The latest bee find, one preserve in amber found in Myanmar, now dates bees to an estimated 100 million years ago, during the Cretaceous. This is the time of dinosaurs, and makes bees even older than the Australian native bees (These were solitary, non-social bees).

Honey storing social bees developed during the Miocene, between 20-10 million years ago. These bees made their nests in hollow trees, caves, crevices, rocks and holes in the ground.
Hives have been in existence since palaeolithic times, about 10.000 years (as depicted in early rock paintings in a Spanish cave in Valencia, in which a ladder was used to reach the nest and a container was used to hold the honeycombs).
Ancient Egyptians used honey in religious rites, for feeding the sacred animals, in many ceremonials, and even for preserving corpses.
Honey has been found in Egyptian tombs, dating back 5.000 years. It was edible, although the colour had changed significantly.
In 3.000 B.C., Egyptians kept written records of bee keeping activities. Egyptian hives were transported down the Nile on barges to access floral sources.
The Rig-Veda, India`s oldest sacred book (between 3.000 – 2.000 B.C.), contains many references to both honey and bees. Here, honey was also used in many rituals and ceremonies, and in pharmacy.


The Greeks have always been closely tied to the world of bees. The first who scientifically worked with those in 300 BC was Aristotle, in his «Historia Animalium».
At that time the Hymettus mountain was the sacred mountain of beekeepers and the local honey was known beyond the borders of ancient Greece.
Aristotle was an ancient Greek philosopher who lived during the time of Alexander the Great. He had been selected by Philip, King of Macedonia, to educate his son Alexander. Thus Aristotle maintained a very close relationship with Alexander the Great and was in position to study much of what existed in Alexander`s Greek empire.
Aristotle`s “Historia Animalium” (History of the Animals) is divided into ten books. Generally they are about the animal kingdom and describe various animals, parts of animals, generative systems, and other useful information observed and studied. It is still worthwhile reading not for exact modern science regarding animals but for the link to the ancient world and the concept of life as Aristotle and his peers saw it. The fourth book discusses animals without blood and in the Fifth book Chapter XVIII he makes a number of remarkable observations about bees. It is characteristic that in over 52 parts of the work History of Animals, Aristotle refers to bees.
Some of the most important parts are:
From Book V chapters 18 and 19
“The king bees never leave the hives, either for food or any other purpose, except with the whole swarm; and they say that, if swarms wander to a distance, they will retrace their steps and return until they find the king by his peculiar scent. They say also that, when the king is unable to fly, he is carried by the swarm; and if he perishes, the whole swarm dies with him.”
From Book Nine, Chapter 27
“It is good for the bees to have a few drones among them, for it makes them more industrious.”
From Book V, Chapter 19
“When the wind is high, they carry (the workers) a stone with them for a balance”
From Book Nine, Chapter 26
“They all have their proper work to perform. Some bring flowers, others water, and others polish and erect the cells. Water is brought when they are rearing their young.”
Aristotle observed bees and wrote about bees but he was no beekeeper. It is certain that he knew beekeepers and honey sellers, observed the work of bees and read about them. Often he repeats, “Other say, other persons affirm, and not all people agreed”. This indicates that much of his material was gathered from others and he is reporting the various points of view then held.
Historia Animalium, “The bee dance”
624A 33-B, D11
In each flight, the bee does not go into a different kind of flowers, but for example, flies in from violet to violet and does not touch any other until she returns to the hive.
When they reach the hive, shake and each other accompanied by two or three bees.

In ancient Greece, Hippocrates recommended honey for many diseases, while Diogenes wrote that «older people who eat only bread and honey, live many years.»
Democritus of Abdera lived 109 years and as he said: «honey inside, olive oil out».
Others, like Zenonas and Pythagoras reached a very old age because they fed exclusively on honey and bread. Also in Ancient Greece, the bee was sacred and was the symbol of fluency.

Bees are insects belonging to Hymenoptera. In the genus Apis, belong four species of bees from Asia, like Apis dorsata, Apis florea and Apis cerana.
The Apis mellifera, located in Europe, Africa, the Middle East and a part of Siberia.
Bees are social insects. This means that they tend to live in colonies where all the individuals are of the same family, often the offspring of one mother. In the more highly organized societies there is a division of labour in which individuals carry out particular duties.
There are three kinds of bee in a colony: in the summer, a few hundred drones or males, one egg-laying female, or queen, and from 20 to 80 thousand sterile females or workers. The mature queen is usually easily recognized by her large abdomen.
A queen bee may live from two to five years and, except for a short period at the end of her life when one of her daughters takes over the colony, she is the only egg-laying female. All the members of the colony, whether drones or workers, are her offspring. She spends all her time laying eggs, perhaps up to 1500 a day, each one being placed in a wax cell made by the workers. The queen can feed herself but in the hive the nearest workers turn towards her, lick her body and feed her by regurgitating a special secretion of their salivary glands, called “royal jelly”, on their probosces from which the queen can absorb it.


The queen usually mates only once in her life (though second and third matings are known to happen) and stores the sperms received from the drone in a sperm sac in her abdomen. This store of sperms lasts her for the two or more years of egg-laying, a small quantity being released with each fertilized egg laid.
When the store of sperms is used up she may continue to lay eggs but they are all unfertilized and will become drones. By this time one of her daughters has been reared as a queen and is ready to take over the egg-laying.
Each egg is laid in one of the hexagonal wax cells and hatches into a tiny, white legless larva. The larva feeds on substances deposited in the cell by the workers; it grows, pupates in the cell, hatches as an adult bee and finally emerges from the cell into the hive. The eggs hatch after three or four days and by nine days are fully grown and ready to pupate. The workers put a capping over the cells at this time. Ten or eleven days later the capping in bitten away and the adult emerges. The times given above vary with changes of temperature and according to whether the bee is becoming a drone, worker or queen.
The drones, who live for about four to five weeks and do not work inside the hive, are fed by the workers or help themselves from the store of pollen and nectar in the combs. Their function is to fertilize a new queen. In the autumn, or when conditions are poor, they are turned out of the hive where, unable to find food for themselves, they soon die.
The workers are female bees whose reproductive organs do not function. Among many other tasks they collect food from outside the hive and store it, make the wax cells and feed the developing larvae.

The wax combs are built hanging vertically with a gap of about half an inch separating each one. The cells in each comb thus lie horizontally. The workers prepare three kinds of cell: worker cells about 5mm across, drone cells about 6mm across, and queen cells quite different from the others. The queen cells are larger and made individually, pointing downwards like small acorns from the surface or bottom of the comb. The relative numbers of these three kinds of cell seem to depend on the time of the year, the temperature , the abundance of food and condition of the colony. Normally, the worker cells predominate.
Eggs are laid by the queen in the brood area. This is where the temperature is about 32oC, kept so by the heat given out by the bees bodies. The area varies, diminishing in the winter and expanding in the summer. The queen moves over the brood area, laying eggs indiscriminately in any of the three types of cell she encounters, by placing her abdomen in the cell and depositing a single egg. The eggs placed in the larger, drone cells, are not fertilized, and this results in the eggs developing into a male bee or drone. In the queen and worker cells, fertilized eggs are laid.
For the first three days after hatching, all the larvae are fed on a protein-rich, milky secretion, called royal jelly, which comes from the salivary glands of workers of a certain age. The grubs in queen cells continue to be fed on royal jelly for the rest of their lives, but those in drone or worker cells are “weaned” onto a mixture of dilute nectar and pollen. If a one to three day old larva is transferred from a worker to a queen cell, it will receive the diet of royal jelly and develop into a queen. Thus, thought there is no difference between the eggs and young larvae in queen and worker cells, their different treatment by the workers results in their becoming quite distinct types of bee.
Exactly what aspects of their feeding cause this is not known for certain. It may be the absence of pollen from the queen`s diet, the cessation of royal jelly in the worker`s diet, the super-abundance of food placed in the queen cells or a vitamin-like chemical fed to the queen larvae in the early stages. After three days, worker grubs cannot be reared as queens, even if they are placed in queen cells and fed on royal jelly.


Drones, then, develop from unfertilized eggs in wide cells, queens and workers from fertilized eggs which are fed differently as larvae.
When a new queen emerges she is fed by the workers. She bites a hole in any other occupied queen cells that she finds and some observers believe she stings the occupants. In any event, the workers usually tear down the other queen cells that have been bitten into and destroy the occupants.
For a few days the queen leaves the hive for shorts flight lasting, at first, only a minute but gradually lengthening to about 15 minutes. During these flights she learns the geography of the district around the hive. On one of these flights she is pursued by drones, but not necessarily from her own hive; in fact, they do not follow her from the hive but are already waiting outside. One of them catches the queen and mates with her, depositing in her vagina sperms which eventually find their way into her sperm sac. She now returns to the hive, and soon after begins to lay eggs.
From glands in her head, the queen produces a mixture of chemicals called pherormones (queen substance). When the workers “lick” her body, the pherormones suppress their fertility. When, at the end of her life, the queen ceases to produce these pherormones, some workers start to lay eggs which, being unfertilized, produce only drones. They do, however, start building new queen cells.
The tasks undertaken by a worker bee depend partly on its age and partly on the immediate needs of colony. Generally speaking, the worker`s life follows the course described below, although the times given are very approximate and in many cases may not apply.
After hatching, she is fed by other workers and spends a good deal of time standing still on the comb. She does, however, clean out cells from which bees have recently hatched by removing the cast larval cuticles. On the fourth day she feeds on honey from the store cells and eats a good deal of pollen. Between the third and fifth day she feeds older larvae by placing nectar, water and pollen in their cells.


The pollen that she eats is rich in protein and helps her salivary, brood food glands to become active, so that by the fifth day they can secrete the brood food or royal jelly which is fed to the younger larvae. After ten or twelve days these glands cease to function effectively but wax glands on the underside of her abdomen begin to secrete wax which the workers uses for comb-building and repair. By this time she is also beginning to leave the hive for short flights during which she learns the position of the hive and the topography of the surroundings.
Between the twelfth and twenty-first days these reconnaissance flights continue; while in the hive, the worker collects pollen and nectar from the incoming field bees and stores it in the cells. She also processes the nectar and begins its conversion to honey, and cleans the hive by removing the dead bees and detritus from its floor.
After three weeks of hive duties the worker becomes a forager and spends the daylight hours collecting water, nectar, pollen and propolis and carrying it back to the hive. This work she may continue for about three weeks before she dies.
The “schedule” given above is by no means rigid, and observers have recorded bees doing several of the duties mentioned at the same time, as well as old bees performing “nurse” duties, and young bees foraging. Some of the duties may be missed out altogether. For example, only a few of the young foragers do duty as guard bees, protecting the hive from invasion by robber bees.
The foraging workers collect nectar from nectaries of flowers. The nectar is drawn off from the nectaries by the long labium. It is pumped up and swallowed into the honey sac, a region of the gut from which it can be regurgitated on reaching the hive. Nectar is a watery sugar solution when collected, but it is processed by the house bees to whom it is passed. These workers repeatedly swallow it, mix it with enzymes and regurgitate it. The enzyme action and the evaporation of water result finally in its conversion to honey. Nectar contains very little protein, and the pollen collected by the foragers makes up this deficiency.
Pollen is collected by combing off with the legs the grains which adhere to the bee`s body after it has visited a flower. The pollen collected on the head, and removed by the front legs, is mixed with a little nectar and passed to the back legs which have combed pollen from the abdomen.. The rows of bristles on the legs assist this combing action. The pollen press, in the joint between the tibia and tarsus of the hind legs, squeezes the pollen which is passed to it from the pollen comb of the opposite hind leg. The pollen and nectar paste is thus pushed by the press into the pollen basket on the tibia, where it is retained by the fringe of setae. All this may be done while the bee hovers in the air or while hanging from the flower. The forager returns to the hive with the two packs of pollen and pushes them off into an empty cell or into one with some pollen already in it.
The younger house bees then break up the pollen masses and pack them down into the cell. When the cell is full it may be cover with a little nectar and sealed over. Both pollen and honey sealed in the store cells are eaten by the bees in the winter months when no other food is available. Water is collected and used to dilute the nectar with which the larvae are fed, but there is no evidence of water being stored. Propolis is a resinous substance that the bees collect from trees and sticky buds. They use it for sealing small cracks and gaps in the hive.
When the size of the colony reaches a certain stage, usually in spring or summer when the nectar flow is at its greatest, the queen and a great many workers, leave the hive in a swarm. The swarm comes to rest in a great cluster on a tree branch or similar situation. Scout bees, who may have left the hive some days before, seek out a suitable situation for a new nest and return to the swarm and communicate this information, whereupon the whole swarm moves off to the new site. In the old hive, one of the new queens hatches out, mates, and takes over the colony that is left.

Honey bees make use of five senses throughout their daily lives; however, honey bees have additional communication aids at their disposal. The methods by which honey bees communicate are of particular interest. One is chemical, the other choreographic. Also sound and vibrations signals exchanged during the performance of wagging dances.

Pheromones are chemical scents that animals produce to trigger behavioral responses from the other members of the same species. Honey bee pheromones provide the “glue” that holds the colony together. The three castes of bees produce various pheromones at various times to stimulate specific behaviors.

Here are just a few basic facts about the ways pheromones help bees communicate:
Certain queen pheromones (known as queen substance) let the entire colony know that the queen is in residence and stimulate many worker bee activities.
Outside of the hive, the queen pheromones act as a sex attractant to potential suitors (male drone bees). They also regulate the drone population in the hive.
Queen pheromones stimulate many worker bee activities, such as comb building, brood rearing, foraging, and food storage.
The worker bees at the hive`s entrance produce pheromones that help guide foraging bees back to their hive. The Nassanoff gland at the tip of the worker bee`s abdomen is responsible for this alluring scent.


Worker bees produce alarm pheromones that can trigger sudden and decisive aggression from the colony.
The colony`s brood (developing bee larvae and pupae) secretes special pheromones that help worker bees recognize the brood`s gender, stage of development, and feeding needs.

Social behavior in bees has a number of advantages. One of the most important of these is the ability to quickly mobilize a large number of foragers to gather floral resources that may only be available for a short period of time. The ability to communicate location with such precision is one of the most interesting behaviors of a very interesting insect.
The recruitment of foragers from a hive begins when a scout bee returns to the hive engorged with nectar from a newly found nectar source. She begins by spending 30-45 seconds regurgitating and distributing nectar to bees waiting in the hive. Once her generosity has garnered an audience, the dancing begins. There are 2 types of bee dances: the round dance and the tail-wagging or waggle dance, with a transitional form known as the sickle dance.
In all cases, the quality and quantity of the food source determine the liveliness of the dances. If the nectar source is of excellent quality, nearly all foragers will dance enthusiastically and at length each time they return from foraging. Food sources of lower quality will produce fewer, shorter, and less vigorous dances; recruiting fewer new foragers.

The round dance is used for food sources 25-100 meters away from the hive or closer. After distributing some of her new-found nectar to waiting bees the scout will begin running in a small circle, switching direction every so often. After the dance ends food is again distributed at this or some other place on the comb and the dance may be repeated three or (rarely) more times.
The round dance does not give directional information. Bees elicited into foraging after a round dance fly out of the hive in all directions searching for the food source they know must be there. Odor helps recruited bees find the new flowers in two ways. Bees watching the dance detect the fragrance of the flower left on the dancing bee. Additionally, the scout bee leaves odor from its scent gland on the flower that helps guide the recruits.

As the food source becomes more distant the round dance is replaced by the waggle dance. There is a gradual transition between the round and waggle dance, taking place through either a figure eight or sickle-shaped pattern.
The waggle dance includes information about the direction and energy required to fly to the goal. Energy expenditure (or distance) is indicated by the length of time it takes to make one circuit. For example, a bee may dance 8-9 circuits in 15 seconds for a food source 200 meters away, 4-5 for a food source 1000 meters away, and 3 circuits in 15 seconds for a food source 2000 meters away.
The direction of the food source is indicated by the direction the dancer faces during the straight portion of the dance when the bee is waggling. If she waggles while facing straight upward, then the food source may be found in the direction of the sun. If she waggles at an angle 60 degrees to the left of upward the food source may be found 60 degrees to the left of the sun.
Similarly, if the dancer waggles 120 degrees to the right of upward, the food source may be found 210 degrees to the right of the sun. The dancer emits sounds during the waggle run that help the recruits determine the direction in the darkness of the hive.


Bibliography:

Aristotle. 1965. Historia Animalium, Loeb Classical Library. Harvard University Press, Massachusetts.

Αριστοτέλης. Των περί τα ζώα ιστοριών. Εκδόσεις Κάκτος. Αθήνα 1992

Frisch, K.V., 1965: Tanzsprache und Orientierung der Bienen. Springer Verlag Berlin/ Heidelberg/ New York, 578 pp.

Frish, K.V. und Lindauer, M., 1954: Himmel und Erde in Konkurrenz bei der Orientierung der Bienen. Naturwiss. 41, 245 – 253

Heymer, A. 1987. Dizionario di etologia. Armando editore, Roma.

Tinbergen, N. 1969. Animal Behaviour. Time Inc., New York

Τσούνης, Γρ. Ιστορίες από τον κόσμο των ζώων, Εκδόσεις Δελφίνι. Αθήνα 1998.