The waggle dance

Ask a non-beekeeper what they know about bees and you’ll probably get answers that involve honey or stings.

Press them a little bit more about what they know about other than honey and stings and some will mention the ‘waggle dance’. 

Karl von Frisch

That the waggle dance is such a well-known feature of honey bee biology is probably explained by two (related) things; it involves a relatively complex form of communication in a non-human animal, and because Karl von Frisch – the scientist who decoded the waggle dance – received the Nobel Prize 1 for his studies in 1973.

Von Frisch did not discover the waggle dance. Nicholas Unhoch described the dance at least a century before Von Frisch decoded the movement, and Ernst Spitzner – 35 years earlier still – observed dancing bees and suggested they were communicating odours of food resources available in the environment.

Inevitably, Aristotle also made a contribution. He described flower constancy 2 and suggested that foragers could communicate this to other bees.

Language and communication are important. The development of language in early humans almost certainly contributed to the evolution of our culture, society and technology. Communication in non-human animals, from the chirping of grasshoppers to the singing of whales, is of interest to scientists and non-scientists alike.

It is therefore unsurprising that the ‘dance language’ of honey bees is also of great interest. Although not a ‘language’ in the true sense of the word, Von Frisch described the symbolic language of bees as “the most astounding example of non-primate communication that we know” over 50 years ago. This still applies.

The waggle dance

The waggle dance usually takes place in the dark on the vertical face of a comb in the brood nest, usually close to the nest entrance. The dance is performed by a successful forager i.e. one that has located a good source of pollen, nectar or water, and provides information on the presence, the quality, identity, direction and distance of the source, so enabling nest-mates to find and exploit it.

The dance consists of two phases:

  1. The figure of eight-shaped ‘return phase’ in which the bee circles back, alternately clockwise and anticlockwise, to the start of …
  2. The ‘waggle phase’, which is a short linear run in which the dancer vigorously waggles her abdomen from side to side.

The direction of the food source is indicated by the angle of the waggle phase from gravity i.e. a vertical line down the face of the comb. This angle (α in the figure below) indicates the bearing from the direction of the sun that needs to be followed to reach the food source. 

For example, if the dancer performs a waggle phase vertically down the face of the comb, the food source must be opposite the current position of the sun.

The waggle dance

The distance information is conveyed by the duration of the waggle phase. The longer this run is, the more distant the source. A run of 1 second duration indicates the food source is about 1 kilometre away.

The quality of the food source is indicated by the vigour of the waggling during the waggle phase and the speed with which the return phase is conducted. 

Surely it can’t be that simple?

Yes, it can.

What I’ve described above allows you to interpret the waggle dance sufficiently well to know where your bees are foraging.

Next time you lift a frame from a hive and see a dancing bee, circling around in a little cleared ‘dance floor’ surrounded by a group of attentive workers, try and decode the dance.

Remember that the dance is performed with relation to gravity in the darkened hive. You’re looking to identify the angle from a vertical line up the face of the brood comb to determine the direction from the sun.

Time a few waggle phases (one elephant, two elephants etc.) and you’ll know how far away the food source is.

Really, it’s that simple?

Of course not 😉

The waggle dance was decoded more than half a century ago and remains an active subject for researchers interested in animal communication.

What you’ll miss in your observations is an indication of the type of nectar or pollen resource that the dancing bee is communicating. The dancing worker carries the odour of the food source and may also regurgitate nectar, presumably helping those ‘watching’ (remember, it’s dark … nothing to see here!) determine the type of resource to look for when they leave the hive.

You will also be unable to detect the pulsed thoracic vibrations that the dancing bee produces. These are also indicators of the quality of the food source; better (e.g. higher sucrose content) resources elicit increased pulse duration, velocity amplitude and duty cycle, though the number of pulses is related to the duration of the waggle phase, and so is another potential indicator of distance.

Inevitably, there are also pheromones involved.

There always are 😉

The dancing bee produces two alkanes, tricosane and pentacosane, and two alkenes, Z-(9)-tricosene and Z-(9)-pentacosene. These appear to stimulate foraging activity 3.

But it’s cloudy … or rain stops play … or nighttime

What happens to dancing bees if foraging is interrupted, for example by poor weather or night? 

The dancing bee continues to change the angle of the waggle phase as the sun moves across the sky. This means that a dancing bee will correctly signal the direction to the food source, even if they have not left the hive for several hours.

During their initial orientation flights they learn the sun’s azimuth as a function of the time of day, and use this to compensate for the sun’s time-dependent movement.

Some bees even dance during the night, in which case the watching workers must presumably make their own compensations for the time that has elapsed since the dance 4.

And what happens if the sun is obscured … by clouds, or buildings or dense woodland? How can those directions be followed?

Under these circumstances the foraging bee detects the position of the sun by the pattern of polarised light in the sky. 

Scout bees

The waggle dance is also performed by scout bees on the surface of a bivouacked swarm. In this instance it is used to communicate the quality, direction and distance of a new potential nest site. 

Swarm of bees

Swarm of bees

The intended audience in this instance are other scout bees, rather than the general forager population 5. These scouts use a quorum decision making process to determine the ‘best’ nest site in the area to which the bivouacked swarm eventually relocates.

The shape of the bivouac often lacks a true vertical surface. However, since it’s in the open the dancing bees can orientate the waggle run directly with relation to the sun’s direction, rather than to gravity.

Under experimental conditions the dancing bee can communicate the presence and quality of a food source on a horizontal comb, but – with no reference to gravity – all directional information is lost 6.

The round dance

The duration of the waggle phase is related to the distance from the nest to the food source. Therefore the recognisable waggle dance tends to get difficult to interpret for sources very close to the nest.

It used to be thought that there was a distinct directionless dance (the ’round dance’) for these nearby i.e. 10-40 metres, food sources. However, more recent study 7 suggests that dancers were able to convey both distance and direction information irrespective of the separation of nest and food source. This indicates that bees have just one type of dance for forager recruitment, the waggle dance.

Do all bees communicate using a waggle dance?

There are a very large number of bee species. In the UK alone there are 270 species, 250 of which are solitary.

There’s a clue.

Solitary bees are like me at a disco … they have no one to dance with 🙁

I’ll cut to the chase to help you erase that vision.

The only bees that use the waggle dance are honey bees. These all belong to the genus Apis.

They include our honey bee, the western honey bee (Apis mellifera), together with a further seven species:

  1. Black dwarf honey bee (Apis andreniformis)
  2. Red dwarf honey bee (Apis florea)
  3. Giant honey bee (Apis dorsata)
  4. Himalayan giant honey bee (Apis laboriosa
  5. Eastern honey bee (Apis cerana)
  6. Koschevnikov’s honey bee (Apis koschevnikovi)
  7. Philippine honey bee (Apis nigrocincta)

Dancing and evolution

Dwarf honey bees nest in the open on a branch and dance on the horizontal surface of the nest. The waggle run is orientated ‘towards’ the food source. Apis dorsata is also an open-nesting bee, but forms large vertically-hanging combs. It dances relative to gravity, and indicates the direction by the angle of the waggle run in the same way that A. mellifera does.

The cavity nesting bees, A. cerana, A. mellifera, A. koschevnikovi, and A. nigrocinta produce the most developed form of the dance.

The dances of A. mellifera and A. cerana are sufficiently similar that they can follow and decode the dance of the other.

The complexity of the nest site and the waggle dance reflects the evolution of these bee species. The earliest to evolve (i.e. the most primitive), A. andreniformis and florea, have the simplest nests and the most basic waggle dance. In contrast, the cavity nesting species evolved most recently, form the most complex brood nests and have the most derived waggle dance.

When and why did the waggle dance evolve?

Assuming that the waggle dance did not independently evolve (there’s no evidence it did, and ample evidence due to its similarity between species that it evolved only once) it must have first appeared at least 20 million years ago, when extant honey bee species diverged during the early Miocene.

The ‘why’ it evolved is a bit more difficult to address.

Behavioural changes often arise in response to the environment in which a species evolves.

Bipedalism in non-human primates (like the australopithecines) is hypothesised to have evolved in part due to a reduction in forest cover and the increase in savannah. Apes had to walk further between clumps of trees and bipedalism offered greater travel efficiency.

Perhaps the waggle dance evolved to exploit a particular type or distribution of food reserves?

In this regard it is interesting that the ‘benefit’ of waggle dance communication varies through the season.

If you turn a hive on its side the combs are horizontal 8. Under these conditions the dancing bees can communicate the presence and quality of a food source. However, they cannot communicate its location (either direction or distance).

No directional or distance information is now available

In landmark studies Sherman and Visscher 9 showed that, at certain periods during the season, the absence of this positional information did not affect the weight gain by the hive i.e. the foraging efficiency of the colony.

They concluded that during these periods forage must be sufficiently abundant that simply stimulating foraging was sufficient. Remember those alkanes and alkenes produced by dancing bees that do exactly that?

Tropical habitats

This observation, and some elegant experimental and modelling studies, suggest that dancing is beneficial when food resources are: 

  • sparsely distributed – therefore difficult (and energetically unfavourable) to find by individual scouting
  • clustered or short-lived resources – when it’s gone, it’s gone
  • distributed with high species richness – if there’s a huge range of flowers, which are the most energetically rewarding (sugar-rich) to collect nectar from?

One of the experimental studies that contributed to these conclusions (though there’s still controversy in this area) was the demonstration that waggle dancing was beneficial in a tropical habitat, but not in two temperate habitats. This makes sense, as food resources have different spatiotemporal distribution in these habitats. Tropical habitats are characterised by clustered and short-lived resources.

Therefore the suggestion is that the waggle dance of Apis species evolved, presumable early in the speciation of the genus, in a tropical region where food resources were patchily distributed, available for only limited period and present alongside a wide variety of other (less good) choices.

For example, like individual trees flowering in a forest …

Finally, it’s worth noting that there is evidence that bees that dance are able to successfully exploit food resources further away than would otherwise be expected from their body size.

This also makes sense.

It’s much less risky flying off over the horizon if you know there’s something to collect once you get there 10.


Notes

If you arrived here from my Twitter feed (@The_Apiarist) you’ll have seen the tweet started with the words “Dance like nobody’s watching”, words that are often attributed to Mark Twain. 

The full quote is something like “Dance like nobody’s watching; love like you’ve never been hurt. Sing like nobody’s listening; live like it’s heaven on earth”.

Pretty sound advice.

But it’s not by Mark Twain. It’s actually from a country music song by Susanna Clark and Richard Leigh. This was first released on the Don Williams album Traces in 1987. So only about 90 years out 😉 

Footnotes

  1. Equally shared with Nikolaas Tinbergen and Konrad Lorenz, for their discoveries concerning organization and elicitation of individual and social behaviour patterns”.
  2. The tendency for a bee to visit a single type of flower, ignoring others in the same environment.
  3. Thom et al., 2007 The scent of the waggle dance. PLoS Biology 5:e228.
  4. I’m not sure if this is formally proven … it might be that the watching workers ignore anything but the most recently ‘watched’ dance.
  5. How do regular foragers know not to ‘watch’ the dance?
  6. This always makes me wonder whether the bees we see dancing on a frame removed from the hive are doing so with relation to the position of the comb in the hive, or the position of the sun when on a frame in our hands. Someone must surely know this? Whatever, it should be easy to test. Next time I find a dancing bee on a frame I’ll rotate the frame through 90° and see if the direction of the waggle run changes.
  7. See Gardner et al., (2008) Do honeybees have two discrete dances to advertise food sources? Animal Behaviour 75:1291-1300.
  8. Formally on two of the four sides, of course.
  9. Sherman & Visscher (2002) Honeybee colonies achieve fitness through dancing. Nature 419:920–922.
  10. And this is a good point to stop. It’s worth noting that waggle dancing has been decoded to map food resources up to 11 km distant … that potentially involves an 11 second waggle run, which I’ll discuss in a separately when I revisit my ageing sphere of influence post.

27 thoughts on “The waggle dance

  1. Richard Elliott

    If the dance is straight down the comb, towards gravity = fly directly AWAY from the sun, do upwards moving dances dictate the angle TOWARDS the sun?

    Is the reference sun direction the angle toward the sun at the time of day that dance happens?

    Reply
    1. David Post author

      Yes Richard … if the waggle run is directly up the face of the comb to the top bar then the food source is in the direction of the sun. The dancing bee adjusts the angle of the waggle run for the time of day she’s doing the dance … she might have discovered a field full of borage at 10 in the morning but then got trapped in the hive by a summer rainstorm. During the rain she’ll still be dancing, but the angle of the waggle run will change over time as the sun moves round.

      Cheers
      David

      Reply
        1. David Post author

          Hi Richard

          Good … though don’t think that all there is to the waggle dance 😉

          Waggle run error

          For example the waggle run isn’t perfectly aligned on every repetition. Is this inaccuracy an adaptive mechanism to make foraging at distance more efficient, or an inherent feature because dancers cannot dance any more accurately if they tried?

          Watch this space …

          Reply
  2. Dorothie Jones

    Fascinating article as usual David.
    It’s always puzzled me….How they can they ‘see’ to interpret the waggle dance if it’s pitch dark inside? do they just do it by vibration?
    Dorothie

    Reply
    1. David Post author

      Hello Dorothie

      There are still some gaps in our understanding of this part of the waggle dance. However, it’s thought that a combination of the ‘dance sounds’ and wing vibrations allow the direction of the waggle run to be determined. The number of pulsed vibrations produced is an indication of distance as indicated above.

      I’ll return to this sometime in the future as there’s other features of the waggle dance that are also interesting.

      Cheers
      David

      Reply
  3. Edward Hill

    Hi
    Any thoughts on Adrian Wenners challange to Von Karl von Frisch’s theory and why they had different views. Curious.

    Reply
    1. David Post author

      Hi Edward

      One of the reason they had different views is that they came from very different scientific backgrounds; that’s not unusual and often results in interpreting the same studies in different ways. Wenner seemed unable to accommodate the inherent (albeit low level) inaccuracy of the system, or things that might impact on the ability of the forager to find the food source after watching the dance. However, Von Frisch’s view prevailed and subsequent harmonic radar studies have supported his interpretations. I’ll return to the waggle dance in a future post to discuss some of the other features of it.

      Cheers
      David

      Reply
          1. David Post author

            Hello Jim

            I can’t find a reference to Bee-L on this page … which link? Or which factoid should be linked? Apologies, it was written best part of 3 months ago and the short-term memory is getting like my long-term memory …

            Cheers
            David

  4. Elaine Robinson

    Good revision for Module 6, David. Interesting about the pheromones involved, thanks for sharing the research reference, the books don’t refer to this, will add to my revision notes for extra brownie points!

    Reply
  5. Stan Thornton

    Similarly, when you mentioned rotating a frame and how the dance would be changed… I wonder how long it takes for the foraging bees to recognize that they need to begin a different dance pattern when the hive is moved or turned 90°? Just saw some eucalyptus blooming here in Northern California today. For my area that is very early by at least a month.

    Reply
    1. David Post author

      Hello Stan

      I don’t know the answer to that I’m afraid. On a related point, not every forager dances … think about what the hive entrance and the frames look like when there’s a really strong nectar flow on. It’s frantically busy at the hive entrance, but only a relatively small proportion of the bees do waggle dances … I also don’t know why some dance and most don’t. More reading to do 🙁

      We’re still a month or two away from spring blossom here … it’s very grey and wet, and I’ve not seen a bee for weeks. Actually, that’s not entirely true. I’ve not seen a flying bee for weeks. I had a peek at some clusters through the clear crownboard on my hives in between the showers and it’s all looking OK, but the worst of the winter is still probably to come.

      Enjoy the eucalyptus 🙂
      David

      Reply
  6. Meryl

    Thanks David, very helpful.
    From which gland do the waggle dancers produce the four named alkanes and alkenes, please?
    As not all returning foragers perform a waggle dance, do these non-dancers also produce these pheromones?

    Reply
    1. David Post author

      Hello Meryl

      I’m not sure it’s known where the alkanes/alkenes originate. Pentacosane is known to be produced by the mandibular gland (though whether this is the pentacosane produced by waggle dancers is not known). The cephalic salivary gland produces the largest amounts of hydrocarbons and esters overll.

      Non-dancing workers produce very little or none of the alkanes/alkenes listed above.

      Good luck with the exams 😉 These are the sorts of detail they often contain.

      Cheers
      David

      Reply
  7. Alex

    I wonder if the dancing foragers just find another appropriate vertical plane (dance floor) if the hive falls over? For example, one of the hive walls.

    Reply
    1. David Post author

      Hi Alex

      Possibly. However, it’s known that bees prefer to dance over empty cells rather than capped brood cells, so they’re clearly influenced by the ‘dance floor’ in some way. In addition, the dance floor is usually close to the hive entrance, so presumably this would influence their choice as well.

      Of course, in the studies looking at the loss of directional information the authors took care to ensure that the bees were dancing where they were supposed to. I actually simplified that part of the description quite a bit. There are some issues with simply reorienting the hive. Instead they provided unidirectional or diffuse lighting as the reference. They then demonstrated that the dances were truly disoriented, by confirming that foragers ‘dancing’ for a known source of forage (a feeder) generated dances in which the waggle runs were essentially randomly oriented.

      If you want more gory details have a read of the Sherman and Visscher reference I cited … in particular Figure 1.

      Cheers
      David

      Reply
      1. Alex

        Thanks. I’ll have to add that paper to my list to look up in the Moir Library when life returns to normal – i’m hoping they get copies of Nature??!?!?! I got an IBRA account for xmas so that’s keeping me occupied at the moment!!! 🙂

        Reply
        1. David Post author

          I can also recommend the American Bee Journal … very reasonable as a digital download and some good writing from people like Jamie Ellis, Randy Oliver and Pete Borst. Lots of adverts, but it’s always interesting to see what other people are buying (or being encouraged to buy). You have to be cautious as it’s (obviously) US-centric, but it’s a big country and in many places their climate is similar to some of ours … and bees are bees.

          Except those in the UK aren’t Africanized 😉

          Cheers
          David

          Reply
  8. Paul Lindstrom

    “Dance like nobody’s watching” is often quoted, but rarely is the source/author mentioned. Thanks for that. One of my daughters used it on a birthday card to me, and added – “why care if you look like a fool”. 😉

    BTW – Scout bees can also be told off if they continues to dance to promote a new site for the swarming colony that has been rejected by the majority of the other scouts (according to a lecture by Tom Seeley fairly recently). They go up and head butt the annoying colleague, sort of telling her to “shut up and sit down – we have decided on where to go”. For some reason I smiled at that (and secretly thought that I should perhaps try it in a meeting that just goes on and on). 😉

    Reply
    1. David Post author

      Hello Paul

      It’s always good to see where quotes originate.

      Interesting comment about scout bees from Tom Seeley. He’s also reported part of the evidence for a quorum decision making process is swarms that take off, split and set off in opposite directions. If that’s what they do it’s clear that some don’t get the headbutted message …

      Cheers
      David

      Reply

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