Queen mating behavior as an example of basic science observation in beekeeping technology development

Janko Bozic1 and Ida Gnilsak2
1 Department of Biology, University of Ljubljana, Slovenia
2 Apicultural Museum, Radovljica, Slovenia

Ljubljana, 2000 

Contents:

Abstract
Economic background
Major nectar flow favored swarm production in the 18th century
Queen rearing as a part of artificial swarm production
Detailed description of queen mating behavior
Spread of the knowledge on queen mating behavior
Conclusion
References
Beekeeping in Slovenia
Foraging behavior of honeybees and hormones

Abstract

More then two hundred years ago beekeepers of Austrian empire county Carniola had adopted management with bees to the nectar flow of buckwheat. Beekeepers exploited buckwheat blooming in late August and early September with the swarms made and caught during summer or they simply bought swarms from other beekeepers who were specialized on swarm production. Many specialized beekeepers managed to prepare artificial swarms. They observed in detail behaviors related to swarming and reproduction. Natural scientist and physician Anton Scopoli published first printed report of queen mating outside the hive in his book Entomologia carniolica in 1763. Scopoli got most of the information about honeybee biology from beekeepers. Peter Pavel Glavar (1768, 1776), Anton Janša (1771, 1775) and Huml (1773) wrote also about queen mating outside the hive. All written resources show that queen mating and swarming were well understood among Slovenian beekeepers. They achieved this knowledge by searching of more efficient technology of artificial swarm production. Since Anton Janša was court teacher of beekeeping in Wiene and Peter Pavel Glavar had written correspondence with many beekeepers in Europe, this knowledge was spread across the Europe. A decade later queen mating behavior was also described correctly by Fr. Huber (1788).
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Economic background

More then two hundred years ago beekeepers of Austrian empire county Carniola understood well queen mating behavior and reproduction biology of honeybees which enabled them to develop several different methods of artificial swarm production. They were interested in swarm production to exploit late summer nectar flow on buckwheat.
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Major nectar flow favored swarm production in the 18th century

The most important source of honey was nectar of buckwheat (Miheliè 1970). This crop was planted at the one third of fields. Actually such a big proportion was a result of regular cycling of different crops on the fields. Each third year farmers seeded buckwheat after harvesting of regular wheat crop usually at the end of June. In two months after seeding buckwheat starts blooming and producing nectar. Blooming on the different fields usually lasted for a month. Usual blooming time is in the late August and early September. This is the time when bees usually don’t swarm anymore and they stop building drone comb.
In the 18th century beekeepers use simple wooden box as a hive. Bees built a lot of comb and fill it with honey on buckwheat nectar flow. Since the most important flow was late in the summer, beekeepers tend to produce as many swarms as possible in the spring. In that way they were able to destroy a large part of hives in the fall.  It was common to destroy also bees to get honey  even though that advanced beekeepers like Janša (1771) and Glavar (1976) taught to drive bees out of hive with smoke and knocking.  They joined together expelled bees in strong colonies, which over winter better and also produce more swarms in the spring. Some advanced beekeepers found that it is easier to make artificial swarms then to wait for swarms and collect them in the trees.
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Queen rearing as a part of artificial swarm production

Janša (1771) and Glavar (1776) suggested to use queens which can be collected during natural swarming or prior artificial swarm production. If there were no available queens they reared new one out of the brood left in old hive or they put a piece of comb with small larvae into the hive with the swarm. They know that such artificial colonies can swarm later. If they wanted some queens, they collected them before swarming otherwise they destroyed surplus queen cells and queens. They were able to manipulate bees to rear queens for their needs because they observed bees carefully and learned reproduction of bees. Part of that was also observation of queen mating signs as a part of wider understanding of queen mating behavior.
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Detailed description of queen mating behavior

First report of queen mating outside the hive was in 1763. At that time Anton Scopoli published book Entomologia Carniolica. He repeated his note latter in Dissertatio de Apibus (1770), which was also translated into German (1787) and Italian (1779). In 1771 Anton Janša published Dissertation on Bee swarming (original title: Abhandlung vom Schwärmen der Bienen). Janša also wrote about mating of queens outside the hive. He nicely described how virgin queen leave the hive and come back:
"On a nice day from 9AM to 3 or 4 PM  queen comes with numerous court of bees and drones out of the hive. She turns around in all directions at the hive entrance to memorize hive, that she can find it later when she comes back from mating. For a while she flies in circles in front of the hive, watches it and elevates higher and higher. When she turns back from mating, she doesn’t go immediately into the hive, but  flies in front of it for a while. At this time we have to be careful on following signs after which we can recognize that queen has been copulated. If the abdomen is opened at the place where bees have sting, or something white is hanging from the abdomen, like a thin thread, and if it looks like that abdomen is broken and rent then queen has copulated for sure."
Janša noticed that queen flies out several time, but he didn’t point out in his book that queen copulates also several times and he didn’t describe actual mating in the air. Detailed description of queen mating with the drones was reported by Anton Humel (1775). This report was written at list four years before printing. Agricultural association in Ljubljana got it from Anton Humel. In 1771 the association asked well known beekeeper and priest Peter Pavel Glavar to write some comments. Glavar wrote a long comment with more details of queen mating (Miheliè 1976). He wrote that matted queen is staying in the hive and fly out later only with the swarm. He also pointed out that the old queen goes with the first swarm. He noticed also that queen is fertilized by several drones and not only one as was generally excepted at that time. His note was published together with Humel text in 1775.
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Spread of the knowledge on queen mating behavior

Many different sources show that many beekeepers of Carniola understood well queen mating behavior in the 18th century (Miheliè 1976). We can expect that beekeepers in Austrian empire learned that from Anton Janša, who was a court teacher of beekeeping in Wien. On the other hand, Peter Pavel Glavar was very active in teaching of beekeeping in Carniola. He tried to spread his knowledge among poor people. He knew that they can improve their living with beekeeping. On the other hand he had a big library and he had many correspondence with different kind of people around the Europe. Anton Scopoli, who first published observation on queen mating, was not  a beekeeper. He was well known naturalist, who collects and determine many plant and animal species. Most likely he got information about queen mating from beekeepers, and that he copied this information from Janša and Glavar. Observation of queen mating behavior was published later also by Huber (1792). We have no evidences if he got any information from Carniola.
Even though that beekeepers had a good knowledge about bee reproduction biology in 18th century and that there were several reports on that, this knowledge was lost in 19th century. It was replaced by the dogma of queen monogamy supported with the romantic stories created by the poets like Maeterlinck. It was very hard to break this dogma in this century. Ruttner wrote in 1985: "We experienced ourselves 30 years ago how much evidences was needed to break this dogma". In fact, this result in a high volume of scientifically collected data by many scientists in the middle of that century (see review by Ruttner 1956).
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Conclusion

More then two hundred years ago technological needs for artificial swarming production initiated detailed and careful naturalistic observation of reproduction behavior of honeybees. Beekeepers from this age gave us good descriptions of queen mating behavior. This is a nice example in beekeeping science how technological and economical needs effects understanding of fundamental phenomena in bee biology. Something similar happens nowadays, when spread of Varroa mites initiated many aplicative and also fundamental research of honeybees.
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References:

Glavar,  P.P. 1976*. Dissertation on bee swarms (Pogovor o èebelnih rojih). In: Ob 200 letnici pisane besede o slovenskem èebelarstvu, p.p. 79-260. Zveza èebelarskih društev Slovenije - Ljubljana.
Huber, F. 1792. New observations on bees. Transl. (1926), Datant, Hamilton, Ill.
Humel, A. 1775. Praktische Eröffnung daß der Weiser wirklich von den Thränen außer den Bienenstock befruchtet wird. In: Wöchentlichen Kundschaftsblatt des Herzogthum Krain (21).
Janša, A. 1771. Abhandlung vom Schwärmen der Bienen. Wien.
Miheliè, S. 1970. Beekeeping (Èebelarstvo). In: Gospodarska in družbena zgodovina Slovencev I, p.p.395-408. DZS, Ljubljana.
Miheliè, S. 1976. Peter Pavel Glavar, beekeeper, beekeeping writer, teacher and organization worker (Peter Pavel Glavar, èebelar, èebelarski pisec, uèitelj in organizator). In: Ob 200 letnici pisane besede o slovenskem èebelarstvu, p.p. 20-65. Zveza èebelarskig društev Slovenije - Ljubljana.
Ruttner, F. 1956. The mating of honey bee. Bee World (3) p.p. 2-15, 23-24.
Ruttner, F. 1985. Reproductive behavior in honeybees. In: Fortschritte der Zoologie Vol.31: Experimental Behavioral Ecology and Sociobiology, Edds. B. Holldobler and M. Lindauer, p.p.225-236. Gustav Fischer Verlag, Stuttgart, New York.

* Original text was written in 1776 but not printed. This was the first written beekeeping book in Slovene language. Glavar translated Janša’s (1771) book and added a lot of his own comments and descriptions. Glavar’s original text was put in modern Slovene by Stane Miheliè and printed in 1976.
 

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