European honey bee
Encyclopedia
The Western honey bee or European honey bee (Apis mellifera) is a species of honey bee
. The genus
Apis is Latin
for "bee", and mellifera comes from Latin melli- "honey" and ferre "to bear"—hence the scientific name means "honey-bearing bee". The name was coined in 1758 by Carolus Linnaeus
who, realizing that the bees do not bear honey, but nectar, tried later to correct it to Apis mellifica ("honey-making bee") in a subsequent publication. However, according to the rules of synonymy
in zoological nomenclature, the older name has precedence. As of October 28, 2006, the Honey Bee Genome Sequencing Consortium
fully sequenced and analyzed the genome of Apis mellifera.
In 2007 media attention focused on colony collapse disorder
, a decline in European honey bee colonies in a minority of regions of North America.
Honeybees differentiated into geographic races as they spread from Africa into Europe and Asia. There are currently 28 recognized subspecies based on these geographic variations. All species are cross fertile though reproductive adaptations may make crossing unlikely. The 28 subspecies can be assigned to one of four major branches based on work by Ruttner and subsequently confirmed by analysis of mitochondrial DNA. African subspecies are assigned to branch A, Northwest European subspecies to branch M, Southwest European subspecies to branch C, and Mideast subspecies to branch O. The subspecies are grouped and listed in the sidebar.
Geographic isolation led to numerous adaptations as honeybees spread after the last ice age. These adaptations include brood cycles synchronized with the bloom period of local flora, forming a winter cluster in colder climates, migratory swarming in Africa, enhanced foraging behavior in desert areas, and numerous other inherited traits.
s survive winter as a colony, and the queen begins egg
laying in mid to late winter, to prepare for spring. This is most likely triggered by longer day length
. She is the only fertile female
, and deposits all the eggs from which the other bees are produced. Except for a brief mating period when she may make several flights to mate with drones, or if she leaves in later life with a swarm to establish a new colony
, the queen rarely leaves the hive after the larva
e have become full grown bees. The queen deposits each egg in a cell prepared by the worker bees. The egg hatches into a small larva which is fed by nurse bees (worker bees who maintain the interior of the colony). After about a week, the larva is sealed up in its cell by the nurse bees and begins the pupal stage. After another week, it will emerge an adult bee.
For the first ten days of their lives, the female worker bees clean the hive and feed the larvae. After this, they begin building comb cells. On days 16 through 20, a worker receives nectar and pollen from older workers and stores it. After the 20th day, a worker leaves the hive and spends the remainder of its life as a forager. The population of a healthy hive in mid-summer can average between 40,000 and 80,000 bees.
The larvae and pupa
e in a frame of honeycomb are referred to as frames of brood and are often sold (with adhering bees) by beekeepers to other beekeepers to start new beehives.
Both workers and queens are fed "royal jelly
" during the first three days of the larval stage. Then workers are switched to a diet of pollen and nectar or diluted honey, while those intended for queens will continue to receive royal jelly. This causes the larva to develop to the pupa stage more quickly, while being also larger and fully developed sexually. Queen breeders consider good nutrition during the larval stage to be of critical importance to the quality of the queens raised, good genetics and sufficient number of mating
s also being factors. During the larval and pupal stages, various parasites can attack the pupa/larva and destroy or damage it.
Queens are not raised in the typical horizontal brood cells of the honeycomb
. The typical queen cell is specially constructed to be much larger, and has a vertical orientation. However, should the workers sense that the old queen is weakening, they will produce emergency cells known as supersedure cells. These cells are made from a cell with an egg or very young larva. These cells protrude from the comb. As the queen finishes her larval feeding, and pupates, she moves into a head downward position, from which she will later chew her way out of the cell. At pupation the workers cap or seal the cell. Just prior to emerging from their cells, young queens can often be heard "piping". The purpose of this sound is not yet fully understood.
Worker bees are infertile females, but in some circumstances, generally only in times of severe stress or with the loss or injury or declining health of the queen, they may lay
infertile eggs, and in some subspecies these eggs may actually be fertile. However, since the worker bees are 'imperfect' females (not fully sexually developed), they do not mate with drones. Any fertile eggs that they lay would be haploid, having only the genetic contribution of their mother, and in honey bees these haploid eggs will always develop into drones. Worker bees also secrete the wax
used to build the hive, clean and maintain the hive, raise the young, guard the hive and forage for nectar and pollen.
In honey bees, the worker bees have a modified ovipositor
called a stinger
with which they can sting to defend the hive, but unlike other bees of any other genus (and even unlike the queens of their own species), the stinger is barbed. Contrary to popular belief, the bee will not always die soon after stinging: this is a misconception based on the fact that a bee will usually die after stinging a human
or other mammal
. The sting and associated venom sac are modified so as to pull free of the body once lodged (autotomy
), and the sting apparatus has its own musculature and ganglion which allow it to keep delivering venom
once detached. It is presumed that this complex apparatus, including the barbs on the sting, evolved specifically in response to predation by vertebrates, as the barbs do not function (and the sting apparatus does not detach) unless the sting is embedded in elastic material. Even then, the barbs do not always "catch", so a bee may occasionally pull the sting free and either fly off unharmed, or sting again.
bees of the colony. Since they do not have ovipositors, they also do not have stingers. Drone honey bees do not forage for nectar or pollen. In some species, drones are suspected of playing a contributing role in the temperature regulation
of the hive. The primary purpose of a drone bee is to fertilize a new queen. Multiple drones will mate with any given queen in flight, and each drone will die immediately after mating; the process of insemination requires a lethally convulsive effort. Drone honey bees are haploid (having single, unpaired chromosomes) in their genetic structure and are descended only from their mother, the queen. They truly do not have a father
. In essence, drones are the equivalent of flying gametes. In regions of temperate climate, the drones are generally expelled from the hive before winter and left to die of cold and starvation, since they are unable to forage or produce honey or take care of themselves.
of the queen in most subspecies is three to four years. However, there are reports that in the German/European Black Bee subspecies that was previously used for beekeeping, the queen was said to live up to 8 years. Because queens deplete their store of sperm, towards the end of their life they start laying more and more unfertilized eggs. Beekeepers therefore frequently change queens every year or every other year.
The lifespan of the workers varies drastically over the year in places with an extended winter. Workers born in the spring and summer will work hard and live only a few weeks, whereas those born in the autumn will stay inside for several months as the colony clusters. On average during the year about one percent of a colony's worker bees die naturally per day. Except for the queen, all of the colony's workers are therefore exchanged about every four months.
Bees produce honey
by collecting nectar, which is a clear liquid consisting of nearly 80% water with complex sugars. The collecting bees store the nectar in a second stomach
and return to the hive where worker bees remove the nectar. The worker bees digest the raw nectar for about 30 minutes using enzyme
s to break up the complex sugars into simpler ones. Raw honey is then spread out in empty honeycomb cells to dry, which reduces the water content to less than 20%. When nectar is being processed, honey bees create a draft through the hive by fanning with their wings
. Once dried, the cells of the honeycomb are sealed (capped) with wax to preserve the honey.
When a hive detects smoke, many bees become remarkably non-aggressive; it is speculated that this is a defense mechanism. Wild colonies generally live in hollow trees, and when bees detect smoke it is presumed that they prepare to evacuate from a forest fire, carrying as much food reserve as they can. In order to do this, they will go to the nearest honey storage cells and gorge on honey. In this state they are quite docile since defense from predation is less important than saving as much food as possible.
, the internal temperature may be as low as 20–22 °C (68–71.6 F).
Honey bees are able to forage over a 30 C-change range of air temperature, largely because they have behavioural and physiological mechanisms for regulating the temperature of their flight muscles. From very low to very high air temperatures, the successive mechanisms are; shivering before flight and stopping flight for additional shivering, passive body temperature regulation in a comfort range that is a function of work effort, and finally, active heat dissipation by evaporative cooling from regurgitated honey sac contents. The body temperatures maintained differ depending on caste and expected foraging rewards.
The optimal air temperature for foraging is 22–25 °C (71.6–77 F). During flight, the rather large flight muscles create heat, which must dissipate. The honey bee uses a form of evaporative cooling to release heat through its mouth. Under hot conditions, heat from the thorax is dissipated through the head; the bee regurgitates a droplet of hot internal fluid — a "honeycrop droplet" – which immediately cools the head temperature by 10 C-change.
Below 7–10 °C (44.6–50 F) bees become immobile and above 38 °C (100.4 °F) bee activity slows. Honey bees can tolerate temperatures up to 50 °C (122 °F) for short periods.
Regardless of the trigger, the workers develop the larvae into queens by continuing to feed them royal jelly which triggers an extended development as a pupa.
When the virgin queen emerges, she is commonly thought to seek out other queen cells and sting the infant queens within. It is also thought that, should two queens emerge simultaneously, they will fight to the death. Recent studies, however, have indicated that as many as 10% of Apis mellifera colonies may maintain two queens. The mechanism by which this occurs is not yet known, but it has been reported to occur more frequently in some South African subspecies of Apis mellifera. Regardless, the queen asserts her control over the worker bees through the release of a complex suite of pheromones called queen scent.
After several days of orientation within and around the hive, the young queen flies to a drone congregation point (a site near a clearing and generally about 30 feet (9.1 m) above the ground) where the drones from different hives tend to congregate in a swirling aerial mass. Drones detect the presence of a queen in their congregation area by her smell, and then find her by sight and mate with her in mid air (drones can be induced to mate with "dummy" queens if they have the queen pheromone applied). A queen will mate multiple times and may leave to mate several days in a row, weather permitting, until her spermatheca
is full.
The queen lays all the eggs in a healthy colony. The number and pace of egg-laying is controlled by weather, availability of resources and the characteristics of the specific race of honey bee. Honey bee queens generally begin to slow egg-laying in the early fall and may even stop during the winter. Egg-laying will generally resume in late winter as soon as the days begin to get longer and peak in the spring. At the height of the season, the queen may lay over 2500 eggs per day – more than her own body mass.
The queen fertilizes each egg as it is being laid into worker size cell using stored sperm from the spermatheca. Eggs, laid into drone size (larger) cells are left unfertilized. The unfertilized eggs have only half as many genes as the queen or worker eggs and develop into drones.
and the mosquito
, to have its genome mapped. According to the scientists who analysed its genetic code, the honey bee originated in Africa and spread to Europe in two ancient migrations. They have also discovered that the number of genes in the honey bees related to smell outnumber those for taste, and they have fewer genes for immunity than the fruit fly and the mosquito. The genome sequence revealed several groups of genes, particularly the genes related to circadian rhythms, were closer to vertebrates than other insects. Genes related to enzymes that control other genes were also vertebrate-like.
The genome is unusual in having very few transposon
s, while they have been present in the evolutionary past (inactive remains were found) and in general evolved slower than in Diptera species.
, alarm, defense
, orientation, kin
and colony
recognition, food production, and integration of colony activities. Pheromones are thus essential to honey bees for their survival.
Honey bees are an excellent animal to study with regard to behavior because they are abundant and familiar to most people. An animal that is disregarded every day has very specific behaviors that go unnoticed by the normal person. Karl von Frisch
, who was awarded the Nobel Prize
for physiology and medicine in 1973 for his study of honey bee communication, noticed that bees communicate through the language of dance. Honey bees are able to direct other bees to food sources through the round dance and the waggle dance
. The round dance tells the other foragers that food is within 50 meters of the hive, but it does not provide much information regarding direction. The waggle dance, which may be vertical or horizontal, provides more detail about both the distance and the direction of the located food source. It is also hypothesized that the bees rely on their olfactory sense
to help locate the food source once the foragers are given directions from the dances.
Another signal for communication is the shaking signal, also known as the jerking dance, vibration dance, or vibration signal. It is a modulatory communication signal because it appears to manipulate the overall arousal or activity of behaviors. The shaking signal is most common in worker communication, but it is also evident in reproductive swarming. A worker bee vibrates its body dorsoventrally while holding another honey bee with its front legs. Jacobus Biesmeijer examined the incidence of shaking signals in a forager’s life and the conditions that led to its performance to investigate why the shaking signal is used in communication for food sources. Biesmeijer found that the experienced foragers executed 92.1% of the observed shaking signals. He also observed that 64% of the shaking signals were executed by experienced foragers after they had discovered a food source. About 71% of the shaking signal sessions occurred after the first five foraging success within one day. Then other communication signals, such as the waggle dance, were performed more often after the first five successes. Biesmeijer proved that most shakers are foragers and that the shaking signal is most often executed by foraging bees over pre-foraging bees. Beismeijer concluded that the shaking signal presents the overall message of transfer work for various activities or activity levels. Sometimes the signal serves to increase activity, when bees shake inactive bees. At other times, the signal serves as an inhibitory mechanism such as the shaking signal at the end of the day. However, the shaking signal is preferentially directed towards inactive bees. All three types of communication between honey bees are effective in their jobs with regards to foraging and task managing.
For a discussion of bees' cognition, response to training, varieties of dance, and use of odors, see Bee learning and communication
.
insect
that is often maintained, fed, and transported by beekeepers
. Honey bees do not survive individually, but rather as part of the colony. Reproduction is also accomplished at the colony level. Colonies are often referred to as superorganism
s.
Honey bees collect flower nectar and convert it to honey
which is stored in their hives. The nectar is transported in the stomach of the bees, and is converted to honey through the addition of various digestive enzyme
s, and by being stored in a "honey cell" and then partially dehydrated. Nectar and honey provide the energy for the bees' flight muscles and for heating the hive during the winter period. Honey bees also collect pollen
which supplies protein and fat for bee brood to grow. Centuries of selective breeding
by humans have created honey bees that produce far more honey than the colony needs. Beekeepers
, also known as "apiarists," harvest the honey.
Beekeepers often provide a place for the colony to live and to store honey. There are seven basic types of beehive
: skeps, Langstroth hive
s, top-bar hive
s, box hives, log gums, D.E. hives and miller hives. All U.S. states require beekeepers to use movable frames to allow bee inspectors to check the brood for disease. This allows beekeepers to keep the Langstroth, top-bar, and D.E. hives freely, but other types of hives require special permitting, such as for museum use. The type of beehive used significantly impacts colony health and wax and honey production.
Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide.
In cold climates some beekeepers have kept colonies alive (with varying success) by moving them indoors for winter. While this can protect the colonies from extremes of temperature and make winter care and feeding more convenient for the beekeeper, it can increase the risk of dysentery
(see the Nosema section of diseases of the honey bee) and can create an excessive buildup of carbon dioxide
from the respiration of the bees. Recently, inside wintering has been refined by Canadian beekeepers, who build large barns just for wintering bees. Automated ventilation systems assist in the control of carbon dioxide build-up.
1. Egg Laying Rate – Queen can lay a few hundred eggs per day up to about 5000 eggs per day.
2. Egg Viability Rate – Can range from 0 up to 100% of eggs that hatch and develop into bees.
3. Brood Cycle Length – For worker bees ranges from 17 days up to 21 days.
4. Brood Nurturing – This is a measure of how intent the nurse bees are at nurturing brood.
5. Foraging Aggressiveness – This trait determines potential for honey production.
6. Time Of Foraging – Some bees forage earlier in the day and later in the evening than others.
7. Disease Resistance – This is a measure of innate tolerance to brood diseases and adult diseases.
8. Pest Resistance – This is a measure of innate tolerance to pests such as tracheal and varroa mites.
9. Defensive Behavior – Determines aggressiveness and stinging propensity.
10. Swarming Tendency – This determines the timing and success of colony reproduction.
11. Winter Hardiness – Clustering behavior and ability to survive extended low temperatures.
12. Life Span – Ranges from a 22 days up to 305 days for workers with an average of 36 days.
13. Body Size – Small bees are typical in Africa with larger bees typical in colder climates.
14. Sense Of Smell – Ability to detect flower odors and respond to nectar availability.
15. Hygienic Cleaning Behavior – This is a major component of both disease and pest resistance.
16. Time Of Brood Development – Bees must begin brood rearing at least 8 weeks before nectar flows.
17. Thrift – Adjustment of brood production to available food sources for efficient use of resources.
18. Honey Arrangement – Storing honey closer to or further away from the brood next.
19. Pollen Collection – Amount and floral source of pollen are under direct genetic control.
20. Type Of nectar Collected – This has a major impact on honey quality and value.
21. Comb Building – Willingness to build comb and expand the colony.
22. Capping Structure – Ranges from flat and watery to gray to white dome shaped cappings.
23. Propolis Collection – Associated with wintering success, ranges from none to covering all hive surfaces.
24. Brace Comb Construction – Ranges from very little to cross comb throughout the colony.
25. Abdominal Color – Ranges from black to various color stripes in tan, yellow, and orange.
26. Antennae Structure – Number and placement of sensors is inherited, associated with sense of smell.
27. Number of Drones Produced – Ranges from very few up to ¼ of brood combs.
28. Number of Queens Produced – Ranges from a few up to several hundred.
The honey bee's primary commercial value is as a pollinator
of crops. Orchards and fields have grown larger; at the same time wild pollinators have dwindled
. In several areas of the world the pollination shortage is compensated by migratory beekeeping, with beekeepers supplying the hives during the crop bloom and moving them after bloom is complete. In many higher latitude locations it is difficult or impossible to winter over enough bees, or at least to have them ready for early blooming plants, so much of the migration is seasonal, with many hives wintering in warmer climates and moving to follow the bloom to higher latitudes.
As an example, in California
, the pollination of almond
s occurs in February, early in the growing season
, before local hives have built up their populations. Almond orchard
s require two hives per acre
(2,000 m² per hive) for maximum yield and so the pollination is highly dependent upon the importation of hives from warmer climates. Almond pollination, which occurs in February and March, is the largest managed pollination event in the world, requiring more than one third of all the managed honey bees in the United States. Massive movement of honey bee are also made for apples in New York
, Michigan
, and Washington. And despite the inefficiency of honey bees in pollinating blueberries, huge numbers are also moved to Maine for blueberries, because they are the only pollinators that can be relatively easily moved and concentrated for this and other monoculture
crops.
Commercial beekeepers plan their movements and their wintering locations with prime reference to the pollination services they plan to perform.
Bees, as well as some other insects, are particularly beneficial as pollinators to most plants, as they maintain flower constancy
, which means they are more likely to transfer pollen to other conspecific plants. Also, flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species.
s and seals
.
and carry it back to the hive. There, pollen is used as a protein
source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives. It is often eaten as a health supplement.
n and European populations were severely depleted by varroa mite infestations in the early 1990s, and US
beekeepers were further affected by colony collapse disorder
in 2006 and 2007. Chemical treatments against Varroa mites saved most commercial operations and improved cultural practices. New bee breeds are starting to reduce the dependency on miticides (acaracides) by beekeepers. Feral bee populations were greatly reduced during this period but now are slowly recovering, mostly in areas of mild climate, owing to natural selection
for Varroa resistance and repopulation by resistant breeds. Further, insecticides, particularly when used in violation of label directions, have also depleted bee populations, while various bee pests and diseases are becoming resistant to medications (e.g. American foulbrood, tracheal mites and Varroa mites
).
s have spread across the southern United States
where they pose a small danger to humans, although they may make beekeeping difficult and potentially dangerous (particularly hobby beekeeping).
As an invasive species
, feral honey bees have become a significant environmental problem in places where they are not native. Imported bees may compete with and displace native bees and birds, and may also promote the reproduction of invasive plants
that native pollinators do not visit. Also, unlike native bees, they do not properly extract or transfer pollen from plants with poricidal anthers
(anthers that only release pollen through tiny apical pores), as this requires buzz pollination
, a behavior which honey bees rarely exhibit. For example, honey bees reduce fruiting in Melastoma affine
(a plant with poricidal anthers) by robbing stigmas of previously deposited pollen.
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
. The genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Apis is Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
for "bee", and mellifera comes from Latin melli- "honey" and ferre "to bear"—hence the scientific name means "honey-bearing bee". The name was coined in 1758 by Carolus Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...
who, realizing that the bees do not bear honey, but nectar, tried later to correct it to Apis mellifica ("honey-making bee") in a subsequent publication. However, according to the rules of synonymy
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
in zoological nomenclature, the older name has precedence. As of October 28, 2006, the Honey Bee Genome Sequencing Consortium
Honey Bee Genome Sequencing Consortium
The Honey Bee Genome Sequencing Consortium is an international collaborative group of genomics scientists, scientific organisations and universities who are trying to decipher the genome sequences of the honey bee . It was formed in 2001 by American scientists...
fully sequenced and analyzed the genome of Apis mellifera.
In 2007 media attention focused on colony collapse disorder
Colony Collapse Disorder
Colony collapse disorder is a phenomenon in which worker bees from a beehive or European honey bee colony abruptly disappear. While such disappearances have occurred throughout the history of apiculture, the term colony collapse disorder was first applied to a drastic rise in the number of...
, a decline in European honey bee colonies in a minority of regions of North America.
Geographic distribution
The honey bee is native to the continents of Europe, Asia, and Africa. As of the early 1600s, the insect was introduced to North America, with subsequent introductions of other European subspecies two centuries later. Since then, they have spread throughout the Americas.Honeybees differentiated into geographic races as they spread from Africa into Europe and Asia. There are currently 28 recognized subspecies based on these geographic variations. All species are cross fertile though reproductive adaptations may make crossing unlikely. The 28 subspecies can be assigned to one of four major branches based on work by Ruttner and subsequently confirmed by analysis of mitochondrial DNA. African subspecies are assigned to branch A, Northwest European subspecies to branch M, Southwest European subspecies to branch C, and Mideast subspecies to branch O. The subspecies are grouped and listed in the sidebar.
Geographic isolation led to numerous adaptations as honeybees spread after the last ice age. These adaptations include brood cycles synchronized with the bloom period of local flora, forming a winter cluster in colder climates, migratory swarming in Africa, enhanced foraging behavior in desert areas, and numerous other inherited traits.
Biology, life cycle
In the temperate zone, honey beeHoney bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
s survive winter as a colony, and the queen begins egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
laying in mid to late winter, to prepare for spring. This is most likely triggered by longer day length
Day length
Day length, or length of day, or length of daytime, refers to the time each day from the moment the upper limb of the sun's disk appears above the horizon during sunrise to the moment when the upper limb disappears below the horizon during sunset...
. She is the only fertile female
Female
Female is the sex of an organism, or a part of an organism, which produces non-mobile ova .- Defining characteristics :The ova are defined as the larger gametes in a heterogamous reproduction system, while the smaller, usually motile gamete, the spermatozoon, is produced by the male...
, and deposits all the eggs from which the other bees are produced. Except for a brief mating period when she may make several flights to mate with drones, or if she leaves in later life with a swarm to establish a new colony
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...
, the queen rarely leaves the hive after the larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e have become full grown bees. The queen deposits each egg in a cell prepared by the worker bees. The egg hatches into a small larva which is fed by nurse bees (worker bees who maintain the interior of the colony). After about a week, the larva is sealed up in its cell by the nurse bees and begins the pupal stage. After another week, it will emerge an adult bee.
The larvae and pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
e in a frame of honeycomb are referred to as frames of brood and are often sold (with adhering bees) by beekeepers to other beekeepers to start new beehives.
Both workers and queens are fed "royal jelly
Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae, as well as adult queens. It is secreted from the glands in the hypopharynx of worker bees, and fed to all larvae in the colony....
" during the first three days of the larval stage. Then workers are switched to a diet of pollen and nectar or diluted honey, while those intended for queens will continue to receive royal jelly. This causes the larva to develop to the pupa stage more quickly, while being also larger and fully developed sexually. Queen breeders consider good nutrition during the larval stage to be of critical importance to the quality of the queens raised, good genetics and sufficient number of mating
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...
s also being factors. During the larval and pupal stages, various parasites can attack the pupa/larva and destroy or damage it.
Honeycomb
A honeycomb is a mass of hexagonal waxcells built by honey bees in their nests to contain their larvae and stores of honey and pollen.Beekeepers may remove the entire honeycomb to harvest honey...
. The typical queen cell is specially constructed to be much larger, and has a vertical orientation. However, should the workers sense that the old queen is weakening, they will produce emergency cells known as supersedure cells. These cells are made from a cell with an egg or very young larva. These cells protrude from the comb. As the queen finishes her larval feeding, and pupates, she moves into a head downward position, from which she will later chew her way out of the cell. At pupation the workers cap or seal the cell. Just prior to emerging from their cells, young queens can often be heard "piping". The purpose of this sound is not yet fully understood.
Worker bees are infertile females, but in some circumstances, generally only in times of severe stress or with the loss or injury or declining health of the queen, they may lay
Laying worker bee
A laying worker bee is a worker bee that lays unfertilized eggs usually in the absence of a queen bee. Only drones develop from the eggs of laying worker bees . A beehive cannot survive with only a laying worker bee....
infertile eggs, and in some subspecies these eggs may actually be fertile. However, since the worker bees are 'imperfect' females (not fully sexually developed), they do not mate with drones. Any fertile eggs that they lay would be haploid, having only the genetic contribution of their mother, and in honey bees these haploid eggs will always develop into drones. Worker bees also secrete the wax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...
used to build the hive, clean and maintain the hive, raise the young, guard the hive and forage for nectar and pollen.
Ovipositor
The ovipositor is an organ used by some animals for oviposition, i.e., the laying of eggs. It consists of a maximum of three pairs of appendages formed to transmit the egg, to prepare a place for it, and to place it properly...
called a stinger
Stinger (organ)
A sting, sometimes called a stinger in the US, is a sharp organ or body part found in various animals that delivers some kind of venom . A true sting differs from other piercing structures in that it pierces by its own action and injects venom, as opposed to teeth, which pierce by the force of...
with which they can sting to defend the hive, but unlike other bees of any other genus (and even unlike the queens of their own species), the stinger is barbed. Contrary to popular belief, the bee will not always die soon after stinging: this is a misconception based on the fact that a bee will usually die after stinging a human
Human
Humans are the only living species in the Homo genus...
or other mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
. The sting and associated venom sac are modified so as to pull free of the body once lodged (autotomy
Autotomy
Autotomy or self amputation is the act whereby an animal severs one or more of its own appendages, usually as a self-defense mechanism designed to elude a predator's grasp...
), and the sting apparatus has its own musculature and ganglion which allow it to keep delivering venom
Venom
Venom is the general term referring to any variety of toxins used by certain types of animals that inject it into their victims by the means of a bite or a sting...
once detached. It is presumed that this complex apparatus, including the barbs on the sting, evolved specifically in response to predation by vertebrates, as the barbs do not function (and the sting apparatus does not detach) unless the sting is embedded in elastic material. Even then, the barbs do not always "catch", so a bee may occasionally pull the sting free and either fly off unharmed, or sting again.
Drones
Drone bees are the maleMale
Male refers to the biological sex of an organism, or part of an organism, which produces small mobile gametes, called spermatozoa. Each spermatozoon can fuse with a larger female gamete or ovum, in the process of fertilization...
bees of the colony. Since they do not have ovipositors, they also do not have stingers. Drone honey bees do not forage for nectar or pollen. In some species, drones are suspected of playing a contributing role in the temperature regulation
Thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different...
of the hive. The primary purpose of a drone bee is to fertilize a new queen. Multiple drones will mate with any given queen in flight, and each drone will die immediately after mating; the process of insemination requires a lethally convulsive effort. Drone honey bees are haploid (having single, unpaired chromosomes) in their genetic structure and are descended only from their mother, the queen. They truly do not have a father
Father
A father, Pop, Dad, or Papa, is defined as a male parent of any type of offspring. The adjective "paternal" refers to father, parallel to "maternal" for mother...
. In essence, drones are the equivalent of flying gametes. In regions of temperate climate, the drones are generally expelled from the hive before winter and left to die of cold and starvation, since they are unable to forage or produce honey or take care of themselves.
Life expectancy
The average lifespanLife expectancy
Life expectancy is the expected number of years of life remaining at a given age. It is denoted by ex, which means the average number of subsequent years of life for someone now aged x, according to a particular mortality experience...
of the queen in most subspecies is three to four years. However, there are reports that in the German/European Black Bee subspecies that was previously used for beekeeping, the queen was said to live up to 8 years. Because queens deplete their store of sperm, towards the end of their life they start laying more and more unfertilized eggs. Beekeepers therefore frequently change queens every year or every other year.
The lifespan of the workers varies drastically over the year in places with an extended winter. Workers born in the spring and summer will work hard and live only a few weeks, whereas those born in the autumn will stay inside for several months as the colony clusters. On average during the year about one percent of a colony's worker bees die naturally per day. Except for the queen, all of the colony's workers are therefore exchanged about every four months.
Honey production
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
by collecting nectar, which is a clear liquid consisting of nearly 80% water with complex sugars. The collecting bees store the nectar in a second stomach
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
and return to the hive where worker bees remove the nectar. The worker bees digest the raw nectar for about 30 minutes using enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s to break up the complex sugars into simpler ones. Raw honey is then spread out in empty honeycomb cells to dry, which reduces the water content to less than 20%. When nectar is being processed, honey bees create a draft through the hive by fanning with their wings
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
. Once dried, the cells of the honeycomb are sealed (capped) with wax to preserve the honey.
When a hive detects smoke, many bees become remarkably non-aggressive; it is speculated that this is a defense mechanism. Wild colonies generally live in hollow trees, and when bees detect smoke it is presumed that they prepare to evacuate from a forest fire, carrying as much food reserve as they can. In order to do this, they will go to the nearest honey storage cells and gorge on honey. In this state they are quite docile since defense from predation is less important than saving as much food as possible.
Thermoregulation
The honey bee needs an internal body temperature of 35 °C (95 °F) to fly — which is also the temperature maintained within the cluster. The same temperature is required in the brood nest over a long period to develop the brood, and it is the optimal temperature for the creation of wax. The temperature on the periphery of the cluster varies with the outside air temperature and in the winter clusterWinter cluster
In beekeeping, a winter cluster is a well-defined cluster of honey bees that forms inside a beehive when the air temperature dips below 54 to 57 °F . Honey bees are but a few insects that survive the winter as a hive. As the outside air temperature decreases the winter cluster becomes tighter and...
, the internal temperature may be as low as 20–22 °C (68–71.6 F).
Honey bees are able to forage over a 30 C-change range of air temperature, largely because they have behavioural and physiological mechanisms for regulating the temperature of their flight muscles. From very low to very high air temperatures, the successive mechanisms are; shivering before flight and stopping flight for additional shivering, passive body temperature regulation in a comfort range that is a function of work effort, and finally, active heat dissipation by evaporative cooling from regurgitated honey sac contents. The body temperatures maintained differ depending on caste and expected foraging rewards.
The optimal air temperature for foraging is 22–25 °C (71.6–77 F). During flight, the rather large flight muscles create heat, which must dissipate. The honey bee uses a form of evaporative cooling to release heat through its mouth. Under hot conditions, heat from the thorax is dissipated through the head; the bee regurgitates a droplet of hot internal fluid — a "honeycrop droplet" – which immediately cools the head temperature by 10 C-change.
Below 7–10 °C (44.6–50 F) bees become immobile and above 38 °C (100.4 °F) bee activity slows. Honey bees can tolerate temperatures up to 50 °C (122 °F) for short periods.
Queens
Periodically, the colony determines that a new queen is needed. There are three general triggers.- The colony becomes space-constrained because the hive is filled with honey, leaving little room for new eggs. This will trigger a swarm where the old queen will take about half the worker bees to found a new colony, leaving the new queen with the other half of worker bees to continue the old colony.
- The old queen begins to fail — this is thought to be recognized by a decrease in queen pheromones throughout the hive. This situation is called 'supersedure; at the end of the supersedure, the old queen is generally killed.
- The old queen dies suddenly — this situation is known as 'emergency supersedure'. The worker bees will find several eggs or larvae of the right age-range and attempt to develop them into queens. Emergency supersedure can generally be recognized because the new queen cells are built out from regular cells of the comb rather than hanging from the bottom of a frame.
Regardless of the trigger, the workers develop the larvae into queens by continuing to feed them royal jelly which triggers an extended development as a pupa.
When the virgin queen emerges, she is commonly thought to seek out other queen cells and sting the infant queens within. It is also thought that, should two queens emerge simultaneously, they will fight to the death. Recent studies, however, have indicated that as many as 10% of Apis mellifera colonies may maintain two queens. The mechanism by which this occurs is not yet known, but it has been reported to occur more frequently in some South African subspecies of Apis mellifera. Regardless, the queen asserts her control over the worker bees through the release of a complex suite of pheromones called queen scent.
Spermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...
is full.
The queen lays all the eggs in a healthy colony. The number and pace of egg-laying is controlled by weather, availability of resources and the characteristics of the specific race of honey bee. Honey bee queens generally begin to slow egg-laying in the early fall and may even stop during the winter. Egg-laying will generally resume in late winter as soon as the days begin to get longer and peak in the spring. At the height of the season, the queen may lay over 2500 eggs per day – more than her own body mass.
The queen fertilizes each egg as it is being laid into worker size cell using stored sperm from the spermatheca. Eggs, laid into drone size (larger) cells are left unfertilized. The unfertilized eggs have only half as many genes as the queen or worker eggs and develop into drones.
Genome
The European honey bee is the third insect, after the fruit flyDrosophila melanogaster
Drosophila melanogaster is a species of Diptera, or the order of flies, in the family Drosophilidae. The species is known generally as the common fruit fly or vinegar fly. Starting from Charles W...
and the mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...
, to have its genome mapped. According to the scientists who analysed its genetic code, the honey bee originated in Africa and spread to Europe in two ancient migrations. They have also discovered that the number of genes in the honey bees related to smell outnumber those for taste, and they have fewer genes for immunity than the fruit fly and the mosquito. The genome sequence revealed several groups of genes, particularly the genes related to circadian rhythms, were closer to vertebrates than other insects. Genes related to enzymes that control other genes were also vertebrate-like.
The genome is unusual in having very few transposon
Transposon
Transposable elements are sequences of DNA that can move or transpose themselves to new positions within the genome of a single cell. The mechanism of transposition can be either "copy and paste" or "cut and paste". Transposition can create phenotypically significant mutations and alter the cell's...
s, while they have been present in the evolutionary past (inactive remains were found) and in general evolved slower than in Diptera species.
Pheromones
Honey bees use special pheromones, or chemical communication, for almost all behaviors of life. Such uses include (but are not limited to): matingMating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...
, alarm, defense
Self-defense
Self-defense, self-defence or private defense is a countermeasure that involves defending oneself, one's property or the well-being of another from physical harm. The use of the right of self-defense as a legal justification for the use of force in times of danger is available in many...
, orientation, kin
Kin
-Places:* Kin, Okinawa, a town in Okinawa, Japan* Kin, Pakistan, a village along the Indus in Pakistan* Kin, Mogok, a village in Mogok Township, Burma * Kin, Ye, a village in Ye Township, Burma...
and colony
Colony
In politics and history, a colony is a territory under the immediate political control of a state. For colonies in antiquity, city-states would often found their own colonies. Some colonies were historically countries, while others were territories without definite statehood from their inception....
recognition, food production, and integration of colony activities. Pheromones are thus essential to honey bees for their survival.
Communication
Karl von Frisch
Karl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....
, who was awarded the Nobel Prize
Nobel Prize
The Nobel Prizes are annual international awards bestowed by Scandinavian committees in recognition of cultural and scientific advances. The will of the Swedish chemist Alfred Nobel, the inventor of dynamite, established the prizes in 1895...
for physiology and medicine in 1973 for his study of honey bee communication, noticed that bees communicate through the language of dance. Honey bees are able to direct other bees to food sources through the round dance and the waggle dance
Waggle dance
Waggle dance is a term used in beekeeping and ethology for a particular figure-eight dance of the honey bee. By performing this dance, successful foragers can share with their hive mates information about the direction and distance to patches of flowers yielding nectar and pollen, to water...
. The round dance tells the other foragers that food is within 50 meters of the hive, but it does not provide much information regarding direction. The waggle dance, which may be vertical or horizontal, provides more detail about both the distance and the direction of the located food source. It is also hypothesized that the bees rely on their olfactory sense
Olfaction
Olfaction is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates...
to help locate the food source once the foragers are given directions from the dances.
Another signal for communication is the shaking signal, also known as the jerking dance, vibration dance, or vibration signal. It is a modulatory communication signal because it appears to manipulate the overall arousal or activity of behaviors. The shaking signal is most common in worker communication, but it is also evident in reproductive swarming. A worker bee vibrates its body dorsoventrally while holding another honey bee with its front legs. Jacobus Biesmeijer examined the incidence of shaking signals in a forager’s life and the conditions that led to its performance to investigate why the shaking signal is used in communication for food sources. Biesmeijer found that the experienced foragers executed 92.1% of the observed shaking signals. He also observed that 64% of the shaking signals were executed by experienced foragers after they had discovered a food source. About 71% of the shaking signal sessions occurred after the first five foraging success within one day. Then other communication signals, such as the waggle dance, were performed more often after the first five successes. Biesmeijer proved that most shakers are foragers and that the shaking signal is most often executed by foraging bees over pre-foraging bees. Beismeijer concluded that the shaking signal presents the overall message of transfer work for various activities or activity levels. Sometimes the signal serves to increase activity, when bees shake inactive bees. At other times, the signal serves as an inhibitory mechanism such as the shaking signal at the end of the day. However, the shaking signal is preferentially directed towards inactive bees. All three types of communication between honey bees are effective in their jobs with regards to foraging and task managing.
- "The general story of the communication of the distance, the situation, and the direction of a food source by the dances of the returning (honey bee) worker bee on the vertical comb of the hive, has been known in general outline from the work of Karl von FrischKarl von FrischKarl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....
in the middle 1950s."
For a discussion of bees' cognition, response to training, varieties of dance, and use of odors, see Bee learning and communication
Bee learning and communication
Honey bees learn and communicate in order to find food sources and for other means.-Learning:Learning is essential for efficient foraging. Honey bees are unlikely to make many repeat visits if a plant provides little in the way of reward...
.
Beekeeping
The honey bee is a colonialColony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...
insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
that is often maintained, fed, and transported by beekeepers
Beekeeping
Beekeeping is the maintenance of honey bee colonies, commonly in hives, by humans. A beekeeper keeps bees in order to collect honey and other products of the hive , to pollinate crops, or to produce bees for sale to other beekeepers...
. Honey bees do not survive individually, but rather as part of the colony. Reproduction is also accomplished at the colony level. Colonies are often referred to as superorganism
Superorganism
A superorganism is an organism consisting of many organisms. This is usually meant to be a social unit of eusocial animals, where division of labour is highly specialised and where individuals are not able to survive by themselves for extended periods of time. Ants are the best-known example of...
s.
Honey bees collect flower nectar and convert it to honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
which is stored in their hives. The nectar is transported in the stomach of the bees, and is converted to honey through the addition of various digestive enzyme
Digestive enzyme
'Digestive enzymes' are enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption by the body. Digestive enzymes are found in the digestive tract of animals where they aid in the digestion of food as well as inside the cells,...
s, and by being stored in a "honey cell" and then partially dehydrated. Nectar and honey provide the energy for the bees' flight muscles and for heating the hive during the winter period. Honey bees also collect pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
which supplies protein and fat for bee brood to grow. Centuries of selective breeding
Selective breeding
Selective breeding is the process of breeding plants and animals for particular genetic traits. Typically, strains that are selectively bred are domesticated, and the breeding is sometimes done by a professional breeder. Bred animals are known as breeds, while bred plants are known as varieties,...
by humans have created honey bees that produce far more honey than the colony needs. Beekeepers
Beekeeping
Beekeeping is the maintenance of honey bee colonies, commonly in hives, by humans. A beekeeper keeps bees in order to collect honey and other products of the hive , to pollinate crops, or to produce bees for sale to other beekeepers...
, also known as "apiarists," harvest the honey.
Beekeepers often provide a place for the colony to live and to store honey. There are seven basic types of beehive
Beehive (beekeeping)
A beehive is an enclosed structure in which some honey bee species of the subgenus Apis live and raise their young. Natural beehives are naturally occurring structures occupied by honeybee colonies, while domesticated honeybees live in man-made beehives, often in an apiary. These man-made...
: skeps, Langstroth hive
Langstroth hive
The Langstroth bee hive, patented in October 1852, is the standard beehive used in many parts of the world for beekeeping. The advantage of this hive is that the bees build honeycomb into frames, which can be moved with ease. The frames are designed to prevent bees from attaching honeycombs where...
s, top-bar hive
Top-bar hive
Top-bar hives are a style of beehive used for beekeeping. They are especially useful in areas where resources are limited, but are also increasingly popular among hobby beekeepers in industrialized nations...
s, box hives, log gums, D.E. hives and miller hives. All U.S. states require beekeepers to use movable frames to allow bee inspectors to check the brood for disease. This allows beekeepers to keep the Langstroth, top-bar, and D.E. hives freely, but other types of hives require special permitting, such as for museum use. The type of beehive used significantly impacts colony health and wax and honey production.
Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide.
In cold climates some beekeepers have kept colonies alive (with varying success) by moving them indoors for winter. While this can protect the colonies from extremes of temperature and make winter care and feeding more convenient for the beekeeper, it can increase the risk of dysentery
Dysentery
Dysentery is an inflammatory disorder of the intestine, especially of the colon, that results in severe diarrhea containing mucus and/or blood in the faeces with fever and abdominal pain. If left untreated, dysentery can be fatal.There are differences between dysentery and normal bloody diarrhoea...
(see the Nosema section of diseases of the honey bee) and can create an excessive buildup of carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
from the respiration of the bees. Recently, inside wintering has been refined by Canadian beekeepers, who build large barns just for wintering bees. Automated ventilation systems assist in the control of carbon dioxide build-up.
Breeding
Numerous traits are present in the various subspecies of honeybees that are amenable to selection via breeding. This listing covers a small part of the variation present in the honeybee genome.1. Egg Laying Rate – Queen can lay a few hundred eggs per day up to about 5000 eggs per day.
2. Egg Viability Rate – Can range from 0 up to 100% of eggs that hatch and develop into bees.
3. Brood Cycle Length – For worker bees ranges from 17 days up to 21 days.
4. Brood Nurturing – This is a measure of how intent the nurse bees are at nurturing brood.
5. Foraging Aggressiveness – This trait determines potential for honey production.
6. Time Of Foraging – Some bees forage earlier in the day and later in the evening than others.
7. Disease Resistance – This is a measure of innate tolerance to brood diseases and adult diseases.
8. Pest Resistance – This is a measure of innate tolerance to pests such as tracheal and varroa mites.
9. Defensive Behavior – Determines aggressiveness and stinging propensity.
10. Swarming Tendency – This determines the timing and success of colony reproduction.
11. Winter Hardiness – Clustering behavior and ability to survive extended low temperatures.
12. Life Span – Ranges from a 22 days up to 305 days for workers with an average of 36 days.
13. Body Size – Small bees are typical in Africa with larger bees typical in colder climates.
14. Sense Of Smell – Ability to detect flower odors and respond to nectar availability.
15. Hygienic Cleaning Behavior – This is a major component of both disease and pest resistance.
16. Time Of Brood Development – Bees must begin brood rearing at least 8 weeks before nectar flows.
17. Thrift – Adjustment of brood production to available food sources for efficient use of resources.
18. Honey Arrangement – Storing honey closer to or further away from the brood next.
19. Pollen Collection – Amount and floral source of pollen are under direct genetic control.
20. Type Of nectar Collected – This has a major impact on honey quality and value.
21. Comb Building – Willingness to build comb and expand the colony.
22. Capping Structure – Ranges from flat and watery to gray to white dome shaped cappings.
23. Propolis Collection – Associated with wintering success, ranges from none to covering all hive surfaces.
24. Brace Comb Construction – Ranges from very little to cross comb throughout the colony.
25. Abdominal Color – Ranges from black to various color stripes in tan, yellow, and orange.
26. Antennae Structure – Number and placement of sensors is inherited, associated with sense of smell.
27. Number of Drones Produced – Ranges from very few up to ¼ of brood combs.
28. Number of Queens Produced – Ranges from a few up to several hundred.
Pollination
Pollinator
A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain...
of crops. Orchards and fields have grown larger; at the same time wild pollinators have dwindled
Pollinator decline
The term pollinator decline refers to the reduction in abundance of pollinators in many ecosystems worldwide during the end of the twentieth century....
. In several areas of the world the pollination shortage is compensated by migratory beekeeping, with beekeepers supplying the hives during the crop bloom and moving them after bloom is complete. In many higher latitude locations it is difficult or impossible to winter over enough bees, or at least to have them ready for early blooming plants, so much of the migration is seasonal, with many hives wintering in warmer climates and moving to follow the bloom to higher latitudes.
As an example, in California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...
, the pollination of almond
Almond
The almond , is a species of tree native to the Middle East and South Asia. Almond is also the name of the edible and widely cultivated seed of this tree...
s occurs in February, early in the growing season
Growing season
In botany, horticulture, and agriculture the growing season is the period of each year when native plants and ornamental plants grow; and when crops can be grown....
, before local hives have built up their populations. Almond orchard
Orchard
An orchard is an intentional planting of trees or shrubs that is maintained for food production. Orchards comprise fruit or nut-producing trees which are grown for commercial production. Orchards are also sometimes a feature of large gardens, where they serve an aesthetic as well as a productive...
s require two hives per acre
Acre
The acre is a unit of area in a number of different systems, including the imperial and U.S. customary systems. The most commonly used acres today are the international acre and, in the United States, the survey acre. The most common use of the acre is to measure tracts of land.The acre is related...
(2,000 m² per hive) for maximum yield and so the pollination is highly dependent upon the importation of hives from warmer climates. Almond pollination, which occurs in February and March, is the largest managed pollination event in the world, requiring more than one third of all the managed honey bees in the United States. Massive movement of honey bee are also made for apples in New York
New York
New York is a state in the Northeastern region of the United States. It is the nation's third most populous state. New York is bordered by New Jersey and Pennsylvania to the south, and by Connecticut, Massachusetts and Vermont to the east...
, Michigan
Michigan
Michigan is a U.S. state located in the Great Lakes Region of the United States of America. The name Michigan is the French form of the Ojibwa word mishigamaa, meaning "large water" or "large lake"....
, and Washington. And despite the inefficiency of honey bees in pollinating blueberries, huge numbers are also moved to Maine for blueberries, because they are the only pollinators that can be relatively easily moved and concentrated for this and other monoculture
Monoculture
Monoculture is the agricultural practice of producing or growing one single crop over a wide area. It is also known as a way of farming practice of growing large stands of a single species. It is widely used in modern industrial agriculture and its implementation has allowed for large harvests from...
crops.
Commercial beekeepers plan their movements and their wintering locations with prime reference to the pollination services they plan to perform.
Bees, as well as some other insects, are particularly beneficial as pollinators to most plants, as they maintain flower constancy
Flower constancy
Flower constancy or pollinator constancy is defined as the tendency of individual pollinators to exclusively visit certain flower species or morphs within a species, bypassing other available flower species that could potentially be more rewarding...
, which means they are more likely to transfer pollen to other conspecific plants. Also, flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species.
Honey
Honey is the complex substance made when the nectar and sweet deposits from plants and trees are gathered, modified and stored in the honeycomb by honey bees. Honey is a complex biological mixture that consists mostly of inverted sugars, primarily glucose and fructose. It has antibacterial and antifungal properties and will not rot or ferment when stored under normal conditions. However, honey will crystallize with time. Crystallized honey is not damaged or defective in any way, for human use, but bees will automatically remove crystallized honey from their hive and discard it, since they can only use liquid honey.Beeswax
Worker bees of a certain age will secrete beeswax from a series of glands on their abdomen. They use the wax to form the walls and caps of the comb. When honey is harvested, the wax can be gathered to be used in various wax products like candleCandle
A candle is a solid block or cylinder of wax with an embedded wick, which is lit to provide light, and sometimes heat.Today, most candles are made from paraffin. Candles can also be made from beeswax, soy, other plant waxes, and tallow...
s and seals
Seal (device)
A seal can be a figure impressed in wax, clay, or some other medium, or embossed on paper, with the purpose of authenticating a document ; but the term can also mean the device for making such impressions, being essentially a mould with the mirror image of the design carved in sunken- relief or...
.
Pollen
Bees collect pollen in the pollen basketPollen basket
The pollen basket or corbicula is part of the tibia on the hind legs of the four related lineages of apid bees that used to comprise the family Apidae: the honey bees, bumblebees, stingless bees, and orchid bees...
and carry it back to the hive. There, pollen is used as a protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives. It is often eaten as a health supplement.
Propolis
Propolis is a resinous mixture that honey bees collect from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the hive. Propolis is marketed for its alleged health benefits, but may cause severe allergic reactions in some individuals.Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of the larvae. It is marketed for its alleged health benefits, but may cause severe allergic reactions in some individuals.Hazards and survival
European honey bee populations have recently faced threats to their survival. North AmericaNorth America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
n and European populations were severely depleted by varroa mite infestations in the early 1990s, and US
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
beekeepers were further affected by colony collapse disorder
Colony Collapse Disorder
Colony collapse disorder is a phenomenon in which worker bees from a beehive or European honey bee colony abruptly disappear. While such disappearances have occurred throughout the history of apiculture, the term colony collapse disorder was first applied to a drastic rise in the number of...
in 2006 and 2007. Chemical treatments against Varroa mites saved most commercial operations and improved cultural practices. New bee breeds are starting to reduce the dependency on miticides (acaracides) by beekeepers. Feral bee populations were greatly reduced during this period but now are slowly recovering, mostly in areas of mild climate, owing to natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
for Varroa resistance and repopulation by resistant breeds. Further, insecticides, particularly when used in violation of label directions, have also depleted bee populations, while various bee pests and diseases are becoming resistant to medications (e.g. American foulbrood, tracheal mites and Varroa mites
Varroa
Varroa is a genus of parasitic mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar who was also a beekeeper.-History and behavior:...
).
Environmental hazards
In North America, Africanized beeAfricanized bee
Africanized honey bees, known colloquially as "killer bees", are a hybrid variety of the European honeybee , generated by a man-made breeding of the African honey bee, A. m. scutellata, with various European honey bees such as the Italian bee A. m. ligustica and A. m. iberiensis. These bees are far...
s have spread across the southern United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
where they pose a small danger to humans, although they may make beekeeping difficult and potentially dangerous (particularly hobby beekeeping).
As an invasive species
Invasive species
"Invasive species", or invasive exotics, is a nomenclature term and categorization phrase used for flora and fauna, and for specific restoration-preservation processes in native habitats, with several definitions....
, feral honey bees have become a significant environmental problem in places where they are not native. Imported bees may compete with and displace native bees and birds, and may also promote the reproduction of invasive plants
Invasive species
"Invasive species", or invasive exotics, is a nomenclature term and categorization phrase used for flora and fauna, and for specific restoration-preservation processes in native habitats, with several definitions....
that native pollinators do not visit. Also, unlike native bees, they do not properly extract or transfer pollen from plants with poricidal anthers
Stamen
The stamen is the pollen producing reproductive organ of a flower...
(anthers that only release pollen through tiny apical pores), as this requires buzz pollination
Buzz pollination
Sonication or buzz pollination is a technique used by some bees to release pollen which is more or less firmly held by the anthers, which makes pollination more efficient. The anther of buzz-pollinated species of plants is typically tubular, with an opening at only one end, and the pollen is inside...
, a behavior which honey bees rarely exhibit. For example, honey bees reduce fruiting in Melastoma affine
Melastoma affine
Melastoma affine, also known by the common names Blue Tongue or Native Lassiandra ,Nekkarika in Kannada, is a shrub of the Melastomataceae family. Distributed in tropical and sub-tropical forests of India, South-east Asia and Australia, it is a plant of rainforest margins...
(a plant with poricidal anthers) by robbing stigmas of previously deposited pollen.
Insects
- Asian giant hornetAsian giant hornetThe Asian giant hornet , including the subspecies Japanese giant hornet , colloquially known as the yak-killer hornet, is the world's largest hornet, native to temperate and tropical Eastern Asia...
- Southeast Asia - Robber fliesAsilidaeInsects in the Diptera family Asilidae are commonly called robber flies. The family Asilidae contains about 7,100 described species worldwide....
- DragonflyDragonflyA dragonfly is a winged insect belonging to the order Odonata, the suborder Epiprocta or, in the strict sense, the infraorder Anisoptera . It is characterized by large multifaceted eyes, two pairs of strong transparent wings, and an elongated body...
- Green DarnerGreen DarnerThe Green Darner or Common Green Darner , after its resemblance to a darning-needle, is a species of dragonfly in the family Aeshnidae. One of the most common and abundant species throughout North America and its range south to Panama. It is well known for its great migration distance from the...
- Green Darner
- European beewolf - Europe and North Africa
- Chinese mantis
- Bald-faced hornetBald-faced hornetDolichovespula maculata is a North American insect commonly called the bald-faced hornet...
- Common water striderWater striderGerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
- YellowjacketYellowjacketYellowjacket is the common name in North America for predatory wasps of the genera Vespula and Dolichovespula. Members of these genera are known simply as "wasps" in other English-speaking countries...
Spiders
- Six-spotted fishing spider
- Green lynx spiderLynx spiderLynx Spiders are the members of the family Oxyopidae. They all are hunting spiders that spend their lives on plants, flowers and shrubs. At least one species has been identified as exhibiting social behaviour.-Description and habits:...
- Goldenrod spiderMisumenaFlower crab spiders are crab spiders of the genus Misumena with holarctic distribution.The goldenrod crab spider is the largest and most well-known of the North American flower spiders, commonly seen hunting in goldenrod sprays in autumn. It can change its color between white and yellow to match...
- Black argiope
Reptiles and amphibians
- American toadAmerican toadThe American Toad is a common species of toad found throughout the eastern United States and Canada. It is divided into three subspecies—the Eastern American Toad , the Dwarf American Toad , and the rare Hudson Bay Toad...
- AnolesAnolisAnolis is a genus of lizards belonging to the family Polychrotidae. With nearly 400 species, Anolis represents the world's most species rich amniote genus. Several species of Anolis are occasionally ascribed to the genus Norops, but the validity of the Norops genus is not widely accepted...
- BullfrogBullfrogThe American bullfrog , often simply known as the bullfrog in Canada and the United States, is an aquatic frog, a member of the family Ranidae, or “true frogs”, native to much of North America. This is a frog of larger, permanent water bodies, swamps, ponds, and lakes, where it is usually found...
- Wood frogWood FrogThe Wood Frog has a broad distribution over North America, extending from the southern Appalachians to the boreal forest with several notable disjunct populations including lowland eastern North Carolina...
Birds
- Bee-eaterBee-eaterThe bee-eaters are a group of near-passerine birds in the family Meropidae. Most species are found in Africa and Asia but others occur in southern Europe, Australia, and New Guinea. They are characterised by richly coloured plumage, slender bodies, and usually elongated central tail feathers...
- Common grackleCommon GrackleThe Common Grackle, Quiscalus quiscula, is a large icterid.-Description:The long adult has a long dark bill, pale yellowish eyes and a long tail; its feathers appear black with purple, green or blue iridescence on the head, and primarily bronze shine in the body plumage...
- Ruby-throated hummingbirdRuby-throated HummingbirdThe Ruby-throated Hummingbird , is a small hummingbird. It is the only species of hummingbird that regularly nests east of the Mississippi River in North America.- Description :...
- Tyrant flycatcherTyrant flycatcherThe tyrant flycatchers are a family of passerine birds which occur throughout North and South America. They are considered the largest family of birds on Earth, with more than 400 species. They are the most diverse avian family in every country in the Americas, except for the United States and...
- Great crested flycatcherGreat Crested FlycatcherThe Great Crested Flycatcher is a large insect-eating bird of the tyrant flycatcher family. It is the most widespread member of the genus, Myiarchus, in North America and is found over most of the eastern and mid-western portions of the continent...
Mammals
Contrary to popular perception, bears and honey badgers are brood predators; honey is only of secondary interest.- BearBearBears are mammals of the family Ursidae. Bears are classified as caniforms, or doglike carnivorans, with the pinnipeds being their closest living relatives. Although there are only eight living species of bear, they are widespread, appearing in a wide variety of habitats throughout the Northern...
- HumanHumanHumans are the only living species in the Homo genus...
- Least shrewLeast ShrewThe North American Least Shrew is one of the smallest mammals, growing to be only up to 3 inches long. The North American Least Shrew has a long pointed snout and a tail never more than twice the length of their hind foot. It has a dense fur coat that is either grayish-brown or reddish-brown...
- Opossum
- RaccoonRaccoonProcyon is a genus of nocturnal mammals, comprising three species commonly known as raccoons, in the family Procyonidae. The most familiar species, the common raccoon , is often known simply as "the" raccoon, as the two other raccoon species in the genus are native only to the tropics and are...
- Honey badgerRatelThe honey badger , also known as the ratel, is a species of mustelid native to Africa, the Middle East and the Indian Subcontinent. Despite its name, the honey badger does not closely resemble other badger species, instead bearing more anatomical similarities to weasels...
- SkunkSkunkSkunks are mammals best known for their ability to secrete a liquid with a strong, foul odor. General appearance varies from species to species, from black-and-white to brown or cream colored. Skunks belong to the family Mephitidae and to the order Carnivora...
Designated state insect
- ArkansasArkansasArkansas is a state located in the southern region of the United States. Its name is an Algonquian name of the Quapaw Indians. Arkansas shares borders with six states , and its eastern border is largely defined by the Mississippi River...
(1973) - North CarolinaNorth CarolinaNorth Carolina is a state located in the southeastern United States. The state borders South Carolina and Georgia to the south, Tennessee to the west and Virginia to the north. North Carolina contains 100 counties. Its capital is Raleigh, and its largest city is Charlotte...
(1973) - New JerseyNew JerseyNew Jersey is a state in the Northeastern and Middle Atlantic regions of the United States. , its population was 8,791,894. It is bordered on the north and east by the state of New York, on the southeast and south by the Atlantic Ocean, on the west by Pennsylvania and on the southwest by Delaware...
(1974) – state bug - GeorgiaGeorgia (U.S. state)Georgia is a state located in the southeastern United States. It was established in 1732, the last of the original Thirteen Colonies. The state is named after King George II of Great Britain. Georgia was the fourth state to ratify the United States Constitution, on January 2, 1788...
(1975) - MaineMaineMaine is a state in the New England region of the northeastern United States, bordered by the Atlantic Ocean to the east and south, New Hampshire to the west, and the Canadian provinces of Quebec to the northwest and New Brunswick to the northeast. Maine is both the northernmost and easternmost...
(1975) - NebraskaNebraskaNebraska is a state on the Great Plains of the Midwestern United States. The state's capital is Lincoln and its largest city is Omaha, on the Missouri River....
(1975) - KansasKansasKansas is a US state located in the Midwestern United States. It is named after the Kansas River which flows through it, which in turn was named after the Kansa Native American tribe, which inhabited the area. The tribe's name is often said to mean "people of the wind" or "people of the south...
(1976) - LouisianaLouisianaLouisiana is a state located in the southern region of the United States of America. Its capital is Baton Rouge and largest city is New Orleans. Louisiana is the only state in the U.S. with political subdivisions termed parishes, which are local governments equivalent to counties...
(1977) - VermontVermontVermont is a state in the New England region of the northeastern United States of America. The state ranks 43rd in land area, , and 45th in total area. Its population according to the 2010 census, 630,337, is the second smallest in the country, larger only than Wyoming. It is the only New England...
(1977) - WisconsinWisconsinWisconsin is a U.S. state located in the north-central United States and is part of the Midwest. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake Michigan to the east, Michigan to the northeast, and Lake Superior to the north. Wisconsin's capital is...
(1977) - South DakotaSouth DakotaSouth Dakota is a state located in the Midwestern region of the United States. It is named after the Lakota and Dakota Sioux American Indian tribes. Once a part of Dakota Territory, South Dakota became a state on November 2, 1889. The state has an area of and an estimated population of just over...
(1978) - MississippiMississippiMississippi is a U.S. state located in the Southern United States. Jackson is the state capital and largest city. The name of the state derives from the Mississippi River, which flows along its western boundary, whose name comes from the Ojibwe word misi-ziibi...
(1980) - UtahUtahUtah is a state in the Western United States. It was the 45th state to join the Union, on January 4, 1896. Approximately 80% of Utah's 2,763,885 people live along the Wasatch Front, centering on Salt Lake City. This leaves vast expanses of the state nearly uninhabited, making the population the...
(1983) - MissouriMissouriMissouri is a US state located in the Midwestern United States, bordered by Iowa, Illinois, Kentucky, Tennessee, Arkansas, Oklahoma, Kansas and Nebraska. With a 2010 population of 5,988,927, Missouri is the 18th most populous state in the nation and the fifth most populous in the Midwest. It...
(1985) - TennesseeTennesseeTennessee is a U.S. state located in the Southeastern United States. It has a population of 6,346,105, making it the nation's 17th-largest state by population, and covers , making it the 36th-largest by total land area...
(1990) – official agricultural insect - OklahomaOklahomaOklahoma is a state located in the South Central region of the United States of America. With an estimated 3,751,351 residents as of the 2010 census and a land area of 68,667 square miles , Oklahoma is the 28th most populous and 20th-largest state...
(1992) - West VirginiaWest VirginiaWest Virginia is a state in the Appalachian and Southeastern regions of the United States, bordered by Virginia to the southeast, Kentucky to the southwest, Ohio to the northwest, Pennsylvania to the northeast and Maryland to the east...
(2002)
See also
- Bee sting therapy
- BeelineBeelineBeelining is an ancient art used to locate feral bee colonies by capturing and marking foraging worker bees, then releasing them from various points to establish the direction and distance of the colony's home...
- Bee beardingBee beardingBee bearding is the practice of wearing several hundred thousand honey bees on the face, usually as a sideshow-type demonstration at agricultural shows...
- Ecological importance of bees
External links
- Flight of Honey Bee finally understood
- Honey Bee Pheromones
- Cornwall Honey Beepedia
- BeeSource
- Honey Bee Apis mellifera diagnostic pictures, descriptions, natural history
- Brother Adams On-line book
- Beekeeping explained by the Food and Agriculture OrganizationFood and Agriculture OrganizationThe Food and Agriculture Organization of the United Nations is a specialised agency of the United Nations that leads international efforts to defeat hunger. Serving both developed and developing countries, FAO acts as a neutral forum where all nations meet as equals to negotiate agreements and...
- Types of hives by FAO
- Interactive 3D flash presentation of Honey beehive from PBS
- Brainy Bees Think Abstractly
- Digital library of Beekeeping
- Anatomy of the honeybee by the FAO
- Apis mellifera pomonella, a new honey bee subspecies from Central Asia; Apidologie 34 (2003) 367-375; Sheppard and Meixner
- Origin of honeybees from the Yucatan peninsula inferred from mitochondrial DNA analysis; Molecular Ecology (2001)10, 1347-1355
- Cyclopedia of American Agriculture ed. by L. H. Bailey (1911), Vol. III—Animals, "Bees," including a history of the literature related to European honey bees in the United States.