HONEY and honey of infected colonies be destroyed by BEE. In Grout, R. A., ed.. The Hive and the Honey Bee. Rev., 556 pp. burning. Beekeepers should consult their local Dadant & Sons, Hamilton, 111. apiary inspector for instructions on the disposal (4) GREAT BRITAIN MINISTRY OF AGRICULTURE, FISHERIES of diseased hives and the use of drugs. AND FOOD. 1959. DISEASES OF BEES. Bulletin 100, 31 pp. (5) TowNSEND, G. F., BURKE, P. W., and SMITH, N. V. Sending Samples for Laboratory 1965. BEE DISEASES AND PESTS OF THE APIARY. Ontario Dept. Agr. Pub. 429, 36 pp. Examination (6) U.S. AGRICULTURAL RESEARCH SERVICE, ENTOMOLOGY RESEARCH DIVISION. If only a small amount of the brood or a few 1967. IDENTIFYING BEE DISEASES IN THE APIARY. bees are affected or if the symptoms are unusual. U.S. Dept. Agr. Inform. Bui. 313, 19 pp. DISEASES AND PESTS OF ADULT HONEY BEES

By M. DIANE ROBERTS, parasitologist, Entomology Research Division, Agricultural Research Service ^ Nosema Disease search for a new host. They transfer to the body hairs of a new host and enter its thoracic spiracles. Nosema disease is a major disease of adult honey Symptoms of acariñe disease include inability bees and can be found wherever bees are found. It to ñy, distended abdomens, disjointed wings, and is responsible for heavy economic losses in the bee- dwindling colonies. The spread of the disease keeping industry. The causative organism of the within the colony depends partly on the presence disease is a protozoan parasite, of newly emerged bees. The thoracic spiracles are Zander. The disease is transmitted by ingestion protected by a fringe of hairs, which are soft and of its spores, which germinate soon after reaching flexible on the young bee but soon harden to form the ventriculus. a barrier against the entering mite. Therefore, bees The external symptoms of nosema disease are that escape infestation during the first week of not specific. Infected bees may have disjointed life do not become infected. wings and swollen abdomens. The infected ventri- Positive diagnosis is possible only by dissection culus may be swollen, soft, and grayish white. of the thorax and observation under a low-power However, the disease may be present with no vis- microscope of the exposed tracheae. If healthy, ible symptoms. A heavily infected colony may the tracheae appear uniformly white or flesh collect less food or rear less brood. The ovaries of colored; if heavily infested, they are blackened. an infected queen soon degenerate, and the result Intermediate stages between noninfested and may be a queenless colony or supersedure. heavily infested can be recognized by a bronzing The ventriculus and Malpighian tubules are the or streaking or both of the tracheae. organs principally injured by the organism; how- ever, it may spread to other tissues and organs. The infection greatly interferes with normal ab- Septicemia sorption of food. Workers, drones, and queens are affected. Septicemia is an infectious bacterial disease of Many questions remain unanswered concerning the adult honey bee, in which the causative agent the life cycle of N. apis. This disease appears to multiplies and spreads in the body cavity. The have an annual cycle. The incidence of infection causal organism was first classified as Bacillus reaches a peak in the spring, gradually declines apisepticus Burnside and later reclassified as during the summer, and may display a small peak Pseudomonas apiseptica (Burnside). Other orga- in the fall. nisms, classified as Serratia sp., cause a septicemia- Diagnosis of nosema disease is based on the like disease in Switzerland. spores seen in a microscopic examination of the The method by which these pathogenic bacteria crushed ventriculus or midgut. Identification of reach the blood is not understood, but once in the spores in droppings of living bees is also valid and blood they multiply rapidly and cause death of useful. The absence of spores does not insure that the bee in a short time. other stages are not present. The principal symptom of septicemia is the disintegration of the bee. The slightest handling soon after death causes the legs, wings, and other Acariñe Disease body parts to fall apart. Bees with septicemia Although acariñe disease has not been found in have a characteristic foul odor. The blood of in- North America, it is a potential threat to American fected bees is often a milky color rather than clear beekeeping. This disease is caused by a mite, Aca- or slightly opalescent as in healthy bees. rapis woodi (Rennie), which spends most of its life Septicemia can be diagnosed by the typical dis- cycle in the anterior thoracic tracheae of the adult bee. Eggs are laid and hatched, and the mites ma- memberment of the bee's body soon after death. ture within the tracheae. Mature female mites More positive diagnosis is made by microscopic leave the tracheae and migrate to the body hairs to examination of the hemolymph, where short rod- shaped bacteria indicate the presence of the 1 In cooperation with Louisiana Agricultural Experiment causative organism. Station. No treatment is presently available. 2Ô1-907 0—^7—7 93 94 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE Bee Paralysis Wax Moth Bee paralysis is a name given to several maladies The greater wax moth (Galleria mellonella having similar symptoms. Suspected causes of (L.)) causes a substantial loss of combs, honey, paralysis are viruses, pollen and nectar from such and bee equipment. It is found almost every- plants as^ buttercup, rhododendron, laurel, and where bees are raised. Since it is most active some species of lime, deficient pollen during brood at higher temperatures, its greatest damage occurs rearing in the early spring, and consumption of in the Southern United States. stored fermented pollen. Masses of small white eggs (fig. 1, A) are laid Two morphologically different viruses, chronic on combs or in cracks between hive parts. The bee paralysis virus (CBPV) and acute bee paralysis larva (fig. 1, B) receives its nourishment from virus (ABPV), have been isolated from paralytic the impurities in the wax. In obtaining these bees. CBPV kills the bees more slowly than does impurities it ingests the wax itself and in so ABPV. Paralytic bees are feeble and shaky. Their doing destroys the comb. Brood combs and legs and wings are slightly spread and their honey combs containing pollen are attacked more stomachs are abnormally distended with honey. readily than new combs or foundation. In the chronically paralyzed bees, microscopic When fully grown, the larva spins a rough examination of hindgut tissue sections shows silken cocoon, which is usually attached to some certain cell inclusions that do not occur in healthy solid support, such as the frame or the inside or acutely paralyzed bees. The mouth parts of of the hive. Frequently the larva cements the ABPV-infected bees are permanently extruded and cocoon inside a cavity chewed in the wood. disjointed, so that it is impossible for them to eat. Frames chewed in this way are weakened and The virus is spread by infected nurse bees and easily broken. by contaminated feces. ABPV can be transmitted Within the cocoon the larva changes to the through the egg or transovarially. Susceptibility pupa and overwinters in the pupal stage. Under to CBPV appears to be inherited from the queen. warm^ conditions adults may emerge at almost Tentative diagnosis can be made on the basis any time of year. of symptoms alone. Positive diagnosis requires histological examination. No treatment is known The adult female moth (fig. 1, C) is about for bee paralysis; however, requeening sometimes % inch long with a wingspan of 1 to IK inches. The males are smaller and have a scalloped eliminates the disease. In the United States, losses outer margin on the fore wing. due to bee paralysis are considered relatively insignificant. The female begins to deposit eggs 4 to 10 days after emergence from the cocoon. The total number of eggs laid by a female varies, but is Other Adult Diseases usually fewer than 300. The adults may live as Amoeba disease is caused by Vahlkampßa long as 3 weeks. (Malpighamoeba) mellificae Prell, a micro-orga- The larva is most destructive to combs stored nism that invades the Malpighian tubules. Para- in dark, warm, poorly ventilated places. It some- sites develop and multiply in the excretory cells times attacks combs within the active hive. The lining the tubules, forming spherical cysts, which damage consists of small tunnels and borings are passed in the excreta of the bee. Cysts are through the wax caps of the honey cells (fig. 2). swallowed by other bees to begin another phase The most effective enemies of the greater wax of growth and multiplication of the disease. Its moth are bees themselves. Bees in a strong colony synaptoms and effect on the individual bee are will remove the moths and prevent damage. similar to those of nosema disease. ^ The diagnosis of the disease depends on the Therefore, beekeeping methods that help main- discovery of the cysts under the microscope tain strong colonies will help prevent wax moth either in saniples of bees or in their excreta. damage. For protection, combs can be held in In the United States, bee losses from this cold rooms or stacked in such a A\'ay that a strong disease are relatively mild. Combs from diseased draft of air flows over them. colonies can be made safe for use by fumigation The larvae of the lesser wax moth (Achroia with acetic acid. grisella (F.)) cause damage to stored combs Gregarines are protozoan parasites found in similar to that of the greater \\'ax moth. The the digestive tract of adult honey bees. They larvae of the Mediterranean flour moth {Anagasta occur in the United States, France, Italy, Switzer- kuehniella (Zeller)) feed on pollen in the hive and land, Canada, and South America. Gregarine spores are eliminated from the bee in the excreta cause some damage by tunneling into the midrib and are ingested by new hosts in water or food. and the brood cells. These two moths are con- The pathological importance of this parasite has trolled by the same methods used for the greater not been determined. wax moth. BEEKEEPING IN TîIE UNITED STATES 95

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BN-30036, BN-30035, BN-30034

FIGURE 1.—Greater wax moth: A, Eggs; B, larvae (left, dorsal; right, lateral); C, adult.

has a tremendous appetite for bees and can se- Other Pests verely weaken a colony. They may be controlled Ants sometimes invade the hive and disrupt by trapping and poisoning. Bears eat bees, brood, the work of the colony, but generally they are and honey and destroy the hive in the process. Electric fences are helpful in discouraging such more of a pest to the beekeeper. Termites can raids. damage the hive parts placed on the soil. Other Mice may enter the hive in winter if the entrance insects such as dragonflies, robber flies, praying is not reduced or protected by a mouse guard. mantises, ambush bugs, and certain wasps are They che\\' the combs and frames and construct natural enemies of the honey bee. their nest in the deserted area of the hive. Skunks and bears cause extensive damage to Birds, toads, and frogs are also minor pests of apiaries in many parts of the country. A skunk the honey bee. 96 AGRICULTURIE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

FIGURE 2.—Brood comb infested with greater wax moth larvae.

External mites closely resembling Acarapis mainly infests queen bees. The adult louse gen- woodi, the cause of acariñe disease, infest honey erally does little damage. It is not a true parasite, bees and are widely distributed in the United but feeds on the nectar or honey from the mouth States. Apparently they have no adverse effect on their hosts. parts of its host. The greatest damage is from the The bee louse Braula coeca Nitzsch is occasion- burrowing of the larvae in the cappings of ally fomid on worker bees and drones, but it honeycomb. PESTICIDES

By PHILIP F. TORCHIO, apiculturist, Entomology Research Division, Agricultural Research Service'-

One of the major problems faced by beekeepers in the United States and in most other countries with highly developed agriculture is the poisoning of bees by pesticides. By nature, bees from a colony roam and visit flowers over an area of several square miles. The intensity of visitation in any one part of the area is determined by the relative attractiveness of the flowers. The extent of damage to the colony caused by a pesticide application is affected by the number of bees from the colony working in the treated area, the type of food (nectar and pollen) collected, the time of day the pesticide is applied, the method and number of applications, and the relative toxicity of the material to bees. The damage can occur when the bee collects food from treated or drift-contaminated plants or by contact with the pesticide on the plant or in the air. The brood can be damaged if fed contaminated pollen. Symptoms of Pesticide Poisoning The following are some usual symptoms of pesticide poisoning. Not all of them are likely to be seen at any one time. Nor are they conclusive as pesticide poisoning, as they can also be the result of other causes. None of these symptoms indicate which material may have affected the bees.

(1) An excessive number of dead bees in front BN-30058 of the colony (fig. 1). (2) An unusual number of dead colonies at one FIGURE 1.—Dead bees at hive entrance- -a symptom time, particularly if they contain honey. of pesticide poisoning. (3) A depleted population when the colony should be strong. (4) Sudden cessation of food storage. (12) Incoming nectar- or pollen-laden bees at- (5) Dead or deserted brood, with honey in tacked at the hive entrance by other bees. the hive. (13) An unusual number of bees emerging (6) Dead bees on the floor of the hive during from the entrance carrying dead bees. (The mild weather. normal daily death rate inside the colony is (7) A severe break in the brood rearing cycle. about 100 bees.) (8) A cessation in flower visitation (of sig- (14) Paralyzed, stupefied, or preening bees nificance especially where pollination is desired). on weeds or other objects in the apiary. (9) Bees crawling from the entrance to die nearby. (10) Dead bees in the hive—on the tops of Chemical Analysis of Bee Samples frames or on the bottom board. (11) The absence of the usual "hum" of Establishing proof of bee poisoning by chemical workers in the air. analysis is difficidt even when the type of pesticide is known. Many pesticides break down rapidly ■ In cooperation with Utah Agricultural Experiment when exposed to the elements. For a sample to Station. be of any value to the analytical chemist, it should 97 98 AGRICULTURE HANDBOOK 3 3 5, U.S. DEBT. OF AGRICULTURE be collected immediately after exposure and kept search workers. Hundreds of pesticide formulations under deep-freeze conditions until analyzed. have been tested in both dusts and sprays. These There is no Federal laboratory equipped for tests have not been conducted on low-volume or routine analysis of bee samples for all pesticide resi- ultra-low-volume applications. Limited experience dues. Some State experiment stations are equipped indicates that with these methods of application to determine certain residues. If analysis of a sam- the hazard to bees increases with certain materials. ple of bees is desired, the State experiment station Extra precautions should be taken until the hazard or extension service should be consulted before the of pesticides applied by these new methods has sample is submitted to determine whether an anal- been determined. The results of field and labora- ysis can be made. Some commercial laboratories tory tests using diluted pesticides are incorporated analyze for residues on a fee basis. in table 1. Wild bees are also damaged not only by con- taminated food but also by contaminated leaf Determining Toxicity Levels of material, florets, soil, and other articles used in Pesticides nesting. Furthermore, pesticide residues in soil can remain toxic to ground-nesting bees for The problem of determining toxicity levels on several years. Table 2 shows the comparative honey bees in the laboratory and field has been toxicity of 29 pesticides to the alkali bee and the studied for years by many Federal and State re- leafcutting bee.

TABLE 1.—Summary of toxicity and poisoning hazard oj pesticides to honey bees

Field application in dust Field application in spray Laboratory Pesticide 2 Type 3 toxicity Toxicity Residual Use Toxicity Residual Use effect class 4 effect class ^

Abate P Low 3 hours II aldrin__ C Very high__ Very high I Very high allethrin B Low III Aramite_ M _-_do Moderate II-III_. Low III azinphosethyl (Ethyl P Very high Very high__ 1 day +___ Guthion). azinphosmethyl (Guth- P ___do do 2-4 days ion). BanoL _ Ca __do >7 hours. _- Bay 39007 (Baygon) Ca High High >1 day____ Bay 39007 (Baygon G)__ Ca Low Ill Bay 41831 P Very high Bidrin_ P do_ Very high__ 5 hours-1 day+. binapaycryl (Morocide) D Low Low <2>^ hours. III Bomyl _ P Very high Low-high_ - _ 2 days calcium arsenate I High Very high_ _ Long I__ _ carbaryl (Sevin) Ca Low-high _ High 3 days + --- I Moderate- 7-12days+- high. Carbaryl (Seyin G) Ca Low <2 houis -_ III carbophenothion (Tri- P Moderate __ High >1 day____ I High <5 hours--- II thion). chlorbenside (Mitox) M Low Low <2 hours-__ III chlordane C Very high__ High-yery 2-3 days___ I - High high. chlorobenzilatc M Moderate Low III chloropropylate M do

TABLE 1.—Summary of toxicity and poisoning hazard of pesticides to honey bees ^—Contbued

Field application in dust Field application in spray Laboratory Pesticide 2 Type toxicity Toxicity Residual Use Toxicity Residual Use effect effect class *

Dilan C Low .Low-high. .3 hours- .II Low-high. 3 hours. __ II dimethoate P Very high. Very high- 1-2 days.. I dimetilan Ca Moderate.. Low 3 hours II Dimite (DMC) M Low ..do III dinitrobutylphenol D Very high. Very high. lday + . I (DNOSBP). dinitrocresol (DNOC)___ D High ..do I dinocap (Karathane) D Low III dioxathion (Delnav) P ...do Low-high- 2 hours. II disulfoton (Di-Syston)___ P Very high- Low 3 hours- II Di-Syston G P Low. <2 hours .. III DN-111 D Low III endosulfan (Thiodan) __ C Moderate.. Low <5 hours. _ II endrin C Very high. Moderate.. <2 hours.. II EPN P -.do High. 1 day + Very high.. I ethion P Low Low-high.. <2 hours. II famphur (Famophos) P Very high. I fenson M Low <2 hours _. III fenthion (Baytex) P Very high. Very high.. 2-3 days+. I Genite 923 M Low Moderate II-III Low <2 hours __ III heptachlor C Very high. ... Very high.. I heptachlor G C Moderate <2 hours Hooker HRS-16 (Pentac) M Low <1 day.. III Imidan P Very high. Very high. 1-4 days.. I isobenzan (Telodrin) C do High >2 hours. II isodrin C Moderate- II Isolan Ca High Low. 3 hours.. II isopropyl parathion P Low III(?) Kepone C do Low. <Í'd¿y- Ill Kepone bait C Low_ III. lead arsenate I Very high. Very high_. Long I lime sulfur I Low Moderate. III lindane and benzene C Very high. Very high. 2 days + _ I_. High I hexachloride (BHC). malathion do_.-- do 1 day + .. I.. Moderate- 2 hours-2. I very high. days + . malathion G P Low None 11. Matacil Ca High. Very high. >3 days.. I menazon P Moderate. <2 hours- II methoxy chlor C Low.. do 7 hours- do >3 hours. II Naugatuck D-014 (Omite) M Low <3 hours- III Nemacide (V-C 13)___ P High. do 2 hours II Neotran M Low_ do-... III nicotine sulfate B .--do- Low Few hours. ilí Nissol A Low 3 hours. II ovex M Low.. Low-high. II-III .-do ox y demetonmethy 1 P High. Moderate. None... II (Metasystox R). paraoxon P Very high. I parathion P .--do Very high. 1 day+._ I High lday+. I Perthane C Moderate. Moderate. lday+-. II Low <1 day. II phorate (Thimet E)___ P .--do Very high. 5 hours.. II phorate (Thimet G) P Moderate- <2 hours. II phosphamidon P Very high- Very high. 2 hours-2 days. Footnotes at end of table. 100 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

TABLE 1.—Summary of toxicity and poisoning hazard of pesticides to honey bees ^—Continued

Field application in dust Field application in spray Laboratory Pe&ticide 2 Type toxicity Toxicity Residual Use Toxicity Residual Use effect class ^ effect class ^

Phostex P Moderate. High. 2 hours II propyl thiopyrophos- P Very high, phate (NPD). Low. 2}^ hours II Pyramat Ca __do I pyrethrum B Low. Low_ 3 hours. III ronnel (Korlan) Low III P Moderate.. 3 hours II rotenone B Low_ Low-high. _ 3 hours... II P Low-very Low 1 day III high. sodium hexafluorosili- Low. cate bait. Ill Str obane C Low III sulfotepp P Very high_ sulfur I I Low III Low III Sulphenone M Moderate. Moderate. II-III TDE (Rhothane)___ ...do III C ._do --do II Moderate. II Temik G Ca Low None III tepp P Very high. Very high- <3 hours. II tetradifon (Tedion) Very high. 3 hours... II M Low Low <2 hours. III Tetram P _.do Thiocron Moderate. II P High Low 3 hours. II thioquinox (Eradex) Co Moderate. toxaphene II C Low Low-high. 3 hours. I terex). Low-high.. 2-5 hours. II Zectran Ca Very high, Zinophos Very high. . 1-2 days. I P .-do I

1 Arrangement and much of data from Johansen, C. A., hydrocarbon, Ca=carbamate, Co=carbonate, D=dinitro Summary of the Toxicity and Poisoning Hazard of Insecti- compound, I=inorganic compound, M=specific miticide, cides to Honey Bees, Gleanings Bee Cult. 94: 474-475 and P=organophosphorus compound. 1966. ' ^ Classification of materials: I=hazardous to bees at any 2 Chemical names given at end of section on Pesticides. time, II=not hazardous if applied when bees are not 3 A=acetamide, B=botanical or derivative, C=chlorinated foraging, III=not hazardous to bees at any time.

Laboratory tests of pesticide formulations on for a particular period. In addition, the exami- honey bees determine toxicity levels on individ- nation of colonies in treated and untreated areas uals but do not indicate seriousness of damage and counting dead bees in front of hives before to the field force and its pollinating or honey- and after apphcations provide a basis for evalu- production potential. The field testing of insec-^ ating the effect of the material on the colony. The ticides on the honey bee is especially diflacult, combined data obtained from these testing because individuals visit fields briefly, the social techniques provide for an intelligent estimate of organization normally prevents all members of pesticide effects on honey bees in the field. the colony from being equally exposed to pesti- cides, and a part of the field force of any colony may be visiting areas outside the confines of the Reduction in Bee Losses experiment. Any or all of these factors can Observation of the following precautions can drastically affect results. Furthermore, there is no significantly reduce bee losses from pesticide accurate measurement of sublethal effects on a poisoning. colony exposed to a pesticide application. Techniques are used that determine (1) toxic Grower Cooperation effects of direct apphcations, by placing caged The grower should use a pesticide only when bees in the field as the field is being treated; (2) needed. The benefit of the material should out- fuming effect, by placing additional caged bees weigh the harm it does to bees. The value of the m the field at hourly intervals after treatment bees as polHnators should be considered, as well until mortaUty ceases; and (3) residual effects as the effect of the pesticide on them. The effect of on the field force, by counting floral visitors before the pesticide on the pollinators of other crops and after exposure to treated and untreated areas in the area should also be considered. A pesticide BEEKEEPING IN THE UNITED STATES 101

TABLE 2.—Comparative toxicity oj 29 'pesticides the night is sufficiently cool for the bees to move to alkali bee and alfalfa leaf cutting bee ^ inside. Colonies can be damaged by fumes of some pesticides, such as parathion, azinphos- Alfalfa methyl, malathion, and benzene hexachloride. Pesticide 2 Alkali bee leaf cut- Notify the beekeepers near areas to be treated so ting bee that they may move or otherwise protect the colonies. However, notification is not a release of Aramite III responsibility. Bidrin I I carbaryl (Sevin) I I Beekeeper Cooperation carbophenothion (Trithion)_ II I DDT III I Select safe bee locations. Place colonies away demeton (Systox) III III from agricultural areas if possible, away froni fields diazinon I routinely treated, or at least where they will not dicofol (Kelthane) III III be subject to drift of the material from the treated dimethoate I I dioxathion (Delnav) III field. endosulfan (Thiodan) I I Identify the colonies. Post owner's name, ad- endrin II I dress, and telephone number in a conspicuous EPN I place in the apiary. Let the nearby growers know ethion I malathion II I where the bees are located so the beekeeper can menazon III be notified. methoxychlor III Know the pesticides. Be acquainted with pesti- mevinphos (Phosdrin) I cides likely to be used in the area and their lulled (Dibrom) II II oxydemetonmethvl (meta-Svstox potential hazard to bees. R) ^^ III Confine the bees when hazardous ^ materials parathion I I are applied. Beehives can be covered with plastic phosphamidon I I sheeting that will confine the bees and exclude Phostex III ronnel (Korlan) I pesticide sprays, dusts, or fumes. Since heat builds schradan (OMPA) III up rapidly under plastic exposed to the sun, con- tepp III finement can only last for a few hours after dawn tetradifon (Tedion) III on warm days. This may be long enough to protect toxaphenc III trichlorfon (Dylox, Dipterex) the bees from some materials. Hives can also be covered with wet burlap for a day or more, even during the hottest weather, 1 Classification of materials: I = hazardous to bees at any time, 11 = not hazardous if appHed when bees are not and the bees will not suffer from lack of air or foraging, 111 = not hazardous to bees at any time. water. The hives should be covered at night when all 2 Chemical names given at end of section on Pesticides. the bees are in the hives. During the day the bur- lap should be soaked with water at least once every hour. aiding one crop could seriously reduce production Relocate the colonies if they are likely to be of another one in the area. repeatedly exposed to hazardous pesticides. Select the right pesticide. All pesticides are not Bees are valuable to the grower. Try to con- equally hazardous to bees. Some pesticides will vince him of their value to him and of the kill an entire colony, some will weaken it, but still importance of protecting them. others are safe. Select the pesticide that is least hazardous to pollinators and that will control the harmful pests. References Apply granules or sprays rather than dust. (1) ANDERSON, L. D., and ATKINS, E. L., JR. Granules are, in general, harmless to bees. Sprays 1958. EFFECT OF PESTICIDES ON BEES. Calif. drift less than dusts. Agr. 12: 3-4. Use ground equipment. Airplanes discharge (2) and ATKINS, E. L., JR. 1960. HAZARDS TO HONEY BEES AND OTHER BENE- pesticides at higher altitude and with greater FICIAL INSECTS. 2 pp. Calif. Univ. Citrus turbulence than ground machines. This increases Expt. Sta. [Processed.] the likelihood that bees in flight will come in (3) ATKINS, E. L., JR., and ANDERSON, L. D. 1954 TOXICITY OF PESTICIDE DUSTS TO HONEY contact with the pesticide or that it will drift onto BEES. Jour. Econ. Ent. 47: 969-972. adjacent crops or into apiaries. Time the pesticide (4) BoHART, G. E., and LIEBERMAN, F. V. application. The safest time to apply pesticides is 1949. EFFECT OF AN EXPERIMENTAL FIELD APPLI- when bees are not working plants. Treat at night CATION OF DUST ON NOMIA MELANDERI. or at a time of day when bees are not in the field. Jour. Econ. Ent. 43: 519-522. (5) JOHANSEN, C. A. Avoid drift of the pesticide. Bees cluster on the 1961. LABORATORY TOXICITY OF SEVERAL INSEC- hive entrance on hot days and nights where they TICIDES TO THE HONEY BEE. Jour. ECOU. can be exposed to drifting pesticides. Wait until Ent. 54: 1008-1009. 102 AGRICULTUREi HANDBOOK 335, U.S. I>EPT. OF AGRI-CUUTURE

(6) JAYCOX, E. R., and HUTT, R. (10) TODD, F. E., LIEBERMAN, F. V., NYE, W. P., and 1963. THE EFFECT OF PESTICIDES ON THE ALFALFA KNOWLTON, G. F. LEAFCUTTING BEE MEGACHILE ROTUNDATA. 1949. THE EFFECT OF FIELD APPLICATIONS OF IN- Wash. Agr. Expt. Sta. Cir. 418, 12 pp. SECTICIDES ON BEES. Agr. Chem. 4: (7) LIEBERMAN, F. V., BOHART, G. E., KNOWLTON, G. F., 27-29. and NYE, W. P. (11) and MCGREGOR, S. E. 1954. ADDITIONAL STUDIES ON THE EFFECT OF ^^^^- '^^^ ^^^ ^^ HONEY BEES IN THE PRODUC- FIELD APPLICATIONS OF INSECTICIDES ON ,,^, ^^^ , TiON OF CROPS. Ann. Rev. Ent. 5: 265-278. HONEY BEES. Jour. Econ. Ent. 47: (i^-^ ^'^- AGRICULTURAL RESEARCH SERVICE, ENTOMO- 316-320. ... LOGY RESEARCH DIVISION. (8) LiNSLEY, E. G., MACSWAIN, J. W., and SMITH, R. F. ^^^^' PROTECTING HONEY BEES FROM PESTICIDES. 1950. COMPARATIVE SUSCEPTIBILITY OF WILD (13) WAITE M f ^^ ^' ' ^^' Fcnn Fn? ^''•''^^ a^^^ ^"^ ''''''' ^'''''' 1895. POLLINATION OF PEAR FLOWERS. U.S. íQ\ Mnr^^nr.^ T'^ 7^T n rp Dept. AgF. Div. Vcg. Path. Bui. 5, 86 pp. (9) MCGREGOR, S. E., and VORHIES, C. T. (14) WAY, M. J., and SYNGE, A. D. 1947. BEEKEEPING NEAR COTTON FIELDS DUSTED 1948. THE EFFECTS OF DDT AND BENZENE HEX- wiTH DDT. Ariz. Agr. Expt. Sta. Bui. 207, ACHLORIDE ON BEES. Ann. Appl. Biol. 19 pp. 35: 94-109. Chemicals Referred to in Tables 1 and 2

Name used Chemical name Abate ® 0,0-diinethyl phosphorothioate 0,0-diester with 4,4'-thiodiphenol aldrm not less than 95 percent of l,2,3,4,10,10-hexachloro-l,4,4a,5,8,8a- hexahydro-l,4-e7i¿o-6a;o-5,8-diinethanonaphthalene allethrm 2-allyl-4-hydroxy-3-methyl-2-cyclopenten-l-one ester of 2,2-di- . methyl-3-(2-methylpropenyl)+ cyclopropanecarboxylic acid Aramite ® 2-(^-¿6r¿-butylphenoxy)-l-methylethyl 2-chloroethyl suKite azmphosethyl (Ethyl Guthion ®)_ 0,0-diethyl phosphorodithioate S-ester with 3-(mercaptomethyl)- 1,2,3-benzotriazin-4 (3iZ) -one azinphosmethyl (Guthion (g)) 0,0-dimethyl phosphorodithioate AS-ester with 3-(mercaptomethyl)- l,2,3-benzotriazin-4(3fí)-one Banol (R) 6-chloro-3,4-xylyl methylcarbamate Bay 39007 (Baygon (g)) o-isopropoxyphenyl methylcarbamate Bay 41831 0,0-dimethyl 0-4-nitro-m-tolyl phosphorothioate benzene hexachloride 1,2,3,4,5,6-hexachlorocyclohexane, consisting of several isomers and containing a specified percentage of gamma isomer Bidrin (g) 3-hydroxy-A^,Ar-dimethyl-cis-crotonamide dimethyl phosphate bmapacryl (Morocide ®) 2-sec-butyl-4,6-dinitrophenyl 3-methyl-2-butenoate Bomyl ® dimethyl 3-hydroxyglutaconate dimethyl phosphate calcium arsenate calcium arsenate carbaryl (Sevin (§)) 1 -naphthyl methylcarbamate carbophenothion (Trithion (g)) Ä'-[i(p-chlorophenyl)thio]methyl] 0,0-diethyl phosphorodithioate chlorbenside (Mitox ®) p-chlorobenzyl ^-chlorophenyl sulfide chlordane. l,2,4,5,6,7,8,8-octachloro-3a,4,7,7a-tetrahydro-4,7-methanoindane chlorobenzilate ethyl 4,4'-dichlorobenzilate Chloropropylate ® isopropyl 4,4'-dichlorobenzilate Chlorthion ® (now unavailable) _ _ ö-(3-chloro-4-nitrophenyl) 0,0-dimethyl phosphorothioate Ciodrin ® aZ^Aa-methylbenzyl 3-hydroxycrotonate dimethyl phosphate cryolite sodium hexafluoroaluminate DDT 1,1 ,l-trichloro-2,2-bis(p-chlorophenyl)ethane demeton (Systox ®) mixture of 0,0-diethyl AS'(and 0)-[2-(ethylthio)ethyl] phosphorothio- ates diazinon 0,0-diethyl 0-(2-isopropyl-4-methyl-6-p3n:'imidinyl) phosphorothioate dicapthon 0-(2-chloro-4-nitrophenyl) 0,0-dimethyl phosphorothioate dichlorvos (Vapona ®) 2,2-dichlorovinyl dimethyl phosphate dicofol (Kelthane ®) 4,4'-dichloro-aZpAa-(trichloromethyl)benzhydrol dieldrin not less than 85 percent of l,2,3,4,10,10-hexachloro-6,7-epoxy- l,4,4a,5,6,7,8,8a-octahydro-l,4-e7i¿o-gíco-5,8-dimethanonaphtha- lene Dilan ® a mixture of 1 part of 1,1 -bis (p-chlorophenyl) -2-nitropropane (Prolan ®) and 2 parts of l,l-bis(^-chlorophenyl)-2-nitrobutane (Bulan ®) BEEKEEPING IN THE UNITED STATES 103

Name used Chemical name dimethoate 0,0-dimethyl ^-(A^-niethylcarbamoylmethyl) phosphorodithioate dimetilan 1 -dimethylcarbamoyl-5-inethyl-3-pyrazolyl dimethylcarbamate Dimite ® (DMC) 4,4'-dichloro-aZ^Aa-methylbenzhydrol dinitrobutylphenol (DNOSBP) 2-s6C-butyl-4,6-dinitrophenol dinitrocresol (DNOC) 4,6-dinitro-o-cresol dinocap (Karathane ®) 2-(l-methylheptyl)-4,6-dinitrophenyl crotonate dioxathion (Delnav (g)) 2,3-p-dioxanedithiol S,S-his (0,0-diethyl phosphodithioate) disulfoton (Di-Syston ®) 0,0-diethyl S-[2-(ethylthio) ethyl] phosphorodithioate DN-111 (R) dicyclohexylamine salt of 2-cyclohexyl-4,6-dinitrophenol endosulfan (Thiodan ®) 6,7,8,9,10,10-hexachloro-l,5,5a,6,9,9a-hexahydro-6,9-methano- 2,4,3-benzodioxathiepin 3-oxide endrin 1,2,3,4,10,10-hexachloro-6,7-epoxy-l,4,4a,5,6,7,8,8a-octahydro- l,4-67i¿o-6n¿o-5,8-dimethanonaphthalene EPN 0-ethyl 0-p-nitrophenyl phenylphosphonothioate ethion 0,0,0',O'-tetraethyl S,iS"-inethylenebisphosphorodithioate famphur (Famophos) 0,0-dimethyl 0-[^-(dimethylsuKamoyl)phenyl] phosphorothioate fenson _p-chlorophenyl benzenesulfonate fenthion (Baytex ®) 0,0-dimethyl 0-[4-(methylthio)-m-tolyl] phosphorothioate Genite 923 ® 2,4-dichlorophenyl benzenesulf onate heptachlor 1,4,5,6,7,8,8-heptachloro-3a,4,7,7a-tetrahydro-4,7-methanoindene Hooker HRS-16 (Pentac) decachlorobi-2,4-cyclopentadien-l-yl Imidan ® 0,0-dimethyl S-phthalimidomethyl phosphorodithioate isobensan (telodrin ®) l,3,4,5,6,7,8,8-octachloro-l,3,3a,4,7,7a-hexahydro-4,7- methanoisobenzofuran isodrin 1,2,3,4,10,10-hexachloro-l,4,4a,5,8,8a-hexahydro-l,4-endo-e7i¿o-5, 8-dimethanonaphthalene Isolan ® 1 -isopropyl-3-methylpyrazol-5-yl dimethylcarbamate isopropyl parathion 0,0-diisopropyl 0-^-nitrophenyl phosphorothioate Kepone ® decachlorooctahydro-1,3,4-metheno-2íZ"-cyclobuta[cá]pentalen-2-one lead arsenate acid lead arsenate lime sulfur 30 percent calcium polysulfide and various small amounts of calcium thiosulfate plus water and free suKur lindane 1,2,3,4,5,6-hexachlorocyclohexane, gamma isomer of not less than 99 percent purity malathion 0,0-dimethyl dithiophosphate of diethyl mercaptosuccinate Matacil ® 4-(dimethylamino)-m-tolyl methylcarbamate menazon S-[(4,6-diamino-s-triazin-2-yl)methyl] 0,0-dimethyl phosphoro- dithioate methoxychlor 1,1 ,l-trichloro-2,2-bis(;p-methoxyphenyl)ethane methyl demeton (Metasystox ®)_ mixture of 0,0-dimethyl ^(and 0)-[2-(ethylthio)ethyl] phosphoro- thioates methyl parathion 0,0-dimethyl 0-^-nitrophenyl phosphorothioate Methyl Trithion ® S-[[(^-chlorophenyl) thio]methyl] 0,0-dimethyl phosphorodithioate mevinphos (Phosdrin ®) methyl 3-hydroxy-a¿^Aa-crotonate dimethyl phosphate mirex dodecachlorooctahydro-1,3,4-metheno-2íZ'-cyclobuta[ccí]pentalene Morestan ® 6-methyl-2,3-quinoxalinedithiol cyclic SjS-dithiocarbonate naled (Dibrom ®) 1,2-dibromo-2,2-dichloroethyl dimethyl phosphate Naugatuck D-014 (Omite) 2-(^-Í6rí-butylphenoxy)cyclohexyl 2-propynyl sulfite Nemacide ® 0-2,4-dichlorophenyl 0,0-diethyl phosphorothioate Neotran ® bis(^-chlorophenoxy)methane nicotine sulfate nicotine sulfate Nissol 2-fluoro-iV-methyl-A^-l -naphthylacetamide ovex ^-chlorophenyl p-chlorobenzenesulfonate oxydemetonmethyl (meta-Systox E S'-[2-(ethylsulfinyl) ethyl] 0,0-dimethyl phosphorothioate ®). paraoxon diethyl ;p-nitrophenyl phosphate parathion 0,0-diethyl O-^^-nitrophenyl phosphorothioate Perthane® a mixture of l,l-dichloro-2,2-bîs(2?-ethylphenyl)ethane (95 per- cent) and related reaction products (5 percent) phorate (Thimet ®) 0,0-diethyl AS-[(ethylthio)methyl] phosphorodithioate 104 AGRICULTURE, HANDBOOK 3 3 5, U.S. DEPT. OF AGRICULTURE

Name used Chemical name phosphamidon 2-chloro-2-diethylcarbamoyl-l-methylvinyl dimethyl phosphate rnostex® a mixture of bis(dialkyloxyphosphinothioyl) disulfides (alkyl , ^, . ratio 75 percent ethyl, 25 percent isopropyl) propyl thiopyro- propyl thiopyrophosphate phosphate (NPD ®). i^J i F Pyramat ® 6-methyl-2-propyl-4-pyrimidinyl dimethylcarbamate ronnel (Korlan ®) 0 "O-dimethyl 0-24,54richÎo7ophVnyTphosphorothïoate rotenone 1.2,12,12a-tetrahydro-2-isopropenyl-8,9-dimethoxy[l]benzopyrano [3,4-6]furo[2,3-A][l]benzopyran-6(6aíí)-one ryania __ schradan (OMPA) octamethylpyrophosphoramide sodium hexafluorosihcate bait sodium hexafluorosiUcate Strobane® terpene polychlorinates (65 percent chlorine) sulfotepp _ 0,0,0,0-tetraethyl dithiopyropho^phate sultur sulfur Sulphenone ® i^-chlorophenyl phenyl sulf one iDJ^ (Rhothane ®) l,l-dichloro-2,2-bis(^-chlorophenyl)ethane 1 emik ® 2-methyl-2-(methylthio)propionaldehyde O-(methylcarbamoyl) oxime tepp ethyl pyrophosphate tetradif on (Tedion ®) p-chlorophenyl 2,4,5-trichlorophenyl sulf one Tetram ® '_ AS-[2-(diethylamino)ethyl] 0,0-diethyl phosphorothioate hydrogen oxalate Thiocron 0,0-dimethyl phosphorodithioate S-ester with 2-mercapto-A'-(2- methoxy ethyl) acetamide thioqumox (Eradex ®) 2,3-quinoxaUnedithiol cycUc trithiocarbonate toxaphene.. chlorinated camphene containing 67-69 percent chlorine trichlorfon(Dylox®,Dipterex®)_ dimethyl (2,2,2-trichloro-l-hydroxyethyl)phosphonate Zectran ® _ _ _ _ 4-(dimethylamino)-3,5-xylyl methylcarbamate Zmophos ® 0,0-diethyl o-2-pyrazinyí phosphorothioate

FOLLOW THE LABEL

U.S. DEPARTMENT OF AGRICULTURE HYMENOPTEROUS INSECT STINGS

By THOR LEHNERT, apiculturist, Entomology Research Division, Agricultural Research Service

The honey bee is the most common single spe- powerful depressant, causing a rapid fall in blood cies of stinging insect in the United States. Bees pressure. and the related wasps, hornets, yellow jackets, Melittin, a protein having a molecular weight and ants usually do not sting unless stepped on, of 33,000 to 35,000, is thought to be respon- touched, or molested. They are usually not active sible for the general local toxicity of the venom. at temperatures below 55° F. or on rainy days. Melittin in high concentrations has also caused The highest incidence of stings is in August. Yel- hemolysis of red blood cells. low jackets, honey bees, and wasps are the worst Bee venom contains at least two enzymes—a offenders in that order. Yellow j ackets cause more hyaluronidase and phospholipase A. The hyalu- moderate and also more severe reactions than bees ronidase is believed to be the ''spreading^^ factor. or wasps. By breaking down the cell-cementing substance, hyaluronidase allows the toxic principles of bee Bee Venom venom to infiltrate the tissues. Phospholipase A apparently has no general The poison gland system of the bee consists of toxicity. However, through indirect action on a small alkaline gland and a larger acid gland. the unsaturated fatty acids, it causes hemolysis The venom comes from these two glands. The of red blood cells. Phospholipase A also causes stinger is made up of two lancets with sharp barbs inactivation of thrombokinase, inhibits oxidative pointing backward, similar to a harpoon. When a phosphorylation, and attacks enzymes involved honey bee stings, its stinger is pulled from its with metabolic dehydrogenation. The pain expe- body. Even after the stinger is separated from rienced after being stung may well be the result the bee, its muscular mechanism can continue to of these last three actions. force its venom into the wound. Because venom continues to be injected by the stinging mech- For many years formic acid was erroneously anism, the stinger should be removed immedi- believed to be the major component of venom ately. Most people make the mistake of pulling produced by the honey bee, and this belief is it out with the thumb and index finger and thereby still held by many. The action of venom is much squeeze more venom into the wound. The stinger more complex than the simple concept of direct should be scraped or scratched out with a action on the tissue by formic acid. fingernail. Other stinging Hymenoptera, such as yellow Sting Reactions jackets, wasps, and hornets, retain their stingers. Bee venom is a water-clear liquid with a sharp In most cases insect stings cause only a local bitter taste and a distinct acid reaction. The spe- reaction, with pain lasting for several minutes cific gravity is 1.1313. The venom is easily soluble after penetration of the stinger. A redness and in water and acid, but almost insoluble in alcohol. slight swelling at the sting site may also occur. The three toxic effects of bee venom are neuro- Until recently some people believed severe symp- toxic (paralysis of the nervous system), hemor- toms and death from stinging insects were due rhagic (increase of the permeability of the blood to venom being introduced directly into a small capillaries), and hemolytic (destruction of red blood vessel. Severe reactions are now considered blood cells). to be due to sensitivity to bee protein not venom. Recent work reveals that bee venom is a very In about 2 percent of persons a hypersen- complicated substance with several active bio- sitivity develops in which each additional sting chemical components. By 1956 at least eight ac- produces a more severe reaction. Hypersen- tive components plus several biological inactive sitivity may appear after a varying number^ of components had been identified. The substances stings, usually each sting making the reaction showing activity are histamine, melittin (a pro- progressively worse. Some develop sensitivity tein), a hyaluronidase, and phospholipase A. after one sting, whereas others after a series of The histamine recovered was shown not to be normal reactions. In a few individuals hypersen- a major pharmacological factor in bee venom, sitivity appears to be inborn. The first sting has although histamine is commonly known to be a resulted in death. 105 106 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

Symptoms in an allergic person usually appear one can take several precautions to prevent stings. within a few minutes after the sting, but may For example, the type of clothing worn affects the not appear for 24 hours. Local swelling may be probability of being stung. Beekeepers consider excessive. A hivelike condition may break out clothing color to be one of the most important over the body. There is a sensation of choking, factors. Light-colored clothes should be worn in difficult breathing, asthma, and the lips turn preference to dark, rough, or woolly clothing. blue. Shocklike symptoms, vomiting, and loss of Suede or leather materials, particularly horsehide, consciousness may follow in rapid succession. seem to be especially irritating to bees. If around bees, women should wear some type of head Treatment and Precautions covering to keep them out of their hair. The odor of perspiration has little effect on bees. Treatment is divided into three stages. (1) Im- Certain hair oils and perfumes should be avoided mediate treatment for anaphylaxis is epinephrine because they seem to be irritating to honey bees, 1:1,000, 0.3 cc. to 0.5 cc. given intramuscularly which when once attracted are likely to sting. or by deep subcutaneous injection. (2) Second- Bees will more quickly attack a moving object stage treatment includes a sympathomimetic than an immobile object. This can easily be demon- agent, such as metaraminol, 100 mg. in 500-cc. strated by making fast jerky hand movements isotonic saline solution given intravenously, a when working with a hive of bees. Striking, tourniquet above the sting site if practical, anti- swatting, and swinging at bees will increase the histamine given intramuscularly, and cortico- chance of being stung. steroids. (3) Long-term care involves immunization No success has been attained in developing an with the appropriate insect antigen. effective repellent against stinging insects. Di- The long-term treatment undertaken to relieve methyl phthalate, which has been very effective the allergic condition to stings is known as de- in repelling mosquitoes and biting flies, is not sensitization. This treatment consists of a grad- effective as a repellent against stinging insects. uated series of injections of an extract made from Only a small percentage of the population the body of the offending type of insect. If the suffers serious reactions from hymenopterous stinging insect is not identified, the sensitive insect stings. However, when reactions other than person should be treated with an antigen com- local irritation and swelling do occur, a physician posed of honey bee, hornet, paper wasp, and should be called for immediate treatment. He yellow jacket. should be consulted for long-range protection with The degree of relief from allergic reactions is desensitizing shots. not the same for all persons after desensitizing treatments. The frequency of treatment also varies. In some cases one series of treatments is References effective. In other cases the treatment must be repeated with booster injections once a month. (1) ANONYMOUS. 1965. INSECT-STING ALLERGY. Amer. Med. Assoc. Another important point is that of cross-pro- Jour. 193: 115-120. tection. The injection of stinging insect extract of (2) HODGSON, N. B. one species will protect some persons against all 1955. BEE VENOM: ITS COMPONENTS AND THEIR stinging insects. Some will need a combination of PROPERTIES. Bee World 36: 217-222. (3) MARSHALL, T. K. extracts to achieve complete protection. This 1957. WASP AND BEE STINGS. Practitioner [Lon- niakes the procedure of desensitization an indi- don] 178: 712-722. vidual process. (4) MORSE, R. A., and GHENT, R. L. 1959. PROTECTIVE MEASURES AGAINST STINGING Desensitization seems to be helpful to about INSECTS. N.Y. State Jour. Med. 59: three-fourths of persons with severe reactions, in 1546-1548. that they report lessened reactions to subsequent (5) PECK, G. A. stings. Desensitization should be considered by 1963. INSECT STING ANAPHYLAXIS. Calif. Med. 99: 166-172. persons who have severe reactions. (6) SCHOFIELD, F. W. Such persons should have an emergency kit 1957. PREVENTION OF SEVERE REACTIONS FOL- available at all times. This kit should contain a LOWING STINGS OF THE HONEY BEE. tourniquet, ampoules of epinephrine 1:1,000, and Ganad. Bee Jour. 65: 11-13. (7) SHAFFER, J. H. an injectable antihistamine. Instructions on the 1964. HYPOSENSITIZATION WITH INSECT EX- use of remedies in the kit should be obtained from TRACTS. Amer. Med. Assoc. Jour. 187: a physician. 968. Immunization of a sensitive person should (8) STIER, R. A., and STIER, R. F. 1959. THE RESULTS OF DESENSITIZATION IN always be carried out, since there is no way of ALLERGY TO INSECT STINGS. Amer. Acad. completely avoiding stinging insects. However, Gen. Pract. 19: 103-108. OBSERVATION BEEHIVE

By THOR LEHNERT, apiculturist, Entomology Research Division, Agricultural Research Service

The observation hive offers many useful pos- the outside should be located so that they wiU sibilities to the beekeeper, teacher, and researcher. leave the house above the heads of passers-by. AH types of observation hives are in use today, The location of the hive should be weU lighted ranging from single frame models to full-scale so that activities of the bees can be observed. If hives enclosed in glass. Although observation a permanent outside exit cannot be provided, hives are available commercially from many of the hive must be taken outside every second or the bee supply houses, most people get more third day so the bees can fly. They should be satisfaction from designing and building their own. allowed to fly from late afternoon to sundown. Several points must be considered when con- The hive can then be closed again and returned structing the hive. A screened hole should be indoors. provided in the hive for adequate ventilation. A The hive may be stocked with bees from a local bee space of one-fourth to three-eighths inch beekeeper who can supply a queen, a frame of between the frames and glass sidewalls, top, brood, a frame or two of honey, and worker bees. bottom, and ends of the hive will allow the bees If bees are not available from a local source, a to move around freely. If more space is provided, queen and 2-pound package may be bought from bees will quickly build burr comb on the glass a package bee producer. and make observations difficult. In figure 1 are given the design and list of References materials necessary to construct the three-frame (1) ECKERT, J. E. observation hive that is used at the Bee Disease 1958. THE OBSERVATION HIVE. Calif. Univ. Research Laboratory in Beltsville, Md. Agr. Ext. Leaflet 102, 2 pp. (2) SHAW, F. R. The hive may be set up anywhere in the house 1952. OBSERVATION BEEHIVES: CONSTRUCTION- except where the bees will get too hot, since they USE. Mass. Agr. Ext. Ser v. Spec. Cir. cannot tolerate extreme heat. The flight hole to 191,2 pp. 107 108 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

MATERIALS

TOP A 1- '/2x2'/i6 x'/î'CHANNEL, 20" LONG B CUT r X 8-VENT AND COVER WITH HARDWARE CLOTH c SECURE WITH MASKING TAPE

FRAME

D 2-]8'/2x9V4" DOUBLE-STRENGTH GLASS PANELS E 2- 10V4x2i/2'>< '/)6" ALUMINUM SIDES F 2-5/¿x 2 V2"x S/g'STACKING CLIPS, 3" LONG G 4- IOVA'SIDE RAILS (REYNOLDS WINDOW MOLD #30A) H 4- 19%'TOP AND BOTTOM RAILS (REYNOLDS WINDOW MOLD #30A) I 8- CORNER CLIPS J 2-3/4"x3/4'ANGLE FRAME SUPPORTS IV2"L0NG (1/16'ALUMINUM SHEET- RIVET TO SIDE PANELS) 20-'A'SHEET METAL SCREWS

BASE

K 1- I93/4'x2 7/i6-x ]5/8'WOODEN FRAME SUPPORT (CUT TO SHAPE) L 1- 3/4'x 53/4X20' PLYWOOD BASE (ASSEMBLE AND DRILL 1'/^DIAMETER HOLE AT CENTER) M I- 41/2'FLANGE (2"PIPE) N 1- 1V2'NIPPLE (2'PIPE,) O 1- TEE (2'PIPE) P 1- 5V2'FLANGE (WELD TO BOTTOM OF TEE) 7- rWOOD SCREWS Q 1- 2'PIPE , LENGTH TO SUIT (18 SHOWN W/CAP) R I- P/8x7"x7"WOOD BLOCK (CUT TO FIT) S 1- V'x 6'/2"x4"WOOD BLOCK (CUT TO FIT) T 1 - MASON JAR WITH FLANGED LID DRILLED WITH EIGHT l/32'HOLES

ASSEMBLED HIVE

FIGURE 1.—Design and list of materials for three-frame observation hive. MANAGEMENT OF WILD BEES

By GEORGE E. BOHART, ayiciUturist, Entomology Research Division, Agricultural Research Service '

The term 'Vild bees'^ includes all bees except bees. If these bees can be managed, they may the three species of honey bees Apis mellifera contribute greatly to the complete pollination L., indica Fabricius, and cerana Fabricius ^ man- of our crops. aged by man. Generally bees are hymenopterous insects that provide their young with a diet of pollen and honey. They are characterized by the Bumble Bees presence of plumose body hairs and usually by somewhat expanded hindlegs. Most species con- Bumble bees (Bombus) (fig. 1) are important struct and provision nests, but perhaps 5 percent pollinators of many plants. However, their small lay their eggs in nests constructed by other bees. colonies do not make them well suited for large- Most bees are solitary (no cooperation between scale pollination. females and no contact between mother and Bumble bee colonies are obtained by placing adult offspring), but perhaps 10 percent display domiciles in the field to attract queens or by various degrees of social development. Many of inducing queens to establish colonies in dom- the solitary species are neighborly and develop iciles kept in cages or greenhouses. The queens nesting aggregations numbering into the thousands can forage and seek out the domiciles, or they or even millions. can be confined to the domiciles until they start According to C. D. Michener, there are at to nest (fig. 2). least 20,000 species of bees in the world. These In areas where bumble bees are abundant, are distributed among eight recognized families attracting searching queens in the spring to dom- and perhaps 400 genera. In the United States iciles placed in the field is often the most success- there are seven families, about 100 genera, and ful method of obtaining colonies. A colony between 3,000 and 4,000 species. In favorable established in this manner can be moved just locations only a few square miles may be in- before the first workers emerge. It should be habited by several hundred species. placed on a post under a shelter and the entrance Probably the high point in wild bee populations tube reduced to confine the queen. Honey should in the United States was reached about 1915, be placed in the honeypot when this is done. when hedgerows and unused fields provided ideal Colonies of Bombus terrestris (L.) and B. habitats and insecticides were rarely used. Today lapidarius (L.) have produced as many as 1,000 intensive land use, elimination of weeds, and bees, although never more than about one-third widespread and frequent broadcasting of poisons of this number is present at one time. The queens are undoubtedly taking their toll. However, seed produced in such colonies may number as many growers in some areas are protecting wild bees as 300, but these are not very useful for pollina- and propagating a few species for pollination. tion until the following year. For most species, Furthermore, there is increasing emphasis nation- 50 to 100 workers during the peak population ally on the development of control measures are exceptionally high. Some rarely have more that minimize the use of insecticides. than 10 to 15. With such small colonies it is questionable whether bumble bees could be Pollination profitably maintained for pollination in the field, although interest in doing so persists. Crops visited freely and pollinated efficiently by sufficient numbers of honey bees have no need A more practical result of bumble bee culture for wild bees. However, under some circumstances may be the accumulation of knowledge about either certain ^'problem" crops are not visited these bees on which to base recommendations freely by honey bees or their visits do not usually for their ])rotection and increase. We know that bring about the thorough pollination desired. Some a continuity of suitable forage is important, but of these crops are efficiently pollinated by wild we know httle about how to provide suitable nesting conditions on a large scale or how to 1 111 cooperation with Utah Agricultural Experiment effect i)ractical control of such enemies as Station. 2 Often known as Apis indica japónica Radoszkowski. Psithyrus, ants, wax moths, and rodents. 109 261-907 0—167- no AGRICULTURE. HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

FIGURE 1.—Species of bumble bees: A, Bombus nevadensis Cresson; B, fervidus (Fabricius); C, huntii Greene; D, griseocollis (De Geer).

Alfalfa Leaf cutting Bee limited by soil conditions and is present in most farming areas within its range, at least in small The alfalfa leafcutting bee Megachile rotun- numbers. data (Fabricius) was accidentally introduced This bee is the smallest leafcutting bee in the into the Eastern United States in the 1930's United States (one-fourth to three-eighths inch and has since spread westward across the northern long). Like most other leaf cutters, it is charcoal two-thirds of the country, reaching Utah and gray and carries its pollen on the underside of California in the early 1950's. the abdomen. Its preferred host range is limited, Alfalfa seed growers have been aware of the consisting primarily of small-flowered forage potential value of this bee since 1950 (fig. 3). legumes, some mints, and a few crucifers. In the Throughout the 1960's, growers in Utah, Idaho, West, sweetclover is the principal host plant, Oregon, Washington, Alberta, Montana, South competing with alfalfa. Unlike other leafcutting Dakota, and California have been following bees, the alfalfa leafcutting bee takes its leaf recommendations of experiment stations in their pieces from herbaceous plants (fig. 4), such as alfalfa, areas and also developing methods of their own rather than shrub and tree leaves, but it occa- for culturing it. Since it nests in many kinds of sionally damages gardens by cutting flower petals. manmade structures, as well as holes in banks, Depending somewhat on their size, the female pores in lava, and other natural cavities, it is not bees choose %î- and %2-inch-diameter holes in BEEKEEPING IN THE UNITED STATES 111

FIGURE 2.—Nest of Bomhus morrisoni Cresson: A, Honey pots; B, pollen tube; C, egg baskets on ooooons (one opened); D, young brood in wax cells. which to nest. Since brood mortaUty is usually The most generally satisfactory nesting units lower in the larger holes, many growers use only devised so far are made of grooved boards stacked large holes. The best hole depth seems to be from together (fig. 5). Planer blades are modified to 4 to 5 inches. cut a series of Ke-inch grooves in boards, which are then strapped tightly together. The principal Nesting Materials advantages of the grooved units are clean holes, Of the three materials most commonly placed resulting from grooving with the grain, and the in the field for their use, alfalfa leafcutting bees ease with which the units can be taken apart to usually prefer holes in wood to bundles of soda clean otit scavenger beetles and pollen-blocked straws, and soda straws to stacks or rolls of holes. corrugated cardboard. Holes cast in foam plastic (polyurethane) are also sold for leafcutter nests Populations in the Northwest, but these are not yet well tested. These preferences are usually demon- One female per five square yards appears to be strated when different nesting materials are adequate for good pollination of alfalfa seed fields. combined near a natural nesting population. If this figure is doubled to allow for the bees not However, since newly emerged females tend to actually foraging at any one time, 100,000 nesting renest in the same kind of nest hole from which females should be able to pollinate a 50-acre field. they emerged, the preferences observed may Since the bees tend to forage within a few hundred largely represent the type of nest inhabited by a natural population. It is not surprising that feet of their nests, they should be placed in shelters growers often have difficulty changing materials distributed throughout the field at about 500-foot and hole sizes from one season to the next. intervals. 112 AGRICULTURE HANDBOOK 3 3 5, U.S. DEPT. OF AGRICULTURE

FIGURE 3.—Alfalfa leaf cutting bee "tripping'' alfalfa florets.

Shelter Designs The shelter should be oriented to receive the early morning sun but protect the nests from direct rays after about 10:00 a.m. It should afford some protection from wind and rain, provide good ventilation, and be conspicuous for the bees. It should accommodate at least 10,000 nesting bees, be placed at least 2}^ feet above the ground, and be built so that covers or screens can be added for protection against pesticides or birds. A box or cupboard supported on legs and pro- tected by a shade board roof is generally satis- factory. It is also advisable to insulate the inner walls of the shelter and to paint the outside white or cover it on the top and the south and west sides with aluminum foil. At the back near the top, screened slots should be provided for ventilation. If birds are a problem, the open side of the shelter should be covered with rabbit wire. Cloth can be used to confine the bees for a few hours after insecticide applications, or until the temperature FiGUKE 4.—Cell of alfalfa leafcutting bee opened to show inside builds up to about 95° F. For longer pro- construction. tection, the shelter box can be placed in a cool, dark room. cellar for overwintering and protection from mice. To prevent a buildup of scavenger beetles Overwintering during storage and to guard against excessive By mid-September in most areas the nesting temperature fluctuations, it is better to keep the materials can be moved to an unheated room or nests (or cells taken from the nests) in cold- BEEKEEPING IN THE UNITED STATES 113 their shelters in the field. Emergence time will ^^^^ßm vary by several weeks, depending on climatic conditions. For a more predictable emergence date, it is better to incubate the nests or loose cells at a nearly constant temperature between 86° and 90° F. until the males begin to emerge (usually about 15 days) before placing them in the field. Water should be added to the incubator to provide humidity. To prevent bees from renesting in boxes with loose cells or in unwanted nesting materials, the nests or cells should be placed in a tight box with one or more long screen cones attached to one side. Bees trying to reenter the box will usually go to the base of the cones. Before placing loose cells in a shelter, it is important that the bees be ready to emerge. Otherwise the pupae are likely to desic- cate. If the bees are being used for the first crop seed and the temperatures are rather low, the shelters should be oriented to the south until shade temperatures reach the high 80's. Moving Adult Bees in Field If the bees are moved after they begin nesting, they will return to their original location. If nothing but alfalfa remains, most of them will return to the new location, especially if it is marked by a large shelter to which they can orient. Several seed growers have successfully used large shelters mounted on trailers, which can be moved to areas needing additional pollination. Protection From Parasites The most serious parasite of the alfalfa leaf- cvitting bee is a metallic-colored chalcid wasp with a long ovipositor (Monodontomerus). The female parasites lay eggs in the host cocoons during the nesting season and also in the spring before the bees emerge. The best time to control them is during incubation. Since the wasps emerge earlier than the bees under constant temperature incubation, a light can be placed over a tray of water in the incubator to attract and then drown them. Unless control measures are used during incubation, a light degree of parasitism in the fall may become almost total

BN-30070 before the bees can emerge in the spring.

FIGURE 5.—Cells of alfalfa leafcutting bee in grooved Protection From Scavenger Beetles board unit (leaf pieces at lower right). A species of grain beetle, Tribolium madens (Charpentier), and several species of carpet storage facilities at 30° to 40° F. Before nests beetles (Trogoderma) infest leaf cutter nests. Both brought from the field are placed in cold storage, groups of beetles can be baited from the nests they should be held at room temperature for and killed with DDT-treated pollen pellets taken about 12 days. from pollen traps on honej^ bee hives. The pellets can be placed under the nesting materials where Incubation the bees cannot contact them or in boards with Approximately 1 month before the bees are shallow grooves or corrugated cardboard with needed for pollination, the nests can be placed in flutes too narrow for the bees to enter. This 114 AGRICULTURE. HANDBOOK 335, U.S. DEPT. OF AGRICULTURE

method of control can be used during incubation gumweed (Grindelia squarrosa (Pursh) Dunal), or nesting and also during cold storage if the sunflower (Helianthus annuus L.), and rabbit- temperature is above 40° F. Studies in Alberta, brush (Chrysothamnus spp.). Canada, indicate that carpet beetles can also be In Utah, Idaho, and Wyoming, adult males controlled during incubation by using the method usually appear in the fields near the first of July. described previously for chalcid wasp control. Females often do not appear in any large numbers until the middle or even the end of July. Males in the field are restless and stop only occasionally to Megachile concinna Smith feed on nectar from flowers. However, they do This species is closely related to the alfalfa trip a high percentage of the flowers visited, and leafcutting bee Megachile rotundata (Fabricius) we have observed fields in which they have been and apparently arrived on the east coast of the the most important pollinators. Females work United States at about the same time. It took a much more efficiently than males. The grower, in southerly route across the country, or possibly order to take full advantage of alkali bees, must time his bloom for late July or early August. Since was independently introduced to Arizona, from there is a partial second generation, some adults which it spread to California. M. concinna differs may be found until September. from M. rotundata as follows: The adults cut Alkali bees construct nest burrows in the soil leaves of shrubs and trees as well as herbaceous from 6 to 10 inches deep (figs. 6 and 7). These bees leaves and flower petals. It presumably has are highly gregarious, but each female builds and greater tolerance for high temperatures. It seems provisions her own nest without help from her to prefer bamboo stems over straws or holes in neighbors. A successful nest contains from 15 to 20 wood. In Arizona its nests are infested with a brood cells, in each of which is placed a ball of small chalcid wasp (Tetrastichus), which seems to honey-moistened pollen and a single egg (fig. 8). be a serious limiting factor. As soon as the egg is laid, the mother bee seals the cell and has no further contact with her offspring. Alkali Bee Most of the nests are constructed and provisioned during July and early August. The larva matures The alkali bee Nomia melanderi Cockerell is by late August and becomes a pupa during the found in scattered localities west of the eastern following June or early July. escarpment of the Rocky Mountains. In Nevada, Wyoming, Idaho, Utah, and the intermountain Characteristics of Ideal Nesting Site areas of Washington and Oregon it ranks with the alfalfa leafcutting bee in importance as an An ideal nesting site is composed of a fine sandy alfalfa pollinator. In Washington, Oregon, Idaho, loam, a well-drained surface, a constant under- and Nevada, seed growers have successfully ground supply of moisture extending upward to established and maintained nesting sites for the surface, a bare or only sparsely vegetated sur- several years by following recommendations of face, and a salty crust. This crust should not be their State experiment stations. thick or hard, and no fluffy dry layer should be Alkali bees are nearly as large as honey bees and under the crust. can be readily recognized by the pale-green or A good nesting site may contain more than a greenish-bronze highly polished bands across the million nests and remain populous for many years. rear part of their bodies. They are found princi- However, after a few years, the population may ])ally in the larger valleys where poorly drained, decline or disappear altogether. The causes, except alkaline areas are prevalent. They are abundant for untimely rains, are at least partially under the in some areas and totally absent in others. They grower's control. Many farming practices that are may appear suddenly in large numbers in one ordinarily desirable, if done without regard for season and disap])ear almost as quickly several alkali bees, can sharply reduce seed yields by years later. damaging the bees' nesting sites. Growers should The host range of alkah bees is rather limited. remember that land occupied by an alkali bee site In the Delta area of Utah, only sweetclover is worth many times the same acreage devoted to (Melilotus si)p.) ^ and Russian-thistle (Salsola crop production. kali L.) ofl^er serious com])etition to alfalfa, al- though a few other less abundant i)lants such as Protection of Nesting Sites saltcedar (Tamarix gallica L.), morning-glory (Convolvulus arvensis L.), and various clovers are Seed growers should find all the nesting sites attractive. Common competitors in the area for within several miles of their fields and take what- honey bees present no attraction for alkah bees. ever steps are needed to protect them. Usually These include greasewood (Sarcobatus vermiculatus nesting sites can be protected most effectively (Hook.) Torr.), i)()vertyweed (Iva axillaris Pursh), on a community basis, since the value of the bees BEEKEEPING IN THE UNITED STATES 115

FIGURE 6.—Female alkali bee at nest entrance. extends beyond property lines, as does the effect Some growers have found that "a penny a fly" of many farming practices, such as spraying and is expensive but worthwhile. Blackbirds and other drainage. Most of the measures involve simple birds are also a serious problem in some areas. control over farming practices. Simply fencing off Scarecrows and firecracker devices soon lose their a nesting site does not necessarily protect it. effectiveness. ... ,-,1 This may permit excessive growth of vegetation, which removes soil moisture and shades the Building and Maintaining Nesting Sites surface. Weeds can be eliminated by mowing or by spraying with herbicides. Skunks are readily Where a hardpan layer exists a foot or more controlled by baiting or trapping. Seed growers under the soil surface, a natural nesting site can can organize and offer a bounty for skunks on an often be built by gently crowning the area and pro- areawide basis. viding it with water in a series of blind, parallel A decline in nest density often results from a irrigation ditches close enough for moisture to year-to-year buildup of fungus (Aspergillus) in seep to the surface from one ditch to the next. the soil. By systematically removing about 20 If the water is run through draintiles laid above percent of the nesting bed to a depth of about 10 the hardpan and the tile has open joints covered inches each year and backfilling with fresh soil, by gravel, a better site is likely to result. it should be possible to maintain relatively "clean" Where a natural hardpan is not present or beds. The removed soil can often be taken out as is insufficiently impervious, plastic film can be "plugs" with overwintering larvae and used or substituted. To construct sites with plastic sold for stocking newly prepared beds. film, a 3}^-foot-deep excavation should be pre- Controlling the bee fly parasite Heterostylum pared with a level bottom and 1-to-l or 2-to-l robustum (Osten Sacken) is difficult (fig. 9). On slope on the walls. The entire excavation should newly established nesting sites the adult parasites be lined with 0.008-inch polyethylene or 0.006- are simply swatted by teams of "vigilantes." inch vinyl film overlaid with 4 to 6 inches of soil 116 AGRICULTURE HANDBOOK 3 3 5, U.S. DEPT. OP AGRICULTURE

Water should be run into the pipes until moisture appears uniformly across the surface of the site. Where the soil is low in salt (sodium chloride), it is necessary to add stock salt at the

-■'■> rate of a pound per square foot to help draw moisture to the surface. The salt can be raked into the top few inches as crystals or dissolved y^i^ in water and sprinkled on the surface. More salt may be required, but it is easier to add it than to remove an excess. The soil should be ST moist when added to the site. The moisture should be kept up to the surface in the site during the entire nesting season. Sometimes only one watering is required before the bees are active. The size of the site depends on the acreage of alfalfa to be pollinated and the resources of the grower. Since it is unlikely that a surplus of wild pollinators will ever be developed in artificial sites, the best recommendation is to continue building and expanding them as time and resources permit. It would be advantageous for groups of farmers to pool their equipment and labor to build sites on an areawide basis. Newly created sites can be colonized by install- ing blocks of undisturbed soil removed from existing sites in the spring. Steel cylinders about 1 foot long made from 12- to 18-inch well casing ^■■^'^ can be sharpened on one end and welded to a /« protective rim on the other. The cyHnder should be cut down one side and provided with clamps to hold it together. It can be driven into a densely populated part of the site with a mallet or pref- erably with the hydraulic drawbar of a tractor. It can then be pried loose against an existing FIGURE 7.—Nest mounds (tumuli) of alkali bees. trench and the soil plug released from it by opening the clamps. The plug can be protected from break- for protection. If the excavation is larger than age by wrapping it in bottle-wrap cardboard the sheets of film, adjacent pieces can be over- held in place with bands made from truck or lapped on berms about 1 foot high placed where tractor inner tubes. Another way to obtain good needed. This procedure creates two or more soil blocks is to saw them out with a chain saw or a water-holding compartments. A 6- to 8-inch layer disk saw attached to a tractor. The sawed blocks of clean gravel should be spread over the protect- can be pried loose with cardboard cartons slipped ing soil. Some site builders lay a grid of draintile over them. in the gravel layer to insure rapid and uniform distribution of water. At 3- or 4-foot intervals, References footwide holes should be dug into the gravel (1) BOHART, G. E. and filled with soil to serve as wicks. The gravel 1952. POLLINATION BY NATIVE INSECTS. In In- layer should then be covered with an inch or two sects. U.S. Dept. Agr. Ybk. 1952: 107- of straw or with plastic screen to keep dirt from 121. plugging the gravel. One-foot-wide strips of (2) 1962. INTRODUCTION OP FOREIGN POLLINATORS, plastic film a foot apart can also be used. The PROSPECTS AND PROBLEMS. 1st Internatl. excavation should then be filled with fine sandy Symposium Pollination, Aug. 1960, Copen- loam until the surface is strongly crowned. At hagen, Proc, Pub. Commun. 7, Swedish each end of the site or of its compartments a Seed Assoc, pp. 181-188. (3) large diameter pipe or tile should extend from 1963. HOW TO MANAGE THE LEAF-CUTTING BEE above the soil surface to several inches deep in FOR ALFALFA POLLINATION. Utah Agr. the gravel layer. The gravel should be mounded Expt. Sta. Cir. 144, 7 pp. around the base of the tile. If a grid of draintile (4) and KNOWLTON, G. F. 1962. MANAGING ALKALI BEES FOR ALFALFA has been laid in the gravel, the standpipe should POLLINATION. Utah State Univ. Ext. connect with it. Serv. Leaflet 78, 2 pp. BEEKEEPING IN THE UNITED STATES 117

FIGURE 8.—Alkali bcc nest (horizontal section at cell level) : A, Egg on pollen ball; B, growing larva; C, diseased larva ; D, adult in main burrow.

(5) and NYE, W. P. LEAF-CUTTER BEE. Canad. Dept. Agr. 1956. BEES. FORAGING FOB NECTJIR AND POLLEN. Pub. 1209, 11 pp. Gleanings Bee Cult. 84: 602-606. (14) NuMMi, W. O., and VIROSTEK, J. F. (6) and NYE, W. P. 1962. MANAGING COLONIES OF BUMBLE BEES 1956. BEES, NESTS AND NESTING SITES. Glean- (HYMENOPTEBA-APOIDEA) FOB POLLINATION ings Bee Cult. 84: 468-472. PURPOSES. Canad. Ent. 94: 1121-1132. (7) and NYE, W. P. (15) HOLM, S. N. 1956. BEES, THEIR TLACE IN THE WORLD OF 1960. EXPERIMENTS ON THE DOMESTICATION OF INSECTS. Gleanings Bee Cult. 84: 265- BUMBLEBEES (BOMBUS LATR.) IN PAR- 268. TICULAR B. LAPIDABNIS L. AND B. TER- (8) and NYE, W. P. BESTRis L. Roy. Vet. and Agr. Col., 1956. BEES. THE TOOLS OF THEIR TRADE. Copenhagen, Ybk. 1960: 1-19. Gleanings Bee Cult. 84: 400-405. (16) KNUTH, P. (9) and NYE, W. P. 1906. HANDBOOK OF FLOWEB POLLINATION. 1956. KINDS OF BEES. Gleanings Bee Cult. (Transi, by J. R. Davis.) 2 v. Clarendon 84: 331-333. Press, Oxford. (10) and NYE, W. P. 1957. BEES. THE NURSERY. Gleanings Bee (17) MEEUSE, J. D. Cult. 85: 82-87. 1961. THE STOBY OF POLLINATION. 243 pp. Ronald Press, New York. (11) and PEDERSEN, M. 1963. THE ALFALFA LEAF-CUTTING BEE, MEGA- (18) MiCHENEB, C. D. OHILE ROTUNDATA (F.), FOR POLLINATION 1944. COMPARATIVE EXTERNAL MORPHOLOGY, OF ALFALFA IN CAGES. Crop Soi. 3: 183- PHYLOGENY, AND CLASSIFICATION OF THE 184. BEES (HYMENOPTERA). Amer. Mus. Nat. (12) and ToDD, F. E. Hist. Bui. 82, pp. 151-326. 1961. POLLINATION OF SEED CROPS BY INSECTS. (19) MuESEBECK, C. F. W., KBOMBEIN, K. V., TOWNES) In Seeds. U.S. Dopt. Agr. Ybk. 1961: H. K., and others. 240-246. 1951. HYMENOPTERA OF AMERICA NORTH OF (13) HoBBS, G. A. MEXICO SYNOPTIC CATALOG. U.S. Dcpt. 1965. IMPORTING AND MANAGING THE ALFALFA Agr. Agr. Monog. 2, 1420 pp. 118 AGRICULTURE HANDBOOK 33 5, U.S. DEFT. OF AGRICULTURE

(20) NOGUEIBA-NETO, P. 1953. A CRIAÇlo DE ABELHAS INDÍGENAS SEM ♦ Hni * FEBHAo (MELIPONINAE). 280 pp. Chá- caras c Quintais, Sao Paulo, Brazil. (21) PLATH, O. E. ""*-*-'& EII^HSIHâ i^ttk 1934. BUMBLEBEES AND THEIB WAYS. 210 pp. Maomillan Co., New York. S'M .'^H (22) SLADEN, F. W. L. HHII 1912. THE BUMBLE-BEE. 283 pp. Macmillan - "-^^I Co., London. (23) STEPHEN, W. P., and EVANS, D. D. 1960. STUDIES ON THE ALKALI BEE (NOMIA MELANDEBI CKLL.). Oreg. Agr. Expt. Sta. Tech. Bui. 52, 39 pp. Í M '"^9H '' ^H^^L

£^^^^1 L „„„.a&^^^Wl^ m^■pr ■■B 1^Ml IF^ ^^

FIGURE 9.—Third-instar larva of bee fly parasite feeding on alkali bee prcpupa. (Artificial cell—fecal pellets not in natural position.) FEDERAL AND STATE BEE LAWS AND REGULATIONS

By A. S. MICHAEL, Assistant Chief, Apiculture Research Branch, Entomology Research Division, Agricultural Research Service

The Federal Government has no laws or passed by the California legislature, and by 1906, regulations pertaining to honey bees or beekeep- 12 States had laws relating to foulbrood. At ing within the United States. However, on August present, almost all States have laws regulating 31, 1922, Congress passed a law, popularly known honey bees and beekeeping. as the Honeybee Act, restricting the importation State laws and regulations relating to honey of living adult honey bees into the United States. bees and beekeeping are designed primarily to This act was amended in 1947 and again in 1962. control bee diseases. Therefore, they usually This last amendment, Public Law 87-539, dated attempt to regulate movement and entry of bees, July 19, 1962, resulted in the establishment of issuances of permits and certificates, apiary loca- the following regulations covering all living tion control and quarantine, inspection, and adults of the genus Apis: methods of treating diseased colonies. These laws and regulations are summarized in tables 1 and The importation into the United States of adult honey bees is prohibited and all adult 2. Table 1 is a compilation of the bee laws for honey bees offered for entry shall be destroyed intrastate regulation; table 2 is a compilation of if not immediately exported, with the following bee laws regulating interstate movement of bees exceptions. and used bee equipment in the United States. No disease dangerous to adult honey bees exists in Canada other than those already present As will be noted from these tables, there is in the United States and adequate precautions a lack of uniformity in State bee laws and regu- have been taken to prevent the importation of lations, but considerable agreement on specific adult honey bees into Canada from countries points of law. Most of the States require regis- where such dangerous diseases are known to exist. Therefore, the unrestricted importation tration of apiaries, permits for movement of bees of adult honey bees from Canada by any person and equipment interstate, certificates of inspec- is authorized. tion, right of entry of the inspector, movable- Importation of adult honey bees from any frame hives, quarantine of diseased apiaries, country other than Canada shall be conditioned notification of the owner upon finding disease, upon a determination that no such diseases dangerous to adult honey bees exist in the prohibition of sale or transfer of diseased material, country in question and that adequate pre- and use of penalties in the form of fines or jail cautions have been taken by that country to or both. Although the destruction of American prevent the importation or entry of adult honey foulbrood diseased colonies is included in most bees from countries where such dangerous diseases exist. The determination shall be based State laws, table 1 shows that most States also upon adequate investigations by the Agricultural allow the use of drugs for control or preventive Research Service of the United States Department treatment of this disease. of Agriculture. In the absence of substantial The key figure in the enforcement of bee laws evidence that diseases dangerous to adult honey bees do not exist in the country in question or and regulations is the apiary inspector. He may that adequate precautions have been taken by have the entire State, a county, or a community the country in question to prevent the importa- under his jurisdiction. His efforts are directed tion or entry of adult honey bees from countries toward locating American foulbrood and elimi- where such dangerous diseases exist, importa- tion is prohibited. If under the conditions existing nating sources of it whenever found. in areas surrounding the country in question, The effectiveness of bee laws and regulations adequate precautions cannot be taken by such is based on the compliance of the beekeepers. country to prevent the entry of adult honey In the final analysis, responsibility for disease bees from countries where diseases dangerous to adult honey bees exist, importation is pro- control remains with the beekeeper, who should hibited. routinely examine colonies for disease as a regular The Agricultural Research Service, United part of his management program and take the States Department of Agriculture, may import necessary steps when disease is found. into the United States from any country adult honey bees for experimental or scientific purposes. Copies of State apiary laws and regulations are available from the various State departments The first apiary inspection law in the United of agriculture. State apiary inspectors can also States was established in San Bernardino County, be consulted in care of their State department Calif., in 1877. By 1883, a statewide law was of agriculture. 119 120 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

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By S. E. MCGREGOR, Chief, Apiculture Research Branch, Entomology Research Division, Agricultural Research Service

or cooperative agreements (about 3 years) at Federal Research State universities. These institutions and studies Research on honey bees by the U.S. Department are as follows : of Agriculture was initiated in 1885 and has been University of Arizona, Tucson.—Innate and continuous since 1891. It is presently under the acquired immunity in honey bees. direction of the Entomology Research Division, University of Illinois, Urbana.—Behavioral study Agricultural Research Service, with headquarters of effect of hormonal secretions of queen honey bee at Plant Industry Station, Beltsville, Md. This {Apis mellifera L.) on industriousness of worker agency often acts as technical adviser to bee- honey bees. related industries and is a source of scientific Michigan State University, East Lansing.— and popular publications on bees. The research Honey bee pollination requirements of hybrid studies conducted in six field laboratories are as cucumbers. follows : Ohio State University, Columbus.—Pathogenesis Arizona, University of Arizona,^ Tucson.—Pol- and diagnosis of nosema disease in Apis mellifera lination of various crops, bee nutrition and L. physiology, bee behavior, and effect of pesticides Utah State University, Logan.—Glands in bees, on bees; cooperative study between Entomology their topography, innervation, morphology, his- Research Division and Agricultural Engineering tology, and physiology. Research Division on mechanization of beekeeping Research is also supported abroad with Public methods and equipment. Law 480 funds, popularly known as P.L. 480 or Louisiana, Louisiana State University,^ Baton the Food for Peace Program. This research in- Rouge.—Bee breeding and bee genetics, nosema cludes the following studies : disease, nectar and pollen plants, and whiteclover Egypt.—Biology, ecology, and utilization of in- pollination. sects other than honey bees in pollination of Maryland, Agricultural Research Center, U.S. agricultural crops. Department of Agriculture, Beltsville.—Bee dis- India.—Biology, ecology, and utilization of in- eases, their control, and relationship of bee nu- sects other than honey bees in pollination of trition to bee diseases; disease diagnostic serv- agricultural crops. ice for beekeeping industry and Bee Culture Italy.—Acariñe disease of honey bees. Branch of National Agricultural Library located Poland.—Distance of mating flights of honey here (see p. 128). bee queens and drones, necessary isolation of Utah, Utah State University,^ Logan.—Commer- mating stations to prevent mismatings, and cial use of wild bees as pollinators, effect of biology and utilization of diploid drone honey bees. pesticides on both honey bees and wild bees, The agencies of the U.S. Department of Agricul- and honey bee pollination. ture concerned in some way with bees, honey, or Wisconsin, University of Wisconsin,^ Madison.— wax and their areas of interest are as follows : Productive management of colonies for high Agricultural Research Service, Agricultural En- yields, use of two-queen colonies, bee disease gineering Research Division, Farm Electrification control, queen rearing, feeding of pollen supple- Research Branch.—Mechanization of beekeeping ments; cooperative study between Entomology and bee equipment, honey processing, and handling Research Division and Agricultural Engineering (see pp. 37-51); Entomology Research Division, Research Division on honey handling and proces- Agriculture Research Branch.—Bee research inves- sing. tigations ; Pestócíáe Chemicals Research Branch.— Wyoming, University of Wyoming,^ Lar amie.— Determines toxicity of pesticides to bees under Bee diseases in open colonies, mode of transmis- laboratory conditions; Plant Pest Control Division, sion of bee diseases within and between colonies, Pesticide Safety and Monitoring.—Monitoring long- disease resistance, and bee poisoning. term effect of pesticides on bees; Plant Quarantine The U.S. Department of Agriculture also Division, Program Development and Service Staß, supports research on bees in the form of grants Technical Services.—Enforces Federal Honey Bee Act, which restricts entry of adult honejr bees to prevent introduction and spread of certain bee 1 Cooperator. diseases (seep. 119). 125 '261-907 0—67- -9 126 AGRICULTURE HANDBOOK 335, U.S. DEFT. OF AGRICULTURE

Agricultural Stabilization and Conservation Georgia, Coastal Plain Experiment Station, Service, Sugar Policy Staff.—Focal point in Tifton.—Controlling pests of honey bee and mis- formulation and recommendation on honey price cellaneous bee problems. support program; Policy and Program Appraisal Illinois, University of Illinois, Urbana.—Bee Division, Price Program Development Staff.— behavior and value of bees in crop pollination. Prepares price support and other national pro- Maryland, University of Maryland, College grams for honey (see p. 136); Farmer Programs Park.—Nutrition and caste determination in honey Division, Price Support Branch.—Writes program bees. regulations and provisions under which county New Jersey, Rutgers—The State University, ASC committees administer program; Procure- New Brunswick.—Blueberry pollination and bee ment and Sales Division.—Supervises taking title diseases. to unredeemed honey under Commodity Credit New York, Cornell University, Ithaca.—Bee Corporation price support loan, and processing, behavior, pollination, pesticides, and honey. sales, and other disposition of such honey as may be necessary; Minneapolis (Minn.) ASCS Com- Ohio, Ohio State University, Columbus.—Genetics modity Office.—Responsible for acquiring title to of American foulbrood resistance in honey bees. price supported honey and for its storage and Oregon, Oregon State University, Corvallis.— disposition. Bee behavior and pollination. Consumer and Marketing Service, Fruit and Pennsylvania, Pennsylvania State University, Vegetable Division, Market News Branch.—Pre- University Park.—Bee venom and bee diseases. pares monthly Honey Market News report; Washington, Washington State University, Pull- Processed Products Standardization and Inspection man.—Pollination and effects of pesticides on bees. Branch.—Grade standards and inspection of Occasional research, as situations arise, is con- honej', Specialty Crops Branch.—Honey Marketing ducted on bees in the following States: Aids (see p. 134). Alaska, University of Alaska, College.—Improv- Federal Extension Service, Agricultural Science, ing efficiency of Alaska's insect pollinators. Technology and Management Division.—Extension Arizona, University of Arizona, Tucson.—Pollen entomologist advises State entomologists on api- substitutes. culture matters. Arkansas, University of Arkansas, Fayetteville.— Foreign Agricultural Service, General Sales, Honey and pollen plants. Export Sales and Pricing, Tobacco, Peanuts, Naval Idaho, University of Idaho, Moscow.—Pollina- Stores, and Honey.—Foreign sales of surplus prod- tion of vegetable seed crops. ucts; Commodity Programs, Sugar and Tropical Indiana, Purdue University, Lafayette.—Bee Products Division, Commodity and Products Analy- behavior in flight room. sis Branch.—Production and world supply of honey Massachusetts, University of Massachusetts, (seep. 143). Amherst.—Effect of pesticides on honey bees. National Agricultural Library, Field and Special Michigan, Michigan State University, East Services, Division of Field Services, Bee Culture Lansing.—Pollination. Branch (see p. 128). Minnesota, University of Minnesota, St. Paul.— Statistical Reporting Service, Agricultural Esti- Bee nutrition and methods of bee management. mates Division, Livestock, Dairy and Poultry Rhode Island, University of Rhode Island, Branch.—Statistics on bees and honey (see p. 139). Kingston.—Bee diseases. Utah, Utah State University, Logan.—Bee anat- omy, pesticides, wild bees, and pollination. State Research Wisconsin, University of Wisconsin, Madison.— Biology of bumble bees, pollination, and bee- The States with one or more full-time apicul- management studies. turists and their areas of research are as follows : One or more courses in beekeeping are taught California, University of California, Davis.— in universities or colleges in Alabama, Arizona, Breeding and genetics of honey bees, mechanism Arkansas, California, Colorado, Florida, Illinois, of bee behavior, pollination, and bee mites; River- Kansas, Louisiana, Maryland, Massachusetts, side.—Laboratory and seasonal field tests on effect Michigan, Minnesota, Mississippi, Missouri, Ne- of pesticides on bees; Santa Barbara.—Communi- braska, New Jersey, New York, North Carolina, cation in honey bees. North Dakota, Ohio, Pennsylvania, South Caro- Florida, University of Florida, Gainesville.— lina, South Dakota, Utah, Virginia, and Wisconsin. Pollination, pollen substitutes, and bee disease California, Florida, Ohio, and Pennsylvania control; Vineland.—Watermelon pollination by have full-time extension bee specialists. Part-time honey bees. extension work by apiculturists is conducted in BEEKEEPING IN THE UNITED STATES 127

Illinois, Maryland, Michigan, Minnesota, New Service employs one extension entomologist, who Jersey, and New York. Attention in varying devotes part time to apiculture (see p. 126). degrees is given to apiculture by extension ento- Almost every State has some form of apiary mologists in other States. The Federal Extension inspection service (see p. 119). COLLECTIONS OF BEE LITERATURE

By JULIA S. MERRILL.^ librarian, Bee Culture Branch, National Agricultural Library

The United States is fortunate to possess a With this index, information on almost any good share of the bee literature of the world. subject relating to bees can be located. For The best known collections in the United States instance, there are over 2,200 references on the are at the U.S. Department of Agriculture, Cornell role of the honey bee in plant pollination, 1,300 University, and the Universities of California, cards on the toxicity of insecticides to bees, and Minnesota, and Wisconsin. Canada has a sizable numerous references on such diversified subjects collection at the University of Guelph in Ontario. as bee anatomy, behavior, genetics, diseases, honey, beeswax, venom, colony management, and U.S. Department of Agriculture State and Federal laws relating to bees. In addition to the bee literature, references The Bee Culture Branch of the Department's for inclusion in the Beekeeping Bibliography are National Agricultural Library at Beltsville, Md., obtained from medical, biological, agricultural, has one of the largest collections of bee literature and other nonbeekeeping books and periodicals. in the United States. It was started in the 1920's Abstract journals are covered as well as the by James I. Hambleton of the former U.S. Bibliography of Agriculture and Pesticides Docu- Bureau of Entomology. mentation Bulletin, which are monthly publica- This library now has about 6,500 bound tions of the National Agricultural Library. Each volumes, over 2,000 scientific reprints, and about reference is filed by author and subject under 500 translations. It currently receives 121 periodi- one or more of the 28 major classifications and cals relating to beekeeping, 79 of which are in over 200 subheadings. Special bibliographies are foreign languages. In addition to current books prepared on request for Department of Agri- and periodicals relating to fundamental or applied culture personnel and for others as time permits. research, this library has several volumes of The Bee Culture Branch has a translator. historic value. Most of the older books added in Translations from Dutch, French, German, Rus- recent years have been obtained through gift or sian, and Spanish are prepared. The original exchange. The library receives $1,000 annually remains in the library, and a copy is sent to for the purchase of books and periodicals. Ad- each apiculture laboratory of the Department. ditional funds are sometimes made available. Copies are also sent to the National Agricultural To facilitate reference service and literature Library, Washington, D.C., and the Clearing searches, an annotated card index of selected House for Federal Scientific and Technical Infor- references from the world's literature on apicul- mation, Springfield, Va. Translations are ex- ture and closely related subjects was started in changed with the Department of Apiculture, the 1920's. This Beekeeping Bibliography, as it University of Guelph, Ontario, Canada; the is familiarly known, is currently maintained as Apiculture Section, Canada Department of Agri- a joint project of the Bee Culture Branch of the culture, Ottawa; and the Bee Research Associ- National Agricultural Library and the Entomology ation, an international organization for the Research Division, Agricultural Research Service. furtherance of bee science, Gerrards Cross, Buck- References are added monthly, and copies are inghampshire, England. No list of translations sent to the apiculture field laboratories of the is available, but each is noted in the Department's Department of Agriculture in Tucson, Ariz., Baton Bibliography of Agriculture. Rouge, La., Logan, Utah, Madison, Wis., and Though the essential function of this library Laramie, Wyo. is to serve the U.S. Department of Agriculture bee research staff, anyone is welcome to use the ^ Grateful acknowledgment is made to the following library. Publications may be borrowed on inter- people for supplying statistical and historical information: library loan, but no individual loans are made J. Richard Blanchard, librarian, University of California, Davis; Mrs. Evelyn L. Gish, entomology Hbrarian, Uni- to other than Department personnel. Photoprint versity of Minnesota, MinneapoUs; Emory M. Pittenger, and microfilm copies of articles or translations agricultural Hbrarian, University of Wisconsin, Madison; may be ordered from the Photoduplication Sec- Whiton Powell, librarian, Albert R. Mann Library, Cornell tion, National Agricultural Library, U.S. Depart- University, Ithaca, N.Y.; and Gordon F. Townsend, Head, Department of Apiculture, University of Guelph, ment of Agriculture, Washington, D.C. 20250. Ontario, Canada. Photoprints cost $1 for each four pages, or fraction. 128 BEEKEEPING IN THE UNITED STATES 129 from a single book or article, and microfilm costs because of their age or because they are out of $1 for each 30 pages, or fraction. print. Over 600 bound periodicals, representing 55 beekeeping titles, are on file, and six American Cornell University and 18 foreign bee journals are currently received. The beekeeping collection at this library con- The Phillips Memorial Beekeeping Library at tains an extensive group of books from eastern Cornell University, Ithaca, N.Y., has valuable Europe. No actual count has been made, but and beautifully bound books acquired b}^ the such countries as Austria, Czechoslovakia, and late Everett F. Phillips when he was in charge Russia are well represented. of apiculture research at the U.S. Department of Agriculture and w^hile he was Professor of University of Wisconsin Apiculture at Cornell University. Included in this collection are such items as the original The C. C. Miller Memorial Beekeeping Library patent granted to Lorenzo Lorraine Langstroth, at the University of Wisconsin in Madison was 'Hhe father of modern beekeeping,'^ for his started in 1923, 3 years after the death of movable-frame hive, his diary and letters written Charles C. Miller, as a memorial to that nationally in his own hand, and original manuscripts of known beekeeper, teacher, investigator, and several editions of Moses Quinby's Mysteries of writer. In 1922, his admirers presented to the Beekeeping Explained. These, along with manu- regents of this university a fund of $1,957, the scripts of Dr. Phillips' own books and 1,000 more income of which was to be used for the mainte- volumes printed before 1900, are kept in locked nance of a beekeeping library. The current income cases on a balcony overlooking the reference amounts to about $250 per year and is used only room of the Albert R. Mann Library at Cornell tu augment the library, not for maintenance or University. The bee periodicals, books, and pam- other library expenses. phlets published in the 20th century are shelved In this library is the Walker Collection, which with the general collection in the stacks. was obtained with funds given by the late Sigurd The budget for sustaining the Phillips collection L. Odegard of Madison. The nucleus of this col- currently amounts to $700. This is the income lection was a group of late 19th century books from an endowment fund derived partly by gifts owned by Alfred Neighbour, an English bee from beekeepers and partly from the Dyce Honey appliance manufacturer and author. Upon his Crystallization process patent. The patent was death in 1890, they were bought by H. J. O. given to Cornell University by its owner, Elton Walker of Devon, England. Colonel Walker went J. Dyce, now retired, who was Dr. Phillips' to great length to add to this collection every successor. important item on bees up until 1930, when he gave up the project and sold the collection. University of California From the time the books arrived for the Miller The University of California at Davis has 1,630 Library until his retirement in 1948, H. F. Wilson books and bound periodicals on bees, 66 of which of the Department of Economic Entomology was are rare volumes. It currently receives 32 foreign custodian. Under his guidance the collection grew and seven American bee journals. Noteworthy to what it is today. As early as 1930, a conservative is the John S. Harbison manuscript collection estimate of its value was $12,000. For niuch of dating from 1857 to 1912. Harbison was one of the the material. Professor Wilson used his own first beekeepers to import bees into California. money. He kept records of money received from This collection includes his records of these ship- beekeepers who donated a 10-pound pail of honey, ments from New York to Sacramento by boat, or its equivalent in cash, each year for a fund he freight receipts, records of other business trans- sponsored to buy books. These records are actions, and patent grants. preserved as a part of the collection. The Miller Library is housed in a special locked area of the Agricultural Library of the University University of Minnesota of Wisconsin. It consists now of approximately The collection of bee literature at the University 5,500 volumes, most of them beautifully bound, of Minnesota in Minneapolis was begun about gold tooled, and in excellent condition. Section 4 1900 by Father Jager, a Catholic priest. The of the agreement between the University and Division of Beekeeping, of which he was head, the Committee of the C. C. Miller Memorial was then a one-man department. In 1928, the Beekeeping Library, dated September 16, 1922, Division was consolidated with the Department states: 'Tn the case of rare books or journals, of Entomology and Economic Zoology, and Father the University of Wisconsin agrees to retain two Jager's books were cataloged for the Entomology copies of such books or journals to secure the Library. Approximately 850 bee books are now on Library in case of loss of one copy. This shall hand, 150 of which are kept under lock and key apply to books or journals published in the United 130 AGRICULTURE, HANDBOOK 33 5, U.S. DEFT. OF AGRICULTURE

States more than 50 years previous to the time loan service may be arranged through the main they are received, or to books and journals library of the University of Guelph. pubHshed in foreign countries more than 20 The extensive reprint collection serves as th- years previous to their receipt/' Within the scope ^^American Branch" of the Bee Research Ase of this definition, 2,000 vokimes are considered sociation Library. Reprints are available for loan rare. Over 50 bee periodicals are received currently, to American members of the Bee Research three-fourths of which are foreign. Association at no cost other than the return The Miller Library has a historical collection postage. of note, which may be used by anyone with the A full-time employee is responsible for main- permission of the librarian. Journals and rare taining the Guelph collection. The funds for books are not loaned, but copies of most items are salary and books are included in the overall available. budget for the Apiculture Department. Also a trust fund is available for book purchase, and additional money can be allocated for special University of Guelph purchases. The University of Guelph, Ontario, Canada, Though there is duplication among the literature has a collection of bee literature. It consists of collections mentioned here, items in each are approximately 1,700 books, several of which are unique. No serious effort has been made to obtain of historical value; 800 volumes of bound and a ''Union List'' of the holdings of all the significant current periodicals, 46 of which are currently collections of bee literature on this continent, subscribed to; and over 5,000 beekeeping reprints but this might be a goal for the future. Such a and translations. These are housed in the Aj^icul- project, when completed, would be of inestimable ture Building at the University and are available value to apiculture students, research personnel, to interested persons for consultation. Interlibrary and libraries throughout the world. NONGOVERNMENT BEEKEEPING ORGANIZATIONS

By J. D. HITCHCOCK, apiculturistj Entomology Research Division, Agricultural Research Service

National Organizations 1953: Adopted a honey house sanitation schedule. 1954: Obtained congressional appropriations for The first national organization of beekeepers in research on pollination and on mechanization of the United States was the American Bee Asso- beekeeping equipment. ciation established at a convention in Cleveland, 1959: Sponsored a national honey queen contest Ohio, in 1860, but it was soon disrupted by the to be held annually, based on beauty and person- Civil War. In 1871, two beekeepers' organizations, ality, the winning girl then traveling across the both having the Rev. L. L. Langstroth as their Nation on a honey promotion tour. president, merged at a meeting, also in Cleveland, The federation through its Honey and Pollen to form the North American Bee Keepers' Society. Plants Committee has sponsored eight pollination Some type of national organization, including the conferences as follows: 1945 and 1946: Atlantic, American Honey Producer's League (1924-42), Iowa; 1947: Amherst, Mass.; 1948: Lincoln, Nebr.; has been in existence continuously since then, 1949: Seattle, Wash.; 1950: Tucson, Ariz.; 1951: although the name has been changed eight times. Ardmore, Okla.; 1965: College Station, Tex. The present national organization of honey The federation has also cooperated with the producers essentially began with the National U.S. Department of Agriculture in exhibiting Federation of State Beekeepers' Associations in honey at various international food fairs in foreign 1943. The present name, American Beekeeping countries, such as those at Hamburg, Germany, Federation^ was first applied in 1949, and the or- in 1961; Brussels, Belgium, in 1962; Amsterdam, ganization was incorporated in 1954. Annual Netherlands, in 1963; and London, England, in meetings have been held every year since 1871, 1965. Such promotion helps the honey export except during 1 or 2 war years. Membership in the trade. It has also sponsored national honey shows national organization exceeded 5,000 in 1911, but in the United States annually since 1953. dropped to less than 200 during World War I. In The American Bee Breeders^ Association is a 1965, there were 1,372 members in the federation. national organization of queen breeders, formed Its purpose is to promote the common interests in 1948 at a meeting in Meridian, Miss. Its pri- and general welfare of the beekeeping industry. mary objective is to improve the quality of honey The first national publication was the Beekeepers' bees in use in this country. It also has established Review, published in 1912. The present bimonthly standards so that its members can advertise publication is the Federation News Letter, now under the association's emblem as a sign of in its 24th year. reliability. It has about 70 members. Some of the outstanding accomplishments of The American Honey Institute was organized the national beekeepers' organization are as follows: in 1928. Its purpose is to promote the use of 1879: Persuaded the U.S. Post Office to permit honey. To achieve this it has often cooperated sending queen bees through the mails. with other industries for the mutual promotion 1906: Lobbied for congressional passage of the of honey with other foods. Its headquarters were Pure Food and Drug Act, to prevent adulteration first in Indianapolis, Ind., then transferred to of honey. Madison, Wis., in 1932, and to Chicago, 111., in 1922: Lobbied for congressional passage of the 1965. In addition to thousands of leaflets, book- Honeybee Act, which prohibited importation of lets, streamers, and other articles giving honey adult honey bees into North America, in order to recipes tested in its own kitchen, the institute prevent introduction of acariñe disease. has published three books: Old Favorite Honey 1924: Published a treatise on the law pertaining Recipes (1941), New Favorite Honey Recipes to the honey bee. (1947), and More Favorite Honey Recipes (1956). 1936: Urged establishment of a Federal bee It has maintained close contact with leading research laboratory in the whiteclover region, re- home economists and thus has increased distri- sulting in the North Central States laboratory, bution of its honey recipes. Attractive photo- now the Bee Management Investigations labor- graphs and eye-catching color pictures form an atory at Madison, Wis. important part of its advertising campaign. In 1949: Obtained a Federal Honey Price Support addition to a number of sustaining organizations, Program (see pp.136-138). headed by the Honey Industry Council of Amer- 131 132 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

ica, many individual beekeepers contribute shire, New Jersey, New York, Ohio, Pennsylvania, toward its finances. Rhode Island, Vermont, Virginia, West Virginia, The Apiary Inspectors of America held national and Ontario, Canada. AH members in these as- meetings at various times, at least as early as sociations, over 2,600 in 1966, are considered 1906, when E. F. Phillips presided at a meeting members of the Eastern Apicultural Society. in San Antonio, Tex. However, their present The Northwest Beekeepers' Association, primarily organization began in 1928, under the leadership beekeepers of Idaho, Oregon, and Washington of R. L. Parker of Kansas, at a meeting of the State, has had biennial conferences since 1957. American Honey Producers' League—then the These are usually attended by about 50 to 100 national beekeepers' organization—in Sioux City, beekeepers and, like the Eastern Apicultural Iowa. The purposes of this organization are (1) Society, it obtains nationally prominent speakers to promote better beekeeping conditions through from both industry and science. uniform, effective laws and methods for the Similarly beekeepers from Arkansas, Colorado, suppression of bee diseases and (2) to seek mutual Iowa, Kansas, Missouri, Nebraska, and Oklahoma cooperation between apiary inspection officials of have joined in annual Seven States Beekeepers different States. Most State laws apply to Amer- regional meetings since 1962. ican foulbrood and require certification that bee The Southern States Beekeepers' Federation was colonies are free of disease before they can be organized at a meeting in Texarkana, Ark., in moved across State lines, as in migratory bee- 1928. It now has about 600 members from about keeping. There were 36 members in 1966. The 20 States. Since its purpose is to promote southern organization publishes proceedings of its annual beekeeping, it is involved primarily with problems meetings. of the package-shipping and queen-rearing phases The Bee Industries Association is a national of the beekeeping industry. It has supported organization of manufacturers of beekeeping research on such problems at universities in supplies and equipment. It was started in 1941 Mississippi and Louisiana. The organization is and has about a dozen member firms. Its purpose afBliated with the American Beekeeping Fed- is to try to standardize beekeeping equipment eration. and to promote the welfare of the beekeeping The Tidewater Beekeepers' Association, com- industry. Outstanding services include its inter- prised of beekeepers from the Southeastern cession in Washington for high priority materials Atlantic Seaboard States, was organized in 1939. for the beekeeping trade during World War II Another example of several adjacent States and lending substantial financial aid to beekeeping getting together for beekeepers meetings is the groups. Four States' Beekeepers of Iowa, Minnesota, The Honey Industry Council of America was Nebraska, and South Dakota (1952). established in 1953. It coordinates the overall plans of the beekeeping industry as a whole. Its Check-Off Plan, whereby both producers and Representative State and Local processors allocate a certain amount of money Organizations per unit volume of honey, provides funds used to advertise honey and promote research. The California Bee Breeders, Inc., was or- The national Honey Packers^ and Dealers^ ganized in 1933 to promote the production of Association was organized in about 1950 and re- high quality queens and package bees in California. organized in 1953. Its members are primarily It has about 50 members. large-scale honey processors. Its purpose is to The California Honey Advisory Board is a unique improve methods of processing and packaging State organization operating under the California honey and to increase its sale. Department of Agriculture. It was organized in The Royal Jelly Research Foundation was 1952. Its purpose is to promote the use of honey organized in 1958 at a meeting of 27 royal jelly by research and advertising. It appears to be producers and dealers in Valdosta, Ga. Its purpose doing on a State scale what the American Honey is to support research on the nutritional and Institute does on a national scale. It has its own therapeutic properties of royal jelly. recipe-testing kitchen. It also supports research programs at various branches of the University Some Regional Organizations of California. It prints numerous honey recipe leaflets and other educational information about The Eastern Apicultural Society was founded honey. These are provided free of charge to the at the University of Maryland in 1955, and State's honey producers and packers for distri- incorporated in 1962. It was an outgrowth of bution with their honey sales. It has published associations of beekeepers from Connecticut, three honey recipe booklets, Honey's Nifty Fifty Massachusetts, and Rhode Island. It now includes Recipes (1958), The Best From the West With associations in Connecticut, Delaware, Kentucky, Honey (1962), and Honey: It's Good Everyway Maine, Maryland, Massachusetts, New Hamp- Every Day (1965). BEEKEEPING IN THE UNITED STATES 133

There are State Beekeepers^ Associations in (6) DADANT, M. G. 1964. THE SOUTHERN STATES BEEKEEPING FED- almost every State in the United States. The ERATION. Amer. Bee Jour. 104: 294-295. oldest is in Michigan established in 1865. At (7) DEYELL, A. least 15 other States organized beekeepers^ as- 1956. THE AMERICAN BEEKEEPING FEDERATION sociations before 1900, including California, Colo- (HISTORY). Gleanings Bee Cult. 84: 161. (8) GROUT, R. rado, Connecticut, Illinois, Kentucky, Maine, 1949. THE AMERICAN BEEKEEPING FEDERATION: Minnesota, New York, Ohio, Tennessee, Texas, OBJECTIVES AND ACCOMPLISHMENTS. Vermont, Washington, and Wisconsin. Perhaps Gleanings Bee Cult. 77: 144-145, 149, 173. the most recent is in Nevada organized in 1959. [Also anonymously in Amer. Bee Jour. 89: 132-133.] Most State associations hold annual nieetings (9) HAEGER, P. during the winter. Several also hold summer 1965. FIRST STEPS TOWARD CONSUMER EDUCATION picnic meetings. Most associations distribute (BY THE AMERICAN HONEY INSTITUTE) . some type of newsletter to their members. Gleanings Bee Cult. 93: 749. (10) JONES, A. 1965. EASTERN APICULTURAL SOCIETY TEN YEARS OLD. Amer. Bee Jour. 105: 260-261. (11) JONES, G. O. References 1948. THE PROGRAM AND PURPOSES OF THE NATIONAL FEDERATION (OF STATE BEE- (1) ANONYMOUS. KEEPERS' ASSOCIATIONS) . Amer. Bee Jour. 1929. THE LEAGUE MEETING (AN ORGANIZATION 88: 496, 505, 543, 594-595. OF THE APIARY INSPECTORS OF AMERICA) . (12) KREBS, H. M., DAVIS, J. H., FLOYD, C. D., BAILEY, Gleanings Bee Cult. 57: 139. S. E. (2) 1950. RESOLUTIONS ADOPTED BY THE APIARY 1943. BEE INDUSTRIES ASSOCIATION OF AMERICA. INSPECTORS OF AMERICA. Gleaniugs Bee Amer. Bee Jour. 83: 105. Cult. 78: 155. (3) (13) MiLUM, V. G. 1953. HONEY INDUSTRY COUNCIL OF AMERICA. 1964. HISTORY OF OUR NATIONAL BEEKEEPING Amer. Bee Jour. 93: 223. ORGANIZATIONS. 88 pp. American Bee- (4) keeping Federation, Urbana, 111. 1965. MICHIGAN BEEKEEPERS' ASSOCIATION CELE- (14) PuETT, G., and WILLSON, R. B. BRATES 100 YEARS. Gleanings Bee Cult. 1958. ROYAL JELLY RESEARCH FOUNDATION OR- 93: 534-535, 570. GANIZED. Amer. Bee Jour. 98: 193-194. (5) BESSONET, E. C. (15) RASMUSSEN, P. 1948. AMERICAN BEE BREEDERS' ASSOCIATION. 1948. THE AMERICAN HONEY INSTITUTE. Glean- Amer. Bee Jour. 88: 184-185. ings Bee Cult. 76: 609-613. HONEY MARKETING AIDS

By J. S. MILLER, marketing specialist, Fruit and Vegetable Division, Consumer and Marketing Service The Consumer and Marketing Service (C&MS) Quality Standardization serves as the focal point for honey marketing activities of the U.S. Department of Agriculture. QuaUty standardization provides a means of This agency frequently serves as liaison between uniformly classing quality of the honey being the honey industry and the various other agencies marketed. Furthermore, it is important since of the Department. C&MS provides important consumers prefer to buy on a quality basis. The marketing aids for use of the honey industry. U.S. Department of Agriculture has issued quality grade standards for extracted and comb honey. Current Marketing Information Use of the standards is not compulsory. They are designed to provide a convenient basis for sales, Marketing of honey is nationwide since pro- for establishing quality control programs, and for duction occurs in all States. It is estimated that determining loan values. The standards also serve approximately one-half of each year's honey as a basis for inspection of honey by C&MS and production is sold by the producer direct to con- as a quality guide for processors. sumers through roadside stands, by house-to- There are four designated U.S. grade levels for house selling, through mail-order sales, or from extracted honey: U.S. grade A or U.S. Fancy, the producer's home. Some of these sales are made U.S. grade B or U.S. Choice, U.S. grade C or by the small or part-time producers, who have no U.S. Standard, and U.S. grade D or Substandard. real means of determining a true market price for Quality factors considered in ascertaining the their product. Often such sales are made on hear- grade of extracted honey are flavor, absence of say rather than on factual information. defects, clarity, and moisture. Honey color is C&MS has for many years published Honey classed by means of the Department's permanent Market News to make current marketing infor- glass color standards or by the Pfund honey color mation available to producers and others interested. grader. This unbiased monthly report is national in scope. U.S. standards also have been developed for It contains factual information on supply, demand, the grades and types of comb honey. Grades for market prices, beeswax, colony and honey plant comb-section honey are U.S. Fancy, U.S. No. 1, U.S. No. 1 Mixed Color, U.S. No. 2, and Un- conditions, and crop production material on a classified. Grades for shallow frame comb, wrapped State and national basis. cut-comb, and chunk or bulk comb honey packed Various means are used to obtain the infor- in tin or glass are U.S. Fancy, U.S. No. 1, and mation included in the monthly report. Data on Unclassified. Quality factors used in determining producer and packer sales of honey in the various grades are appearance of cappings, presence of size containers are obtained directly from the pollen grains, uniformity of honey, attachment individuals or firms. Each month a questionnaire of comb to section, absence of granulation, listing specific information needed is mailed from presence of honey dew, and weight. Washington, D.C., to approximately 300 members C&MS offers inspection services to the honey of the industry. Information voluntarily furnished industry on a fee-for-service basis. Such services include certification of grades, which helps the by these contacts is summarized in the market industry assure consumers of a clean and whole- report each month. Additional market information some product. Four general types of inspection is obtained from wholesale dealers in many of the service are available: Lot Inspection—examination large cities throughout the United States by of a number of containers sampled from specific trained C&MS market news reporters. lots; Continuous Inspection—one or more in- Honey import and export data are included in spectors assigned to a processing plant to observe the monthly report to help provide a complete and check operations continuously; Plant In- picture of the supply situation. Likewise, foreign spection-Pack Certification—similar to continuous honey crop reports, furnished by the Department's inspection, except that the inspector is not re- Foreign Agricultural Service, are included to give quired to be present at all times; Unofficially some insight on the world honey market. Submitted Samples—inspection of samples sub- 134 BEEKEEPING IN THE UNITED STATES 135 mitted by applicant to determine quality, con- food editors, and to persons doing educational dition, and grade. work in home economics and nutrition. C&MS program specialists select the foods each month for designation as '^plentiful,'' basing their Plentiful Foods Program selections on the Department's crop reporting The C&MS Plentiful Foods Program is a part and outlook services. of the broad policy of the U.S. Department of Agriculture to provide food distributors and con- Surplus Removal sumers with information on food supplies and prices. Its purpose is to expand the market for When supplies outrun demand and farm prices foods that are in peak seasonal supply or are fall drastically, C&MS can sometimes help out otherwise plentiful. with surplus removal programs. These programs It is essentially a program to enlist concerted serve to widen the market for farm products by action by the Nation's grocers and food-service encouraging domestic consumption, exports, or establishments to focus consumer attention on development of new outlets and uses. They are '^plentifuls.'^ This is done through stimulating carried out under authority of section 32 of creative merchandising, with simultaneous pub- PubHc Law 320. ''Section 32 programs'' help to licity in the press and on radio and television. stabilize prices for farmers and make it possible The trade elements are alerted to current plentiful to find useful outlets for many surplus farm foods through a monthly periodical, the Plentiful commodities that might otherwise be wasted. Foods List, and through other special bulletins Products purchased are used to improve the and personal contact by field personnel. The diets of school children and needy persons. material is aimed at the grocery trade and all Inquiries and requests for information con- important feeding operations, including restau- cerning any of these marketing aids or for inspec- rants, industrial and institutional feeders, and tion service and copies of the standards for grades schools participating in the National School of honey may be addressed to Fruit and Vegetable Lunch Program. Division, Consumer and Marketing Service, U.S. Information about plentiful foods also is Department of Agriculture, Washington, D.C. supplied regularly to mass media, especially to 20250, or to field offices of the agency. HONEY PRICE SUPPORT PROGRAM

By HARRY A. SULLIVAN, agricultural economist, Policy and Program Appraisal Division, Agricultural Stabilization and Conservation Service

Price support programs for agricultural com- (6) Ability to dispose of stocks acquired through modities were undertaken as a result of the de- price support operations. pression of the 1930's. For some years, price (7) The need for offsetting temporary losses of support operations were restricted to what were export rñarkets. called the ^^basic^' commodities (corn, cotton, (8) The ability and willingness of producers to peanuts, rice, tobacco, and wheat), but gradually keep supply in line with demand. other commodities were added. Parity prices are a measure of the price levels needed to give agricultural commodities a pur- Factors Leading to Honey Price chasing power with respect to articles that farmers buy, equivalent to the purchasing power of those Support Program agricultural commodities in a base period. The technique used to determine parity prices for Sugar rationing during World War II and the agricultural commodities has been outlined by requests by the Government to increase the Congress. The parity price calculations and production of honey led to a large increase in determinations are made by the Statistical colony numbers and a proportionate increase in Reporting Service of the U.S. Department of honey production. With the end of sugar rationing, Agriculture. prices for honey dropped close to prewar levels. Due to the depressed economic situation facing them, representatives of the beekeeping industry Operating Features of Price Support requested assistance from Congress. In taking note of the industry's request, the House Com- Program mittee on Agriculture had this to say: The price of honey is presently supported Since the close of the war, the price of honey has through a farm-storage loan or purchase program dropped to the point where beekeepers are finding or both. Loans at the applicable price support it impossible to obtain their costs of production. It appears obvious to the committee that, if these rate on farm-stored honey are made available vitally important insects are to be maintained to beekeepers during the crop year on any or in sufficient numbers to pollinate our crops, the all of the honey produced during that year. beekeeping industry must have immediate assist- By providing immediate cash for his crop, this ance. Until the time comes when beekeepers can receive an adequate return from pollination allows the beekeeper to hold his honey and to services, the committee believes that a price market it later during the crop year when he support program for honey, as provided in this thinks it is most advantageous to him. However, bill, is the only answer to this problem. if the market prices fail to rise above the support price, he may cancel his loan by delivering honey Honey Price Support Legislation of value equal to the loan value at the end of the year unless arrangements for earlier delivery The Agricultural Act of 1949 required that have been made. honey, along with several other commodities If the beekeeper has not made use of the loan under the heading ''Designated Nonbasic Agri- feature of the program for a part or all of his cultural Commodities,'' be supported at a level production, he can then use the purchase option. between 60 and 90 percent of parity. In deter- This means that the Commodity Credit Corpora- mining the actual level of support within the tion (CCC) stands ready to buy at the applicable prescribed limits, the Secretary of Agriculture is support price any of his production he wishes directed to consider the following factors: to sell and which is not already obligated to CCC (1) Supply in relation to demand. as loan collateral. (2) Price levels at which other commodities are The beekeeper, along with producers of other being supported. price-supported commodities, obtains loans or (3) Availability of funds. payment from a CCC purchase of his honey (4) Perishability of honey. at his Agricultural Stabilization and Conserva- (5) Importance of honey to agriculture and the tion Service county office. The U.S. Department national economy. of Agriculture carries out price support operations 136 BEEKEEPING IN THE UNITED STATES 137 through the Agricultural Stabilization and Con- ment, under which they agreed to pay beekeepers 9 servation Service (ASCS), which has field offices cents per pound dehvered to their packing plants in each of the more than 3,000 counties in the for all the honey acquired from them that met the country. The county office also issues delivery requirements of the program. These requirements instructions as to time and location for honey were especially concerned with the cleanliness of deliveries being made to CCC. the honey, its moisture content, and the flavor. The Department, in turn, agreed to accept from the contracting packer all the honey the packer Other Provisions of Loan and offered and to pay the support price, plus estab- Purchase Program Hshed charges for handhng, storage, and any processing requested by the Department. Quality and Quantity Determination In the 1951 season a similar program was For loan purposes, the beekeeper^s statement is operated, except a price support differential was accepted as to the quaUty of the honey offered introduced related to the degree of acceptabihty as collateral. He then receives 90 percent of the of honey for table use. The differential was 1.1 value (price support rate times quantity) of cents per pound between honeys of ''general the loan honey. When honey is actually acquired national acceptabihty'' and 'limited acceptabil- by CCC through purchase or loan repayment, ity'' for table use, reflecting to a degree the dif- quantity is determined by the actual weight ference in market value for variations in this of honey dehvered. QuaUty determination for regard. this purpose is made by the Processed Products The type of price support program in operation Standardization and Inspection Branch, Fruit during these first 2 years was not giving universal and Vegetable Division, Consumer and Market- satisfaction. The 1952 season saw the producer ing Service, in accordance with U.S. standards (i.e., beekeeper) loan and purchase agreement type for grades of extracted honey based on samples of program developed, which is now in use for drawn by ASCS representatives supervising de- honey and for most of the other agricultural Hvery. CCC bears the cost of this quahty and commodities. color determination. Summary of Price Support Activity Color and Area Differential Structure As indicated in table 1, activity under the price Honey is supported on the basis of color and support program has been relatively modest, class with an East-West area differential also with smaU quantities put under loan and even appHed. The color and class, along with the —Honey: Price support activity, 1950-66 area differential for the 1966 crop, are illustrated TABLE 1 as follows: West East Quantity placed Class and color (cents per (cents per Support under loan pound) pound) Price as per- Quan- Table honey : Year support cent of tity White and light 12. 0 12. 9 rate parity As per- acquired Extra light amber 11. 0 11. 9 Amount cent of by CCC Light amber 10. 0 10. 9 produc- Other table honey 8. 0 8. 9 tion Nontable honey 8. 0 8. 9

The national average support rate on a 60-pound Million Million and larger container basis is 11.4 cents per pound Cents Percent poimds Percent pounds for the 1966 crop. 1950 9. 0 60 0) 7.4 1951 10. 0 60 0) 17.8 Fees and Charges 1952 11. 4 70 9. 3 3.4 7. 0 1953 10. 5 70 3. 1 1.4 . 5 A producer pays a nonrefundable loan fee of $4 1954 10. 2 70 1. 5 . 7 0 for each loan disbursed. He pays a dehvery charge 1955 9. 9 70 1.9 . 8 0 0 of 1 cent per hundredweight on the quantity of 1956 9. 7 70 1. 6 . 7 1957 9. 7 70 2.9 1.2 . 1 honey delivered to CCC. The producer also pays 1958 9. 6 70 5.6 2. 1 2. 0 all charges relative to insurance premiums, storage, 1959 8. 3 60 1. 3 . 5 0 and handling. 1960 8.6 60 1. 1 . 4 0 1961 11.2 75 4.2 1.5 1. 1 1962 11. 2 71 3. 4 1. 2 0 Early Support Program 1963 11.2 67 3.2 1. 1 0 2. 2 The Department first decided that niandatory 1964 11.2 65 9.6 3.4 1965 11. 2 63 17. 3 6.2 3. 3 honey price support could be most widely and 1966 11.4 61 (') (2) e) effectively assured by working through existing marketing machinery. Under the 1950 program, 1 Direct Packer Purchase Program. packers of honey signed contracts with the Depart- 2 Data not available. 138 AGRICULTURE HANDBOOK 3 3 5, U.S. DEPT. OF AGRICULTURE smaller quantities acquired by CCC. The honey seem to indicate that the honey industry's greatest acquired by CCC has been disposed of primarily need for this type of program had been in these through the National School Lunch Program. two periods. However, there seems to be a general The greatest activity in terms of CCC acquisition consensus that the price support program has or loan placement took place in the initial 3 years provided the honey market with a significant after implementation of the program and in the degree of stabiHty during the period it has been in 1964 and 1965 crop years. This pattern would operation. STATISTICS ON BEES AND HONEY

By E. B. HANNAWALD, Chief, Livestock, Dairy and Poultry Branch, Agricultural Estimates Division, Statistical Reporting Service

Statistics concerning bees and honey were honey, and extracted honey; producers' stocks of first issued by the U.S. Department of Agriculture honey as of December 15; and beeswax production in 1940. Its Statistical Reporting Service, through and price. Estimates are made for each State; U.S. its Crop Reporting Board, issues three reports estimates are the sum of the State estimates each year. The first report, in July, carries an (tables 1 and 2). estimate of the number of colonies on hand and the condition of nectar plants and colonies. The second, in October, includes preliminary estimates Number of Colonies of the number of colonies, yield per colony, and The first estimates in 1940 were based on production of honey. This report also shows studies of the Census of Agriculture enumerations, producers^ stocks of honey for sale as of September State inspection records, estimates by State 15. The last report, in January, estimates the apiary inspectors and entomologists, and reports number of colonies, yield per colony, and total from commercial honey buyers and packers. production for the previous year. The report also Since then the yearly change in the number of carries wholesale and retail prices received by colonies has been based primarily on surveys and producers for all honey, comb honey, chunk State inspection records. No questions regarding

TABLE 1.—Honey: Number oj colonies, production, value, and stocks. United States, 1940-66 ^

Number of Yield Honey Price per Stocks on Year colonies per colony production pound Total value hand Dec. 15 for sale

Thousand Thousand Thousands Pounds pounds Cents Thousands pounds 1940 4,350 47.3 205, 767 6. 1 $12, 584 1941 4,477 49.6 221, 959 7.2 15, 973 1942 4,893 36.3 177, 672 13.8 24, 481 40, 512 1943 4,887 38.9 189, 867 16.8 31, 740 31, 361 1944 5,217 36. 2 188, 917 17.7 33, 433 26, 237 1945 5,460 42. 7 233, 070 18.6 43, 339 27, 133 1946 5,787 36.9 213, 814 24.4 52, 137 10, 787 1947 5,916 38.6 228, 582 24. 9 56, 878 62, 408 1948 5,721 36.0 206, 185 17.9 36, 852 70, 774 1949 5,578 40. 6 226, 334 15.0 34, 064 82, 893 1950 5,601 41. 5 232, 431 15.3 35, 615 83, 119 1951 5,546 46.4 257, 522 16.0 41, 234 71, 284 1952 5,493 49.5 272, Oil 16. 2 44, 063 77, 047 1953 5,520 40.5 223, 770 16. 5 36, 997 53, 210 1954 5,451 39.7 216, 419 17.0 36, 737 40, 965 1955 5,252 48.6 255, 222 14. 8 45, 424 58, 618 1956 5, 195 41.2 214, 035 19.0 40, 621 49, 489 1957 5, 199 46. 4 241, 218 18.7 45, 049 64, 024 1958 5, 152 50. 6 260, 522 17.4 45, 216 71, 084 1959 5, 109 46. 3 236, 578 17.0 40, 135 59, 969 1960 5,005 48.5 242, 802 17. 9 43, 500 52, 206 1961 4,992 51. 3 255, 868 18. 0 46, 072 68, 119 1962 4,900 50.9 249, 608 17.4 43, 494 55, 933 1963 4,849 55.0 266, 778 18.0 48, 096 55, 082 1964 4,840 51. 9 251, 188 18.6 46, 638 65, 776 1965 4,783 51. 1 244, 549 17.8 43, 475 58, 327 1966 4,770 51.8 246, 972 17.4 42, 927 56, 854

1 Excludes Hawaii 1940-54 for all items and 1956 and 1957 for Dec. 15 stocks. 2 Data not available. 139 140 AGRICULTURE HANDBOOK 33 5, U.S. DEPT. OF AGRICULTURE

TABLE 2.—Beeswax: Production and value, United variable on charts, whereas the dependent variable States, 1942-66' is the Crop Reporting Board's percentage change in number of colonies from the preceding year. Year Produc- Price per Total (For an example, see figure 1.) From these indica- tion pound value tions a percentage change from the previous year is adopted, which is multiplied by the estimated Thou- number of colonies for the previous year. This sand Thou- product is the estimated number of colonies for pounds Cents sands the current year. In some States, data on numbers 1942. 3,344 40. 3 $1, 343 of inspected colonies are available. Such records 1943. 3,743 41.4 551 1944. 3,921 41.5 627 are considered in making the estimates. 1945. 4,543 41.3 888 The estimate of number of colonies is again 1946. 4,381 44.4 947 reviewed for the October and December surveys 1947 4,500 43. 8 969 and revised in December if necessary. Most 1948. 4,041 43.2 746 1949. 4,139 37.6 554 consideration is given to the July survey, since 1950. 4,261 42.8 824 the estimate relates to the number of colonies on 1951. 4,692 50.4 368 hand at the beginning of the honey flow, and 1952. 4,812 43. 1 075 this date is the closest to the beginning of the 1953. 4,081 41. 0 673 1954. 4,011 44. 1 764 flow. 1955. 4,620 51.2 367 The estimate of colony numbers can also be 1956. 4,096 54. 6 237 revised the following year in July and December. 1957. 4,452 57.0 537 When revisions are made, they are based on 1958. 4,695 46.0 159 1959. 4,215 44.4 872 State inspection records and other data not 1960. 4,372 44. 0 922 earlier available. 1961 4,720 44. 1 080 1962 4,805 44. 1 117 1963 4,828 44.2 133 Honey Production 1964 4,672 44. 3 070 1965 4,749 44. 9 132 The first or preliminary estimate of the yield 1966 4,728 46.6 202 of honey per colony is made in October. This estimate is based on the amount of honey that ^ Excludes Hawaii 1942-54. producers have already taken and expect to take from the hives. the number of colonies of bees have been asked A final estimate is made in January, when on the census enumeration since 1950. producers are asked the amount of honey that Questionnaires are sent to producers for each was taken from the hives during the past year. survey. Each questionnaire asks the number of The total pounds of honey, as reported by pro- colonies on hand at the beginning of the honey ducers returning questionnaires, are divided by flow. The questionnaires are tabulated by size the sample number of colonies to obtain an aver- groups based on the number of colonies. Normally age yield per colony. The yields for each size the size groups used are 1-9 colonies, 10-39, and group are then weighted together to compute 40 or more. In States with large commercial a State average. The weighting is particularly producers, a further breakdown of the large important because yield per colony increases group is made. markedly as the number of colonies increases. The July inquiry asks for the number of colonies As colonies increase, management tends to im- on hand a year earlier as well as the current prove; large producers move colonies from one number. These two questions provide an indica- location to another to obtain higher yields. tion of the annual change in the number of The reported yields are plotted on a chart colonies. This indication is referred to as the C/H with Board final yields. The chart is then used (current/historic) percentage. The current number to determine the yield per colony. This derived reported on the July returns is also matched with yield per colony is multiplied by the Board the number reported on the July inquiry a year estimate of number of colonies to obtain the earlier for the same producers. This provides an estimate of total honey production. indication of change from the previous year, called C/C (current/current). Stocks of Honey Indications from each size group on these two measurements are combined by weighting each Stocks of producers^ honey for sale are esti- size group indication by percentages that reflect mated and published in the October and Jan- the amount of the colonies in the universe in each uary reports. A percentage indication of honey size group. Each of these weighted indications stocks on hand is derived from the sample by for a series of years is plotted as the independent dividing the reported stocks by the reported BEEKEEPING IN THE UNITED STATES 141 110

UJ LU / CÛ U. O 105 Xoi958

y^1964 100

3 196H /•1960 95 O US: CD LU 1959#/^ • 1965

Z i/^ r- < rO 1 LINE O 90

O

85 90 95 100 105 110

REPORTED PERCENT OF PREVIOUS YEAR JULY HONEY INQUIRY-IDENTICALS (C/C)

FIGURE 1.—Colonies of bees, Iowa: Official estimates of number of colonies of bees as percentages of previous year's estimates in relation to percentage relation between number reported on same farm in current year to number r reported in previous year on successive July honey surveys.

total honey produced. After the percentage of indication of beeswax output is derived from the honey on hand for sale is determined for each sample by dividing the reported pounds of bees- size group, a weighted percentage is obtained. wax by the reported total honey production. This weighted percentage is then expanded The percentage is then expanded into an estimate into a stocks estimate by multiplying it by the of beeswax production by multiplying the total total honey production for the year. honey production for the year by this percentage. Beeswax Production Prices An estimate is made in January of the amount Detailed estimates of prices received by pro- of beeswax produced during the previous year. ducers for honey were first published by the The estimate is based on the pounds of beeswax Department in 1940. The questionnaire has since in relation to honey production. A percentage been changed several times to furnish the more 2i61-907 0-^67^—10 142 AGRICULTURE HANDBOOK 335, U.S. DEFT. OE AGRICULTURE comprehensive price data needed as price support for each size container sold, both wholesale legislation for honey became law. In 1948, the and retail. Estimated quantities of honey sold, questionnaire was changed to provide a break- by categories for each State, are used in this down of both wholesale and retail sales of ex- process to obtain a price for all honey. The tracted honey, by size of containers most com- quantities used as weights are derived from monly sold. Sizes included were 60-pound, 5- pounds of honey sold in each category as reported pound, 1-pound, and ''other'' containers. In 1959, on the questionnaire and from other available questions on prices and quantity of wholesale sales data. The weights also are used to compute sales in 55-gallon drums were added. regional and U.S. average prices for wholesale The price estimate of all honey sold, pubUshed and retail sales of extracted, comb, chunk, or in January, is obtained by combining prices bulk honey and for all honey. WORLD PRODUCTION AND TRADE IN HONEY

By LESLIE C. HURT, Acting Chief, Commodity and Products Analysis Branch, Sugar and Tropical Products Division, Foreign Agricultural Service

Honey is produced in all States of the United half of their crops. Canada exports a much States and in all continents of the world. World smaller amount both in total volume and as a production of honey, excluding mainland China, percentage of production. However, the Canadian is currently about 900 million pounds. All con- market has had substantial growth in the last tinents contribute to this total. North America few years. The bulk of the sales is made to the is by far the largest producer, whereas only small United Kingdom, and the trend has been to amounts are produced in Africa. The United smaller containers, predominantly the 1-pound States accounts for about one-third of total variety. world production. The U.S.S.R. ranks second in Ranking second in imports in 1965, despite production, with about 200 million pounds. Other its leading position in world production, was the sizable producers are Argentina, Mexico, Canada, United States. Although a large share of U.S. Australia, France, Turkey, Spain, and West imports of honey is for baking and manufacturing Germany. Smaller producers include Brazil, Chile, industries, there are also imports for specialty New Zealand, Italy, and Japan. The production purposes. Exports from the United States since trend is upward throughout the world. World War II have substantially exceeded im- World trade in honey also is generally upward ports in most years. During 1946-65, imports and amounts to about 150 million pounds annu- averaged 9.2 million pounds, whereas exports ally. The largest exporters in recent years have averaged 14.4 million pounds. The level of exports been Mexico, Argentina, Australia, and in some was highest during the 1950's; however, 1963 years the United States. In recent years, Europe exports of 25.1 million pounds reflected the all- has often accounted for about 95 percent of all time record production. Practically all exports world imports. West Germany imports about 65 have been to European countries, with West percent of the world total. Major suppliers in Germany the leading buyer and France usually 1965 continued to be Argentina and Mexico, second. followed by mainland China in terms of quantity World per capita consumption of honey is at and the United States in terms of value. Several a rather low level. In the United States it amounts European countries import less than 10 million to about 1.5 pounds and has not varied much pounds of honey annually. in recent years. Argentina in 1965 furnished a Of the largest exporters, Mexico in recent years good example of what can be done by promoting has been shipping the biggest ])roportion of its consumption. Local consumption rose to 35 mil- production. Much of this is shipped to West lion pounds, triple that of the previous year. Germany, but a substantial amount has also been The success of this promotion is referred to as "^ shipped to the United States. Argentina and 'The Argentine Miracle,'' and illustrates the Australia for several years have exported over dormant potential for honey consumption. 143 GLOSSARY

Abdomen: Segmented posterior part of bee con- Bee moth: (See Wax moth.) taining heart, honey stomachy intestines, Bee paralysis : Condition of bee, sometimes caused reproductive organs, and sting. by virus, that prevents it from flying or per- Acarapis ivoodi (Rennie): Scientific name of forming other functions normally. acariñe mite, which infests tracheae of bees. Bee plants: Vegetation visited by bees for nectar Acariñe disease: Disease caused by acariñe mite. or pollen. Alighting board: Extended entrance of beehive Bee space: Amount of space acceptable to bees, on which incoming bees land. neither too narrow nor too wide; discovered American foulbrood (AFB): Contagious disease by great American beekeeper Langstroth. of bee larvae caused by Bacillus larvae White. Beeswax : Wax secreted from glands on underside Antennae: Slender jointed feelers, which bear of bee abdomen; molded to form honeycomb certain sense organs, on head of insects. and can be melted into solid block. Anther : Part of plant that develops and contains Bee tree: Hollow tree in which bees live. pollen. Bee veil: Screen or net worn over head and face Apiarist: Beekeeper. for protection from bee stings. Apiary: Group of bee colonies. Bee venom: Poison injected by bee sting. Apiculture: Science of beekeeping. Bee yard: (See Apiary.) Apis: Genus to which honey bees belong. Bottom board: Floor of beehive. Apis dor sata Fabricius: Scientific name for giant Brace comb: Section of comb built between and bee of India; largest of all honey bees. attached to other combs. Artificial cell cup: (See Cell cup.) Braula coeca Nitzsch: (See Bee louse.) Artificial insemination: Instrumental impregna- Breathing pores : (S'^e Spiracles.) • tion of confined queen bee with sperm. Brood: Immature or developing stages of bees; Bacillus larvae White: Bacterial organism caus- includes eggs, larvae (unsealed brood), and ing American foulbrood. pupae (sealed brood). Balling a queen: Clustering around unacceptable Brood chamber: Section of hive in which brood is queen by worker bees to form a tight ball; reared and food may be stored. usually queen dies or is killed in this way. Brood comb : Wax comb from brood chamber of Bee bread: Stored pollen in comb. hive containing brood. Bee dance: Movement of bee on comb as means Brood nest: Area of hive where bees are densely of communication; usually same movement clustered and brood is reared. is repeated over and over. Brood rearing : Raising bees. Bee escape: Device to let bees pass in only one Bumble bee: Large hairy bee in genus Bombus. direction; usually inserted between combs of Burr comb: Comb built out from wood frame or honey and brood nest when removal of bees comb, but usually unattached on one end. from honey is desired. Cap: Covering of cell. Bee gum: Usually hollow log hive; occasionally Capped brood: (&e Sealed brood.) refers to any beehive. Capped honey: Honey stored in sealed cells. Beehive: Domicile prepared for colony of honey bees. Carniolan bee: Gentle grayish-black bee originally from Carniolan Mountains in or near Austria. Bee louse: Relatively harmless insect that gets on honey bees, but larvae can damage honey- Caucasian bee: Gentle black bee originally from comb; scientific name is Braula coeca Nitzsch. Caucasus area of Russia; noted for its heavy propolizing characteristic. Bee metamorphosis: Stages in development of honey bee from egg to adult. Cell: Single unit of space in comb in which honey is stored or bee can be raised; worker cells Period (days) are about 25 cells per square inch of comb, Stage - Queen Worker Drone drone cells are about 18 per square inch. Egg 3 3 Cell cup: Queen cell base and part of sides; Larva 8 10 13 Pupa 4 8 artificial cell cups are about as wide as deep. Chilled brood: Immature stages in life of bee Emerge as adult 15 21 24 that have been exposed to cold too long. 144 BEEKEEPING IN THE UNITED STATES 145 Chunk comb honey : Type of honey pack in which Dwindling: Rapid or unusual depletion of hive piece of honeycomb is placed in container of population. liquid honey or wrapped ''dry'' in plastic Dysentery: Unusual watery discharge of bee feces, container. often associated with nosema disease. Orcadian rhythm: Biological rhythm with period Emerging brood: Young bees first coming out of length of about 1 day. their cells. Clarified honey: Honey that has been heated, Enzyme: Material produced by both man and then filtered to remove all wax or other animals that acts on another material to particles. change it without changing itself. Cleansing flight: Flight bees take after days of Escape board: Board with one or more bee es- confinement, during which they void their capes on it to permit bees to pass one way. feces. European foulbrood: Infectious disease of larval Clipped queen: Queen whose wing (or wings) has brood, caused by Streptococcus pluton (White). been clipped for identification purposes. Excluder: (See Queen excluder.) Cluster: Collection of bees in colony gathered Extracted honey: Honey extracted from comb. into limited area. Extractor: Machine that rotates honeycombs at Colony: Social community of several thousand sufficient speed to remove honey from them. worker bees, usually containing queen with Feces: Bee droppings or excreta. or without drones. Fecundate: To inseminate or implant sperm into Comb: {See Honeycomb.) female. Comb foundation: Thin sheet of beeswax im- Fertilize: To make fertile, as by implanting pressed by mill to form bases of cells; some sperm into ova. foundation is.also made of plastic and metal. Field bees: Bees 2^ to 3 weeks old that collect Comb honey: Edible comb containing honey; food for hive. usually all cells are filled with honey and Flash heater: Device for heating and cooling sealed by bees with beeswax. honey within few minutes. Commercial beekeeper: One who operates suffi- Food chamber: Hive body containing honey- ciently large number of colonies so that his filled combs on which bees are expected to entire time is devoted to beekeeping. live. Cross-pollination: Transfer of pollen from anther Foulbrood: Common name of two brood diseases; of one plant to stigma of different plant or usually applied to American foulbrood. clone of same species. Foundation: (/S^e Comb foundation.) Crystallization: {See Granulated honey.) Frame: Wood case for holding honeycomb. Cut-comb honey: Comb honey cut into appropri- Fructose : {See Lévulose.) ate sizes and packed in plastic. Fumagillin: Antibiotic given bees to control no- Demaree: Method of swarm control, by which sema disease. queen is separated from most of brood; de- Gallería mellonella (L.): Scientific name of vised by man of that name. greater wax moth. Dequeen: Remove queen from colony. Giant bee: {See Apis dorsata Fabricius.) Dextrin: Soluble carbohydrate of poor nutritive Glucose: {See Dextrose.) value to bee. Grafting: Transfer of larvae from worker cells into Dextrose: Also known as glucose; one of prin- queen cells. cipal sugars of honey. Granulated honey: Crystallized or candied honey. Diastase: Enzyme that aids in converting starch Gynandromorph: Bee having both male and fe- ß to sugar. male characters. Division board: Flat board used to separate two Half-depth super: Super only half as deep as colonies or colony into two parts. standard 10-frame Langstroth super. Division board feeder: Feeder to hold sirup; Heterosis: Greater vigor displayed by crossbred usually size of frame in hive. animals. Drawn comb: Foundation covered with completed Hive: Man-constructed home for bees. cells. : Metal tool for prying supers or frames Drifting bees: Tendency of bees to shift from apart. their own colony to adjacent ones. Hobbyist beekeeper: One who keeps bees for Drone: Male bee. pleasure or occasional income. Drone brood: Area of brood in hive consisting of Hoffman frame: Self-spacing wood frame of type drone larvae or pupae. customarily used in Langstroth hives. Drone comb: Comb having cells measuring about Honey: Sweet viscous fluid elaborated by bees four to the inch and in which drones are from nectar obtained from plant nectaries, reared. chiefly floral. Drone egg: Unimpregnated ^g^. Honey bee: Genus Apis^ family Apidae, order Drone layer: Queen that lays only infertile eggs. Hymenoptera. 146 AGRICULTURE HANDBOOK 335, U.S. DEPT. OF AGRICULTURE Honeycomb: Comb built by honey bees with Nucleus (nuclei) : Miniature hives. hexagonal back-to-back cells on median mid- Nurse bees: Young worker bees that feed larvae. rib. Observation hive: Hive with glass sides so bees Honeydew: Sweet secretion from aphids and can be observed. scale insects. Ocellus (ocelli) : Simple eye(s) of bees. Honey extractor: {See Extractor.) Package bees: Screen wire and wood container Honey flow: Period when bees are collecting nec- with 2 or 3 pounds of live bees. tar from plants in plentiful amounts. Parafoulbrood: Relatively rare bee disease similar Honey house: Building in which honey is ex- to European foulbrood; caused by bacterium tracted and handled. Bacillus para-alvei Burnside. Honey pump: Pump for transferring liquid honey Paralysis: {See Bee paralysis.) from one container to another. Parthenogenesis: Production of offspring from Honey stomach : Area inside bee abdomen between virgin female. esophagus and true stomach. Pheromone: Chemicals secreted by animals to Honey sump : Temporary honey-holding area with convey information to or affect behavior of baffles; tends to hold back sizable pieces of other animals of same species. wax and comb. Pistil: Part of flower extending from ovary to Hormone: Substance produced in small quantity stigma. in one part of body (usually in gland of Play flight: Short orientation flight taken by internal secretion) and transported to other young bees, usually by large numbers at one parts, where it exerts its action. time and during warm part of day. Hymenoptera: Order to which all bees belong, as Pollen: Dustlike material produced in flower and well as ants, wasps, and certain parasites. necessary on stigma of female flower for seed Introducing cage: Small wooden and wire cage production; also collected in pellets on hindlegs used to ship queens and also to release them of bees. quietly into cluster. Pollen basket: Area on hindleg of bee adapted for Invertase: Enzyme produced by bee that speeds carrying pellet of pollen. inversion of sucrose to glucose and fructose. Pollen cake: Cake of sugar, water, and pollen or Italian bees: Bees originally from Italy; most pollen substitute for bee feed. popular race in United States. Pollen substitute: Mixture of water, sugar, and Jumbo hive : Hive 2)^ inches deeper than standard other material, such as soy flour, brewer's Langstroth hive. yeast, and egg yolk, used for bee feed. Langstroth frame: 9)^- by 17%-inch frame. Pollen supplement: Mixture, usually of six parts Langstroth hive: Hive with movable frames; each (by weight) pollen, 18 parts soy flour, 16 frame usually 9)á by 17% inches. parts water, and 32 parts sugar. Larva: Stage in life of bee between egg and pupa; Pollen trap: Device installed over colony en- ^'grub'^ stage. trance that scrapes pollen from legs of Laying worker: Worker bee that lays eggs after entering bees. colony has been queenless for many days. PolMnation : Transfer of pollen from male to Legume: One of Leguminosae, or plants such as female element of flower. clover, alfalfa, peas, or beans. Pollinator: Agent that transfers pollen. Lévulose: Fructose or fruit sugar; one of sugars, PoUinizer: Plant that furnishes pollen for another. with glucose, into which sucrose is changed. Proboscis: Tongue of bee. Mandibles: Jaws of insects. Mating flight: Flight taken by virgin queen when Propolis: Resinous material of plants collected she mates with drone in air. and utilized by bees within hive to close Metamorphosis: Changes of insect from egg to small openings or cover objectionable objects adult. within hive. Migratory beekeeping: Movement of apiaries Pupa: Stage in life of developing bee after larva from one area to another to take advantage and before maturity. of honey flows from different crops. Queen: Sexually developed female bee. Mite: {See Acarapis woodi (Rennie).) Queen cell: Cell in which queen develops. Movable frame: Frame bees are not inclined to Queen excluder: Device that lets workers pass attach to hive because it allows proper bee through but restricts queen. space around it. Nectar: Sweet exúdate from nectaries of plants. Queenless: Without queen. Nectaries: Special cells on plants from which Queen rearing: Producing queens. nectar exudes. Queenright: With queen. Nosema disease: Disease of bees caused by Queen substance: Material produced from glands protozoan spore-forming parasite. Nosema in head of queen; has strong effect on colony apis Zander. behavior.

V BEEKEEPING IN THE UNITED STATES 147

Ripe honey: Honey from which bees have evapo- Sucrose: Cane sugar; main solid ingredient of rated sufficient moisture so that it contains nectar before inversion into other sugars. no more than 18.6 percent water. Super: Extra division of hive above brood nest Robbing: Bees of one hive taking honey from area. another. Supersedure: Replacement of one queen by Royal jelly: Food secreted by worker bees and another while first is still alive. placed in queen cells for larval food. Swarm: Natural division of colony of bees. Sacbrood: Minor disease of bees caused by Tarsus: Fifth segment of bee leg. filterable virus. Thorax: Middle part of bee. Sealed brood: Brood in pupal stage with cells Tracheae: Breathing tubes of insects. sealed. Tumuli: Nest mounds. Self-pollination: Transfer of pollen from male to Uncapping knife: Knife used to remove honey cell female element within same fiower. caps so honey can be extracted. Septicemia: Usually minor disease of adult bees Unite: Combine one colony with another. - caused by Pseudomonas apiseptica (Burnside). Unsealed brood: Brood in egg and larval stages / Shallow super: Super less than 9%6 inches deep. only. > Shipping cage: Screen and wood container used , V to ship bees. Virgin queen: Unmated queen. Skep: Beehive made of straw. Wax glands: Glands on underside of bee abdomen jf Smoker: Device used to blow smoke on bees to from which wax is secreted after bee has been reduce stinging. gorged with food. Solar wax extractor: Glass-covered box in which Wax moth: Lepidopterous insect whose larvae wax combs are melted by sun's rays and wax destroy wax combs. is recovered in cake form. Wild bees: Any insects that provision their nests Spermatheca: Small saclike area in queen in with pollen, but do not store surplus edible which sperms are stored. honey. Spermatozoan : Male reproductive cell. Winter cluster: Closely packed colony of bees in Spiracles: External openings of tracheae. winter. Stamen: Male part of flower on which pollen- Wired foundation: Foundation with strengthen- producing anthers are borne. ing wires embedded in it. Stigma: Receptive part of style where pollen Wired frames: Frames with wires holding sheets germinates. of foundation in place. Sting: Modified ovipositor of female Hymenop- Worker bee : Sexually undeveloped female bee. tera developed into organ of defense. Worker comb: Honeycomb with about 25 cells Streptococcus pluton (White) : Causative agent per square inch. of European foulbrood. Worker egg: Fertilized bee egg.

U.S. GOVERNMENT PRINTING OFFICE: 1967—O261-907

NAHONAL AGRICULTURAL LIBRARYil 022209584 1022209584

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