How does the honeybee waggle dance work?

A forager that has found good food returns to the hive and dances on the wax comb. The dance tells other workers how far away the food is and in which direction to fly. Recruits follow the dancer, then leave the hive and travel to the place she indicates. The dance is also used by scouts to recruit bees to a new nest site during swarming. NC State Extension describes it as the only known abstract language in nature other than human language. [1]

According to NC State Extension, the dance changes form with distance. Food closer than 50 meters brings a round dance. Food at 50 to 150 meters brings a crescent-shaped sickle dance. Food beyond 150 meters brings the waggle dance, the only one of the three that carries both distance and direction. Kew similarly describes the round dance as signaling a rich source nearby and the waggle dance as telling nest-mates how far to go and in what direction. [1] [2]

In the waggle dance the bee runs straight ahead while wagging her abdomen from side to side and buzzing with her wings. She then circles back to the start and repeats the run, alternating sides to trace a figure eight. The duration of the straight waggle run is the simplest and most reliable indicator of distance, and longer runs mean farther food in a roughly linear relationship. NC State gives the example that a 2.5-second run signals a source about 2,625 meters away. [1]

Direction is given relative to the sun. The hive comb hangs vertically, and straight up stands for the direction of the sun. The angle of the waggle run from vertical matches the angle of the food from the sun's direction. If the food lies 60 degrees left of the sun, the run is 60 degrees left of vertical. If the food lies directly away from the sun, the run points straight down. NC State describes this as transposing the solar angle into a gravitational angle. [1] [3] [4]

Because the sun moves, the dance for the same food source changes through the day. A source due east is danced roughly straight up in the morning and roughly straight down in the late afternoon. NC State notes that reading the dance therefore requires the run's angle plus the sun's compass direction, which depends on place, date and time. [1]

NOVA reports that a dancer signals how rich a source is by how long or vigorously she dances, and that she shares the flowers' odor, which nearby bees sample with their antennae. [3]

A recent study in Behavioral Ecology and Sociobiology used video of dances and harmonic radar tracking of recruits' flights. It found that recruits search more precisely than the spread of the dancers' encoded vectors would predict. The authors suggest this could be explained by recruits averaging over about eight waggle runs, though they say other factors may be involved. [5]

Some scientists, including Adrian Wenner, argued that the dance's existence does not show it communicates location. They held that floral odors on the forager's body are the main cues recruits use, and experiments confirmed that odor matters. NC State notes that von Frisch himself once held this view before changing his mind in favor of the dance language. [1]

NC State says the most commonly accepted view lies between the extremes: recruits go to the area the dance indicates, then home in on the flowers using odor. A robotic bee that performed the dance recruited novices to specific locations, but only when it carried an odor cue. The source concludes that bees clearly use the dance's distance and direction information. [1]