A waggle dance consists of one to 100 or more circuits, each of which consists of two phases: the waggle phase and the return phase. A worker bee's waggle dance involves running through a small figure-eight pattern: a waggle run (aka waggle phase) followed by a turn to the right to circle back to the starting point (aka return phase), another waggle run, followed by a turn and circle to the left, and so on in a regular alternation between right and left turns after waggle runs. Waggle-dancing bees produce and release two alkanes, tricosane and pentacosane, and two alkenes, Z-(9)-tricosene and Z-(9)-pentacosene, onto their abdomens and into the air.

The direction and duration of waggle runs are closely correlated with the direction and distance of the patch of flowers being advertised by the dancing bee. Flowers located directly in line with the sun are represented by waggle runs in an upward direction on the vertical combs, and any angle to the right or left of the sun is coded by a corresponding angle to the right or left of the upward direction. The distance between hive and recruitment target is encoded in the duration of the waggle runs. The farther the target, the longer the waggle phase, with a rate of increase of about 75 milliseconds per 100 meters.

Waggle dancing bees that have been in the hive for an extended time adjust the angles of their dances to accommodate the changing direction of the sun. Therefore, bees that follow the waggle run of the dance are still correctly led to the food source even though its angle relative to the sun has changed.

The consumption of ethanol by foraging bees has been shown to reduce waggle dance activity and increase occurrence of the tremble dance.

When scientists placed a deadApis melliferabee on flowers they discovered that bees performed far fewer waggle dances upon returning to their nest. This is likely to be because they associate the dead bee with the presence of a predator on the flower and so it is better for other bees to not forage there.

Though first decoded by Karl von Frisch, dancing behavior in bees had been observed and described multiple times prior. 100 years before Frisch’s discovery, Nicholas Unhoch described dancing behavior of bees as being an indulgence “in certain pleasures and jollity”. He did, however, admit ignorance as to purpose of the dancing. 35 years before that, Ernst Spitzner observed bees dancing and interpreted it as transmitting forage resource odors to other nestmates. Even Aristotle, in addition to describing flower constancy behavior, suspected that some form of communication occurred between foragers within a nest:

“On each trip the bee does not fly from a flower of one kind to a flower of another, but flies from one violet, say, to another violet, and never meddles with another flower until it has got back to the hive; on reaching the hive they throw off their load, and each bee on her return is followed by three or four companions. What it is that they gather is hard to see, and how they do it has not been observed.”

The Dance Language vs. The Waggle Dance

The dance language, as defined by von Frisch, is the information about direction, distance, and quality of a resource (such as food or nesting sites) contained within the waggle dance. Karl von Frisch named the dance language “Tanzsprache” in his native tongue.

Though von Frisch insisted on the direct connection between the dance language and the waggle dance, recent criticism holds that potential foragers need not correctly translate the dance language from the waggle dance to successfully forage.

In an experiment on the honeybeeApis mellifera, most individuals who thoroughly followed a waggle dance ignored the resource direction and location information. Instead, 93% of the foragers returned to foraging areas they had previous knowledge of.

Bees that follow a waggle dance can successfully forage without decoding the dance language information in several ways Dance follower may use olfactory information from the dancer and find either the same resource or a different one with a similar scent. Following a dance may simply trigger foraging behavior. A forager may then search randomly for resources. Following a dance may reactivate private knowledge of a resource. After reactivation, the forager may return the known resource. Using information communicated in the waggle dance is more useful to foragers when private information about resources is lacking. The use of the word “language” may lead to misrepresentations of the waggle dance. The Swiss linguist Ferdinand de Saussure proposed a system of language a sign is made up two chief components. The signifier is the physical or phonetic representation of a sign. The signified is the conceptual component. If the dance language followed the Saussurian dyadic model of semiotics, the signifier would be the waggle dance and the signified would be the location of the foraging resource. Though the dance language may or may not follow this sort of pattern, it is not considered to be a language with syntactical grammar or a set of symbols.

Efficiency and Adaptation

The waggle dance may be less efficient than once thought. Some bees observe over 50 waggle runs without successfully foraging, while others will forage successfully after observing 5 runs.

The waggle dance may be adaptive in some environments and not in others. In temperate habitats, honey bee colonies routinely perform the waggle dance, but can still successfully forage when the dance is experimentally obscured. In tropical habitats, honey bee foraging is severely impaired if waggle dancing is prevented. This is thought to be due to the patchiness of resources in tropical environment versus the homogeneity of resources in temperate environments. In the tropics, food resources can come in the form of flowering trees which are rich in nectar but sparsely scattered and bloom for short periods of time. Thus, in tropical zones information about forage location might be more valuable than in temperate zones.