Western honey bee (Apis mellifera)
Honey bee foraging on African Oil Palm flowers
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Apidae
Genus:	Apis
Species:	A. mellifera
Binomial name
Apis mellifera Linnaeus, 1758
Subspecies
North-west of Europe A. m. iberica A. m. intermissa A. m. lihzeni A. m. mellifera A. m. sahariensis South-west of Europe A. m. carnica A. m. cecropia A. m. ligustica A. m. macedonica A.m. ruttneri A. m. sicula Middle East A. m. adamii A. m. anatoliaca A. m. armeniaca A. m. caucasica A. m. cypria A. m. meda Africa A. m. adansonii A. m. capensis A. m. intermissa A. m. lamarckii A. m. litorea A. m. major A. m. monticola A. m. sahariensis A. m. scutellata A. m. unicolor A. m. jemenitica
Synonyms
Apis mellifica Linnaeus, 1761

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, upon realizing 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.

Geographic distribution[link]

The western 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.^[1] Since then, they have spread throughout the Americas.

Western honey bees differentiated into geographic races as they spread from Africa into Eurasia.^[2] Currently, 28 subspecies based on these geographic variations are recognized.^{[citation needed]} All races 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. Regions with localized variations may become identified subspecies in the near future, such as A. m. pomonella from the Tian Shan Mountains, which would be included in the Mideast subspecies branch.

Geographic isolation led to numerous adaptations as honey bees 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[link]

In the temperate zone, honey bees 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 larvae 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 which 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.

Larvae to the left and eggs to the right

For the first 10 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 pupae 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, with good genetics and sufficient number of matings also being factors. During the larval and pupal stages, various parasites can attack the pupa/larva and destroy or damage it.

Pupae of drones

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 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 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.

Stages of development of the drone pupae

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 misconception is 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. This complex apparatus, including the barbs on the sting, is presumed to have 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 [link]

Drones are the male 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, they 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 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, produce honey, or take care of themselves.

Life expectancy [link]

The average lifespan of the queen in most subspecies is three to four years. However, reports from the German/European black bee subspecies previously used for beekeeping, indicate the queen could live up to eight years.^[3] Because queens deplete their store of sperm, towards the end of their lives, 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.^[4] Except for the queen, all of the colony's workers are therefore exchanged about every four months.

Honey production [link]

Honey bee with tongue partially extended

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 enzymes 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 nonaggressive; this is speculated to be a defense mechanism. Wild colonies generally live in hollow trees, and when bees detect smoke, they are assumed to prepare to evacuate from a forest fire, carrying as much food reserve as they can. 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 [link]

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 cluster, the internal temperature may be as low as 20–22 °C (68–72 °F).

Honey bees are able to forage over a 30 °C (54.0 °F) 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.^[5]

The optimal air temperature for foraging is 22–25 °C (72–77 °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 (18.0 °F).^[6]

Below 7–10 °C (45–50 °F), bees become immobile, and above 38 °C (100 °F), their activity slows. Honey bees can tolerate temperatures up to 50 °C (122 °F) for short periods.

Queens [link]

Periodically, the colony determines a new queen is needed. There are three general triggers:

1. 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.
2. 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.
3. 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. Should two queens emerge simultaneously, they are thought to fight to the death. Recent studies, however, have indicated 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.^{[citation needed]} Regardless, the queen asserts her control over the worker bees through the release of a complex suite of pheromones called queen scent.

Peanut-like queen brood cells are extended outward from the brood comb.

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 midair (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. 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 a worker-sized cell using stored sperm from the spermatheca. Eggs, laid into drone-sized (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[link]

The European honey bee is the third insect, after the fruit fly 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.^[7] They have also discovered the number of genes related to smell outnumber those for taste, and they have fewer genes for immunity than the fruit fly and the mosquito.^[8] 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.^[9]

The genome is unusual in having very few transposons, while they have been present in the evolutionary past (inactive remains were found) and in general evolved slower than in Diptera species.^[8]

Pheromones [link]

Honey bees use special pheromones, or chemical communication, for almost all behaviors of life. Such uses include (but are not limited to): mating, alarm, defense, orientation, kin and colony recognition, food production, and integration of colony activities. Pheromones are thus essential to honey bees for their survival.

Communication [link]

A large honey bee swarm on a fallen tree trunk

Honey bees are an excellent animal species 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 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. Bees also are thought to 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.

"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 Frisch 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.

Beekeeping[link]

A queen bee with workers

The honey bee is a colonial 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 superorganisms.

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 enzymes, 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 hives, top-bar hives, 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.

Breeding [link]

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.

Products [link]

Honey bees [link]

One of the primary products is more honeybees. Honey bees are bought and sold as mated queens, in spring packages of a queen with 2 to 5 pounds of bees, as nucleus colonies which include frames of brood, and as full colonies of bees. Commercial sale of bees dates to prehistory. Modern methods of producing queens and dividing colonies for increase date to the late 1800's. Bees are typically produced in temperate to tropical regions and sold to colder areas, as for example packages of bees produced in Florida and sold to beekeepers in Michigan.

Pollination[link]

Beehives set up for pollination

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 almonds occurs in February, early in the growing season, before local hives have built up their populations. Almond orchards 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,^[10] 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.

California High Desert Honey Bees Immersed in Yellow Beavertail Cactus Flower Pollen

^[12]

Honey[link]

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.

The bee carries pollen in a pollen basket back to the hive

Beeswax[link]

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 candles and seals.

See movie of bee at work collecting pollen. Requires an Ogg Theora enabled browser or player, such as Firefox or VideoLAN.

Pollen[link]

Bees collect pollen in the pollen basket 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.

Propolis[link]

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[link]

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[link]

European honey bee populations have recently faced threats to their survival. North American 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.^[13] 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^{[citation needed]}, while various bee pests and diseases are becoming resistant to medications (e.g. American foulbrood, tracheal mites and Varroa mites).

Environmental hazards[link]

A bee swarm. Bees are remarkably non-aggressive in this state as they have no hive to protect, and can be captured with ease.

In North America, Africanized bees 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.^[14]

Predators [link]

Insects[link]

• Asian giant hornet - Southeast Asia
• Robber flies
• Dragonfly
  • Green Darner
• European beewolf - Europe and North Africa
• Chinese mantis
• Bald-faced hornet
• Common water strider
• Yellowjacket

Spiders[link]

• Six-spotted fishing spider
• Green lynx spider
• Goldenrod spider ^[15]
• Black argiope

Reptiles and amphibians[link]

• American toad
• Anoles
• Bullfrog
• Wood frog

Birds[link]

• Bee-eater
• Common grackle
• Ruby-throated hummingbird
• Tyrant flycatcher
• Great Crested Flycatcher

Mammals[link]

Contrary to popular perception, bears and honey badgers are brood predators; honey is only of secondary interest.

• Bear
• Human
• Least shrew
• Opossum
• Raccoon
• Honey badger
• Skunk

Designated state insect[link]

See also [link]

• Bee sting therapy
• Beeline
• Bee bearding
• Ecological importance of bees

References [link]

Bibliography[link]

This article includes a list of references, but its sources remain unclear because it has insufficient inline citations. Please help to improve this article by introducing more precise citations. (February 2010)

External links [link]

Wikimedia Commons has media related to: Apis mellifera

• 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 Organization
• 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.

This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see European honey bee.

Honeybees Temporal range: Oligocene–Recent PreЄ Є O S D C P T J K Pg N
European honey bee carrying pollen back to the hive
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Apidae
Subfamily:	Apinae
Tribe:	Apini Latreille, 1802
Genus:	Apis Linnaeus, 1758
Species
Subgenus Micrapis: Apis andreniformis Apis florea Subgenus Megapis: Apis dorsata Subgenus Apis: Apis cerana Apis koschevnikovi Apis mellifera Apis nigrocincta

Honey bees (or honeybees) 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. Currently, there are only seven recognised species of honey bee with a total of 44 subspecies,^[1] though historically, anywhere from six to eleven species have been recognised. Honey bees represent only a small fraction of the approximately 20,000 known species of bees. Some other types of related bees produce and store honey, but only members of the genus Apis are true honey bees.

Origin, systematics and distribution[link]

Morphology of a female honey bee

Honey bees as a group appear to have their centre of origin in South and South East Asia (including the Philippines), as all but one (i.e. Apis mellifera), of the extant species are native to that region. Notably the most plesiomorphic living species (Apis florea and Apis andreniformis) has the center of origin there.^[2]

The first Apis bees appear in the fossil record at the Eocene–Oligocene (23-56 Mya) boundary, in European deposits. The origin of these prehistoric honey bees does not necessarily indicate that Europe is where the genus originated, only that it occurred there at that time. There are few known fossil deposits in South Asia, the suspected region of honey bee origin, and fewer still have been thoroughly studied.

No Apis species existed in New World in human time before introduction of Apis melifera by Europeans. There is only one fossil species documented from the New World, Apis nearctica, known from a single 14-million-year old specimen from Nevada.^[3]

The close relatives of modern honey bees –- e.g. bumblebees and stingless bees –- are also social to some degree, and social behavior seems a plesiomorphic trait that predates the origin of the genus. Among the extant members of Apis, the more basal species make single, exposed combs, while the more recently evolved species nest in cavities and have multiple combs, which has greatly facilitated their domestication.

Most species have historically been cultured or at least exploited for honey and beeswax by humans indigenous to their native ranges. Only two of these species have been truly domesticated, one (Apis mellifera) at least since the time of the building of the Egyptian pyramids, and only that species has been moved extensively beyond its native range.

Today's honey bees constitute three clades.^[1][4]

Apis dorsata on Tribulus terrestris in Hyderabad, India

[edit] Genetics

The chromosomes counts of female bees for the three clades are: Micrapis 2N=16, Megapis 2N=16, Apis 2N=32. Drones of all species have 1N chromosome counts. The genome of Apis has been mapped.

Drones are produced from unfertilized eggs and therefore represent only the DNA of the queen that laid the eggs, i.e. have only a mother. Workers and queens result from fertilized eggs and therefore have both a mother and a father. A modified form of parthenogenesis controls sex differentiation. The sex allele is polymorphic and so long as two different variants are present, a female bee results. If both sex alleles are identical, diploid drones are produced. Honeybees detect and destroy diploid drones after the eggs hatch.

Queens typically mate with multiple drones on more than one mating flight. Once mated, they lay eggs and fertilize them as needed from sperm stored in the spermatheca. Since the number of sex alleles is limited - about 18 are known in Apis - there is a high probability that a queen will mate with one or more drones having sex alleles identical with one of the sex alleles in the queen. It is therefore typical for a queen to produce a percentage of diploid drone eggs.

[edit] Micrapis

Apis florea and Apis andreniformis are small honey bees of southern and southeastern Asia. They make very small, exposed nests in trees and shrubs. Their stings are often incapable of penetrating human skin, so the hive and swarms can be handled with minimal protection. They occur largely sympatrically though they are very distinct evolutionarily and are probably the result of allopatric speciation, their distribution later converging. Given that A. florea is more widely distributed and A. andreniformis is considerably more aggressive, honey is – if at all – usually harvested from the former only. They are the most ancient extant lineage of honey bees, maybe diverging in the Bartonian (some 40 million years ago or slightly later) from the other lineages, but among themselves do not seem to have diverged a long time before the Neogene.^[4]

[edit] Megapis

There is one recognised species in subgenus Megapis. It usually builds single or a few exposed combs on high tree limbs, on cliffs, and sometimes on buildings. They can be very fierce. Periodically robbed of their honey by human "honey hunters", colonies are easily capable of stinging a human being to death when provoked.

Apis dorsata on comb

• Apis dorsata, the giant honey bee, is native and widespread across most of South and Southeast Asia.
• Apis dorsata binghami, the Indonesian honey bee, is classified as the Indonesian subspecies of the giant honey bee or a distinct species; in the latter case, A. d. breviligula and/or other lineages would probably also have to be considered species.^[5]
• Apis dorsata laboriosa, the Himalayan honey bee, was initially described as a distinct species. Later, it was included in A. dorsata as a subspecies^[1] based on the biological species concept, though authors applying a genetic species concept have suggested it should be considered a species.^[4] Essentially restricted to the Himalayas, it differs little from the giant honey bee in appearance, but has extensive behavioral adaptations which enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.

[edit] Apis

Eastern species

Eastern honey bee (Apis cerana) from Hong Kong

These are three or four species. The reddish Koschevnikov's bee (Apis koschevnikovi) from Borneo is well distinct; it probably derives from the first colonization of the island by cave-nesting honey bees. Apis cerana, the Eastern honey bee proper, is the traditional honey bee of southern and eastern Asia, kept in hives in a similar fashion to Apis mellifera, though on a much smaller and regionalised scale. It has not been possible yet to resolve its relationship to the Bornean Apis cerana nuluensis and Apis nigrocincta from the Philippines to satisfaction; the most recent hypothesis is that these are indeed distinct species but that A. cerana is still paraphyletic, consisting of several good species.^[4]

European/Western/Common honey bee

Apis mellifera, the most commonly domesticated species, was the third insect to have its genome mapped. It seems to have originated in eastern tropical Africa and spread from there to Northern Europe and eastwards into Asia to the Tien Shan range. It is variously called the European, Western or Common honey bee in different parts of the world. There are many subspecies that have adapted to the local geographic and climatic environment, and in addition, hybrid strains such as the Buckfast bee have been bred. Behavior, color and anatomy can be quite different from one subspecies or even strain to another.

The European honey bee originated from eastern Africa. This bee is pictured in Tanzania.

Regarding phylogeny, this is the most enigmatic honey bee species. It seems to have diverged from its Eastern relatives only during the Late Miocene. This would fit the hypothesis that the ancestral stock of cave-nesting honey bees was separated into the Western group of E Africa and the Eastern group of tropical Asia by desertification in the Middle East and adjacent regions, which caused declines of foodplants and trees which provided nest sites, eventually causing gene flow to cease. The diversity of subspecies is probably the product of a largely Early Pleistocene radiation aided by climate and habitat changes during the last ice age. That the Western honey bee has been intensively managed by humans since many millennia – including hybridization and introductions – has apparently increased the speed of its evolution and confounded the DNA sequence data to a point where little of substance can be said about the exact relationships of many A. mellifera subspecies.^[4]

Apis mellifera is not native to the Americas and therefore were not present upon the arrival of the European explorers and colonists. There were, however, other native honey bee species kept and traded by indigenous peoples.^[6] In 1622, European colonists brought the dark bee (A. m. mellifera) to the Americas, followed later by Italian bees (A. m. ligustica) and others. Many of the crops that depend on honey bees for pollination have also been imported since colonial times. Escaped swarms (known as "wild" bees, but actually feral) spread rapidly as far as the Great Plains, usually preceding the colonists. Honey bees did not naturally cross the Rocky Mountains; they were transported by the Mormon pioneers^[7] to Utah in the late 1840s, and by ship to California in the early 1850s.

Africanized bee[link]

Africanized bees (known colloquially as "killer bees") are hybrids between European stock and one of the African subspecies A. m. scutellata; they are often more aggressive than and do not create as much of a surplus as European bees, but are more resistant to disease and are better foragers^{[citation needed]}. Originating by accident in Brazil, they have spread to North America and constitute a pest in some regions. However, these strains do not overwinter well, and so are not often found in the colder, more northern parts of North America. On the other hand, the original breeding experiment for which the African bees were brought to Brazil in the first place has continued (though not as intended). Novel hybrid strains of domestic and re-domesticated Africanized bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.

Beekeeping[link]

Two species of honey bee, A. mellifera and A. cerana indica, are often maintained, fed, and transported by beekeepers. 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, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations.

Colony Collapse Disorder (CCD)[link]

Frame removed from Langstroth hive

Beekeepers in Western countries have been reporting slow declines of stocks for many years, apparently due to impaired protein production, changes in agricultural practice, or unpredictable weather. In early 2007, abnormally high die-offs (30-70% of hives) of European honey bee colonies occurred in North America; such a decline seems unprecedented in recent history. This has been dubbed "Colony collapse disorder" (CCD); it is unclear whether this is simply an accelerated phase of the general decline due to stochastically more adverse conditions in 2006, or a novel phenomenon. Research has so far failed to determine what causes it, but the weight of evidence is tentatively leaning towards CCD being a syndrome rather than a disease as it seems to be caused by a combination of various contributing factors rather than a single pathogen or poison.

Life cycle[link]

A queen bee: a coloured dot, in this case yellow, is added to assist the beekeeper in identifying the queen.

Honey bee eggs shown in cut open wax cells

Pupae drones

Emergence of a black bee (Apis mellifera mellifera)

Eggs and larvae

As in a few other types of eusocial bees, a colony generally contains one queen bee, a fertile female; seasonally up to a few thousand drone bees or fertile males;^[8] and a large seasonally variable population of sterile female worker bees. Details vary among the different species of honey bees, but common features include:

1. Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the worker bees. Using her spermatheca, the queen actually can choose to fertilize the egg she is laying, usually depending on what cell she is laying in. Drones develop from unfertilised eggs and are haploid, while females (queens and worker bees) develop from fertilised eggs and are diploid. Larvae are initially fed with royal jelly produced by worker bees, later switching to honey and pollen. The exception is a larva fed solely on royal jelly, which will develop into a queen bee. The larva undergoes several moltings before spinning a cocoon within the cell, and pupating.

2. Young worker bees clean the hive and feed the larvae. When their royal jelly producing glands begin to atrophy, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers, and guarding the hive. Later still, a worker takes her first orientation flights and finally leaves the hive and typically spends the remainder of her life as a forager.

3. Worker bees cooperate to find food and use a pattern of "dancing" (known as the bee dance or waggle dance) to communicate information regarding resources with each other; this dance varies from species to species, but all living species of Apis exhibit some form of the behavior. If the resources are very close to the hive, they may also exhibit a less specific dance commonly known as the "Round Dance".

4. Honey bees also perform tremble dances which recruit receiver bees to collect nectar from returning foragers.

5. Virgin queens go on mating flights away from their home colony, and mate with multiple drones before returning. The drones die in the act of mating.

6. Colonies are established not by solitary queens, as in most bees, but by groups known as "swarms", which consist of a mated queen and a large contingent of worker bees. This group moves en masse to a nest site that has been scouted by worker bees beforehand. Once they arrive, they immediately construct a new wax comb and begin to raise new worker brood. This type of nest founding is not seen in any other living bee genus, though there are several groups of Vespid wasps which also found new nests via swarming (sometimes including multiple queens). Also, stingless bees will start new nests with large numbers of worker bees, but the nest is constructed before a queen is escorted to the site, and this worker force is not a true "swarm".

Winter survival[link]

In cold climates honey bees stop flying when the temperature drops below about 10 °C (50 °F) and crowd into the central area of the hive to form a "winter cluster". The worker bees huddle around the queen bee at the center of the cluster, shivering in order to keep the center between 27 °C (81 °F) at the start of winter (during the broodless period) and 34 °C (93 °F) once the queen resumes laying. The worker bees rotate through the cluster from the outside to the inside so that no bee gets too cold. The outside edges of the cluster stay at about 8–9 °C (46–48 °F). The colder the weather is outside, the more compact the cluster becomes. During winter, they consume their stored honey to produce body heat. The amount of honey consumed during the winter is a function of winter length and severity but ranges in temperate climates from 30 to 100 lbs.^[9]

Pollination[link]

Species of Apis are generalist floral visitors, and will pollinate a large variety of plants, but by no means all plants. Of all the honey bee species, only Apis mellifera has been used extensively for commercial pollination of crops and other plants. The value of these pollination services is commonly measured in the billions of dollars.

Pollination by a honey bee

Honey[link]

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 as a food source for the colony. All living species of Apis have had their honey gathered by indigenous peoples for consumption, though for commercial purposes only Apis mellifera and Apis cerana have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various stingless bees.

Honeycombs

Beeswax[link]

Worker bees of a certain age will secrete beeswax from a series of glands on their abdomens. They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered for various purposes.

Bee (apis)

A forager collecting pollen

Pollen[link]

Bees collect pollen in the pollen basket and carry it back to the hive. In the hive, pollen is used as a protein source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of A. mellifera and A. cerana. It is often eaten as a health supplement.

Propolis[link]

Propolis or bee glue is created from resins, balsams and tree saps. Those species of honey bees which nest in tree cavities use propolis to seal cracks in the hive. Dwarf honey bees use propolis to defend against ants by coating the branch from which their nest is suspended to create a sticky moat. Propolis is consumed by humans as a health supplement in various ways and also used in some cosmetics.

Defense[link]

Apis cerana japonica forming a ball around two hornets. The body heat trapped by the ball will overheat and kill the hornets.

All honey bees live in colonies where the worker bees will sting intruders as a form of defense, and alarmed bees will release a pheromone that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species (and virtually all other Hymenoptera) by the possession of small barbs on the sting, but these barbs are found only in the worker bees. The sting and associated venom sac are also 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. The worker bee dies after the stinger is torn from its body. As with other forms of life, warnings are given before an attack is launched. In the case of some honey bee species in the wild, this takes the form of a 'Mexican wave' which spreads as a ripple across a layer of bees densely packed on the surface of a comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings.^[10]

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 usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the flexible exoskeletal joints in appendages of other insects (and is used in fights between queens), in the case of Apis cerana defense against other insects such as predatory wasps is usually performed by surrounding the intruder with a mass of defending worker bees, who vibrate their muscles so vigorously that it raises the temperature of the intruder to a lethal level.^[11] It was previously thought that the heat alone was responsible for killing intruding wasps, but recent experiments have demonstrated that it is the increased temperature in combination with increased carbon dioxide levels within the ball that produces the lethal effect.^[12][13] This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as balling the queen, named for the ball of bees formed.

Communication[link]

Honey bees are known to communicate through many different chemicals and odours, as is common in insects, but also using specific behaviours that convey information about the quality and type of resources in the environment, and where these resources are located. The details of the signalling being used vary from species to species; for example, the two smallest species, Apis andreniformis and Apis florea, dance on the upper surface of the comb, which is horizontal (not vertical, as in other species), and worker bees orient the dance in the actual compass direction of the resource to which they are recruiting.

Symbolism[link]

Both the Atharva Veda^[14] and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at Delphi was the "Delphic Bee".

A community of honey bees has often been employed throughout history by political theorists as a model of human society:

"This image occurs in Aristotle and Plato; in Virgil^[15] and Seneca; in Erasmus and Shakespeare; in Marx and Tolstoy."^[16]

Honey bees, signifying immortality and resurrection, were royal emblems of the Merovingians, revived by Napoleon.^[17] The bee is the heraldic emblem too of the Barberini.

See also[link]

Wikimedia Commons has media related to: Apis (insect)

Look up honey bee in Wiktionary, the free dictionary.

• Apiology
• Bees and toxic chemicals
• Honey bee life cycle
• Bee sting

References[link]

Further reading[link]

• Engel, Michael S. & Grimaldi, David (2005): Evolution of the Insects. Cambridge University Press.
• Kak, Subhash C. (1991): The Honey Bee Dance Language Controversy. The Mankind Quarterly Summer 1991: 357-365. HTML fulltext
• Lindauer, Martin (1971): Communication among social bees. Harvard University Press.

Blake Shelton
Shelton in April 2010
Background information
Birth name	Blake Tollison Shelton
Born	(1976-06-18) June 18, 1976 (age 36)[1] Ada, Oklahoma, United States
Genres	Country
Occupations	Singer
Instruments	Vocals, guitar
Years active	2001–present
Labels	Giant, Warner Bros. Nashville, Reprise Nashville
Associated acts	Bobby Braddock, Miranda Lambert, Trace Adkins, Adam Levine, Christina Aguilera, Cee Lo Green
Website	blakeshelton.com

Blake Tollison Shelton (born June 18, 1976) is an American country music artist. In 2001, he made his debut with the single "Austin". Released as the lead-off single from his self-titled debut album, "Austin" went on to spend five weeks at Number One on the Billboard Hot Country Songs charts. This song was the first single from his gold-certified debut album, which also produced two more Top 20 hits. Although the album was released on Giant Records Nashville, Shelton was transferred to Warner Bros. Records Nashville after Giant closed in late 2001.

His second and third albums, 2003's The Dreamer (his first for Warner Bros. proper) and 2004's Blake Shelton's Barn & Grill, were each certified gold as well. Shelton's fourth album, Pure BS, was issued in 2007, and re-issued in 2008 with a cover of Michael Bublé's pop hit "Home" as one of the bonus tracks. This cover was also that album's third single. A fifth album, Startin' Fires, was released in November 2008. It was followed by the extended plays Hillbilly Bone and All About Tonight in 2010, and the album Red River Blue in 2011. He is currently a vocal coach on the NBC reality talent show The Voice with Adam Levine, Christina Aguilera, and Cee Lo Green.

Overall, Shelton has charted 17 singles on the country charts, including eleven Number One hits: "Austin" (2001), "The Baby" (2003), "Some Beach" (2004–2005), "Home" (2008), "She Wouldn't Be Gone" (2009), "Hillbilly Bone" (2010), a duet with Trace Adkins, "All About Tonight" (2010), "Who Are You When I'm Not Looking" (2011)", "Honey Bee" (2011), "God Gave Me You" (2011), and "Drink on It" (2011-2012). Additionally, three more of his singles have reached Top Ten: a cover version of Conway Twitty's "Goodbye Time", "Nobody but Me", and "I'll Just Hold On".

He also covered "Footloose" for the movie remake in 2011.

Early life[link]

Shelton was born in Ada, Oklahoma, to Dick, a used car dealer, and Dorothy, a beauty salon owner. He began singing at an early age and by age 12, he learned to play guitar. By age sixteen, Shelton had received a Denbo Diamond Award in his home state.^[1] On November 13, 1990, his older brother Richie, who was 24 at the time, died in a car accident. He has an older sister, Endy. At age 17, he moved to Nashville, Tennessee, in pursuit of a singing career.

Musical career[link]

Shelton performing at the Crawford City Fair in August 2005

[edit] 1998–2002: Blake Shelton

After several years in Nashville, he signed to Giant Records in 1998. In 2001, he was slated to release a song entitled "I Wanna Talk About Me" as a single. Staff at the label considered the song unsuitable for a lead-off single, however, and the song was eventually recorded by Toby Keith, whose version was a Number One hit.^[2]

Instead, Giant released "Austin" as Shelton's debut single. Shortly after its release, however, Giant Records was dissolved, and Shelton was transferred to Warner Bros. Records, the parent company. "Austin" went on to spend five weeks at Number One on the Billboard Hot Country Singles & Tracks (now Hot Country Songs) charts. His self-titled debut album also produced the No.18 "All Over Me" (which Shelton co-wrote with fellow country artist Earl Thomas Conley) and the No.14 "Ol' Red", which was a cover version of a George Jones song. Although Shelton's rendition of "Ol' Red" was not a major radio hit, he considers it his signature song, and it has become popular in concert.^[3] Blake Shelton is certified gold by the RIAA.^[1]

[edit] 2002–2003: The Dreamer

Shelton's second album, The Dreamer, was first released on February 4, 2003 on Warner Bros. Records. Its lead-off single, "The Baby", reached Number One on the country charts, holding that position for three weeks. Although the second and third singles ("Heavy Liftin'" and "Playboys of the Southwestern World", respectively) only reached No.32 and #24, The Dreamer earned gold certification as well. He, along with Andy Griggs and Montgomery Gentry, sang guest vocals on Tracy Byrd's mid-2003 single "The Truth About Men".

[edit] 2004–2006: Blake Shelton's Barn & Grill

Blake Shelton's Barn & Grill was the title of Shelton's third studio album, released in 2004.^[1] Its lead-off single, the Harley Allen co-write "When Somebody Knows You That Well", peaked at No.37 on the country charts, while the follow-up "Some Beach" became his third No.1 hit, holding that position for four weeks. It was followed by a cover of Conway Twitty's 1988 single "Goodbye Time".^[4] Both this cover and its followup, "Nobody but Me", reached Top Ten for Shelton as well. As with his first two albums, Blake Shelton's Barn & Grill was certified gold. Accompanying the album's release was a video collection entitled Blake Shelton's Barn & Grill: A Video Collection.^[1]

On December 18, 2005, several of Shelton's songs, including "Nobody but Me", appeared on the TV movie The Christmas Blessing, starring Neil Patrick Harris, Rebecca Gayheart, Angus T. Jones, and Rob Lowe. Shelton had a small role at the end of the movie, playing himself at a benefit concert, singing "Nobody but Me".

[edit] 2006–2008: Pure BS

Shelton issued his fourth studio album, Pure BS, in early 2007.^[1] Unlike with his first three albums, which were produced entirely by Bobby Braddock, Shelton worked with Braddock, Brent Rowan, and Paul Worley as producers for this album. Its first two singles—"Don't Make Me" and "The More I Drink"—were both Top 20 hits on the country charts, respectively reaching No.12 and #19. Also in late 2007, Shelton made appearances on television shows: first as a judge on the talent competition Nashville Star, and later on Clash of the Choirs.

Pure BS was re-released in 2008 with three bonus tracks, including a cover of Michael Bublé's hit single "Home". This cover, released in early 2008 as the album's third single, became his fourth No.1 hit in July.^[5]

[edit] 2008–2009: Startin' Fires

"Home" was followed in August 2008 by the single "She Wouldn't Be Gone", his 14th chart entry and his fifth No. 1 hit. It was the first time in his career that he had two consecutive Number Ones. "She Wouldn't Be Gone" is the lead-off to Shelton's fifth studio album, Startin' Fires, which has also produced the single "I'll Just Hold On." This album also includes "Bare Skin Rug," a duet with Lambert.^[6] It was produced by Scott Hendricks, except for one track which was produced by Brent Rowan, and another ("I Don't Care", which carried over from Pure BS) was produced by Braddock.^[7]

[edit] 2009–2011: Hillbilly Bone and All About Tonight

After "I'll Just Hold On" fell off the chart in October 2009, Blake Shelton released a duet with Trace Adkins titled "Hillbilly Bone." It was the lead-off single on Shelton's EP, Hillbilly Bone, released on March 2, 2010 through Reprise Records Nashville. After "Hillbilly Bone" reached No.1 in February, came the August release of "All About Tonight", the lead-off single to his EP of the same name. In September, he released the second single from All About Tonight, "Who Are You When I'm Not Looking", his 18th single release. He released his first greatest hits album, Loaded: The Best of Blake Shelton in November 2010.

Shelton was invited to join the Grand Ole Opry during the September 28, 2010 "Country Comes Home" concert celebrating reopening of the Grand Ole Opry House after the Cumberland River flooded the Opry House in May 2010. The formal invitation was extended on his Twitter account and was announced by Opry star Trace Adkins.^[8] He was formally inducted by Adkins at the Saturday, October 23, 2010 Opry performance.

Shelton recorded the title track for the "Footloose" remake film released on October 14th, 2011.

Blake Shelton appeared on the 44th Annual Country Music Awards on November 10, 2010 where he performed "All About Tonight" and won Male Vocalist of the Year.^[9]

Shelton will serve as a judge/coach to singers on the 2011 NBC reality television series The Voice.^[10]

[edit] 2011–present: Red River Blue

Blake Shelton released the album Red River Blue on July 12, 2011, led by the single, "Honey Bee". The song received 138,000 downloads in its first week and was certified gold in its seventh week, setting a new record for the fastest gold certification by a male country singer.^[11] On June 13, 2011, in its tenth chart week, "Honey Bee" went to #1 on the Billboard Hot Country Songs chart, becoming his ninth #1 and his fastest-climbing. The album was expected to debut at #1 on the Billboard 200 with around 110,000 copies sold.^[12] "God Gave Me You", a cover of a Dave Barnes song, was the album's second single; it also reached #1. "Drink On It", the fifth song on the album, hit #1 in April 2012, giving him his 11th #1 song. On April 30, 2012, Blake performed "Over" on the Semi-finals of the second season of The Voice. "Over" later debuted at #53 on the Billboard Hot Country Songs on 19 May, 2012.^[13]

Blake Shelton is a coach on the NBC show The Voice. His finalist Dia Frampton came in second. He returned for a second season of the show with Jermaine Paul as the champion ^[14]. Frampton also recorded a track with Shelton titled "I Will" on her album Red, released December 6, 2011.

Blake Shelton took the stage with Miranda Lambert at Super Bowl XLVI in February 2012 to open the event by singing a duet version of “America the Beautiful”. It marked their first TV performance since their headlining nuptials in May 2011.

Television[link]

[edit] 2011–present: The Voice

In 2011, Shelton became a coach on NBC's new show, The Voice, along with Christina Aguilera, Cee Lo Green and Adam Levine.

In season 1, Shelton became the runner-up coach as his final team member, Dia Frampton, won second place of the season.

In season 2, Shelton became the winning coach as his final team member, Jermaine Paul, was declared winner of the season.

Season 3 of The Voice will air in the fall of 2012. All four judges have signed on to return. ^[15]

Personal life[link]

Shelton married longtime girlfriend Kaynette Gern on November 17, 2003. They divorced in 2006.

Shelton is now married to fellow country singer Miranda Lambert, whom he met at the CMT's 100 Greatest Duets Concert.^[16] Lambert also sang background vocals on his cover of Michael Bublé's song, "Home", which became a Number 1 single on the Hot Country Songs chart. On May 9, 2010, after dating for four years,^[17] Shelton proposed to Miranda, after seeking (and receiving) her father's blessing.^[17][18]

Shelton and Lambert were married on May 14, 2011 at Don Strange Ranch in Boerne, Texas.^[17] 550 people were in attendance, including many celebrities, such as Reba McEntire, Cee Lo Green, Martina McBride, Kelly Clarkson, Dierks Bentley, Charles Kelley, and the Bellamy Brothers. After the wedding, Shelton expressed his joy stating, "I wanted to be married to Miranda the day I proposed. Finally, it's official!"^[19] Blake and Miranda reside in the small town of Tishomingo, OK, located in the Texoma area.

On January 17, 2012, Shelton's father, Dick, 71, died in Oklahoma, following a period of declining health.^[20][21]

Discography[link]

Studio albums

• Blake Shelton (2001)
• The Dreamer (2003)
• Blake Shelton's Barn & Grill (2004)
• Pure BS (2007)
• Startin' Fires (2008)
• Red River Blue (2011)

Extended plays

• Blake Shelton: Collector's Edition (2008)
• Hillbilly Bone (2010)
• All About Tonight (2010)

Compilation albums

• Blake Shelton – The Essentials (2009)
• Loaded: The Best of Blake Shelton (2010)

Awards and nominations[link]

References[link]

External links[link]

• Official Website
• Blake Shelton at the Internet Movie Database