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1993  Wenner, A.M. and R.W. Thorp.  The  honey bees of Santa Cruz Island.  Bee Culture.  121 (5):272-275.
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Our familiar honey bee, not native to the United States, has been intentionally transported on a larger scale than might be true for any other animal or plant species (ably reviewed in 1989 by Walter Sheppard). During the colonial period, the dark European bee (”German” bee), Apis mellifera mellifera, was the primary bee of commerce (1500-1850). In fact, in Sheppard’s words, “Prior to the introduction of the Italian race…, A. m. mellifera was the sole race of honey bees present in the United States.”

The Virginia Colony was the first to import honey bees (in 1622); by 1654 honey bees had been established in New England. The frequent swarming habit of dark European bees, favored by beekeepers before moveable frame hives were available, resulted in their rapid spread throughout the eastern United States during the next 200 years.

Up to a century and a half ago, overland shipment of bees to California was considered impossible - travel through the Great Basin and deserts of the Southwest was too arduous. A botanist, Christopher A. Shelton, first breached that geographical barrier by bringing bees in by ship. An unknown beekeeper had transported 12 colonies down to Panama in early 1853 and sold them to Shelton, who at that time was introducing various plant species to California. Shelton managed to get the colonies to the Robert F. Stockton Ranch, just north of San Jose, California, but only one colony survived.

Other shipments followed, imported by John Harbison during the mid- 1850s. Harbison had abandoned gold mining to start the first nursery of fruit and ornamental trees in the Sacramento Valley but soon turned to beekeeping on a large scale. Known as the “Bee King of California,” he invented the comb honey section box still in use today and published The Beekeeper’s Directory. Soon honey bees were dispersed throughout California - before 1860 a thousand colonies were already present in San Jose.

In 1856 Southern California got its first bees from some of those original importations, and Ventura County - nearest point to the Northern Channel Islands off Santa Barbara - had its first commercial apiary in 1873. About that time an unknown beekeeper brought bees out to Santa Cruz Island but abandoned them well before 1880. Bees from that original introduction multiplied and spread over the entire island, with apparently no introductions since that time. The adjacent Northern Channel Islands had no such importations and have never had honeybees, but Santa Catalina Island in the southern group does have bees.

While dark European bees were being introduced into California, another development began in 1851, one that changed both beekeeping and the bee of commerce. Lorenzo Langstroth determined dimensions of the correct “bee space” and invented the first practical moveable frame hive. Beekeepers began replacing dark European bees, noted for their excessive use of propolis and rather poor hivekeeping behavior, with Italian bees. In Ohio, Langstroth obtained his earliest shipment of Italian bees from Germany in 1863, but the first successful direct shipment of Italian bees had been into New York three years earlier.

Completion of the Transcontinental Railroad in 1869 permitted rapid transportation of bees from the East to the West with the first Italian colonies on record reaching Los Angeles, California in 1875. As in the East, California beekeepers gradually replaced dark European bees.

There are two sides to this coin, however. Whereas we can laud diligent beekeepers for their transportation of colonies nearly everywhere in the world, the degree to which those bees have affected various native bee species remains a question. During our visits to Santa Cruz Island and Santa Rosa Island (six miles to the west and an Island on which European bees have never existed), we noticed remarkable differences in insect visitation on flowering plants.

Nearly all insects visiting the more prominent Santa Cruz Island plants were honey bees, but a wide spectrum of bees, wasps, flies, and moths visited the same types of flowers on Santa Rosa Island.

Several years ago all five of the Northern Channel Islands were designated a U.S. National Park. Santa Barbara Island, Anacapa Island, and San Miguel Island were already administered by the Park Service at that time. Subsequently, Santa Rosa Island was purchased and placed under their control. Most of Santa Cruz Island remains an inholding of the National Park under ownership by The Nature Conservancy, with the Park Service continuing negotiations to obtain the remainder of the eastern tip of that island.

An opportunity arose. Since Santa Cruz Island is the only one of the five Islands that has ever had honeybees, is essentially uninhabited, and has well defined limits, we reasoned that it should be possible to locate and remove all honey bee colonies. Such an effort would provide us with at least one National Park where native bees would prevail and where a pre-European ecosystem could be studied, once native insects on that island had again achieved somewhat of a balance. Furthermore, with the apparently inevitable influx of Africanized bees into our local area, we could have a Southern California habitat free of that hostile intruder.

Accordingly, five years ago Robbin Thorp of the University of California, Davis, campus started a study of plant visitation to determine diversity and abundance of native bees and potential competition for pollen and nectar with honey bees while Adrian Wenner of the UC, Santa Barbara, campus searched for feral honey bee colonies. After two years of study, elimination of colonies from only the eastern half of the island began. This two-stage removal process would then permit studies of flower visitation and pollination on the eastern vs western halves of the island - as well as permit similar studies between Santa Cruz Island and honey bee-free Santa Rosa Island. Fortunately, our area has a Mediterranean climate, active foraging occurs all year, and studies can be conducted year-round.

Work on the project has been proceeding much on schedule, with the eastern half of the island largely free of feral bee colonies. More than 160 colonies have been located (see table), of which about 130 have been removed. Laying queens have been recovered and provided to bee researchers and beekeepers so they can learn more about the characteristics of these bees after 110 years of isolation. Inspection of colony structure and behavior indicates that the island feral bees appear to be vely similar to the dark European strain (except for color) and remarkably uniform over the entire island.

Records are also being kept of both colony location and cavity type (see table below). About two-thirds of the colonies found have been in cliff faces, either in discrete cavities, within rocky crevices, or under rock shelves. Other common sites are eroded cavities under the boles of scrub oak trees. Rarely are colonies found in the classic bee tree cavity, even though many such trees exist all over the island.
Dozens of the Schmidt-Thoenes bait hives have been installed at various points around the island. Catch frequencies by those hives permit a comparison with how well cavities formerly occupied by colonies will attract swarms (see table below) - to date, formerly occupied cavities seem more attractive.

To determine whether native bees have been forced to small refuges by the more dominant honey bees, we measure diversity and abundance of all bees at selected flower species as honey bees are being removed. Since honey bee removal began, the numerous species of native bees have been increasing in numbers rapidly and now outnumber honey bees at blossoms on much of the eastern half of the island (see figure).

The isolation and minimal human habitation of the 96-square-mile island since honey bees were introduced 110 years ago also permits an unparalleled opportunity for studies of natural colony distribution, foraging patterns of colonies, and competition between colonies. The pressure to find feral colonies quickly has also led to the first major changes in those techniques in hundreds of years (see Sources at end of article). Colony locations can now often be found within a few hours after finding bees at blossoms or at water - the record (held by Dan Meade of UCSB) is 24 minutes.

Other bee researchers have become involved in the project to varying degrees. From the Tucson USDA bee research Laboratory, Justin Schmidt and Steve Thoenes have furnished bait hives and pheromone lures for our use, while Gerry Loper has studied drone aggregation sites. Steve Buchmann from that laboratory has started an analysis of pollen grains to determine how far bees might range while foraging. Howell Daly from the UC Berkeley campus is conducting a measurement of wing patterns to determine which strain the Santa Cruz Island bees might belong to, while Rob Page of the UC Davis campus is conducting allozyme and DNA analyses toward that same end.

Dr. Wenner recently retired as Professor of Natural History at the University of CA, Santa Barbara but continues his honey bee research on Santa Cruz Island. Dr. Thorp is Professor of Entomology and Apiculturist at the University of CA, Davis.

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The information for this article was obtained from several sources, most of which are listed here.

Caron, D.M. The Harbison California hive has a place in our beekeeping heritage. American Bee Journal 128:29-31. 1988.

Crane, E. The Archeology of Beekeeping. Cornell Univ. Press, Ithaca, NY. 1983.

Harbison, F.R. Flood Tides along the Allegheny. Massy Harbison Chapter, DAR (reprint), New Kensington, PA. 1941.

Mason, J.D. History of Santa Barbara and Ventura Counties, California. Howell North Books, Berkeley. 1883.

Root, A.I., E.R. Root, H.H. Root, and M.J. Deyell. The ABC and XYZ of Bee Culture. The A.I. Root Co., Medina, Ohio. 1947.

Ruttner, F., E. Milner, and J.E. Dews. The Dark European Honey Bee: Apis mellifera mellifera Linnaeus 1758. The British Isles Bee Breeders Association. 1989.

Schmidt, J.O. Swarms traps: An example of research and technology transfer. American Bee Journal 130: 333-334.

Schmidt, J.O. and S.C. Thoenes. The efficiency of swarm traps: What percent of swarms are captured and at what distance from the hive? American Bee Journal 130: 811-812. 1985.

Sheppard, W.S. A history of the introduction of honey bee races into the United States. American Bee Journal 129:617-619; 664-667. 1989.

Watkins, L.H. California’s first honey bees. Amer. Bee Journal 108:190-191. 1968.

Watkins, L.H. First honey bees in New England - 1638? American Bee Journal 108: 19. 1968.

Watkins, L.H. On the transportation of honey bees to California, 1853-1861. American Bee Journal 109: 468-470.

Wenner. AM., D.E. Meade, and L.J. Friesen. Recruitment, search behavior, and flight ranges of honey bees. American Zoologist 31: 768-782. 1991.

Wenner, A.M., J.E. Alcock, and D.E. Meade. Efficient hunting of feral colonies. Bee Science 2: 64-70. 1992.

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1961 Wenner, A.M. A method of training bees to visit a feeding station. Bee World. 42:8-11.
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ADRAIN M. WENNER*
Department of Biological Sciences,
University of California,
Santa Barbara, California, USA

INTRODUCTION

This paper describes a method for training bees to visit a feeding station, a subject which has been inadequately treated in the literature. Von Frish (1950, 1954), Ribbands (1953) and various Russian workers Have included descriptions of training bees. These methods, however, have often proved ineffective; Rothenbuhler (1959) and Murie (1960) have mentioned that prospective researchers in bee behaviour have been unable to proceed because they had no effective method for training bees. Difficulties were also experienced by the author (Wenner, 1961), even during periods of nectar shortage.

The following method was suggested in part by Boch (1959) and was refined and tested in 1960.

METHODS AND RESULTS

In southern Michigan the clover nectar flow normally lasts until the middle of July, at which time the nectar supply slackens (Martin, 1959). This period was chosen to take advantage of a probable nectar source. An unusually cool and moist spring and early summer promoted a greater supply of midsummer wild flowers than normal, however, which might have been expected to reduce the possibility of success in training bees during this period. Nevertheless, the method proved successful.

In testing this method, a seven-frame hive was located on a stand with its entrance approximately 1/2 m. from the ground. The box on which the feeder sat (at the same height as the hive entrance) was placed in front of the hive. On a cool (60º-70ºF.), rainy day in the middle of July the hive was opened and the top bars and bees were liberally sprinkled with a 2-molar solution of peppermint-flavoured sugar syrup (5 drops of peppermint flavoring to 1 litre of syrup). After the bees had had time to store this syrup, drops of syrup were continually offered at the hive entrance until 10-20 bees were regularly taking it. Then drops of syrup were placed on an 8 X 13 cm. (3 X 5 in.) card which was held at the hive entrance, forcing bees to walk 4-5 cm. before they were able to imbibe syrup. Drops of syrup were added to the card as fast as bees removed them. When 5-10 bees were on the card, the card was carefully transferred to the box, at the same height as the hive entrance, but leaving a gap of 3-4 cm. between it and the card. Bees would not normally fly across the gap, however, and this transferring of the card to the stand had to be repeated 20-30 times before any bees flew from the hive entrance to the stand. The preceding step, forcing the bees to fly, was the most difficult step encountered.

After the bees were regularly flying from the entrance of the hive to the stand, the stand was moved in the direction the station would eventually be from the hive. This movement was only 5-10 cm. each 5-10 min. until the foragers were flying about 2 m. At this distance a feeder, consisting of an inverted jar of syrup in a saucer (with a gap between jar and saucer, providing an automatic dispensing of the syrup), was used instead of syrup on a card. Bees were then allowed to become accustomed to imbibing at the feeder before the station was moved further from the hive.

The moving of the station away from the hive consisted of a set procedure which was repeated until the station was at the desired distance. During a period of decreased activity, an opaque can was placed over the feeder and any remaining bees. As the remaining bees became full, they were allowed to escape and return to the hive; newly arrived bees were not allowed access too the syrup. In 1/2 - 1 min., all bees which had been feeding had left for the hive. The can was then left in place about 3 min. until all bees had had time to return from the hive. (This time varied according to the distance of the feeding station from the hive; 2-3 min. was the minimum time needed.) As a result, all bees which had previously visited the station were now flying in its vicinity. The opaque can was then lifted from the feeder, allowing the bees to settle and imbibe the syrup. When all but 1-2 bees were settled, the entire feeding station was lifted and carefully carried in a straight line from the hive. Movement away from the hive was continued until the first bee was filled and flew back towards the hive. The station was then placed on the ground. The procedure given in this paragraph was repeated each time all bees had averaged 2-3 round trips to a new location, until the station was at the desired distance from the hive.

Initially the station was moved 210 m. from the hive and kept there 7-8 days for an experiment. At the end of that time the number of visitors dropped from 10-15 to 3-4 regular visitors, probably because of inadequate hive ventilation. After providing more ventilation to the hive, I decided to begin at the hive and re-train the bees to fly 420 m. to a station in the same direction. Meanwhile, however, all available foragers had reverted to visiting wild flowers, and bees did not imbibe sugar syrup at the hive entrance. The first attempt at training bees had been made on a cool rainy day. During this second attempt, the weather was generally warm (75º-85º F.) and clear. Apparently the first attempt had succeeded, at least in part, because of a temporary nectar shortage brought about by inclement weather.

The method which has been previously described succeeded, however, after pepperment-flavoured honey was substituted for flavoured sugar syrup. The bees could then be trained to fly to a station at 420 m., at which time the honey was replaced by a 2-molar solution of flavoured sugar syrup.

This success at training bees during a good nectar flow implies that it may be possible to train bees to visit feeding stations during most of the summer. The sugar concentration of the syrup can be adjusted to provide adequate competition to the natural nectar supply. The concentration should probably be kept as low as possible, however, in order to reduce its attractiveness to scout bees from other hives.

DISCUSSION

The basic difference between the method described above and one which consists of moving the station a short distance each time, appears to be in forcing bees to fly for some time before allowing then to alight. An experience the previous summer included attempts to move the station while bees were feeding on the syrup, but without having kept them away from it for a short time before permitting them to alight. These attempts all met with failure; bees so moved did not generally find the station in the new location.

The findings of von Frisch (1948), Shaposhnikova (1958) and Wenner (1961) indicate that foraging bees depend on their outgoing flight for the information they give to other bees about the distance of food from the hive. Thus, keeping bees in the air for a longer period of time than is necessary for flight to the food source may act in resetting the bee’s sensory mechanism. This, in turn, may allow a longer flight on the the next trip out from the hive. This does not mean, however, that every bee is certain to find the station in its new location. The success of the method apparently depends on the majority of bees being able to find the station after it has been moved. If these bees are allowed to make 2-3 round trips, they are likely to produce a signal in the hive which other recruits can follow (Shaposhnikova, 1958).

ACKNOWLEDGEMENTS

Dr. Francis C. Evans, director of the Edwin S. George Reserve, kindly permitted this study on the Reserve. The Edwin S. George Reserve Committee provided living quarters and financial assistance during the study. Mr. Harold E. Losey generously furnished bees, equipment, and time.

REFERENCES

BOCH, R. (1959) Personal communication

FRISCH, K. von (1948) Geloste und ungeloste Ratsel der Bienensprache Naturwissenschaften 35 : 12-23, 38-43

__________ (1950) Bees, their vision, chemical senses, and language Ithaca, N.Y. : Cornell University Press

__________ (1954) The dancing bees London : Methuen

MARTIN, E. C. (1959) Michigan nectar flows [Mimeographed summary of data for 1951-1957] E. Lansing : Michigan State University

MURIE, M. L. (1960) Personal communication

RIBBANDS, C. R. (1953) The behaviour and social life of honeybees London : Bee Research Association

ROTHENBUHLER, W. C. (1959) Personal communication

SHAPOSHNIKOVA, N. G. (1958) [The factors determining the formation of the recruitment signal in the honeybee, Apis mellifera carnica] Rev. Ent. U.R.S.S. 37(3) : 473-481

WENNER, A. M. (1961) Sound production during the waggle dance of the honey bee Anim. Behav. (in press)

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* This study was done at the University of Michigan’s Edwin S. George Reserve at Pinckney, Michigan.

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A nuc (nucleus) hive has all the features of a standard 10 frame hive except on a reduced scale. The nuc hive is used for making splits, swarm control, queen introduction, pollen/nectar monitoring, to name a few. This version is put out by the U.S.D.A.

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1992 Wenner, A.M., J.E. Alcock, and D.E. Meade. Efficient hunting of feral colonies. Bee Science 2:64-70.

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Adrian M. Wenner, Joe E. Alcock and Daniel E. Meade

Department of Biological Sciences
University of California,
Santa Barbara, California

ABSTRACT

In locating feral honey bee colonies, the various updated versions of an ancient “bee box” technique can require several days to locate each colony. A feral colony can be found more efficiently (within only a few hours) by following a given general protocol and then exploiting one or more specific techniques of varying effectiveness. The general protocol includes the initiation of beelines, marking and timing bees, sensitively interpreting round trip times, providing adequate rewards to insure rapid recruitment, using an appropriate marking scent, and exploiting wind direction and wind borne odors. Effective specific techniques include interpreting the behavior of water gatherers (allowing recruitment of others), following natural bee lines (”aerial pathways”) downwind to a colony, and mass conversion of bees from blossoms to feeding stations.

Contact address:
Adrian M. Wenner,
Dept. of Biological Sciences,
University of California, Santa Barbara,
Santa Barbara, CA 93106
Phone: 805-893-2838
Fax: 805-893-8062

Manuscript received: January 26, 1992
Manuscript accepted: March 13, 1992
Keywords: honey bees, Apis mellifera, feral colonies, bee hunting, conservation/restoration

Adrian M. Wenner, Professor of Natural History and Provost of the College of Creative Studies at University of California at Santa Barbara, has worked with honey bees since 1946, both commercially and in research. He has also studied crustacean growth and reproduction, island biogeography, monarch butterflies, and the philosophy and sociology of science. As a graduate student in neurobiology and behavior at Cornell, Joe F. Alcock has begun a study of fiddler crab behavior. Daniel E. Meade is a graduate student at UCSB, where he studies foraging ecology.

INTRODUCTION

Honey bee colonies have been sought in the wild for thousands of years, likely long before written history (e.g. Crane 1983). The first lucid and complete account known (Columella ~50 A.D. [1954]) was based in part on information obtained from Virgil.

The United States had no Apis sp. honey bees before European settlers repeatedly introduced bees into the eastern United States from various parts of Europe. These introductions began early in the 17th century (Sheppard 1989). In fact, the wave of feral colonies moved westward faster than the settlers themselves, leading native Americans to label them “white man’s fly” - knowing that settlers would not be far behind (e.g. Barton 1793). Before the establishment of domestic beekeeping in any newly inhabitated area, settlers started their apiaries by finding feral colonies derived from escaped swarms.

With the rise of domestic beekeeping in the U.S., beehunting evolved from the simple foraging exercise (e.g. Dudley 1723; Duden 1826) it had remained in other parts of the world into somewhat of a sport (see Thoreau 1852 [1906]; Burroughs 1875; Plumley 1919; Scoville 1927; Edgell 1949; Parsons 1949; Billings 1961; Morse 1969; Chapman 1970; Donovan 1980). Native Americans participated also and sometimes did better than the settlers themselves (e.g. Duden 1826 [1980]).

Several years ago we relied on earlier accounts of how one might find colonies by use of the “bee box” technique and embarked upon a formidable task (Wenner 1989). We agreed to help The Nature Conservancy and the National Park Service in their Northern Channel Islands conservation/restoration program by locating and removing all feral (wild) European honey bee colonies from Santa Cruz Island, off the coast of Santa Barbara, California. Preliminary results (unpublished) indicate that the European bees on the island visit primarily introduced European flora, while the more than a hundred species of solitary bees visit mostly native flora. We work under the assumption that removing the honey bees should help restore the island to a pre-European ecology.

Santa Cruz Island is a rather barren island (25,000 hectares, 96 square miles of mountainous terrrain). Because of the severe seasonal drought (usually no rain between April and November), we estimated that the vegetation could not support more than 30 colonies. The colonies are apparently all descendents of one or two colonies introduced before 1880. According to ranch records kept since 1880, no other honey bees have been introduced since that time. By late 1991, we had already located about 125 colonies, with many more still to find.

Within only a few months after starting, we found that the traditional bee box technique (e.g. Edgell 1949; Donovan 1980; Visscher and Seeley 1989) was too slow for the task ahead of us (see below). Instead, published accounts from further back in history (e.g. Columella 50 A.D. [1954]; Dudley 1723; Burroughs 1875) led us to more effective techniques. We also learned that various techniques did not work equally well in different situations - to be efficient, one may have to use a variety of techniques in any given circumstance. We subsequently increased the speed at which we located colonies; experienced volunteers now usually need less than half a day to find a target colony on this difficult terrain. The best approach is to assume a treasure hunt or scavenger hunt attitude and to exploit each available clue as it is revealed by the behavior of foragers and searching bees.

METHODS

For successful bee hunting one must be fully aware of basic bee biology, including bee orientation behavior, sugar and honey characteristics, the role of scent, and the effects of wind direction.

Beelining - In most cases, one needs to ascertain both direction of the homeward flight line and round trip times for bees visiting food or water sources. Such information is normally acquired after one converts foragers from flowers or water to an alternative and more convenient source (e.g. Thoreau 1852; Burroughs 1875; Edgell 1949; Chapman 1970). With bearing and distance determined, one can eventually narrow down colony location. The entire process is usually known as “beelining” (e.g. Dudley 1723; Billings 1961; Morse 1973; Visscherand Seeley 1989). We have found, as others before us (e.g. Dudley 1723; Thoreau 1852; Chapman 1970), that the first few round trips by a newly converted forager do not provide reliable bearing and round trip times. Neither does knowledge of bearing alone provide one with sufficient information; the colony can be any distance away; but, if one can get bearings for a colony from feeding stations located in different directions, triangulation becomes possible (e.g. Dudley 1723; Burroughs 1875; Parsons 1949; Visscher and Seeley 1989).

Recruitment of new foragers from the parent colony can be exceedingly slow, even during a dearth of natural sources of nectar, unless one exploits wind direction in the manner discussed below. In fact, obtaining new recruits is often the most difficult part of the hunt and nearly impossible with only one or two foragers when food is more than 300m downwind from the colony (Friesen 1973; Wenner, Meade and Friesen 1991).

Marking and Timing Bees - Once round trips have become routine for a few bees, individuals must be distinguished from one another to obtain round trip times. Water foragers often settle on the same spot each time they return for a load, permitting timing of flights without a need to mark individuals. After several trips, foragers visiting sugar solution or diluted honey can be marked with paint (model airplane enamel, in our case), colored dust (as Columella used), or even cooking lime (as used by Mexican cowboys: C. H. Muller, personal communication). That is because nectar foragers may alight anywhere at a dish where there is room. If one anaesthetizes foragers, numbered disks or Christmas-tree tinsel can be glued in place.

With foragers now recognizable as individuals, the time of arrival of each marked bee is tallied (one needs no more than 10 round trips per bee), while recording at the same time the homeward bearing of marked individuals. One normally needs at least three (but no more than six) bees coming from each colony to counter error due to between-bee variation. The third or fourth shortest time for each bee (rather than the mean or median time) provides a usable estimate of round trip time (Figure 1). The median time for several such estimates usually fit our formula closely (see below), but times were shorter than normal for the Frazer Point colony due to exceptionally light winds all day (see Wenner 1963.)

Interpreting Round Trip Time Data - Earlier accounts ranged from vague statements about colony distance (e.g. Edgell 1949:20) to more specific flight time equivalents (e.g. Donovan 1980:85). Such accounts are not particularly helpful; during field work we use a simple formula: x = 150y - 500 (straight line in Figure 1). That is, to estimate distance (x = meters or yards) to each colony, we multiply complete round trip time (y = time between arrivals) by 150 and subtract 500 from the result. (The constant value of 500 represents the time spent filling at the station and unloading in the colony - see Wenner 1963). Error can be considerable, because several bees (marked and unmarked) can be landing and departing each minute, markings are not always clearly distinguishable, some individual foragers are not consistent, wind (as well as uphill or downwind flight paths) alters time, and foragers from more than one colony can be traveling to the feeding station.

Two examples of multiple-colony visitation are shown in Figure 1. At our Prisoners Stream feeding station, some round trip times by foragers indicated that the target colony was located about 250m east of our feeding station (Pris. East-1). However, round-trip times of other foragers heading in the same direction suggested that a second colony might be about 700m away. By treating the data as a bimodal distribution, we eventually located both colonies (Pris. East-2, also). The same circumstance prevailed for a colony we found at a 755m distance; a second colony was later located at about 1700m from our station in the same direction. These examples illustrate the reliability of estimates derived from the formula.

Reward - Regardless of technique used, one must provide an appropriate reward or foragers will not switch from their customary routine. In earlier and more elaborate efforts, a piece of dark honeycomb, filled previously with sugar syrup by means of a medicine dropper, served as a substitute for normal forage. Others have used diluted honey. Some people have even heated beeswax or old comb in further efforts to attract bees (e.g. Edgell 1949). As a substitute for honeycomb, we found that a piece of cellulose sponge, moistened with water, placed on an inverted coffee can, and then saturated with scented sugar solution or a 1:1 honey:water mixture, provided an adequate substitute for honeycomb.

Recipes abound for the composition of the sugar solution or sugar syrup used. Edgell (1949:9) suggested that “…white sugar one-third, and water two-thirds, boiled for fifteen minutes and then cooled, seems to be as tempting to bees as real honey.” In cool weather, pure honey may be too thick (see Billings 1961). Since the nectar concentration in flowers varies widely, no one concentration need be used. If nectar flow is moderate (i.e. when nectar sources are readily available to colonies), one must use a fairly concentrated solution to compete with nature. During heavy honey flows in spring or early summer, competition with nature may be impossible.

In our earlier work, we settled on one part sugar to one part hot water by volume, with scent added (see below). We now most often use diluted honey (1:1 honey:water, by volume) or undiluted honey instead of sugar solution or sugar syrup. Additional scent is then unnecessary (e.g. Thoreau 1852 [1906]). The use of scented sugar solution instead of honey has one advantage: if recruitment is too rapid and too many bees interfere with attempts to obtain round trip times and bearings, one can reduce the scent level and thereby reduce the number of new arrivals per unit time (Wells and Wenner 1971). Refrigeration of the stock solution between uses prevents the growth of mold.

Scent - Table sugar (sucrose) is one of the purest subtances one can buy and has a vapor pressure of zero. That means that sucrose solution has no odor of its own, hence, some suitable marker odor must be provided. Fructose has a pleasant odor but is difficult to locate and expensive. Bees often ignore glucose (personal observation; Root et al. 1947). Besides the reward provided in any device used to catch foragers, one must establish a feeder station with similar scent nearby for those foragers who return on their own.

Although scent is necessary (e.g. von Frisch 1939), just about any scent will do (e.g. Edgell 1949:9), since there seems to be no such thing as an inherently “attractive” odor to bees (e.g. Wenner and Wells 1990: Excursus NG). Edgell advocated the use of anise, but on Santa Cruz Island we cannot use that scent; introduced sweet fennel (”anise”) plants cover large regions. Ten droplets of clove oil per liter (quart) of sugar solution works fine; such an amount is strong and helps overpower the scent of natural forage in the colony. That is, the more odor accumulation in the colony, the more likely that other bees will find the newly established station (e.g. Wenner, Wells and Johnson 1969).

After establishing a steady stream of bees traveling between station and colony and then determining both bearing and distance, one can often find the colony rather quickly. Any one of several techniques, listed below in generally decreasing order of effectiveness, will suffice. However, circumstances dictate which technique might be best under given conditions.

SPECIFIC TECHNIQUES

Water Foragers - As Columella noted 2000 years ago, most bee colonies are located near water. That makes sense, if only because bees have little energy left to fly once they have filled their “fuel tanks” (honey stomachs) with water. Water sources can be few and far between in the southwestern United States (but not so few as one might suppose). On Santa Cruz Island, we study topographic maps to locate likely outcroppings of springs, often situated at junctions of streams. Slowly hiking up water courses may reveal bees collecting water; their home colony is usually nearby (Columella 50 AD. [1954]).

Finding bees at water is not necessarily easy, since each bee has its own particular site and requires 4-5 minutes per round trip. Furthermore, individuals blend in all too well with the background while drinking, and only a few of the bees from isolated colonies are water carriers at any one time. If one walks up a stream course too fast, water collectors can be easily missed, especially if colonies are small.

As indicated above, the fidelity of each bee to a rather precise point at a water collecting site allows one to estimate round trip times without disturbing individuals by marking them. Furthermore, these regular water foragers already head on a “bee line” to their colony without the preliminary circling flights characteristic of converted foragers or inexperienced recruits. Our record time to locate a colony with this technique is 24 minutes.

The disadvantage of this technique is that no new recruits arrive, and one can only work with the few water foragers at hand. Neither can one move closer to the colony to get another bearing or a shorter round trip time except by going along the stream and finding another water forager from the same colony.

Columella advocated capturing bees at water, slipping them into a reed one by one until a dozen or more had been obtained, and then letting them out one at a time (Butler republished that recipe in 1609). By successive moves toward the colony, while noting new vanishing bearings (just as Mexican cowboys have done after dusting foragers with cooking lime), one gets ever closer to the colony. However, such a disturbance may disorient bees, and they may not head directly home.

Water to Honey Conversion - Water foragers can be exploited in a different way - by simply converting them from water to honey. After finding a bee imbibing water, we dip a thin stick into undiluted honey and ease this honey drop to a point upslope from the bee’s mouth parts as it imbibes water. With care, a steady hand, and close observation, one can see the bee’s tongue suddenly switch from water to the honey. Soon foragers and recruits will visit nearby sponges soaked with honey water, and an immediate round trip pattern results.

The switch from water-to-honey technique works especially well during hot weather and/or when natural nectar sources are scarce. That is both because colonies must be cooled by evaporation of water and because colonies must dilute honey in their stores before it can be used for colony functions. Attempting to switch bees to a point source of food by putting honey on a blossom fed upon by bees is more difficult. To date, we have been able to get foragers already feeding on blossoms to switch to honey only when their parent colonies are near starvation levels or when they are foraging on nectar-poor plants.

Following Bee Lines - In an area with stable wind patterns, foragers primarily visit plants located upwind from their colonies (see Friesen 1973). After positioning oneself in a patch of flowers and looking downwind or slightly crosswind with binoculars (at different times of day to exploit various light conditions) one can locate major aerial pathways between that patch and target colonies. By repositioning oneself at a vanishing point toward the colony along one such pathway and repeating the operation, one can get ever closer to that colony and gain an ever better estimate of colony bearing.

Aerial flyways can also be located if one is positioned at right angles to that pathway and has good backlighting (e.g. a woods in shadow behind the flyway), because honey bees can be distinguished from other flying insects by their characteristic (deliberate) bee line flight. Also, sprinkling cooking lime or other powder on foragers often causes them to stop foraging and head for home. The white dust provides greater visibility for tracking the homeward flight path.

The visual technique is difficult for the novice beehunter, since it requires a thorough understanding of foraging patterns, topographical constraints, and characteristic flight patterns. Many hours may be needed before a full deductive skill is developed; we find some people are more intuitive in this exercise than others. Further suggestions include the following: 1) Several bees flying in the same direction are usually traveling to a colony; 2) Bees flying in several directions from a patch suggest the presence of more than one colony, if so, bee lines must be distinguished from one another before proceeding; 3) Proceed along the bee line, stopping often to refine the bearing and to establish new landmarks; 4) Patience pays - perhaps no more than one or two bees from a colony will pass along the aerial pathway each minute. As one gets closer to the target colony, bees will pass by more frequently.

Mass Conversion - With only one or two bees routinely traveling between station and colony, as occurs commonly with the bee box technique (new recruits may not arrive for hours or days - see below), odor accumulation in the colony and recruitment at the station occur only slowly (see Wenner and Wells 1990: Ch. 5, Excursus OS). Searching recruits that attempt to follow the drifting odor trail left in the air by only one or a few foragers continually lose odor cues and must retrace their course repeatedly; that may explain why an inordinate amount of time is needed by recruits as they search for the station frequented by foragers (e.g. Gould et al. 1970: Table 4; Wenner and Wells 1990: Excursus NEG).

To expedite odor accumulation in the colony and to increase dramatically the odor trail left in the air between station and colony by foragers during their round trips, we invented a crude technique effective in areas of medium to heavy bee visitation on flowers. We sweep the blossoms with an insect net until up to 25 foragers have been captured and are in the net. The net is then inverted into a large coffee can and the opening covered with stiff cardboard.

The can has been previously prepared: we place a dampened cellulose sponge laced with honey water (with its own scent) in the bottom of the can and cover the sponge with a coarse plastic screen. That screen reduces the chance that bees will become wetted down (fouled) with the reward while attempting to escape. Three or four ten-penny nail holes punched in the side of the can near the bottom provide some light and insure that foragers in the can move to the bottom as they attempt to escape. They then touch the reward with their legs and begin feeding.

After a sufficient time (at least 4 min) has elapsed for all bees to become engorged with the honey-water, the lid is removed and bees permitted to escape. All of them then return to the colony and provide a large input of odor in the dance area at approximately the same time. In the meantime, another series of sweeps made with the net provides a second batch of potential foragers, etc. Even though only a small percentage returns after each attempt, the large number handled insures rather rapid success in establishing a bee line, an aerial pathway that may provide odor cues between station and colony (e.g. Wenner and Wells: Ch. 5, Excursus OS).

Bee Boxes - Use of specially constructed “bee boxes” to locate colonies has gained the most attention the past few hundred years. In fact, the effort of constructing such a box seems to impart almost a mystical nature to the practice of bee lining (e.g. Dudley 1723; Thoreau 1852 [1906]; Plumley 1919; Edgell 1949; Chapman 1970; Morse 1973).

Most traditional bee boxes share common features, including two (or more) compartments with a sliding panel between two of them. One chamber has a closeable door to the outside which a bee can be knocked into from off a flower. The second chamber has a window that can be covered from outside and a port through which a bee can be released after it has had its fill of reward. If such boxes are small enough, they can be held in one hand, leaving the other hand free to tap bees into the first chamber.

Once a bee has been caught and the first chamber darkened by closure of the door, raising the panel between the two chambers allows the bee to pass into the lighted second chamber. After passage of the bee into the lighted chamber, the panel between the two compartments is closed, the window to the outside of the second chamber is closed, and the bee has little else to do but fill up on the scented sugar solution or diluted honey previously placed in the comb or on a sponge. In the meantime, one can knock another bee into the first chamber and repeat the process; a dozen or more bees can be caught in sequence.

We found that the traditional bee-box technique was too inefficient for the Santa Cruz Island beehunt project, that of locating all the feral colonies in the 25,000 hectare (96 sq. mi.) area. In our experience, most often at least half a day passed before we had even a single bee making round trips. Frequently, more than two days had passed before we had obtained reliable bearings and round trip times; sometimes more than four days were required to locate a single colony. We no longer bother with this technique.

Indicator Plants - The distance bees will travel apparently depends on the abundance of nectar and/or pollen that they can obtain from the various species of plants they visit and the intensity of competition from other colonies (e.g. Peer 1955; Lee 1961). On Santa Cruz Island we have found bees as far as 2 km upwind but only a short distance downwind from their colony when they visit introduced horehound (Marrubium vulgare) and yellow mustard (Brassica spp.). Since both these plants and the honey bees themselves are European species, one might expect that this visitation reflects a co-evolution in this rather long distance exploitation of food sources.

The situation differs in the case of bee visitation on Santa Cruz Island’s native plants. We have found honey bees foraging no more than 1 km directly upwind from their colonies on mule fat (Baccharis glutinosa), while those gathering pollen from coyote bush (Baccharis pilularis) have exceeded that distance. Bees visiting deerweed (Lotus scoparius) and island buckwheat (Erioginum arborescens) have been found no more than 500m upwind and those on doveweed (Eremocarpus setigerus) no more than 100m from their colonies.

The disadvantage of using indicator plants is somewhat obvious; each locality will have a different circumstance of visitation, one that will change according to wind direction, time of day, season, competition, and colony condition. One must become somewhat of an insect-plant expert to exploit this technique fully. Fortunately, conditions are remarkably simple on Santa Cruz Island; wind comes past each colony from only one direction throughout the summer on much of the island, and the total number of plant species visited by honey bees is quite small. Accordingly, we have had some success with this technique.

Satellite Stations - Once we have an approximate bearing and distance for colony location, we place one or more satellite feeding stations a hundred or more meters upwind or crosswind from the suspected location. If the estimate is fairly accurate, recruits begin arriving in great numbers immediately, since they first search areas close to their own colony (e.g. von Frisch 1939).

COLONY CAVITIES

Colonies need not be in “bee trees,” and one must examine possible locations carefully. Of the first 122 colonies we located on Santa Cruz Island (nearly all of those that occur on the eastern half of the island), cliff holes, rock crevices, and rock overhangs sheltered 33, 30, and 13 of them, respectively (62%). Six were found in cavities in clay banks, and 19 were found in cavities under the tree boles of scrub oak and island cherry. Only 17 colonies (14%) were located in what could be considered bee trees, even though we have seen dozens of large oaks with apparently suitable cavities on the island. By contrast, in a study of colonies encountered by California residents (Gambino et al. 1990), nearly half of the 193 colonies reported had been found in trees; somewhat fewer were found in structures. (See their paper for a review of other such studies.)

DISCUSSION

Now that African(ized) bees are here in the United States, a method to locate feral colonies with dispatch could be a great asset for bee researchers, beekeepers, and the cities in which they live. A joint and concerted community beehunting effort can be instituted to locate any colonies that may be unknown but that may pose a threat to hikers. This effort can supplement a campaign such as that practiced in the Panama Canal Zone (Boreham and Roubik 1987; Caron and Gray 1991), where colonies were reported when found (as in the Gambino et al. 1990 study). These joint efforts could well lead to an impressive reduction of the African bee problem in limited areas.

Similarly, national parks, state parks, recreation areas, and other agencies interested in conservation/restoration of their holdings could repeat our experience on The Nature Conservancy’s Santa Cruz Island Preserve (Channel Islands National Park). Swarms normally move a geometric mean distance of only 800m (lognormal) from their parent colonies (Wenner, Meade, and Friesen 1991), hence, repopulation would be rather slow and could be monitored. The benefits would be numerous, the costs rather minimal.

ACKNOWLEDGMENTS

This project has been supported by a grant from The Nature Conservancy and a faculty research grant from the University of California. The College of Creative Studies (UC Santa Barbara) provided financial support for interns. Technical input and assistance was provided throughout the project by R. W. Thorp (UC Davis), by S. L. Buchmann, J. 0. Schmidt, and S. C. Thoenes (USDA, Tucson), and by P. K. Visscher (UC Riverside), who provided an extensive bibliography of bee hunting publications. We also thank S. L. Buchmann and R. W. Thorp for their review of the manuscript. Special thanks go to H. Carlson, J. Sulentich, and R. Klinger (The Nature Conservancy), to Lyndal Laughrin (UC Santa Cruz Island Reserve), to A. H. Schuyler and B. Fagan for assistance in transportation, and to the many student volunteers who have assisted us these past four years.

REFERENCES

Barton, B. S. 1793. An inquiry into the question, whether the Apis mellifica, or true honey-bee, is a native of America. Trans. of the Amer. Phil. Soc. 3:241-261.
Billings, H. 1961 (July). A-beein’ in the Ozarks. Am. For. 67: 28-29.
Boreham, M. M. and D. W. Roubik. 1987. Population change and control of Africanized honey bees (Hymenoptera: Apidae) in the Panama Canal area. Bull. Entomol. Soc. Am. 33: 34-39.
Burroughs, J. 1875. An idyl of the honey-bee. Birds and Bees and other Studies in Nature. Houghton Mifflin Co., New York.
Butler, C. 1609 (1969). The Feminine Monarchie. Da Capo Press, New York.
Caron, D. M. and H. Baltazar Gray. 1991. The impact of the Africanized bee on beekeeping in Panama. BeeScience. 1:139-143.
Chapman, A. 1970 (Aug). Hunting the bee tree. Am. For. 76: 16-19.
Columella, L. J. M. ~50 AD. (1954). Lucius Junius Moderatus Columella on Agriculture. Translation by E. S. Forster and E. H. Heffner. Harvard Univ. Press, Cambridge, MA.
Crane, E. 1983. The Archaeology of Beekeeping. Cornell Univ. Press, Ithaca, NY.
Donovan, R. E. 1980. Hunting Wild Bees. Winchester Press, Tulsa, OK.
Duden, G. 1826. (1980). Report on a Journey to the Western States of North America and a Stay of Several Years Along the Missouri (During the Years 1824, ‘25, ‘26, and 1827). An English Translation. The State Historical Society of Missouri and University of Missouri Press, Columbia, MO.
Dudely, P. 1723. An account of a method lately found out in New-England, for discovering where the bees hive in the woods, in order to get their honey. Phil. Trans. R. Soc. London, B. 31 (367): 148-150.
Edgell, G. H. 1949. The Bee Hunter. Harvard Univ. Press, Cambridge, MA.
Friesen, L. J. 1973. The search dynamics of recruited honey bees, Apis mellifera ligustica Spinola. Biol. Bull. 144: 107-131.
Frisch, K. von. 1939. The language of bees. In Annual Report of the Smithsonian Institution for the Year Ended June 30, 1938, pp. 423-431. Publication 3491. Washington, D.C.: U.S. Government Printing Office.
Gambino, P., K. Hoelmer, and H. V. Daly. 1990. Nest sites of feral honey bees in California, USA. Apidologie. 21: 34-45.
Gould, J. L., M. Henerey, and M. C. MacLeod. 1970. Communication of direction by the honey bee. Science. 169:544-554.
Lee, W. R. 1961. The nonrandom distribution of foraging honey bees between apiaries. J. Econ. Entomol. 54: 928-933.
Morse, R. A. 1969 (Aug). Uncle Perk’s bee tree. Field and Stream. 74: 8+ _________.1973 (June-July). Tracking the wild bee. Conservationist. 27:14-17.
Parsons, C. 1949 (5 Sept). Like honey? Hunting the wild bee. Time. 54: 47. (about Edgell)
Peer, D. F. 1955. THE FORAGING RANGE OF THE HONEY BEE. PART 1, PH.D. Thesis, University of Wisconsin.
Plumley, L. 1919 (Jan). Wild bee hunting. Country Life. 35: 81.
Root, A.I., E.R. Root, H.H. Root, and M.J. Deyell. 1947. The ABC and XYZ of Bee Culture. A.I. Root, Medina, OH.
Scoville, S., JR. 1927 (23 July). The bee tree. Independent. 119: 89-90.
Sheppard, W. S. 1989. A history of the introduction of honey bee races into the United States. Am. Bee J. 121: 617-619, 664-667.
Thoreau, H. D. 1852. (1906). Bee hunting. In The Writings of Henry David Thoreau; pp. 368-375 in Journal IV (Bradford Torrey, ed.). Houghton Mifflin and Co., Boston.
Visscher, P. K. and T. D. Seeley. 1989. Bee-lining as a research technique in ecological studies of honey bees. Amer. Bee J. 129: 536-539.
Wells, P. H. and A. M. Wenner. 1971. The influence of food scent on behavior of foraging honey bees. Physiol. Zool. 44: 191-209.
Wenner, A. M. 1963. The flight speed of honeybees: A quantitative approach. J. Apic. Res. 2: 25-32.
__________. 1989. “Bee-lining” and ecological research on Santa Cruz Island. Am. Bee J. 129: 808-809.
Wenner, A. M., D. E. Meade, and L. J. Friesen. 1991. Recruitment, search behavior, and flight ranges of honey bees. Amer. Zool. 31 (6): (in press)
Wenner, A. M. and P. H. Wells. 1990. Anatomy of a Controversy: The Question of a “Language” Among Bees. Columbia Univ. Press, New York.
Wenner, A. M., P. H. Wells, and D. L. Johnson. 1969. Honey bee recruitment to food sources: Olfaction or language? Science. 164: 84-86.

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1990 Wenner, A.M. and P.H. Wells. Anatomy of a Controversy: The Question of a “Language” Among Bees. Columbia University Press. Pages 274-284.
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“[The] development becomes known to the public. Popular science books . . . spread the basic postulates of the theory; applications are made in distant fields, money is given to the orthodox and is withheld from the rebels. More than ever the theory seems to possess tremendous empirical support.”

- Paul Feyerabend 1975:43

Scientists reputedly welcome challenge and readily adopt new interpretation whenever evidence no longer conforms to observed phenomena (Kneller 1978). Both Mahoney (1976) and Veldink (1989) found otherwise. Some comments can clarify why this discrepancy in attitudes about scientific objectivity exists.

Collectively, and over varying time spans, Kneller’s assessment appears to be correct; scientific interpretation changes constantly and relatively rapidly. That change in commitment to interpretation is one feature that distinguishes science from religion.

When new evidence arises that conflicts with “conventional wisdom” on relatively important issues in science, however, individual scientists may momentarily rush to the defense of an existing paradigm with religious fervor. Feyerabend deplored this defensive behavior of scientists, and wrote: “To sum up: Unanimity of opinion may be fitting for a church. . . . Variety of opinion is necessary for objective knowledge” (1975:46; emphasis Feyerabend’s).

Something akin to religious fervor was what we encountered when we challenged the dance language hypothesis. That was because that hypothesis had already evolved into a full-fledged paradigm spanning several fields. As might be expected, some felt that our impression - that the defense of the dance language hypothesis was extreme - was a gross exaggeration, since “everyone knows that” scientists shun “censorship” and suppression of new ideas. We were even considered by some scientists to be “paranoid” (Veldink 1989).

Prior to our challenge, and while we were still respected members of the animal language research community, our manuscripts received favorable reviews and were published without undue delay. Subsequent to that challenge, we encountered increasing hostility (as noted in excursus SI). Within a very few years, our chance of getting material into print or obtaining grant support went from difficult to nearly impossible. The negative evidence we had gathered relative to the dance language hypothesis seemed to be accorded little significance in this development (e.g., Veldink 1989).

Due to the extreme adverse reaction we encountered, we feel that this volume would not be complete - dealing as it does with the human side of scientists - unless we included some documentation for the comments made above. Accordingly, we provide here a chronological sequence of some pertinent material spanning those first few critical years.

ATTEMPTED REBUTTAL TO A LETTER BY RICHARD DAWKINS IN THE JOURNAL SCIENCE

Richard Dawkins, in a letter to Science, exemplified the confusion existing among some ethologists on the significance of our crucial experiment (Wenner, Wells, and Johnson 1969; see also our chapter 10). He objected to the interpretation we provided for the results of that experiment, in part, as follows:

Suppose a man tells me there is a bar three blocks down the street on the right. I set off thirsty, but on the way a strong smell of beer distracts me to another bar hidden up a side alley. Does this prove that human language does not communicate information? . . . it is entirely reasonable to suppose that bees have alternative ways of finding food - among them, the dance, smell, and the presence of other bees - and that each of these cues may predominate under different circumstances. For example, the artificial use of strong scent might cause olfactory cues to prevail. (1969:751)

Dawkins concluded his letter:

“In brief, bees are easily distracted. This modest and uncontroversial conclusion is all that can be drawn
from the experiments purporting to disprove von Frisch’s classic work” (1969:751).

Dawkins thereby adhered close to the “party line” of those committed to the dance language paradigm. From our perspective, on the other hand, Dawkins’ ad hoc rationalization merely confused the issue, because he ignored the potential influence of wind direction, an essential element when one considers odor movement and resultant insect flight (see chapter 5 and excursus OS). Furthermore, Dawkins contradicted von Frisch’s earlier assertions that recruit bees would not be distracted during their outgoing flight. For example, von Frisch had written:

There is no doubt that the bees understand the message of the dance. When they fly out, they search only in the neighborhood of the indicated range, ignoring dishes set closer in or farther away. Not only that, they search only in the direction in which the original feeding dish is located. (1962:78; see also our chapter 9)

Scent (notice in the above passage Dawkins’ use of the word strong), strong or otherwise, if of the right kind, can “distract” searching bees (see chapter 8), but only if searching bees are already downwind from such a source. In that sense odors are quite a different stimulus than either sound or light stimuli.

Dawkins, for example, could have been distracted by the smell of beer during his tavern hunt only if the second tavern had been upwind of his original path. But he could have been distracted by the sound or sight of the tavern, regardless of its direction from his original path. It is unlikely that Dawkins would have been distracted by the odor of a bakery under the circumstances of his example.

Other dance language proponents began citing Dawkins’ letter as refutation of our work, even though it was obviously nothing more than an expression of opinion, faulted at that by Dawkins’ failure to consider wind direction in his polemical statement. This curious turn of events led one of us (Wells) to draft a reply to that Dawkins letter. We present that short draft here in its entirety so that the referee’s comments, which follow and which formed the basis for rejection by Science, will fall into perspective.

Reply to Dawkins

Subsequent to our comparison of the predictive values of the “olfaction” and “language” hypotheses of honey bee forager recruitment [Wenner, Wells, and Johnson 1969], Richard Dawkins generated a challenge [Dawkins 1969] which confuses key issues. I am surprised to find that some ethologists have [now] accepted Dawkins’ discussion as an excuse for disregarding our experimental work. It is necessary, therefore, to reply to his three objections to our paper.

First, Dawkins asserted that we “presume to challenge the findings of a great biologist” (Karl von Frisch). To this we plead guilty; but, the charge isn’t serious enough. In so doing we also presume to challenge a host of subsequent investigators who have uncritically accepted any and all of von Frisch’s interpretations of available data; and who have used these interpretations as a foundation on which their own work is based. This may account for the emotion occasionally engendered by our papers.

Next, he objected to our citing (in our paper’s introductory paragraphs) earlier studies which yielded data inconsistent with the hypothesis that bees use linguistic communication [Wenner, Wells, and Rohlf 1967; Wenner and Johnson 1966; Johnson 1967a; Lopatina 1964; Wells and Giacchino 1968; Wenner 1967] on grounds that the failure of bees to use language does not prove they have none. With this we heartily agree! Experimentation does not “prove” or “disprove” hypotheses, for formally this can never be done. Rather, it tests their usefulness in the a priori prediction of events. Usually one retains as his working hypotheses those with high predictive success under the most rigorously controlled conditions.

Rather than “purporting to disprove von Frisch’s classic work” (Dawkins’ misinterpretation of the intent of our experiments) our experimental results show that, under well controlled conditions, the olfactory hypothesis is successful while the language hypothesis fails to predict the distribution of recruit foragers in the field.

As a substantive objection to our paper, Dawkins suggested in an eloquent but anthropomorphic analogy that crowds of bees or strong scent might distract recruits just as “the smell of beer distracts me” from a prior destination. We purposely inserted controls against these possibilities in our experiments. These controls preclude his interpretation. As the legend to our Fig. 2 [in Wenner, Wells, and Johnson 1969] indicates, one experimental feeding station always was upwind and the other one downwind from a control station visited by no bees.

According to Dawkins’ line of reasoning, bees would have had to be simultaneously distracted to the scent at that control site both upwind and downwind from their “intended” destinations. Either that, or recruited bees would have failed to verify von Frisch’s contention that “if no clues are provided by scent the bees use information conveyed by the dance” [von Frisch 1968:532].

Furthermore, on days nine and fourteen of our experimental series [table 1 of Wenner, Wells, and Johnson 1969] no scent was placed at the control station. In the absence of “distracting” odor cues, recruit bees still failed to find the unscented experimental stations about which they might have been linguistically informed, even though “crowds of bees” were there.

But back to Richard Dawkins’ delightful analogy. I feel confident that he and I would agree that scientific questions are more likely to be resolved by experimentation than by polemics such as our exchange of letters in Science. If I am right, and we can avoid being distracted from that one key point of agreement, next time he is in Los Angeles, I’ll buy!

The letter by Wells attempting to reply to Dawkins was rejected by the editor of Science. The rejection was accompanied by a set of comments by an anonymous referee. We include that text, also in its entirety.

Referee’s Comments

This paper is uncomplimentary to all scientists who do not accept the author’s theory. Specifically the author accuses ethologists of accepting Dawkins’ discussion as “an excuse for disregarding our experimental work.” Can’t he accept the thought that they reject his work because after critical study they do not consider it irrefutable?

And if that were not infuriating enough he accuses a “host of investigators for uncritically accepting any and all of von Frisch’s interpretations.” How can he presume to know that? Is any new theory accepted without critical study? Perhaps if his own work were as thorough, complete and convincing as that of von Frisch he would not need to berate other scientists for not accepting it. The use of polemics is a poor substitute for irrefutable facts.

THE FIRST ATTEMPT TO REBUT THE GOULD, HENEREY, AND MACLEOD 1970 ARTICLE

In 1970 Gould, Henerey, and MacLeod, three undergraduate students at the California Institute of Technology, published their ten-page lead article in Science (see chapter 12). By then we were becoming aware of the sociological implications of the dance language controversy. Nevertheless, we were still ill prepared for what transpired next.

A study of the paper by Gould and co-workers revealed that these undergraduates had gathered data for only a few hours at the end of the summer in 1969, just after the publication of our crucial experiment paper (Wenner, Wells, and Johnson 1969). It was immediately apparent to us that the results they had obtained were completely at variance with the expectations of the dance language hypothesis (see chapters 1 and 13). At the same time their results supported our interpetation (see excursus NEG).

One of us (Wenner) submitted a letter to Science that stressed the fact that their results, in fact, supported our position. In the cover letter to that manuscript Wenner included the following comments:

After carefully studying that article, I have concluded that it is not really a challenge of our work but a substantiation of our earlier findings. Unfortunately, this aspect of their study is clouded by the investigators efforts to discredit our earlier work and by their conclusion which does not necessarily follow from their data [see our chapter 12].

I think that your readers should have an opportunity to read this divergent opinion. Toward this end I have written the enclosed comment, “A Divergence from the Expected,” and hope you will publish it. I have kept my statement short and in the nature of opinion so that it can appear as a Letter in your journal.

Wenner also included a list of names of reputable people who could serve as referees, as follows: Bernard Abbott, Kenneth Armitage, Vincent Dethier, Jerry Downhower, W. George Evans, and Edward Glassman. We do not know if any of them were used.

The manuscript was rejected, and comments from only a single referee were appended, with the last portion of those comments removed. Furthermore, the number 2 circled at the top of the page (in the manner of Science editors in those days) indicated that the comments of referee number 1 were not furnished to us. We reproduce that one referee’s comments verbatim:

Dr. Wenner should read the paper he is criticizing! The average time spent by successful recruit bees was 3 to 4 minutes as clearly stated in the second paragraph on p. 551 of Gould et al and in their Table 2. It can only be perversity which led Wenner to utilize, instead, the data in Table 7, which contains (clearly stated!) data on delay between dance attendance and arrival at a feeding site. To repeat, Gould et al. clearly show the successful recruit takes 3 to 4 minutes to fly to the feeding site not “40 times the usual flight time.”

Contrary to Dr. Wenner’s incredible “interpretation” of Gould et al. all 225 recruits arrived at a station. Only 37 were marked, the 188 not listed in Table 4 were unmarked recruits, not unsuccessful recruits! Thus, we have 25 marked bees that arrived at the “correct” station and 12 that arrived at a station +/- 180 degrees from the correct direction. No marked recruits arrived at stations +/- 90 degrees from the correct direction. This may be an interesting comment on bee language. I have performed bee-language experiments with my animal behavior classes and we have occasionally seen: (a) foragers which indicated both the training direction and the opposite direction in a single bout of dancing, (b) foragers which indicated only the opposite to the training direction [sic]. In both of these cases, it has been my impression that these errors were the result of interference in the dances by too many attendants to the dance. However, occasional bees may be misled by such errors, or as some of our data also suggest, there may be some ambiguity or confusion in the dance which can lead recruits to seek food in a direction 180 degrees from the dance direction. If the editors of Science would like such a comment on the excellent paper by Gould et al. I should be happy to write one. (unpublished comments by an anonymous referee; emphasis his)

THE FATE OF AN APPEAL TO REBUT THE SAME ARTICLE

When Wenner showed the above comments to Larry Friesen, Friesen replied, “But virtually everything the referee wrote is wrong in point of fact!” We then sat down and listed some of the errors in the referee’s comments as follows:

1. The “3 to 4 minutes as clearly stated” applied to forager bees, not recruits.
2. There are no data on flight times in table 2.
3. There is no table 7 in the Gould, Henerey, and MacLeod paper.
4. From page 550 one can conclude that 225 recruits arrived, but on page 552 Gould and co-workers clearly stated: “Only 37 (that is, 13 percent) of these 277 attendants were successfully recruited and later caught at a feeding station. . . .”
5. There were no stations at locations 90 degrees from the correct direction.
6. The original dance language hypothesis does not permit recruits to arrive at a station located 180 degrees from the “correct” direction.

At that point Friesen insisted to Wenner, “if you point out the errors made by the referee, they will have to publish your letter.” Wenner replied, “Larry, you don’t know what’s going on, do you?”

Nevertheless, Wenner wrote a detailed comparison of the comments made by the referee to the actual facts in the case, revised the manuscript slightly, and resubmitted it with a cover letter. Some of the points made in the cover letter are as follows:

Normally I would not appeal such a decision on your part (This is the third consecutive rejection of such a manuscript by Science [on this issue]); but in this latest case I feel the referee has done both Science and me a disservice. The referee has made some serious errors in fact, a point which will undoubtedly make a difference to you as editor. (Attached herewith please find documentation for this claim.)

Even if the referee had been correct, my opinion should be printed in Science. In this regard I have always admired Science’s stated policy:

“Science serves its readers as a forum for the presentation and discussion of important issues related to the advancement of science, including the presentation of minority or conflicting points of view, rather than by publishing only material on which a consensus has been reached.”

I would hope that this particular controversy does not fall outside that general policy. That is, if Science can print a 10-page challenge of our work, surely space can be provided for a few paragraphs of divergent opinion. Even if my facts had been wrong (which they are not), I do not see why the referee should object to an airing of my opinion. If I had been as incorrect as he implied, supporters of the language hypothesis should be pleased to see such gross errors on my part in print.

On the positive side, the referee’s comments did help reveal two minor errors in my manuscript (neither of which he pinpointed and neither of which makes any difference in my argument). For purposes of clarification, however, I have modified the manuscript slightly. The revised manuscript is enclosed.

I trust you will pass favorably on my manuscript this time around. Thank you for considering the topic once again.

We provide here the entire manuscript, as submitted the second time to Science.

A Divergence from the Expected

A recent article in Science, “Communication of direction by the honey bee,” is of more than incidental interest, as it contains an extensive amount of data not available earlier. However, some important aspects of their data do not fall into line with what one might expect from the classic honey bee “dance language” hypothesis. The series 1 experiments were especially pertinent in this regard, since they were: “. . . designed to examine the behavior of individual recruits as each attended a dance and subsequently arrived at a feeding station.”

The first item of interest is that approximately a third of those marked recruits which succeeded in finding a station arrived at one in a direction opposite to that “indicated” in the dance maneuver. This result clearly contradicts the expectations of the classic hypothesis [von Frisch 1947, 1950; Wenner, Wells, and Johnson 1969]. All 37 of the successful bees should have arrived at the “correct” station.

In their study these investigators also found that successful recruit bees spend a considerable amount of time in flight, on the average, before reaching a food source. The direct line flight time between hive and feeding stations in their experiments would normally be only about 24 sec [Wenner 1963]; and recruited bees generally fly from the hive within a minute and a half after leaving a dancing bee (50% leave within 30 sec) [Wenner 1963]. With these facts in mind, an examination of the data in Table 4 of the Gould, et al. paper reveals that the 25 bees which arrived at the “correct” station flew an average of about 30 times longer than necessary if they were to “fly directly out” to the food source. (Interestingly enough, the 12 bees which ended up at a station in the opposite direction averaged only 36 times as long as necessary.)

Clearly the above results do not match the expectation that “. . . recruits alarmed by these dancers . . . find the feeding place with surprising speed and precision” [von Frisch 1967].

Another interesting point is that Gould and co-workers have confirmed earlier findings of my co-workers and myself [Johnson and Wenner 1970; Wenner, Wells, and Rohlf 1967]; that is, that bees which leave the hive are not likely to find a food source, even under the best of circumstances [Note: Some confusion results here because Gould and co-workers apparently define "recruits" as those bees which succeed in reaching a station. I prefer to consider recruits as those bees which leave the hive after contact with a successful forager]. [For example, they stated]: “Only 37 of 277 attendants were successfully recruited and later caught at a feeding station, even though the high molarity of the sucrose used reportedly produces maximum dancing and recruitment.

If one works in unscented localities, rather than in the highly pungent area such as that chosen by the authors, the recruitment efficiency becomes even worse. As we reported earlier in Science, while working with unscented sucrose in a relatively odor-free locality, “. . . in the absence of a major nectar source for the colony, we received only five recruits from a hive of approximately 60,000 bees after ten bees had foraged at each of four stations for a total of 1374 round trips during a 3-hour period” [Wenner, Wells, and Johnson 1969]. This result is especially interesting since we have subsequently found that ever smaller amounts of odor in the food and in the locality result in an ever higher frequency of dancing, all other factors being equal [unpublished data, available upon request].

In summary, these workers have obtained results which generally agree with what we have published earlier and which differ markedly from those obtained earlier by von Frisch and co-workers or with those expected on the basis of the classic language hypothesis [von Frisch 1967]. And, in contradistinction to the authors’ conclusion, the results are generally in excellent agreement with what we would expect under the circumstances. The final distribution of successful recruits does differ significantly from what one might obtain if these animals had flown at random about the countryside, it is true, but there is no compelling reason for assuming that the dance language hypothesis is a necessary explanation for this divergence from randomness [Wenner, Wells, and Johnson 1969]. I feel that a slight difference in composite location odor near the experimental sites, perceptible to bees but not to investigators, could have been responsible [Johnson and Wenner 1970].

This second time around, the manuscript was again rejected. The comments of only one referee were enclosed, as follows:

As one of the referees who urged Science to publish earlier research reports by Dr. Wenner and his associates, I now recommend that you do not publish this particular critique of the article by Gould, Henery, and MacLeod. Enough is enough! His critique, the referee’s criticism of the critique, and his countercritique of the referee’s criticism, have become labyrinthine. It is conceivable that the Gould et al. paper should not have been published; however, you would require a jury discussing all this material for hours to decide. I no longer believe it is worth the effort, because Dr. Wenner’s own criticisms of the von Frisch explanation are mostly beside the point, and they consistently fail to mention a growing body of positive evidence strongly favoring the von Frisch explanation. To publish an increasingly complex and ambiguous debate on but one aspect of negative evidence is an inefficient - and misleading - use of space in Science. If and when Dr. Wenner comes up with solid evidence to support his views, he will find an interested and sympathetic audience waiting.

(Note in the above referee comments the reliance on the verification approach.)

REBUFF OF THE ATTEMPT TO REBUT THE 1975 GOULD SCIENCE ARTICLE

Five years after the Gould, Henerey, and MacLeod lead article, Science published an eight-page lead article by Gould, based upon work he did for his doctoral dissertation (see our chapter 12). Again, this article was a challenge of our work. Once again, Wenner “tested the water” to see if Science was by now receptive to the publication of a rebuttal of the paper by Gould (actually a clarification). The letter of inquiry included the following statements:

Anyone well-versed in the work of my colleagues and myself who carefully reads the Gould article will certainly recognize a convergence of viewpoints, with Gould now coming very close to our position in the bee language controversy. Unfortunately, he does not mention the existence of that convergence, and those not overly familiar with our work will likely miss that important point.

As evidence of the foregoing, Wenner provided some appendices to the letter, documentation that juxtaposed quotations from our earlier work and quotations from the Gould paper so that the editor could compare the statements. Wenner continued:

Dr. Wells and I would like to know whether Science might publish something similar to that which is enclosed. While it is true that there is a normal process which should be followed (submitting a manuscript), our recent attempts at getting material into print in Science have failed because of an intensely hostile peer review. . . . Dr. Wells and I eventually got our message into Nature ([1973] 241:171-175), but only after a considerable delay. Ironically, Gould has now gotten that same message into print in Science - See the two starred items in the enclosed Appendix A.

Gould’s publication of the same train of thought as we have expressed earlier would indicate that the climate has now changed at Science. Is this true? Do our comments now have a chance of getting into print? We realize that space is at a premium and would be happy to have only the cover statement considered [two manuscript pages], provided some reference would be made to the availability of the two appendixes for those who might wish to obtain them directly from us.

The cover statement included two quotations from the Gould paper, as follows:

Excepting the experiments reported here, the locale-odor hypothesis can effectively account for all the results achieved to date [including those of von Frisch] without recourse to the dance-language theory. . . . recruitment to odors alone might be the usual system in honey bee colonies not under stress. (1975b:686, 691)

The same assistant editor again rebuffed our attempt, and wrote:

When you have an experimental paper we would be glad to consider it. However, we would not be interested in publishing your cover statement, with or without the appendices. . . . As for the circumstances that prevail here, bee classes [sic] are not among the controversies on which we have a position.

In the years that followed, however, Science continued to publish papers supportive of the dance language hypothesis and peripheral research areas, but one finds no research papers reporting results that do not fit within the dance language paradigm.

By J. W. WHITE, JR. AND LANDIS W. DONER(1)
BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 82 - 91

Honey is essentially a highly concentrated water solution of two sugars, dextrose and levulose, with small amounts of at least 22 other more complex sugars. Many other substances also occur in honey, but the sugars are by far the major components. The principal physical characteristics and behavior of honey are due to its sugars, but the minor constituents - such as flavoring materials, pigments, acids, and minerals - are largely responsible for the differences among individual honey types.

Honey, as it is found in the hive, is a truly remarkable material, elaborated by bees with floral nectar, and less often with honeydew. Nectar is a thin, easily spoiled sweet liquid that is changed (”ripened”) by the honey bee to a stable, high-density, high-energy food. The earlier U.S. Food and Drug Act defined honey as “the nectar and saccharine exudation of plants, gathered, modified, and stored in the comb by honey bees (Apis mellifera and A. dorsata); is levorotatory; contains not more than 25% water, not more than 0.25% ash, and not more than 8% sucrose.” The limits established in this definition were largely based on a survey published in 1908. Today, this definition has an advisory status only, but is not totally correct, as it allows too high a content of water and sucrose, is too low in ash, and makes no mention of honeydew.

Colors of honey form a continuous range from very pale yellow through ambers to a darkish red amber to nearly black. The variations are almost entirely due to the plant source of the honey, although climate may modify the color somewhat through the darkening action of heat.

The flavor and aroma of honey vary even more than the color. Although there seems to be a characteristic “honey flavor,” almost an infinite number of aroma and flavor variations can exist. As with color, the variations appear to be governed by the floral source. In general, light-colored honey is mild in flavor and a darker honey has a more pronounced flavor. Exceptions to the rule sometimes endow a light honey with very definite specific flavors. Since flavor and aroma judgments are personal, individual preference will vary, but with the tremendous variety available, everyone should be able to find a favorite honey.

—-
(1) Research leader and research chemist, respectively, Science and Education Administration, Eastern Regional Research Center, Philadelphia, Pa. 19118.

Composition of Honey

By far, the largest portion of the dry matter in honey consists of the sugars. This very concentrated solution of several sugars results in the characteristic physical properties of honey - high viscosity, “stickiness,” high density, granulation tendencies, tendency to absorb moisture from the air, and immunity from some types of spoilage. Because of its unique character and its considerable difference from other sweeteners, chemists have long been interested in its composition and food technologists sometimes have been frustrated in attempts to include honey in prepared food formulas or products. Limitations of methods available to earlier researchers made their results only approximate in regard to the true sugar composition of honey. Although recent research has greatly improved analytical procedures for sugars, even now some compromises are required to make possible accurate analysis of large numbers of honey samples for sugars.

An analytical survey of U.S. honey is reported in Composition of American Honeys, Technical Bulletin 1261, published by the U.S. Department of Agriculture in 1962. In this survey, considerable effort was made to obtain honey samples from all over the United States and to include enough samples of the commercially significant floral types that the results, averaged by floral type, would be useful to the beekeeper and packer and also to the food technologist. In addition to providing tables of composition of U.S. honeys, some general conclusions were reached in the bulletin on various factors affected by honey composition.

Where comparisons were made of the composition of the same types of honey from 2 crop years, relatively small or no differences were found. The same was true for the same type of honey from various locations. As previously known, dark honey is higher than light honey in ash (mineral) and nitrogen content. Averaging results by regions showed that eastern and southern honeys were darker than average, whereas north-central and intermountain honeys were lighter. The north-central honey was higher than average in moisture, and the intermountain honey was more heavy bodied. Honey from the South Atlantic States showed the least tendency to granulate, whereas the intermountain honey had the greatest tendency.

The technical bulletin includes complete analyses of 490 samples of U.S. floral honey and 14 samples of honeydew honey gathered from 47 of the 50 States and representing 82 “single” floral types and 93 blends of “known” composition. For the more common honey types, many samples were available and averages were calculated by computer for many floral types and plant families. Also given in this bulletin are the average honey composition for each State and region and detailed discussions of the effects of crop year, storage, area of production, granulation, and color on composition. Some of the tabular data are included in this handbook.

Table 1 gives the average value for all of the constituents analyzed in the survey and also lists the range of values for each constitutent. The range shows the great variability for all honey constituents. Most of the constituents listed are familiar. Levulose and dextrose are the simple sugars making up most of the honey. Fructose and glucose are other commonly used names for these sugars. Sucrose (table sugar) also is present in honey, and is one of the main sugars in nectar, along with levulose and dextrose. “Maltose” is actually a mixture of several complex sugars, which are analyzed collectively and reported as maltose. Higher sugars is a more descriptive term for the material formerly called honey dextrin.

The undetermined value is found by adding all the sugar percentages to the moisture value and subtracting from 100. The active acidity of a material is expressed as pH; the larger the number the lower is the active acidity. The lactone is a newly found component of honey. Lactones may be considered to be a reserve acidity, since by chemically adding water to them (hydrolysis) an acid is formed. The ash is, of course, the material remaining after the honey is burned and represents mineral matter. The nitrogen is a measure of the protein material, including the enzymes, and diastase is a specific starch-digesting enzyme.

Most of these constituents are expressed in percent, that is, parts per hundred of honey. The acidity is reported differently. In earlier times, acidity was reported as percent formic acid. We now know that there are many acids in honey, with formic acid being one of the least important. Since a sugar acid, gluconic acid, has been found to be the principal one in honey, these results could be expressed as “percent gluconic acid” by multiplying the numbers in the table by 0.0196. Since actually there are many acids in honey, the term “milliequivalents per kilogram” is used to avoid implying that only one acid is found in honey. This figure is such that it properly expresses the acidity of a honey sample independently of the kind or kinds of acids present.

In table 1, the differences between floral honey and honeydew honey(2) can be seen. Floral honey is higher in simple sugars (levulose and dextrose), lower in disaccharides and higher sugars (dextrins), and contains much less acid. The higher amount of mineral salts (ash) in honeydew gives it a less active acidity (higher PH). The nitrogen content reflecting the amino acids and protein content is also higher in honeydew.

The main sugars in the common types of honey are shown in table 2. Levulose is the major sugar in all the samples, but there are a few types, not on the list, that contain more dextrose than levulose (dandelion and the blue curls). This excess of levulose over dextrose is one way that honey differs from commercial invert sugar. Even though honey has less dextrose than levulose, it is dextrose that crystallizes when honey granulates, because it is less soluble in water than is levulose. Even though honey contains an active sucrose-splitting enzyme, the sucrose level in honey never reaches zero.

Honey varies tremendously in color and flavor, depending largely on its floral source. Its composition also varies widely, depending on its floral sources (table 2). Although hundreds of kinds of honey are produced in this country, only about 25 or 30 are commercially important and available in large quantities. Until the comprehensive survey of honey composition was published in 1962, the degree of compositional variation was not known. This lack of information hindered the widespread use of honey by the food industry.

Water Content

The natural moisture of honey in the comb is that remaining from the nectar after ripening. The amount of moisture is a function of the factors involved in ripening, including weather conditions and original moisture of the nectar. After extraction of the honey, its moisture content may change, depending on conditions of storage. It is one of the most important characteristics of honey influencing keeping quality, granulation, and body.

Beekeepers as well as honey buyers know that the water content of honey varies greatly. It may range between 13 and 25 percent. According to the United States Standards for Grades of Extracted Honey, honey may not contain more than 18.6 percent moisture to qualify for U.S. grade A (U.S. Fancy) and U.S. grade B (U.S. Choice). Grade C (U.S. Standard) honey may contain up to 20 percent water; any higher amount places a honey in U.S. grade D (Substandard).

These values represent limits and do not indicate the preferred or proper moisture content for honey. If honey has more than 17 percent moisture and contains a sufficient number of yeast spores, it will ferment. Such honey should be pasteurized, that is, heated sufficiently to kill such organisms. This is particularly important if the honey is to be “creamed” or granulated, since this process results in a slightly higher moisture level in the liquid part. On the other hand, it is possible for honey to be too low in moisture from some points of view. In the West, honey may have a moisture content as low as 13 to 14 percent. Such honey is somewhat difficult to handle, though it is most useful in blending to reduce moisture content. It contains over 6 percent more honey solids than a product of 18.6 percent moisture.

In the 490 samples of honey analyzed in the Department’s Technical Bulletin 1261, the average moisture content was 17.2 percent. Samples ranged between 13.4 and 22.9 percent, and the standard deviation was 1.46. This means that 68 percent of the samples (or of all U.S. honey) will fall within the limits of 17.2 ± 1.46 percent moisture (15.7 - 18.7); 95.5 percent of all U.S. honey will fall within the limits of 17.2 ± 2.92 percent moisture (14.3 - 20.1).

In the same bulletin, a breakdown of average moisture contents by geographic regions is shown. These values (percent) are North Atlantic, 17.3; East North Central, 18.0; West North Central, 18.2; South Atlantic, 17.7; South Central, 17.5; Intermountain West, 16.0; and West, 16.1.

Sugars

Honey is above all a carbohydrate material, with 95 to 99.9 percent of the solids being sugars, and the identity of these sugars has been studied for many years. Sugars are classified according to their size or the complexity of the molecules of which they are made. Dextrose (glucose) and levulose (fructose), the main sugars in honey, are simple sugars, or monosaccharides, and are the building blocks for the more complex honey sugars. Dextrose and levulose account for about 85 percent of the solids in honey.

Until the middle of this century, the sugars of honey were thought to be a simple mixture of dextrose, levulose, sucrose (table sugar), and an ill-defined carbohydrate material called “honey dextrin.” With the advent of new methods for separating and analyzing sugars, workers in Europe, the United States, and Japan have identified many sugars in honey after separating them from the complex honey mixture. This task has been accomplished using a variety of physical and chemical methods.

Dextrose and levulose are still by far the major sugars in honey, but 22 others have been found. All of these sugars are more complex than the monosaccharides, dextrose and levulose. Ten disaccharides have been identified: sucrose, maltose, isomaltose, maltulose, nigerose, turanose, kojibiose, laminaribiose, a, B-trehalose, and gentiobiose. Ten trisaccharides are present: melezitose, 3-a-isomaltosylglucose, maltotriose, l-kestose, panose, isomaltotriose, erlose, theanderose, centose, and isopanose. Two more complex sugars, isomaltotetraose and isomaltopentaose, have been identified. Most of these sugars are present in quite small quantities.

Most of these sugars do not occur in nectar, but are formed either as a result of enzymes added by the honeybee during the ripening of honey or by chemical action in the concentrated, somewhat acid sugar mixture we know as honey.

Acids

The flavor of honey results from the blending of many “notes,” not the least being a slight tartness or acidity. The acids of honey account for less than 0.5 percent of the solids, but this level contributes not only to the flavor, but is in part responsible for the excellent stability of honey against microorganisms. Several acids have been found in honey, gluconic acid being the major one. It arises from dextrose through the action of an enzyme called glucose oxidase. Other acids in honey are formic, acetic, butyric, lactic, oxalic, succinic, tartaric, maleic, pyruvic, pyroglutamic, a-ketoglutaric, glycollic, citric, malic, 2- or 3-phosphoglyceric acid, a- or B-glycerophosphate, and glucose 6-phosphate.

Proteins and Amino Acids

It will be noted in table 1 that the amount of nitrogen in honey is low, 0.04 percent on the average, though it may range to 0.1 percent. Recent work has shown that only 40 to 65 percent of the total nitrogen in honey is in protein, and some nitrogen resides in substances other than proteins, namely the amino acids. Of the 8 to 11 proteins found in various honeys, 4 are common to all, and appear to originate in the bee, rather than the nectar. Little is known of many proteins in honey, except that the enzymes fall into this class.

The presence of proteins causes honey to have a lower surface tension than it would have otherwise, which produces a marked tendency to foam and form scum and encourages formation of fine air bubbles. Beekeepers familiar with buckwheat honey know how readily it tends to foam and produce surface scum, which is largely due to its relatively high protein content.

The amino acids are simple compounds obtained when proteins are broken down by chemical or digestive processes. They are the “building blocks” of the proteins. Several of them are essential to life and must be obtained in the diet. The quantity of free amino acids in honey is small and of no nutritional significance. Breakthroughs in the separation and analysis of minute quantities of material (chromatography) have revealed that various honeys contain 11 to 21 free amino acids. Proline, glutamic acid, alanine, phenylalanine, tyrosine, leucine, and isoleucine are the most common, with proline predominating.

Amino acids are known to react slowly, or more rapidly by heating, with sugars to produce yellow or brown materials. Part of the darkening of honey with age or heating may be due to this.

Minerals

When honey is dried and burned, a small residue of ash invariably remains, which is the mineral content. As shown in table 1, it varies from 0.02 to slightly over 1 percent for a floral honey, averaging about 0.17 percent for the 490 samples analyzed.

Honeydew honey is richer in minerals, so much so that its mineral content is said to be a prime cause of its unsuitability for winter stores. Schuette and his colleagues at the University of Wisconsin have examined the mineral content of light and dark honey. They reported the following average values:
Enzymes

One of the characteristics that sets honey apart from all other sweetening agents is the presence of enzymes. These conceivably arise from the bee, pollen, nectar, or even yeasts or micro-organisms in the