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in his lab placed the sugar water on a boat in the middle of a small lake. When scouts returned to the hive to communicate their find, Gould claimed that other bees refused to go with them, even though they frequently flew over the lake to reach pollen sources on the opposite shore. To Gould, these observations suggested that "bees somehow consider information to see if it makes sense before they act on it". No one observed behavior of bees in the hive during this experiment, however, and when the experiment was replicated by Wray et al. in 2008, the researchers found no evidence for Gould's assertion that followers of waggle dances advertising "implausible" sites refused to respond to these dances. The authors' conclusion: "Contrary to prior findings, our results provide no evidence that honeybees assess the plausibility of information contained in waggle dances or reject dances for locations that are unlikely to yield food. Thus, we conclude that the original Lake
Experiment should no longer be cited as evidence that honeybees possess cognitive maps, ‘insight’ or ‘imagination’."
180:. He performed an elegant experiment that showed not only that the bees could discriminate colors but that they demonstrated associative learning. He first trained his bees to feed from a small dish filled with a nectar-like sugar water. This dish was placed on a piece of blue colored cardboard so that the color was visible to the bees as they arrived at the dish and fed. Next, von Frisch placed identically sized pieces of cardboard in varying shades of grey, each with a dish, all around the blue piece. Lacking color vision, the bees should visit one or more of the gray pieces as often as the blue piece, but he found the vast majority of the bees flew directly to the blue piece of cardboard on which they had previously obtained their reward. The bees largely ignored the gray pieces which had not been rewarded. Von Frisch repeated the experiment with other colors like violet and yellow and got the same results. Later other researchers used this experimental design to test the color vision of vertebrates.
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choice between two patterns (matching and non-matching) placed at different distances (Can bees trained in the task continue to perform correctly when the matching pattern as well as a non-matching pattern are presented in the tunnel?); and a choice between two patterns (Can bees learn which of two sequentially encountered patterns in the tunnel is the pattern to be matched in the decision cylinder?). The researchers found that working memory in the honey bee is both robust and flexible. The experiments demonstrated that bees can choose between alternatives, determine if a stimulus is the same or different than one seen earlier, remember the earlier one for a short period, and generalize this performance to new pairs of stimuli. The bees retained information in working memory for about 5 seconds, and they might have been simultaneously learning a matching and a nonmatching task; further research was needed.
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386:(the "dwarf honeybees") the dance is performed on the dorsal, horizontal portion of the nest, which is exposed. The runs and dances point directly toward the resource in these species. Each honey bee species has a characteristically different correlation of "waggling" to distance, as well. Such species-specific behavior suggests that this form of communication does not depend on learning but is rather determined genetically. It also suggests how the dance may have
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429:. Significant to the argument are the experiments of William F. Towne, of the Kutztown University in Pennsylvania, in which hives are moved to "mirror image" terrain settings, and the bees are thereby fooled into dancing about the wrong location for a nectar source. Foragers were successfully recruited to the wrong location, but only when the sun was obscured by clouds, forcing them to rely on terrain-based navigation rather than "
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pigments which have maximal absorbance at wavelengths of 350 nm, 440 nm, and 540 nm. Menzel also found that most of the cells he studied had secondary sensitivities that corresponded to wavelength regions at which the other two receptor types were maximally active. He used spectral efficiency experiments to suggest that these secondary sensitivities result from electric coupling between the receptors.
406:. They argue that bees instead are primarily recruited by odor. The purpose of the dance is simply to gain attention to the returning worker bee so she can share the odor of the nectar with other workers who will then follow the odor trail to the source. Most scientists agree that odor is used in recruitment to resources, but they differ strongly in opinion as to the information content of the dance.
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mate to the resource. Once an individual finds a good food source, they will return to the same source for many days. If an individual detects the scent of a rival bee, they will avoid the plant in order to avoid conflict and to save time. It has also been shown that pheromones are a method of sexual selection between male drones and queens.
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a different colored background. One of the colors was the same as that used during the initial trial, and the others were novel, unrewarded colors. Remarkably, after a single trial and several days without exposure to the rewarded color, bees correctly chose to explore the color used in the first trial more than fifty-percent of the time.
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dish could not be seen but the hive was still visible. When released one by one, the bees appeared disoriented for a few seconds, then flew directly to the dish, 73 of 75 bees reaching it in about 28 seconds. They apparently accomplished this feat by devising a new flight path based on a cognitive map of visible landmarks.
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by marking individual cells with a fluorescent dye and recording from these cells as single units. From this analysis he determined that there are three types of receptors in the honey bee eye: 1) UV receptors, 2) blue receptors, and 3) green receptors The three receptors contain three rhodopsin-like
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in the stomach. The forager bees feed this primer pheromone to the worker bees, and the chemical keeps them in a nurse bee state. The pheromone prevents the nurse bees from maturing too early to become forager bees. As forager bees die off, less of the ethyl oleate is available and nurse bees more
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Menzel then repeated this experiment with another group of bees, keeping all factors the same except that in the second round of testing he gave the bees three initial trials with the rewarded color instead of just one. When, after several days in confinement, the bees were presented with a choice of
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Menzel did several experiments to answer these questions. First, he gave individual bees a single sugar reward on a colored background. He then kept these bees in small cages for several days without any further trials. After a few days, he presented each bee with an array of several dishes, each on
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Interpretations of another test suggested not only the use of a map, but also an ability to remember and combine relevant information. Gould moved a supply of sugar water 25% farther away from a hive each day. The bees communicated the location of the water to each other as usual. Then a researcher
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Neither of these points invalidate the dance theory, but simply suggest that odor might be involved, which is indeed conceded by all proponents of dance theory. Critics of the odor plume theory counter that most natural nectar sources are relatively large—orchards or entire fields— so, precision may
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Other experiments further document the communicative nature of the waggle dance. For example, dances by robotic dummy bees induced some recruitment. Research has also shown that the dance may vary with the environmental context, a finding that may explain why the results of some earlier studies were
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theory and the "odor plume" theory. The dance theory is far more widely accepted, and has far more empirical support than the odor theory. Supporters of the dance theory often grant odor a significant role in recruitment, while supporters of the odor theory have claimed that the dance is essentially
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To test bees ability to distinguish between two different colors, Menzel placed a small dish containing sugar-water in one circle and a second empty dish some distance away on a differently colored circle. A single bee was placed equidistant between the two circles and allowed to choose between the
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The German scientist
Randolf Menzel continued the study of color vision in honey bees with more detailed tests. He was curious about whether bees would learn certain colors faster than others. He used lights of various color and intensity to project circles of light on a surface, a set-up like that
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for information they have learned, and utilize it when foraging. In an experimental demonstration, Gould lured some bees to a dish of artificial nectar, then gradually moved it farther from the hive. He marked the trained bees, placed them in a darkened jar, and relocated them to a spot where the
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adjusts its distribution of labor most beneficially. In order to survive as a bee colony of sometimes 50,000–100,000 individual bees, the communal structure has to be adaptable to seasonal changes and the availability of food. The division of labor has to adjust itself to the resources available
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also showed that they discriminated between rewarding and unrewarding patterns. This knowledge persisted, both after 24 hours of learning and after 7 days. Dr. Muth's studies have also shown that bumblebees do not prefer nor choose against a flower based on its complexity. However, they will learn
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In still other experiments, Menzel explored the timing of bee color learning by testing whether bees register color before, during, or after receiving their sugar-water reward. For this purpose Menzel displayed the color beneath a rewarded dish at different stages of the honey bee feeding process:
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of honey bees, after learning to associate a certain pattern with a reward (delayed matching-to-sample). Bees were shown a pattern at the beginning of a tunnel, and then subjected to a series of variations: in the length of the tunnel (How long can bees retain the pattern in working memory?), in a
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requires such learning. For example, honey bees make few repeat visits to a plant if it provides little in the way of reward. A single forager will visit different flowers in the morning and, if there is sufficient reward in a particular kind of flower, she will make visits to that type of flower
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rely on pheromones for much of their communication with nest mates and rivals. They produce pheromones from their labial glands. The function of signaling depends on the profitability, but they commonly will scent mark a food source either for self-orientation, to deter rivals or to direct a nest
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After his work on color preferences, Menzel extended his experiments to study aspects of color learning and memory. He wanted to know how many trials bees need to reliably choose a previously rewarded color when they are presented with several alternatives, and how long they would remember the
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correlated the runs and turns of the dance to the distance and direction of the food source from the hive. He reported that the orientation of the dance is correlated with the relative position of the sun to the food source, and the length of the waggle portion of the run is correlated to the
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logical difficulties of a small-scale dance (a few centimeters across) giving directions precise enough to hold the other bees on course during a flight that could be several kilometers long. Misreading by even a few degrees would lead the bee off course by hundreds of meters at the far end.
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Menzel's results showed that bees do not learn to discriminate between all color pairs equally well. Bees learned the fastest when violet light was rewarded, and the slowest when the light was green; the other colors fell somewhere in between. This evidence of inherent bias is evolutionarily
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Menzel found that bees register color during both approach and feeding, and that they had to see the color for about a total of about 5 seconds, with best performance usually coming with about three seconds exposure during the approach and two seconds after landing and beginning to feed.
257:. Bumblebees were shown to be able to learn multiple color-food associations and tended to continue to apply what they learned. In another study, Dr. Felicity Muth continued to learn more about these associations. Bumblebees initially preferred yellow
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from foraging. While the division of labor in a bee colony is quite complex, the work can be roughly seen as work inside the hive and outside the hive. Younger bees play a role inside the hive while older bees play a role outside the hive mostly as
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Testing for color vision in honey bees. The majority of bees flew directly to the dish with the blue background as they had been trained to do. Thus, they were able to discriminate between gray and blue backgrounds, showing their capability for color
328:). A waggle run oriented 45° to the right of ‘up' on the vertical comb indicates a food source 45° to the right of the direction of the sun outside the hive. The abdomen of the dancer appears blurred because of the rapid motion from side to side.
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This ability to retain information about color-linked rewards for several days after minimal exposure to the rewarded color demonstrates the remarkable facility with which bees learn and retain color information.
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Odor is essential and even necessary at various stages of the recruitment process, including once a recruited forager reaches the vicinity of the resource while some scientists think that dancing may be a simple
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and retain this information for several days at least; they communicate the location and nature of sources of food; they adjust their foraging to the times at which food is available; they may even form
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Wray MK, Klein BA, Seeley TD. 2011. Honey bees use the social information in waggle dances more fully when foraging errors are more costly. Behavioral
Ecology. 23:125-131. doi:10.1093/beheco/arr165
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distance of the food from the hive. Von Frisch also reported that the more vigorous the display is, the better the food. Von Frish published these and many other observations in his 1967 book
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Menzel, R. and
Backhaus, W. 1989. Color vision in honey bees: Phenomena and physiological mechanisms. In D. Stavenga and R. Hardie (eds): Facets of vision. Berlin-Heidelberg-New York: 281-297
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dishes. The bees quickly learned to choose the color signaling the dish with the reward, and Menzel was able to measure how quickly the bees learned this task with various color differences.
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While many researchers believe that bee dances give enough information to locate resources, proponents of the odor plume theory argue that the dance gives little, or no actual guidance to a
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Muth, Felicity; Keasar, Tamar; Dornhaus, Anna (March 2015). "Trading off short-term costs for long-term gains: how do bumblebees decide to learn morphologically complex flowers?".
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A seminal paper by Menzel (1975) described the morphology and spectral sensitivity of the honey bee eye that underlie their color vision. He examined color-coding in the honey bee
1054:. in: Hartfelder, K.H, De Jong, D. et al. eds. (2004) Proceedings of the 8th IBRA International Conference on Tropical Bees and VI Encontro sobre Abelhas. Ribierao Preto: USP/FM
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Menzel, R. and
Backhaus, W. 1991. Colour Vision in Insects. In P. Gouras (ed): Vision and Visual Dysfunction. The Perception of Colour. London: MacMillan Press, 262-288.
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movement that conveys no information. Others see the dance as conveying information, but doing it poorly compared to other means and potentially used backup approach.
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cells were found to have long visual nerve fibers that penetrated the lamina with deep tree-like branchings. Blue and green receptor cells had more shallow fibers.
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reasonable, given that bees forage for differently-colored nectar-bearing flowers, many of which are to be found in green foliage which does not signal reward.
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to forage in the same area. The factors that determine recruiting success are not completely known but probably include evaluations of the quality of
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269:. After this initial test, they began associating floral color with pollen success. The bumblebees' association between pollen and features of the
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Leoncini I, Le Conte Y, Costagliola G, Plettner E, Toth AL, Wang M, Huang Z, Becard JM, Crauser D, Slessor KN, Robinson GE (December 14, 2004).
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used by von Frisch except that, by using light instead of cardboard, Menzel was able to easily change the intensity and color of the circles.
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The
Austrian zoologist Karl von Frisch began the exploration of color vision in honey bees when, in 1919, he asked whether or not bees have
1307:"The use of food source scent marks by the stingless bee Trigona corvina (Hymenoptera: Apidae): the importance of the depositor's identity"
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Michelsen, A.; Anderson, B. B.; Kirchner, W. H.; Lindauer, M. (1989). "Honeybees can be recruited by a mechanical model of a dancing bee".
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Riley, J.R.; Greggers, U.; Smith, A.D.; Reynolds, D.R.; Menzel, R. (2005). "The flight paths of honeybees recruited by the waggle dance".
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quickly mature to become foragers. It appears that this control system is an example of decentralized decision making in the bee colony.
470:, can be used to communicate the quality of a food source, temperature, a need for water, and the condition of the queen (Sebeok, 1990).
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Later work has supported Von Frisch's observations and added many details. It appears that all of the known species and races of
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Munz, T. (November 2005). "The Bee
Battles: Karl von Frisch, Adrian Wenner and the Honey Bee Dance Language Controversy".
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An
American specialist in bee cognition, Dr. Felicity Muth, has studied the mechanism behind the associative learning in
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1359:"The Trail Pheromone of a Stingless Bee, Trigona corvina (Hymenoptera, Apidae, Meliponini), Varies between Populations"
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not be necessary or even desirable. They have also challenged the reproducibility of the odorless source experiment.
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and later
Randolf Menzel began asking questions about color vision and various aspects of color learning in bees.
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Genetic
Control of the Honey Bee (apis mellifera) Dance Language: Segregating Dance Forms in a Backcrossed Colony
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Jarau, Stefan; Dambacher, Jochen; Twele, Robert; Aguilar, Ingrid; Francke, Wittko; Ayasse, Manfred (2010-09-01).
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Frisch, K. von. 1956. Bees; their vision, chemical senses, and language. Ithaca, N.Y., Cornell University Press.
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irrelevant to recruitment. The academic debate between these theories has been polarized and sometimes hostile.
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experiments with odorless sugar sources which show that worker bees are unable to recruit to those sources and
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Menzel, R; Blakers, M (1975). "Colour receptors in the bee eye — Morphology and spectral sensitivity".
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Visscher, P.K. and Tanner, D.A. (2004). Sensory aspects of recruitment-dance performance in honey bees
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Dyer, F.C.; Seeley, T.D. (1991). "Dance dialects and foraging range in three Asian honey bee species".
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exhibit the behavior, but details of its execution vary among the different species. For example, in
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A number of experiments have demonstrated color recognition, discrimination and memory in honey bees
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Note: much of the research on the two competing hypotheses of communication has been restricted to
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354:. This waggle pattern of movement was thought to attract the attention of other bees. In 1947,
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Currently the group is working on four projects in the area of mid-air collision avoidance...
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for most of the day, unless the plants stop producing nectar or weather conditions change.
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https://www.apiservices.biz/en/articles/sort-by-popularity/562-why-do-honeybees-dance-2000
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Sebeok (1990). Essays in Zoosemiotics. Toronto: Toronto Semiotic Circle. ISSN 0838-5858 .
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Sherman, G.; Visscher, P.K. (2002). "Honeybee colonies achieve fitness through dancing".
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Boogert, Neeltje Janna; Hofstede, Frouke Elisabeth; Monge, Ingrid Aguilar (2006-05-01).
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Foragers communicate their floral findings in order to recruit other worker bees of the
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Anthophila. Some species have been studied more extensively than others, in particular
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823:"Bees remember flowers for more than one reason: pollen mediates associative learning"
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Research that was published in November 2004, by scientists under the leadership of
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colors they almost always chose the color that was used on the first three trials.
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Gould, J L "A honey of a question, are bees intelligent?" Discover, 1986, August
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There are two main hypotheses to explain how foragers recruit other workers—the
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Miller, J A "Biology" Science News; 4/23/1983, Vol. 123 Issue 17, p271, 1/6p
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Swarming bees require good communication to all congregate in the same place
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of their surroundings. They also communicate with each other by means of a "
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506:. Huang's team found that forager bees gather and carry a chemical called
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Miller, Julie Ann (April 23, 1983). "Do Bees Plan Ahead Intelligently?".
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74:. They can demonstrate capabilities such as color discrimination through
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Behavioral Neurobiology: The Cellular Organization of Natural Behavior
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The primary lines of evidence used by the odor plume advocates are
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1581:"Neural basis of forward flight control and landing in honeybees"
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Muth, Felicity; Papaj, Daniel; Leonard, Anne (1 September 2015).
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Carew, Thomas J. (2000). "9. Associative Learning in Honeybees".
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56:, or European honey bee. Color learning has also been studied in
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those unique traits if the reward, the pollen, is great enough.
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Giurfa, Martin; Zhang, Shaowu; Jenett, Arnim; Menzel, Randolf;
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Muth, Felicity; Papaj, Daniel; Leonard, Anne (January 2016).
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1727:"The concepts of 'sameness' and 'difference' in an insect"
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634:"Visual working memory in decision making by honey bees"
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Certain morphologies distinguished the receptor types.
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1148:(23). Kutztown University of Pennsylvania: 3737–3743.
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448:(see the work of F.C. Dyer though). Other species of
1781:"The Sensory Basis of the Honeybee's Dance Language"
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Ibbotson, M.R.; Hung, Y.; Meffin, H.; N. Boeddeker;
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1686:"The Elusive Honey Bee Dance "Language" Hypothesis"
454:use variants on the same theme, and other types of
119:take the same form in honey bees as they do in the
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332:It has long been known that successfully foraging
324:Figure-Eight-Shaped waggle dance of the honeybee (
1779:Kirchner, Wolfgang H.; Towne, William F. (2000).
1566:(On the idiothetic hypothesis) O'Dea, JD (2000) "
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23:Bees learn and communicate in a variety of ways.
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425:Odor learning in bees is usually tested by the
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1116:"Experiments on Directing Bee Flight by Odors"
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2769:Association for the Study of Animal Behaviour
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155:. Beginning in the early 1900s, scientists
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497:play an important part in how a honey bee
361:The Dance Language and Orientation of Bees
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1537:(17). Society for Science &: 271.
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1255:Williams, Caroline (18 September 2009).
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949:The Honey Bee Dance Language Controversy
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478:For more background on this topic, see
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16:Cognitive and sensory processes in bees
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1802:"Pleasure chemical controls bee dance"
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1133:Towne, William F. (1 December 2008).
1132:
963:. Cambridge, MA: Harvard Univ. Press.
738:
705:
365:Nobel Prize in Physiology or Medicine
888:
473:
184:Color learning rates and preferences
1684:Wenner, Adrian M. (November 2002).
1440:Journal of Comparative Physiology A
1257:"Rethinking the bee's waggle dance"
975:Behavioral Ecology and Sociobiology
533:suggest that honey bees may have a
136:A 2005 three-part study tested the
42:, that is the insects in the seven
13:
1561:
1347:
1295:
309:"waggle dance" or "dance language"
66:are sensitive to odors (including
14:
2880:
1923:Subspecies, Breeds and Phenotypes
1800:Hayes, Jacqui (16 January 2007).
891:Journal of the History of Biology
628:Zhang, S; Bock F; Si A; Tautz J;
70:), tastes, and colors, including
2833:
2832:
916:
493:indicates that so called primer
350:described this behaviour in his
281:
1469:
1417:
1408:
1399:
1276:
1248:
1234:
1183:
1142:Journal of Experimental Biology
1126:
1108:
1057:
1044:
1001:
966:
953:
942:
924:
461:
363:and in 1973 he was awarded the
208:
111:, and many of the phenomena of
2478:Bee learning and communication
1909:Bee learning and communication
1677:Johnson, R. N. et al. (2002)
882:
847:
814:
763:
621:
458:use other methods altogether.
28:Bee learning and communication
1:
868:10.1016/j.anbehav.2014.12.024
840:10.1016/j.anbehav.2015.09.029
615:
397:
2427:
1283:Publications of Fred C. Dyer
546:Neurobiology of color vision
524:
480:List of honey bee pheromones
243:Color learning in bumblebees
7:
2053:Apis mellifera sinisxinyuan
1289:September 14, 2006, at the
573:
196:Honey bee collecting pollen
145:Color learning in honeybees
94:
10:
2887:
2864:Western honey bee behavior
1690:Journal of Insect Behavior
1657:Queensland Brain Institute
1637:Queensland Brain Institute
1607:10.1038/s41598-017-14954-0
477:
427:proboscis extension reflex
285:
38:processes in all kinds of
2828:
2782:
2761:
2640:
2535:Evolutionary neuroscience
2435:
2387:Honeybee Discovery Center
2364:
2306:
2298:Diseases of the honey bee
2285:
2237:
2146:
2095:
2032:Apis mellifera scutellata
1922:
1886:
1855:
1850:types and characteristics
1242:"Why do honeybees dance?"
903:10.1007/s10739-005-0552-1
491:Michigan State University
342:upon their return to the
131:
2488:Behavioral endocrinology
2365:Museums and insectariums
2277:Colony collapse disorder
2252:Varroa sensitive hygiene
2039:Apis mellifera siciliana
2018:Apis mellifera monticola
2004:Apis mellifera pomonella
1990:Apis mellifera artemisia
1983:Apis mellifera adansonii
1667:"Honeybee Communication"
1661:University of Queensland
230:Timing in color learning
107:Honey bees are adept at
2683:Irenäus Eibl-Eibesfeldt
2463:Animal sexual behaviour
2171:Horizontal top-bar hive
2081:Apis mellifera unicolor
2060:Apis mellifera sossimai
2046:Apis mellifera simensis
1702:10.1023/A:1021131725124
1503:10.1073/pnas.0407652101
1272:(subscription required)
659:10.1073/pnas.0501440102
580:Bumblebee communication
288:Bumblebee communication
2622:Tool use by non-humans
2575:Philosophical ethology
2520:Comparative psychology
2468:Animal welfare science
2238:Parasites and diseases
2074:Apis mellifera taurica
2067:Apis mellifera syriaca
2025:Apis mellifera remipes
1997:Apis mellifera litorea
1723:Srinivasan, Mandyam V.
1265:(2726). Archived from
959:von Frisch, K. (1967)
789:10.1098/rsbl.2015.0628
710:. Sinauer Associates.
555:
466:The exchange of food,
329:
197:
173:
117:classical conditioning
104:
24:
1376:10.1093/chemse/bjq057
1324:10.1051/apido:2006001
553:
367:for his discoveries.
323:
195:
170:
102:
91:" and in other ways.
22:
2859:Animal communication
2728:William Homan Thorpe
2493:Behavioural genetics
2453:Animal consciousness
2448:Animal communication
2377:Bee Museum of Rhodes
2293:Topics in beekeeping
1899:Honey bee life cycle
932:"HISTORIA ANIMALIUM"
163:Color discrimination
109:associative learning
80:operant conditioning
2483:Behavioural ecology
2372:Malacca Bee Gallery
2272:Deformed wing virus
2163:BS National Beehive
2011:Apis mellifera meda
1945:Carniolan honey bee
1812:on 19 January 2007.
1792:Scientific American
1743:2001Natur.410..930G
1716:on 3 December 2003.
1673:on 22 January 2016.
1627:– via Nature.
1599:2017NatSR...714591I
1579:(6 November 2017).
1494:2004PNAS..10117559L
1212:10.1038/nature03526
1204:2005Natur.435..205R
1122:. 24 February 2016.
1086:10.1038/nature01127
1078:2002Natur.419..920S
1022:1989NW.....76..277M
1010:Naturwissenschaften
938:. 6 September 2008.
650:2005PNAS..102.5250Z
595:Trained hymenoptera
316:Dance communication
2812:Behavioral Ecology
2733:Nikolaas Tinbergen
2525:Emotion in animals
2503:Cognitive ethology
2267:American foulbrood
1586:Scientific Reports
1452:10.1007/bf00625437
1154:10.1242/jeb.003640
1030:10.1007/BF00368642
987:10.1007/bf00175094
556:
446:Western honey bees
383:Apis andreniformis
352:Historia Animalium
334:Western honey bees
330:
265:when foraging for
198:
174:
105:
25:
2846:
2845:
2738:Jakob von Uexküll
2508:Comfort behaviour
2395:
2394:
2257:Small hive beetle
2246:Varroa destructor
1970:Western honey bee
1965:Russian honey bee
1960:Maltese honey bee
1950:European dark bee
1873:Laying worker bee
1737:(6831): 930–933.
1725:(19 April 2001).
1198:(7039): 205–207.
1072:(6910): 920–922.
717:978-0-87893-084-5
632:(April 5, 2005).
554:Western honey bee
474:Primer pheromones
2876:
2836:
2835:
2798:Animal Cognition
2791:Animal Behaviour
2743:Wolfgang Wickler
2443:Animal cognition
2422:
2415:
2408:
2399:
2398:
2123:Honey extraction
1841:
1834:
1827:
1818:
1817:
1813:
1808:. Archived from
1788:
1770:
1751:10.1038/35073582
1717:
1712:. Archived from
1674:
1669:. Archived from
1655:'s group at the
1650:
1645:
1643:
1628:
1618:
1593:(14591): 14591.
1577:M. V. Srinivasan
1554:
1525:
1515:
1505:
1488:(50): 17559–64.
1464:
1463:
1435:
1424:
1421:
1415:
1412:
1406:
1403:
1397:
1396:
1378:
1354:
1345:
1344:
1326:
1302:
1293:
1280:
1274:
1273:
1270:
1252:
1246:
1245:
1238:
1232:
1231:
1187:
1181:
1180:
1178:
1176:
1139:
1130:
1124:
1123:
1112:
1106:
1105:
1061:
1055:
1052:(Apis mellifera)
1048:
1042:
1041:
1005:
999:
998:
970:
964:
957:
951:
946:
940:
939:
928:
922:
921:
920:
914:
886:
880:
879:
856:Animal Behaviour
851:
845:
844:
842:
827:Animal Behaviour
818:
812:
811:
801:
791:
767:
761:
758:
752:
749:
736:
733:
722:
721:
703:
682:
681:
671:
661:
625:
514:Other bees like
254:Bombus impatiens
214:rewarded color.
2886:
2885:
2879:
2878:
2877:
2875:
2874:
2873:
2849:
2848:
2847:
2842:
2824:
2778:
2757:
2753:Solly Zuckerman
2693:Karl von Frisch
2678:Richard Dawkins
2663:John B. Calhoun
2648:Patrick Bateson
2636:
2570:Pain in animals
2431:
2426:
2396:
2391:
2360:
2308:
2302:
2281:
2233:
2219:Honey extractor
2175:Langstroth hive
2142:
2091:
1930:Africanized bee
1918:
1882:
1851:
1845:
1806:COSMOS magazine
1665:
1653:M.V. Srinivasan
1641:
1639:
1631:
1564:
1562:Further reading
1543:10.2307/3967590
1472:
1467:
1436:
1427:
1422:
1418:
1413:
1409:
1404:
1400:
1363:Chemical Senses
1355:
1348:
1303:
1296:
1291:Wayback Machine
1281:
1277:
1271:
1253:
1249:
1240:
1239:
1235:
1188:
1184:
1174:
1172:
1137:
1131:
1127:
1114:
1113:
1109:
1062:
1058:
1049:
1045:
1006:
1002:
971:
967:
958:
954:
947:
943:
930:
929:
925:
915:
887:
883:
852:
848:
819:
815:
776:Biology Letters
768:
764:
759:
755:
750:
739:
734:
725:
718:
704:
685:
626:
622:
618:
576:
548:
529:Experiments by
527:
517:Trigona corvina
483:
476:
464:
431:solar ephemeris
400:
356:Karl von Frisch
318:
290:
284:
251:, specifically
245:
232:
211:
186:
165:
157:Karl von Frisch
147:
134:
97:
17:
12:
11:
5:
2884:
2883:
2872:
2871:
2866:
2861:
2844:
2843:
2841:
2840:
2829:
2826:
2825:
2823:
2822:
2815:
2808:
2805:Animal Welfare
2801:
2794:
2786:
2784:
2780:
2779:
2777:
2776:
2771:
2765:
2763:
2759:
2758:
2756:
2755:
2750:
2745:
2740:
2735:
2730:
2725:
2720:
2718:Desmond Morris
2715:
2710:
2705:
2700:
2695:
2690:
2685:
2680:
2675:
2673:Marian Dawkins
2670:
2668:Charles Darwin
2665:
2660:
2655:
2650:
2644:
2642:
2638:
2637:
2635:
2634:
2629:
2624:
2619:
2614:
2613:
2612:
2607:
2602:
2597:
2587:
2582:
2577:
2572:
2567:
2562:
2557:
2552:
2550:Human ethology
2547:
2542:
2537:
2532:
2527:
2522:
2517:
2516:
2515:
2505:
2500:
2495:
2490:
2485:
2480:
2475:
2470:
2465:
2460:
2458:Animal culture
2455:
2450:
2445:
2439:
2437:
2433:
2432:
2425:
2424:
2417:
2410:
2402:
2393:
2392:
2390:
2389:
2384:
2379:
2374:
2368:
2366:
2362:
2361:
2359:
2358:
2353:
2351:United Kingdom
2348:
2343:
2338:
2333:
2328:
2323:
2318:
2312:
2310:
2304:
2303:
2301:
2300:
2295:
2289:
2287:
2283:
2282:
2280:
2279:
2274:
2269:
2264:
2259:
2254:
2249:
2241:
2239:
2235:
2234:
2232:
2231:
2226:
2224:Queen excluder
2221:
2216:
2211:
2206:
2201:
2199:Wax foundation
2196:
2191:
2186:
2183:Stewarton hive
2156:
2150:
2148:
2144:
2143:
2141:
2140:
2135:
2130:
2125:
2120:
2115:
2110:
2105:
2099:
2097:
2093:
2092:
2090:
2089:
2087:Honey bee race
2084:
2077:
2070:
2063:
2056:
2049:
2042:
2035:
2028:
2021:
2014:
2007:
2000:
1993:
1986:
1979:
1976:Apis laboriosa
1972:
1967:
1962:
1957:
1952:
1947:
1942:
1940:Cape honey bee
1937:
1932:
1926:
1924:
1920:
1919:
1917:
1916:
1911:
1906:
1901:
1896:
1890:
1888:
1884:
1883:
1881:
1880:
1875:
1870:
1865:
1859:
1857:
1853:
1852:
1844:
1843:
1836:
1829:
1821:
1815:
1814:
1797:
1776:
1718:
1696:(6): 859–878.
1681:
1675:
1663:
1629:
1563:
1560:
1559:
1558:
1555:
1526:
1471:
1468:
1466:
1465:
1425:
1416:
1407:
1398:
1369:(7): 593–601.
1346:
1317:(3): 366–375.
1294:
1275:
1269:on 2009-09-23.
1247:
1233:
1182:
1125:
1107:
1056:
1043:
1016:(6): 277–280.
1000:
981:(4): 227–233.
965:
952:
941:
923:
897:(3): 535–570.
881:
846:
813:
762:
753:
737:
723:
716:
683:
644:(14): 5250–5.
619:
617:
614:
613:
612:
607:
602:
597:
592:
590:Grooming dance
587:
582:
575:
572:
547:
544:
526:
523:
475:
472:
463:
460:
419:
418:
414:
399:
396:
394:inconsistent.
326:Apis mellifera
317:
314:
283:
280:
244:
241:
231:
228:
210:
207:
185:
182:
164:
161:
152:Apis mellifera
146:
143:
138:working memory
133:
130:
96:
93:
85:cognitive maps
53:Apis mellifera
46:making up the
15:
9:
6:
4:
3:
2:
2882:
2881:
2870:
2867:
2865:
2862:
2860:
2857:
2856:
2854:
2839:
2831:
2830:
2827:
2821:
2820:
2816:
2814:
2813:
2809:
2807:
2806:
2802:
2800:
2799:
2795:
2793:
2792:
2788:
2787:
2785:
2781:
2775:
2772:
2770:
2767:
2766:
2764:
2760:
2754:
2751:
2749:
2746:
2744:
2741:
2739:
2736:
2734:
2731:
2729:
2726:
2724:
2723:Thomas Sebeok
2721:
2719:
2716:
2714:
2713:Konrad Lorenz
2711:
2709:
2708:Julian Huxley
2706:
2704:
2703:Heini Hediger
2701:
2699:
2696:
2694:
2691:
2689:
2686:
2684:
2681:
2679:
2676:
2674:
2671:
2669:
2666:
2664:
2661:
2659:
2656:
2654:
2651:
2649:
2646:
2645:
2643:
2639:
2633:
2632:Zoomusicology
2630:
2628:
2625:
2623:
2620:
2618:
2615:
2611:
2608:
2606:
2603:
2601:
2598:
2596:
2593:
2592:
2591:
2588:
2586:
2583:
2581:
2578:
2576:
2573:
2571:
2568:
2566:
2565:Neuroethology
2563:
2561:
2558:
2556:
2553:
2551:
2548:
2546:
2543:
2541:
2538:
2536:
2533:
2531:
2528:
2526:
2523:
2521:
2518:
2514:
2511:
2510:
2509:
2506:
2504:
2501:
2499:
2496:
2494:
2491:
2489:
2486:
2484:
2481:
2479:
2476:
2474:
2473:Anthrozoology
2471:
2469:
2466:
2464:
2461:
2459:
2456:
2454:
2451:
2449:
2446:
2444:
2441:
2440:
2438:
2434:
2430:
2423:
2418:
2416:
2411:
2409:
2404:
2403:
2400:
2388:
2385:
2383:
2380:
2378:
2375:
2373:
2370:
2369:
2367:
2363:
2357:
2356:United States
2354:
2352:
2349:
2347:
2344:
2342:
2339:
2337:
2334:
2332:
2329:
2327:
2324:
2322:
2319:
2317:
2314:
2313:
2311:
2305:
2299:
2296:
2294:
2291:
2290:
2288:
2284:
2278:
2275:
2273:
2270:
2268:
2265:
2263:
2260:
2258:
2255:
2253:
2250:
2248:
2247:
2243:
2242:
2240:
2236:
2230:
2227:
2225:
2222:
2220:
2217:
2215:
2212:
2210:
2207:
2205:
2202:
2200:
2197:
2195:
2192:
2190:
2187:
2184:
2180:
2176:
2172:
2168:
2164:
2160:
2157:
2155:
2152:
2151:
2149:
2145:
2139:
2136:
2134:
2131:
2129:
2126:
2124:
2121:
2119:
2116:
2114:
2111:
2109:
2106:
2104:
2101:
2100:
2098:
2094:
2088:
2085:
2083:
2082:
2078:
2076:
2075:
2071:
2069:
2068:
2064:
2062:
2061:
2057:
2055:
2054:
2050:
2048:
2047:
2043:
2041:
2040:
2036:
2034:
2033:
2029:
2027:
2026:
2022:
2020:
2019:
2015:
2013:
2012:
2008:
2006:
2005:
2001:
1999:
1998:
1994:
1992:
1991:
1987:
1985:
1984:
1980:
1978:
1977:
1973:
1971:
1968:
1966:
1963:
1961:
1958:
1956:
1953:
1951:
1948:
1946:
1943:
1941:
1938:
1936:
1933:
1931:
1928:
1927:
1925:
1921:
1915:
1912:
1910:
1907:
1905:
1902:
1900:
1897:
1895:
1892:
1891:
1889:
1885:
1879:
1876:
1874:
1871:
1869:
1866:
1864:
1861:
1860:
1858:
1854:
1849:
1842:
1837:
1835:
1830:
1828:
1823:
1822:
1819:
1811:
1807:
1803:
1798:
1795:
1793:
1786:
1782:
1777:
1774:
1768:
1764:
1760:
1756:
1752:
1748:
1744:
1740:
1736:
1732:
1728:
1724:
1719:
1715:
1711:
1707:
1703:
1699:
1695:
1691:
1687:
1682:
1680:
1676:
1672:
1668:
1664:
1662:
1658:
1654:
1649:
1638:
1634:
1630:
1626:
1622:
1617:
1612:
1608:
1604:
1600:
1596:
1592:
1588:
1587:
1582:
1578:
1573:
1572:
1571:
1569:
1556:
1552:
1548:
1544:
1540:
1536:
1532:
1527:
1523:
1519:
1514:
1509:
1504:
1499:
1495:
1491:
1487:
1483:
1479:
1474:
1473:
1461:
1457:
1453:
1449:
1445:
1441:
1434:
1432:
1430:
1420:
1411:
1402:
1394:
1390:
1386:
1382:
1377:
1372:
1368:
1364:
1360:
1353:
1351:
1342:
1338:
1334:
1330:
1325:
1320:
1316:
1312:
1308:
1301:
1299:
1292:
1288:
1284:
1279:
1268:
1264:
1263:
1262:New Scientist
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2698:Jane Goodall
2658:Donald Broom
2627:Zoosemiotics
2580:Sociobiology
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2382:Honey Museum
2309:by countries
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1935:Buckfast bee
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1651:The work of
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605:Waggle dance
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508:ethyl oleate
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123:. Efficient
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89:waggle dance
62:
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2688:Dian Fossey
2653:Marc Bekoff
2641:Ethologists
2341:New Zealand
2307:Beekeeping
2194:Honey super
2138:Royal jelly
2096:Cultivation
1955:Italian bee
1794:, June 1994
862:: 191–199.
585:Eusociality
531:James Gould
377:Apis florea
121:vertebrates
72:ultraviolet
2853:Categories
2590:Structures
2585:Stereotypy
2229:Jenter kit
2214:Bee smoker
2209:Queen clip
2189:Hive frame
2103:Beekeeping
1894:Bee colony
1887:Life cycle
1868:Worker bee
1856:Bee castes
1785:Apiculture
1311:Apidologie
1175:3 November
833:: 93–100.
616:References
495:pheromones
439:idiothetic
398:Odor plume
372:honey bees
336:perform a
286:See also:
249:bumblebees
68:pheromones
64:Honey bees
58:bumblebees
2819:Behaviour
2762:Societies
2600:Honeycomb
2316:Australia
2204:Hive tool
2167:Flow Hive
2147:Equipment
2128:Honeycomb
1878:Drone bee
1863:Queen bee
1848:Honey bee
1789:(Also in
1771:(Also on
1446:: 11–13.
1385:0379-864X
1333:0044-8435
911:120244044
525:Cognition
348:Aristotle
261:and blue
76:classical
32:cognitive
30:includes
2838:Category
2783:Journals
2610:Instinct
2560:Learning
2555:Instinct
2530:Ethogram
2513:Grooming
2436:Branches
2429:Ethology
2133:Propolis
2108:Apiology
1914:Swarming
1759:11309617
1710:41167065
1625:29109404
1522:15572455
1393:20534775
1287:Archived
1220:15889092
1170:11997761
1162:19011214
1094:12410309
1038:22396565
995:23263610
876:53200007
808:26423070
678:15795382
574:See also
504:foragers
263:corollas
125:foraging
95:Learning
44:families
2540:Feeding
2346:Ukraine
2331:Ireland
2321:Hungary
2262:Waxworm
2159:Beehive
2113:Beeswax
1767:1753638
1739:Bibcode
1642:10 July
1616:5673959
1595:Bibcode
1551:3967590
1490:Bibcode
1460:2676212
1341:4873234
1228:4413962
1200:Bibcode
1102:4431247
1074:Bibcode
1018:Bibcode
799:4614436
646:Bibcode
388:evolved
300:and/or
259:anthers
172:vision.
113:operant
36:sensory
2154:Apiary
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669:555688
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560:retina
499:colony
302:pollen
298:nectar
271:anther
267:pollen
132:Memory
2617:Swarm
2545:Hover
2498:Breed
2336:Nepal
2326:India
2286:Lists
2118:Honey
1904:Brood
1763:S2CID
1706:S2CID
1547:JSTOR
1456:S2CID
1337:S2CID
1224:S2CID
1166:S2CID
1138:(PDF)
1098:S2CID
1034:S2CID
991:S2CID
907:S2CID
872:S2CID
782:(9).
275:petal
48:clade
2869:Bees
2605:Nest
2595:Hive
1755:PMID
1644:2020
1621:PMID
1518:PMID
1389:PMID
1381:ISSN
1329:ISSN
1216:PMID
1177:2022
1158:PMID
1090:PMID
804:PMID
712:ISBN
674:PMID
456:bees
451:Apis
380:and
344:hive
294:hive
273:and
115:and
78:and
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2179:Nuc
1747:doi
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1698:doi
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1603:doi
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