Apivox Varroa Eliminator Project

The project APIVOX VARROA ELIMINATOR, like previous project APIVOX SMART MONITOR, at the beginning did not assume that it will be necessary to study the behavior of Varroa mites in bee colonies. Having studied dozens of scientific works of world-famous scientists over the 20th century and in recent years, we realized that there is an opportunity not to destroy the mites, but to introduce their life into a frameworks, living in which, they will cease to destroy bee colonies. All scientists' studies confirm this, but all scientists afraid to go against the tide, and to say, that especially this factor can save the bees from the mites ... I decided to confirm experimentally that this is possible.

Basing on these ideas, I developed the hive and a method for its use, which used together, should create such conditions for the mites, in which they should stop uncontrolled reproduction. Naturally, the hive had to be cheap, compatible with standard hive models, and non-volatile. In laboratory conditions, all this is simple to do, but in a real apiary, everything turned out to be not so simple as it seemed at the beginning.

In addition to the fact, that we needed to test our hive, we were faced with the fact, that there is no unambiguous criterion by which it would be possible to evaluate the effectiveness of its work, without interfering with the very process of bee life and the life of the mites population. All methods of control of quantity of the mites in the bee colonies do not give 100% accurate results. And every method has its own shortcomings.

We have chosen the most gentle control method - the method based on the analysis of free fall of the mites on sticky board. And as it turned out, even here there is no exact scientific data. We have been observing and analyzing the results for a year and a half. I had to create working hypotheses and test them in practice. In the process of work, some conclusions and ideas appeared, which we presented for review in the form of articles to the beekeeping community in our blog, which you can always read here - apivox.blogspot.com.

Unfortunately, we have not yet solved the main problem. The behavior of the bees turned out to be much more complex than what the scientists described. Not all of our actions had the expected effect due to the behavior of the bees at different times of the day and the year. But there is a certain results ... We managed to slow down the development of the mite's population in the experimental hive compared to the control group. We managed to understand the features of the development of the mite's population in a bee colony during the year. We managed to create a method for predicting the size of the mite's population, a method for predicting their further development, to determine which moments in the life of bee colony are the most critical from the point of view of a sharp increase in the mite's population. We believe, that we have partially understood the mechanisms of growth of the mite's population and the possible mechanisms of influence on them ...

Our work continues despite all the difficulties and lack of assistance from the world beekeeping community...


In the frameworks of our Apivox Varroa Eliminator project, we continue to review and analyze the state of the bee's colonies and the mite's population living in this colonies, using the method of counting free fall of the mites on sticky board, and we believe that this method, although it does not yet allow one hundred percent accurate numerical data, can help to determine key points in the development of bee's colonies and the population of mites parasitizing on them.

For a long time scientists tried to adapt such a parameter as the mites drop, for prediction of the degree of mites, parasitizing on the bees in the colonies ... But everyone wanted to get specific numerical data, even by simply adjusting the numbers to the needed results ...

On the basis of this method, the ministries of agriculture of the United States, Britain, Spain and other countries distribute, their own recommendations and methods of Varroa mites' control. On the basis of the average number of mites, falling on sticky board per day, it is proposed to build the strategy for treating bees with acaricidal drugs! Representatives of the agricultural industry, trusting scientists, made the method of calculating the free fall of Varroa mites, the main one for estimating of infestation of bees' colonies, as the least laborious.

It is believed, that it gives fairly accurate results. But sometimes it's not quite right, and sometimes it's not right at all .... We came to this conclusion for two reasons: The first reason - is that in different works of scientists, who studied and evaluated the accuracy of all known methods of calculating of the quantity of Varroa mites in bees colonies, it is written, that all methods are equally not accurate at the beginning and at the end of season, when the number of mites in the colony is too small. And that in these works, scientists use at least three different coefficients to translate the amount of daily free fall of mites, into the total number of mites in the colony of bees. We know that scientists use this special coefficient, by which, the average daily fall of mites is multiplied ... It is equal to :

1) 20 in spring, and to 40 in summer and to 250-500 during broodless season. ( Population model for the ectoparasitic mite Varroa jacobsoni in honey bee (Apis mellifera) colonies Stephen Martin National Bee Unit, Central Science Laboratory, Sand Hutton, York, YO4 1LZ, UK )

2) 120 throughout the season. (Development of the mite, Varroa jacobsoni oud., in the honeybee, Apis mellifera L., in Michigan, USA, and a comparison of diagnostic methods for detection of the mites Ahmad Al Ghamdi and Roger Hoopingarner )


This already suggests, that each researcher adjusted the coefficient to the situation in his apiary, or in his zone. And even this fitting does not work if used in mathematical models, which are compared with the real results of counting of Varroa mites in the brood and on adult bees... (Population growth of Varroa destructor (Acari: Varroidae) in honey bee colonies is affected by the number of foragers with mites. Gloria DeGrandi-Hoffman, Fabiana Ahumada, Victor Zazueta, Mona Chambers, Geoffrey Hidalgo, Emily Watkins deJong)

It must be said, that counting of a quantity of mites, using another methods is not much better ... Counting the quantity of mites, parasityzing on adult bees is not indicative since in summer, up to 70% of the mites are situated in the sealed brood... analysis of the sealed brood are not accurate due to the uneven distribution of the mites in the brood and due to the presence of a part of the mites on adult bees...

Calculation of the percentage of mites in any form is also not an indicator... With different strengths of bee colonies, the same percentage can mean a multiple difference in the real number of mites, which, in the process of development of bee colonies and reproduction of mites, leads to a multiple difference in the percentage of mites when the strength of bee colonies will become equal...

For our project we invented our own method of calculating of mites drop. It consists in a separate count of dark - old, already participated in reproduction females, and pale-colored young female mites and deutonymphs, which died or did not fertilized by a male mite inside the cells, and were thrown out by the bees. We call this the two-graph method. So, what is its meaning ...

The point is that in addition to the fall of an old dark mites, that is clearly visible to everyone, there is another process - the process of death of young females that have not yet participated in the growth of the mite's population. Pay attention to the photo ... It shows the mites, which fell down on sticky board of our experimental hive in different seasons of the year.

They all have a different color !!! Mature old fertilized females are deep red and burgundy! Young females have a bright red color that is darker or lighter depending on maturity, and the paler ones are not mature females and transparent individuals are deutonymphs ... In order to find this facts, we had to turn to authoritative documents ....

On the upper diagram we see once again the confirmation of our practical results - 1) Varroa mites in the summer live 2-3 months. In practice, we see about 2.5-3 months. 2) Immature and not fertilized young females do not survive. 3) The final red-brown color of Varroa mites is gained after the last molt ot the mite, when it become fully grown and capable to lay eggs. Young females does not have such a color. They are paler. The color is closer to brigh-red. Confirmation of this item we took from another document... (A deadly honey bee parasite The Varroa Mite PUBLISHED BY Bayer AG, beecare@bayer.com www.beecare.bayer.com)

"...Graphic showing Varroa mite development stages The first development stage of the Varroa mite is a six-legged larva that develops inside its closed egg. The eight-legged protonymph emerges in the post-hatching second stage. In the third phase this protonymph develops into a deutonymph, which then becomes the adult mite. In the final nymph phase, the growing mite undergoes two immobile transitional chrysalis stages (protoand deutochrysalis). Young female nymphs are white in color and, after molting several times during the development process, turn dark reddish-brown during the last immobile stage – the deutochrysalis – and maintain this color as adult Varroa mites..."

So, the method of two graphs that we use, assumes separate counting of dead old females of the mites, dying during or after breeding, and dead young females, that did not participate in breeding. Shifting back the graph of mites drop of mature dark mites by 2.5-3 months, or by the approximate lifespan of the tick, we can talk about the approximate time of their birth, and we can analyze the events, that led to their appearance and development in the bee colony. The appearance of the mite's drop, consists of pale mites is, according to our long-term observations, a very bad sign, and speaks of an over-intensive reproduction of the mites, when it uses all brood zones for reproduction, even that are not too suitable for reproduction in terms of their parameters. Thus, using two graphs, we can learn about the past of the mite's population, about the prospects for its increase in future, and about the intensity of the process of growth of the mite's population at the moment.

In general, free fall of the mites at the time of testing is not always indicative. There are intervals, as we can see from the graphs, when there is practically no mite's drop or it is small ... This does not mean that there is no mites or its number is small... It is just mean, that the mites are waiting for suitable conditions for mass reproduction.

From the graphs we can see, that the mass death of the mites of all ages is directly related to the cycles of enhanced reproduction, which apparently takes away the last strength from the old individuals and leads to their death... In this case, the idea that the mites live for about 2-3 months and the method of shifting the graph of free fall of the mites by 2-3 months back shows the time of their birth, allows us to see the real time of birth of this mass of falling mites ... And these are the months, when it seemed to us, that there was almost no mites or very few... In particular, so was in January-February 2020, as can be seen from the graphs below ...

(The graph of free fall of the mites (mites per day) in 4 colonies of our experimental apiary during 2019-2020 years)

An analysis of the graphs of free fall of the mites obtained by us during 1.5 years of continuous observations of 4 colonies of bees, shows the following...

Almost throughout the winter, except November and December, the mite population had the opportunity to reproduce and maintain its existence, since the bees had brood due to the warm winter and wintering in a pavilion heating by the sun. This led to the fact, that the first massive release of new mites and the fall of old and immature mites coincided with the honey harvest from the willow on about April 23rd. The influx of large volumes of nectar and pollen led to the first mass release of young bees from the brood, and accordingly, to the first powerful release of the mites. The second release took place at pre-swarming time from 11 to 30 May, when the number of brood at least doubled, and was laid drone brood. All these led to the next outbreak of the mites population.

In colonies 5-1 and 5-2, the growth of the mite's population during the main honey gathering stopped due to a reduction in brood quantity due to the filling of the nest with honey, but at the end of it, the mites began to reproduce again. The colony 5-1 lost its queen and was left without open brood for a couple of weeks. The mites drop in it has grown quite insignificantly, and then it sharply decreased. But in the 5-2 colony, at the end of honey harvest, free fall of the mites of all ages again sharply increased to close to critical values.

In the E-1 hive, the growth of the mite's population was so strong that, when the swarming began, it led to a complete loss of the colony. The reason is that this colony with an old queen had already laid swarming queen cells by May 7. The sharp increase in the amount of well-heated brood resulted in a powerful increase in the mite's population. As a result, in the colony, there were a lot of sick and dead larvae in the brood, and sick young bees. All infected brood was abandoned. The colony, despite the removal of swarming queen cells and the division in half, flew away in the form of two swarms.

In the hive 8-1, a similar picture began with some lag behind E-1. The bees of this colony also laid swarming queen cells, but the queen in this colony was young, and the removal of queen cells led to the termination of the swarming mood of the colony. Despite this, it was laid a lot of drone brood , which caused the subsequent increase in the mite's population. The growth of the mite's population began in this colony too, but a little bit later, on the background of honey collection, since the colony was not quite ready for it and began to bring marketable honey later than 5-1 and 5-2 ... In order not to lose the colony, were taken a measures, as you can see on the graph, on June 23. One frame of drone brood was removed and was completely destroyed drone brood on all other frames. The fall of immature and mature mites dropped sharply. This indicates the possibility of temporarily suppressing of the development of mite's population in pre-swarming time by removing the maximum amount of drone brood. But at the end of the honey harvest, the mites were able to develop vigorously again, and the fall of the mites of all types increased sharply to close to critical values.

So, what did our observations show in general...

1) Observations show on the example of all colonies, that each next burst of the mite's reproduction is at least 2-3 times stronger than the previous one. This difference roughly corresponds to the amount of mature mites fallen free during the present reproductive cycle, divided by the amount of fallen free immature mites of the previous reproductive cycle... The value is approximate, but it is confirmed in all experimental colonies. You can see it on the graph.

2) Observations show that swarming state of bees should be avoided by all means. It is the massive laying of drone and bee brood before swarming and, most importantly, its thorough heating by the bees, that leads to a catastrophic increase in the mite's population due to a multiple increase in the reproductive territory, comfortable for it.

What happens in this case with the bee colony? Let's have a look...

In the course of our scientific work and acoustic monitoring of bee colonies using our Apivox Smart Monitor device, we came to understanding, of the changes, that occur in the bee nest in pre-swarming period...

The fact is that at the beginning of the process of preparing for the swarming, the task of the colony and the queen is to build up the maximum possible strength of the colony before the swarm leaves. Then, dividing in two, the colony will remain large enough to survive and collect honey for the wintering. To do this, the queen begins to fill all the cells in the hive suitable for this with brood ... But the nest is not the whole hive, and the bees do the following !! They start powerful heating of the ENTIRE hive ! The whole hive turns into a nest! The earliest queen cells appear on the periphery of the nest - on the second frames after the extreme honey frames ...

It was at this moment when a specific acoustic signal appears in the hive, which we use as a sign of an impending swarming, which Woods considered to be a signal of young bees, and prof. Eskov mistakenly considered to be a signal of passive bees .... A signal, that accompanies heating of the hive and a signal that accompanies heating and aeration of open brood. These signals becomes so powerful, that they dominates in the hive even in the daytime, suppressing the signals associated with the work of bees on honey harvest!

This event in the life of a colony of bees becomes the "golden" time for Varroa mites and the cause of the subsequent troubles of bees ... down to their death .... Now the mites can actively reproduce in a huge amount of brood, and not only in brood of worker bees, which used to be warm enough even earlier, and also to reproduce actively in drone brood... This at least doubles their breeding potential, which ultimately leads to exponential growth of the mite's population in April-June.

Such a leap in the growth of the mite's population occurs primarily in the colonies, which are initially ready for swarming - that is, in the colonies with old queens. It is possible, that this was the reason for the confidence of many beekeepers that bee colonies with young queens are better able to cope with the mites. But it is normal thing, in the colonies with young queens, bees are less prone to swarming, which means, that they are less prone to heating the entire hive and to the massive construction of drone cells, which become a "bomb" placed under the colony during swarming time. In normal times, drone brood located in cool areas of the hive does not create such an "incubator" for the mites, and colonies that do not enter the swarming state suffer from the mites much less.

The use of anti-mite's techniques associated with expansion and, as a consequence, with cooling of the nest, will worsen the mite's reproductive capabilities and, possibly, save the bees from death in the event of a high infection of the bee's colonies in spring.

Basing on this, we can say, that in springtime, early diagnosis of swarming state of bee colonies become a very important point for a beekeeper who want to get ecologically pure honey. Using Apivox Smart Monitor application, you will be able to know about it at least 2 weeks before the start of the swarming process, or even earlier, and will be able to understand, that it is high time to take measures, which will not only help to avoid losing of swarms and honey, but will also dramatically reduce the number of mites in bee colonies.

3) Observations show that if, during the next cycle of enhanced reproduction, free fall of an immature mites significantly exceeds free fall of an old mature dark mites, and has sufficiently large values, then this is a very alarming signal. It can be assumed, that not only a large number of young mites were released, but the founding females did not die ! Thanks to this, accumulates the mass of live mites, ready to participate in the new reproductive cycle, and exactly this make the next flash of reproduction so powerful.

During the main honey flow period, when the mites have less bee's brood in the nest for reproduction and worsening climatic conditions in the nest, overall reproductive opportunities of the mites reduces. Free fall of the mites sharply reduces or stops altogether ... The mites of reproductive age are waiting for the optimal moment to start laying eggs ... With a decrease in the flow of honey at the end of the main honey harvest, as well as with removal of honey by beekeepers at the end of summer, the volume of free space for laying eggs by the queen increases ... The weather conditions at this time, in August-September, are still favorable for the bee's brood rearing and for the mite's reproduction. At this time, beekeepers are trying to get as many young bees as possible, because exactly these bees will winter and form a colony next year. Under these conditions, starts new powerful laying of brood by the mites, which, as a rule, leads to a powerful autumn surge of the mites population, and during release of young bees from the cells, to a strong fall of old mature females, and immature young female mites.

All this can be seen in the diagrams for colonies E-1 and 5-1.5-2. It was in the E-1 colony, which showed the greatest growthof free fall of pale immature mites, and as a result, a sharp increase in the mites population till a critical numbers. This led to the loss of the colony. In colonies 5-1 and 5-2, the number of immature mites falling free on sticky board was significantly less, and it needed about two months for the mite's population, to grow to almost the same size as in the E-1 colony.

4) Observations show, that the state of a bee colony when a quantity of falling mature mites reaches quantity up to 5 mites per day, in principle, is not too bad for a colony of bees of almost any strength. The bees of a strong colony can even withstand situation, when a quantity of falling mature mites reaches 60-70 mites per day, for up to a month. They are also can withstand situation when a quantity of falling mature mites reaches from 20 to 40 ticks per day, for 2-3 months. Of course, in the bee's colony appear many damaged bees that are unable to fly, and this weakens the colony and reduces its honey productivity... We can see such a situation in the graphs of three colonies of our experimental apiary - 5-1, 5-2, 8-1. At the same time, 8-1 and 5-1 produced about 40 kilograms of marketable honey, and a 5-2 colony - 60 kilograms, while being in a stationary apiary. Thus, we can say, that the state of a bee colony when a quantity of falling mature mites reaches quantity up to 10 mites per day, in principle, is not terrible in the short term for a strong colony of bees. The danger appears when, in addition to mature dark-colored mites, immature pale-colored mites appear on sticky board. This is a signal of alarm. This is a sign of an incipient reproductive "outbreak" of the mites. So, as you can imagine, the main task of beekeeper is to combat these periodic reproductive outbreaks! Their suppression should be the main task.

What methods can be used for this... Let's consider these methods in chronological order in accordance with the life of bees...

Wintering: The first step in keeping bees properly in terms of Varroa mites control - is cold wintering. This method has been known in Europe for a long time. Carnica - is a European bee breed and it is in Germany that they know how important it is to suppress its natural tendency to constant brood rearing. It is very important to make bees to stop brood rearing early enough, and to start brood rearing as late as possible. It is very bad when the bees finish growing the brood in November and start growing it again in January. It is thanks to this that the mites can slowly and imperceptibly increase their quantity during winter, and with the first abundant spring bees' brood, can sharply increase size of their population to critical values. This is how our E-1 colony died. The graph clearly shows that exactly in this colony we can see fairly strong fall of old mites throughout the winter, and already in April we can see the appearance of a peak in free fall of immature and mature females - that is, a powerful surge in the reproductive activity of the mites.

It is ideal, if bees, especially Carnica bees, begin to actively rear brood with the arrival of stable heat and appearance of pollen sources in nature. It is from this moment and not earlier begins the active development of bee colonies . At this time, if the bees have warm nest and are supplied with nectar and pollen, they build up their strength surprisingly quickly, especially in the case of the bees of Carnica breed.

In other case, exactly population of Varroa mites, rather than the population of the bees, develops and renews better. That is why in Europe and especially in Germany, cold wintering has acquired particular importance in the fight against varroatosis. It is exactly cold wintering should not allow the mites to multiply during the winter, when beekeeper cannot help the bees to get rid of them. It is this event that will help to avoid the first spring peak in the growth of the mite's population. And here we can say, that design of our new hive, or rather its inner cover, makes it easy to provide the bees with both - cold in winter and with warm in spring.

Spring development till the main honey flow: The main methods of influencing the mites population in this period are: the reduction of the space suitable for the reproduction of the mites and the deterioration of the conditions, required for their reproduction. Several techniques can be involved here together or separately ...

1) Creation and maintenance of young colonies. Creating young colonies in the apiary to replace old working colonies by forming artificial swarms, by forming offshoots with mature queen cells, the beekeeper "kills two birds with one stone at once." Firstly - when creating such colonies, the brood which at this time is the main repository of the mites, is not used or is used minimally. Secondly - such colonies are easy to process in any way, including using of formic acid, which is a method, that minimally contaminates frames and honey. Thirdly - the colonies will have young queens, which is an excellent anti-swarming technique. Such colonies, as a rule, have a good start in development and begin wintering healthy and clean.

2) Suppression of swarming state of the bees in order to deprive the mites possibility of super-active reproduction in a huge amount of well-heated brood of worker bees and drones. Before the start of the main honey collection, possible methods are: the presence of young queens, expansion of the nest, removal of maximum possible amount of drone brood. Firstly, as we have already said, colonies with young queens are, in principle, much less prone to swarming. Secondly, the expansion of the nest reduces swarming mood of worker bees and deprives mites of a large, well-heated brood zone and, most importantly, a well-heated drone brood. Thirdly, the use of building frames allows not only to reduce swarming mood of the bees, but also to remove from the hive and destroy almost ninety percent of drone brood with mites in it, with a minimum expenditure of efforts and without injuring the bee colony.

3) Removal of sealed brood. Removal of frames with sealed brood, which can be carried out during the period of bee's colonies growth, till the main honey collection, in order to prevent swarming and to create offshoots, also sharply reduces the number of mites in bee colonies, and significantly contributes to the fact, that the disease does not take on a threatening character in the main colonies until autumn, even in the absence of other mites control measures. In addition, holding such an event in June can, under certain conditions, increase honey productivity of bee's colonies by reducing number of bees engaged in brood rearing. At the same time, the colonies from which the brood was removed, can be treated with formic acid, and this treatment will be very effective at this stage.

In the process of honey collection, bees themselves usually create conditions, which are not very good for the mites reproduction ... This can be seen from the graphs of free fall of the mites in all four colonies. ( June-mid of July) But even at this time, it is possible to influence the mite population using methods of temporary limitation of queen egg laying, and removal of sealed brood. Unfortunately, such measures during the period of honey collection cannot be radical, otherwise there will be losses of marketable honey due to weakening of the strength of bee colonies.

Autumn preparation of bee colonies for wintering. It is possible that you can again use the method of removing the sealed brood, and then proceed to cooling the nest, in order to stimulate the end of brood rearing season. In this case, the first task is to interrupt the mite reproduction chain in working colony. Using this method, beekeeper can remove most young mites and founding females into a separate colony, which will be a subject of radical anti-mites treatment after the entire brood will release. At the same time, in all colonies from which the brood will be taken, should be given new combs, in which the queens will begin to lay eggs, from which will born the bees, which will go wintering strong and healthy. At the same time, a huge number of mites will be removed from working bee colonies, and will not be able to start a new reproduction cycle. This should be done at the very beginning of the growth of peaks of free fall of the immature mites, which means, that a lot of mites already entered the brood cells for reproduction. The second task is to ensure a smooth transition of colonies to cold wintering with corresponding decrease and further termination of brood rearing.

The use of our new Apivox Varroa Eliminator project and a new method of keeping bees takes a special place. Using this method, we managed, although not to completely suppress the development of the mites population in the experimental colony, but to reduce significantly their growth rates in summer ... Today we believe, that the design of the hive, although correct, is still not optimal. But even today it has certain advantages ...

- Its design makes it easy to control the mites drop, in order to understand the processes occurring in the mites population.

- Its design allows to ensure the autumn transition to the winter regime and cold wintering of bees, as well as their warming in the spring and summer.

- Its design allows to accelerate the development of colonies in spring and summer, while reducing the intensity of mites reproduction.

An analysis of data from the past 2019 showed, that if we can consider free fall of the mites proportional to their number in the bee colony, or to the size of the mites population, then, the growth rate of the mites population in the experimental hive number E-1 did not exceed six times, despite the fact that the colony had an old queen bee and a lot brood, while control colonies, some of which had young queens born in this year, and which had less mites, had growth rates from ten to one hundred times... In this case, we are talking about relative values. The absolute values of free fall of the mites in experimental colony were initially higher than in the control colonies, but by the fall these values were almost equal ... In the end, not the design of the hive led to the death of experimental colony, but the need to conduct observations of the development of the mites population in it even till its death!

In general, new hive showed that the bees can live and develop in it, and we can select its parameters in order to suppress the development of Varroa mites to one degree or another. But this will have to be done again next season, in order to obtain more statistically reliable results.

So, we are confident that application of the above rules of keeping bees throughout the year and the use of our new hive will be able to jointly solve the problem of preserving bees and of suppressing the development of Varroa mites in their colonies.

Here we used diagrams and drawings from the following works:

1. Population model for the ectoparasitic mite Varroa jacobsoni in honey bee (Apis mellifera) colonies Stephen Martin National Bee Unit, Central Science Laboratory, Sand Hutton, York, YO4 1LZ, UK
2. Development of the mite, Varroa jacobsoni oud., in the honeybee, Apis mellifera L., in Michigan, USA, and a comparison of diagnostic methods for detection of the mites Ahmad Al Ghamdi and Roger Hoopingarner
4. Population growth of Varroa destructor (Acari: Varroidae) in honey bee colonies is affected by the number of foragers with mites Gloria DeGrandi-Hoffman Fabiana Ahumada Victor Zazueta Mona Chambers Geoffrey Hidalgo Emily Watkins deJong
5. A deadly honey bee parasite, the Varroa mite. PUBLISHED BY Bayer AG


APIVOX PROJECT. A new theory of communication in bees and robotic systems. What is common.
Examples of usage of communicative and marker signals by bees.