[1992 Wenner, A.M., J.E. Alcock, and D.E. Meade. Efficient hunting of feral colonies. Bee Science 2:64-70.]
Adrian M. Wenner, Joe E. Alcock and Daniel E. Meade
Department of Biological Sciences
University of California,
Santa Barbara, California
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.
Adrian M. Wenner,
Dept. of Biological Sciences,
University of California, Santa Barbara,
Santa Barbara, CA 93106
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.
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. ) 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 ; 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 ).
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. ; 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.
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 ). 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.
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. ).
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 ; 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).
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.)
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.
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.
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.